Showing papers in "Journal of Biogeography in 2004"

Animal species diversity driven by habitat heterogeneity/diversity: the importance of keystone structures

Abstract: Aim In a selected literature survey we reviewed studies on the habitat heterogeneity–animal species diversity relationship and evaluated whether there are uncertainties and biases in its empirical support. Location World-wide. Methods We reviewed 85 publications for the period 1960–2003. We screened each publication for terms that were used to define habitat heterogeneity, the animal species group and ecosystem studied, the definition of the structural variable, the measurement of vegetation structure and the temporal and spatial scale of the study. Main conclusions The majority of studies found a positive correlation between habitat heterogeneity/diversity and animal species diversity. However, empirical support for this relationship is drastically biased towards studies of vertebrates and habitats under anthropogenic influence. In this paper, we show that ecological effects of habitat heterogeneity may vary considerably between species groups depending on whether structural attributes are perceived as heterogeneity or fragmentation. Possible effects may also vary relative to the structural variable measured. Based upon this, we introduce a classification framework that may be used for across-studies comparisons. Moreover, the effect of habitat heterogeneity for one species group may differ in relation to the spatial scale. In several studies, however, different species groups are closely linked to ‘keystone structures’ that determine animal species diversity by their presence. Detecting crucial keystone structures of the vegetation has profound implications for nature conservation and biodiversity management.

A world-wide study of high altitude treeline temperatures

Abstract: Aim At a coarse scale, the treelines of the world`s mountains seem to follow a common isotherm, but the evidence for this has been indirect so far. Here we aim at underpinning this with facts. Location We present the results of a data-logging campaign at 46 treeline sites between 68degrees N and 42degrees S. Methods We measured root-zone temperatures with an hourly resolution over 1-3 years per site between 1996 and 2003. Results Disregarding taxon-, landuse- or fire-driven tree limits, high altitude climatic treelines are associated with a seasonal mean ground temperature of 6.7 degreesC (+/-0.8 SD; 2.2 K amplitude of means for different climatic zones), a surprisingly narrow range. Temperatures are higher (7-8 degreesC) in the temperate and Mediterranean zone treelines, and are lower in equatorial treelines (5-6 degreesC) and in the subarctic and boreal zone (6-7 degreesC). While air temperatures are higher than soil temperatures in warm periods, and are lower than soil temperatures in cold periods, daily means of air and soil temperature are almost the same at 6-7 degreesC, a physics driven coincidence with the global mean temperature at treeline. The length of the growing season, thermal extremes or thermal sums have no predictive value for treeline altitude on a global scale. Some Mediterranean (Fagus spp.) and temperate South Hemisphere treelines (Nothofagus spp.) and the native treeline in Hawaii (Metrosideros) are located at substantially higher isotherms and represent genus-specific boundaries rather than boundaries of the life-form tree. In seasonal climates, ground temperatures in winter (absolute minima) reflect local snow pack and seem uncritical. Main conclusions The data support the hypothesis of a common thermal threshold for forest growth at high elevation, but also reflect a moderate region and substantial taxonomic influence.

An evaluation of methods for modelling species distributions

Abstract: Aim Various statistical techniques have been used to model species probabilities of occurrence in response to environmental conditions. This paper provides a comprehensive assessment of methods and investigates whether errors in model predictions are associated to specific kinds of geographical and environmental distributions of species. Location Portugal, Western Europe. Methods Probabilities of occurrence for 44 species of amphibians and reptiles in Portugal were modelled using seven modelling techniques: Gower metric, Ecological Niche Factor Analysis, classification trees, neural networks, generalized linear models, generalized additive models and spatial interpolators. Generalized linear and additive models were constructed with and without a term accounting for spatial autocorrelation. Model performance was measured using two methods: sensitivity and Kappa index. Species were grouped according to their spatial (area of occupancy and extent of occurrence) and environmental (marginality and tolerance) distributions. Two-way comparison tests were performed to detect significant interactions between models and species groups. Results Interaction between model and species groups was significant for both sensitivity and Kappa index. This indicates that model performance varied for species with different geographical and environmental distributions. Artificial neural networks performed generally better, immediately followed by generalized additive models including a covariate term for spatial autocorrelation. Non-parametric methods were preferred to parametric approaches, especially when modelling distributions of species with a greater area of occupancy, a larger extent of occurrence, lower marginality and higher tolerance. Main conclusions This is a first attempt to relate performance of modelling techniques with species spatial and environmental distributions. Results indicate a strong relationship between model performance and the kinds of species distributions being modelled. Some methods performed generally better, but no method was superior in all circumstances. A suggestion is made that choice of the appropriate method should be contingent on the goals and kinds of distributions being modelled.

A new land-cover map of Africa for the year 2000

Abstract: Aim In the framework of the Global Land Cover 2000 (GLC 2000), a land-cover map of Africa has been produced at a spatial resolution of 1 km using data from four sensors on-board four different Earth observing satellites. Location The map documents the location and distribution of major vegetation types and non-vegetated land surface formations for the entire African continent plus Madagascar and the other surrounding islands. Methods The bulk of these data were acquired on a daily basis throughout the year 2000 by the VEGETATION sensor on-board the SPOT-4 satellite. The map of vegetation cover has been produced based upon the spectral response and the temporal profile of the vegetation cover. Digital image processing and geographical information systems techniques were employed, together with local knowledge, high resolution imagery and expert consultation, to compile a cartographic map product. Radar data and thermal sensors were also used for specific land-cover classes. Results A total of 27 land cover categories are documented, which has more thematic classes than previously published land cover maps of Africa contain. Systematic comparison with existing land cover data and 30-m resolution imagery from Landsat are presented, and the map is also compared with other pan-continental land cover maps. The map and digital data base are freely available for non-commercial uses from http://www.gvm.jrc.it/tem/africa/products.htm Main conclusions The map improves our state of knowledge of the land-cover of Africa and presents the most spatially detailed view yet published at this scale. This first version of the map should provide an important input for regional stratification and planning purposes for natural resources, biodiversity and climate studies.

Do we need land‐cover data to model species distributions in Europe?

Abstract: Aim To assess the influence of land cover and climate on species distributions across Europe. To quantify the importance of land cover to describe and predict species distributions after using climate as the main driver. Location The study area is Europe. Methods (1) A multivariate analysis was applied to describe land-cover distribution across Europe and assess if the land cover is determined by climate at large spatial scales. (2) To evaluate the importance of land cover to predict species distributions, we implemented a spatially explicit iterative procedure to predict species distributions of plants (2603 species), mammals (186 species), breeding birds (440 species), amphibian and reptiles (143 species). First, we ran bioclimatic models using stepwise generalized additive models using bioclimatic variables. Secondly, we carried out a regression of land cover (LC) variables against residuals from the bioclimatic models to select the most relevant LC variables. Finally, we produced mixed models including climatic variables and those LC variables selected as decreasing the residual of bioclimatic models. Then we compared the explanatory and predictive power of the pure bioclimatic against the mixed model. Results (1) At the European coarse resolution, land cover is mainly driven by climate. Two bioclimatic axes representing a gradient of temperature and a gradient of precipitation explained most variation of land-cover distribution. (2) The inclusion of land cover improved significantly the explanatory power of bioclimatic models and the most relevant variables across groups were those not explained or poorly explained by climate. However, the predictive power of bioclimatic model was not improved by the inclusion of LC variables in the iterative model selection process. Main conclusion Climate is the major driver of both species and land-cover distributions over Europe. Yet, LC variables that are not explained or weakly associated with climate (inland water, sea or arable land) are interesting to describe particular mammal, bird and tree distributions. However, the addition of LC variables to pure bioclimatic models does not improve their predictive accuracy.

