Last straw versus Blitzkrieg overkill: Climate-driven changes in the Arctic Siberian mammoth population and the Late Pleistocene extinction problem

A set of radiocarbon dates on woolly mammoth were obtained from several regions of Arctic Siberia: the New Siberian Islands (n = 68), north of the Yana-Indigirka Lowland (n = 43), and the Taimyr Peninsula (n = 18). Based on these and earlier published dates (n = 201) from the East Arctic, a comparative analysis of the time-related density distribution of ¹⁴C dates was conducted. It was shown that the frequencies of ¹⁴C dates under certain conditions reflect temporal fluctuations in mammoth numbers. At the end of the Pleistocene the number of mammoths in the East Arctic changed in a cyclic manner in keeping with a general “Milankovitch-like” trend. The fluctuations in numbers at the end of the Pleistocene occurred synchronously with paleoenvironmental changes controlled by global climatic change. There were three minima of relative mammoth numbers during the last 50 000 years: 22 000, 14 500–19 000, and 9500 radiocarbon years ago, or around 26 000, 16–20 000, and 10 500 calendar years respectively. The last mammoths lived on the New Siberian Islands, which were connected to the continent at that time, 9470 ± 40 radiocarbon years ago (10 700 ± 70 calendar years BP). This new youngest date approximates the extinction time of mammoths in the last continental refugium of the Holarctic. The adverse combination of environmental parameters was apparently a major factor in the critical reduction in mammoth numbers. The dispersal of humans into the Arctic areas of Siberia no later than 28 000 radiocarbon years ago did not overtly influence animal numbers. Humans were not responsible for the destruction of a sustainable mammoth population. The expanding human population could have become fatal to mammoths during strong the minima of their numbers, one of which occurred at the very beginning of the Holocene.     

Woolly mammoth (Mammuthus primigenius Blum.) and its environment in northern Europe during the last glaciation

Woolly mammoths were large, herbivorous, cold-adapted mammals of the Late Pleistocene. The diet and habitat requirements of the species set certain constraints on the palaeoenvironments it could occupy. The relationship between the mammoth’s shifting range and changing environments can be explored using independent data on ice sheet configuration, temperature, and vegetation, provided the locality and age of the fossil remains can be validated. Here we present a comprehensive record of occurrence of the woolly mammoth in the circum-Baltic region of northern Europe during the last glaciation, based on a compilation of radiocarbon-dated remains. The record shows that the mammoth was widespread in northern and north-eastern Europe during Marine Isotope Stage 3 (MIS 3), at 50,000–30,000 calibrated years ago (50–30 ka). The presence of the species up to 65°N latitude supports the restriction of the Scandinavian Ice Sheet (SIS) during MIS 3. The widest distribution range round 30 ka was followed by a decline that led to the disappearance of mammoths from the area during the maximum extent of the SIS, from 22 to 18 ka. The woolly mammoth re-colonized the Baltic region and southern Scandinavia after the onset of the late-glacial deglaciation at 17 ka. The late-glacial record suggests a markedly fluctuating population changing its range in tune with the rapid environmental changes. The last appearance of mammoth in our study region was in Estonia during the Younger Dryas (Greenland Stadial 1; GS1) at about 12 ka. The two major periods of occurrence during MIS 3 and the late-glacial stadial suggest that mammoth had a wide tolerance of open to semi-open tundra and steppe-tundra habitats with intermediately cold climate, whereas the 22–18 ka disappearance suggests a major southward and/or eastward retreat in response to extremely cold, glacial conditions near the SIS margin. The final regional extinction correlates with the re-forestation during the rapid warming at the Younger Dryas–Holocene boundary.     

