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Species area relationships in mediterranean-climate plant communities

Aim To determine the best-fit model of species–area relationships for Mediterranean-type plant communities and evaluate how community structure affects these species–area models. Location Data were collected from California shrublands and woodlands and compared with literature reports for other Mediterranean-climate regions. Methods The number of species was recorded from 1, 100 and 1000 m 2 nested plots. Best fit to the power model or exponential model was determined by comparing adjusted r 2 values from the least squares regression, pattern of residuals, homoscedasticity across scales, and semi-log slopes at 1–100 m 2 and 100–1000 m 2 . Dominance–diversity curves were tested for fit to the lognormal model, MacArthur's broken stick model, and the geometric and harmonic series. Results Early successional Western Australia and California shrublands represented the extremes and provide an interesting contrast as the exponential model was the best fit for the former, and the power model for the latter, despite similar total species richness. We hypothesize that structural differences in these communities account for the different species–area curves and are tied to patterns of dominance, equitability and life form distribution. Dominance–diversity relationships for Western Australian heathlands exhibited a close fit to MacArthur's broken stick model, indicating more equitable distribution of species. In contrast, Californian shrublands, both postfire and mature stands, were best fit by the geometric model indicating strong dominance and many minor subordinate species. These regions differ in life form distribution, with annuals being a major component of diversity in early successional Californian shrublands although they are largely lacking in mature stands. Both young and old Australian heathlands are dominated by perennials, and annuals are largely absent. Inherent in all of these ecosystems is cyclical disequilibrium caused by periodic fires. The potential for community reassembly is greater in Californian shrublands where only a quarter of the flora resprout, whereas three quarters resprout in Australian heathlands. Other Californian vegetation types sampled include coniferous forests, oak savannas and desert scrub, and demonstrate that different community structures may lead to a similar species–area relationship. Dominance–diversity relationships for coniferous forests closely follow a geometric model whereas associated oak savannas show a close fit to the lognormal model. However, for both communities, species–area curves fit a power model. The primary driver appears to be the presence of annuals. Desert scrub communities illustrate dramatic changes in both species diversity and dominance–diversity relationships in high and low rainfall years, because of the disappearance of annuals in drought years. Main conclusions Species–area curves for immature shrublands in California and the majority of Mediterranean plant communities fit a power function model. Exceptions that fit the exponential model are not because of sampling error or scaling effects, rather structural differences in these communities provide plausible explanations. The exponential species–area model may arise in more than one way. In the highly diverse Australian heathlands it results from a rapid increase in species richness at small scales. In mature California shrublands it results from very depauperate richness at the community scale. In both instances the exponential model is tied to a preponderance of perennials and paucity of annuals. For communities fit by a power model, coefficients z and log c exhibit a number of significant correlations with other diversity parameters, suggesting that they have some predictive value in ecological communities.

Journal of Biogeography↗

Twentieth century demographic changes in cirio and cardón in Baja California, Mexico

Aim&ensp; Our purpose was to discern long-term large-scale patterns of survivorship and recruitment of two dominant, charismatic and useful desert plants, cirio [Fouquieriaceae; Fouquieria columnaris (Kell.) Kell. ex Curran] and card&oacute;n [Cactaceae; Pachycereus pringlei (S.Watson) Britton & Rose], and to evaluate the effects of physical and human factors. Location&ensp; The study included 77 sites distributed over c. 13,200 km 2 in the northern Vizca&iacute;no Region of the Sonoran Desert, state of Baja California, M&eacute;xico. Method&ensp; Censuses used identified individuals ( n = 3780 cirio and 2246 card&oacute;n) and were based on repeat photography. Time intervals between censuses ranged from 29 to 96 years, ending in 1996&ndash;2002. Earlier repetition (1963) of old photographs at 14 sites allowed analysis of temporal variation. The effect of elapsed time was modelled with Weibull functions for survivorship and linear functions for recruitment. To explain the residual variations, we tested categorical and continuous variables related to substrate, geomorphology, climate and geography. Human impacts were weighed with historical and quantitative analysis. Results&ensp; Elapsed time accounted for 45% of the variation of survivorship in cirio but only 35% in card&oacute;n. The fitted Weibull curves were used to estimate longevity: all-size cohorts of 1000 individuals would die out in 388 (cirio) and 723 years (card&oacute;n). Recruitment in cirio was linearly related to time ( r 2 = 0.71) and averaged < 0.003 year &minus;1 but was less than mortality at 90% of the sites. For card&oacute;n, recruitment averaged > 0.005 year &minus;1 , was linearly related to time ( r 2 = 0.66) and was less than mortality at only 33% of the sites. Vital rates were not strongly correlated between mid- and late-century but were mostly similar to the long-term regional rates. Removing the factor of elapsed time, the residual variations showed that survivorship was greater on sites protected from winds for cirio and was less on very coarse soils for card&oacute;n. Recruitment increased with latitude in cirio and was greater on southern exposures for both species. The residual variations were not clearly affected, at a regional scale, by other factors thought to be important, including elevation, distance to the Pacific Ocean, geology, slope gradient, soil stability, older vs. young alluvial soils, and soil Ca : Mg and Na : K ratios. Human impacts have been sporadic and heterogeneous but locally strong; our quantitative indices of accessibility did not show regionally significant effects. Blowdown by hurricanes is a sporadic cause of major mortality for cirio but not necessarily for card&oacute;n. Main conclusions&ensp; At our scales, effects of time outweigh those of substrate, and human impact was scant. Large patterns were pronounced: cirio experienced a prolonged, widespread decline in the 1900s, while card&oacute;n fluctuated in different directions and degree among local populations. Cirio was found to be inherently much shorter-lived than card&oacute;n. We also suggest that recruitment in cirio was low due to long periods with relatively dry winters that affected the entire region, while spatial heterogeneity of card&oacute;n recruitment seemed more related to the variation in summer rains.

