Species Richness and Evolutionary Niche Dynamics: A Spatial Pattern-Oriented Simulation Experiment

Multiple determinants of Australian tropical frog biodiversity

For optimal planning of conservation and monitoring measures, it is important to know the spatial pattern of species richness and especially areas with high species richness. A spatial pattern of the species richness of ants in Georgia (Caucasus) was modeled, areas with the highest number of ant’s species were inferred, and climatic factors that influence the pattern of ant diversity were identified. A database was created by accumulating occurrences for 63 ant species, including 256 localities and 2,018 species/occurrences. Species richness was positively correlated with variables associated with temperature and negatively correlated with variables associated with precipitation. Species richness reaches a maximum at the elevations 800–1,200 m a.s.l. and declines at both lower and higher altitudes. The role of climatic variables and geography of the study area in determining the observed pattern of species richness is discussed.

We introduce a novel framework for conceptualising, quantifying and unifying discordant patterns of species richness along geographical gradients. While not itself explicitly mechanistic, this approach offers a path towards understanding mechanisms. In this study, we focused on the diverse patterns of species richness on mountainsides. We conjectured that elevational range midpoints of species may be drawn towards a single midpoint attractor - a unimodal gradient of environmental favourability. The midpoint attractor interacts with geometric constraints imposed by sea level and the mountaintop to produce taxon-specific patterns of species richness. We developed a Bayesian simulation model to estimate the location and strength of the midpoint attractor from species occurrence data sampled along mountainsides. We also constructed midpoint predictor models to test whether environmental variables could directly account for the observed patterns of species range midpoints. We challenged these models with 16 elevational data sets, comprising 4500 species of insects, vertebrates and plants. The midpoint predictor models generally failed to predict the pattern of species midpoints. In contrast, the midpoint attractor model closely reproduced empirical spatial patterns of species richness and range midpoints. Gradients of environmental favourability, subject to geometric constraints, may parsimoniously account for elevational and other patterns of species richness.

Most studies examining continental-to-global patterns of species richness rely on the overlaying of extent-of-occurrence range maps. Because a species does not occur at all locations within its geographic range, range-map-derived data represent actual distributional patterns only at some relatively coarse and undefined resolution. With the increasing availability of high-resolution climate and land-cover data, broad-scale studies are increasingly likely to estimate richness at high resolutions. Because of the scale dependence of most ecological phenomena, a significant mismatch between the presumed and actual scale of ecological data may arise. This may affect conclusions regarding basic drivers of diversity and may lead to errors in the identification of diversity hotspots. Here, we examine avian range maps of 834 bird species in conjunction with geographically extensive survey data sets on two continents to determine the spatial resolutions at which range-map data actually characterize species occurrences and patterns of species richness. At resolutions less than 2 degrees ( approximately 200 km), range maps overestimate the area of occupancy of individual species and mis-characterize spatial patterns of species richness, resulting in up to two-thirds of biodiversity hotspots being misidentified. The scale dependence of range-map accuracy poses clear limitations on broad-scale ecological analyses and conservation assessments. We suggest that range-map data contain less information than is generally assumed and provide guidance about the appropriate scale of their use.

Quantifying spatial patterns of species richness and determining the processes that give rise to these patterns are core problems in biodiversity theory. The aim of the present paper was to more accurately detect patterns of vascular species richness at different scales along altitudinal gradients in order to further our understanding of biodiversity patterns and to facilitate studies on relationships between biodiversity and environmental factors. Species richness patterns of total vascular plants species, including trees, shrubs, and herbs, were measured along an altitudinal gradient on one transect on a shady slope in the Dongling Mountains, near Beijing, China. Direct gradient analysis, regression analysis, and geostatistics were applied to describe the spatial patterns of species richness. We found that total vascular species richness did not exhibit a linear pattern of change with altitude, although species groups with different ecological features showed strong elevational patterns different from total species richness. In addition to total vascular plants, analysis of trees, shrubs, and herbs demonstrated remarkable hierarchical structures of species richness with altitude (i.e. patchy structures at small scales and gradients at large scales). Species richness for trees and shrubs had similar spatial characteristics at different scales, but differed from herbs. These results indicated that species groups with similar ecological features exhibit similar biodiversity patterns with altitude, and studies of biodiversity based on species groups with similar ecological properties or life forms would advance our understanding of variations in species diversity. Furthermore, the gradients or trends appeared to be due mainly to local variations in species richness means with altitude. We also found that the range of spatial scale dependencies of species richness for total vascular plants, trees, shrubs, and herbs was relatively large. Thus, to detect the relationships between species richness with environmental factors along altitudinal gradients, it was necessary to quantify the scale dependencies of environmental factors in the sampling design or when establishing non‐linear models.(Managing editor: Ya‐Qin Han)