Collection of radiocarbon dates on the mammoths (Mammuthus primigenius) and other genera of Wrangel Island, northeast Siberia, Russia

We present and discuss a full list of radiocarbon dates for woolly mammoth and other species of the Mammoth fauna available from Wrangel Island, northeast Siberia, Russia. Most of the radiocarbon dates are published here for the first time. Of the124 radiocarbon dates on mammoth bone, 106 fall between 3700 and 9000 yr ago. We believe these dates bracket the period of mammoth isolation on Wrangel Island and their ultimate extinction, which we attribute to natural causes. The absence of dates between 9–12 ka probably indicates a period when mammoths were absent from Wrangel Island. Long bone dimensions of Holocene mammoths from Wrangel Island indicate that these animals were comparable in size to those on the mainland; although they were not large animals, neither can they be classified as dwarfs. Occurrence of mammoth Holocene refugia on the mainland is suggested. Based on other species of the Mammoth fauna that have also been radiocarbon on Wrangel Island, including horse, bison, musk ox and woolly rhinoceros, it appears that the mammoth was the only species of that fauna that inhabited Wrangel Island in the mid-Holocene.     

Taphonomy of two last glacial maximum mammoth sites in the central Great Plains of North America: A preliminary report on La Sena and Lovewell

Two mammoth sites from the central Great Plains of North America, each containing one adult Columbian mammoth (Mammuthus columbi), were excavated from Last Glacial Maximum (LGM) loess and fine-grained alluvial deposits, respectively. Taphonomic data from both sites indicate that the mammoth skeletons exhibit numerous spirally fractured limb elements. Dynamic loading points are present on midshafts of large limb bones. Bone flakes produced from the partial thickness of thick cortical bone are also present. Hypotheses of carnivore activity, mammoth trampling, and human-induced fracturing are evaluated as possible causes of the fractured limb bone. Testing the hypotheses using modern data from actualistic taphonomic studies of elephant skeletons, paleontological data from two proboscidean natural death sites, experimental data from elephant bone fracturing, and archaeologically derived data concerning late Pleistocene human modification of mammoth limb bone indicates that the first two hypotheses can be rejected, while the third hypothesis is supported.     

The taphonomy of mammoth localities in southeastern Wisconsin (USA)

Four southeastern Wisconsin mammoth localities located within a glacial landscape had well-preserved remains found in inter-morainal depressions filled with lacustrine clays covered with peat. Numerous radiocarbon ages dated the mammoths to the late Pleistocene. The taphonomic analysis focused on determining the agency or agencies involved in site formation and the agency or agencies involved in modification to the bones. Statistical approaches to bone orientation data underscored that water transport was not a factor in the formation of the bone beds nor was water movement a disturbance factor. The bone modification profile of the overall assemblage was dominated by chemical weathering, followed by other chemical and microbiological processes. Root etching was the most common biological modification, and the most frequent modifications are involved with the microenvironments of the surface to burial substrate. Potentially cultural modification accounted for only a small portion of the cortical damage to the bones. This general profile indicated an assemblage far more influenced by the immediate environment than by passing animals or people.     

Calibration of mammoth (Mammuthus) dispersal into North America using rare earth elements of Plio-Pleistocene mammals from Florida

The first appearance of mammoth (Mammuthus) is currently used to define the beginning of the Irvingtonian North American Land Mammal Age at about 1.4 Ma. Thereafter, mammoth fossils are common and widespread in North America until the end of the Pleistocene. In contrast to this generally accepted biochronology, recent reports have asserted that mammoth occurs in late Pliocene (ca. 2.5 Ma) alluvium from the Santa Fe River of northern Florida. The supposedly contemporaneous late Pliocene fossil assemblage from the Santa Fe River that produced the mammoth specimens actually consists of a mixture of diagnostic Blancan (late Pliocene) and late Rancholabrean (latest Pleistocene) species. Fossil bones and teeth of the two mammalian faunas mixed together along the Santa Fe River have significantly different rare earth element (REE) signatures. The REE signatures of mammoth are indistinguishable from those of Rancholabrean mammals, yet they are different from those of diagnostic Blancan vertebrates from these same temporally mixed faunas of the Santa Fe River. Thus, no evidence for late Pliocene mammoth exists in Florida, and mammoth fossils remain reliable biochronological indicators for Irvingtonian and Rancholabrean terrestrial sequences throughout mid- and lower-latitude North America.     