Journal of Biogeography↗

Analysis of vegetation distribution in Interior Alaska and sensitivity to climate change using a logistic regression approach

Aim: To understand drivers of vegetation type distribution and sensitivity to climate change. Location: Interior Alaska. Methods: A logistic regression model was developed that predicts the potential equilibrium distribution of four major vegetation types: tundra, deciduous forest, black spruce forest and white spruce forest based on elevation, aspect, slope, drainage type, fire interval, average growing season temperature and total growing season precipitation. The model was run in three consecutive steps. The hierarchical logistic regression model was used to evaluate how scenarios of changes in temperature, precipitation and fire interval may influence the distribution of the four major vegetation types found in this region. Results: At the first step, tundra was distinguished from forest, which was mostly driven by elevation, precipitation and south to north aspect. At the second step, forest was separated into deciduous and spruce forest, a distinction that was primarily driven by fire interval and elevation. At the third step, the identification of black vs. white spruce was driven mainly by fire interval and elevation. The model was verified for Interior Alaska, the region used to develop the model, where it predicted vegetation distribution among the steps with an accuracy of 60-83%. When the model was independently validated for north-west Canada, it predicted vegetation distribution among the steps with an accuracy of 53-85%. Black spruce remains the dominant vegetation type under all scenarios, potentially expanding most under warming coupled with increasing fire interval. White spruce is clearly limited by moisture once average growing season temperatures exceeded a critical limit (+2 ??C). Deciduous forests expand their range the most when any two of the following scenarios are combined: decreasing fire interval, warming and increasing precipitation. Tundra can be replaced by forest under warming but expands under precipitation increase. Main conclusion: The model analyses agree with current knowledge of the responses of vegetation types to climate change and provide further insight into drivers of vegetation change. ?? 2005 Blackwell Publishing Ltd.

Journal of Biogeography↗

A 40,000-year woodrat-midden record of vegetational and biogeographical dynamics in north-eastern Utah