Summary Climate and topography are among the most fundamental drivers of plant diversity. Here, we assessed the importance of climate and topography in explaining diversity patterns of species richness, endemic richness and endemicity on the landscape scale of an oceanic island and evaluated the independent contribution of climatic and topographic variables to spatial diversity patterns. We constructed a presence/absence matrix of perennial endemic and native vascular plant species (including subspecies) in 890 plots on the environmentally very heterogeneous island of La Palma, Canary Islands. Species richness, endemic richness and endemicity were recorded, interpolated and related to climate (i.e. variables describing temperature, precipitation, variability and climatic rarity) and topography (i.e. topographic complexity, solar radiation, geologic age, slope and aspect). We used multimodel inference, spatial autoregressive models, variance partitioning and linear regression kriging as statistical methods. Species richness is best explained by both climatic and topographic variables. Topographic variables (esp. topographic complexity and solar radiation) explain endemic richness, and climatic variables (esp. elevation/temperature and rainfall seasonality) explain endemicity. Spatial patterns of species richness, endemic richness and endemicity were in part geographically decoupled from each other. Synthesis. We identified several topography‐dependent processes ranging from evolutionary processes (micro‐refugia, in situ speciation, pre‐adaptation to rupicolous conditions, dispersal limitations) to human‐induced influences (introduced herbivores, fire, land use) that possibly shape the endemic richness pattern of La Palma. In contrast, climate mainly drives endemicity, which is connected to ecological speciation and specialization to local conditions. We highlight the importance of incorporating climatic variability into future studies of plant species diversity and endemism. The spatial incongruence in hot spots of species richness, endemic richness and endemicity emphasizes the need for an integrated conservation approach acknowledging different diversity measures to protect the complete spectrum of diversity. High‐elevation islands such as La Palma are highly suitable to study drivers of diversity and endemism, as they offer environmental gradients of continental magnitude on the landscape scale of a single climatic mini‐continent and a large array of in situ‐speciated endemics.

AimsThe herb–shrub flora has been widely neglected in science and conservation policy throughout the world, so that this biodiversity component remains largely unknown. The objective of this study was to elucidate the spatial patterns of species richness and phytogeographic regions of the Cerrado herb–shrub flora, and to estimate the percentage of areas with high species richness that is still covered by natural vegetation and is located in protected areas or priority areas for conservation.LocationCentral Brazil and surrounding territories.MethodsTo estimate the pattern of species richness we produced species distribution models (SDMs) for 5362 species using botanical records combined with 17 least correlated environmental variables. All SDMs were tested against a bias‐corrected null model and the 5039 significant SDMs were stacked to generate the pattern of botanical richness. The resulting presence–absence matrix was subjected to a partitioning around medoids (PAM) cluster analysis to delineate phytogeographic regions.ResultsThe pattern of species richness indicated that highest levels of richness are in the central–south and west parts of the Cerrado. The study recognized two floristically distinct clusters at the core of the Cerrado region, and seven additional regions that share floristic elements with the two core clusters. Many areas with high levels of modelled species richness are located outside protected and priority areas for conservation.ConclusionsWe recognized patterns of regional variation, reflected in the phytogeographic division. Furthermore, we show that the current protected areas in Brazil do not effectively protect high richness areas of the herb–shrub flora.

