赛罕乌拉国家级自然保护区红角鸮(Otus scops)的繁殖生态观察

为了解红角鸮(Otus scops)的繁殖状态,2009年5～8月,在内蒙古赛罕乌拉国家级自然保护区,以直接观察法对红角鸮的繁殖生态进行了研究。结果表明,所观察的4个巢址全部是以柳树为主干的树洞巢,巢内铺垫物较少或无铺垫,巢距地面的平均高度为91.3cm,巢内径平均为13.5cm,巢外径平均为15.0cm,巢深平均32.0cm(n=4)。平均窝卵数为3.75,对其中一窝进行测量,4枚卵平均量度是28.9mm×24.6mm,重量9.5g。雏鸟孵出后,雌性亲鸟承担主要育雏任务,雄性亲鸟日间停歇于离巢不远的树上负责警戒工作,夜间与雌鸟共同哺育雏鸟。小型鼬科动物的捕食可能是干扰红角鸮繁殖成功率的主要因素。     

Long‐term decline despite conservation efforts questions Eurasian Stone‐curlew population viability in intensive farmlands

Agricultural intensification over the past decades has led to a generalized decline in farmland biodiversity. Farmland birds are particularly exposed to rapid changes in habitat and reduced food resources or availability. Understanding how farmland specialists can be preserved and their populations enhanced are major challenges for this century. Based on a long‐term (19‐year) study of a Eurasian Stone‐curlew Burhinus oedicnemus population, we estimated the demographic parameters, including clutch size, egg volume, hatching success, survival rate and apparent population size. Demographic rates found for this French population were, on average, comparable to those found elsewhere in Europe. However, all demographic parameters showed negative trends, including a dramatic decline in the local population (26% decline over 14 years) and a 10% decline in adult survival rate over 11 years. Such a long‐term decline, despite on‐going conservation efforts, calls into question the overall sustainability of arable Stone‐curlew populations. We infer some of the possible causes of this decline, in particular food shortage, and discuss how this pattern could be reversed through conservation measures applicable at very large spatial scales.     

Food and habitats requirements of the Eurasian Scops Owl (Otus scops) in Switzerland revealed by very high-resolution multi-scale models

In Europe, agricultural practices have progressively evolved towards high productivity leading either to the intensification of productive and accessible areas or to the abandonment of less profitable sites. Both processes have led to the degradation of semi-natural habitats like extensive grasslands, threatening species such as the Eurasian Scops Owl Otus scops that rely on extensively managed agricultural landscapes. In this work, we aimed to assess the habitat preferences of the Scops Owl using habitat suitability models combined with a multi-scale approach. We generated a set of multi-scale predictors, considering both biotic and abiotic variables, built on two newly developed vegetation management and orthopteran abundance models. To select the variables to incorporate in a ‘best multi-scale model’, we chose the best spatial scale for each variable using univariate models and by calculating their relative importance through multi-model inference. Next, we built ensembles of small models (ESMs) at 10 different scales from 50 to 1000 m, and an additional model with each variable at its best scale (‘best multi-scale model’). The latter performed better than most of the other ESMs and allowed the creation of a high-resolution habitat suitability map for the species. Scops Owls showed a preference for dry sites with extensive and well-structured habitats with 30–40% bush cover, and relied strongly on semi-extensive grasslands covering at least 30% of the surface within 300 m of the territory centre and with high orthopteran availability near the centre (50-m radius), revealing a need for good foraging grounds near the nest. At a larger spatial scale within a radius of 1000 m, the habitat suitability of Scops Owls was negatively related to forest cover. The resulting ESM predictions provide valuable tools for conservation planning, highlighting sites in need of particular conservation efforts together with offering estimates of the percentage of habitat types and necessary prey abundance that could be used as targets in future management plans to ensure the persistence of the population.     

Disease prevalence in an urban raptor related to pest species: The case of Eurasian Scops Owl Otus scops infection by Gongylonema sp.

