Invasion of North American drainages by alien fish species

1. Data from the literature were used to document colonization patterns by introduced freshwater fishes in 125 drainages across temperate North America. We analysed this data set to quantify susceptibility to invasion, success of the invaders and changes in species richness.
2. Drainages with a high number of impoundments, large basin area and low native species diversity had the greatest number of introduced species. Those drainages containing few native fishes exhibited great variation in the number of invaders, while waters with a rich native fauna contained few introduced species. However, this pattern did not differ significantly from random simulations because the pool of potential invaders is greater for drainages with low species richness.
3. In most drainages, there were more introduced than imperilled or extirpated species, suggesting that invaders tend to increase overall species richness.
4. These patterns suggest that North American fish communities are not saturated with species, but instead, are capable of supporting higher levels of diversity if the pool of potential colonists and the rate of colonization from that pool is increased.     

A comparison of methods for mapping species ranges and species richness

Aim  Maps of species richness are the basis for applied research and conservation planning as well as for theoretical research investigating patterns of richness and the processes shaping these patterns. The method used to create a richness map could influence the results of such studies, but differences between these methods have been insufficiently evaluated. We investigate how different methods of mapping species ranges can influence patterns of richness, at three spatial resolutions.
Location  California, USA.
Methods  We created richness maps by overlaying individual species range maps for terrestrial amphibians and reptiles. The methods we used to create ranges included: point-to-grid maps, obtained by overlaying point observations of species occurrences with a grid and determining presence or absence for each cell; expert-drawn maps; and maps obtained through species distribution modelling. We also used a hybrid method that incorporated data from all three methods. We assessed the correlation and similarity of the spatial patterns of richness maps created with each of these four methods at three different resolutions.
Results  Richness maps created with different methods were more correlated at lower spatial resolutions than at higher resolutions. At all resolutions, point-to-grid richness maps estimated the lowest species richness and those derived from species distribution models the highest. Expert-drawn maps and hybrid maps showed intermediate levels of richness but had different spatial patterns of species richness from those derived with the other methods.
Main conclusions  Even in relatively well-studied areas such as California, different data sources can lead to rather dissimilar maps of species richness. Evaluating the strengths and weaknesses of different methods for creating a richness map can provide guidance for selecting the approach that is most appropriate for a given application and region.     

Elevational species richness patterns for vascular plants on Mount Kinabalu, Borneo

Aim  We quantify the elevational patterns of species richness for all vascular plants and some functional and taxonomic groups on a regional scale on a tropical mountain and discuss some possible causes for the observed patterns.
Location  Mount Kinabalu, Sabah, Borneo.
Methods  A data base containing elevational information on more than 28,000 specimens was analysed for vascular plant distribution, taking into account sampling effort. The total species richness pattern was estimated per 300-m elevational interval by rarefaction analyses. The same methods were also applied to quantify species richness patterns of trees, epiphytes, and ferns.
Results  Total species richness has a humped relationship with elevation, and a maximum species richness in the interval between 900 and 1200 m. For ferns and epiphytes the maximum species richness is found at slightly higher elevations, whereas tree species did not have a statistically significant peak in richness above the lowest interval analysed.
Main conclusions  For the first time a rigorous estimate of an elevational pattern in species richness of the whole vascular plant flora of a tropical mountain has been quantified. The pattern observed depends on the group studied. We discuss the differences between the groups and compare the results with previous studies of elevational patterns of species richness from other tropical areas. We also discuss the methods used to quantify the richness pattern and conclude that rarefaction gives an appropriate estimate of the species richness pattern.     

Climate-based models of spatial patterns of species richness in Egypt's butterfly and mammal fauna

Aim  Identifying areas of high species richness is an important goal of conservation biogeography. In this study we compared alternative methods for generating climate-based estimates of spatial patterns of butterfly and mammal species richness.
Location  Egypt.
Methods  Data on the occurrence of butterflies and mammals in Egypt were taken from an electronic database compiled from museum records and the literature. Using M axent , species distribution models were built with these data and with variables describing climate and habitat. Species richness predictions were made by summing distribution models for individual species and by modelling observed species richness directly using the same environmental variables.
Results  Estimates of species richness from both methods correlated positively with each other and with observed species richness. Protected areas had higher species richness (both predicted and actual) than unprotected areas.
Main conclusions  Our results suggest that climate-based models of species richness could provide a rapid method for selecting potential areas for protection and thus have important implications for biodiversity conservation.     

