Guatemalan forest synthesis after Pleistocene aridity

Sediments from two lakes in the Peten Department, Guatemala, provide palynological evidence from Central America of late Pleistocene aridity and subsequent synthesis of mesic forests. Late Glacial vegetation consisted of marsh, savanna, and juniper scrub. An early Holocene temperate forest preceded a mesic tropical forest with Brosimum (ramon). Thus “primeval” rain forests of Guatemala are no older than 10,000 to 11,000 years and are considerably younger in the Peten due to Mayan disturbances. Among dated Neotropical sites, the Peten has the most mesic vegetation yet shown to have supplanted xeric vegetation present during the Pleistocene. The arid late Glacial-humid early Holocene transition appears to have been pantropical in the lowlands. The Peten was not a Pleistocene refugium for mesophytic taxa, as has been suggested. Thus genesis of extant rain forests in northern Central America and southern Mexico remains unexplained.     

Late quaternary extinction of a tree species in eastern North America

Widespread species- and genus-level extinctions of mammals in North America and Europe occurred during the last deglaciation [16,000-9,000 yr B.P. (by (14)C)], a period of rapid and often abrupt climatic and vegetational change. These extinctions are variously ascribed to environmental change and overkill by human hunters. By contrast, plant extinctions since the Middle Pleistocene are undocumented, suggesting that plant species have been able to respond to environmental changes of the past several glacial/interglacial cycles by migration. We provide evidence from morphological studies of fossil cones and anatomical studies of fossil needles that a now-extinct species of spruce (Picea critchfieldii sp. nov.) was widespread in eastern North America during the Last Glacial Maximum. P. critchfieldii was dominant in vegetation of the Lower Mississippi Valley, and extended at least as far east as western Georgia. P. critchfieldii disappeared during the last deglaciation, and its extinction is not directly attributable to human activities. Similarly widespread plant species may be at risk of extinction in the face of future climate change.     

Abrupt tropical climate change: past and present

Three lines of evidence for abrupt tropical climate change, both past and present, are presented. First, annually and decadally averaged delta(18)O and net mass-balance histories for the last 400 and 2,000 yr, respectively, demonstrate that the current warming at high elevations in the mid- to low latitudes is unprecedented for at least the last 2 millennia. Second, the continuing retreat of most mid- to low-latitude glaciers, many having persisted for thousands of years, signals a recent and abrupt change in the Earth's climate system. Finally, rooted, soft-bodied wetland plants, now exposed along the margins as the Quelccaya ice cap (Peru) retreats, have been radiocarbon dated and, when coupled with other widespread proxy evidence, provide strong evidence for an abrupt mid-Holocene climate event that marked the transition from early Holocene (pre-5,000-yr-B.P.) conditions to cooler, late Holocene (post-5,000-yr-B.P.) conditions. This abrupt event, approximately 5,200 yr ago, was widespread and spatially coherent through much of the tropics and was coincident with structural changes in several civilizations. These three lines of evidence argue that the present warming and associated glacier retreat are unprecedented in some areas for at least 5,200 yr. The ongoing global-scale, rapid retreat of mountain glaciers is not only contributing to global sea-level rise but also threatening freshwater supplies in many of the world's most populous regions.     

Nonlinear response of summer temperature to Holocene insolation forcing in Alaska

