The causes and scope of political egalitarianism during the Last Glacial: a multi-disciplinary perspective

This paper reviews and synthesizes emerging multi-disciplinary evidence toward understanding the development of social and political organization in the Last Glacial. Evidence for the prevalence and scope of political egalitarianism is reviewed and the biological, social, and environmental influences on this mode of human organization are further explored. Viewing social and political organization in the Last Glacial in a much wider, multi-disciplinary context provides the footing for coherent theory building and hypothesis testing by which to further explore human political systems. We aim to overcome the claim that our ancestors’ form of social organization is untestable, as well as counter a degree of exaggeration regarding possibilities for sedentism, population densities, and hierarchical structures prior to the Holocene with crucial advances from disparate disciplines.     

After Last Glacial Maximum: The third migration

A critical analysis of “Real World” data concerning the genetic origins of people, archaeology and palaeoclimatic conditions, demonstrates possibilities of population migration for third time in ancient history, from East to West after Last Glacial Maximum (LGM), which gave the foundations of modern human civilization.     

Quantitative paleo-estimation: hypothetical experiments with extrapolation and the no-analog problem

We experiment with artificial data to test the response of five numerical techniques in extrapolating paleo-environments for no-analog conditions. No-analog conditions are those beyond the technique calibration (modern) data set and will be encountered in applications to the geologic past, though they may not be easy to recognize. In the ideal, a numerical technique will correctly extrapolate to no-analog conditions. Failing this, the technique will have a consistent, predictable error response to increasing no-analog conditions, as these are measured by a reliable index. The no-analog conditions that we used are a natural extension of the calibration conditions we created. Thus we test techniques for their response to shifting environmental conditions rather than for factors unrelated to the ecology of the taxa (e.g. post-depositional fossil preservation). Five numerical techniques we test with our hypothetical data are (1) multivariate regression of species percents, (2) correlation-based principal components with linear regression, (3) covariance-based principal components with linear regression, (4) correlation-based principal components with non-linear regression, and (5) the Imbrie and Kipp technique. All the techniques show increasing estimation error as conditions depart from those of the calibration data set. There are two main causes of error in our estimates: (1) the distorting effects of matrix closure on taxon abundances; and (2) generation of ratio no-analogs among species abundances because of non-linear responses to conditions departing progressively from the calibration range. With all the techniques, the distribution of error for no-analog conditions is complex. Non-linear regression with factors shows the least predictable error response. We found that currently developed no-analog indicators do not have a good correlation to estimation error. This means that better indicators, more closely linked to the accuracy of estimates, need to be developed.     

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。     

Evidence for a lacustrine faunal refuge in the Larsemann Hills, East Antarctica, during the Last Glacial Maximum

Aim  There is no previous direct evidence for the occurrence of lacustrine refuges for invertebrate fauna in Antarctica spanning the Last Glacial Maximum (LGM). In the absence of verified LGM lacustrine refuges many species are believed to result from Holocene dispersal from sub-Antarctic islands and continents further north. If freshwater lake environments were present throughout the LGM, extant freshwater species may have been associated with Antarctica prior to this glacial period. This study looked at faunal microfossils in a sediment core from an Antarctic freshwater lake. This lake is unusual in that, unlike most Antarctic lakes, the sediment record extends to c . 130,000 yr bp , i.e. prior to the LGM.
Location  Lake Reid, Larsemann Hills, East Antarctica (76°23' E; 69°23' S).
Methods  Palaeofaunal communities in Lake Reid were identified through examination of faunal microfossils in a sediment core that extended to c . 130,000 yr bp .
Results  Ephippia and mandibles from the cladoceran Daphniopsis studeri and loricae of the rotifer Notholca sp. were found at all depths in the sediment, indicating that these two species have been present in the lake for up to 130,000 years. Copepod mandibles were also present in the older section of the core, yet were absent from the most recent sediments, indicating extinction of this species from Lake Reid during the LGM.
Main conclusion  The presence of D. studeri and Notholca sp. microfossils throughout the entire Lake Reid core is the first direct evidence of a glacial lacustrine refugium for invertebrate animals in Antarctica, and indicates the presence of a relict fauna on the Antarctic continent.     

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.     