Distribution of rotifer assemblages in North Island, New Zealand, lakes: relationships to environmental and historical factors

1. The geographical distribution of planktonic rotifer species was investigated in 31 lakes in the North Island of New Zealand.
2. A total of 78 species was recorded. Species richness, previously thought to be low in New Zealand, was found to be comparable with that of northern temperate lakes with an average of 21.1 species found per lake. The large, deep oligotrophic Lake Taupo had the lowest richness, and the artificial reservoirs the highest.
3. The distribution of species, investigated using multivariate techniques [Cluster analysis, canonical correspondence analysis (CCA)], was most strongly associated with trophic state gradients. For example, Conochilus unicornis, C. dossuarius and Ascomorpha ovalis were associated with more oligotrophic conditions, and Brachionus budapestinensis, B. calyciflorus and Keratella tropica with more eutrophic conditions. Inorganic turbidity was also of importance in determining rotifer distributions in some shallower lakes.
4. Some species, for example K. australis and C. exiguus , appear to be limited in distribution by poor dispersal abilities.     

Climate and history explain the species richness peak at mid-elevation for Schizothorax fishes (Cypriniformes: Cyprinidae) distributed in the Tibetan Plateau and its adjacent regions

Aim  We studied elevational species richness patterns of Schizothorax fishes and identified the roles of ecological and evolutionary factors in shaping the patterns of elevational diversity.
Location  The Tibetan Plateau and its adjacent regions.
Methods  We assembled distribution and altitude data for all Schizothorax species using the literature. We merged ecological and evolutionary approaches to test the relationships between species richness and ecological factors (climate, area, the mid-domain effect) or evolutionary factors (diversification rates and time of colonization).
Results  We found that species richness of Schizothorax fishes peaked at mid-elevations. Rainfall, area, the mid-domain effect and diversification rate were weak predictors of the richness pattern. Temperature showed a nonlinear relationship with species richness. Temperature and time of colonization were the most important variables in explaining the elevational diversity pattern.
Main conclusion  Our findings indicate that the time-for-speciation effect and niche conservatism play important roles in variation of species richness.     

Amphibian species richness and distribution in relation to pond water chemistry in south-western Ontario, Canada

1. We assessed the patterns of amphibian species richness and distribution in relation to water chemistry over a large geographical area in 1992–94.
2. Thirteen amphibian species were observed at 180 ponds, with mean species richness 3.5 ± 0.13 species per pond (range zero to nine). Water samples were collected from 143 ponds, analysed for fifteen chemical variables, and further analysed by multivariate statistical techniques.
3. Water in the study area was hard, alkaline and well-buffered against pH change, and most ponds were eutrophic. Amphibian species richness was negatively correlated with five chemical variables (chloride, conductivity, magnesium, total hardness, turbidity).
4. Principle components analysis reduced the data set to four chemical components that explained 65.4% of the variance in the original variables. Principle component scores were retained for use in further multivariate tests. Multiple regression accounted for only 19.0% of the variance in amphibian species richness. Discriminant function analysis (DFA) was used to determine if water chemistry variables discriminated among species, but it was only able to classify 17.5% of cases correctly. DFA was also used to determine if water chemistry distinguished between used and unused sites for individual species. DFA was moderately successful, classifying 61–77% of cases correctly.
5. General water chemistry appears to play only a minor part in affecting amphibian species richness in south-western, Ontario. However, chemical variables may be helpful to distinguish between used and unused sites for some species.     

Amphibian species richness and distribution in relation to pond water chemistry in south-western Ontario, Canada

