末次盛冰期以来云南异龙湖的环境演变——来自地球化学沉积记录的证据

通过对云南异龙湖4.56 m沉积岩芯进行X射线荧光光谱分析(XRF),结合13个AMS14C测年结果,获得了高分辨率元素地球化学序列。异龙湖岩芯中元素的因子分析显示,F1因子揭示了沉积地球化学元素的来源,其正载荷代表了流域外源物质输入加强,而负载荷则指示了湖泊自生碳酸盐沉淀的增加,F2因子指示了湖泊氧化还原状态,F3因子可能与人类活动有关。在此基础上结合磁化率、矿物组成及有机质含量等指标,对异龙湖沉积环境演化过程进行探讨。结果表明,在末次盛冰期期间(26000—17000 cal. yr. BP),区域气候特征是气温低、降水少;在17000—14500 cal. yr. BP期间,异龙湖经历了显著的干旱气候,可能与亚洲夏季风突然减弱有关; 14500—9000 cal. yr. BP,异龙湖经历了显著的高湖面阶段,与滇池高水位时期基本一致,表明流域降水丰沛,外源物质输入增加; 9000—6000 cal. yr. BP期间,区域温度增加导致流域干旱化加剧,此阶段与西南地区全新世大暖期(全新世适宜期)相对应; 6000 cal. yr. BP以后外援物质输入锐减可能指示了区域降水的降低。异龙湖地球化学沉积记录揭示的环境演变过程与区域气候环境变化具有一致性,也具有明显的特殊性。     

末次盛冰期环境恶化对中国北方旧石器文化的影响

末次冰期间冰段时,中国北方地区水热条件普遍较好,旧石器文化也比较发达。然而,进入末次盛冰期后,随着气温的普遍降低和降水量的明显减少,草原和沙漠带急速向南扩张,动物群也大举南迁。对于这种严酷的生存环境,中国北方的旧石器文化是如何响应的呢?本文以宁夏彭阳PY01-02,PY03和PY05等地点的地层剖面和石制品的研究为基础,首先提出彭阳地点群地层沉积相由河湖相沉积迅速转变为黄土堆积说明由末次冰期间冰段到末次盛冰期自然环境发生了剧烈变化。其次,通过对上述两种不同沉积相中出土石制品的大小、重量、原料以及技术上的比较说明随着环境的变化石制品本身也发生了相应的变化。下层河湖相堆积中出土的石制品粗大笨重,主要以各种石英岩为原料,与同期泾水上游旧石器文化的特征相近,应是当地文化特征的一种具体体现。上层马兰黄土中的石制品的普遍较小,主要以石英和燧石为原料,其中用燧石制成的细石器具有宁夏灵武水洞沟下文化层同类制品的特征,而用石英制成的小石器则同海原一带的史前文化有着较为密切的联系,据此作者推测这可能与史前人类在环境日益恶劣的情况下向南迁移有关。山西南部地区的下川和柿子滩遗址的地层和石制品证据亦表明,末次盛冰期前后该地区旧石器文化的明显差异可能也与末次盛冰期环境恶化时人类的迁徙有关。最后,中国北方地区考古遗址的空间分布变化为上述史前人类南迁的推测提供了有力证据。因此,作者认为末次冰期的环境恶化对中国北方旧石器文化的影响具有普遍性。     

A Matrilineal Genetic Legacy from the Last Glacial Maximum Confers Susceptibility to Schizophrenia in Han Chinese

Mitochondrial dysfunction has been widely reported in schizophrenia patients. To dissect the matrilineal structure of Han Chinese with or without schizophrenia and to decipher the maternal influence and evolutionary history of schizophrenia, a total of 1212 schizophrenia patients and 1005 matched healthy controls, all of Han Chinese origin, were recruited in Hunan Province, China. We classified haplogroup for each individual based on mitochondrial DNA （mtDNA） sequence variations and compared the haplogroup distribution pattern between cases and controls. Haplogroup B5a presented a higher frequency in cases than in controls （P = 0.02, OR = 1.67, 95% CI = [1.09, 2.56]）, and this result could be confirmed by permutation analysis. Age estimation of haplogroup B5a in cases revealed a much younger age than that of controls, which was coincident with the Northern Hemisphere deglaciation at the end of the Last Glacial Maximum. Analysis of complete mtDNA in five patients belonging to haplogroup B5a showed that this background effect might be caused by haplogroup- defining variants m.8584G〉A and m.10398A〉G. Our results showed that matrilineal risk factor for schizophrenia had an ancient origin and might acquire a predisposing effect on schizophrenia due to the environment change and/or orchestration with other nuclear genetic factors appeared recently in human evolutionary history.     