The mid-domain effect applied to elevational gradients: species richness of small mammals in Costa Rica

Abstract: Aim The objective of this study was to comprehensively document and examine the alpha and gamma patterns of species richness in non-volant, small mammals (rodents, shrews and mouse opossums) along a tropical elevational gradient. These data were used to determine the support for existing hypotheses of species richness encompassing mid-domain null models, as well as climatic, and community overlap hypotheses. Location Field studies were conducted along a Caribbean slope of the Rio Penas Blancas watershed in the north-eastern region of Costa Rica between 750 and 1850 m at 10 sampling sites. Methods Species richness and abundances of small mammals were surveyed for four seasons including three temporal replicates at each of five elevational sites: late wet season (2000), early wet season (2001), and dry season (2002), and one spatial replicate at five different sites within the same elevations during the late wet season (2001). Species richness at elevations below 700 m was compiled from specimen records from 23 US national and international collections. Predictions of a null model based solely on geometric constraints were examined using a Monte Carlo simulation program, Mid-Domain Null. Results In 16,900 trap nights, 1561 individuals from 16 species were captured. Both alpha and gamma species richness peaked at mid-elevation between 1000 and 1300 m, with richness declining both at higher and lower elevations. Most of the empirical curves of species richness occur within 95% prediction curves of the mid-domain model, although deviations from the null model exist. Regression of the empirical richness on the null model predictions explained nearly half of the variation observed (r2 = 0.45, P = 0.002). Main conclusions The geometric constraints of montane topography appear to influence the diversity pattern of small mammals, although climatic conditions including an intermediate rainfall and temperature regime, and distance from the persistent cloud cap also are correlated with the pattern of species richness. The predictions of productivity, and community overlap hypotheses are not supported with the empirical data.

Trophic structure patterns of Brazilian reef fishes: a latitudinal comparison

Abstract: Aim To investigate how reef fish trophic structure responds to latitudinal changes, using a simple model: the extensive Brazilian coast. Location Six Brazilian tropical and subtropical coral and rocky coastal reefs, and the oceanic island of Atol das Rocas, between latitudes 0° and 27° S. Methods Underwater visual census data collected by the authors (five locations) or obtained from the literature (two locations) were used to estimate the relative abundance of 123 fish species belonging to 33 reef-associated families. Cryptic species were excluded from the analysis. Fishes were grouped in eight trophic categories: roving herbivores, territorial herbivores, mobile invertebrate feeders, sessile invertebrate feeders, omnivores, planktivores, piscivores and carnivores. After a series of detailed predictions based on phylogeny, physiological constraints and anthropogenic impacts was established, the community trophic structure was analysed along a latitudinal gradient and among coastal, mid-shore and oceanic sites. Results The trophic structure of Brazilian reef fish assemblages clearly changed with latitude. Roving herbivores such as scarids and acanthurids were proportionally more abundant at low latitudes. The browsing herbivores kyphosids followed an opposite latitudinal pattern. The parrotfish genus Sparisoma, more plastic in its feeding habits than Scarus, presented wider distribution. The relative abundance of territorial herbivores did not decrease towards higher latitudes. Mobile invertebrate feeders were the most important (in low latitudes) or the second most important trophic guild (in high latitudes) at all coastal sites. Sessile invertebrate feeders did not show any clear latitudinal trend, despite an expected increase in abundance towards low latitudes. Omnivores dominated high latitude reefs (27° S) and planktivores the oceanic island Atol das Rocas. Piscivores and carnivores were proportionally better represented in high latitudes. Main conclusions Latitudinal patterns seem to be influenced by phylogeny, physiological constraints (mainly related to temperature), and also by anthropogenic impacts. Grazing scarids and acanthurids are largely restricted to tropical reefs and show an abrupt decline beyond 23° S. This does not reflect the amount of algae present, but probably temperature-dependent physiological constraints. Other herbivores seem to overcome this through symbiotic microbial digestive processes (kyphosids), manipulating the structure of algal turfs or increasing animal protein from within the territory (pomacentrids). Omnivores dominate the southern sites Arraial do Cabo and Arvoredo, being more adapted to environment constraints related to seasonal and/or stochastic shifts. Large carnivores (including piscivores) extend farther into high-latitude habitats, apparently not constrained by thermal thresholds that limit the herbivores. Overfishing and/or ornamental harvesting certainly has been modifying local fish communities, but could not be detected properly at the large-scale patterns found in this study. The data presented put in evidence for the first time how reef fish trophic structure behave in the extensive south-western Atlantic latitudinal gradient.

Fern species richness along a central Himalayan elevational gradient, Nepal

Abstract: Aim The study explores fern species richness patterns along a central Himalayan elevational gradient (100–4800 m a.s.l.) and evaluates factors influencing the spatial increase and decrease of fern richness. Location The Himalayas stretch from west to east by 20°, i.e. 75–95° east, and Nepal is located from 80 to 88° east in this range. Methods We used published data of the distribution of ferns and fern allies to interpolate species elevational ranges. Defining species presence between upper and lower elevation limit is the basis for richness estimates. The richness pattern was regressed against the total number of rainy days, and gradients that are linearly related to elevation, such as length of the growing season, potential evapotranspiration (PET, energy), and a moisture index (MI = PET/mean annual rainfall). The regressions were performed by generalized linear models. Results A unimodal relationship between species richness and elevation was observed, with maximum species richness at 2000 m. Fern richness has a unimodal response along the energy gradients, and a linear response with moisture gradients. Main conclusions The study confirms the importance of moisture on fern distributions as the peak coincides spatially with climatic factors that enhance moisture levels; the maximum number of rainy days and the cloud zone. Energy-related variables probably control species richness directly at higher elevations but at the lower end the effect is more probably related to moisture.

Expansion of human settlement in Kenya's Maasai Mara: what future for pastoralism and wildlife?

Abstract: Aim Wildlife and pastoral peoples have lived side-by-side in the Mara ecosystem of south-western Kenya for at least 2000 years. Recent changes in human population and landuse are jeopardizing this co-existence. The aim of the study is to determine the viability of pastoralism and wildlife conservation in Maasai ranches around the Maasai Mara National Reserve (MMNR). Location A study area of 2250 km2 was selected in the northern part of the Serengeti-Mara ecosystem, encompassing group ranches adjoining the MMNR. Emphasis is placed on Koyake Group Ranch, a rangeland area owned by Maasai pastoralists, and one of Kenya's major wildlife tourism areas. Methods Maasai settlement patterns, vegetation, livestock numbers and wildlife numbers were analysed over a 50-year period. Settlement distributions and vegetation changes were determined from aerial photography and aerial surveys of 1950, 1961, 1967, 1974, 1983 and 1999. Livestock and wildlife numbers were determined from re-analysis of systematic reconnaissance flights conducted by the Kenya Government from 1977 to 2000, and from ground counts in 2002. Corroborating data on livestock numbers were obtained from aerial photography of Maasai settlements in 2001. Trends in livestock were related to rainfall, and to vegetation production as indicated by the seasonal Normalized Difference Vegetation Index. With these data sets, per capita livestock holdings were determined for the period 1980–2000, a period of fluctuating rainfall and primary production. Results For the first half of the twentieth century, the Mara was infested with tsetse-flies, and the Maasai were confined to the Lemek Valley area to the north of the MMNR. During the early 1960s, active tsetse-control measures by both government and the Maasai led to the destruction of woodlands across the Mara and the retreat of tsetse flies. The Maasai were then able to expand their settlement area south towards MMNR. Meanwhile, wildebeest (Connochaetes taurinus) from the increasing Serengeti population began to spill into the Mara rangelands each dry season, leading to direct competition between livestock and wildlife. Group ranches were established in the area in 1970 to formalize land tenure for the Maasai. By the late 1980s, with rapid population growth, new settlement areas had been established at Talek and other parts adjacent to the MMNR. Over the period 1983–99, the number of Maasai bomas in Koyake has increased at 6.4% per annum (pa), and the human population at 4.4% pa. Over the same period, cattle numbers on Koyake varied from 20,000 to 45,000 (average 25,000), in relation to total rainfall received over the previous 2 years. The rangelands of the Mara cannot support a greater cattle population under current pastoral practices. Conclusions With the rapid increase in human settlement in the Mara, and with imminent land privatization, it is probable that wildlife populations on Koyake will decline significantly in the next 3–5 years. Per capita livestock holdings on the ranch have now fallen to three livestock units/reference adult, well below minimum pastoral subsistence requirements. During the 1980s and 90s the Maasai diversified their livelihoods to generate revenues from tourism, small-scale agriculture and land-leases for mechanized cultivation. However, there is a massive imbalance in tourism incomes in favour of a small elite. In 1999 the membership of Koyake voted to subdivide the ranch into individual holdings. In 2003 the subdivision survey allocated plots of 60 ha average size to 1020 ranch members. This land privatization may result in increased cultivation and fencing, the exclusion of wildlife, and the decline of tourism as a revenue generator. This unique pastoral/wildlife system will shortly be lost unless land holdings can be managed to maintain the free movement of livestock and wildlife.