Urbanization is often associated with homogenization, including the homogenization of biodiversity and overpopulation by generalist species that can change community and disease dynamics. In Madrid, nestlings of Eurasian Scops Owl Otus scops frequently suffer necrotic oropharyngeal disease as a result of infection by Gongylonema sp., a parasitic nematode transmitted by adult owls to their offspring through the diet, more specifically through consumption of a pest species: the oriental cockroach Blatta orientalis. We studied how the presence of cockroaches and owl infection rates are affected by urban features such as green areas. We found that a higher number of birds were affected by the parasite in the initial period of the breeding season and in areas with greater surface area of green spaces. We suggest that urban management should promote the diversity of insects in green areas so that the owls diversify their dietary offer to their offspring, to reduce the prevalence of the disease and improve their breeding success in the city of Madrid, as this problem has not yet been described in other cities.     

The consistency and stability of abundance-occupancy relationships in large-scale population dynamics

1. Abundance-occupancy relationships comprise some of the most general and well-explored patterns in macro-ecology. The theory governing these relationships predicts that species will exhibit a positive interspecific and intraspecific relationship between regional occupancy and local abundance. Abundance-occupancy relationships have important implications in using distributional surveys, such as atlases, to understand and document large-scale population dynamics and the consequences of environmental change. A basic need for interpreting such data bases is a better understanding of whether changes in regional occupancy reflect changes in local abundance across species of varying life-history characteristics. 2. Our objective was to test the predictions of the abundance-occupancy rule using two independent data sets, the New York State Breeding Bird Atlas and the North American Breeding Bird Survey. The New York State Breeding Bird Atlas consists of 5332 25-km(2) survey blocks and is one of the first atlases in the USA to be completed for two time periods (1980-85 and 2000-05). The North American Breeding Survey is a large-scale annual survey intended to document the relative abundance and population change of songbirds throughout the USA. 3. We found that regional occupancy was positively correlated with relative abundance across 98 (beta = 0.60 +/- 0.11 SE, P < 0.001, R(2) = 0.60) and 85 species (beta = 0.67 +/- 0.06 SE, P < 0.001, R(2) = 0.57) in two separate time periods. This relationship proved stable over time and was notably consistent between breeding habitat groups and migratory guilds. 4. Between 1980 and 2005, changes in regional occupancy were highly correlated with long-term abundance trend estimates for 75 species (beta = 5.73 +/- 0.24 SE, P < 0.001, R(2) = 0.88). Over a 20-year period, woodland and resident birds showed an increase in occupancy while grassland species showed the greatest decline; these patterns were mirrored by changes in local abundance. 5. Although exceptions existed, we found most changes in occupancy parallel changes in local abundance. These findings support the basic predictions of the abundance-occupancy rule and demonstrate its consistency and stability in species and groups of varying life-history characteristics.     

Mark–recapture may reveal more about ecology than about population trends: Demography of a threatened ghost bat (Macroderma gigas) population

Since European settlement in Australia, the geographical range of ghost bats (Macroderma gigas) has contracted northwards. Ghost bats are thought to occur in disjunct populations with little interpopulation migration, raising concerns over the current status and future viability of the southernmost colony, which has also been threatened by mining activity. To address these concerns, demographic parameters of the southernmost colony were estimated from a mark–recapture study conducted during 1975–1981. Female bats gave birth to a single young in late spring, but only 40% (22–70%, 95% CI) of females bred in their second year, increasing to 93% (87–97%, 95% CI) for females ≥ 2 years old. Sixty‐five percent of juveniles caught were female. Annual adult survival ranged between 0.57–0.77 for females and 0.43–0.66 for males, and was lowest over winter–spring and greatest in autumn–winter. Juvenile survival for the first year ranged between 0.35–0.46 for females and 0.29–0.42 for males. Adult survival varied among seasons, was negatively associated with rainfall, but was not associated with temperature beyond being lower in late winter. Poor survival may result from the inferior daytime roosts that bats must use if water seepage forces them to leave their normal roosts. Although these age‐specific rates of fecundity and survival suggested a declining population, mark–recapture estimates of the population trend indicated stability over the study period. Counts at daytime roosts also suggested a population decline, but were considered unreliable because of an increasing tendency of bats to avoid detection. It is therefore likely that some assumptions in estimating survival were violated. These results provide a caution against the uncritical use of population projections derived from mark–recapture estimates of demographic parameters, and the use of untested indices as the basis for conservation decisions.     