Aim A conspicuous climatic and biogeographical transition occurs at 40-45° N in western North America. This pivot point marks a north–south opposition of wet and dry conditions at interannual and decadal time-scales, as well as the northern and southern limits of many dominant western plant species. Palaeoecologists have yet to focus on past climatic and biotic shifts along this transition, in part because it requires comparisons across dissimilar records [i.e. pollen from lacustrine sediments to the north and plant macrofossils from woodrat ( Neotoma ) middens to the south]. To overcome these limitations, we are extending the woodrat-midden record northward into the lowlands of the central Rocky Mountains. Location Woodrat middens were collected from crevices and rock shelters on south-facing slopes of Dutch John Mountain (2000-2200 m, 40°57′ N, 109°25′ W), situated on the eastern flanks of the Uinta Mountains in north-eastern Utah. The site is near the regional limits for Pinus ponderosa, P. edulis, P. contorta, Cercocarpus ledifolius var. intricatus, Abies concolor, Ephedra viridis and other important western species. Methods We analysed pollen and plant macrofossils from the 40,000-year midden sequence. The middens represent brief, depositional episodes (mostly years to decades). Four middens represent the early to full-glacial period (40,000–18,000 cal-yr bp), eight middens are from the late-glacial/early Holocene transition (13,500–9000 cal yr bp), and 33 middens span the mid-to-late Holocene (last 7500 years). Temporal density of our Holocene middens (one every c. 210 years) is comparable with typical Holocene pollen sequences from lake sediments. Results Early to full-glacial assemblages are characterized by low diversity and occurrence of montane conifers ( Picea pungens, Pseudotsuga menziesii, P. flexilis, Juniperus communis ) absent from the site today. Diversity increases in the late-glacial samples with the addition of J. scopulorum, J. horizontalis, C. montanus, C. ledifolius var. intricatus and mesic understory species. The coniferous trees and J. communis declined and J. osteosperma appeared during the late-glacial/Holocene transition. Juniperus osteosperma populations have occupied the site throughout the Holocene. Pinus ponderosa was established by 7500 cal-yr bp, and has occurred at least locally ever since. Montane conifers and J. horizontalis persisted until c. 5500 cal-yr bp. The signature events of the late Holocene were the invasions of P. edulis and Ephedra viridis and establishment of pinyon–juniper woodland in the last 800 years. Main conclusions The Dutch John Mountain midden record adds to an emerging picture in which mid-elevation conifers ( P. flexilis, Pseudotsuga menziesii, Picea pungens, J. scopulorum, J. communis ) dominated vegetation over a wide area of the Colorado Plateau and adjacent Rocky Mountains. Rather than being fragmented, as often assumed in phylogeographical studies, these species had broader and more-connected distributions than they do in the region today. Paradoxically, subalpine conifers ( Picea engelmannii, A. lasiocarpa ) occurred at higher elevations to the south, possibly representing declining precipitation from south to north owing to southward displacement of the polar jet stream. The Dutch John Mountain record displays a series of extinction and invasion events. Most of the extinctions were local in scale; nearly all constituents of fossil midden assemblages occur within a few kilometres of Dutch John Mountain, and some occur at least locally on its slopes. The sole exception is J. horizontalis , which is regionally extinct. In contrast to extinctions, Holocene invasions were regional in scale; J. osteosperma, P. ponderosa, P. edulis and Ephedra viridis immigrated from glacial-age source populations far to the south.

Utah↗

A land-cover map for South and Southeast Asia derived from SPOT-VEGETATION data

Aim Our aim was to produce a uniform ‘regional’ land-cover map of South and Southeast Asia based on ‘sub-regional’ mapping results generated in the context of the Global Land Cover 2000 project. Location The ‘region’ of tropical and sub-tropical South and Southeast Asia stretches from the Himalayas and the southern border of China in the north, to Sri Lanka and Indonesia in the south, and from Pakistan in the west to the islands of New Guinea in the far east. Methods The regional land-cover map is based on sub-regional digital mapping results derived from SPOT-VEGETATION satellite data for the years 1998–2000. Image processing, digital classification and thematic mapping were performed separately for the three sub-regions of South Asia, continental Southeast Asia, and insular Southeast Asia. Landsat TM images, field data and existing national maps served as references. We used the FAO (Food and Agriculture Organization) Land Cover Classification System (LCCS) for coding the sub-regional land-cover classes and for aggregating the latter to a uniform regional legend. A validation was performed based on a systematic grid of sample points, referring to visual interpretation from high-resolution Landsat imagery. Regional land-cover area estimates were obtained and compared with FAO statistics for the categories ‘forest’ and ‘cropland’. Results The regional map displays 26 land-cover classes. The LCCS coding provided a standardized class description, independent from local class names; it also allowed us to maintain the link to the detailed sub-regional land-cover classes. The validation of the map displayed a mapping accuracy of 72% for the dominant classes of ‘forest’ and ‘cropland’; regional area estimates for these classes correspond reasonably well to existing regional statistics. Main conclusions The land-cover map of South and Southeast Asia provides a synoptic view of the distribution of land cover of tropical and sub-tropical Asia, and it delivers reasonable thematic detail and quantitative estimates of the main land-cover proportions. The map may therefore serve for regional stratification or modelling of vegetation cover, but could also support the implementation of forest policies, watershed management or conservation strategies at regional scales.

Journal of Biogeography↗

Geographical and climatic limits of needle types of one- and two-needled pinyon pines