Palaeoenvironment and palaeoclimate in the western Liguria region (northwestern Italy) during the Last Glacial. The small mammal sequence of Riparo Mochi (Balzi Rossi,Ventimiglia)

Riparo Mochi is considered a key site for the Middle–Upper Palaeolithic chrono-cultural sequence in the Mediterranean area. Climatic and environmental conditions have been reconstructed after the study of the small mammal assemblage by means of linear regression method and the Habitat Weighting method. The climate proxies for the late Mousterian and Proto-Aurignacian (cultural units I to G) indicate an environment with a relative high percentage of forest component during two cold and one relative warm oscillations. Open environments with low temperatures are registered for the Gravettian (cultural Unit D) allowing a possible relation to the Heinrich 3 Event. The final Gravettian (cultural Unit C) shows a relative warm peak, tentatively related to Greenland Interstadial 2. The late Epigravettian (cultural Unit A) is characterized by a high percentage of forest-related environment suggesting that this horizon formed during the Bølling–Allerød Interstadial. Finally, the Riparo Mochi sequence contributes to reconstruct the processes of faunal changes along the northern Tyrrhenian coast in a period of climatic instability, especially during the end of Marine Isotope Stage 3 when foraging groups adopted different mobility strategies between southern France and the Italian peninsula.     

The biogeography of aquatic macrophytes in North America since the Last Glacial Maximum

Aim To document the post‐glacial migration of the major aquatic macrophytes of North America. Location North America north of Mexico. Methods Aquatic macrophyte pollen were extracted from the North American Pollen Database. The modern pollen distribution was mapped and related to the climate to document the geographical and climatic constraints on these taxa. The fossil pollen were mapped at 2‐ka intervals for the past 21 ka. Results Numerous genera were present in ice‐free Alaska during the Last Glacial Maximum, and south of the Laurentide Ice Sheet in the southeast. Those taxa with the widest modern climatic ranges migrated rapidly into ice‐marginal areas, first in the west and then in the east of North America. Subsequent changes in the range and abundance were smaller. Main conclusions There were four migration routes of aquatic macrophytes during the late‐glacial and post‐glacial periods: a southward migration from Alaska between 14–13 and ka, a northern migration in the west at the same time into the ice‐free Cordilleran region, and movements east and west of Appalachia as early as 19 ka for some taxa into the lower Mississippi and into the upper Mississippi and Great Lakes by 11 ka. As the Laurentide ice sheet wasted, aquatic taxa with the broadest contemporary temperature tolerances rapidly occupied ice‐marginal environments.     

Predictive modelling of tree species distributions on the Iberian Peninsula during the Last Glacial Maximum and Mid-Holocene

This paper reports a bioclimatic envelope model study of the potential distribution of 19 tree species in the Iberian Peninsula during the Last Glacial Maximum (LGM; 21 000 yr BP) and the Mid-Holocene (6000 yr BP). Current patterns of tree species richness and distributions are believed to have been strongly influenced by the climate during these periods. The modelling employed novel machine learning techniques, and its accuracy was evaluated using a threshold-independent method. Two atmospheric general circulation models, UGAMP and ECHAM3 (generated by the Palaeoclimate Modelling Intercomparison Project, PMIP), were used to provide climate scenarios under which the distributions of the 19 tree species were modelled. The results obtained for these scenarios were assessed by agreement measure analysis; they differed significantly for the LGM, but were more similar for the Mid-Holocene.
The results for the LGM support the inferred importance of pines in the Iberian Peninsula at this time, and the presence of evergreen Quercus in the south. Important differences in the altitude at which the modelled species grew were also predicted. During the LGM, some normally higher mountain species potentially became re-established in the foothills of the Pyrenees. The warm Mid-Holocene climate is clearly reflected in the predicted expansion of broad-leaved forests during this period, including the colonization of the northern part of the Iberian Peninsula by evergreen Quercus species.     

Glacial refugia influence plant diversity patterns in the Mediterranean Basin

Aim The aims of this study were to assess the distribution of putative Mediterranean refugia of plants, to compare the locations of refugia and those of regional hotspots of plant biodiversity, and to provide a critical analysis of the Mediterranean refugium paradigm. Furthermore, we consider how biogeographical and genetic results can be combined to guide global conservation strategies. Location The Mediterranean region. Methods We started from a detailed analysis of the scientific literature (1993–2007) in order to identify refugia in the Mediterranean region, based on intra‐specific phylogeographical studies of plant species. We used population locations together with gene‐pool identity to establish the database, comparing patterns of phylogeographical concordance with the locations of Mediterranean refugia. We then tested the biogeographical congruence between two biodiversity components, namely phylogeographical refugia and regional hotspots. Results We identified 52 refugia in the Mediterranean bioclimatic region and confirmed the role played by the three major peninsulas, with a shared total of 25 refugia. We emphasize the importance of areas that have previously been attributed a lesser role (large Mediterranean islands, North Africa, Turkey, Catalonia). Of the 52 refugia identified, 33 are situated in the western Mediterranean Basin and 19 in the eastern part. The locations of the phylogeographically defined refugia are significantly associated with the 10 regional hotspots of plant biodiversity, with 26 of these refugia (i.e. 50%) occurring within the hotspots. Main conclusions The locations of refugia are determined by complex historical and environmental factors, the cumulative effects of which need to be considered because they have occurred since the Tertiary, rather than solely during the last glacial period. Refugia represent climatically stable areas and constitute a high conservation priority as key areas for the long‐term persistence of species and genetic diversity, especially given the threat posed by the extensive environmental change processes operating in the Mediterranean region. The refugia defined here represent ‘phylogeographical hotspots’; that is, significant reservoirs of unique genetic diversity favourable to the evolutionary processes of Mediterranean plant species.     