Regional climate responses to large-scale forcings, such as precessional changes in solar irradiation and increases in anthropogenic greenhouse gases, may be nonlinear as a result of complex interactions among earth system components. Such nonlinear behaviors constitute a major source of climate “surprises” with important socioeconomic and ecological implications. Paleorecords are key for elucidating patterns and mechanisms of nonlinear responses to radiative forcing, but their utility has been greatly limited by the paucity of quantitative temperature reconstructions. Here we present Holocene July temperature reconstructions on the basis of midge analysis of sediment cores from three Alaskan lakes. Results show that summer temperatures during 10,000–5,500 calibrated years (cal) B.P. were generally lower than modern and that peak summer temperatures around 5,000 were followed by a decreasing trend toward the present. These patterns stand in stark contrast with the trend of precessional insolation, which decreased by ∼10% from 10,000 y ago to the present. Cool summers before 5,500 cal B.P. coincided with extensive summer ice cover in the western Arctic Ocean, persistence of a positive phase of the Arctic Oscillation, predominantly La Niña-like conditions, and variation in the position of the Alaskan treeline. These results illustrate nonlinear responses of summer temperatures to Holocene insolation radiative forcing in the Alaskan sub-Arctic, possibly because of state changes in the Arctic Oscillation and El Niño-Southern Oscillation and associated land–atmosphere–ocean feedbacks.     

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.     

Climate change, adaptive cycles, and the persistence of foraging economies during the late Pleistocene/Holocene transition in the Levant

Climatic forcing during the Younger Dryas (～12.9-11.5 ky B.P.) event has become the theoretical basis to explain the origins of agricultural lifestyles in the Levant by suggesting a failure of foraging societies to adjust. This explanation however, does not fit the scarcity of data for predomestication cultivation in the Natufian Period. The resilience of Younger Dryas foragers is better illustrated by a concept of adaptive cycles within a theory of adaptive change (resilience theory). Such cycles consist of four phases: release/collapse (Ω); reorganization (α), when the system restructures itself after a catastrophic stimulus through innovation and social memory--a period of greater resilience and less vulnerability; exploitation (r); and conservation (K), representing an increasingly rigid system that loses flexibility to change. The Kebarans and Late Natufians had similar responses to cold and dry conditions vs. Early Natufians and the Pre-Pottery Neolithic A responses to warm and wet climates. Kebarans and Late Natufians (α-phase) shifted to a broader-based diet and increased their mobility. Early Natufian and Pre-Pottery Neolithic A populations (r- and K-phases) had a growing investment in more narrowly focused, high-yield plant resources, but they maintained the broad range of hunted animals because of increased sedentism. These human adaptive cycles interlocked with plant and animal cycles. Forest and grassland vegetation responded to late Pleistocene and early Holocene climatic fluctuations, but prey animal cycles reflected the impact of human hunting pressure. The combination of these three adaptive cycles results in a model of human adaptation, showing potential for great sustainability of Levantine foraging systems even under adverse climatic conditions.     

The protracted Holocene extinction of California's flightless sea duck (Chendytes lawi) and its implications for the Pleistocene overkill hypothesis

Bones of the flightless sea duck (Chendytes lawi) from 14 archaeological sites along the California coast indicate that humans hunted the species for at least 8,000 years before it was driven to extinction. Direct (14)C dates on Chendytes bones show that the duck was exploited on the southern California islands as early as approximately 11,150-10,280 calendar years B.P., and on the mainland by at least 8,500 calendar years B.P. The youngest direct date of 2,720-2,350 calendar years B.P., combined with the absence of Chendytes bones from hundreds of late Holocene sites, suggests that the species was extinct by approximately 2,400 years ago. Although the extinction of Chendytes clearly resulted from human overhunting, its demise raises questions about the Pleistocene overkill model, which suggests that megafauna were driven to extinction in a blitzkrieg fashion by Native Americans approximately 13,000 years ago. That the extermination of Chendytes was so protracted and archaeologically visible suggests that, if the terminal Pleistocene megafauna extinctions were primarily the result of human exploitation, there should also be a long and readily detectable archaeological record of their demise. The brief window now attributed to the Clovis culture ( approximately 13,300-12,900 B.P.) seems inconsistent with an overhunting event.     