Mycological evidence of coprophagy from the feces of an Alaskan Late Glacial mammoth

Dung from a mammoth was preserved under frozen conditions in Alaska. The mammoth lived during the early part of the Late Glacial interstadial (ca 12,300 BP). Microfossils, macroremains and ancient DNA from the dung were studied and the chemical composition was determined to reconstruct both the paleoenvironment and paleobiology of this mammoth. Pollen spectra are dominated by Poaceae, Artemisia and other light-demanding taxa, indicating an open, treeless landscape (‘mammoth steppe’). Fruits and seeds support this conclusion. The dung consists mainly of cyperaceous stems and leaves, with a minor component of vegetative remains of Poaceae. Analyses of fragments of the plastid rbcL gene and trnL intron and nrITS1 region, amplified from DNA extracted from the dung, supplemented the microscopic identifications. Many fruit bodies with ascospores of the coprophilous fungus Podospora conica were found inside the dung ball, indicating that the mammoth had eaten dung. The absence of bile acids points to mammoth dung. This is the second time that evidence for coprophagy of mammoths has been derived from the presence of fruit bodies of coprophilous fungi in frozen dung. Coprophagy might well have been a common habit of mammoths. Therefore, we strongly recommend that particular attention should be given to fungal remains in future fossil dung studies.     

The first radiocarbon data of bone remains of mammoth faunal forms in northwestern Russia

Unlike in the neighboring territories, the distribution and the period of habitation of late Pleistocene mammoth complex animals in the northwestern area of Russia had not been studied until recently. This article fills in this gap using the bone material from the Zoological Institute of the Russian Academy of Sciences and the collections of one of the authors. The samples of 14 bones and teeth of big mammals uncovered in different places of the region were dated. The data obtained by conventional ¹⁴C method and AMS method agree with each other and make it possible to determine two periods of habitation of mammoth complex animals in the region: 39 000–23 000 years ago and 13 000–9800 years ago, which confirms that ice-free landscapes existed here at these time intervals.     

The U.P. Mammoth Site,Carbon County,Wyoming, USA: More Questions than Answers

In the summer of 1960, mammoth bones were discovered by a dragline operator in southern Wyoming at the Union Pacific (U.P.) Mammoth site. Although subsequent archaeological work during 1960 and 1961 identified artifacts in association with the mammoth remains, many authors have since questioned the nature of that association. Also, little has been published about the site other than a brief article in National Geographic Magazine in 1962. In this paper, we present additional information on the U.P. Mammoth site including stratigraphic profiles from the first author's geoarchaeological work in 1961, stratigraphic and spatial location of bones and artifacts derived from the original field notes, and new radiocarbon dates. Although the precise stratigraphic provenance for many artifacts and skeletal elements remains unclear, a compelling argument can be made for spatial and stratigraphic association of the mammoth remains with the artifact assemblage suggesting some kind of human interaction with the animal.     

Palaeo-environmental and dietary analysis of intestinal contents of a mammoth calf (Yamal Peninsula,northwest Siberia)

Intestinal samples from the one-month-old Siberian mammoth calf ‘Lyuba’ were studied using light microscopy and ancient DNA to reconstruct its palaeo-environment and diet. The palynological record indicates a ‘mammoth steppe’. At least some pollen of arboreal taxa was reworked, and thus the presence of trees on the landscape is uncertain. In addition to visual comparison of 11 microfossil spectra, a PCA analysis contributed to diet reconstruction. This yielded two clusters: one of samples from the small intestine and the other of large-intestine samples, indicating compositional differences in food remains along the intestinal tract, possibly reflecting different episodes of ingestion. Based on observed morphological damage we conclude that the cyperaceous plant remains and some remains of dwarf willows were originally eaten by a mature mammoth, most likely Lyuba’s mother. The mammoth calf probably unintentionally swallowed well-preserved mosses and mineral particles while eating fecal material deposited on a soil surface covered with mosses. Coprophagy may have been a common habit for mammoths, and we therefore propose that fecal material should not be used to infer season of death of mammoths. DNA sequences of trnL and rbcL genes amplified from ancient DNA extracted from intestinal samples confirmed and supplemented plant identifications based on microfossils and macro-remains. Results from different extraction methods and barcoding markers complemented each other and show the value of longer protocols in addition to fast and commercially available extraction kits.     

