Disparity between range map- and survey-based analyses of species richness: patterns, processes and implications

Species richness patterns are characterized either by overlaying species range maps or by compiling geographically extensive survey data for multiple local communities. Although, these two approaches are clearly related, they need not produce identical richness patterns because species do not occur everywhere in their geographical range. Using North American breeding birds, we present the first continent‐wide comparison of survey and range map data. On average, bird species were detected on 40.5% of the surveys within their range. As a result of this range porosity, the geographical richness patterns differed markedly, with the greatest disparity in arid regions and at higher elevations. Environmental productivity was a stronger predictor of survey richness, while elevational heterogeneity was more important in determining range map richness. In addition, range map richness exhibited greater spatial autocorrelation and lower estimates of spatial turnover in species composition. Our results highlight the fact that range map richness represents species coexistence at a much coarser scale than survey data, and demonstrate that the conclusions drawn from species richness studies may depend on the data type used for analyses.     

Niche breadth predicts geographical range size: a general ecological pattern

The range of resources that a species uses (i.e. its niche breadth) might determine the geographical area it can occupy, but consensus on whether a niche breadth–range size relationship generally exists among species has been slow to emerge. The validity of this hypothesis is a key question in ecology in that it proposes a mechanism for commonness and rarity, and if true, may help predict species' vulnerability to extinction. We identified 64 studies that measured niche breadth and range size, and we used a meta‐analytic approach to test for the presence of a niche breadth–range size relationship. We found a significant positive relationship between range size and environmental tolerance breadth (z = 0.49), habitat breadth (z = 0.45), and diet breadth (z = 0.28). The overall positive effect persisted even when incorporating sampling effects. Despite significant variability in the strength of the relationship among studies, the general positive relationship suggests that specialist species might be disproportionately vulnerable to habitat loss and climate change due to synergistic effects of a narrow niche and small range size. An understanding of the ecological and evolutionary mechanisms that drive and cause deviations from this niche breadth–range size pattern is an important future research goal.     

Parasite species richness in carnivores: effects of host body mass, latitude, geographical range and population density

Aim  Comparative studies have revealed strong links between ecological factors and the number of parasite species harboured by different hosts, but studies of different taxonomic host groups have produced inconsistent results. As a step towards understanding the general patterns of parasite species richness, we present results from a new comprehensive data base of over 7000 host–parasite combinations representing 146 species of carnivores (Mammalia: Carnivora) and 980 species of parasites.
Methods  We used both phylogenetic and non-phylogenetic comparative methods while controlling for unequal sampling effort within a multivariate framework to ascertain the main determinants of parasite species richness in carnivores.
Results  We found that body mass, population density, geographical range size and distance from the equator are correlated with overall parasite species richness in fissiped carnivores. When parasites are classified by transmission mode, body mass and home range area are the main determinants of the richness of parasites spread by close contact between hosts, and population density, geographical range size and distance from the equator account for the diversity of parasites that are not dependent on close contact. For generalist parasites, population density, geographical range size and latitude are the primary predictors of parasite species richness. We found no significant ecological correlates for the richness of specialist or vector-borne parasites.
Main conclusions  Although we found that parasite species richness increases instead of decreases with distance from the equator, other comparative patterns in carnivores support previous findings in primates, suggesting that similar ecological factors operate in both these independent evolutionary lineages.     

The effect of the taxon and geographic range size of host eucalypt species on the species richness of gall-forming insects

Abstract This field study was designed to test whether the taxonomic group and geographic range size of a host plant species, usually found to influence insect species richness in other parts of the world, affected the number of gall species on Australian eucalypts. We assessed the local and regional species richness of gall-forming insects on five pairs of closely related eucalypt species. One pair belonged to the subgenus Corymbia, one to Monocalyptus, and three to different sections of Symphyomyrtus. Each eucalypt pair comprised a large and a small geographic range species. Species pairs were from coastal or inland regions of eastern Australia. The total number of gall species on eucalypt species with large geographic ranges was greater than on eucalypt species with small ranges, but only after the strong effect of eucalypt taxonomic grouping was taken into account. There was no relationship between the geographic range size of eucalypt species and the size of local assemblages of gall species, but the variation in insect species composition between local sites was higher on eucalypt species with large ranges than on those with small ranges. Thus the effect of host plant range size on insect species richness was due to greater differentiation between more widespread locations, rather than to greater local species richness. This study confirms the role of the geographic range size of a host plant in the determination of insect species richness and provides evidence for the importance of the taxon of a host plant.     