Tracing the effects of the Little Ice Age in the tropical lowlands of eastern Mesoamerica

The causes of late-Holocene centennial to millennial scale climatic variability and the impact that such variability had on tropical ecosystems are still poorly understood. Here, we present a high-resolution, multiproxy record from lowland eastern Mesoamerica, studied to reconstruct climate and vegetation history during the last 2,000 years, in particular to evaluate the response of tropical vegetation to the cooling event of the Little Ice Age (LIA). Our data provide evidence that the densest tropical forest cover and the deepest lake of the last two millennia were coeval with the LIA, with two deep lake phases that follow the Spörer and Maunder minima in solar activity. The high tropical pollen accumulation rates limit LIA''s winter cooling to a maximum of 2°C. Tropical vegetation expansion during the LIA is best explained by a reduction in the extent of the dry season as a consequence of increased meridional flow leading to higher winter precipitation. These results highlight the importance of seasonal responses to climatic variability, a factor that could be of relevance when evaluating the impact of recent climate change.     

Evolution in coyotes (Canis latrans) in response to the megafaunal extinctions

Living coyotes modify their behavior in the presence of larger carnivores, such as wolves. However, little is known about the effects of competitor presence or absence on morphological change in coyotes or wolves over long periods of time. We examined the evolution of coyotes and wolves through time from the late Pleistocene, during which many large carnivorous species coexisted as predators and competitors, to the Recent; this allowed us to investigate evolutionary changes in these species in response to climate change and megafaunal extinctions at the end of the Pleistocene. We measured postcranial skeletal morphologies of wolves (Canis lupus) and coyotes (C. latrans) from Pleistocene-aged tar deposits, as well as early, mid, and recent Holocene populations of both. We found few morphological differences between Pleistocene and Holocene wolf populations. Conversely, we found many differences in coyotes: Pleistocene coyotes were larger and more robust than Holocene populations. However, within 1,000 y of the megafaunal extinctions, coyotes are morphologically indistinguishable from modern populations. We cannot attribute these differences directly to climate change because modern coyotes do not follow Bergmann's rule, which states body size increases with decreasing temperature. Instead, we suggest that Pleistocene coyotes may have been larger and more robust in response to larger competitors and a larger-bodied prey base. Although we cannot separate competition from predator-prey interactions, this study indicates that the effects of biotic interactions can be detected in the fossil record.     

Geophysical, archaeological, and historical evidence support a solar-output model for climate change

Although the processes of climate change are not completely understood, an important causal candidate is variation in total solar output. Reported cycles in various climate-proxy data show a tendency to emulate a fundamental harmonic sequence of a basic solar-cycle length (11 years) multiplied by 2(N) (where N equals a positive or negative integer). A simple additive model for total solar-output variations was developed by superimposing a progression of fundamental harmonic cycles with slightly increasing amplitudes. The timeline of the model was calibrated to the Pleistocene/Holocene boundary at 9,000 years before present. The calibrated model was compared with geophysical, archaeological, and historical evidence of warm or cold climates during the Holocene. The evidence of periods of several centuries of cooler climates worldwide called "little ice ages," similar to the period anno Domini (A.D.) 1280-1860 and reoccurring approximately every 1,300 years, corresponds well with fluctuations in modeled solar output. A more detailed examination of the climate sensitive history of the last 1, 000 years further supports the model. Extrapolation of the model into the future suggests a gradual cooling during the next few centuries with intermittent minor warmups and a return to near little-ice-age conditions within the next 500 years. This cool period then may be followed approximately 1,500 years from now by a return to altithermal conditions similar to the previous Holocene Maximum.     

Ecological consequences of early Late Pleistocene megadroughts in tropical Africa