Aim: The geographical extent and climatic tolerances of one- and two-needled pinyon pines (Pinus subsect. Cembroides) are the focus of questions in taxonomy, palaeoclimatology and modelling of future distributions. The identification of these pines, traditionally classified by one- versus two-needled fascicles, is complicated by populations with both one- and two-needled fascicles on the same tree, and the description of two more recently described one-needled varieties: the fallax-type and californiarum-type. Because previous studies have suggested correlations between needle anatomy and climate, including anatomical plasticity reflecting annual precipitation, we approached this study at the level of the anatomy of individual pine needles rather than species. Location: Western North America. Methods: We synthesized available and new data from field and herbarium collections of needles to compile maps of their current distributions across western North America. Annual frequencies of needle types were compared with local precipitation histories for some stands. Historical North American climates were modelled on a c. 1-km grid using monthly temperature and precipitation values. A geospatial model (ClimLim), which analyses the effect of climate-modulated physiological and ecosystem processes, was used to rank the importance of seasonal climate variables in limiting the distributions of anatomical needle types. Results: The pinyon needles were classified into four distinct types based upon the number of needles per fascicle, needle thickness and the number of stomatal rows and resin canals. The individual needles fit well into four categories of needle types, whereas some trees exhibit a mixture of two needle types. Trees from central Arizona containing a mixture of Pinus edulis and fallax-type needles increased their percentage of fallax-type needles following dry years. All four needle types occupy broader geographical regions with distinctive precipitation regimes. Pinus monophylla and californiarum-type needles occur in regions with high winter precipitation. Pinus edulis and fallax-type needles are found in regions with high monsoon precipitation. Areas supporting californiarum-type and fallax-type needle distributions are additionally characterized by a more extreme May-June drought. Main conclusions: These pinyon needle types seem to reflect the amount and seasonality of precipitation. The single needle fascicle characterizing the fallax type may be an adaptation to early summer or periodic drought, while the single needle of Pinus monophylla may be an adaptation to summer-autumn drought. Although the needles fit into four distinct categories, the parent trees are sometimes less easily classified, especially near their ancestral Pleistocene ranges in the Mojave and northern Sonoran deserts. The abundance of trees with both one- and two-needled fascicles in the zones between P. monophylla, P. edulis and fallax-type populations suggest that needle fascicle number is an unreliable characteristic for species classification. Disregarding needle fascicle number, the fallax-type needles are nearly identical to P. edulis, supporting Little's (1968) initial classification of these trees as P. edulis var. fallax, while the californiarum-type needles have a distinctive morphology supporting Bailey's (1987) classification of this tree as Pinus californiarum.

Journal of Biogeography↗

Mangrove forest distributions and dynamics (1975–2005) of the tsunami-affected region of Asia

Aim We aimed to estimate the present extent of tsunami-affected mangrove forests and determine the rates and causes of deforestation from 1975 to 2005. Location Our study region covers the tsunami-affected coastal areas of Indonesia, Malaysia, Thailand, Burma (Myanmar), Bangladesh, India and Sri Lanka in Asia. Methods We interpreted time-series Landsat data using a hybrid supervised and unsupervised classification approach. Landsat data were geometrically corrected to an accuracy of plus-or-minus half a pixel, an accuracy necessary for change analysis. Each image was normalized for solar irradiance by converting digital number values to the top-of-the atmosphere reflectance. Ground truth data and existing maps and data bases were used to select training samples and also for iterative labelling. We used a post-classification change detection approach. Results were validated with the help of local experts and/or high-resolution commercial satellite data. Results The region lost 12% of its mangrove forests from 1975 to 2005, to a present extent of c . 1,670,000 ha. Rates and causes of deforestation varied both spatially and temporally. Annual deforestation was highest in Burma ( c . 1%) and lowest in Sri Lanka (0.1%). In contrast, mangrove forests in India and Bangladesh remained unchanged or gained a small percentage. Net deforestation peaked at 137,000 ha during 1990–2000, increasing from 97,000 ha during 1975–90, and declining to 14,000 ha during 2000–05. The major causes of deforestation were agricultural expansion (81%), aquaculture (12%) and urban development (2%). Main conclusions We assessed and monitored mangrove forests in the tsunami-affected region of Asia using the historical archive of Landsat data. We also measured the rates of change and determined possible causes. The results of our study can be used to better understand the role of mangrove forests in saving lives and property from natural disasters such as the Indian Ocean tsunami, and to identify possible areas for conservation, restoration and rehabilitation.

Journal of Biogeography↗

Fish assemblages of the Casiquiare River, a corridor and zoogeographical filter for dispersal between the Orinoco and Amazon basins