Determining how ecological and evolutionary processes produce spatial variation in local species richness remains an unresolved challenge. Using mountains as a model system, we outline an integrative research approach to evaluate the influence of ecological and evolutionary mechanisms on the generation and maintenance of patterns of species richness along and among elevational gradients. Biodiversity scientists interested in patterns of species richness typically start by documenting patterns of species richness at regional and local scales, and based on their knowledge of the taxon, and the environmental and historical characteristics of a mountain region, they then ask whether diversity–environment relationships, if they exist, are explained mostly by ecological or evolutionary hypotheses. The final step, and perhaps most challenging one, is to tease apart the relative influence of ecological and evolutionary mechanisms. We propose that elucidating the relative influence of ecological and evolutionary mechanisms can be achieved by taking advantage of the replicated settings afforded by mountains, combined with targeted experiments along elevational gradients. This approach will not only identify potential mechanisms that drive patterns of species richness, but also allow scientists to generate more robust hypotheses about which factors generate and maintain local diversity.

The mechanisms that govern spatial patterns in species richness, community composition, and specia tion are fundamental issues in community and evolu tionary ecology. It has been argued recently (Emerson and Kolm 2005a) that, in the Hawaiian and Canary archipelagos, species diversity itself has been driving the speciation rates of endemic plants and arthropods, in accord with island biogeography theory. Here we show that the observed trends in the distribution of endemics within these two archipelagos are in fact consistent with null predictions. Moreover, we show that similar trends are followed by non-endemic singletons (i.e., species not endemic to the archipelago that occur on a single island), despite the different mode by which they first arose in the archipelago. Together, these findings undermine the validity of the diversity-driven diversi fication hypothesis, at least in the systems considered here. Is biodiversity simply the outcome of ecological and evolutionary processes, or could it also be a causative agent in driving speciation? The idea that species can modify their environment, thereby making it available for additional species, is not new (Jones et al. 1997, Erwin 2005). Emerson and Kolm (2005<2, b) have gone further to suggest that species richness can promote diversification, a prediction they ascribe to the theory of island biogeography (MacArthur and Wilson 1967). Assuming that extinction and speciation rates follow a similar curvilinear dependence on richness, Emerson and Kolm predict that, under equilibrium conditions, the proportional representation of endemic singletons

Aim To test for control of vascular plant species richness in the riparian corridor by exploring three contrasting (although not mutually exclusive) hypotheses: (1) longitudinal patterns in riparian plant species richness are governed by local, river‐related processes independent of the regional species richness, (2) riparian plant species richness is controlled by dispersal along the river (longitudinal control), and (3) the variation in riparian plant species richness mirrors variation in regional richness (lateral control).Location The riparian zones of the free‐flowing Vindel River and its surrounding river valley, northern Sweden.Methods We used data from three surveys, undertaken at 10‐year intervals, of riparian reaches (200‐m stretches of riverbank) spanning the entire river. In addition, we surveyed species richness of vascular plants in the uplands adjacent to the river in 3.75‐km2 large plots along the same regional gradient. We explored the relationship between riparian and upland flora, and various environmental variables. We also evaluated temporal variation in downstream patterns of the riparian flora.Results Our results suggest that local species richness in boreal rivers is mainly a result of local, river‐related processes and dispersal along the corridor. The strongest correlation between species richness and the environment was a negative one between species number and soil pH, but pH varied within a narrow range. We did not find evidence for a correlation between species richness on regional and local scales. We found that the local patterns of species richness for naturally occurring vascular plants were temporally variable, probably in response to large‐scale disturbance caused by extreme floods. Most previous studies have found a unimodal pattern of species richness with peaks in the middle reaches of a river. In contrast, on two of three occasions corresponding to major flooding events, we found that the distribution of species richness of naturally occurring vascular plants resembled that of regional diversity: a monotonic decrease from headwater to coast. We also found high floristic similarity between the riparian corridor and the surrounding landscape.Main conclusions These results suggest that local processes control patterns of riparian species richness, but that species composition is also highly dependent on the regional species pool. We argue that inter‐annual variation in flood disturbance is probably the most important factor producing temporal variability of longitudinal species richness patterns.