Patterns of diversity and range size of selected plant groups along an elevational transect in the Bolivian Andes

Patterns of point, -, and -diversity, and of an index of range size were studied for Acanthaceae, Araceae, Bromeliaceae, Melastomataceae, Palmae, and Pteridophyta in 204 vegetation plots of 400 m² along an elevational gradient in humid montane forest at 220–3950 m in central Bolivia. Zonal forest point-diversity (mean species number per plot) and -diversity (extrapolated total species number) showed either hump-shaped curves or constant values from lowlands to mid-elevations, followed by a steep decline. These patterns correspond to the hypothesis of maximum diversity at intermediate levels of productivity, while for pteridophytes and epiphytic Bromeliaceae they also fitt a null-model of random distribution within bounded geographical ranges. The ratio of point to -diversity was surprisingly constant across study groups and elevation with values of 0.2–0.4. Range size index (mean inverse range size of all species recorded at a given elevation) either increased with elevation, showed hump-shaped patterns, or remained roughly constant. Non-zonal habitats (ravines, ridge-tops, rock faces, roadsides, pastures) contributed significantly to overall species number for terrestrial herbs and shrubs of Bromeliaceae, Melastomataceae, and Pteridophyta, but not for Acanthaceae, Araceae, and epiphytic taxa. With few exceptions, endemism was most pronounced in zonal forests. These results imply that conservation measures should focus on zonal forests in a few large reserves at low and mid-elevations and in many smaller reserves at higher elevations.     

Geographical patterns and determinants of species richness in Mexico across selected families of vascular plants: implications for conservation

Mexico is considered a megadiverse country containing more than 10% of the world's biodiversity. The distribution of this species richness and endemism is different among the different Mexican states. We examined the species richness patterns of 13 families of vascular plants (including ferns, gymnosperms and angiosperms) in Mexico using political divisions (states) as units of analysis. We analysed the species richness values (absolute richness, endemic richness and restrictive richness) of these plant families using stepwise multiple regression analysis, assessing their relationship with a set of 10 environmental variables (expressed as heterogeneity coefficients). A combined cluster analysis with multidimensional scaling analysis (MDS) and an analysis of similarities were also undertaken to define the spatial–geographical patterns. Additionally, we proposed a methodological strategy to determine which states of Mexico have priorities for conservation. Our results suggested that the three species richness values used were significantly predicted by environmental factors, especially by climatic heterogeneity. Notwithstanding that a linear pattern was recognized, the Mexican states were gathered in four groups, which were confirmed by the MDS and the cluster analysis: (1) the Yucatan Peninsula, (2) arid Mexico, (3) the Mexican Transition Zone and (4) the megadiverse states. We proposed that 12 Mexican states include all the environmental conditions and are candidates for developing conservation programmes: (1) Quintana Roo, Tabasco and Yucatán, (2) Baja California, Chihuahua and Sinaloa, (3) Guerrero, Jalisco and Nuevo León and (4) Chiapas, Oaxaca and Veracruz.     