A Short Guide to the Climatic Variables of the Last Glacial Maximum for Biogeographers

Ecological niche models are widely used for mapping the distribution of species during the last glacial maximum (LGM). Although the selection of the variables and General Circulation Models (GCMs) used for constructing those maps determine the model predictions, we still lack a discussion about which variables and which GCM should be included in the analysis and why. Here, we analyzed the climatic predictions for the LGM of 9 different GCMs in order to help biogeographers to select their GCMs and climatic layers for mapping the species ranges in the LGM. We 1) map the discrepancies between the climatic predictions of the nine GCMs available for the LGM, 2) analyze the similarities and differences between the GCMs and group them to help researchers choose the appropriate GCMs for calibrating and projecting their ecological niche models (ENM) during the LGM, and 3) quantify the agreement of the predictions for each bioclimatic variable to help researchers avoid the environmental variables with a poor consensus between models. Our results indicate that, in absolute values, GCMs have a strong disagreement in their temperature predictions for temperate areas, while the uncertainties for the precipitation variables are in the tropics. In spite of the discrepancies between model predictions, temperature variables (BIO1-BIO11) are highly correlated between models. Precipitation variables (BIO12- BIO19) show no correlation between models, and specifically, BIO14 (precipitation of the driest month) and BIO15 (Precipitation Seasonality (Coefficient of Variation)) show the highest level of discrepancy between GCMs. Following our results, we strongly recommend the use of different GCMs for constructing or projecting ENMs, particularly when predicting the distribution of species that inhabit the tropics and the temperate areas of the Northern and Southern Hemispheres, because climatic predictions for those areas vary greatly among GCMs. We also recommend the exclusion of BIO14 and BIO15 from ENMs because those variables show a high level of discrepancy between GCMs. Thus, by excluding them, we decrease the level of uncertainty of our predictions. All the climatic layers produced for this paper are freely available in http://ecoclimate.org/.     

Expansion dating: calibrating molecular clocks in marine species from expansions onto the Sunda Shelf Following the Last Glacial Maximum

The rate of change in DNA is an important parameter for understanding molecular evolution and hence for inferences drawn from studies of phylogeography and phylogenetics. Most rate calibrations for mitochondrial coding regions in marine species have been made from divergence dating for fossils and vicariant events older than 1-2 My and are typically 0.5-2% per lineage per million years. Recently, calibrations made with ancient DNA (aDNA) from younger dates have yielded faster rates, suggesting that estimates of the molecular rate of change depend on the time of calibration, decaying from the instantaneous mutation rate to the phylogenetic substitution rate. aDNA methods for recent calibrations are not available for most marine taxa so instead we use radiometric dates for sea-level rise onto the Sunda Shelf following the Last Glacial Maximum (starting ～18,000 years ago), which led to massive population expansions for marine species. Instead of divergence dating, we use a two-epoch coalescent model of logistic population growth preceded by a constant population size to infer a time in mutational units for the beginning of these expansion events. This model compares favorably to simpler coalescent models of constant population size, and exponential or logistic growth, and is far more precise than estimates from the mismatch distribution. Mean rates estimated with this method for mitochondrial coding genes in three invertebrate species are elevated in comparison to older calibration points (2.3-6.6% per lineage per million years), lending additional support to the hypothesis of calibration time dependency for molecular rates.     