The Holocene spread of Picea abies (L.) Karst. in Fennoscandia and adjacent areas

Abstract: Aim The Holocene spread of Picea abies in Fennoscandia is well established from many sites and thus provides an opportunity for detailed study of the dynamics of tree spread and population expansion. Early- and mid-Holocene macrofossil evidence for presence of P. abies in Fennoscandia has questioned traditional interpretations of the timing and direction of its spread. This paper aims to determine when, from where and by which pathways P. abies spread into Fennoscandia. Understanding the character and dynamics of this spread may give insight into the general understanding of Holocene tree spread. Location The north-western distribution of P. abies in Europe, including Norway, Sweden, Finland, Estonia, Latvia, Lithuania, north-western Russia, parts of Byelorussia and Poland. Methods Holocene pollen diagrams with independent dating control were collected from this region. The timing of the onset of the continuous curve, the timing of the rise of the curve, the first appearance of frequencies of 1%, 3%, 5%, and 10%, as well as timing and the maximum amount of P. abies pollen, was obtained from these pollen diagrams. A GIS analysis was used to display the data and interpolate ages over the area under investigation. Results Maps are presented showing a clear ESE to WNW trend in the spread of P. abies for all characters interpolated. The timing of the rise of the curve was difficult to use as sites east of the Baltic have slowly rising P. abies frequencies while the western sites often show a rapid rise. Main conclusions The spread of P. abies in Fennoscandia and adjacent areas can be separated into two phases: (i) A rapid early Holocene spread out of Byelorussia and northern Russia at low population density giving rise to small outpost populations, possibly as far west as the Scandes Mountains. (ii) A mid- to late Holocene front-like spread at high population densities moving from east to west into the Baltic Republics and Finland, into northern Scandinavia and then moving south and west towards its present-day distributional limits.

Historical biogeography of olive domestication (Olea europaea L.) as revealed by geometrical morphometry applied to biological and archaeological material

Abstract: Aim This study intends to improve our understanding of historical biogeography of olive domestication in the Mediterranean Basin, particularly in the north-western area. Location Investigations were performed simultaneously on olive stones from extant wild populations, extant cultivated varieties from various Mediterranean countries, and archaeological assemblages of Spanish, French and Italian settlements. Methods A combination of morphometrics (traditional and geometrical) allowed us to study both the size and shape of endocarp structure. Concerning shape, a size-standardized method coupled with fitted polynomial regression analysis was performed. Results We found morphological criteria for discriminating between wild and cultivated olive cultivars, and established patterns of morphological variation of olive material according to the geographical origin (for extant material) and to the age of the olive forms (for archaeological material). Levels of morphological convergences and divergences between wild olive populations and cultivated varieties are presented as evidence. Main conclusions Morphological changes of endocarps of olive under domestication at both geographical and chronological scales provide new criteria for the identification of olive cultivars. They allow to determine the origins of cultivated forms created and/or introduced in the north-western Mediterranean regions and to understand how human migrations affected the rest of the Western Mediterranean regions. A model of diffusion of olive cultivation is proposed. It shows evidence of an indigenous origin of the domestication process, which is currently recognized in the north-western area since the Bronze Age.

Butterfly species richness in the north-west Mediterranean Basin: the role of natural and human-induced factors

Abstract: Aim We developed a model enabling us to evaluate the contribution of both natural and human-related factors to butterfly species richness in Catalonia, a Mediterranean area that harbours one of the most diverse butterfly faunas in Europe. Location The study was carried out in Catalonia (north-east Iberian Peninsula), a region of 31,930 km2 lying between the Pyrenees, the Ebro depression and the Mediterranean sea. Methods Data from the Catalan Butterfly Monitoring Scheme were used to assess butterfly species richness from 55 transects spread all over the region. Three groups of environmental variables likely to affect the presence of butterfly species were calculated, above all from geographic information system data: (1) climatology and topography, (2) vegetation structure and (3) human disturbance. Because climatic and topographic variables are expected to be strongly correlated, we first performed a principal component analysis (PCA) to create a summarizing factor that would account for most of the variance within this set of variables. Subsequently, a backward stepwise multiple regression was performed in order to assess the effects of environmental factors on butterfly species richness. Results A total of 131 species were detected in the monitoring transects, representing 75.7% of the butterfly fauna known from Catalonia. Mean species richness per transect and per year was 41.4, although values varied greatly among sites (range: 14–76.8). The final regression model explained more than 80% of the total variance, which indicated a strong association between butterfly species richness and the studied environmental factors. The model revealed the very important contribution of climatic and topographic variables, which were combined into a single factor in the PCA. In contrast to what has been found in other, more northerly countries, species richness was negatively correlated with temperature and positively correlated with rainfall, except for extreme cold and wet conditions. This may be a consequence of the predictably adverse effects of the Mediterranean summer drought on herbivorous insects, and the fact that the limits of distribution of many butterflies correlate well with climatic variables. Human disturbance (defined as the proportion of urban and agricultural landscape cover in buffer areas of 5 km around the transect sites) was the second most important predictor for species richness. We found that a significant decrease in species numbers was associated with an increase in human pressure, a finding that indicates that not only building development, but also modern-day agricultural practices are detrimental to the conservation of Mediterranean butterflies. Surprisingly, vegetation variables had an almost negligible effect on butterfly species richness. Main conclusions Our findings strongly indicate that the current motors of global change will have a negative effect on Mediterranean butterfly assemblages. First, changes in land-use are transforming and fragmenting the landscape into an inhospitable and less permeable matrix for butterflies. Secondly, the negative correlation between species richness and temperature will lead to a predictable loss of diversity over the coming years, as predicted in the most plausible scenarios of climate change. Considering the high butterfly richness characterizing the Mediterranean Basin, this future trend will pose a serious threat to biodiversity.

