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1.  An Abrupt Centennial-Scale Drought Event and Mid-Holocene Climate Change Patterns in Monsoon Marginal Zones of East Asia 
PLoS ONE  2014;9(3):e90241.
The mid-latitudes of East Asia are characterized by the interaction between the Asian summer monsoon and the westerly winds. Understanding long-term climate change in the marginal regions of the Asian monsoon is critical for understanding the millennial-scale interactions between the Asian monsoon and the westerly winds. Abrupt climate events are always associated with changes in large-scale circulation patterns; therefore, investigations into abrupt climate changes provide clues for responses of circulation patterns to extreme climate events. In this paper, we examined the time scale and mid-Holocene climatic background of an abrupt dry mid-Holocene event in the Shiyang River drainage basin in the northwest margin of the Asian monsoon. Mid-Holocene lacustrine records were collected from the middle reaches and the terminal lake of the basin. Using radiocarbon and OSL ages, a centennial-scale drought event, which is characterized by a sand layer in lacustrine sediments both from the middle and lower reaches of the basin, was absolutely dated between 8.0–7.0 cal kyr BP. Grain size data suggest an abrupt decline in lake level and a dry environment in the middle reaches of the basin during the dry interval. Previous studies have shown mid-Holocene drought events in other places of monsoon marginal zones; however, their chronologies are not strong enough to study the mechanism. According to the absolutely dated records, we proposed a new hypothesis that the mid-Holocene dry interval can be related to the weakening Asian summer monsoon and the relatively arid environment in arid Central Asia. Furthermore, abrupt dry climatic events are directly linked to the basin-wide effective moisture change in semi-arid and arid regions. Effective moisture is affected by basin-wide precipitation, evapotranspiration, lake surface evaporation and other geographical settings. As a result, the time scales of the dry interval could vary according to locations due to different geographical features.
PMCID: PMC3943949  PMID: 24599259
2.  High-resolution summer precipitation variations in the western Chinese Loess Plateau during the last glacial 
Scientific Reports  2013;3:2785.
We present a summer precipitation reconstruction for the last glacial (LG) on the western edge of the Chinese Loess Plateau (CLP) using a well-dated organic carbon isotopic dataset together with an independent modern process study results. Our results demonstrate that summer precipitation variations in the CLP during the LG were broadly correlated to the intensity of the Asian summer monsoon (ASM) as recorded by stalagmite oxygen isotopes from southern China. During the last deglaciation, the onset of the increase in temperatures at high latitudes in the Northern Hemisphere and decline in the intensity of the East Asia winter monsoon in mid latitudes was earlier than the increase in ASM intensity and our reconstructed summer precipitation in the western CLP. Quantitative reconstruction of a single paleoclimatic factor provides new insights and opportunities for further understanding of the paleoclimatic variations in monsoonal East Asia and their relation to the global climatic system.
PMCID: PMC3784955  PMID: 24071743
3.  Phylogeographic analysis reveals significant spatial genetic structure of Incarvillea sinensis as a product of mountain building 
BMC Plant Biology  2012;12:58.
Incarvillea sinensis is widely distributed from Southwest China to Northeast China and in the Russian Far East. The distribution of this species was thought to be influenced by the uplift of the Qinghai-Tibet Plateau and Quaternary glaciation. To reveal the imprints of geological events on the spatial genetic structure of Incarvillea sinensis, we examined two cpDNA segments ( trnH- psbA and trnS- trnfM) in 705 individuals from 47 localities.
A total of 16 haplotypes was identified, and significant genetic differentiation was revealed (GST =0.843, NST = 0.975, P < 0.05). The survey detected two highly divergent cpDNA lineages connected by a deep gap with allopatric distributions: the southern lineage with higher genetic diversity and differentiation in the eastern Qinghai-Tibet Plateau, and the northern lineage in the region outside the Qinghai-Tibet Plateau. The divergence between these two lineages was estimated at 4.4 MYA. A correlation between the genetic and the geographic distances indicates that genetic drift was more influential than gene flow in the northern clade with lower diversity and divergence. However, a scenario of regional equilibrium between gene flow and drift was shown for the southern clade. The feature of spatial distribution of the genetic diversity of the southern lineage possibly indicated that allopatric fragmentation was dominant in the collections from the eastern Qinghai-Tibet Plateau.
The results revealed that the uplift of the Qinghai-Tibet Plateau likely resulted in the significant divergence between the lineage in the eastern Qinghai-Tibet Plateau and the other one outside this area. The diverse niches in the eastern Qinghai-Tibet Plateau created a wide spectrum of habitats to accumulate and accommodate new mutations. The features of genetic diversity of populations outside the eastern Qinghai-Tibet Plateau seemed to reveal the imprints of extinction during the Glacial and the interglacial and postglacial recolonization. Our study is a typical case of the significance of the uplift of the Qinghai-Tibet Plateau and the Quaternary Glacial in spatial genetic structure of eastern Asian plants, and sheds new light on the evolution of biodiversity in the Qinghai-Tibet Plateau at the intraspecies level.