Aim: The aim of this study was to determine whether the Casiquiare River functions as a free dispersal corridor or as a partial barrier (i.e. filter) for the interchange of fish species of the Orinoco and Negro/Amazon basins using species assemblage patterns according to geographical location and environmental features. Location: The Casiquiare, Upper Orinoco and Upper Negro rivers in southern Venezuela, South America. Methods: Our study was based on an analysis of species presence/absence data and environmental information (11 habitat characteristics) collected by the authors and colleagues between the years 1984 and 1999. The data set consisted of 269 sampled sites and 452 fish species (> 50,000 specimens). A wide range of habitat types was included in the samples, and the collection sites were located at various points along the entire length of the Casiquiare main channel, at multiple sites on its tributary streams, as well as at various nearby sites outside the Casiquiare drainage, within the Upper Orinoco and Upper Rio Negro river systems. Most specimens and field data used in this analysis are archived in the Museo de Ciencias Naturales in Guanare, Venezuela. We performed canonical correspondence analysis (CCA) based on species presence/absence using two versions of the data set: one that eliminated sites having < 5 species and species occurring at < 5 sites; and another that eliminated sites having < 10 species and species occurring at < 10 sites. Cluster analysis was performed on sites based on species assemblage similarity, and a separate analysis was performed on species based on CCA loadings. Results: The CCA results for the two versions of the data set were qualitatively the same. The dominant environmental axis contrasted assemblages and sites associated with blackwater vs. clearwater conditions. Longitudinal position on the Casiquiare River was correlated (r2 = 0.33) with CCA axis-1 scores, reflecting clearwater conditions nearer to its origin (bifurcation of the Orinoco) and blackwater conditions nearer to its mouth (junction with the Rio Negro). The second CCA axis was most strongly associated with habitat size and structural complexity. Species associations derived from the unweighted pair-group average clustering method and pair-wise squared Euclidean distances calculated from species loadings on CCA axes 1 and 2 showed seven ecological groupings. Cluster analysis of species assemblages according to watershed revealed a stronger influence of local environmental conditions than of geographical proximity. Main conclusions: Fish assemblage composition is more consistently associated with local environmental conditions than with geographical position within the river drainages. Nonetheless, the results support the hypothesis that the mainstem Casiquiare represents a hydrochemical gradient between clearwaters at its origin and blackwaters at its mouth, and as such appears to function as a semi-permeable barrier (environmental filter) to dispersal and faunal exchanges between the partially vicariant fish faunas of the Upper Orinoco and Upper Negro rivers. ?? 2008 The Authors.

Journal of Biogeography↗

Climatic water deficit, tree species ranges, and climate change in Yosemite National Park

Aim (1) To calculate annual potential evapotranspiration (PET), actual evapotranspiration (AET) and climatic water deficit (Deficit) with high spatial resolution; (2) to describe distributions for 17 tree species over a 2300-m elevation gradient in a 3000-km 2 landscape relative to AET and Deficit; (3) to examine changes in AET and Deficit between past ( c. 1700), present (1971–2000) and future (2020–49) climatological means derived from proxies, observations and projections; and (4) to infer how the magnitude of changing Deficit may contribute to changes in forest structure and composition. Location Yosemite National Park, California, USA. Methods We calculated the water balance within Yosemite National Park using a modified Thornthwaite-type method and correlated AET and Deficit with tree species distribution. We used input data sets with different spatial resolutions parameterized for variation in latitude, precipitation, temperature, soil water-holding capacity, slope and aspect. We used climate proxies and climate projections to model AET and Deficit for past and future climate. We compared the modelled future water balance in Yosemite with current species water-balance ranges in North America. Results We calculated species climatic envelopes over broad ranges of environmental gradients – a range of 310 mm for soil water-holding capacity, 48.3°C for mean monthly temperature (January minima to July maxima), and 918 mm yr −1 for annual precipitation. Tree species means were differentiated by AET and Deficit, and at higher levels of Deficit, species means were increasingly differentiated. Modelled Deficit for all species increased by a mean of 5% between past ( c. 1700) and present (1971–2000). Projected increases in Deficit between present and future (2020–49) were 23% across all plots. Main conclusions Modelled changes in Deficit between past, present and future climate scenarios suggest that recent past changes in forest structure and composition may accelerate in the future, with species responding individualistically to further declines in water availability. Declining water availability may disproportionately affect Pinus monticola and Tsuga mertensiana . Fine-scale heterogeneity in soil water-holding capacity, aspect and slope implies that plant water balance may vary considerably within the grid cells of kilometre-scale climate models. Sub-grid-cell soil and topographical data can partially compensate for the lack of spatial heterogeneity in gridded climate data, potentially improving vegetation-change projections in mountainous landscapes with heterogeneous topography.