Holocene vegetation change and the mammal faunas of South America and Africa

Abstract: Aim Although sharing many similarities in their vegetation types, South America and Africa harbour very dissimilar recent mammal faunas, not only taxonomically but also in terms of several faunistic patterns. However late Pleistocene and mid-Holocene faunas, albeit taxonomically distinct, presented many convergent attributes. Here we propose that the effects of the Holocene climatic change on vegetation physiognomy has played a crucial role in shaping the extant mammalian faunistic patterns. Location South America and Africa from the late Pleistocene to the present. Methods Data presented here have been compiled from many distinct sources, including palaeontological and neontological mammalian studies, palaeoclimatology, palynology, and publications on vegetation ecology. Data on Pleistocene, Holocene and extant mammal faunas of South America and Africa allowed us to establish a number of similar and dissimilar faunistic patterns between the two continents across time. We then considered what changes in vegetation physiognomy would have occurred under the late Pleistocene last glacial maximum (LGM) and the Holocene climatic optimum (HCO) climatic regimes. We have ordained these proposed vegetation changes along rough physiognomic seral stages according to assumptions based on current botanical research. Finally, we have associated our hypothesized vegetation changes in South America and Africa with mammalian faunistic patterns, establishing a putative causal relationship between them. Results The extant mammal faunas of South America and Africa differ widely in taxonomical composition; the number of medium and large species they possess; behavioural and ecological characteristics related to herbivore herding, migration and predation; and biogeographical patterns. All such distinctions are mostly related to the open formation faunas, and have been completely established around the mid-Holocene. Considering that the mid-Holocene was a time of greater humidity than the late Pleistocene, vegetation cover in South America and Africa would have been dominated by forest or closed vegetation landscapes, at least for most of their lower altitude tropical regions. We attribute the loss of larger-sized mammal lineages in South America to the decrease of open vegetation area, and their survival in Africa to the existence of vast savannas in formerly steppic or desertic areas in subtropical Africa, north and south of the equator. Alternative explanations, mostly dealing with the disappearance of South American megamammals, are then reviewed and criticized. Main conclusions The reduction of open formation areas during the HCO in South America and Africa explains most of the present distinct faunistic patterns between the two continents. While South America would have lost most of its open formations within the 30� latitudinal belt, Africa would have kept large areas

A modern pollen–climate calibration set from northern Europe: developing and testing a tool for palaeoclimatological reconstructions

Abstract: Aims and location The potential of pollen records in quantitative climate reconstructions has been widely debated but seldom tested. Our aim is to develop a pollen–climate transfer function for northern Europe and test its performance and inference power by numerical cross-validation with modern climate data. Annual mean temperature (Tann) was assessed as the critical climatic variable because Tann has a distinct south–north gradient (5.5 to )4.7 � C) in the study region with a corresponding zonal vegetation gradient from the hemiboreal zone in the south to the northern boreal zone in the north. Methods We collected 137 pollen surface samples from small- to medium size lakes from southern Estonia to northern Finland. The transfer function for Tann was developed with weighted averaging partial least squares (WA-PLS) regression. All 102 terrestrial pollen and spore types were included in the calculation sum and all 137 surface samples and all 102 taxa were included in the transfer function. The performance of the WA-PLS transfer function was evaluated by leave-oneout cross-validation.

Floristic biogeography of the Hawaiian Islands: influences of area, environment and paleogeography

Abstract: Aim A detailed database of distributions and phylogenetic relationships of native Hawaiian flowering plant species is used to weigh the relative influences of environmental and historical factors on species numbers and endemism. Location The Hawaiian Islands are isolated in the North Pacific Ocean nearly 4000 km from the nearest continent and nearly as distant from the closest high islands, the Marquesas. The range of island sizes, environments, and geological histories within an extremely isolated archipelago make the Hawaiian Islands an ideal system in which to study spatial variation in species distributions and diversity. Because the biota is derived from colonization followed by extensive speciation, the role of evolution in shaping the regional species assemblage can be readily examined. Methods For whole islands and regions of each major habitat, species–area relationships were assessed. Residuals of species–area relationships were subjected to correlation analysis with measures of endemism, isolation, elevation and island age. Putative groups of descendents of each colonist from outside the Hawaiian Islands were considered phylogenetic lineages whose distributions were included in analyses. Results The species–area relationship is a prominent pattern among islands and among regions of each given habitat. Species number in each case correlates positively with number of endemics, number of lineages and number of species per lineage. For mesic and wet habitat regions, island age is more influential than area on species numbers, with older islands having more species, more single-island endemics, and higher species : lineage ratios than their areas alone would predict. Main conclusions Because species numbers and endemism are closely tied to speciation in the Hawaiian flora, particularly in the most species-rich phylogenetic lineages, individual islands’ histories are central in shaping their biota. The Maui Nui complex of islands (Maui, Moloka‘i, Lāna‘i and Kaho‘olawe), which formed a single large landmass during most of its history, is best viewed in terms of either the age or area of the complex as a whole, rather than the individual islands existing today.

Diversity and biogeography of vascular epiphytes in Western Amazonia, Yasuní, Ecuador

Abstract: Aim Although vascular epiphytes are important components of species richness and complexity of Neotropical forests, vascular epiphytes are under-represented in large scale biogeographical analyses. We studied the diversity, biogeography and floristic relationships of the epiphytic flora of the Yasuno´ region (Western Amazonia) in a Neotropical context, with special emphasis on the influence of the Andean flora on floristic composition and diversity of surrounding lowland forests. Location Western Amazonian lowland rainforest, Tiputini Biodiversity Station (0� 38¢ S7 6� 09¢ W, 230 m a.s.l., 650 ha), Yasuno´ National Park, Ecuador. Methods We compared the vascular epiphyte flora of Yasuno´ with 16 published Neotropical epiphyte inventories. Secondly, based on a floristic database with records of more than 70,000 specimens of vascular epiphytes from the Neotropics the elevational composition of eight selected inventories was analysed in detail. Results The vascular epiphyte flora of Yasuno´ is characterized by a very high species richness (313 spp.). A moderate portion of species is endemic to the Upper Napo region (c. 10%). However, this figure is much higher than previous analyses primarily based on woody species suggested. Geographical ranges of these species match with a proposed Pleistocene forest refuge. Compared with Northern and Central Amazonian sites, Western Amazonian epiphyte communities are characterized by a higher portion of montane and submontane species. Species richness of vascular epiphytes at the sites was correlated with the amount of rainfall, which is negatively correlated with the number of dry months. Main conclusion Recent and historic patterns of rainfall are the driving forces behind diversity and floristic composition of vascular epiphytes in Western Amazonia: high annual rainfall in combination with low seasonality provides suitable conditions to harbour high species richness. The proximity to the Andes, the most important centre of speciation for most Neotropical epiphytic taxa, in combination with the climatic setting has allowed a continuous supply of species richness to the region. At least for epiphytes, the borderline between the Andean and Amazonian flora is much hazier than previously thought. Moreover, the comparatively moist climate in Western Amazonia during the Pleistocene has probably led to fewer extinctions and/or more speciation than in more affected surrounding lowlands.