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.     

Seasonal and geographic climate variabilities during the Last Glacial Maximum in North America: Applying isotopic analysis and macrophysical climate models

Climate models provide estimates of climatic change over periods of time in the ancient past. Macrophysical climate models (MCM) differ from the more widely used general circulation models (GCM), in that MCMs provide temporally high-resolution (~ 100 years) and site-specific estimations of monthly values of climate variables such as temperature and precipitation. In this paper, seasonal changes in climate variables are modeled for six ¹⁴C-dated fossil localities in North America. Five of these localities represent the time of maximum extent of ice during the most recent glacial episode, the Full Glacial (25 + –15 ka), including one at the peak of the Last Glacial Maximum (17–15 ka). The other locality represents the time as the ice began to recede, the Late Glacial (15–11 ka). Seasonal variations in temperature and precipitation modeled by MCM are herein compared with interpretations of seasonal variation based upon oxygen isotopes from serially sampled hypsodont teeth (mostly Equus and Bison) collected from each of these localities. Additionally, the MCM-modeled seasonal variations are used to predict the expected abundances of different plant functional groups (PFG) during those times, especially C3 and C4 functional groups, using modern relationships. These predictions are compared with carbon isotopic values from the same teeth. The importance of atmospheric pCO² for the relative abundance of plants utilizing the C4 metabolic pathway is discussed, given that glacial episodes are known to have been times of lower atmospheric pCO². Interpretations of seasonal variability and the relative abundance of C3 versus C4 vegetation based upon isotopes from tooth enamel are in broad agreement with predictions using the MCM and the modern distribution of PFGs with climate variables. The influence of pCO² on the distribution of C4 vegetation during glacial times appears to be negligible.     

Spatial variability in species' potential distributions during the Last Glacial Maximum under different Global Circulation Models: Relevance in evolutionary biology

Ecological niche models (ENM) have been used to reconstruct potential distributions during the Last Glacial Maximum (LGM)—or other time periods—and this use is increasingly common in zoological studies. For this reason, we urgently need understanding factors affecting these predictions. Here, we examine how the use of different Global Circulation Models (GCMs) affects the variability in species' potential distributions during the LGM and how the degree of model extrapolation and its associated uncertainty depends on the GCM used. We develop these issues using two North American shrews, Notiosorex crawfordi and Cryptotis alticola, inhabiting two environmentally different regions. First, we compared paleoclimates in these two regions simulated by three GCMs: Community Climate System Model (CCSM), Model for Interdisciplinary Research on Climate (MIROC), and the Max‐Planck‐Institute für Meteorologie model (MPI). Then, we used maxent to estimate the LGM potential distribution of these two mammals under the three GCMs to assess the spatial variability and extrapolation uncertainty associated with idiosyncrasies of GCM. MIROC estimated noticeably more different climatic conditions than CCSM and MPI in the study areas during the LGM, and its pattern of environmental conditions was distributed differently. The MIROC scenario suggested a remarkable different prediction of potential distribution for both species, being more dramatic for the high mountain shrew, C. alticola. In particular, climatic differences among GCMs resulted in differences in the factors that limit and drive the potential distribution of the species during the LGM. Equally dramatic was the disagreement of extrapolation areas among GCMs. MIROC showed a greater number of pixels where extrapolation is required in both regions. Our findings should be taken into consideration when identifying areas of endemism, dynamic geographic barriers, and glacial refugia. When projecting into alternative scenarios of LGM climate, the idiosyncrasies of each GCM should be explicitly taken into account.     

Millennial Climatic Fluctuations Are Key to the Structure of Last Glacial Ecosystems

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.     

