Modelling spatial patterns of biodiversity for conservation prioritization in North-eastern Mexico

Relationships between spatial patterns of bird and mammal species richness in north‐eastern Mexico were analysed in relation to the location of three biosphere reserves (El Abra‐Tanchipa, El Cielo, and Sierra Gorda) and 13 priority areas recently identified for conservation. Ecological niches were modelled and potential distributions delimited for 285 bird and 114 mammal species using a genetic algorithm based on locality information from museum specimens and 15 selected environmental attributes. Potential distributions were transformed into hypothesized current distributions based on species–habitat associations as reflected in a recent land‐use map. Although species richness was lower when distributions were reduced from potential to current, spatial patterns of potential and current richness were similar. Heuristic, complementarity‐based prioritization procedures were used to identify combinations of areas and sites with maximal species representation: the biosphere reserves included 79% of birds and 74% of mammal species; eight priority areas provided an additional 11% of birds and 13% of mammals; the remaining 10% of birds and 13% of mammals were concentrated in new sites across the study area.     

Using patterns of distribution and diversity of Kenyan birds to select and prioritize areas for conservation

Patterns and environmental correlates of species distributions and richness are identified for Kenyan birds at a quarter degree-square scale. This information is used together with iterative complementarity analyses, which employ species richness, taxonomic dispersion and range-restrictedness, to identify priority areas for possible conservation attention. Bird species apparently not conserved by existing protected areas in Kenya are identified. Six avifaunal zones (and one transitional zone) are distinguished based on distributions of suites of bird species. Variation in biotope diversity (the number of forest and aquatic systems) accounts for 79% of the observed variation in Kenyan bird species richness. Although both rainfall and altitudinal range are significantly correlated with species richness, they only explain an additional 3% of the observed variation. The priority areas identified are situated mainly within highlands and coastal lowlands. Although few priority areas are identified in northern Kenya, this region also constitutes a priority, as it contains a suite of xeric species with habitats that are not represented elsewhere in Kenya. The papyrus yellow warbler, Chloropeta gracilirostris, William's bush lark, Mirafra williamsi, white-winged dove, Streptopelia reichenowi, and Jubaland weaver, Ploceus dichrocephalus, are identified as endemics or near-endemics that are probably not adequately conserved in Kenya at present.     

Using Self-Organizing Maps to Explore Patterns in Species Richness and Protection

The combination of species distributions with abiotic and landscape variables using Geographic Information Systems can prioritize areas for biodiversity protection by identifying areas of high richness, although the number of variables and complexity of the relationships between them can prove difficult for traditional statistical methods. The use of these methods, which commonly assume linearity and low correlation between independent variables, can obscure even strong relationships and patterns. Self-Organizing Maps (SOM) is a heuristic statistical tool based on machine learning methods that can be used to explore patterns in large, complex datasets for linear and nonlinear patterns. Here we use SOM to visualize broad patterns in species richness by taxonomic group (birds, mammals, reptiles, and amphibians) and 78 habitat, landscape and environmental variables using data from the Gap analysis project for West Virginia, USA. Soil and habitat variables demonstrated clear relationships with species richness; areas with high species richness occurred in areas with high soil richness. Landscape metrics were less important, although habitat diversity and evenness indices were positively related to species richness in some taxonomic groups. Current coverage of protected areas (e.g., National Forests and state parks) appeared to be insufficient to cover most of the areas of high species richness, especially for reptiles; many of the polygons with the highest richness were not covered by these areas. The identification of polygons with high richness and low protection can be used to focus conservation efforts in those areas.     

Estimating the Spatial and Temporal Distribution of Species Richness within Sequoia and Kings Canyon National Parks