The demography of a newly established Osprey Pandion haliaetus population in France

Using long‐term mark–resighting data acquired over 27 years in continental France, we estimated demographic parameters and modelled the dynamics of a newly established population of Ospreys Pandion haliaetus using a life‐history model. We then performed prospective and retrospective analyses to estimate the sensitivity of the population growth rate to demographic parameters, and to quantify their contribution to the observed variation in abundance. The observed population growth rate was estimated at 1.150 (from one to 38 pairs in the period 1985–2011), and the stochastic population growth rate was estimated at 1.156. The number of fledglings per nest made the largest contribution to the variance of the observed population growth rate. Breeding productivity was stable across years. In contrast, the prospective analysis indicated that the sensitivity of the population growth rate was greatest for immigration and adult survival. Our results suggest that the increase of a new and recently established breeding population of Ospreys was mainly driven by local dynamics (high productivity and high proportion of breeding individuals), with no sign of density‐dependence except for juvenile survival. This probably reflects highly favourable conditions for breeding. Our results show that productivity can be a major driver in recovering raptor populations, and conservation work should aim to protect occupied nest‐sites and their surrounding habitat and to maintain highly favourable foraging areas in the vicinity of breeding sites.     

Effective population size as a driver for divergence of an antimicrobial peptide (Hymenoptaecin) in two common European bumblebee species

Social insects are the target of numerous pathogens. This is because the high density of closely‐related individuals frequently interacting with each other enhances the transmission and establishment of pathogens. This high selective pressure results in the rapid evolution of immune genes, which might be counteracted by a reduced effective population size (N_e) lowering the effectiveness of selection. We tested the effect of N_e on the evolutionary rate of an important immune gene for the antimicrobial peptide Hymenoptaecin in two common central European bumblebee species: Bombus terrestris and Bombus lapidarius. Both species are similar in their biology and are expected to be under similar selective pressures because pathogen prevalence does not differ between species. However, previous studies indicated a higher N_e in B. terrestris compared to B. lapidarius. We found high intraspecific variability in the coding sequence but low variability for silent polymorphisms in B. lapidarius. Estimates of long‐ and short‐term N_e were three‐ to four‐fold higher N_e in B. terrestris, although the species did not differ in census population sizes. The difference in N_e might result in less efficient selection and suboptimal adaptation of immune genes (e.g. hymenoptaecin) in B. lapidarius, and thus this species might become less resistant and more tolerant, turning into a superspreader of diseases.     

Temporal and spatial dynamics of amphioxus population (Branchiostoma belcheri tsingtaneuse) and its influential factors in Luan River Estuary,China

Population distribution of amphioxus (Branchiostoma belcheri tsingtauense) was investigated in the Luan River Estuary from 1999 to 2011, to describe its trends and discover the factors that influence its decline. Amphioxus are distributed on the seabed at 5–10 m depth, between the Xinkai Estuary and the Dapu River Estuary, where the primary sand components of the sediment are medium and fine, with particle size ranging from 0.001 to 1 mm. In recent years, the population density and biomass of amphioxus have sharply decreased. Changes in sediment granularity composition may significantly influence amphioxus distribution. Our data showed that the highest density region of amphioxus occurred where >90% of the sediment particles were between 0.063 and 0.5 mm in size. A reduction in sediment discharge from Luan River and the expansion of raft‐breeding mariculture may be causing the decline in amphioxus through habitat destruction.     

Long‐term breeding demography and density dependence in an increasing population of Golden Eagles Aquila chrysaetos

Few studies have quantified the dynamics of recovering populations of large raptors using long‐term, spatially explicit studies. Using data collected over 37 years in the western Italian Alps, we assessed the trends in distribution, abundance, fecundity and breeding population structure of Golden Eagles Aquila chrysaetos. Using the spatial distribution of territory centroids in 2007, we found that the spatial distribution of eagle territories was over‐dispersed up to 3 km. Although population size and total productivity increased from 1972 to 2008, the proportion of pairs that laid eggs showed a strong decline, falling to no more than 50% after 2003. On average, 15% of successful nests produced two fledglings, and productivity also declined over time. No significant relationship between population growth rate and total population size was detected, but the percentage of pairs that bred and breeding success showed evidence of density dependence, as they declined significantly with increasing density. Our results suggest that density dependence, operating across heterogeneous habitats, is currently regulating this population, while the carrying capacity may still be increasing. This may explain the apparent paradox of reduced breeding effort despite increasing total productivity.     

The population genomic basis of geographic differentiation in North American common ragweed (Ambrosia artemisiifolia L.)