1. We assessed the patterns of amphibian species richness and distribution in relation to water chemistry over a large geographical area in 1992–94.
2. Thirteen amphibian species were observed at 180 ponds, with mean species richness 3.5 ± 0.13 species per pond (range zero to nine). Water samples were collected from 143 ponds, analysed for fifteen chemical variables, and further analysed by multivariate statistical techniques.
3. Water in the study area was hard, alkaline and well-buffered against pH change, and most ponds were eutrophic. Amphibian species richness was negatively correlated with five chemical variables (chloride, conductivity, magnesium, total hardness, turbidity).
4. Principle components analysis reduced the data set to four chemical components that explained 65.4% of the variance in the original variables. Principle component scores were retained for use in further multivariate tests. Multiple regression accounted for only 19.0% of the variance in amphibian species richness. Discriminant function analysis (DFA) was used to determine if water chemistry variables discriminated among species, but it was only able to classify 17.5% of cases correctly. DFA was also used to determine if water chemistry distinguished between used and unused sites for individual species. DFA was moderately successful, classifying 61–77% of cases correctly.
5. General water chemistry appears to play only a minor part in affecting amphibian species richness in south-western, Ontario. However, chemical variables may be helpful to distinguish between used and unused sites for some species.     

Diversity patterns of stygobiotic crustaceans across multiple spatial scales in Europe

1. Using species distribution data from 111 aquifers distributed in nine European regions, we examined the pairwise relationships between local species richness (LSR), dissimilarity in species composition among localities, and regional species richness (RSR). In addition, we quantified the relative contribution of three nested spatial units – aquifers, catchments and regions – to the overall richness of groundwater crustaceans.
2. The average number of species in karst and porous aquifers (LSR) varied significantly among regions and was dependent upon the richness of the regional species pool (RSR). LSR–RSR relationships differed between habitats: species richness in karstic local communities increased linearly with richness of the surrounding region, whereas that of porous local communities levelled off beyond a certain value of RSR.
3. Dissimilarity in species composition among aquifers of a region increased significantly with increasing regional richness because of stronger habitat specialisation and a decrease in the geographic range of species among karst aquifers. Species turnover among karst aquifers was positively related to RSR, whereas this relationship was not significant for porous aquifers.
4. The contribution of a given spatial unit to total richness increased as size of the spatial unit increased, although 72% of the overall richness was attributed to among-region diversity. Differences in community composition between similar habitats in different regions were typically more pronounced than between nearby communities from different habitats.
5. We conclude by calling for biodiversity assessment methods and conservation strategies that explicitly integrate the importance of turnover in community composition and habitat dissimilarity at multiple spatial scales.     

The impact of changing the season in which cereals are sown on the diversity of the weed flora in rotational fields in Denmark

1. Surveys have shown that there has been a dramatic decrease in the weed flora of fields under rotational cultivation during the last 30 years. This trend has been particularly noticeable in winter cereals, a crop of increasing importance in the landscape.
2. The weed flora of spring and winter cereals was compared in 19 unsprayed fields during a 5-year study to test the hypothesis that cereal type exerts no effect on the flora or on the absolute and relative abundance of single species.
3. Plant and species densities, and accumulated species richness, were lower in winter than in spring cereals.
4. Floristic similarity was greater among spring cereal fields and between spring and winter cereals within the same fields than among winter cereal fields.
5. Species that occurred with unequal density in spring and winter cereals occurred at higher densities in the spring cereals; these species germinated mainly in the spring. However, for a few species the relative plant abundance was highest in winter cereals; these species were able to germinate both in the spring and autumn.
6. Some species – on the relative scale – occurred indifferently of season of sowing; all but one of these species were able to germinate both in the spring and autumn.
7. Plant species and taxa that are important food resources for arthropod herbivores occurred at greater densities in spring than in winter cereals and, in addition, occurred with the highest relative abundance in spring cereals.
8. Change in land use from spring to winter cereals involves not only an immediate reduction of more than 25% in the density of plants and species, but also a change and increased uncertainty in the composition of the weed flora. These findings may have serious implications for the ecology of wildlife in the agricultural landscape.     

Are distribution patterns linked to dispersal mechanism? An investigation using pond invertebrate assemblages