Evidence for significant losses of species richness or biodiversity, even within protected natural areas, is mounting. Managers are increasingly being asked to monitor biodiversity, yet estimating biodiversity is often prohibitively expensive. As a cost-effective option, we estimated the spatial and temporal distribution of species richness for four taxonomic groups (birds, mammals, herpetofauna (reptiles and amphibians), and plants) within Sequoia and Kings Canyon National Parks using only existing biological studies undertaken within the Parks and the Parks'' long-term wildlife observation database. We used a rarefaction approach to model species richness for the four taxonomic groups and analyzed those groups by habitat type, elevation zone, and time period. We then mapped the spatial distributions of species richness values for the four taxonomic groups, as well as total species richness, for the Parks. We also estimated changes in species richness for birds, mammals, and herpetofauna since 1980. The modeled patterns of species richness either peaked at mid elevations (mammals, plants, and total species richness) or declined consistently with increasing elevation (herpetofauna and birds). Plants reached maximum species richness values at much higher elevations than did vertebrate taxa, and non-flying mammals reached maximum species richness values at higher elevations than did birds. Alpine plant communities, including sagebrush, had higher species richness values than did subalpine plant communities located below them in elevation. These results are supported by other papers published in the scientific literature. Perhaps reflecting climate change: birds and herpetofauna displayed declines in species richness since 1980 at low and middle elevations and mammals displayed declines in species richness since 1980 at all elevations.     

The pace of past climate change vs. potential bird distributions and land use in the United States

Climate change may drastically alter patterns of species distributions and richness, but predicting future species patterns in occurrence is challenging. Significant shifts in distributions have already been observed, and understanding these recent changes can improve our understanding of potential future changes. We assessed how past climate change affected potential breeding distributions for landbird species in the conterminous United States. We quantified the bioclimatic velocity of potential breeding distributions, that is, the pace and direction of change for each species’ suitable climate space over the past 60 years. We found that potential breeding distributions for landbirds have shifted substantially with an average velocity of 1.27 km yr^?1, about double the pace of prior distribution shift estimates across terrestrial systems globally (0.61 km yr^?1). The direction of shifts was not uniform. The majority of species’ distributions shifted west, northwest, and north. Multidirectional shifts suggest that changes in climate conditions beyond mean temperature were influencing distributional changes. Indeed, precipitation variables that were proxies for extreme conditions were important variables across all models. There were winners and losers in terms of the area of distributions; many species experienced contractions along west and east distribution edges, and expansions along northern distribution edges. Changes were also reflected in the potential species richness, with some regions potentially gaining species (Midwest, East) and other areas potentially losing species (Southwest). However, the degree to which changes in potential breeding distributions are manifested in actual species richness depends on landcover. Areas that have become increasingly suitable for breeding birds due to changing climate are often those attractive to humans for agriculture and development. This suggests that many areas might have supported more breeding bird species had the landscape not been altered. Our study illustrates that climate change is not only a future threat, but something birds are already experiencing.     

Primary productivity and species richness: relationships among functional guilds, residency groups and vagility classes at multiple spatial scales

One of the major determinants of species richness is the amount of energy available, often measured as primary productivity. Heterogeneity of environmental variables has also been found to influence species richness. Predicting species distributions across landscapes and identifying areas that have high species richness, or vulnerable groups of species, is useful for land management. Remotely sensed data may help identify such areas, with the Normalized Difference Vegetation Index (NDVI) providing an estimate of primary productivity. We examined the relationship between maximum productivity (NDVI), heterogeneity of productivity, and species richness of birds and butterflies at multiple spatial scales. We also explored relationships between productivity, functional guilds and residency groups of birds, and vagility classes of butterflies. Positive linear relationships between maximum NDVI and number of functional guilds of birds were found at two spatial scales. We also found positive linear relationships between maximum NDVI and species richness of neotropical migrant birds at two scales. Heterogeneity of NDVI, by contrast, was negatively associated with number of functional guilds of birds and species richness of resident birds. Maximum NDVI was associated with species richness of all butterflies and of the most vagile butterflies. No association was found between heterogeneity of NDVI and species richness of butterflies. In the Great Basin, where high greenness and availability of water correspond to areas of high species richness and maximum NDVI, our results suggest that NDVI can provide a reliable basis for stratifying surveys of biodiversity, by highlighting areas of potentially high biodiversity across large areas. Measures of heterogeneity of NDVI appear to be less useful in explaining species richness.     