Patterns of faunal extinction and paleoclimatic change from mid-Holocene mammoth and polar bear remains, Pribilof Islands, Alaska

Qagnaxˆ Cave, a lava tube cave on St. Paul Island in the Pribilofs, has recently produced a mid-Holocene vertebrate faunal assemblage including woolly mammoth, polar bear, caribou, and Arctic fox. Several dates on the mammoth remains converge on 5700 ¹⁴C yr BP. These dates, ~ 2300 yr younger than mammoth dates previously published from the Pribilof Islands, make these the youngest remains of proboscideans, and of non-extinct Quaternary megafauna, recovered from North America. Persistence of mammoths on the Pribilofs is most parsimoniously explained by the isolation of the Pribilofs and the lack of human presence in pre-Russian contact times, but an additional factor may have been the local existence of high-quality forage in the form of grasses enriched by nutrients derived from local Holocene tephras. This interpretation is reinforced by stable carbon and nitrogen isotope values obtained from the mammoth remains. The endpoint of mammoth survival in the Pribilofs is unknown, but maybe coterminous with the arrival of polar bears whose remains in the cave date to the Neoglacial cold period of ~ 4500 to 3500 ¹⁴C yr BP. The polar bear record corroborates a widespread cooling of the Bering Sea region at that time.     

Geographic and temporal trends in proboscidean and human radiocarbon histories during the late Pleistocene

The causes of large animal extinctions at the end of the Pleistocene remain a hotly debated topic focused primarily on the effects of human over hunting and climate change. Here we examine multiple, large radiocarbon data sets for humans and extinct proboscideans and explore how variation in their temporal and geographic distributions were related prior to proboscidean extinction. These data include 4532 archaeological determinations from Europe and Siberia and 1177 mammoth and mastodont determinations from Europe, Siberia, and North America. All span the period from 45,000 to 12,000 calendar years BP. We show that while the geographic ranges of dated human occupations and proboscidean remains overlap across the terminal Pleistocene of the Old World, the two groups remain largely segregated and increases in the frequency of human occupations do not coincide with declines in proboscidean remains. Prior to the Last Glacial Maximum (LGM; ca 21,000 years BP), archaeological ¹⁴C determinations increase slightly in frequency worldwide while the frequency of dated proboscidean remains varies depending on taxon and location. After the LGM, both sympatric and allopatric groups of humans and proboscideans increase sharply as climatic conditions ameliorate. Post-LGM radiocarbon frequencies among proboscideans peak at different times, also depending upon taxon and location. Woolly mammoths in Beringia reach a maximum and then decline beginning between 16,000 and 15,500 years BP, woolly mammoths in Europe and Siberia ca 14,500 and 13,500 BP, and Columbian mammoth and American mastodont only after 13,000 BP. Declines among woolly mammoths appear to coincide with the restructuring of biotic communities following the Pleistocene–Holocene transition.     

Population-level genotyping of coat colour polymorphism in woolly mammoth (Mammuthus primigenius)

Patterns in the spatial or temporal distribution of genotypes may be indicative of natural selection. Previous work on the woolly mammoth melanocortin-1 receptor (Mc1r) gene identified three polymorphic positions that suggest Pleistocene populations may have harboured both light- and dark-haired mammoths (Rompler et al., 2006, 313: 62). Here, we extend this work and present the first population-level analysis of a functional gene in an extinct species. We genotyped the Mc1r gene in 47 woolly mammoth samples excavated from sites across the central portion of the woolly mammoths’ former range to examine the extent of variation of this polymorphism through time and across space. Only one individual was found to be heterozygous, indicating that the frequency of the ‘light’ mutant allele was very low. We conclude that light-coloured woolly mammoths would have been very rare, and may even have been non-existent if the ‘light’ mutant allele was strongly selected against in its homozygotic form. With the increasing availability of large-scale sequencing technologies, population-level datasets capable of identifying local adaptation will become increasingly attainable.     