The recolonization of Europe by brown bears Ursus arctos Linnaeus, 1758 after the Last Glacial Maximum

1. At the end of the Last Glacial Maximum brown bears Ursus arctos recolonized the glacial landscape of Central and Northern Europe faster than all other carnivorous mammal species of the Holocene fauna. Ursus arctos was recorded in Northern Europe from the beginning of the Late-Glacial. The recolonization of northern Central Europe may have taken place directly after the maximum glaciation. The distribution of the brown bear was restricted to glacial refugia only during the Last Glacial Maximum, for probably no more than 10 000 years. 2. Genetic analyses have suggested three glacial refugia for the brown bear: the Iberian Peninsula, the Italian Peninsula and the Balkans. Subfossil records of Ursus arctos from north-western Moldova as well as reconstructed environmental conditions during the Last Glacial Maximum in this area suggest to us a fourth glacial refuge for the brown bear. Because of its connection to the Carpathians, we designate this as the ‘Carpathian refuge’. 3. Due to the low genetic distance between brown bears of northern Norway, Finland, Estonia, north-eastern Russia and the northern Carpathians (the so-called eastern lineage), the Carpathians were considered the geographical origin of the recolonization of these regions. During the recolonization of northern Europe the brown bear probably reached these areas rapidly from the putative Carpathian refuge.     

Genetic and palaeo-climatic evidence for widespread persistence of the coastal tree species Eucalyptus gomphocephala (Myrtaceae) during the Last Glacial Maximum

Background and Aims

Few phylogeographic studies have been undertaken of species confined to narrow, linear coastal systems where past sea level and geomorphological changes may have had a profound effect on species population sizes and distributions. In this study, a phylogeographic analysis was conducted of Eucalyptus gomphocephala (tuart), a tree species restricted to a 400 × 10 km band of coastal sand-plain in south west Australia. Here, there is little known about the response of coastal vegetation to glacial/interglacial climate change, and a test was made as to whether this species was likely to have persisted widely through the Last Glacial Maximum (LGM), or conforms to a post-LGM dispersal model of recovery from few refugia.

Methods

The genetic structure over the entire range of tuart was assessed using seven nuclear (21 populations; n = 595) and four chloroplast (24 populations; n = 238) microsatellite markers designed for eucalypt species. Correlative palaeodistribution modelling was also conducted based on five climatic variables, within two LGM models.

Key Results

The chloroplast markers generated six haplotypes, which were strongly geographically structured (G_ST = 0·86 and R_ST = 0·75). Nuclear microsatellite diversity was high (overall mean H_E 0·75) and uniformly distributed (F_ST = 0·05), with a strong pattern of isolation by distance (r² = 0·362, P = 0·001). Distribution models of E. gomphocephala during the LGM showed a wide distribution that extended at least 30 km westward from the current distribution to the palaeo-coastline.

Conclusions

The chloroplast and nuclear data suggest wide persistence of E. gomphocephala during the LGM. Palaeodistribution modelling supports the conclusions drawn from genetic data and indicates a widespread westward shift of E. gomphocephala onto the exposed continental shelf during the LGM. This study highlights the importance of the inclusion of complementary, non-genetic data (information on geomorphology and palaeoclimate) to interpret phylogeographic patterns.     

50,000 years of ice and seals: Impacts of the Last Glacial Maximum on Antarctic fur seals

Ice is one of the most important drivers of population dynamics in polar organisms, influencing the locations, sizes, and connectivity of populations. Antarctic fur seals, Arctocephalus gazella, are particularly interesting in this regard, as they are concomitantly reliant on both ice‐associated prey and ice‐free coastal breeding areas. We reconstructed the history of this species through the Last Glacial Maximum (LGM) using genomic sequence data from seals across their range. Population size trends and divergence events were investigated using continuous‐time size estimation analysis and divergence time estimation models. The combined results indicated that a panmictic population present prior to the LGM split into two small refugial populations during peak ice extent. Following ice decline, the western refugial population founded colonies at the South Shetlands, South Georgia, and Bouvetøya, while the eastern refugial population founded the colony on Iles Kerguelen. Postglacial population divergence times closely match geological estimates of when these coastal breeding areas became ice free. Given the predictions regarding continued future warming in polar oceans, these responses of Antarctic fur seals to past climate variation suggest it may be worthwhile giving conservation consideration to potential future breeding locations, such as areas further south along the Antarctic Peninsula, in addition to present colony areas.     