Common ragweed (Ambrosia artemisiifolia L.) is an invasive, wind‐pollinated plant nearly ubiquitous in disturbed sites in its eastern North American native range and present across growing portions of Europe, Africa, Asia, and Australia. Phenotypic divergence between European and native‐range populations has been described as rapid evolution. However, a recent study demonstrated major human‐mediated shifts in ragweed genetic structure before introduction to Europe and suggested that native‐range genetic structure and local adaptation might fully explain accelerated growth and other invasive characteristics of introduced populations. Genomic differentiation that potentially influenced this structure has not yet been investigated, and it remains unclear whether substantial admixture during historical disturbance of the native range contributed to the development of invasiveness in introduced European ragweed populations. To investigate fine‐scale population genetic structure across the species' native range, we characterized diallelic SNP loci via a reduced‐representation genotyping‐by‐sequencing (GBS) approach. We corroborate phylogeographic domains previously discovered using traditional sequencing methods, while demonstrating increased power to resolve weak genetic structure in this highly admixed plant species. By identifying exome polymorphisms underlying genetic differentiation, we suggest that geographic differentiation of this important invasive species has occurred more often within pathways that regulate growth and response to defense and stress, which may be associated with survival in North America's diverse climatic regions.     

The evolution of microendemism in a reef fish (Hypoplectrus maya)

Marine species tend to have extensive distributions, which are commonly attributed to the dispersal potential provided by planktonic larvae and the rarity of absolute barriers to dispersal in the ocean. Under this paradigm, the occurrence of marine microendemism without geographic isolation in species with planktonic larvae poses a dilemma. The recently described Maya hamlet (Hypoplectrus maya, Serranidae) is exactly such a case, being endemic to a 50‐km segment of the Mesoamerican Barrier Reef System (MBRS). We use whole‐genome analysis to infer the demographic history of the Maya hamlet and contrast it with the sympatric and pan‐Caribbean black (H. nigricans), barred (H. puella) and butter (H. unicolor) hamlets, as well as the allopatric but phenotypically similar blue hamlet (H. gemma). We show that H. maya is indeed a distinct evolutionary lineage, with genomic signatures of inbreeding and a unique demographic history of continuous decrease in effective population size since it diverged from congeners just ~3,000 generations ago. We suggest that this case of microendemism may be driven by the combination of a narrow ecological niche and restrictive oceanographic conditions in the southern MBRS, which is consistent with the occurrence of an unusually high number of marine microendemics in this region. The restricted distribution of the Maya hamlet, its decline in both census and effective population sizes, and the degradation of its habitat place it at risk of extinction. We conclude that the evolution of marine microendemism can be a fast and dynamic process, with extinction possibly occurring before speciation is complete.     

Food quantity–quality co‐limitation: Interactive effects of dietary carbon and essential lipid supply on population growth of a freshwater rotifer

相似文献   

Life history and population dynamics of southern Alaska resident killer whales (Orcinus orca)

Resident (fish eating) killer whales (Orcinus orca) in the North Pacific have been the subject of long‐term studies in several geographical regions. The current study examines population parameters in the southern Alaska resident population from 1984 to 2010 and develops a population model. The southern Alaska resident population ranges from southeastern Alaska through the Kodiak archipelago and contains over 700 individuals. We follow the life histories of 343 identifiable whales in 10 pods from two clans born before and during the study. Population parameters were comparable to those of the British Columbia northern resident population during the 1970s and 1980s, except that age of maturity was approximately one year earlier. The average annual rate of increase was slightly higher in Alaska (3.5%) than for the British Columbia northern residents (2.9%) and probably represents a population at r‐max (maximum rate of growth). Reasons for the high growth rate in Alaska could be a recovery following past anthropogenic mortalities, or more likely, a response to increasing salmon returns in recent decades, resulting in an increase in carrying capacity. The slow maturation and low rate of reproductive response makes these whales slow to recover from natural or anthropogenic catastrophes.     