Canyon Creek: A late Pleistocene vertebrate locality in interior Alaska

The Canyon Creek vertebrate-fossil locality is an extensive road cut near Fairbanks that exposes sediments that range in age from early Wisconsin to late Holocene. Tanana River gravel at the base of the section evidently formed during the Delta Glaciation of the north-central Alaska Range. Younger layers and lenses of fluvial sand are interbedded with arkosic gravel from Canyon Creek that contains tephra as well as fossil bones of an interstadial fauna about 40,000 years old. Solifluction deposits containing ventifacts, wedge casts, and rodent burrows formed during a subsequent period of periglacial activity that took place during the maximum phase of Donnelly Glaciation about 25,000–17,000 years ago. Overlying sheets of eolian sand are separated by a 9500-year-old paleosol that may correlate with a phase of early Holocene spruce expansion through central Alaska. The Pleistocene fauna from Canyon Creek consists of rodents (indicated by burrows), Mammuthus primigenius (woolly mammoth), Equus lambei (Yukon wild ass), Camelops hesternus (western camel), Bison sp. cf. B. crassicornis (large-horned bison), Ovis sp. cf.O. dalli (mountain sheep), Canis sp. cf. C. lupus (wolf), Lepus sp. cf. L. othus or L. arcticus (tundra hare), and Rangifer sp. (caribou). This assemblage suggests an open landscape in which trees and tall shrubs were either absent or confined to sheltered and moist sites. Camelops evidently was present in eastern Beringia during the middle Wisconsin interstadial interval but may have disappeared during the following glacial episode. The stratigraphic section at Canyon Creek appears to demonstrate that the Delta Glaciation of the north-central Alaska Range is at least in part of early Wisconsin age and was separated from the succeeding Donnelly Glaciation by an interstadial rather than interglacial episode.     

Holes in the spinous processes of woolly mammoth vertebrae: spatial and temporal distribution,and the causes of pathology formation

The latest data on holes in the spinous processes of the vertebrae of woolly mammoths, a rare pathology, are presented. This was identified at 19 sites of northern Eurasia. Such destructive changes are recorded ca. 34–12k ¹⁴C a bp , and only two sites dated to >50k and >41k ¹⁴C a bp. The main hypotheses about hole formation are: vertebral abnormalities; bone infections; genetic traits; and unfavourable geochemical environment. The pathology occurred in mammoths of all age groups, and could have arisen at the embryonic stage. There are two types: classic holes associated with osteolytic changes; and very rarely tumour-like lesions. The most likely cause of the lesions is alimentary osteodystrophy caused by chronic mineral starvation. The aetiology of this disease is usually associated with a deficiency or excess of macro- and microelements in the geochemical landscape, and through forage and water this leads to a severe metabolic disorder. Analysis of palaeopathological data shows two waves of geochemical stress in animals, ca. 26–18k and ca. 16–12k ¹⁴C a bp. Therefore, the woolly mammoth extinction can be viewed as a non-linear function, with two peaks of high mortality corresponding to the Last Glacial Maximum and the Lateglacial.     

AMS 14C chronology of the world's southernmost woolly mammoth (Mammuthus primigenius Blum.)

The world's definite southernmost woolly mammoth record is a molar from Ji’nan (around 36°N), Shandong Province, China. AMS ¹⁴C dating of the specimen, gave a conventional ¹⁴C age of 33,150±250 BP. The period of 40–30 ka BP corresponds to the later phase of the Marine Oxygen Isotope Stage 3 (MIS 3a), recognized as the global interstadial of the last glacial period. However, it is known that the winter monsoon strengthened in Asia during the period 35–33 ka BP, and the age of the woolly mammoth specimen from Ji’nan corresponds to that age. The specimen suggests that this area became cold and dry in 33 ka BP, and grassland or open forest, suitable habitat for woolly mammoths, developed during this short time span. This age is similar to the age of the southernmost woolly mammoth record in Europe, therefore supporting a hypothesis by Porter and An [1995. Correlation between climate events in the North Atlantic and China during the last glaciation. Nature 375, 305–308] that an important component of Chinese palaeoclimates may be linked to changes in North Atlantic oceanic conditions.     