Journal of Biogeography↗

Predicting species distributions from checklist data using site-occupancy models

Aim: (1) To increase awareness of the challenges induced by imperfect detection, which is a fundamental issue in species distribution modelling; (2) to emphasize the value of replicate observations for species distribution modelling; and (3) to show how 'cheap' checklist data in faunal/floral databases may be used for the rigorous modelling of distributions by site-occupancy models. Location: Switzerland. Methods: We used checklist data collected by volunteers during 1999 and 2000 to analyse the distribution of the blue hawker, Aeshna cyanea (Odonata, Aeshnidae), a common dragonfly in Switzerland. We used data from repeated visits to 1-ha pixels to derive 'detection histories' and apply site-occupancy models to estimate the 'true' species distribution, i.e. corrected for imperfect detection. We modelled blue hawker distribution as a function of elevation and year and its detection probability of elevation, year and season. Results: The best model contained cubic polynomial elevation effects for distribution and quadratic effects of elevation and season for detectability. We compared the site-occupancy model with a conventional distribution model based on a generalized linear model, which assumes perfect detectability (p = 1). The conventional distribution map looked very different from the distribution map obtained using site-occupancy models that accounted for the imperfect detection. The conventional model underestimated the species distribution by 60%, and the slope parameters of the occurrence-elevation relationship were also underestimated when assuming p = 1. Elevation was not only an important predictor of blue hawker occurrence, but also of the detection probability, with a bell-shaped relationship. Furthermore, detectability increased over the season. The average detection probability was estimated at only 0.19 per survey. Main conclusions: Conventional species distribution models do not model species distributions per se but rather the apparent distribution, i.e. an unknown proportion of species distributions. That unknown proportion is equivalent to detectability. Imperfect detection in conventional species distribution models yields underestimates of the extent of distributions and covariate effects that are biased towards zero. In addition, patterns in detectability will erroneously be ascribed to species distributions. In contrast, site-occupancy models applied to replicated detection/non-detection data offer a powerful framework for making inferences about species distributions corrected for imperfect detection. The use of 'cheap' checklist data greatly enhances the scope of applications of this useful class of models. ?? 2010 Blackwell Publishing Ltd.

Journal of Biogeography↗

Co-occurrence patterns of trees along macro-climatic gradients and their potential influence on the present and future distribution of Fagus sylvatica L.

During recent and future climate change, shifts in large-scale species ranges are expected due to the hypothesized major role of climatic factors in regulating species distributions. The stress-gradient hypothesis suggests that biotic interactions may act as major constraints on species distributions under more favourable growing conditions, while climatic constraints may dominate under unfavourable conditions. We tested this hypothesis for one focal tree species having three major competitors using broad-scale environmental data. We evaluated the variation of species co-occurrence patterns in climate space and estimated the influence of these patterns on the distribution of the focal species for current and projected future climates. Location: Europe. Methods: We used ICP Forest Level 1 data as well as climatic, topographic and edaphic variables. First, correlations between the relative abundance of European beech (Fagus sylvatica) and three major competitor species (Picea abies, Pinus sylvestris and Quercus robur) were analysed in environmental space, and then projected to geographic space. Second, a sensitivity analysis was performed using generalized additive models (GAM) to evaluate where and how much the predicted F. sylvatica distribution varied under current and future climates if potential competitor species were included or excluded. We evaluated if these areas coincide with current species co-occurrence patterns. Results: Correlation analyses supported the stress-gradient hypothesis: towards favourable growing conditions of F. sylvatica, its abundance was strongly linked to the abundance of its competitors, while this link weakened towards unfavourable growing conditions, with stronger correlations in the south and at low elevations than in the north and at high elevations. The sensitivity analysis showed a potential spatial segregation of species with changing climate and a pronounced shift of zones where co-occurrence patterns may play a major role. Main conclusions: Our Results: demonstrate the importance of species co-occurrence patterns for calibrating improved species distribution models for use in projections of climate effects. The correlation approach is able to localize European areas where inclusion of biotic predictors is effective. The climateinduced spatial segregation of the major tree species could have ecological and economic consequences.

Journal of Biogeography↗

Multilocus phylogeography and population structure of common eiders breeding in North America and Scandinavia

Aim Glacial refugia during the Pleistocene had major impacts on the levels and spatial apportionment of genetic diversity of species in northern latitude ecosystems. We characterized patterns of population subdivision, and tested hypotheses associated with locations of potential Pleistocene refugia and the relative contribution of these refugia to the post-glacial colonization of North America and Scandinavia by common eiders ( Somateria mollissima ). Specifically, we evaluated localities hypothesized as ice-free areas or glacial refugia for other Arctic vertebrates, including Beringia, the High Arctic Canadian Archipelago, Newfoundland Bank, Spitsbergen Bank and north-west Norway. Location Alaska, Canada, Norway and Sweden. Methods Molecular data from 12 microsatellite loci, the mitochondrial DNA (mtDNA) control region, and two nuclear introns were collected and analysed for 15 populations of common eiders ( n = 716) breeding throughout North America and Scandinavia. Population genetic structure, historical population fluctuations and gene flow were inferred using F -statistics, analyses of molecular variance, and multilocus coalescent analyses. Results Significant inter-population variation in allelic and haplotypic frequencies were observed (nuclear DNA F ST = 0.004–0.290; mtDNA Φ ST = 0.051–0.927). Whereas spatial differentiation in nuclear genes was concordant with subspecific designations, geographic proximity was more predictive of inter-population variance in mitochondrial DNA haplotype frequency. Inferences of historical population demography were consistent with restriction of common eiders to four geographic areas during the Last Glacial Maximum: Belcher Islands, Newfoundland Bank, northern Alaska and Svalbard. Three of these areas coincide with previously identified glacial refugia: Newfoundland Bank, Beringia and Spitsbergen Bank. Gene-flow and clustering analyses indicated that the Beringian refugium contributed little to common eider post-glacial colonization of North America, whereas Canadian, Scandinavian and southern Alaskan post-glacial colonization is likely to have occurred in a stepwise fashion from the same glacial refugium. Main conclusions Concordance of proposed glacial refugia used by common eiders and other Arctic species indicates that Arctic and subarctic refugia were important reservoirs of genetic diversity during the Pleistocene. Furthermore, suture zones identified at MacKenzie River, western Alaska/Aleutians and Scandinavia coincide with those identified for other Arctic vertebrates, suggesting that these regions were strong geographic barriers limiting dispersal from Pleistocene refugia.