Spatiotemporal population dynamics of the banana rind thrips,Elixothrips brevisetis (Bagnall) (Thysanoptera: Thripidae)

The understanding of how environmental factors and agricultural practices affect population dynamics of insect pests is necessary for pest management. Here, we provide insight into the ecology of the banana rind thrips Elixothrips brevisetis (Bagnall) (Thysanoptera: Thripidae) by collecting and analysing a spatiotemporal database of population estimates in Martinique (West French Indies). We assessed the influence of climatic variables (which were rainfall and temperature) and biotic variables (which were banana and three weed species) on the adult thrips abundance for different components of the banana plant (sucker, mother plant and bunch) and evaluated the effect of thrips abundance and standard bunch covers on damages. The abundance of thrips on the sucker, the mother plant, and the bunch was significantly related to the abundance on neighbouring banana plants, and spatial autocorrelation indicated that E. brevisetis dispersed for only short distances. The number of thrips on the mother plant and on the bunch was positively related to the number of thrips on the sucker, suggesting that the thrips may disperse from the sucker to the mother plant and then to the bunch. The abundance of thrips on the sucker increased with sucker height and was positively correlated with the mean daily rainfall during the 17 days before sampling; the length of that period might correspond with the time required for an individual to complete its life cycle. Covered bunches had 98% fewer thrips than non‐covered bunches, and the damage caused by thrips was linearly related to the number of thrips present between the 2nd and 4th week after flowering. Finally, we found that the presence of Alocasia cucullata, Dieffenbachia seguine and Peperomia pellucida is significantly related with a decrease in thrips abundance on banana plants, suggesting the use of these weeds as potential trap plants.     

Evaluating the existence and benefit of major histocompatibility complex‐based mate choice in an isolated owl population

How mate preferences evolve in the first place has been a major conundrum for sexual selection. Some hypotheses explaining this assume fitness benefit derived from subsequent generations. Major histocompatibility complex (MHC)‐based mate choice is a representative example of the mate choice that is associated with such trans‐generational mechanisms. To provide evidences for fitness benefit of MHC‐based mate choice, previous studies assessed the association between own MHC genotype and own fitness components. However, the association between MHC‐based mate choice in the parental generation and fitness components in the resultant offspring generation has only rarely been measured in wild populations. Focusing on the isolated population of the monogamous Ryukyu Scops Owl (Otus elegans interpositus) on Minami‐daito Island, Japan, we found evidence of MHC‐based mate choice. However, we found no evidence of MHC‐based mate choice increasing own reproductive success or offspring survival. This is a rare case study that directly examines the existence of the trans‐generational indirect benefit of MHC‐based mate choice for genetic compatibility from trans‐generational data in a wild bird population. By investigating the fitness benefits of mate choice, this study serves to facilitate our understanding of the evolution of MHC‐based mate choice.     

Variations in age‐ and sex‐specific survival rates help explain population trend in a discrete marine mammal population

Understanding the drivers underlying fluctuations in the size of animal populations is central to ecology, conservation biology, and wildlife management. Reliable estimates of survival probabilities are key to population viability assessments, and patterns of variation in survival can help inferring the causal factors behind detected changes in population size. We investigated whether variation in age‐ and sex‐specific survival probabilities could help explain the increasing trend in population size detected in a small, discrete population of bottlenose dolphins Tursiops truncatus off the east coast of Scotland. To estimate annual survival probabilities, we applied capture–recapture models to photoidentification data collected from 1989 to 2015. We used robust design models accounting for temporary emigration to estimate juvenile and adult survival, multistate models to estimate sex‐specific survival, and age models to estimate calf survival. We found strong support for an increase in juvenile/adult annual survival from 93.1% to 96.0% over the study period, most likely caused by a change in juvenile survival. Examination of sex‐specific variation showed weaker support for this trend being a result of increasing female survival, which was overall higher than for males and animals of unknown sex. Calf survival was lower in the first than second year; a bias in estimating third‐year survival will likely exist in similar studies. There was some support first‐born calf survival being lower than for calves born subsequently. Coastal marine mammal populations are subject to the impacts of environmental change, increasing anthropogenic disturbance and the effects of management measures. Survival estimates are essential to improve our understanding of population dynamics and help predict how future pressures may impact populations, but obtaining robust information on the life history of long‐lived species is challenging. Our study illustrates how knowledge of survival can be increased by applying a robust analytical framework to photoidentification data.     