Global patterns in the shape of species geographical ranges reveal range determinants

Aim Do species range shapes follow general patterns? If so, what mechanisms underlie those patterns? We show for 11,582 species from a variety of taxa across the world that most species have similar latitudinal and longitudinal ranges. We then seek to disentangle the roles of climate, extrinsic dispersal limitation (e.g. barriers) and intrinsic dispersal limitation (reflecting a species’ ability to disperse) as constraints of species range shape. We also assess the relationship between range size and shape. Location Global. Methods Range shape patterns were measured as the slope of the regression of latitudinal species ranges against longitudinal ranges for each taxon and continent, and as the coefficient of determination measuring the degree of scattering of species ranges from the 1:1 line (i.e. latitudinal range = longitudinal range). Two major competing hypotheses explaining species distributions (i.e. dispersal or climatic determinism) were explored. To this end, we compared the observed slopes and coefficients of determination with those predicted by a climatic null model that estimates the potential range shapes in the absence of dispersal limitation. The predictions compared were that species distribution shapes are determined purely by (1) intrinsic dispersal limitation, (2) extrinsic dispersal limitations such as topographic barriers, and (3) climate. Results  Using this methodology, we show for a wide variety of taxa across the globe that species generally have very similar latitudinal and longitudinal ranges. However, neither neutral models assuming random but spatially constrained dispersal, nor models assuming climatic control of species distributions describe range shapes adequately. The empirical relationship between the latitudinal and longitudinal ranges of species falls between the predictions of these competing models. Main conclusions We propose that this pattern arises from the combined effect of macroclimate and intrinsic dispersal limitation, the latter being the major determinant among restricted‐range species. Hence, accurately projecting the impact of climate change onto species ranges will require a solid understanding of how climate and dispersal jointly control species ranges.     

Modelling geographical patterns in species richness using eigenvector-based spatial filters

Aim To test the mechanisms driving bird species richness at broad spatial scales using eigenvector‐based spatial filtering. Location South America. Methods An eigenvector‐based spatial filtering was applied to evaluate spatial patterns in South American bird species richness, taking into account spatial autocorrelation in the data. The method consists of using the geographical coordinates of a region, based on eigenanalyses of geographical distances, to establish a set of spatial filters (eigenvectors) expressing the spatial structure of the region at different spatial scales. These filters can then be used as predictors in multiple and partial regression analyses, taking into account spatial autocorrelation. Autocorrelation in filters and in the regression residuals can be used as stopping rules to define which filters will be used in the analyses. Results Environmental component alone explained 8% of variation in richness, whereas 77% of the variation could be attributed to an interaction between environment and geography expressed by the filters (which include mainly broad‐scale climatic factors). Regression coefficients of environmental component were highest for AET. These results were unbiased by short‐scale spatial autocorrelation. Also, there was a significant interaction between topographic heterogeneity and minimum temperature. Conclusion Eigenvector‐based spatial filtering is a simple and suitable statistical protocol that can be used to analyse patterns in species richness taking into account spatial autocorrelation at different spatial scales. The results for South American birds are consistent with the climatic hypothesis, in general, and energy hypothesis, in particular. Habitat heterogeneity also has a significant effect on variation in species richness in warm tropical regions.     

Gradual changes in range size accompany long‐term trends in species richness

Species richness has long been used as an indicator of ecosystem functioning and health. Global richness is declining, but it is unclear whether sub‐global trends differ. Regional trends are especially understudied, with most focused on island regions where richness is strongly impacted by novel colonisations. We addressed this knowledge gap by testing for multi‐decade trends in species richness in nine open marine regions around North America (197 region‐years) while accounting for imperfect observations and grounding our findings in species‐level range dynamics. We found positive richness trends in eight of nine regions, four of which were statistically significant. Species' range sizes generally contracted pre‐extinction and expanded post‐colonisation, but the ranges of transient species expanded over the long‐term, slowly increasing their regional retention and driving increasing richness. These results provide more evidence that sub‐global richness trends are stable or increasing, and highlight the utility of range size for understanding richness dynamics.     