Journal of Biogeography↗

Land use and climate influences on waterbirds in the Prairie Potholes

Aim We examined the influences of regional climate and land‐use variables on mallard ( Anas platyrhynchos ), blue‐winged teal ( Anas discors ), ruddy duck ( Oxyura jamaicensis ) and pied‐billed grebe ( Podilymbus podiceps ) abundances to inform conservation planning in the Prairie Pothole Region of the United States. Location The US portion of Bird Conservation Region 11 (US‐BCR11, the Prairie Potholes), which encompasses six states within the United States: Montana, North Dakota, South Dakota, Nebraska, Minnesota and Iowa. Methods We used data from the North American Breeding Bird Survey (NABBS), the National Land Cover Data Set, and the National Climatic Data Center to model the effects of environmental variables on waterbird abundance. We evaluated land‐use covariates at three logarithmically related spatial scales (1000, 10,000 and 100,000 ha), and constructed hierarchical spatial count models a priori using information from published habitat associations. Model fitting was performed using a hierarchical modelling approach within a Bayesian framework. Results Models with the same variables expressed at different scales were often in the best model subset, indicating that the influence of spatial scale was small. Both land‐use and climate variables contributed strongly to predicting waterbird abundance in US‐BCR11. The strongest positive influences on waterbird abundance were the percentage of wetland area across all three spatial scales, herbaceous vegetation and precipitation variables. Other variables that we included in our models did not appear to influence waterbirds in this study. Main conclusions Understanding the relationships of waterbird abundance to climate and land use may allow us to make predictions of future distribution and abundance as environmental factors change. Additionally, results from this study can suggest locations where conservation and management efforts should be focused.

Iowa, Minnesota, Montana, Nebraska, North Dakota, ↗

Evolution of natural history information in the 21st century – developing an integrated framework for biological and geographical data

Threats to marine and estuarine species operate over many spatial scales, from nutrient enrichment at the watershed/estuarine scale to invasive species and climate change at regional and global scales. To help address research questions across these scales, we provide here a standardized framework for a biogeographical information system containing queriable biological data that allows extraction of information on multiple species, across a variety of spatial scales based on species distributions, natural history attributes and habitat requirements. As scientists shift from research on localized impacts on individual species to regional and global scale threats, macroecological approaches of studying multiple species over broad geographical areas are becoming increasingly important. The standardized framework described here for capturing and integrating biological and geographical data is a critical first step towards addressing these macroecological questions and we urge organizations capturing biogeoinformatics data to consider adopting this framework.

Journal of Biogeography↗

The human dimension of fire regimes on Earth

Humans and their ancestors are unique in being a fire‐making species, but ‘natural’ (i.e. independent of humans) fires have an ancient, geological history on Earth. Natural fires have influenced biological evolution and global biogeochemical cycles, making fire integral to the functioning of some biomes. Globally, debate rages about the impact on ecosystems of prehistoric human‐set fires, with views ranging from catastrophic to negligible. Understanding of the diversity of human fire regimes on Earth in the past, present and future remains rudimentary. It remains uncertain how humans have caused a departure from ‘natural’ background levels that vary with climate change. Available evidence shows that modern humans can increase or decrease background levels of natural fire activity by clearing forests, promoting grazing, dispersing plants, altering ignition patterns and actively suppressing fires, thereby causing substantial ecosystem changes and loss of biodiversity. Some of these contemporary fire regimes cause substantial economic disruptions owing to the destruction of infrastructure, degradation of ecosystem services, loss of life, and smoke‐related health effects. These episodic disasters help frame negative public attitudes towards landscape fires, despite the need for burning to sustain some ecosystems. Greenhouse gas‐induced warming and changes in the hydrological cycle may increase the occurrence of large, severe fires, with potentially significant feedbacks to the Earth system. Improved understanding of human fire regimes demands: (1) better data on past and current human influences on fire regimes to enable global comparative analyses, (2) a greater understanding of different cultural traditions of landscape burning and their positive and negative social, economic and ecological effects, and (3) more realistic representations of anthropogenic fire in global vegetation and climate change models. We provide an historical framework to promote understanding of the development and diversification of fire regimes, covering the pre‐human period, human domestication of fire, and the subsequent transition from subsistence agriculture to industrial economies. All of these phases still occur on Earth, providing opportunities for comparative research.