The Amazon River as a dispersal barrier to passerine birds: effects of river width, habitat and taxonomy

Abstract: Aim To evaluate the relative effectiveness of the lower and upper sections, respectively, of the Amazon River as a barrier to bird distribution, and to evaluate ecological and taxonomic factors affecting the efficacy of the river barrier. Location Amazon River of South America between its confluence with the Napo River in the west and its delta in the east. Methods Using published distribution maps for 448 species of passerine birds occurring along the Amazon River, we evaluated whether each was distributed along one bank only (river presumed to be a barrier) or both banks (no barrier) to test the predictions that the river was more effective as a dispersal barrier: (1) along the lower, wider portion of the river than the upper, narrower portion; (2) for species inhabiting forests than open country; (3) for species inhabiting forest understorey than forest canopy; (4) for species restricted to terra firme (never inundated upland forest) than those not restricted to terra firme and (5) for certain taxonomic groups. Results Our analyses demonstrated that the Amazon River was most effective as a dispersal barrier along its lower portion and for species restricted to forests and terra firme. However, the river was not significantly more of a barrier for species inhabiting forest understorey than forest canopy. The river was most significant as a barrier to dispersal for the antbirds (Thamnophilidae) and was less significant as a barrier to species belonging to several large families including woodcreepers (Dendrocolaptidae), ovenbirds (Furnariidae), flycatchers (Tyrannidae), cotingids (Cotingidae), tanagers (Thraupidae), seed-eating finches (Emberizidae) and blackbirds (Icteridae). Main conclusions The robust widths of Amazonian rivers are widely considered to represent impediments to dispersal and gene flow for many taxa of birds and other animals, and may have represented agents of vicariance in the diversification of species. Our study reaffirms the effectiveness of the lower Amazon River as a current barrier to bird dispersal for forest birds and provides new insights into the effects of habitat and taxonomy on the efficacy of the river barrier. Although supportive of several predictions of the river hypothesis of biological diversification, our study is limited in addressing the historical impact of river barriers as agents of vicariance in the process of diversification.

Regional biogeography of shallow reef fish and macro-invertebrate communities in the Galapagos archipelago

Abstract: Aim To delineate biogeographical patterns in Galapagos shallow-water reef fauna at regional scales. Location Galapagos Islands. Methods Fishes and macro-invertebrates were quantitatively censused using underwater visual techniques along more than 500 transects at defined depth strata across the Galapagos archipelago. Data were analysed using multivariate techniques to define regional patterns and identify species typical of different regions. Results Subtidal communities of fishes and macro-invertebrates on shallow reefs differed consistently in species composition across the Galapagos archipelago, with three major biogeographical groupings: (1) the ‘far-northern area’ containing the islands of Darwin and Wolf, (2) the ‘central/south-eastern area’, including the east coast of Isabela, and (3) the ‘western area’, encompassing Fernandina and western Isabela. In addition, the northern islands of Pinta, Marchena and Genovesa form a separate region in the central/south-eastern area, and Bahia Elizabeth and Canal Bolivar separate from other parts of the western area. The far-northern bioregion is characterized by high fish species richness overall, including a high proportion of species of Indo-Pacific origin. However, very few endemic fishes or species with distributions extending south from Ecuador (‘Peruvian’ species) are present, and the bioregion also possesses relatively low species richness of mobile macro-invertebrate taxa. By contrast, the ‘western’ bioregion possesses disproportionately high numbers of endemic fish taxa, high numbers of cool-temperate Peruvian fish species, and high invertebrate species richness, but very few species of Indo-Pacific origin. The Bahia Elizabeth/Canal Bolivar bioregion possesses more endemic species and fewer species with Peruvian affinities than coasts within the western bioregion. The northern bioregion of Pinta, Marchena and Genovesa represents an overlap zone with affinities to both the far-northern and south-eastern islands. The south-eastern bioregion includes species from a variety of different sources, particularly ‘Panamic’ species with distributions extending north to Central America. Main conclusions On the basis of congruent divisions for reef fish and macro-invertebrate communities, the Galapagos archipelago can be separated into three major biogeographical areas, two of which can be further subdivided into two regions. Each of these five bioregions possesses communities characterized by a distinctive mix of species derived from Indo-Pacific, Panamic, Peruvian and endemic source areas. The conservation significance of different regions is not reflected in counts of total species richness. The regions with the lowest overall fish species richness possess a temperate rather than tropical climate and highest levels of endemism.

Use of arboreal and terrestrial space by a small mammal community in a tropical rain forest in Borneo, Malaysia

Abstract: Aim: Small mammals were live-trapped in a primary rain forest to evaluate the relative distribution of species to each other and to microhabitat properties on the ground and in the canopy. Location: Kinabalu National Park in Borneo, Sabah, Malaysia. Methods: Seven trapping sessions were conducted along two grids with 31 trap points at distances of 20 m on the ground and in the lower canopy at an average height of 13.5 m. Results: Species diversity and abundance of small mammals proved to be high: 20 species of the families Muridae, Sciuridae, Tupaiidae, Hystricidae, Viverridae and Lorisidae were trapped, with murids being dominant in both habitat layers. The terrestrial community was significantly more diverse with 16 captured species (Shannon-Wiener's diversity index Hterr′ = 2.47), while 11 species were trapped in the canopy (Harb′ = 1.59). The Whitehead's rat, Maxomys whiteheadi, and the red spiny rat, Maxomys surifer, dominated the terrestrial community whereas the large pencil-tailed tree mouse, Chiropodomys major, was by far the most abundant species in the canopy. Other abundant species of the canopy community, the dark-tailed tree rat, Niviventer cremoriventer, and the lesser treeshrew, Tupaia minor, were also abundant on the ground, and there was no clear boundary between arboreal and terrestrial species occurrences. Main conclusions: As most species were not confined to specific microhabitats or habitat layers, species seemed to rely on resources not necessarily restricted to certain microhabitats or habitat layers, and separation of species probably resulted mainly from a species' concentrated activity in a preferred microhabitat rather than from principal adaptations to certain habitats. Ecological segregation was stronger in the more diverse terrestrial community, though microhabitat selection was generally not sufficient to explain the co-occurrences of species and the variability between local species assemblages. Constraints on small mammal foraging efficiency in the three-dimensional more complex canopy may be responsible for the similarity of microhabitat use of all common arboreal species. Community composition was characterized by mobile species with low persistence rates, resulting in a high degree of variability in local species assemblages with similar turnover rates in both habitats.

Large‐scale diversity patterns of vascular epiphytes in Neotropical montane rain forests

Abstract: Aim Epiphytes contribute up to 30% to the number of vascular plant species in certain global biodiversity hotspots, e.g. the Ecuadorian Andes. However, their large scale diversity patterns are still discussed on the base of results from a few, local epiphyte inventories. Consequently, explanatory models on epiphyte diversity concentrate on the impact of local climate on small scale epiphyte species richness. Our aim was to analyse large scale elevational patterns of epiphyte diversity integrating data from different geographic scales. Location Tropical America, with special emphasis on the Ecuadorian Andes. Methods Our study is based on two data sources. First, we analysed the elevational patterns of epiphyte diversity based on the Catalogue of the Vascular Plants of Ecuador and the Libro Rojo de las Plantas Endemicas del Ecuador. Secondly, the floristic turnover between the epiphyte inventories of seven montane and four lowland study sites in the Neotropics was analysed. Results The floristic turnover between Neotropical montane epiphyte floras is higher than the one between lowland epiphyte floras. Montane study sites located only a few kilometres apart from each other show considerable differences in their epiphyte species inventories. Irrespectively of their similar dispersal mode, the floristic turnover is much higher for orchids than for Pteridophyta. The Orchidaceae are the species richest group in all of the examined 11 Neotropical epiphyte floras. At the larger scale of the Ecuadorian Flora, c. 50% of the species in the elevational zone with maximum epiphyte diversity (between 1000 and 1500 m) are orchids. Elevational patterns of epiphyte diversity strongly reflect patterns of Orchidaceae. Main conclusions Our results support the observation of a 'mid-elevation bulge' of epiphyte diversity by Gentry and Dodson. It has been frequently shown that the high humidity in mid-elevations is suitable to maintan a high epiphyte species richness. Our findings show that in addition, large scale epiphyte diversity in montane rain forest is increased by the high floristic turnover at local and regional scale. Based on the importance of Orchidaceae for epiphyte diversity, we discuss that speciation processes corresponding to the highly diverse environment are a driving force for endemism, floristic heterogeneity and consequently for large scale epiphyte species richness in montane forests.