Ecological correlates of geographical range occupancy in North American birds

Aim  The degree to which a species is predictably encountered within its range varies tremendously across species. Understanding why some species occur less frequently within their range than others has important consequences for conservation and for analyses of ecological patterns based on range maps. We examined whether patterns in geographical range occupancy can be explained by species-level traits.
Location  North America.
Methods  We used survey data from 1993 to 2002 from the North American Breeding Bird Survey along with digital range maps produced by NatureServe to calculate range occupancy for 298 species of terrestrial birds. We tested whether species traits explained variation in range occupancy values using linear regression techniques.
Results  We found three species traits that together explained more than half of the variation in range occupancy. Population density and niche breadth were positively correlated with occupancy, while niche position was negatively correlated with occupancy.
Main conclusions  Our results suggest that high range occupancy will occur in species that are common at sites on which they occur, that tolerate a relatively wide range of ecological conditions and that tend to have ranges centred on areas with common environmental conditions. Furthermore, it appears that niche-based characteristics may explain patterns of distribution and abundance from local habitats up to the scale of geographical ranges.     

The relationships between abundance, range size and niche breadth in Central European tree species

Aim  Range size and niche breadth have been found to be positively related to abundance in many plant and animal groups. We tested these two relationships for the tree species flora of Central Europe; that is, for all 25 species that have their distribution centre in this region.
Location  Eurasia, with a focus on Central Europe.
Methods  We devised an abundance and niche variable classification system to transform the existing literature data into a semi-quantitative assessment of abundance and niche breadth (in terms of soil chemical and physical variables, and temperature) for each of the 25 tree species. Regression analyses between abundance, range size and niche breadth were conducted for the entire species sample and for subsets of species defined by their ecology or phylogeny.
Results  The relationship between abundance in the distribution centre and range size was weak for the Central European tree species. However, significant abundance–range size relationships were found for phylogenetically or ecologically more homogenous species groups (for example for trees of the order Rosales and for mid-successional tree species). Realized niche breadth was positively related to range size in the case of temperature, but not for soil-related variables. No relationship existed between niche breadth and abundance in the distribution centre.
Main conclusions  We hypothesize that the weak relationship between abundance and range size is primarily a consequence of substantial ecological and phylogenetic heterogeneity within this rather species-poor assemblage. The positive relationship between realized temperature niche breadth and range size emphasizes the strong influence of climatic variables on plant distribution patterns over continental or global scales.     

中国蔷薇科植物多样性格局及其资源植物保护现状

蔷薇科(Rosaceae)是在中国广泛分布并具有重要经济价值的植物类群, 但蔷薇科资源植物的物种多样性格局及其保护状况尚缺乏较系统的评估。该文旨在: 1)整理中国蔷薇科资源植物名录, 显示其物种多样性格局及热点地区, 并探究这一格局的形成机制。2)评估中国蔷薇科资源植物的保护状况, 为其保护规划提供基础数据。通过广泛收集整理《中国植物志》、省级植物志等资料中关于蔷薇科的记录, 建立了中国蔷薇科物种名录(共914种), 确定了物种的主要经济用途(包括食用植物、园林绿化植物、药用植物和水果种质资源), 并建立了每种植物的高精度分布图。在此基础上, 估算了蔷薇科全部物种及主要资源植物类别的物种多样性格局, 并利用广义线性模型和冗余分析探讨了蔷薇科物种多样性格局与环境的关系。最后将物种分布与中国国家级和省级自然保护区进行叠加分析, 评估了蔷薇科植物的保护现状。结果显示: 1)四川盆地北部、东部和西部山区以及横断山区是中国蔷薇科植物的热点地区。2)蔷薇科植物多样性主要受水分因子影响。3)横断山区、云南东南部和西藏东南部等地是保护薄弱物种集中的区域, 而悬钩子属(Rubus)等类群的保护不足。     

Geographical expansion and increased prevalence of common species in avian assemblages: implications for large-scale patterns of species richness