Journal of Biogeography↗

Rodent middens reveal episodic, long-distance plant colonizations across the hyperarid Atacama Desert over the last 34,000 years

Aim To document the impact of late Quaternary pluvial events on plant movements between the coast and the Andes across the Atacama Desert, northern Chile. Location Sites are located along the lower and upper fringes of absolute desert (1100&ndash;2800 m a.s.l.), between the western slope of the Andes and the Coastal Ranges of northern Chile (24&ndash;26&deg; S). Methods We collected and individually radiocarbon dated 21 rodent middens. Plant macrofossils (fruits, seeds, flowers and leaves) were identified and pollen content analysed. Midden assemblages afford brief snapshots of local plant communities that existed within the rodents' limited foraging range during the several years to decades that it took the midden to accumulate. These assemblages were then compared with modern floras to determine the presence of extralocal species and species provenance. Results Five middens span the last glacial period (34&ndash;21 ka) and three middens are from the last glacial&ndash;interglacial transition (19&ndash;11 ka). The remaining 13 middens span the last 7000 years. Coastal hyperarid sites exhibit low taxonomic richness in middens at 19.3, 1.1, 1.0, 0.9, 0.5 ka and a modern sample. Middens are also dominated by the same plants that occur today. In contrast, middens dated to 28.1, 21.3, 17.3, 3.7 and 0.5 ka contain more species, including Andean extralocals. Precordillera middens (c. 2700 m) show a prominent increase in plant macrofossil richness, along with the appearance of Andean extralocals and sedges at 34.5 and 18.9 ka. Six younger middens dated to 6.1&ndash;0.1 ka are similar to the modern local vegetation. Main conclusions Increased species richness and Andean extralocal plants occurred along the current lower fringes of absolute desert during the last glacial&ndash;interglacial transition and late Holocene. The absence of soil carbonates indicates the persistence of absolute desert throughout the Quaternary. Colonization by Andean plants could have been accomplished through long-distance seed dispersal either by animals or floods that originated in the Andes. We postulate that dispersal would have been most frequent during regional pluvial events and associated increases in groundwater levels, forming local wetlands in the absolute desert, and generating large floods capable of crossing the Central Depression.

Atacama Desert↗

Patterns and controlling factors of species diversity in the Arctic Ocean

Aim The Arctic Ocean is one of the last near-pristine regions on Earth, and, although human activities are expected to impact on Arctic ecosystems, we know very little about baseline patterns of Arctic Ocean biodiversity. This paper aims to describe Arctic Ocean-wide patterns of benthic biodiversity and to explore factors related to the large-scale species diversity patterns. Location Arctic Ocean. Methods We used large ostracode and foraminiferal datasets to describe the biodiversity patterns and applied comprehensive ecological modelling to test the degree to which these patterns are potentially governed by environmental factors, such as temperature, productivity, seasonality, ice cover and others. To test environmental control of the observed diversity patterns, subsets of samples for which all environmental parameters were available were analysed with multiple regression and model averaging. Results Well-known negative latitudinal species diversity gradients (LSDGs) were found in metazoan Ostracoda, but the LSDGs were unimodal with an intermediate maximum with respect to latitude in protozoan foraminifera. Depth species diversity gradients were unimodal, with peaks in diversity shallower than those in other oceans. Our modelling results showed that several factors are significant predictors of diversity, but the significant predictors were different among shallow marine ostracodes, deep-sea ostracodes and deep-sea foraminifera. Main conclusions On the basis of these Arctic Ocean-wide comprehensive datasets, we document large-scale diversity patterns with respect to latitude and depth. Our modelling results suggest that the underlying mechanisms causing these species diversity patterns are unexpectedly complex. The environmental parameters of temperature, surface productivity, seasonality of productivity, salinity and ice cover can all play a role in shaping large-scale diversity patterns, but their relative importance may depend on the ecological preferences of taxa and the oceanographic context of regions. These results suggest that a multiplicity of variables appear to be related to community structure in this system.

Journal of Biogeography↗

Temporal, spatial and ecological dynamics of speciation among amphi-Beringian small mammals

Quaternary climate cycles played an important role in promoting diversification across the Northern Hemisphere, although details of the mechanisms driving evolutionary change are still poorly resolved. In a comparative phylogeographical framework, we investigate temporal, spatial and ecological components of evolution within a suite of Holarctic small mammals. We test a hypothesis of simultaneous divergence among multiple taxon pairs, investigating time to coalescence and demographic change for each taxon in response to a combination of climate and geography.

Beringia↗