Small-scale distribution of a cnidarian population in the western Mediterranean

The neritic area of the coast of Catalonia (north–westMediterranean) has a continental shelf narrower in the northernhalf and broader in the southern half. This area is crossedfrom north to south by a coastal current whose intensity decreasesto the south. Both conditions, together with other local geographicaland hydrographical features, are thought to be the key factorsin determining the distribution of planktonic cnidarian assemblages,along two main variational axes: a north-south axis and a coast-opensea axis. Aiming to find support to this hypothesis, the planktoniccnidarians collected in two seasons of the year 1983 have beenstudied. The first season (April–May) corresponds to thehighest abundance and heterogeneity, and the second (September–October),to the epoch when the cnidarian population is lower, but constant,along the coastal area. In a global principal component analysisthe factor that best explains the group distribution in thearea is the coastal character, as given by the bottom depthof the sampling station. The number of individuals is greaternear the coast (due to the dominance of a euryhaline species,Muggiaea atlantica), but the species number increases towardsthe open sea (due to the addition of more oceanic species, suchas Narcomedusae and Trachymedusae and the effect of a permanenthydrographic front at the shelf-slope boundary). The secondfactor is the latitudinal axis. The northern half of the coastis under the influence of waters favouring a greater numberof species and individuals. On the contrary, the southern halfshows a less marked influence of the north–south current,their cnidarian assemblages being more constant over time. Althoughthe global cnidarian population follows this general pattern,the most abundant and frequent species (M.atlantica and Aglaurahemistoma) show a broad distribution over all the studied areabecause they are rather insensitive to hydrographical variations.     

Geochemical and Microbiological Studies of Nitrous Oxide Variations within the New NEEM Greenland Ice Core during the Last Glacial Period

Deep polar ice cores provide atmospheric records of nitrous oxide (N₂O) and other trace gases reflecting climate history along with a parallel archive of microbial cells transported with mineral dust, marine and volcanic aerosols from around the globe. Our interdisciplinary study of 32 samples from different depths of the recently drilled NEEM Greenland ice core addressed the question whether the identified microorganisms were capable of post-depositional biological production of N₂O in situ. We used high-resolution geochemical and microbiological approaches to examine the N₂O concentrations, the quantitative distributions of dust, Ca⁺², NH₄⁺ and NO₃^? ions related to N cycle pathways, the microbial abundance and diversity at specific NEEM core depths from 1758 m to 1867.8 m. Results showed varying concentrations of N₂O (220–271.5 ppb). Microbial abundance fluctuated between 3.3 × 10⁴ and 3.3 × 10⁶ cells mL^?1 in direct correlation with dust and Ca²⁺ concentrations with higher cell numbers deposited during colder periods. The average values of NH₄⁺ and NO₃^? indicated that substrates were available for the microorganisms capable of utilizing them. PCR amplification of selected functional genes involved in bacterial and archaeal nitrification and denitrification was not successful. Sanger and Illumina MiSeq sequence analyses of SSU rRNA genes showed variable representation of Alpha-, Beta- and Gammaproteobacteria, Firmicutes, Actinobacteria, chloroplasts and fungi. The metabolic potential of the dominant genera of Proteobacteria and Firmicutes as possible N₂O producers suggested that denitrification activity may have led to in-situ production and accumulation of N₂O.     

Phylogeographic analyses suggest that a deciduous species (Ostryopsis davidiana Decne., Betulaceae) survived in northern China during the Last Glacial Maximum

Aim Palaeontologial data suggest that all temperate forest species in northern China migrated southwards during the Last Glacial Maximum (LGM) and recolonized post‐glacially within the Holocene. We tested this assumption using phylogeographical studies of a temperate deciduous shrub species (Ostryopsis davidiana Decne., Betulaceae), which has a wide distribution in northern China. Location Northern China. Methods We sequenced two chloroplast DNA (cpDNA) fragments (trnL–trnF and psbA–trnH, together about 1300 bp in length) of 294 plants from 21 populations across the total distribution range of this species. We used maximum parsimony and haplotype network methods to construct phylogenetic relationships among haplotypes. Results The analysis of cpDNA variation identified eight haplotypes. A single haplotype was fixed in all populations except for one population that was polymorphic, having two haplotypes. The population subdivisions were extremely high (G_ST = 0.972 and N_ST = 0.974), suggesting very low gene flow between populations. Haplotypes clustered into two tentative clades, both of which occur in the southern region of the species’ range but only one of which occurs in the northern region. Across the sampled populations, the haplotype distributions were differentiated geographically. Main conclusions Our analyses suggest that multiple refugia were maintained across the range of O. davidiana in both northern (north of the Qing Mountains) and southern (south of the Qing Mountains) regions during the LGM rather than that the species survived only in the south and subsequently colonized northwards. The extremely low within‐population diversity of this species suggests strong bottleneck or founder effects within each fragmented region during the Quaternary climatic oscillations. These findings provide important clues for understanding range shifts and changes in within‐ and/or between‐population genetic diversity of temperate forests in response to past climatic oscillations in northern China.