Extremely arid conditions in tropical Africa occurred in several discrete episodes between 135 and 90 ka, as demonstrated by lake core and seismic records from multiple basins [Scholz CA, Johnson TC, Cohen AS, King JW, Peck J, Overpeck JT, Talbot MR, Brown ET, Kalindekafe L, Amoako PYO, et al. (2007) Proc Natl Acad Sci USA 104:16416-16421]. This resulted in extraordinarily low lake levels, even in Africa's deepest lakes. On the basis of well dated paleoecological records from Lake Malawi, which reflect both local and regional conditions, we show that this aridity had severe consequences for terrestrial and aquatic ecosystems. During the most arid phase, there was extremely low pollen production and limited charred-particle deposition, indicating insufficient vegetation to maintain substantial fires, and the Lake Malawi watershed experienced cool, semidesert conditions (<400 mm/yr precipitation). Fossil and sedimentological data show that Lake Malawi itself, currently 706 m deep, was reduced to an approximately 125 m deep saline, alkaline, well mixed lake. This episode of aridity was far more extreme than any experienced in the Afrotropics during the Last Glacial Maximum (approximately 35-15 ka). Aridity diminished after 95 ka, lake levels rose erratically, and salinity/alkalinity declined, reaching near-modern conditions after 60 ka. This record of lake levels and changing limnological conditions provides a framework for interpreting the evolution of the Lake Malawi fish and invertebrate species flocks. Moreover, this record, coupled with other regional records of early Late Pleistocene aridity, places new constraints on models of Afrotropical biogeographic refugia and early modern human population expansion into and out of tropical Africa.     

Native Americans experienced a strong population bottleneck coincident with European contact

The genetic and demographic impact of European contact with Native Americans has remained unclear despite recent interest. Whereas archeological and historical records indicate that European contact resulted in widespread mortality from various sources, genetic studies have found little evidence of a recent contraction in Native American population size. In this study we use a large dataset including both ancient and contemporary mitochondrial DNA to construct a high-resolution portrait of the Holocene and late Pleistocene population size of indigenous Americans. Our reconstruction suggests that Native Americans suffered a significant, although transient, contraction in population size some 500 y before the present, during which female effective size was reduced by ～50%. These results support analyses of historical records indicating that European colonization induced widespread mortality among indigenous Americans.     

Two terrestrial records of rapid climatic change during the glacial-Holocene transition (14,000- 9,000 calendar years B.P.) from Europe

Two independent multidisciplinary studies of climatic change during the glacial-Holocene transition (ca. 14,000-9,000 calendar yr B.P.) from Norway and Switzerland have assessed organism responses to the rapid climatic changes and made quantitative temperature reconstructions with modern calibration data sets (transfer functions). Chronology at Krakenes, western Norway, was derived from calibration of a high-resolution series of 14C dates. Chronologies at Gerzensee and Leysin, Switzerland, were derived by comparison of delta18O in lake carbonates with the delta18O record from the Greenland Ice Core Project. Both studies demonstrate the sensitivity of terrestrial and aquatic organisms to rapid temperature changes and their value for quantitative reconstruction of the magnitudes and rates of the climatic changes. The rates in these two terrestrial records are comparable to those in Greenland ice cores, but the actual temperatures inferred apply to the terrestrial environments of the two regions.     