Aim The assumption that ecological patterns at large spatial scales originate exclusively from non‐anthropogenic processes is growing more questionable with the increasing domination of the biosphere by humans. Because common and rare species are known to respond differently to anthropogenic activities at local scales these differential responses could, over time, be reflected in distributional patterns of species richness at larger spatial scales. This work tests the hypothesis that modern processes have played a role in shaping these patterns, by examining recent changes in the structure and composition of assemblages of breeding avifauna over a large geographical extent. Location The portion of North America containing the contiguous United States and southern Canada. Methods Changes in the geographical range structure of breeding avifauna in North America from 1968 to 2003 were analysed in regions containing historically moderate levels of anthropogenic activities. Two geographical measures, extent of occurrence and area of occupancy, were used to identify the level of rarity or commonality of individual species and to estimate, based on a vector analysis, patterns of change in geographical range structure for individual species and avian assemblages. Results More species experienced patterns of geographical range expansion (51%) than contraction (28%). The majority of avian assemblages (43%) displayed patterns of geographical range expansion: common species increased in number and proportion (6%) in association with reciprocal losses in rare and moderately rare species, resulting in a constant level of species richness. The minority of avian assemblages (21%) displayed patterns of geographical range contraction: gains occurred for common species as well as for rare and moderately rare species, resulting in substantial increases in species richness and a decline in the proportion of common species (4%). The remaining avian assemblages presented equivocal patterns characterized by gains in the number and proportion (2%) of common species and gains in species richness. Main conclusions Modern processes have played a role in shaping the distribution patterns of species richness at large spatial scales based on the composition of common and rare species. This suggests that anthropogenic activities cannot be ignored as a possible causal factor when considering ecological patterns at large spatial scales.     

Phylogenetic diversity and the conservation biogeography of African primates

Aim Phylogenetics has an important role in conservation biogeography. However, there are few data on the phylogenetic diversity of African primates. The phylogenetic diversity (PD) of a species is a measure of its taxonomic distinctness and can be estimated by looking at the phylogenetic relationships among taxa. Species‐specific metrics on PD can then be used to determine conservation priorities at various biogeographical scales. We used PD metrics to rank 55 African primate species according to their conservation priorities at the country level and within six African biogeographical regions. We also addressed the following question: are there differences in conservation rankings between the IUCN Red List and our PD metrics? Location Africa. Methods We created a consensus phylogeny for all African primate clades based on genetic studies. Analyses of species distributions were determined using presence/absence scores at two levels: country and biogeographical region. A node‐based method that standardizes for widespread taxa and endemicity was used to calculate PD indices. Hierarchical cluster analysis was used to convert one of the standardized, phylogenetic indices into three clusters that could be ranked and compared with the main IUCN conservation rankings of endangered, vulnerable, and lower risk. Results At the country and region levels, the top‐priority species in terms of PD are Pan paniscus, Macaca sylvanus, Arctocebus calabarensis, Gorilla beringei, Arctocebus aureus, Allenopithecus nigroviridis, Gorilla gorilla, Procolobus verus, Cercopithecus solatus, Cercocebus galeritus, Colobus angolensis, Theropithecus gelada, Galagoides zanzibaricus, Galagoides granti, and Procolobus (Piliocolobus) badius. Geographic rankings were highest for the Democratic Republic of the Congo (country level) and Central Africa (region level). Although there were no overall differences between IUCN conservation ranks and the PD rankings, there were significant differences between the two systems for vulnerable and endangered primate taxa. Main conclusions There are few ecological and behavioural data on populations of some of the African primates that represent the highest levels of phylogenetic diversity. Studies of primate taxa with high PD rankings should focus on identifying sites suitable for intensive studies of population densities, feeding ecology, and reproductive behaviour. We suggest that PD metrics can serve as an important, complementary data set in the IUCN ranking system for primates.     