1. Dispersal ability influences the distribution and abundance of organisms, but empirical investigations of the relationship between dispersal ability and the composition of ecological assemblages are scarce. Here, we compare between-site variation in the species richness and community composition of actively and passively dispersing pond invertebrates.
2. Coleoptera (active dispersers) and microcrustacea (passive dispersers) were sampled over a season from 16 ponds within a 4-km radius in south-west England. Species richness and community composition were related to environmental variables using regression and Canonical Correspondence Analysis (CCA), respectively.
3. Coleopteran species richness was significantly and positively correlated with pond permanence and maximum area, whereas microcrustacean species richness was relatively equal across sites and did not correlate with environmental variables. The frequency of species' occurrence between sites was the same for both groups, which suggests that active and passive dispersers exhibited the same degree of dispersal.
4. Between-site variation in community composition was non-random for both groups, with pond permanence and area, together, explaining similar proportions of between-site variation for Coleoptera. Permanence was correlated most strongly with microcrustacean community composition and a high proportion (25%) of microcrustacean species were more numerous in smaller, more ephemeral ponds.
5. These data suggest that, at small spatial scales, Coleoptera which can undertake multiple dispersal events, are more likely to colonise large, more permanent ponds than passively dispersing microcrustacea. For microcrustacea, other traits (in this case those permitting existence in ephemeral habitats) may over-ride the influences of dispersal in driving between-site variation in species composition.     

Can the tropical conservatism hypothesis explain temperate species richness patterns? An inverse latitudinal biodiversity gradient in the New World snake tribe Lampropeltini

Aim  A latitudinal gradient in species richness, defined as a decrease in biodiversity away from the equator, is one of the oldest known patterns in ecology and evolutionary biology. However, there are also many known cases of increasing poleward diversity, forming inverse latitudinal biodiversity gradients. As only three processes (speciation, extinction and dispersal) can directly affect species richness in areas, similar factors may be responsible for both classical (high tropical diversity) and inverse (high temperate diversity) gradients. Thus, a modified explanation for differential species richness which accounts for both patterns would be preferable to one which only explains high tropical biodiversity.
Location  The New World.
Methods  We test several proposed ecological, temporal, evolutionary and spatial explanations for latitudinal diversity gradients in the New World snake tribe Lampropeltini, which exhibits its highest biodiversity in temperate regions.
Results  We find that an extratropical peak in species richness is not explained by latitudinal variation in diversification rate, the mid-domain effect, or Rapoport's rule. Rather, earlier colonization and longer duration in the temperate zones allowing more time for speciation to increase biodiversity, phylogenetic niche conservatism limiting tropical dispersal and the expansion of the temperate zones in the Tertiary better explain inverse diversity gradients in this group.
Main conclusions  Our conclusions are the inverse of the predictions made by the tropical conservatism hypothesis to explain higher biodiversity near the equator. Therefore, we suggest that the processes invoked are not intrinsic to the tropics but are dependent on historical biogeography to determine the distribution of species richness, which we refer to as the 'biogeographical conservatism hypothesis'.     

Differences in insect species richness and faunal composition of birch seedlings, saplings and trees: the importance of plant architecture

Abstract. 1. Data are presented on the species richness and faunal composition of herbivorous insects on birch seedlings, saplings and trees at one site in Northern England.
2. Species richness of insect herbivores in equal-sized samples from birch seedlings and trees was similar through most of the season.
3. Effects of plant architecture were confined to the first sampling date, when seedling faunas were species poor compared with trees – possibly due to safe overwintering sites on the extensive bark, twigs and buds of trees.
4. The faunal composition of birch seedlings, saplings and trees was also similar. Out of a total of 112 recorded species of herbivores, only one aphid species was confined to seedlings.
5. Similarly, no evidence for clear-cut vertical stratification of insects within trees was found.
6. Species turnover as host plants mature ('horizontal' stratification) and vertical stratification within trees add little to the high overall species richness of birch-feeding insects in Britain, contrary to the predictions of Lawton (1983).     

Using potential distributions to explore determinants of Western Palaearctic migratory songbird species richness in sub-Saharan Africa

Aim  Although the breeding ranges of most Western Palaearctic migratory passerines are well documented in Europe, their overwintering ranges and patterns of species richness in Africa remain poorly understood. To illustrate potential patterns of species richness despite severely limited data, we extrapolated species ranges from a new and unique data bank of locality records that documents overwintering locations of these birds in Africa.
Location  Sub-Saharan Africa.
Methods  We predicted potential geographical distributions of 60 species of passerine birds based on overwintering records using bioclimatic models. We then combined these predictions to estimate potential species richness and explored response shapes using spatial linear regression. We also evaluated the evidence for a mid-domain effect using a one-dimensional null model.
Results  Spatial linear regression analyses of the species richness pattern revealed non-linear relationships to seasonality in precipitation, minimum net primary productivity, minimum average temperature, habitat heterogeneity, percentage of tree cover, distance from the Sahara Desert and inter-annual variability in net primary productivity. The explanatory power of these variables decreased with geographic range size. The one-dimensional null model of species richness based on distance from the Sahara Desert did not show evidence of a mid-domain effect.
Main conclusions  Distributions of migrants seem generally strongly determined by distance from the Sahara Desert working in concert with climatic effects, but this cannot adequately explain richness patterns of species with small ranges in Africa, many of which are of substantial conservation concern.     