Vertebrate community on an ice-age Caribbean island

We report 95 vertebrate taxa (13 fishes, 11 reptiles, 63 birds, 8 mammals) from late Pleistocene bone deposits in Sawmill Sink, Abaco, The Bahamas. The >5,000 fossils were recovered by scuba divers on ledges at depths of 27–35 m below sea level. Of the 95 species, 39 (41%) no longer occur on Abaco (4 reptiles, 31 birds, 4 mammals). We estimate that 17 of the 39 losses (all of them birds) are linked to changes during the Pleistocene–Holocene Transition (PHT) (∼15–9 ka) in climate (becoming more warm and moist), habitat (expansion of broadleaf forest at the expense of pine woodland), sea level (rising from −80 m to nearly modern levels), and island area (receding from ∼17,000 km² to 1,214 km²). The remaining 22 losses likely are related to the presence of humans on Abaco for the past 1,000 y. Thus, the late Holocene arrival of people probably depleted more populations than the dramatic physical and biological changes associated with the PHT.Interpreting the late Quaternary vertebrate fossil record on West Indian islands has been limited by the vague chronological resolution of most sites. Insular fossil faunas often have been assumed to be Pleistocene (rather than Holocene) in age without direct radiometric or other nonfaunal evidence such as sea level (1–3) (see Site Setting). This practice has been widespread with noncultural (paleontological) sites bearing extinct species of mammals, in part because extinct late Quaternary mammals from nearby North America (ground sloths, sabertooth cats, proboscideans, horses, camels, etc.) are indeed from Pleistocene contexts (4). Through direct radiocarbon (¹⁴C) dating using accelerator-mass spectrometer (AMS) technology, we now know that at least some of the large, extinct West Indian mammals, such as sloths, survived well into the Holocene (5–7). Although AMS ¹⁴C dates on insular sloth fossils range from the mid-Holocene to the late Pleistocene (7–9), all successful AMS ¹⁴C dates done thus far on extirpated West Indian reptiles, birds, or micromammals (bats) are Holocene rather than Pleistocene (10–12).Developing a sound chronology from cultural (archaeological) sites is often facilitated by ¹⁴C dating charcoal, by stratigraphic association of the bones with temporally diagnostic ceramic or lithic artifacts, or by AMS ¹⁴C dating the identified bones (e.g., refs. 13–15). In all such cases, the cultural sites on Caribbean islands are found to be mid-Holocene to late Holocene in age, not late Pleistocene. For Bahamian islands in particular, human arrival took place only about a millennium ago (∼1 ka) (16, 17).A West Indian fossil vertebrate community (38 taxa of reptiles, birds, and mammals) assigned to the Pleistocene rather than the Holocene was reported from the underwater Owl Roost deposits in Sawmill Sink, Abaco, The Bahamas (18). Our subsequent field and laboratory research at this flooded sinkhole has more than doubled the Pleistocene fauna to 95 species, by far the richest and most taxonomically diverse set of vertebrate fossils from the West Indies.Here, we describe this mainly predator-accumulated fossil assemblage, which was deposited in glacial times when sea level was much lower than today. We focus on the late Pleistocene Owl Roost fossils rather than the younger fossils from the Sawmill Sink peat deposit, which have been AMS ¹⁴C dated to the mid-Holocene to late Holocene (18, 19). The Pleistocene fossils allow us to evaluate which species were able to withstand the major changes in climate, sea level, land area, and habitat during the glacial–interglacial transition. These data thus provide long-term context for projecting how future climate change might affect West Indian biodiversity.     

Population genetics of ice age brown bears

The Pleistocene was a dynamic period for Holarctic mammal species, complicated by episodes of glaciation, local extinctions, and intercontinental migration. The genetic consequences of these events are difficult to resolve from the study of present-day populations. To provide a direct view of population genetics in the late Pleistocene, we measured mitochondrial DNA sequence variation in seven permafrost-preserved brown bear (Ursus arctos) specimens, dated from 14,000 to 42,000 years ago. Approximately 36,000 years ago, the Beringian brown bear population had a higher genetic diversity than any extant North American population, but by 15,000 years ago genetic diversity appears similar to the modern day. The older, genetically diverse, Beringian population contained sequences from three clades now restricted to local regions within North America, indicating that current phylogeographic patterns may provide misleading data for evolutionary studies and conservation management. The late Pleistocene phylogeographic data also indicate possible colonization routes to areas south of the Cordilleran ice sheet.     

Middle East coastal ecosystem response to middle-to-late Holocene abrupt climate changes