Quantifying the links between land use and population growth rate in a declining farmland bird

Land use is likely to be a key driver of population dynamics of species inhabiting anthropogenic landscapes, such as farmlands. Understanding the relationships between land use and variation in population growth rates is therefore critical for the management of many farmland species. Using 24 years of data of a declining farmland bird in an integrated population model, we examined how spatiotemporal variation in land use (defined as habitats with “Short” and “Tall” ground vegetation during the breeding season) and habitat‐specific demographic parameters relates to variation in population growth taking into account individual movements between habitats. We also evaluated contributions to population growth using transient life table response experiments which gives information on contribution of past variation of parameters and real‐time elasticities which suggests future scenarios to change growth rates. LTRE analyses revealed a clear contribution of Short habitats to the annual variation in population growth rate that was mostly due to fledgling recruitment, whereas there was no evidence for a contribution of Tall habitats. Only 18% of the variation in population growth was explained by the modeled local demography, the remaining variation being explained by apparent immigration (i.e., the residual variation). We discuss potential biological and methodological reasons for high contributions of apparent immigration in open populations. In line with LTRE analysis, real‐time elasticity analysis revealed that demographic parameters linked to Short habitats had a stronger potential to influence population growth rate than those of Tall habitats. Most particularly, an increase of the proportion of Short sites occupied by Old breeders could have a distinct positive impact on population growth. High‐quality Short habitats such as grazed pastures have been declining in southern Sweden. Converting low‐quality to high‐quality habitats could therefore change the present negative population trend of this, and other species with similar habitat requirements.     

Phylogeography and demographic history of Gyrodactylus konovalovi (Monogenoidea: Gyrodactylidae), an ectoparasite on the East Asia Amur minnow (Cyprinidae) in Central China

Gyrodactylus konovalovi is an ectoparasite on the Amur minnow (Rhynchocypris lagowskii) that is widely distributed in the cold fresh waters of East Asia. In the present study, the phylogeography and demographic history of G. konovalovi and the distribution of its host in the Qinling Mountains are examined. A total of 79 individual parasites was sequenced for a 528 bp region of the mitochondrial NADH dehydrogenase subunit 5 (ND5) gene, and 25 haplotypes were obtained. The substitution rate (dN/dS) was 0.068 and indicated purifying selection. Haplotype diversity (h) and nucleotide diversity (π) varied widely in the Qinling Mountains. Phylogenetic trees based on Bayesian inference (BI), maximum likelihood (ML), and maximum parsimony (MP) methods and network analysis revealed that all haplotypes were consistently well‐supported in three different lineages, indicating a significant geographic distribution pattern. There was a significant positive correlation between genetic differentiation (F_st) and geographic distance. The results of mismatch distribution, neutrality test and Bayesian skyline plot analyses showed that whole populations underwent population contraction during the Pleistocene. Based on the molecular clock calibration, the most common ancestor was estimated to have emerged in the middle Pleistocene. Our study suggests for the first time that a clearly phylogeography of G. konovalovi was shaped by geological events and climate fluctuations, such as orogenesis, drainage capture changes, and vicariance, during the Pleistocene in the Qinling Mountains.     

Genetic structure and demographic history of Lymantria dispar (Linnaeus, 1758) (Lepidoptera: Erebidae) in its area of origin and adjacent areas

We analyzed the population genetic structure and demographic history of 20 Lymantria dispar populations from Far East Asia using microsatellite loci and mitochondrial genes. In the microsatellite analysis, the genetic distances based on pairwise F_ST values ranged from 0.0087 to 0.1171. A NeighborNet network based on pairwise F_ST genetic distances showed that the 20 regional populations were divided into five groups. Bayesian clustering analysis (K = 3) demonstrated the same groupings. The populations in the Korean Peninsula and adjacent regions, in particular, showed a mixed genetic pattern. In the mitochondrial genetic analysis based on 98 haplotypes, the median‐joining network exhibited a star shape that was focused on three high‐frequency haplotypes (Haplotype 1: central Korea and adjacent regions, Group 1; Haplotype 37: southern Korea, Group 2; and Haplotype 90: Hokkaido area, Group 3) connected by low‐frequency haplotypes. The mismatch distribution dividing the three groups was unimodal. In the neutral test, Tajima's D and Fu's FS tests were negative. We can thus infer that the Far East Asian populations of L. dispar underwent a sudden population expansion. Based on the age expansion parameter, the expansion time was inferred to be approximately 53,652 years before present (ybp) for Group 1, approximately 65,043 ybp for Group 2, and approximately 76,086 ybp for Group 3. We propose that the mixed genetic pattern of the inland populations of Far East Asia is due to these expansions and that the inland populations of the region should be treated as valid subspecies that are distinguishable from other subspecies by genetic traits.