Distribution, endemism and threat status of freshwater fishes in the Western Ghats of India

Abstract: Aim To study (1) the large-scale distribution patterns of freshwater fishes in the Western Ghats of India; (2) the endemism and uniqueness of the fishes in various zones; and (3) the threat status of fishes by categorizing them under low risk (LR), vulnerable (VU), endangered (EN) and critically endangered (CR).

Topographic controls on the regional-scale biodiversity of the south-western USA

Abstract: Aim Topography is a fundamental geophysical observable that contains valuable information about the geodynamic, tectonic and climatic history of a region. Here, we extend the traditional uses of topographic analysis to evaluate the role played by topography in the distribution of regional-scale biodiversity in the south-western USA. An important aspect of our study is its ability to provide a way to quantify characteristics of the topographic fabric and to construct predictive models that can be used to test hypotheses that relate topography and biodiversity. Location South-western USA region of the North American Cordillera. Methods Our approach begins with a quantitative analysis of the topography and the construction of a predicted biodiversity map based on measurable topographic quantities: organization, roughness, slope aspect, grain orientation and mean elevation. We then make a quantitative comparison between the predicted and observed biodiversity, based on the assumption that land-cover diversity is a plausible measure of regional-scale biodiversity. Land-cover information used for this study was collected as part of the U.S.G.S. global land cover characteristics (GLCC) project and is derived from satellite (AVHRR) imagery. Results To a first order, the predicted regional-scale biodiversity based on our topographic model shows a good correlation with the observed biodiversity (as estimated from the land-cover diversity). Our model overestimates the biodiversity in many parts of the Colorado Plateau, Rio Grande Rift, and the low desert regions of the Southern Basin and Range, suggesting that in these provinces a biodiversity estimate based solely on topography is an oversimplification. However, much of the Madrean Archipelago and Sierra Madre provinces, which are centres of high biodiversity in this region, show excellent agreement between the observed and predicted biodiversity. Main conclusions While we acknowledge that many other factors in addition to topography have an important influence on biodiversity (particularly on a local scale), we conclude that topography plays a primary role in the regional to continental-scale biodiversity, particularly in regions characterized by insular mountain fabrics.

The distribution of South American galaxiid fishes: the role of biological traits and post-glacial history

Abstract: Aim The aim of this work is to update the distribution data of Galaxiidae in South America, relating extant distribution to physiological and reproductive characteristics of the species, latitude, temperature, and post-glacial opportunities for colonization. Location Lakes and rivers of Patagonia. Methods We compared, and eventually reconsidered, general data about distribution based on the original literature about capture sites, incorporating several published and unpublished data to the analysis of the biological traits and distribution of Galaxiidae. Results The more consistent issue in the comprehension of galaxiid biogeography in South America is the ability to establish landlocked populations. Different founding events in landlocked populations of Galaxias maculatus suggest the possible existence of older and younger landlocked populations. This difference in the time since the establishment of lacustrine populations could have been expressed in their ability for colonization of post-glacial areas. Galaxias maculatus, Aplochiton and Brachygalaxias are more clearly excluded from the post-glacial area than G. platei. For all the species we could note a more abundant record of lake populations at the area of glacial refuges. It could be noted that the most successful species, Galaxias platei, is a specialized deep bottom dweller. Deep bottom dwelling helps to endure winter constraints and it appears to be an alternative to the colonization of the littoral and limnetic zones of post-glacial lakes, the prefered habitat of the other Patagonian fish species. Main conclusions At the end of this process of post-glacial colonization, in the beginning of twentieth century, man introduced several salmonid species in Patagonia. In addition, antropogenic actions had its more recent consequences in global warming. Nowadays we were able to observe new localities for Brazilian fishes into the Austral Subregion and expect some changes in the distribution of Galaxiidae. Northern limits for all species and southern limits for landlocked G. maculatus, Brachigalaxias bullocki and Aplochiton zebra, could be displaced southward. Probably, the species less affected by the changes will be G. platei. These predictions could be accurately formulated using the model of B.J. Shuter & J.R. Post (1990) Transactions of the American Fisheries Society119, 314–336, when biological database on these species are completed.

Biological diversification in a complex region: a spatial analysis of faunistic diversity and biogeography of the Andes of Colombia

Abstract: Aim Understanding large-scale patterns of beta diversity and endemism is essential for ecoregional conservation planning We present a study of spatial patterns of faunal diversification and biogeographical relationships in the Andean region of Colombia This region has a great geomorphological complexity, as it is formed by several mountain ranges with different geologic origins We hypothesize that this complexity results in a high turnover in species composition among subregions Location The Andean region of Colombia, including the Santa Marta and Macarena mountain ranges Methods The region was divided into subregions, represented by the eastern and western slopes of each of the three Andean cordilleras, the Cauca and Magdalena valley bottoms, and the peripheral mountain ranges of Perija, Macarena and Sierra Nevada de Santa Marta Species lists for five animal taxa (rodents, bats, birds, frogs and butterflies) were compiled for each subregion and similarities in species composition were determined by cluster analysis To explore biogeographical relationships, species were classified into one of four distributional categories: endemic, tropical Andean, Andean-Central American and wide continental distribution Results The highest species richness in the region was found in the Pacific and eastern versants of the Andes, and the lowest in the Cauca and Magdalena valley bottoms Inter-Andean slopes were intermediate in species richness However, when species richness was calculated per unit area, the most diverse regions were the Santa Marta and Macarena ranges, the Cauca Valley watershed and the Pacific slope Although each taxonomic group had a different branching pattern, dendrograms indicated five common subregional clusterings: (1) Perija-Sierra Nevada, (2) the Pacific slope, (3) the eastern Andean slope, (4) the Cauca and Magdalena valley bottoms, and (5) the inter-Andean slopes Clustering patterns of inter-Andean slopes varied among taxa In birds, bats and rodents, grouping was by opposite slopes of the same valley, whereas frogs were grouped by mountain ranges and butterflies by valleys and their respective slopes Seventy-five per cent of species in all taxa were found in less than five subregions The fauna of the Magdalena and Cauca valley bottoms was composed mostly of lowland species with wide geographical distributions, whereas the cordilleran fauna was mostly restricted to the tropical Andes Main conclusions The western and eastern versants of the Andes have the highest species richness, but are also the largest subregions On a per unit area basis, the peripheral ranges (Santa Marta and Macarena) are the richest, followed by the western portion of the Andes (the Cauca Valley watershed and the Pacific versant) Clustering patterns in dendrograms suggest two major patterns of differentiation of the Andean fauna: one elevational (lowlands vs highlands) and one horizontal (among ranges and/or slopes) Biogeographical affinities of the inter-Andean valley bottoms are with the lowland faunas of tropical America In contrast, Andean faunas diversified locally, resulting in the evolution of a large number of endemic species, particularly among the less vagile taxa Three different main branches of Andean fauna can be recognized, one confined to the Pacific, another to the eastern (Amazonian-Llanos) versant of the Andes, and the third one composed by the inter-Andean slopes of the Cauca and Magdalena valleys The identification of five main biogeographical units in the Andean region of Colombia has important implications for the conservation of the regional biota Conservation initiatives that seek to preserve representative samples of the regional biodiversity should take into account the patterns of diversification described here, and the evolutionary processes that gave rise to these patterns Resumen El conocimiento de los patrones de diversidad beta y endemismo a gran escala espacial es esencial para la planificacion ecorregional En este articulo se presenta un estudio de los patrones espaciales de diversificacion de la fauna y de sus relaciones biogeograficas en la region andina de Colombia y serranias perifericas Esta region es de una gran complejidad geomorfologica, ya que esta conformada por varias cordilleras y serranias de diferentes origenes geologicos Nuestra hipotesis es que esta complejidad genera una alta composicion de especies entre las subregiones La region andina de Colombia (incluyendo la Sierra Nevada de Santa Marta y la serrania de La Macarena), fue dividida en subregiones, constituidas por las vertientes oriental y occidental de cada una de las tres cordilleras, las planicies de los valles del Cauca y Magdalena, las serranias de Perija y La Macarena y la Sierra Nevada de Santa Marta Para cada una de estas subregiones compilamos la lista de especies para cinco grupos taxonomicos: roedores, murcielagos, aves, ranas y mariposas En cada grupo, determinamos la similitud en composicion de especies por medio de analisis de agrupamiento (cluster analysis) Con el objectivo de explorar las relaciones biogeograficas, clasificamos las especies en cuatro categorias de distribucion geografica: endemica, Andes tropicales, andina-centroamericana y distribucion amplia La riqueza de especies mas alta se encontro en las vertientes Pacifica y oriental de los Andes, y la mas baja en los valles del Cauca y Magdalena Las vertientes interandinas presentaron una riqueza intermedia Sin embargo, por unidad de area las subregiones mas diversas fueron La Macarena y Santa Marta, el valle del Cauca y sus vertientes, y la vertiente del Pacifico Aunque el analisis arrojo diferentes patrones de agrupamiento para cada grupo taxonomico, en los dendrogramas se destacan cinco ramas: (1) Perija-Sierra Nevada, (2) la vertiente Pacifico, (3) la vertiente oriental de los Andes, (4) las planicies de los dos valles, y (5) las vertientes interandinas Los patrones de agrupamiento de las vertientes interandinas fueron variables En aves, murcielagos y roedores, las vertientes opuestas de cada valle resultaron muy cercanas en composicion de especies; en cambio, en las ranas se agruparon primero las dos vertientes de cada cordillera; en mariposas, cada valle se agrupo con sus respectivas vertientes La fauna de las planicies del Cauca y Magdalena esta compuesta principalmente por especies de tierras bajas y amplias distribuciones geograficas La fauna de las cordilleras, en cambio, esta compuesta principalmente por especies restringidas a los Andes tropicales Los patrones de agrupamiento en los dendrogramas sugieren dos patrones principales de diferenciacion de la fauna andina: un patron vertical (tierras bajas versus tierras altas) y otro horizontal (entre cordilleras y entre vertientes) Las planicies interandinas del Cauca y Magdalena tienen afinidad biogeografica con la fauna de tierras bajas del tropico americano, mientras que las faunas andinas muestran una diversificacion local, lo cual ha resultado en la evolucion de un gran numero de especies endemicas, particularmente en los taxones menos moviles Se pueden reconocer tres grandes ramas en la fauna propiamente andina: una confinada a la vertiente del Pacifico, otra a la vertiente oriental de los Andes, y una tercera compuesta por las vertientes interandinas de los valles del Cauca y Magdalena La identificacion de cinco grandes unidades biogeograficas en la region andina de Colombia tiene importantes implicaciones para la conservacion de la biota regional Las iniciativas de conservacion que buscan preservar muestras representativas de la diversidad regional deben tener en cuenta no solo los patrones de diversidad descritos en este articulo, sino tambien los procesos evolutivos que originaron estos patrones