Satsurblia: New Insights of Human Response and Survival across the Last Glacial Maximum in the Southern Caucasus

The region of western Georgia (Imereti) has been a major geographic corridor for human migrations during the Middle and Upper Palaeolithic (MP/UP). Knowledge of the MP and UP in this region, however, stems mostly from a small number of recent excavations at the sites of Ortvale Klde, Dzudzuana, Bondi, and Kotias Klde. These provide an absolute chronology for the Late MP and MP–UP transition, but only a partial perspective on the nature and timing of UP occupations, and limited data on how human groups in this region responded to the harsh climatic oscillations between 37,000–11,500 years before present. Here we report new UP archaeological sequences from fieldwork in Satsurblia cavein the same region. A series of living surfaces with combustion features, faunal remains, stone and bone tools, and ornaments provide new information about human occupations in this region (a) prior to the Last Glacial Maximum (LGM) at 25.5–24.4 ka cal. BP and (b) after the LGM at 17.9–16.2 ka cal. BP. The latter provides new evidence in the southern Caucasus for human occupation immediately after the LGM. The results of the campaigns in Satsurblia and Dzudzuana suggest that at present the most plausible scenario is one of a hiatus in the occupation of this region during the LGM (between 24.4–17.9 ka cal. BP). Analysis of the living surfaces at Satsurblia offers information about human activities such as the production and utilisation of lithics and bone tools, butchering, cooking and consumption of meat and wild cereals, the utilisation of fibers, and the use of certain woods. Microfaunal and palynological analyses point to fluctuations in the climate with consequent shifts in vegetation and the faunal spectrum not only before and after the LGM, but also during the two millennia following the end of the LGM.     

The relationship between mammal faunas and climatic instability since the Last Glacial Maximum: A Nearctic vs. Western Palearctic comparison

Climate has played a key role in shaping the geographic patterns of biodiversity. The imprint of Quaternary climatic fluctuations is particularly evident on the geographic distribution of Holarctic faunas, which dramatically shifted their ranges following the alternation of glacial-interglacial cycles during the Pleistocene. Here, we evaluate the existence of differences between climatically stable and unstable regions – defined on the basis of climatic change velocity since the Last Glacial Maximum – in the geographic distribution of several biological attributes of extant terrestrial mammals of the Nearctic and Western Palearctic regions. Specifically, we use a macroecological approach to assess the dissimilarities in species richness, range size, body size, longevity and litter size of species that inhabit regions with contrasting histories of climatic stability. While several studies have documented how the distributional ranges of animals can be affected by long-term historic climatic fluctuations, there is less evidence on the species-specific traits that determine their responsiveness under such climatic instability. We find that climatically unstable areas have more widespread species and lower mammal richness than stable regions in both continents. We detected stronger signatures of historical climatic instability on the geographic distribution of body size in the Nearctic region, possibly reflecting lagged responses to recolonize deglaciated regions. However, the way that animals respond to climatic fluctuations varies widely among species and we were unable to find a relationship between climatic instability and other mammal life-history traits (longevity and litter size) in any of the two biogeographic regions. We, therefore, conclude that beyond some biological traits typical of macroecological analyses such as geographic range size and body size, it is difficult to infer the responsiveness of species distributions to climate change solely based on particular life-history traits.     

Climate and landscape during the Last Glacial Maximum in southwestern Iberia: The small-vertebrate association from the Sala de las Chimeneas,Maltravieso, Extremadura

Here we present the first palaeoenvironmental and palaeoclimatic analysis based on a study of the microvertebates of the Sala de las Chimeneas (Maltravieso Cave, Cáceres). The fauna is ascribed to the end of the Late Pleistocene, as indicated by the presence of Microtus (Iberomys) cabrerae, Microtus agrestis and Arvicola terrestris and by absolute datings of ～ 17 ka BP. The palaeoenvironmental analysis points to a setting within the early part of the Last Glacial Maximum (LGM), with a habitat dominated by woods and humid meadows, a climate (mean annual temperatures) colder than at present (?4 °C), and mean annual precipitation markedly higher (+700 mm) than is currently the case in the area around Cáceres. These data provide new information on the impact of the last cold spells of Marine Isotope Stage 2 (MIS 2) in the Southwest of Europe, in a region where no studies of the microvertebrates of this period had previously been undertaken.     