Global patterns of geographic range size in birds

Large-scale patterns of spatial variation in species geographic range size are central to many fundamental questions in macroecology and conservation biology. However, the global nature of these patterns has remained contentious, since previous studies have been geographically restricted and/or based on small taxonomic groups. Here, using a database on the breeding distributions of birds, we report the first (to our knowledge) global maps of variation in species range sizes for an entire taxonomic class. We show that range area does not follow a simple latitudinal pattern. Instead, the smallest range areas are attained on islands, in mountainous areas, and largely in the southern hemisphere. In contrast, bird species richness peaks around the equator, and towards higher latitudes. Despite these profoundly different latitudinal patterns, spatially explicit models reveal a weak tendency for areas with high species richness to house species with significantly smaller median range area. Taken together, these results show that for birds many spatial patterns in range size described in geographically restricted analyses do not reflect global rules. It remains to be discovered whether global patterns in geographic range size are best interpreted in terms of geographical variation in species assemblage packing, or in the rates of speciation, extinction, and dispersal that ultimately underlie biodiversity.     

On the use by birds of intertidal areas of the Tagus estuary: implications for management

The use of intertidal areas of the Tagus estuary by birds was re-analysed, based on data from 1990 to 1993, to describe (i) the temporal and spatial patterns of use of intertidal areas, (ii) the diet of birds during winter and (iii) the habitat selection patterns of feeding birds, during winter. The most common birds in the estuary were gulls, waders and ducks. Highest densities were recorded for most species in autumn and winter. The spatial distribution of birds in the intertidal areas of the estuary did not vary significantly across seasons, although broader distributions occurred when bird populations were present in high numbers. In autumn, use of intertidal areas was highly variable. Specific areas in the estuary were identified as holding important densities of birds, or having a high species richness. The most abundant species selected feeding areas according to sediment type, although the presence of channels, saltmarsh or humans also influenced the distribution of birds. The invertebrates Carcinus maenas, Hediste diversicolor and Scrobicularia plana were identified as key prey species. Plant material was important as food for ducks. Implications of these findings for the management of the estuary are discussed.     

Ecological versus phylogenetic determinants of helminth parasite community richness

Summary We examine patterns of community richness among intestinal parasitic helminth communities in fishes, herptiles, birds and mammals with respect to the comparative number of component species in a host population. We show that terrestrial hosts have, on average, fewer component species than aquatic hosts. We also show that the mean number of component species in aquatic hosts increases from fishes through herptiles to birds before declining slightly in mammals. For terrestrial hosts, the mean number of component species increases from herptiles, through birds, reaching a maximum in mammals. We conclude that: (i) habitat of the host is more important in determining community richness than is host phylogeny; (ii) the phenomenon of host capture may be largely responsible for increased species richness in some host groups; (iii) aquatic birds harbour the richest intestinal helminth communities; and (iv) as we interpret them, our data refute the time hypothesis, which would predict that fishes as the oldest lifestyle should have the richest helminth communities.Order of authorship determined by random draw and does not imply seniority.     

Contrasting patterns of elevational zonation for birds and mammals in the Andes of southeastern Peru

Abstract. To determine the generality of avian diversity patterns, we investigated patterns of elevational zonation shown by birds and mammals along the eastern slope of the Andes Mountains in southeastern Peru. The strong environmental gradient sampled, entirely within Peru's Manu National Park and Biosphere Reserve, supports highly diverse faunas. Elevational distributions of 901 bird species, 129 bat species, and twenty-eight species of native mice exhibit contrasting patterns in species richness, species composition, and species turnover. Birds and bats showed smooth declines of species richness with elevation, whereas the richness of mouse assemblages was unrelated to elevation. For all three groups, the greatest differences were between lowland and highland faunas, although cutoff points for this contrast varied among groups (≈ 500 m for birds, 750 m for bats, and 1000 m for mice). Differences in composition also separated bird and bat faunas on either side of c. 1400 m (the boundary between montance forest and cloud forest); for mice, this faunal transition may take place nearer to 2000 m. Bird and bat faunas lacked the more discrete zonations suggested for mouse assemblages, as indicated by elevational range profiles and nested subset analyses. Distinct highland assemblages are apparent in two-dimensional histograms of range limits of birds and mice, but not for bats. Highland bat species occupy broader elevational ranges than lowland bat species, but for both birds and mice, species at intermediate elevations had the broadest amplitudes. Finally, clumping of range maxima and minima along the gradient identified zones of pronounced species turnover in each group, but these were generally not strongly associated with the locations of ecotones. Differences in zonation of these groups appear to reflect their different biological attributes and phylogenetic histories. Such differences obviously complicate discussions of ‘general’ diversity patterns, and limit the usefulness of birds to forecast or predict diversity patterns in other more poorly known groups—other groups may show elevated diversity and endemism in areas where avian diversity patterns appear unremarkable. The pronounced contrasts between bats and mice, and the generally intermediate character of avian patterns, suggest that future analyses might profitably partition birds into finer, more homogeneous groups of historically and/or ecologically similar species. Group differences in zonation may ultimately prove explicable with information on both species-abundance patterns and resource distributions.     