The value of georeferenced collection records for predicting patterns of mosquito species richness and endemism in the Neotropics

Abstract.  1. Determining large-scale distribution patterns for mosquitoes could advance knowledge of global mosquito biogeography and inform decisions about where mosquito inventory needs are greatest.
2. Over 43 000 georeferenced records are presented of identified and vouchered mosquitoes from collections undertaken between 1899 and 1982, from 1853 locations in 42 countries throughout the Neotropics. Of 492 species in the data set, 23% were only recorded from one location, and Anopheles albimanus Wiedemann is the most common species.
3. A linear log–log species–area relationship was found for mosquito species number and country area. Chile had the lowest relative density of species and Trinidad-Tobago the highest, followed by Panama and French Guiana.
4. The potential distribution of species was predicted using an Ecological Niche Modelling (ENM) approach. Anopheles species had the largest predicted species ranges, whereas species of Deinocerites and Wyeomyia had the smallest.
5. Species richness was estimated for 1° grids and by summing predicted presence of species from ENM. These methods both showed areas of high species richness in French Guiana, Panama, Trinidad-Tobago, and Colombia. Potential hotspots in endemicity included unsampled areas in Panama, French Guiana, Colombia, Belize, Venezuela, and Brazil.
6. Argentina, The Bahamas, Bermuda, Bolivia, Cuba, and Peru were the most under-represented countries in the database compared with known country species occurrence data. Analysis of species accumulation curves suggested patchiness in the distribution of data points, which may affect estimates of species richness.
7. The data set is a first step towards the development of a global-scale repository of georeferenced mosquito collection records.     

Realized climatic niches of deciduous trees: comparing western Eurasia and eastern North America

Aim  Assessment of cross-continental similarities and differences in climatic limiting values for deciduous tree species and of the possible deterministic influence of past and present climatic differences on the modern tree flora in two regions.
Location  The deciduous forest regions of western Eurasia and eastern North America.
Methods  Based on species distribution data (range maps) and climate site data, the realized climatic niches of 137 deciduous tree species from the two regions were quantified using climatic envelopes. To compare these envelopes on the two continents, a hierarchical cluster analysis was performed, and principal components analysis was used to check cluster consistency.
Results  Significant differences do exist for upper limits of winter temperatures and for lower limits of summer temperatures between Western Eurasia and eastern North America. Lower limits for the annual water balance also appear different, suggesting that the deciduous trees may be more drought-tolerant in western Eurasia than in eastern North America. Climatic range types generated by the cluster analysis can be characterized, according to the distribution of the species, as boreal-temperate, northern temperate, temperate, southern temperate, and Appalachian. Five of the eight clusters contain trees from both regions, but three groups consist only of American species that have no European counterparts.
Main conclusions  Differences in temperature limitations can be explained by location on the east versus west side of the continents and by the almost complete lack of warm moist areas in western Eurasia. The difference in drought tolerance, on the other hand, is more likely to be the product of a deterministic sorting process that occurred during the Plio-Pleistocene.     

Does a ruderal strategy dominate the endemic flora of the West African forests?

Aim  To understand the distribution pattern of endemic plant species in West African rain forests, one of the global priority areas for biodiversity conservation.
Location  Upper Guinean forests, West Africa (Senegal, Guinea-Bissau, Guinea, Sierra Leone, Liberia, Côte d'Ivoire, Ghana and Togo).
Methods  We used herbarium collections from the whole Upper Guinean region ( sensu White 1981) to analyse the distribution patterns of 216 vascular plant species (approximately one-third of the endemic flora ranging from herbs to trees) that are restricted to these rain forests. We related species distribution ranges and species commonness to the main environmental variables and species life-history traits.
Results  We found that most endemic species of West African forests have large distribution ranges and ruderal strategies. Among all plant life-forms, species with small ranges were restricted to very moist places whereas more widespread species were less dependent on the availability of water. These more common species seem able to tolerate drier conditions, indicated by the minimum rainfall conditions where they are found and the positive correlation with local habitat openness. Light-demanding endemics were more common and widespread than shade-tolerant species, and there was a trend towards wind-dispersed species having a higher proportion of records in open places. This suggests that how widespread and common an endemic species is depends on being able to tolerate open and drier conditions.
Main conclusions  A ruderal strategy seems to be key for understanding the success of endemics in the Upper Guinean forests, and indicates the important role of disturbances in shaping the composition of these tropical forests. Our results have large implications for predicting the potential effect of increasing drought on rare endemic plant species of West African forests.     