The distributions of most pollinator species are poorly documented despite their importance in providing ecosystem services. While these and other organisms are threatened by many aspects of the human enterprise, anthropogenic climate change is potentially the most severe threat to pollinator biodiversity. Mounting evidence demonstrates that there have already been biotic responses to the relatively small climate changes that have occurred this century. These include wholesale shifts of relatively well-documented butterfly and bird species in Europe and North America. Although studies of such phenomena are supported by circumstantial evidence, their findings are also consistent with predictions derived from current models of spatial patterns of species richness. Using new GIS methods that are highly precise and accurate, I document spatial patterns of Canadian butterfly diversity. These are strongly related to contemporary climate and particularly to potential evapotranspiration. An even more noteworthy finding is the fact that, for the first time, habitat heterogeneity, measured as the number of land cover types in each study unit, is proven to be an equally strong predictor of butterfly richness in a region where energy alone was thought to be the best predictor of diversity. Although previous studies reveal similar relationships between energy and diversity, they fail to detect the powerful link between richness and habitat heterogeneity. The butterflies of Canada provide a superb baseline for studying the effects of climate on contemporary patterns of species richness and comprise the only complete pollinator taxon for which this sort of analysis is currently possible.

Diversität und Biogeographie der Farne und Vögel Boliviens: Niche Modellierung GIS Applicationen

Rapoport effect predicts that species geographic range sizes will increase toward higher latitudes, probably reflecting adaptations to extreme climatic conditions that increase species tolerance. Recently, studies about spatial patterns in species richness and geographic range size may be associated with the geometry of species' ranges. In this context, null models can be used to search for the causal mechanisms associated with these patterns. In this paper, we analyzed Rapoport effect using a null model to evaluate how phylogenetic structure and geometric constraints simultaneously affect latitudinal extents of 40 species of South American terrestrial Carnivora. The latitudinal extents of Carnivora tended to decrease toward Southern latitudes, in the opposite direction expected under a simple Rapoport effect, but in accordance to geometric expectations of position of midpoints in the continent. Using 5000 simulations, it was possible to show that the null regression coefficients of latitudinal extents against midpoints are positively biased, reflecting the geometric constraints in the latitudinal extents. The results were equivalent in phylogenetic and non-phylogenetic analyses. The observed regression coefficient was significantly smaller (line is less inclined) than expected by chance alone, demonstrating that the geometric constraints in the latitudinal extents exist even after controlling for phylogenetic structure in data using eigenvector regressions. This suggests that the "spirit" of Rapoport effect (sensu Lyons & Willig, 1997) could be maintained, i.e., that latitudinal extents in Southern region of the continent are relatively larger than those in Northern regions, even after controlling for phylogenetic effects.

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.

Knowledge on the spatial distribution of plant species richness and its associated factors is crucial for addressing several ecological problems. Climate factors have long been used to explain species richness patterns across space and time. In this study, we investigated how water and energy variables shape species richness patterns on the alpine Tibetan plateau. An empirical study involving 800 plots of size 1 m2 on 80 different sites was conducted across two transects, covering the arid and semi‐arid alpine region of the western part of the Tibetan plateau to explore plant species composition and richness in relation to climatic factors at a regional scale. We performed detrended correspondence analysis and generalized linear models to explore the effect of energy and water variables on species composition and species richness, respectively. We used a generalized linear mixed model to investigate the interactive effect of energy variables and water variables on species richness. We found that both energy variables and water variables significantly influenced species richness and composition on the alpine Tibetan plateau and supported the water–energy dynamics (WED) hypothesis. Specificall, plant species richness increased with greater liquid water availability. However, it exhibited a unimodal relationship with potential evapotranspiration. Our results also showed that precipitation played a more substantial role in the model than potential evapotranspiration, but their interaction strengthened the model. In conclusion, this study highlights the dominant role of precipitation in explaining species composition and richness patterns on the alpine Tibetan plateau and suggests conducting research on additional environmental factors affecting water availability in this region.