PMCID: PMC3447706  PMID: 22546007
Spatial genetic pattern; cpDNA variations; Phylogeography; Eastern Asian plant
4.  A less or more dusty future in the Northern Qinghai-Tibetan Plateau? 
Scientific Reports  2014;4:6672.
Dust plays an important role in climate changes as it can alter atmospheric circulation, and global biogeochemical and hydrologic cycling. Many studies have investigated the relationship between dust and temperature in an attempt to predict whether global warming in coming decades to centuries can result in a less or more dusty future. However, dust and temperature changes have rarely been simultaneously reconstructed in the same record. Here we present a 1600-yr-long quantitative record of temperature and dust activity inferred simultaneously from varved Kusai Lake sediments in the northern Qinghai–Tibetan Plateau, NW China. At decadal time scale, our temperature reconstructions are generally in agreement with tree-ring records from Karakorum of Pakistan, and temperature reconstructions of China and North Hemisphere based on compilations of proxy records. A less or more dusty future depends on temperature variations in the Northern Qinghai-Tibetan Plateau, i.e. weak and strong dust activities at centennial time scales are well correlated with low and high June–July–August temperature (average JJA temperature), respectively. This correlation means that stronger summer and winter monsoon should occur at the same times in the northern Qinghai–Tibetan Plateau.
PMCID: PMC4205841  PMID: 25335792
5.  Miocene shift of European atmospheric circulation from trade wind to westerlies 
Scientific Reports  2014;4:5660.
The modern European climatic regime is peculiar, due to its unitary winter but diverse summer climates and a pronounced Mediterranean climate in the south. However, little is known on its evolution in the deep time. Here we reconstruct the European summer climate conditions in the Tortonian (11.62–7.246 Ma) using plant fossil assemblages from 75 well-dated sites across Europe. Our results clearly show that the Tortonian Europe mainly had humid to subhumid summers and no arid climate has been conclusively detected, indicating that the summer-dry Mediterranean-type climate has not yet been established along most of the Mediterranean coast at least by the Tortonian. More importantly, the reconstructed distribution pattern of summer precipitation reveals that the Tortonian European must have largely been controlled by westerlies, resulting in higher precipitation in the west and the lower in the east. The Tortonian westerly wind field appears to differ principally from the trade wind pattern of the preceding Serravallian (13.82–11.62 Ma), recently deduced from herpetofaunal fossils. Such a shift in atmospheric circulation, if ever occurred, might result from the development of ice caps and glaciers in the polar region during the Late Miocene global cooling, the then reorganization of oceanic circulation, and/or the Himalayan-Tibetan uplift.
PMCID: PMC4092332  PMID: 25012454
6.  A Southern Ocean trigger for Northwest Pacific ventilation during the Holocene? 
Scientific Reports  2014;4:4046.
Holocene ocean circulation is poorly understood due to sparsity of dateable marine archives with submillennial-scale resolution. Here we present a record of mid-depth water radiocarbon contents in the Northwest (NW) Pacific Ocean over the last 12.000 years, which shows remarkable millennial-scale variations relative to changes in atmospheric radiocarbon inventory. Apparent decoupling of these variations from regional ventilation and mixing processes leads us to the suggestion that the mid-depth NW Pacific may have responded to changes in Southern Ocean overturning forced by latitudinal displacements of the southern westerly winds. By inference, a tendency of in-phase related North Atlantic and Southern Ocean overturning would argue against the development of a steady bipolar seesaw regime during the Holocene.
PMCID: PMC4027855  PMID: 24509792
7.  Fukushima Nuclear Accident Recorded in Tibetan Plateau Snow Pits 
PLoS ONE  2015;10(2):e0116580.
The β radioactivity of snow-pit samples collected in the spring of 2011 on four Tibetan Plateau glaciers demonstrate a remarkable peak in each snow pit profile, with peaks about ten to tens of times higher than background levels. The timing of these peaks suggests that the high radioactivity resulted from the Fukushima nuclear accident that occurred on March 11, 2011 in eastern Japan. Fallout monitoring studies demonstrate that this radioactive material was transported by the westerlies across the middle latitudes of the Northern Hemisphere. The depth of the peak β radioactivity in each snow pit compared with observational precipitation records, suggests that the radioactive fallout reached the Tibetan Plateau and was deposited on glacier surfaces in late March 2011, or approximately 20 days after the nuclear accident. The radioactive fallout existed in the atmosphere over the Tibetan Plateau for about one month.
PMCID: PMC4319914  PMID: 25658094
8.  Millennial Climatic Fluctuations Are Key to the Structure of Last Glacial Ecosystems 
PLoS ONE  2013;8(4):e61963.