The relationship between range size and niche breadth: a test using five species of Gammarus (Amphipoda)

Data for five closely related species of gammarid crustaceans are used to examine interspecific relationships between the breadth of fundamental tolerance or capacity and geographical range size. Gammarus duebeni is, almost without exception, the most tolerant species and that with the best physiological performance. Although there is some limited variation, the remaining species can be ranked broadly in the sequence G. zaddachi  > G. salinus  >  G. oceanicus > G. locusta . The wide tolerance and high performance of G. duebeni is associated with the occupation of a wider range of environmental 'types' than any other of the species. In terms of geographical range size, the species can be ranked from most to least widespread in the sequence G. oceanicus  > G. duebeni  >  G. zaddachi  >  G. salinus  >  G. locusta . This provides little support for Brown's hypothesis, or the argument that the more widely distributed species within a taxonomic assemblage also tend to have the widest fundamental niches. However, if marine ( G. oceanicus and G. locusta ) and estuarine ( G. duebeni , G. zaddachi , G. salinus) species are considered separately, then in each case the species with the largest geographical range is also the most tolerant/best performer. In this sense, the jack-of-all-trades is the master-of-all, rather than the master-of-none.     

The implications of palaeontological evidence for theories of ecological communities and species richness

Abstract Palaeontological evidence raises several questions that relate to current explanations of ecological communities, to the classification of communities and to interpretations of species richness. The first question relates to the stability of species detected in the fossil record. Coupled with that is the issue of incidental association of species on the same trophic level through differential effects of climatic change on the different species. Such observations are seen to support the ‘individualistic’ concept of communities. Recent statements about this concept leave unresolved questions about the acquisition of adaptation, and about the place of adaptation theory in theories of ecological communities and interpretations of ‘regional species richness’. At issue is whether there is justification for continuing to classify communities as a basis for understanding them. There is good reason to reject this approach for one in which questions about communities and ‘local’ and ‘regional’ species richness are replaced by more specific and basic questions about the relationship between adaptation, distribution and abundance, and ecological interactions. Some recent efforts to incorporate species theory into community theory fail because their basis remains the flawed concept of ‘local community’.     

Phytogeography, range size and richness of Australian endemic Sauropus (Euphorbiaceae)

Aim To investigate the distribution of Australian species of Sauropus. The information obtained is used to (1) identify areas of highest richness and centres of endemism, (2) investigate latitudinal gradients of richness and range size, (3) determine the types of rarity shown, and (4) provide hypotheses on historical biogeography of the genus within Australia. Location Australia. Methods Specimens from 17 herbaria and field searches were examined and label and field information collated on distribution, habit and habitat. Distribution information was used to map all species within 784 grid cells of 1° × 1° and within the 97 Australian ‘ecological regions’. Morphometric cluster analysis of species was conducted using Kulczynski association and flexible UPGMA on 23 character states. Simple regression was used to correlate species richness, density and range size to changes in latitude. CLIMEX is used to match the climate of the region of highest richness in Australia with other areas of the world. Results Species richness was highest within the tropical north of Australia, and most species were associated with tropical savanna woodlands. Two areas were identified as centres of endemism and these corresponded closely to areas of high species richness. Four morphological groups were identified. One species (Sauropus trachyspermus) was found to be widespread, however all other species had small geographical ranges. Species richness and range size were significantly correlated with changes in latitude. Ten species were found to be of the rarest type, warranting conservation initiatives. Main conclusions Two regions of high richness and endemism of Sauropus occur, Thailand and Australia. Within Australia, the Kakadu‐Alligator River and the Cairns‐Townsville areas were identified as centres of endemism and high species richness for Sauropus. Australian Sauropus in general occur in similar communities and climates as other members of the genus elsewhere. Ten of the 27 species of Australian endemic Sauropus are extremely rare and warrant conservation initiatives. Correlations of latitude to species richness are potentially due to Sauropus radiating from the climatically stable top end of Australia. Increasing range size in more southern latitudes may also be due to stability of climates in the top end or because there is more available land area at these latitudes. Sauropus micranthus, the only non‐endemic species, is probably a more recent invader from the Tertiary period when tropical rain forests where more extensive and congruent with those of New Guinea.