Mammoth landscapes: good country for hunter-gatherers

Proboscideans, because of their great size and unique behaviors, affect habitats in conspicuous ways, thus leaving information-rich signs about animal health, individual and group movements, and demography. Dung deposits, trail networks, rubbing sites, and excavations in elephant country afford abundant environmental clues to animal health and density. Prehistoric human groups able to subsist by opportunistic exploitation of mammoths or mastodons could have taken advantage of such information to be efficient foragers and rapid dispersers into new ranges. In addition, landscapes containing proboscideans have potential future fossil deposits containing skeletons of many animal taxa, isolated bone elements, fractured and flaked bones and tusks, and plant parts preserved due to proboscidean behavior.     

Chronology and new research on the Schaefer mammoth (?Mammuthus primigenius) site,Kenosha County,Wisconsin, USA

A summary of previous and new research on the Schaefer mammoth site is presented. The Schaefer site in extreme southeastern Wisconsin, USA was excavated in 1992 and 1993. Seventy-five percent of a mammoth ?Mammuthus primigenius, the woolly mammoth (but comparable as well with M. jeffersonii, the Jefferson mammoth) was recovered. Drifted wood specimens and plant macro fossils were also recovered from the site.Twenty-five specimens of wood have been identified as Picea sp., Picea/Larix sp., or unidentified hardwood. Macro fossils in the form of cones have been identified as black spruce (Picea mariana). These preliminary data do not appear to support the traditional environment inferred for the woolly mammoth and points to a need for study of the animal's attribution to species. The animal was a male, 36 years of age at death.Thirteen AMS 14C assays on bone cluster between 12,290 and 12,570 BP. Sixteen dates on wood specimens intimately associated with bone yield a range of dates from 11,980 to 12,940 BP.The remains exhibit multiple cut and wedge marks interpreted as cultural. Non-diagnostic cultural lithics were recovered. A disarticulated bone pile deposited in a low energy environment is consistent with human interaction.     

Mammoth tracks indicate a declining Late Pleistocene population in southwestern Alberta,Canada

Much debate has raged over the role that early humans played in this most recent large extinction. Fossil mammoth (Mammuthus primigenius) footprints were discovered at the St. Mary Reservoir in southwestern Canada (Wally's Beach DhPg-8). They are located in aeolian sediment dated at 11,300–11,000 years BP. By comparing the size distribution of these tracks with those of modern African elephants (Loxodonta africana), the age distribution of this mammoth population was determined. Containing far fewer juveniles than would be expected for an expanding or stable population, these tracks provide the first evidence that a living mammoth population, coexisting with human inhabitants, was in decline. Additionally, the same site provides corroborating evidence of humans hunting megafauna (horse and bovids). This suggests that humans, in addition to climate change, played a role in the end Pleistocene extinctions in North America.     

Pleistocene vertebrates of the Yukon Territory

Unglaciated parts of the Yukon constitute one of the most important areas in North America for yielding Pleistocene vertebrate fossils. Nearly 30 vertebrate faunal localities are reviewed spanning a period of about 1.6 Ma (million years ago) to the close of the Pleistocene some 10 000 BP (radiocarbon years before present, taken as 1950). The vertebrate fossils represent at least 8 species of fishes, 1 amphibian, 41 species of birds and 83 species of mammals. Dominant among the large mammals are: steppe bison (Bison priscus), horse (Equus sp.), woolly mammoth (Mammuthus primigenius), and caribou (Rangifer tarandus) – signature species of the Mammoth Steppe fauna (Fig. 1), which was widespread from the British Isles, through northern Europe, and Siberia to Alaska, Yukon and adjacent Northwest Territories. The Yukon faunas extend from Herschel Island in the north to Revenue Creek in the south and from the Alaskan border in the west to Ketza River in the east. The Yukon holds evidence of the earliest-known people in North America. Artifacts made from bison, mammoth and caribou bones from Bluefish Caves, Old Crow Basin and Dawson City areas show that people had a substantial knowledge of making and using bone tools at least by 25 000 BP, and possibly as early as 40 000 BP. A suggested chronological sequence of Yukon Pleistocene vertebrates (Table 1) facilitates comparison of selected faunas and indicates the known duration of various taxa.