Whereas fossil evidence indicates extensive treeless vegetation and diverse grazing megafauna in Europe and northern Asia during the last glacial, experiments combining vegetation models and climate models have to-date simulated widespread persistence of trees. Resolving this conflict is key to understanding both last glacial ecosystems and extinction of most of the mega-herbivores. Using a dynamic vegetation model (DVM) we explored the implications of the differing climatic conditions generated by a general circulation model (GCM) in “normal” and “hosing” experiments. Whilst the former approximate interstadial conditions, the latter, designed to mimic Heinrich Events, approximate stadial conditions. The “hosing” experiments gave simulated European vegetation much closer in composition to that inferred from fossil evidence than did the “normal” experiments. Given the short duration of interstadials, and the rate at which forest cover expanded during the late-glacial and early Holocene, our results demonstrate the importance of millennial variability in determining the character of last glacial ecosystems.
PMCID: PMC3628573  PMID: 23613985
9.  Phylogeography of Rhodiola kirilowii (Crassulaceae): A Story of Miocene Divergence and Quaternary Expansion 
PLoS ONE  2014;9(11):e112923.
The evolution and current distribution of the Sino-Tibetan flora have been greatly affected by historical geological events, such as the uplift of the Qinghai-Tibetan Plateau (QTP), and Quaternary climatic oscillations. Rhodiola kirilowii, a perennial herb with its distribution ranging from the southeastern QTP and the Hengduan Mountains (HM) to adjacent northern China and central Asia, provides an excellent model to examine and disentangle the effect of both geological orogeny and climatic oscillation on the evolutionary history of species with such distribution patterns. We here conducted a phylogeographic study using sequences of two chloroplast fragments (trnL-F and trnS-G) and internal transcribed spacers in 29 populations of R. kirilowii. A total of 25 plastid haplotypes and 12 ITS ribotypes were found. Molecular clock estimation revealed deep divergence between the central Asian populations and other populations from the HM and northern China; this split occurred ca. 2.84 million year ago. The majority of populations from the mountains of northern China were dominated by a single haplotype or ribotype, while populations of the HM harbored both high genetic diversity and high haplotype diversity. This distribution pattern indicates that HM was either a diversification center or a refugium for R. kirilowii during the Quaternary climatic oscillations. The present distribution of this species on mountains in northern China may have resulted from a rapid glacial population expansion from the HM. This expansion was confirmed by the mismatch distribution analysis and negative Tajima's D and Fu's FS values, and was dated to ca. 168 thousand years ago. High genetic diversity and population differentiation in both plastid and ITS sequences were revealed; these imply restricted gene flow between populations. A distinct isolation-by-distance pattern was suggested by the Mantel test. Our results show that in old lineages, populations may harbour divergent genetic forms that are sufficient to maintain or even increase overall genetic diversity despite fragmentation and low within-population variation.
PMCID: PMC4229298  PMID: 25389750
10.  Oscillators and relaxation phenomena in Pleistocene climate theory 
Ice sheets appeared in the northern hemisphere around 3 Ma (million years) ago and glacial–interglacial cycles have paced Earth's climate since then. Superimposed on these long glacial cycles comes an intricate pattern of millennial and sub-millennial variability, including Dansgaard–Oeschger and Heinrich events. There are numerous theories about these oscillations. Here, we review a number of them in order to draw a parallel between climatic concepts and dynamical system concepts, including, in particular, the relaxation oscillator, excitability, slow–fast dynamics and homoclinic orbits. Namely, almost all theories of ice ages reviewed here feature a phenomenon of synchronization between internal climate dynamics and astronomical forcing. However, these theories differ in their bifurcation structure and this has an effect on the way the ice age phenomenon could grow 3 Ma ago. All theories on rapid events reviewed here rely on the concept of a limit cycle excited by changes in the surface freshwater balance of the ocean. The article also reviews basic effects of stochastic fluctuations on these models, including the phenomenon of phase dispersion, shortening of the limit cycle and stochastic resonance. It concludes with a more personal statement about the potential for inference with simple stochastic dynamical systems in palaeoclimate science.