The Holocene vegetation history of the northern coastal Arabian Peninsula is of long-standing interest, as this Mediterranean/semiarid/arid region is known to be particularly sensitive to climatic changes. Detailed palynological data from an 800-cm alluvial sequence cored in the Jableh plain in northwest Syria have been used to reconstruct the vegetation dynamics in the coastal lowlands and the nearby Jabal an Nuşayriyah mountains for the period 2150 to 550 B.C. Corresponding with the 4.2 to 3.9 and 3.5 to 2.5 cal kyr BP abrupt climate changes (ACCs), two large-scale shifts to a more arid climate have been recorded. These two ACCs had different impacts on the vegetation assemblages in coastal Syria. The 3.5 to 2.5 cal kyr BP ACC is drier and lasted longer than the 4.2 to 3.9 cal kyr BP ACC, and is characterized by the development of a warm steppe pollen-derived biome (1100–800 B.C.) and a peak of hot desert pollen-derived biome at 900 B.C. The 4.2 to 3.9 cal kyr BP ACC is characterized by a xerophytic woods and shrubs pollen-derived biome ca. 2050 B.C. The impact of the 3.5 to 2.5 cal kyr BP ACC on human occupation and cultural development is important along the Syrian coast with the destruction of Ugarit and the collapse of the Ugarit kingdom at ca. 1190 to 1185 B.C.     

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.     

Thieving rodents as substitute dispersers of megafaunal seeds

The Neotropics have many plant species that seem to be adapted for seed dispersal by megafauna that went extinct in the late Pleistocene. Given the crucial importance of seed dispersal for plant persistence, it remains a mystery how these plants have survived more than 10,000 y without their mutualist dispersers. Here we present support for the hypothesis that secondary seed dispersal by scatter-hoarding rodents has facilitated the persistence of these large-seeded species. We used miniature radio transmitters to track the dispersal of reputedly megafaunal seeds by Central American agoutis, which scatter-hoard seeds in shallow caches in the soil throughout the forest. We found that seeds were initially cached at mostly short distances and then quickly dug up again. However, rather than eating the recovered seeds, agoutis continued to move and recache the seeds, up to 36 times. Agoutis dispersed an estimated 35% of seeds for >100 m. An estimated 14% of the cached seeds survived to the next year, when a new fruit crop became available to the rodents. Serial video-monitoring of cached seeds revealed that the stepwise dispersal was caused by agoutis repeatedly stealing and recaching each other's buried seeds. Although previous studies suggest that rodents are poor dispersers, we demonstrate that communities of rodents can in fact provide highly effective long-distance seed dispersal. Our findings suggest that thieving scatter-hoarding rodents could substitute for extinct megafaunal seed dispersers of tropical large-seeded trees.     

Synchronous interhemispheric Holocene climate trends in the tropical Andes

Holocene variations of tropical moisture balance have been ascribed to orbitally forced changes in solar insolation. If this model is correct, millennial-scale climate evolution should be antiphased between the northern and southern hemispheres, producing humid intervals in one hemisphere matched to aridity in the other. Here we show that Holocene climate trends were largely synchronous and in the same direction in the northern and southern hemisphere outer-tropical Andes, providing little support for the dominant role of insolation forcing in these regions. Today, sea-surface temperatures in the equatorial Pacific Ocean modulate rainfall variability in the outer tropical Andes of both hemispheres, and we suggest that this mechanism was pervasive throughout the Holocene. Our findings imply that oceanic forcing plays a larger role in regional South American climate than previously suspected, and that Pacific sea-surface temperatures have the capacity to induce abrupt and sustained shifts in Andean climate.     

Genetic footprints of demographic expansion in North America,but not Amazonia,during the Late Quaternary

The biotic consequences of climate change have attracted considerable attention. In particular, the "refugial debate" centers on the possible retraction of habitats to limited areas that may have served as refuges for many associated species, especially during glaciations of the Quaternary. One prediction of such scenarios is that populations must have experienced substantial growth accompanying climatic amelioration and the occupation of newly expanded habitats. We used coalescence theory to examine the genetic evidence, or lack thereof, for late Pleistocene refugia of boreal North American and tropical Amazonian mammals. We found substantial and concordant evidence of demographic expansion in North American mammals, particularly at higher latitudes. In contrast, small mammals from western Amazonia appear to have experienced limited or no demographic expansion after the Late Pleistocene. Thus, demographic responses to climate change can be tracked genetically and appear to vary substantially across the latitudinal gradient of biotic diversity.