The role of glacial cycles in promoting genetic diversity in the Neotropics: the case of cloud forests during the Last Glacial Maximum

The increasing aridity during the Last Glacial Maximum (LGM) has been proposed as a major factor affecting Neotropical species. The character and intensity of this change, however, remains the subject of ongoing debate. This review proposes an approach to test contrasting paleoecological hypotheses by way of their expected demographic and genetic effects on Neotropical cloud forest species. We reviewed 48 paleoecological records encompassing the LGM in the Neotropics. The records show contrasting evidence regarding the changes in precipitation during this period. Some regions remained fairly moist and others had a significantly reduced precipitation. Many paleoecological records within the same region show apparently conflicting evidence on precipitation and forest stability. From these data, we propose and outline two demographic/genetic scenarios for cloud forests species based on opposite precipitation regimes: the dry refugia and the moist forests hypotheses. We searched for studies dealing with the population genetic structure of cloud forest and other montane taxa and compared their results with the proposed models. To date, the few available molecular studies show insufficient genetic evidence on the predominance of glacial aridity in the Neotropics. In order to disentangle the climatic history of the Neotropics, the present study calls for a general multi‐disciplinary approach to conduct future phylogeographic studies. Given the contradictory paleoecological information, population genetic data on Neotropical cloud forest species should be used to explicitly test the genetic consequences of competing paleoecological models.     

The biostratigraphic and palaeoenvironmental significance of Lower Cambrian (Botomian) trilobites from the Ak-Kaya section of the Altai Mountains (southern Siberia,Russia)

The presence of one of the oldest records of polycystine Radiolaria in the Lower Cambrian sedimentary sequence of the Ak-Kaya section (Gorny Altai) requires a biostratigraphic dating. The trilobites found recently a few tens of meters below the radiolarian-bearing level belong to Calodiscus resimus Repina, Serrodiscus fossuliferus Repina and Alacephalus aff. contortus Repina; they suggest that this part of the Shashkunar Formation can be correlated with the lower part of the Botomian stage. The absence of eyes in the two former species suggests a mode of life buried in the fine pelagic sediments. Indications of the presence of a strongly developed musculature on the third species point to a palaeoenvironment characterized by a relatively high hydrodynamic regime.     

Phylogeography of <Emphasis Type="Italic">Littorina sitkana</Emphasis> in the northwestern Pacific Ocean: evidence of eastward trans-Pacific colonization after the Last Glacial Maximum

We investigated genetic diversity and population structure of the Sitka periwinkle Littorina sitkana along the coastlines of the northwestern Pacific (NWP) to evaluate the possibility of trans-Pacific colonization of this species from the NWP to the northeastern Pacific (NEP) after the Last Glacial Maximum. We sampled L. sitkana from 32 populations in the NWP, and sequenced a region of the mitochondrial cytochrome b oxidase gene for population genetic analyses. The results were compared with those of previous reports from the NEP. The genetic diversity of L. sitkana was much higher in the NWP than in the NEP. Genetic connectivity between the NWP and NEP populations was indicated by an extremely abundant haplotype in the NEP that was also present in eastern Hokkaido and the Kuril Islands. To confirm these results, we compared sequences of the longest intron of the aminopeptidase N gene (APN54) in the nuclear genome in four populations of L. sitkana in the NWP with previous results from the NEP. Again, much higher genetic diversity was found in the NWP than in the NEP and genetic connectivity was supported between the Kuril Islands and the NEP. These results imply postglacial colonization of this species from the NWP to the NEP, probably along the Kuril and Aleutian Island chains. This study is the first report of possible trans-Pacific postglacial colonization of a direct-developing gastropod, inferred from genetic data.     