PMCID: PMC3261435  PMID: 22291227
palaeoclimates; dynamical systems; limit cycle; ice ages; Dansgaard–Oeschger events
11.  Ice-core records of atmospheric sulphur 
Sulphate and methanesulphonate (MSA), the two major sulphur species trapped in polar ice, have been extensivelyh studied in Antarctic and Greenland ice cores spanning the last centuries, as well as the entire last climatic cycle. Data from the cores are used to investigate the past contribution of volcanic and biogenic emissions to the natural sulphur budget in high latitude regions of both Hemispheres. Sulphate concentrations in polar ice very often increased during one or two years after large volcanic eruptions. Sulphate records show that fossil fuel combustion has enhanced sulphate concentrations in Greenland snow by a factor of 4 since the beginning of this century, and that no similar trend has occurred in Antarctica. At present, sulphate in Antarctic snow is mainly marine and biogenic in origin and the rate of dimethyl sulphide (DMS) emissions may have been enhanced during pst developments of El Niño Southern Oscillations (ENSO). Marine biota and non-eruptive volcanic emissions represent the two main contributors to the natural high northern latitude sulphur budget. Whele these two sources have contributed equally to the natural sulphur budget of Greenland ice over the last 9000 years BP, non-eruptive volcanic emissions largely dominated the budget at the beginning of the Holocene. A general negative correlation is observed between surcace air temperatures of the Northern Hemisphere and Greenland snow MSA concentrations over the last two centuries. Positive sea-ice anomalies also seem to strengthen DMS emissions. A steady decrease of MSA is observed in Greenland snow layers deposited since 1945, which may either be related to decreasing DMS emissions from marine biota at high northern latitudes or a changing yield of MSA from DMS oxidation driven by modification of the oxidative capacity of the atmosphere in these regions. Slightly reduced MSA concentrations are obvserved in Greenland glacial ice with respect to interglacial levels. In contrast, sulphate and calcium levels are strongly enhanced during the ice age compared to the present day. These long-term variations in Greenland cores are opposite in sign to those revealed by Antarctic ice cores. Such a difference suggests that climate changes led to a quite different sulphur cycle response in the two Hemispheres.
PMCID: PMC1691926
Ice Cores Greenland Antarctica Methane Sulphonate Ice Age El Niño Southern Oscillations (Enso)
12.  Rapid thinning of the late Pleistocene Patagonian Ice Sheet followed migration of the Southern Westerlies 
Scientific Reports  2013;3:2118.
Here we present the first reconstruction of vertical ice-sheet profile changes from any of the Southern Hemisphere's mid-latitude Pleistocene ice sheets. We use cosmogenic radio-nuclide (CRN) exposure analysis to record the decay of the former Patagonian Ice Sheet (PIS) from the Last Glacial Maximum (LGM) and into the late glacial. Our samples, from mountains along an east-west transect to the east of the present North Patagonian Icefield (NPI), serve as ‘dipsticks' that allow us to reconstruct past changes in ice-sheet thickness, and demonstrates that the former PIS remained extensive and close to its LGM extent in this region until ~19.0 ka. After this time rapid ice-sheet thinning, initiated at ~18.1 ka, saw ice at or near its present dimension by 15.5 ka. We argue this rapid thinning was triggered by a combination of the rapid southward migration of the precipitation bearing Southern Hemisphere (SH) westerlies and regional warming.
PMCID: PMC3698495  PMID: 23817136
13.  Vegetation Controls on Weathering Intensity during the Last Deglacial Transition in Southeast Africa 
PLoS ONE  2014;9(11):e112855.
Tropical climate is rapidly changing, but the effects of these changes on the geosphere are unknown, despite a likelihood of climatically-induced changes on weathering and erosion. The lack of long, continuous paleo-records prevents an examination of terrestrial responses to climate change with sufficient detail to answer questions about how systems behaved in the past and may alter in the future. We use high-resolution records of pollen, clay mineralogy, and particle size from a drill core from Lake Malawi, southeast Africa, to examine atmosphere-biosphere-geosphere interactions during the last deglaciation (∼18–9 ka), a period of dramatic temperature and hydrologic changes. The results demonstrate that climatic controls on Lake Malawi vegetation are critically important to weathering processes and erosion patterns during the deglaciation. At 18 ka, afromontane forests dominated but were progressively replaced by tropical seasonal forest, as summer rainfall increased. Despite indication of decreased rainfall, drought-intolerant forest persisted through the Younger Dryas (YD) resulting from a shorter dry season. Following the YD, an intensified summer monsoon and increased rainfall seasonality were coeval with forest decline and expansion of drought-tolerant miombo woodland. Clay minerals closely track the vegetation record, with high ratios of kaolinite to smectite (K/S) indicating heavy leaching when forest predominates, despite variable rainfall. In the early Holocene, when rainfall and temperature increased (effective moisture remained low), open woodlands expansion resulted in decreased K/S, suggesting a reduction in chemical weathering intensity. Terrigenous sediment mass accumulation rates also increased, suggesting critical linkages among open vegetation and erosion during intervals of enhanced summer rainfall. This study shows a strong, direct influence of vegetation composition on weathering intensity in the tropics. As climate change will likely impact this interplay between the biosphere and geosphere, tropical landscape change could lead to deleterious effects on soil and water quality in regions with little infrastructure for mitigation.
PMCID: PMC4236122  PMID: 25406090
14.  Lake Sediment Records on Climate Change and Human Activities in the Xingyun Lake Catchment, SW China 
PLoS ONE  2014;9(7):e102167.