Using habitat models to map diversity: pan-African species richness of ticks (Acari: Ixodida)

Aims To show how logistic regression models for individual species can be used to produce improved estimates of species richness at a continental scale; to present these data for African ticks (Acari: Ixodida); and to address the question of whether there is a latitudinal gradient in tick species richness. Location Africa. Methods A database of 34,060 collection records for African ticks is used to produce a pan‐African map of known tick species richness at 0.25 × 0.25‐degree resolution. The likely distributions of seventy‐three species are then estimated from environmental factors using logistic regression, and localities where there is a suitably high probability of occurrence for a given species are added to the original data for that species. These augmented data are combined to produce a map of the predicted pan‐African distribution of tick species richness. The relationship of species richness to latitude is considered along a transect placed across some of the more extensively collected areas. Results Maps of known and predicted pan‐African tick species richness are presented, and deficiencies in the available data are highlighted. Correlations using both known and predicted estimates of tick species richness suggest that ticks follow similar species richness patterns to those described for African mammals and birds, with a latitudinal gradient and highest species richness in east equatorial Africa. Tick species ranges are log‐normally distributed. Main conclusions Carefully constructed probability surfaces offer a more powerful approach to mapping species ranges than simple presence‐absence maps. Such models are a useful extension to current biogeographical methods and have a wide range of potential applications in ecology, epidemiology and conservation. Tick species richness at a continental scale follows similar trends to those reported for mammals and birds.     

Biogeographic patterns of the East African coastal forest vertebrate fauna

The archipelago-like coastal forest of East Africa is one of the highest priority ecosystems for biodiversity conservation worldwide. Here we investigate patterns of species richness and biogeographic distribution among birds, mammals and reptiles of these forests, using distribution data obtained from recently published reviews and information collated by the WWF Eastern Africa Coastal Forest Ecoregion Programme. Birds and mammals species were divided into forest specialists and generalists, and forest specialist reptiles into ‘coastal’ and ‘forest’ endemics. The species richness of birds and generalist mammals increased with area, and is probably a result of area-dependent extinction. Only in birds, however, species richness increased with decreasing isolation, suggesting possible isolation-dependent colonization. Forest diversity, associated to altitudinal range, is important for specialist birds and mammals, whose species richness increased with wider altitudinal range. The number of relict coastal endemic and forest endemic reptiles was higher in forests with wider altitudinal ranges and on relatively higher altitude, respectively. Such forests have probably provided a suitable (and perhaps stable) environment for these species through time, thus increasing their persistence. Parsimony analysis of distributions (PAD) and cluster analyses showed geographical distance and general ecological similarity among forests as a determinant factor in bird distribution patterns, with compositional similarity decreasing with increasing inter-forest distance. Compositional similarity patterns of mammals among the forests did not show a strong geographical correspondence or a significant correlation with inter-forest distance, and those of reptiles were not resolved, with very low similarity levels among forest faunas. Our results suggest that the relative importance (and causal relationship) of forest attributes affecting the distribution of the East African coastal forest vertebrate fauna varies depending on life history traits such as dispersal ability and forest specialization. The groupings in PAD are partly congruent with some of the previous classifications of areas of endemism for this region, supporting the ‘naturalness’ of these regions.     