Does host‐plant diversity explain species richness in insects? A test using Coccidae (Hemiptera)

1. The megadiverse herbivores and their host plants are a major component of biodiversity, and their interactions have been hypothesised to drive the diversification of both. 2. If plant diversity influences the diversity of insects, there is an expectation that insect species richness will be strongly correlated with host‐plant species richness. This should be observable at two levels (i) more diverse host‐plant groups should harbour more species of insects, and (ii) the species richness of a group of insects should correlate with the richness of the host groups it uses. However, such a correlation is also consistent with a hypothesis of random host use, in which insects encounter and use hosts in proportion to the diversity of host plants. Neither of these expectations has been widely tested. 3. These expectations were tested using data from a species‐rich group of insects – the Coccidae (Hemiptera). 4. Significant positive correlations were found between the species richness of coccid clades (genera) and the species richness of the host‐plant family or families upon which the clades occur. On a global scale, more closely related plant families have more similar communities of coccid genera but the correlation is weak. 5. Random host use could not be rejected for many coccids but randomisation tests and similarity of coccid communities on closely related plant families show that there is non‐random host use in some taxa. Overall, our results support the idea that plant diversity is a driver of species richness of herbivorous insects, probably via escape‐and‐radiate or oscillation‐type processes.     

The positive correlation between avian species richness and human population density in Britain is not attributable to sampling bias

Aim  To assess whether spatial variation in sampling effort drives positive correlations between human population density and species richness.
Location  British 10 × 10 km squares.
Methods  We calculated three measures of species richness from atlas data of breeding birds in Britain: total species richness, species richness standardised for sampling effort, and the number of species only recorded in supplementary casual records in a manner not standardised for survey effort. We then assessed the form of the relationship between these richness estimates and human population density, both with and without taking spatial autocorrelation into account.
Results  Both total and standardised species richness exhibit similar species richness–human population density relationships; species richness generally increases with human population density, but decreases at the very highest densities. Supplementary species richness is very weakly correlated with human population density.
Main conclusions  In this example, sampling effort only slightly influences the form of species richness–human population density relationships. The positive correlation between species richness and human population density and any resultant conservation conflicts are thus not artefactual patterns generated by confounding human density and sampling effort.     

Effect of landscape structure on the spatial distribution of Mediterranean dung beetle diversity

Aim  Landscape structure influences the distribution of animals, altering their movements and their ability to reach habitat patches. We analysed the spatial patterns of dung beetle species diversity in three differently structured natural landscapes in a Mediterranean protected area in the centre of the Iberian Peninsula.
Location  Cabañeros National Park, Central Spain.
Methods  Diversity components within (α) and among (β) the three main vegetation types in the reserve were compared by using a hierarchical nested design. These were forests, scrublands and grasslands embedded in three different landscapes, where each was the most dominant and structurally connected habitat.
Results  Species richness of grassland habitat did not vary across landscapes, but forest habitat showed lower species richness in the grassland-dominated landscape. Scrubland was the least species-rich habitat, but here again there was no significant variation across landscapes. However, in all cases, there was a significant influence of habitat context (configuration of habitat patches within landscape matrix) on similarity of species composition. These tended to be more similar to the dominant landscape matrix where they were embedded, rather than to the same habitat type in other landscapes. Additive partitioning of diversity showed higher than expected values of β in all landscapes, which indicated a structured response. Highest values of β in the grassland-dominated landscape suggest that this was the least connected landscape for dung beetles.
Main conclusions  Our results suggest that in homogeneous conditions of climate and trophic resources, landscape structure may well be more important than habitat type as a determinant of dung beetle distribution in the Mediterranean.