Sediments from Xinyun Lake in central Yunnan, southwest China, provide a record of environmental history since the Holocene. With the application of multi-proxy indicators (total organic carbon (TOC), total nitrogen (TN), δ13C and δ15N isotopes, C/N ratio, grain size, magnetic susceptibility (MS) and CaCO3 content), as well as accelerator mass spectrometry (AMS) 14C datings, four major climatic stages during the Holocene have been identified in Xingyun′s catchment. A marked increase in lacustrine palaeoproductivity occurred from 11.06 to 9.98 cal. ka BP, which likely resulted from an enhanced Asian southwest monsoon and warm-humid climate. Between 9.98 and 5.93 cal. ka BP, a gradually increased lake level might have reached the optimum water depth, causing a marked decline in coverage by aquatic plants and lake productivity of the lake. This was caused by strong Asian southwest monsoon, and coincided with the global Holocene Optimum. During the period of 5.60–1.35 cal. ka BP, it resulted in a warm and dry climate at this stage, which is comparable to the aridification of India during the mid- and late Holocene. The intensifying human activity and land-use in the lake catchment since the early Tang Dynasty (∼1.35 cal. ka BP) were associated with the ancient Dian culture within Xingyun’s catchment. The extensive deforestation and development of agriculture in the lake catchment caused heavy soil loss. Our study clearly shows that long-term human activities and land-use change have strongly impacted the evolution of the lake environment and therefore modulated the sediment records of the regional climate in central Yunnan for more than one thousand years.
PMCID: PMC4102491  PMID: 25033404
15.  Holocene variations in peatland methane cycling associated with the Asian summer monsoon system 
Nature Communications  2014;5:4631.
Atmospheric methane concentrations decreased during the early to middle Holocene; however, the governing mechanisms remain controversial. Although it has been suggested that the mid-Holocene minimum methane emissions are associated with hydrological change, direct evidence is lacking. Here we report a new independent approach, linking hydrological change in peat sediments from the Tibetan Plateau to changes in archaeal diether concentrations and diploptene δ13C values as tracers for methanogenesis and methanotrophy, respectively. A minimum in inferred methanogenesis occurred during the mid-Holocene, which, locally, corresponds with the driest conditions of the Holocene, reflecting a minimum in Asian monsoon precipitation. The close coupling between precipitation and methanogenesis is validated by climate simulations, which also suggest a regionally widespread impact. Importantly, the minimum in methanogenesis is associated with a maximum in methanotrophy. Therefore, methane emissions in the Tibetan Plateau region were apparently lower during the mid-Holocene and partially controlled by interactions of large-scale atmospheric circulation.
Although it has been widely suggested that the mid-Holocene minimum methane emissions are associated with hydrological change, direct evidence is missing. Here, the authors present evidence from the Tibetan Plateau using tracers of methanogenesis and methanotrophy, in combination with climate simulations.
PMCID: PMC4143914  PMID: 25135106
16.  The ocean's role in polar climate change: asymmetric Arctic and Antarctic responses to greenhouse gas and ozone forcing 
In recent decades, the Arctic has been warming and sea ice disappearing. By contrast, the Southern Ocean around Antarctica has been (mainly) cooling and sea-ice extent growing. We argue here that interhemispheric asymmetries in the mean ocean circulation, with sinking in the northern North Atlantic and upwelling around Antarctica, strongly influence the sea-surface temperature (SST) response to anthropogenic greenhouse gas (GHG) forcing, accelerating warming in the Arctic while delaying it in the Antarctic. Furthermore, while the amplitude of GHG forcing has been similar at the poles, significant ozone depletion only occurs over Antarctica. We suggest that the initial response of SST around Antarctica to ozone depletion is one of cooling and only later adds to the GHG-induced warming trend as upwelling of sub-surface warm water associated with stronger surface westerlies impacts surface properties. We organize our discussion around ‘climate response functions’ (CRFs), i.e. the response of the climate to ‘step’ changes in anthropogenic forcing in which GHG and/or ozone-hole forcing is abruptly turned on and the transient response of the climate revealed and studied. Convolutions of known or postulated GHG and ozone-hole forcing functions with their respective CRFs then yield the transient forced SST response (implied by linear response theory), providing a context for discussion of the differing warming/cooling trends in the Arctic and Antarctic. We speculate that the period through which we are now passing may be one in which the delayed warming of SST associated with GHG forcing around Antarctica is largely cancelled by the cooling effects associated with the ozone hole. By mid-century, however, ozone-hole effects may instead be adding to GHG warming around Antarctica but with diminished amplitude as the ozone hole heals. The Arctic, meanwhile, responding to GHG forcing but in a manner amplified by ocean heat transport, may continue to warm at an accelerating rate.
PMCID: PMC4032509  PMID: 24891392
Southern Ocean; ozone hole; greenhouse gas forcing; climate model
17.  Genomic Analysis of Natural Selection and Phenotypic Variation in High-Altitude Mongolians 
PLoS Genetics  2013;9(7):e1003634.