Brazilian populations of Drosophila maculifrons (Diptera: Drosophilidae): low diversity levels and signals of a population expansion after the Last Glacial Maximum

The Quaternary period was marked by considerable changes in climate. Such palaeoclimatic changes affected the population dynamics of many species, both in the Northern and in the Southern Hemisphere. However, the extent of these impacts on the demographic patterns of Neotropical species presenting different ecological requirements remains unclear. Drosophila maculifrons DUDA 1947 belongs to the guaramunu group of Drosophila and represents a potential indicator of the genetic consequences caused by the climatic fluctuations of the Quaternary, because it seems to be sensitive to temperature and humidity shifts. The aim of this study was to evaluate the evolutionary processes subjacent to the patterns of intraspecific diversity and structure of different populations of D. maculifrons. In total, 152 individuals were collected in the south and south‐east Brazil. Phylogenetic and phylogeographical analyses were performed based on sequences of COI and COII mitochondrial genes. In general, the results pointed to Brazilian populations of D. maculifrons being extremely impoverished in terms of mitochondrial diversity and population structure, which could be explained by a recent population expansion event dated to approximately 12 000 years ago. In fact, with the assistance of species palaeo‐distribution modelling strategies, it was possible to infer that most of the sampled region did not present the D. maculifrons environmental suitability requirements at least during the period of the Last Glacial Maximum. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 112 , 55–66.     

Glacial Holocene environment of the southeastern Okhotsk Sea: evidence from geochemical and palaeontological data

Environmental conditions and productivity changes in the southeastern Okhotsk Sea have been reconstructed for the last 20 ka using planktonic and benthic foraminiferal oxygen isotope records and calcium carbonate, organic carbon and opal content data from two sediment cores. Species variability in benthic foraminiferal and diatom assemblages provides additional palaeoceanographic evidence. AMS radiocarbon dating of the sediments and oxygen isotope stratigraphy serve as the basis for the age models of the cores for the last 20 ¹⁴C kyr and for correlation between environmental variations in the Okhotsk Sea, and regional and global climate changes. Benthic foraminiferal assemblages in the two cores (depth 1590 and 1175 m) varied with time, so that we could recognise seven zones with different species composition. Changes in the benthic foraminiferal assemblages parallel major environmental and productivity variations. During the last glaciation, fluxes of organic matter to the sea floor showed strong seasonal variations, indicated by the presence of abundant A. weddellensis and infaunal Uvigerina spp. Benthic foraminiferal assemblages changed with warming at 12.5–11 and 10–8 ¹⁴C kyr BP, when productivity blooms and high organic fluxes were coeval with global meltwater pulses 1A and 1B. Younger Dryas cooling caused a decline in productivity (11–10 kyr BP) affecting the benthic faunal community. Subsequent warming triggered intensive diatom production, opal accumulation and a strong oxygen deficiency, causing significant changes in benthic fauna assemblages from 5.26–4.4 kyr BP to present time.     

Firm evidence for a post-extinction ichnofauna: earliest Carboniferous Cruziana reticulata assemblage from the Anti-Atlas of Morocco

Cruziana reticulata is an arthropod-related ichnospecies that is characterized by a conspicuous net-like scratch pattern whose initial formation and later preservation require the presence of consolidated substrates in shallow marine fine-grained bottoms. There are two scenarios in which epifaunal to shallow infaunal benthic organisms may access firm siliciclastic substrates: first, by exposure of the compacted bottom after erosion of the upper, water-saturated and usually soft portion of sediment column; and, second, by primary fast substrate stabilization in the absence of biogenic sediment mixing. Whereas the first mode occurs throughout the Phanerozoic, the latter is only prevalent in marine bottoms during the Early Palaeozoic mainly as a consequence of poorly developed infaunal bioturbation. However, by eradicating burrowing organisms, mass extinctions are known (i.e. the end-Permian extinction) to ‘reset’ intensity of ecospace utilization, which entails the return to this anachronistic style of trace fossil preservation in younger times. An earliest Carboniferous ichno-assemblage dominated by Cruziana reticulata from Morocco attests the spread of firm substrates in the aftermath of the Hangenberg Event – a major extinction at the end of the Devonian period. Since evidence for erosion is lacking in accompanying sedimentary rocks and the ichno-assemblage shows characteristics of opportunistic exploitation of the nutritious muddy seabed, we favour the interpretation of this assemblage to represent a post-extinction ichnofauna. It shows that other taxonomically less severe mass extinctions may also exhibit transient but severe ecological effects in open marine ecosystems such as the collapse of vital sediment mixing. Such post-extinction effects may distort the perception of the stratigraphical record as a firmground ichno-assemblage often taken as evidence for submarine erosion and are crucial in highlighting sequence boundaries. We, thus, encourage appreciating macroevolutionary framework of respective ichnofaunas.