Landscape analysis of Aboriginal fire management in Central Arnhem Land, north Australia

Abstract: Aim To describe the spatial and temporal pattern of landscape burning with increasing distance from Aboriginal settlements. Location Central Arnhem Land, a stronghold of traditional Aboriginal culture, in the Australian monsoon tropics. Methods Geographical information system and global positioning system technologies were used to measure spatial and temporal changes in fire patterns over a one decade period in a 100 x 80 km area that included a cluster of Aboriginal settlements and a large uninhabited area. The major vegetation types were mapped and fire activity was assessed by systematic visual interpretation of sequences of cloud-free Landsat satellite images acquired in the first (May to July) and second (August to October) halves of the 7-month dry season. Fire activity in the middle and end of one dry season near an Aboriginal settlement was mapped along a 90-km field traverse. Canopy scorch height was determined by sampling burnt areas beside vehicle tracks. Results Satellite fire mapping was 90% accurate if the satellite pass followed shortly after a fire event, but the reliability decayed dramatically with increasing time since the fire. Thus the satellite mapping provided a conservative index of fire activity that was unable to provide reliable estimates of the spatial extent of individual fires. There was little landscape fire activity in the first half of the dry season, that was mostly restricted to areas immediately surrounding Aboriginal settlements, with burning of both inhabited and uninhabited landscapes concentrated in the second half of the dry season. The mean decadal fire indices for the three dominant vegetation types in the study area were three in the plateau savanna, two in the sandstone and five in the wet savanna. The spatial and temporal variability of Aboriginal burning apparent in the satellite analyses were verified by field traverse surrounding a single settlement. Fires set by Aborigines had low scorch height of tree crowns reflecting low intensity, despite generally occurring late in the dry season. Conclusions Our findings support the idea that Aboriginal burning created a fine-scale mosaic of burnt and unburnt areas but do not support the widely held view that Aboriginal burning was focused primarily in the first half of the dry season (before July). The frequency and scale of burning by Aborigines appears to be lower compared with European fire regimes characterized by fires of annual or biennial frequencies that burn large areas. The European fire regime appears to have triggered a positive feedback cycle between fire frequency and flammable grass fuels. The widely advocated management objective of burning in the first half of the dry season burning provides one of the few options to control fires once heavy grass fuel loads have become established, however we suggest it is erroneous to characterize such a regime as reflecting traditional Aboriginal burning practices. The preservation of Aboriginal fire management regimes should be a high management priority given the difficulty in breaking the grass-fire cycle once it has been initiated.

Pollen-based reconstructions of biome distributions for Australia, Southeast Asia and the Pacific (SEAPAC region) at 0, 6000 and 18,000 14C yr BP

Abstract: Aim: This paper documents reconstructions of the vegetation patterns in Australia, Southeast Asia and the Pacific (SEAPAC region) in the mid- Holocene and at the last glacial maximum (LGM). Methods: Vegetation patterns were reconstructed from pollen data using an objective biomization scheme based on plant functional types. The biomization scheme was first tested using 535 modern pollen samples from 377 sites, and then applied unchanged to fossil pollen samples dating to 6000 ± 500 or 18,000 ± 1000 14C yr BP. Results: 1. Tests using surface pollen sample sites showed that the biomization scheme is capable of reproducing the modern broad-scale patterns of vegetation distribution. The north-south gradient in temperature, reflected in transitions from cool evergreen needleleaf forest in the extreme south through temperate rain forest or wet sclerophyll forest (WSFW) and into tropical forests, is well reconstructed. The transitions from xerophytic through sclerophyll woodlands and open forests to closed-canopy forests, which reflect the gradient in plant available moisture from the continental interior towards the coast, are reconstructed with less geographical precision but nevertheless the broad-scale pattern emerges. 2. Differences between the modern and mid-Holocene vegetation patterns in mainland Australia are comparatively small and reflect changes in moisture availability rather than temperature. In south-eastern Australia some sites show a shift towards more moisture-stressed vegetation in the mid-Holocene with xerophytic woods/scrub and temperate sclerophyll woodland and shrubland at sites characterized today by WSFW or warm- temperate rain forest (WTRF). However, sites in the Snowy Mountains, on the Southern Tablelands and east of the Great Dividing Range have more moisture- demanding vegetation in the mid-Holocene than today. South-western Australia was slightly drier than today. The single site in north-western Australia also shows conditions drier than today in the mid-Holocene. Changes in the tropics are also comparatively small, but the presence of WTRF and tropical deciduous broadleaf forest and woodland in the mid-Holocene, in sites occupied today by cool- temperate rain forest, indicate warmer conditions. 3. Expansion of xerophytic vegetation in the south and tropical deciduous broadleaf forest and woodland in the north indicate drier conditions across mainland Australia at the LGM. None of these changes are informative about the degree of cooling. However the evidence from the tropics, showing lowering of the treeline and forest belts, indicates that conditions were between 1 and 9°C (depending on elevation) colder. The encroachment of tropical deciduous broadleaf forest and woodland into lowland evergreen broadleaf forest implies greater aridity. Main conclusions: This study provides the first continental-scale reconstruction of mid-Holocene and LGM vegetation patterns from Australia, Southeast Asia and the Pacific (SEAPAC region) using an objective biomization scheme. These data will provide a benchmark for evaluation of palaeoclimate simulations within the framework of the Palaeoclimate Modelling Intercomparison Project.