Deedu (DU) Mongolians, who migrated from the Mongolian steppes to the Qinghai-Tibetan Plateau approximately 500 years ago, are challenged by environmental conditions similar to native Tibetan highlanders. Identification of adaptive genetic factors in this population could provide insight into coordinated physiological responses to this environment. Here we examine genomic and phenotypic variation in this unique population and present the first complete analysis of a Mongolian whole-genome sequence. High-density SNP array data demonstrate that DU Mongolians share genetic ancestry with other Mongolian as well as Tibetan populations, specifically in genomic regions related with adaptation to high altitude. Several selection candidate genes identified in DU Mongolians are shared with other Asian groups (e.g., EDAR), neighboring Tibetan populations (including high-altitude candidates EPAS1, PKLR, and CYP2E1), as well as genes previously hypothesized to be associated with metabolic adaptation (e.g., PPARG). Hemoglobin concentration, a trait associated with high-altitude adaptation in Tibetans, is at an intermediate level in DU Mongolians compared to Tibetans and Han Chinese at comparable altitude. Whole-genome sequence from a DU Mongolian (Tianjiao1) shows that about 2% of the genomic variants, including more than 300 protein-coding changes, are specific to this individual. Our analyses of DU Mongolians and the first Mongolian genome provide valuable insight into genetic adaptation to extreme environments.
Author Summary
Throughout history, Mongolians have survived the harsh conditions of northern latitudes, including seasonal cold, drought, and a restricted diet. Approximately 500 years ago, nomadic Deedu (DU; “at high altitude”) Mongolians migrated from the Mongolian steppes to the northeastern highlands of the Qinghai-Tibetan Plateau. Using high-density SNP data, we demonstrate that present-day DU Mongolians share genetic ancestry with other Mongolians and with Tibetans. High-altitude selection candidate genes previously identified in the latter population (EPAS1, PKLR, CYP2E1), and PPARG, a gene long hypothesized to play a role in metabolic adaptation, are among the strongest adaptive signals in DU Mongolians. Furthermore, we show that hemoglobin concentration, associated with high-altitude adaptation in Tibetans, is intermediate in DU Mongolians compared to Tibetans and Han Chinese at comparable altitudes. Whole-genome sequence from a DU Mongolian shows that approximately 300 protein-coding changes are specific to this individual. Our analyses provide new perspectives on genetic variation and adaptation to extreme environments.
PMCID: PMC3715426  PMID: 23874230
18.  Good Days, Bad Days: Wind as a Driver of Foraging Success in a Flightless Seabird, the Southern Rockhopper Penguin 
PLoS ONE  2013;8(11):e79487.
Due to their restricted foraging range, flightless seabirds are ideal models to study the short-term variability in foraging success in response to environmentally driven food availability. Wind can be a driver of upwelling and food abundance in marine ecosystems such as the Southern Ocean, where wind regime changes due to global warming may have important ecological consequences. Southern rockhopper penguins (Eudyptes chrysocome) have undergone a dramatic population decline in the past decades, potentially due to changing environmental conditions. We used a weighbridge system to record daily foraging mass gain (the difference in mean mass of adults leaving the colony in the morning and returning to the colony in the evening) of adult penguins during the chick rearing in two breeding seasons. We related the day-to-day variability in foraging mass gain to ocean wind conditions (wind direction and wind speed) and tested for a relationship between wind speed and sea surface temperature anomaly (SSTA). Foraging mass gain was highly variable among days, but did not differ between breeding seasons, chick rearing stages (guard and crèche) and sexes. It was strongly correlated between males and females, indicating synchronous changes among days. There was a significant interaction of wind direction and wind speed on daily foraging mass gain. Foraging mass gain was highest under moderate to strong winds from westerly directions and under weak winds from easterly directions, while decreasing under stronger easterly winds and storm conditions. Ocean wind speed showed a negative correlation with daily SSTA, suggesting that winds particularly from westerly directions might enhance upwelling and consequently the prey availability in the penguins' foraging areas. Our data emphasize the importance of small-scale, wind-induced patterns in prey availability on foraging success, a widely neglected aspect in seabird foraging studies, which might become more important with increasing changes in climatic variability.
PMCID: PMC3827366  PMID: 24236139
19.  Periodic variability in cetacean strandings: links to large-scale climate events 
Biology Letters  2005;1(2):147-150.
Cetacean strandings elicit much community and scientific interest, but few quantitative analyses have successfully identified environmental correlates to these phenomena. Data spanning 1920–2002, involving a total of 639 stranding events and 39 taxa groups from southeast Australia, were found to demonstrate a clear 11–13- year periodicity in the number of events through time. These data positively correlated with the regional persistence of both zonal (westerly) and meridional (southerly) winds, reflecting general long-term and large-scale shifts in sea-level pressure gradients. Periods of persistent zonal and meridional winds result in colder and presumably nutrient-rich waters being driven closer to southern Australia, resulting in increased biological activity in the water column during the spring months. These observations suggest that large-scale climatic events provide a powerful distal influence on the propensity for whales to strand in this region. These patterns provide a powerful quantitative framework for testing hypotheses regarding environmental links to strandings and provide managers with a potential predictive tool to prepare for years of peak stranding activity.