Current and historical factors influencing patterns of species richness and turnover of birds in the Gulf of Guinea highlands

Aim The aims of this paper are to: examine how current and historical ecological factors affect patterns of species richness, endemism and turnover in the Gulf of Guinea highlands, test theoretical biogeographical predictions and provide information for making informed conservation decisions. Location The Gulf of Guinea highlands in West Africa. Methods We used multivariate and matrix regression models, and cluster analyses to assess the influence of current climate and current and historical isolation on patterns of richness and turnover for montane birds across the highlands. We examined three groups of birds: montane species (including widespread species), montane endemics and endemic subspecies. We applied a complementarity‐based reserve selection algorithm using species richness with irreplaceability measures to identify areas of high conservation concern. Results Environmental factors influenced richness for all groups of birds (species, endemic species and subspecies). Areas with high and consistent annual rainfall showed the highest species and endemic richness. Species clusters for all groups of birds generally differentiated three major montane regions, which are topographically isolated. Multiple mantel tests identified these same regions for endemic species and subspecies. The influence of historical isolation varied by species group; distributions of endemic montane species and subspecies were more associated with historical breaks than were all montane species, which included widespread non‐endemic species. Main conclusions Our analyses indicated important geographical structure amongst the bird assemblages in the highlands and, therefore, conservation prioritization should include mountains from within the geographical subregions identified in these analyses because these regions may harbour evolutionarily distinct populations of birds.     

Species–energy relationships and habitat complexity in bird communities

Species–energy theory is a commonly invoked theory predicting a positive relationship between species richness and available energy. The More Individuals Hypothesis (MIH) attempts to explain this pattern, and assumes that areas with greater food resources support more individuals, and that communities with more individuals include more species. Using a large dataset for North American birds, I tested these predictions of the MIH, and also examined the effect of habitat complexity on community structure. I found qualitative support for the relationships predicted by the MIH, however, the MIH alone was inadequate for fully explaining richness patterns. Communities in more productive sites had more individuals, but they also had more even relative abundance distributions such that a given number of individuals yielded a greater number of species. Richness and evenness were also higher in structurally complex forests compared to structurally more simple grasslands when controlling for available energy.     

Explaining large-scale patterns of vertebrate diversity

The major clades of vertebrates differ dramatically in their current species richness, from 2 to more than 32 000 species each, but the causes of this variation remain poorly understood. For example, a previous study noted that vertebrate clades differ in their diversification rates, but did not explain why they differ. Using a time-calibrated phylogeny and phylogenetic comparative methods, I show that most variation in diversification rates among 12 major vertebrate clades has a simple ecological explanation: predominantly terrestrial clades (i.e. birds, mammals, and lizards and snakes) have higher net diversification rates than predominantly aquatic clades (i.e. amphibians, crocodilians, turtles and all fish clades). These differences in diversification rates are then strongly related to patterns of species richness. Habitat may be more important than other potential explanations for richness patterns in vertebrates (such as climate and metabolic rates) and may also help explain patterns of species richness in many other groups of organisms.     

Species Richness Patterns and Water-Energy Dynamics in the Drylands of Northwest China

Dryland ecosystems are highly vulnerable to climatic and land-use changes, while the mechanisms underlying patterns of dryland species richness are still elusive. With distributions of 3637 native vascular plants, 154 mammals, and 425 birds in Xinjiang, China, we tested the water-energy dynamics hypothesis for species richness patterns in Central Asian drylands. Our results supported the water-energy dynamics hypothesis. We found that species richness of all three groups was a hump-shaped function of energy availability, but a linear function of water availability. We further found that water availability had stronger effects on plant richness, but weaker effects on vertebrate richness than energy availability. We conducted piecewise linear regressions to detect the breakpoints in the relationship between species richness and potential evapotranspiration which divided Xinjiang into low and high energy regions. The concordance between mammal and plant richness was stronger in high than in low energy regions, which was opposite to that between birds and plants. Plant richness had stronger effects than climate on mammal richness regardless of energy levels, but on bird richness only in high energy regions. The changes in the concordance between vertebrate and plant richness along the climatic gradient suggest that cautions are needed when using concordance between taxa in conservation planning.     