Areas of endemism of vascular plants in the Eastern Alps in relation to Pleistocene glaciation

Abstract: Aim The central aim of this paper is to localize areas of endemism of the vascular plant flora in the Eastern Alps. Moreover, causes for location and limits of the areas of endemism are assessed. Location The study area includes the Eastern European Alps and adjacent lowland areas. Methods Analyses were based on 288 vascular plant taxa endemic to the study region, scored in 115 operational geographic units, which were predominantly mountain ranges. For each of these units, number of endemic taxa, range down weighted endemism, environmental variables, elevation of the last glacial maximum ice sheet, and elevation of the snowline during the last glacial maximum, were estimated. To evaluate the relationships between the geographical areas, the presence/absence matrix of endemics in areas was used to calculate phenograms and cladograms. Values from the range down weighted endemism superimposed on phenograms and cladograms were used to indicate areas of endemism. Linear regressions between richness in endemism and environmental as well as palaeo-environmental variables were used to infer causes of the observed patterns of endemism. Results The endemic taxa of the Eastern Alps show a very uneven distribution, the majority being confined to one or few mountain ranges. High levels of endemism are found in the southern, southeastern, easternmost, and northeastern Eastern Alps. Trees obtained from phenetic and cladistic methods were largely congruent. Seven areas of endemism were clearly delimited, all in regions that are regarded as glacial refugia. Two of them are located on siliceous, four on calcareous bedrock. Not all endemic taxa, however, grow in the areas of endemism. Therefore, enlargement areas are suggested. Linear regression showed a significant correlation between high endemism and low glacial ice cover, but not with other environmental variables. Main conclusions Vicariance resulting from Pleistocene glaciations is the most important factor causing distributional patterns of endemic plants and the formation of areas of endemism. Mapping values of weighted endemism onto trees facilitated delimitation and definition of the areas of endemism. Because of their strong dependence on siliceous bedrock and the resulting disjunct distributional patterns of some taxa in the Southern Alps, the inclusion of a few operational geographic units with high levels of endemism was not supported by the analyses. Despite post-vicariant dispersal that causes problems in analyses of areas of endemism, the circumscription of areas of endemism in the Eastern Alps was possible to a satisfactory extent. Other methodological approaches, however, which include concepts of post-vicariant dispersal and that allow overlap of areas of endemism would be highly useful.

Relationships between land snail assemblage patterns and soil properties in temperate‐humid forest ecosystems

Abstract: Aim The objectives were to (1) analyse the combined effects of soil pH, Ca content and soil moisture on total density and species richness of land snails in forest ecosystems, (2) explore relationships between the quantitative composition of snail assemblages and habitat characteristics, (3) investigate the relationships between soil pH and density of some of the most frequent species, and (4) compare the data with those from studies conducted in other temperate-humid regions of Europe. Location Study sites were selected from 15 landscape types including different lithologies within the area of Baden-Wurttemberg (35,000 km2), SW Germany. Methods Snails were recorded quantitatively from 83 study sites, with four plots representing a total of 0.25 m2 per site. Topsoil samples from each site were analysed for pH, exchangeable Ca, and Ca content of carbonates. Three categories of soil moisture (dry, intermediate and wet) were established and defined according to the (climatic) water balance. Numbers of individuals and species were brought in relation to soil moisture and soil pH. Cluster analyses were conducted to identify groups of quantitatively similar snail species assemblages. Results Topsoil pH (2.7–7.5) and soil Ca contents were closely correlated. On dry soils, total snail density and species richness are generally low and do not change with pH, but clearly increase with increasing pH on intermediate moisture soils and on wet soils. On the latter, numbers of individuals and species are generally much higher compared with intermediate moisture sites at the same value of soil pH. Changes of density in relation to soil pH vary between species. Depending on the species, density increases only in the lower or only in the higher range of pH, is not related to pH, or decreases with increasing pH. Furthermore, these patterns vary within the same species depending on the region. This became evident from comparisons with other studies, particularly between sites in SW Germany and southern Scandinavia. From cluster analyses, subgroups of snail assemblages of high quantitative similarity were identified. Group formation is explained by soil pH to some extent, and one subgroup showed a connection with coniferous woodland sites on acidic soils. No further environmental factors available from our data could explain the clustering of snail assemblages more detailed. Main conclusions Soil moisture is the strongest determinant of snail density and species richness at undisturbed woodland sites, but effects of soil moisture and soil pH on these patterns are closely interrelated on intermediate moisture soils and wet soils. However, the quantitative species composition of the land snail assemblages is related to soil properties to a lower degree than snail density and species richness, and other habitat characteristics such as vegetation or litter quality, can be important for species dominance in addition.

Breeding bird species richness in Taiwan: distribution on gradients of elevation, primary productivity and urbanization

Abstract: Aim To examine the richness of breeding bird species in relation to elevation, primary productivity and urbanization. Location The island of Taiwan (120� –122� E, 22� –25� N). Methods We arranged bird species richness (BSR) data from 288 bird censuses undertaken in Taiwan into a 2 · 2 km quadrat system and calculated average values of elevation, primary productivity [surrogated by normalized difference vegetation index (NDVI)], and urbanization (surrogated by road density and percentage of built area) for each 2 · 2 km quadrat. Results Bird species richness showed a hump-shaped relationship with elevation. It increased with elevation from sea level (10–64 species per 2 · 2 km quadrat), peaked around 2000 m (43–76 species), and then decreased with elevation towards its minimum at the highest elevation. Road density and percentage of built area decreased with elevation, and NDVI showed a humpshaped relationship with elevation and inverse relationships with road density and percentage of built area. BSR increased with NDVI and decreased with road density and percentage of built area. Linear and cubic terms of elevation together explained 31.3% of the variance in BSR, and road density explained additional 3.4%. The explanatory power of NDVI on BSR was insignificant after the effects of elevation and road density had been justified. Main conclusions We argue that urbanization plays an important role in the BSR of Taiwan. Urbanization might indirectly decrease BSR through decreasing primary productivity and therefore change the hypothetical inverse relationship between BSR and elevation into a hump-shaped relationship. We also propose a time hypothesis that the biotic communities in the mid-elevation zone of Taiwan had relatively longer periods of existence during the Pleistocene glacial cycles, which might be one underlying process of the observed hump-shaped relationship between species diversity and elevation.

Biogeography of lemurs in the humid forests of Madagascar: the role of elevational distribution and rivers

Abstract: Aim Elucidation of the influence of river systems on the distribution of lemurs. Location Humid forests of Madagascar. Methods We review the elevational distribution of Malagasy lemurs to address hypotheses of the role of rivers as physical barriers. Main conclusions Rivers are barriers for some species. However, several lemurs have altitudinal ranges that are higher than the headwaters of a given river, and in such cases these species are able to disperse across the source.