PMCID: PMC1626231  PMID: 17148151
cetacean strandings; southeast Australia; climate; meridional winds; zonal winds; sea-surface temperature
20.  Glacial vicariance in Eurasia: mitochondrial DNA evidence from Scots pine for a complex heritage involving genetically distinct refugia at mid-northern latitudes and in Asia Minor 
At the last glacial maximum, Fennoscandia was covered by an ice sheet while the tundra occupied most of the rest of northern Eurasia. More or less disjunct refugial populations of plants were dispersed in southern Europe, often trapped between mountain ranges and seas. Genetic and paleobotanical evidences indicate that these populations have contributed much to Holocene recolonization of more northern latitudes. Less supportive evidence has been found for the existence of glacial populations located closer to the ice margin. Scots pine (Pinus sylvestris L.) is a nordic conifer with a wide natural range covering much of Eurasia. Fractures in its extant genetic structure might be indicative of glacial vicariance and how different refugia contributed to the current distribution at the continental level. The population structure of Scots pine was investigated on much of its Eurasian natural range using maternally inherited mitochondrial DNA polymorphisms.
A novel polymorphic region of the Scots pine mitochondrial genome has been identified, the intron 1 of nad7, with three variants caused by insertions-deletions. From 986 trees distributed among 54 populations, four distinct multi-locus mitochondrial haplotypes (mitotypes) were detected based on the three nad7 intron 1 haplotypes and two previously reported size variants for nad1 intron B/C. Population differentiation was high (GST = 0.657) and the distribution of the mitotypes was geographically highly structured, suggesting at least four genetically distinct ancestral lineages. A cosmopolitan lineage was widely distributed in much of Europe throughout eastern Asia. A previously reported lineage limited to the Iberian Peninsula was confirmed. A new geographically restricted lineage was found confined to Asia Minor. A new lineage was restricted to more northern latitudes in northeastern Europe and the Baltic region.
The contribution of the various ancestral lineages to the current distribution of Scots pine was asymmetric and extant endemism reflected the presence of large geographic barriers to migration. The results suggest a complex biogeographical history with glacial refugia shared with temperate plant species in southern European Peninsulas and Asia Minor, and a genetically distinct glacial population located more North. These results confirm recent observations for cold tolerant species about the possible existence of refugial populations at mid-northern latitudes contributing significantly to the recolonization of northern Europe. Thus, Eurasian populations of nordic plant species might not be as genetically homogenous as assumed by simply considering them as offsets of glacial populations located in southern peninsulas. As such, they might have evolved distinctive genetic adaptations during glacial vicariance, worth evaluating and considering for conservation.
PMCID: PMC2222255  PMID: 18034901
21.  The rise of the Himalaya enforced the diversification of SE Asian ferns by altering the monsoon regimes 
BMC Plant Biology  2012;12:210.
The rise of high mountain chains is widely seen as one of the factors driving rapid diversification of land plants and the formation of biodiversity hotspots. Supporting evidence was reported for the impact of the rapid rise of the Andean mountains but this hypothesis has so far been less explored for the impact of the “roof of the world”. The formation of the Himalaya, and especially the rise of the Qinghai–Tibetan Plateau in the recent 20 million years, altered the monsoon regimes that dominate the current climates of South East Asia. Here, we infer the hypothesis that the rise of Himalaya had a strong impact on the plant diversity in the biodiversity hotspot of the Southwest Chinese Mountains.
Our analyses of the diversification pattern of the derived fern genus Lepisorus recovered evidence for changes in plant diversity that correlated with the strengthening of South East Asian monsoon. Southwest China or Southwest China and Japan was recovered as the putative area of origin of Lepisorus and enhancing monsoon regime were found to shape the early diversification of the genus as well as subsequent radiations during the late Miocene and Pliocene.
We report new evidence for a coincidence of plant diversification and changes of the climate caused by the uplift of the Himalaya. These results are discussed in the context of the impact of incomplete taxon sampling, uncertainty of divergence time estimates, and limitations of current methods used to assess diversification rates.
PMCID: PMC3508991  PMID: 23140168
Diversification pattern; East Asian monsoon; Himalaya; LASER; Lepisorus
22.  Recent Changes in Glacial Area and Volume on Tuanjiefeng Peak Region of Qilian Mountains, China 
PLoS ONE  2013;8(8):e70574.