Spatial turnover in the global avifauna

Despite its wide implications for many ecological issues, the global pattern of spatial turnover in the occurrence of species has been little studied, unlike the global pattern of species richness. Here, using a database on the breeding distributions of birds, we present the first global maps of variation in spatial turnover for an entire taxonomic class, a pattern that has to date remained largely a matter of conjecture, based on theoretical expectations and extrapolation of inconsistent patterns from different biogeographic realms. We use these maps to test four predictions from niche theory as to the form that this variation should take, namely that turnover should increase with species richness, towards lower latitudes, and with the steepness of environmental gradients and that variation in turnover is determined principally by rare (restricted) species. Contrary to prediction, we show that turnover is high both in areas of extremely low and high species richness, does not increase strongly towards the tropics, and is related both to average environmental conditions and spatial variation in those conditions. These results are closely associated with a further important and novel finding, namely that global patterns of spatial turnover are driven principally by widespread species rather than the restricted ones. This complements recent demonstrations that spatial patterns of species richness are also driven principally by widespread species, and thus provides an important contribution towards a unified model of how terrestrial biodiversity varies both within and between the Earth's major land masses.     

Do protected areas in urban and rural landscapes differ in species diversity?

Previous studies from Central Europe and North America showed that species richness is higher in urban than in rural landscapes. Do protected areas, which can be found in both city and countryside, reflect this species richness pattern? The impact of urban land-use might reduce conservation success and necessitate special management strategies. We compared species richness and species spatial turnover of selected animal and plant taxa (carabids, butterflies, snails, birds, lichens, mosses, vascular plants) in 30 protected areas in the city of Halle and 56 protected areas in the adjacent rural district of Saalkreis (Central Germany). Species were mapped by experienced biologists within a systematic species inventory. We corrected species numbers for the effects of landscape structure (e.g. size, shape and distance of habitats) which might influence species diversity beyond urbanisation effects. Butterflies, birds and lichens had significantly higher species numbers in the rural protected areas. Species spatial turnover was higher among urban areas than among rural areas or pairs of urban and rural areas for most taxa. Diversity in all taxa depended on the size of a protected area. We discussed these patterns in the context of the general urban-rural species diversity patterns. Our results indicate an increasing isolation of species assemblages with urbanisation and highlight that space for protected areas is even more limited in urban than rural areas. An effective conservation of urban species diversity should include both typical urban and semi-natural habitats to cover the full range of species living in cities.     

Nestedness in fragmented landscapes: birds of the box‐ironbark forests of south‐eastern Australia

Nestedness in biota as a function of species richness – biota of depauperate assemblages being non‐random subsets of richer biotas – has been widely documented in recent years (see Wright et al. 1998 , Oecologia 113: 1–20). Ordering sites by richness maximizes nestedness indices; however, ordering by other criteria such as area or isolation may be more ecologically interpretable. We surveyed birds in true fragments (35 in all), and in “reference areas” in large extant forest blocks (30 locations), of the same range of areas (10, 20, 40, 80 ha). The avifauna was divided into “bush birds”– species dependent on forest and woodland, and “open country” species. We looked at nestedness in four data sets: “bush birds” in fragments and reference areas, and “all birds” in fragments and in reference areas. All data sets were significantly nested. Ordering by area in all cases was not significantly less nested than ordering by richness. Ordering by area in fragments was significantly greater than in reference areas, but the differences in standardized nestedness indices were small (<15%). We identified those birds that had distributions among fragments that conformed strongly with area, those that were more randomly distributed and some species that were more likely to occupy the smallest fragments. Among the latter was a hyperaggressive, invasive, colonial native species (noisy miner Manorina melanocephala). A suite of small, insectivorous birds were more likely to strongly conform with expected distributions in relation to area, which was consistent with observations of their vulnerability to the effects of the noisy miner in smaller fragments.     

Birds and people in Europe

At a regional scale, species richness and human population size are frequently positively correlated across space. Such patterns may arise because both species richness and human density increase with energy availability. If the species-energy relationship is generated through the 'more individuals' hypothesis, then the prediction is that areas with high human densities will also support greater numbers of individuals from other taxa. We use the unique data available for the breeding birds in Europe to test this prediction. Overall regional densities of bird species are higher in areas with more people; species of conservation concern exhibit the same pattern. Avian density also increases faster with human density than does avian biomass, indicating that areas with a higher human density have a higher proportion of small-bodied individuals. The analyses also underline the low numbers of breeding birds in Europe relative to humans, with a median of just three individual birds per person, and 4 g of bird for every kilogram of human.