Glaciers' runoff in the Qilian Mountains serves as a critical water resource in the northern sections of the Gansu province, the northeastern sections of the Qinghai province, and the northeastern fringe of the Tibetan Plateau. Changes in the glacial area and volume around the highest peak of the Qilian Mountains, i.e., Tuanjiefeng Peak, were estimated using multi-temporal remote-sensing images and digital elevation models, and all possible sources of uncertainty were considered in detail. The total glacier area decreased by 16.1±6.34 km2 (9.9±3.9%) during 1966 to 2010. The average annual glacier shrinkage was −0.15% a−1 from 1966 to 1995, −0.61% a−1 from 1995 to 2000, −0.20% a−1 from 2000 to 2006, and −0.45% a−1 from 2006 to 2010. A comparison of glacier surface elevations using digital elevation models derived from topographic maps in 1966 and from the Shuttle Radar Topography Mission in 1999 suggests that 65% of the grid cells has decreased, thereby indicating that the glacier thickness has declined. The average change in glacier thickness was −7.3±1.5 m (−0.21±0.04 m·a−1) from 1966 to 1999. Glaciers with northeastern aspects thinned by 8.3±1.4 m from 1966 to 1999, i.e., almost twice as much as those with southwestern aspects (4.3±1.3 m). The ice volume decreased by 11.72±2.38×108 m3 from 1966 to 1999, which was about 17.4% more than the value calculated from the statistical relationship between glacier area and volume. The relationship between glacier area change and elevation zone indicates that glacier change is not only dominated by climate change but also affected by glacier dynamics, which are related to local topography. The varied response of a single glacier to climate change indicates that the glacier area change scheme used in some models must be improved.
PMCID: PMC3754983  PMID: 24015174
23.  Identification of Photosynthetic Plankton Communities Using Sedimentary Ancient DNA and Their Response to late-Holocene Climate Change on the Tibetan Plateau 
Scientific Reports  2014;4:6648.
Sediments from Tibetan lakes in NW China are potentially sensitive recorders of climate change and its impact on ecosystem function. However, the important plankton members in many Tibetan Lakes do not make and leave microscopically diagnostic features in the sedimentary record. Here we established a taxon-specific molecular approach to specifically identify and quantify sedimentary ancient DNA (sedaDNA) of non-fossilized planktonic organisms preserved in a 5-m sediment core from Kusai Lake spanning the last 3100 years. The reliability of the approach was validated with multiple independent genetic markers. Parallel analyses of the geochemistry of the core and paleo-climate proxies revealed that Monsoon strength-driven changes in nutrient availability, temperature, and salinity as well as orbitally-driven changes in light intensity were all responsible for the observed temporal changes in the abundance of two dominant phytoplankton groups in the lake, Synechococcus (cyanobacteria) and Isochrysis (haptophyte algae). Collectively our data show that global and regional climatic events exhibited a strong influence on the paleoecology of phototrophic plankton in Kusai Lake.
PMCID: PMC4200408  PMID: 25323386
24.  Thermal Controls on the Asian Summer Monsoon 
Scientific Reports  2012;2:404.
The Asian summer monsoon affects more than sixty percent of the world's population; understanding its controlling factors is becoming increasingly important due to the expanding human influence on the environment and climate and the need to adapt to global climate change. Various mechanisms have been suggested; however, an overarching paradigm delineating the dominant factors for its generation and strength remains debated. Here we use observation data and numerical experiments to demonstrates that the Asian summer monsoon systems are controlled mainly by thermal forcing whereas large-scale orographically mechanical forcing is not essential: the South Asian monsoon south of 20°N by land–sea thermal contrast, its northern part by the thermal forcing of the Iranian Plateau, and the East Asian monsoon and the eastern part of the South Asian monsoon by the thermal forcing of the Tibetan Plateau.
PMCID: PMC3349950  PMID: 22582141
25.  Exploring Holocene Changes in Palynological Richness in Northern Europe – Did Postglacial Immigration Matter? 
PLoS ONE  2012;7(12):e51624.
In mid to high latitudes glacial and interglacial cycles have repeatedly changed the area available for plant growth. The speed at which plants are able to colonize areas at the onset of an interglacial is hypothesized to limit their distribution ranges even today (migrational lag). If the spread of plants would have been generally slow then plant diversity in previously glaciated areas would be expected to increase over time. We explore this hypothesis using results from six palynological investigations from two previously glaciated regions: central Sweden and north-eastern Germany. Rarefaction, slope of rank order abundance, and taxa accumulation plots were used to evaluate richness and evenness in pollen data in an attempt to separate richness from evenness. These analyses show little change in palynological richness for the northern sites throughout the Holocene. In contrast, the southern sites show an increase in richness and evenness during the early Holocene; this may be explained by the different initial conditions at the onset of the Holocene. A strong rise in palynological richness around 6000 and 1000 years ago at the southern sites can be attributed to the regional initiation of agriculture and major opening of the forest, respectively. For the northern sites there is no evidence for increased taxonomic diversity through time that could be due to delayed immigration of species.
PMCID: PMC3519870  PMID: 23240048

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