Global patterns of diversity among the specialist birds of riverine landscapes

1. Despite the growing view that biodiversity provides a unifying theme in river ecology, global perspectives on richness in riverine landscapes are limited. As a result, there is little theory or quantitative data on features that might have influenced global patterns in riverine richness, nor are there clear indications of which riverine landscapes are important to conservation at the global scale. As conspicuous elements of the vertebrate fauna of riverine landscapes, we mapped the global distributions of all of the world's specialist riverine birds and assessed their richness in relation to latitude, altitude, primary productivity and geomorphological complexity (surface configuration). 2. Specialist riverine birds, typical of high‐energy riverine landscapes and dependent wholly or partly on production from river ecosystems, occur in 16 families. They are represented by an estimated 60 species divided equally between the passerines and non‐passerines. Major radiation has occurred among different families on different continents, indicating that birds have evolved several times into the niches provided by riverine landscapes. 3. Continental richness varies from four species in Europe to 28 in Asia, with richness on the latter continent disproportionately larger than would be expected from a random distribution with respect to land area. Richness is greatest in mountainous regions at latitudes of 20–40°N in the riverine landscapes of the Himalayan mountains, where 13 species overlap in range. 4. Family, genus and species richness in specialist riverine birds all increase significantly with productivity and surface configuration (i.e. relief). However, family richness was the best single predictor of the numbers of species or genera. In keeping with the effect of surface configuration, river‐bird richness peaks globally at 1300–1400 m altitude, and most species occur typically on small, fast rivers where they feed predominantly on invertebrates. Increased lengths of such streams in areas of high relief and rainfall might have been responsible for species–area effects. 5. We propose the hypothesis that the diversity in channel forms and habitats in riverine landscapes, in addition to high temperature and primary productivity, have been prerequisites to the development of global patterns in the richness of specialist riverine organisms. We advocate tests of this hypothesis in other taxonomic groups. We draw attention, however, to the challenges of categorically defining riverine organisms in such tests because (i) rivers grade into many other habitat types across several different ecotones and (ii) `terrestrialisation' processes in riverine landscapes means that they offer habitat for organisms whose evolutionary origins are not exclusively riverine.     

Cichlid species‐area relationships are shaped by adaptive radiations that scale with area

A positive relationship between species richness and island size is thought to emerge from an equilibrium between immigration and extinction rates, but the influence of species diversification on the form of this relationship is poorly understood. Here, we show that within‐lake adaptive radiation strongly modifies the species‐area relationship for African cichlid fishes. The total number of species derived from in situ speciation increases with lake size, resulting in faunas orders of magnitude higher in species richness than faunas assembled by immigration alone. Multivariate models provide evidence for added influence of lake depth on the species‐area relationship. Diversity of clades representing within‐lake radiations show responses to lake area, depth and energy consistent with limitation by these factors, suggesting that ecological factors influence the species richness of radiating clades within these ecosystems. Together, these processes produce lake fish faunas with highly variable composition, but with diversities that are well predicted by environmental variables.     

The maintenance of a positive spatial correlation between South African bird species richness and human population density

Aim To investigate explanations for the maintenance of a positive spatial species richness–human population density correlation at broad scales, despite the negative impact of humans on species richness. These are (hypotheses 1–4): (1) human activities that create a habitat mosaic and (2) a more favourable climate, and (3) adequate conservation measures (e.g. sufficient natural habitat), maintain the positive species richness–human density correlation; or (4) the full range of human densities decrease the slope of the correlation without changing its form. Location South Africa. Methods Avian species richness data from atlas distribution maps and human population density data derived from 2001 census results were converted to a quarter‐degree resolution. We investigated the number of land transformation types (anthropogenic habitat heterogeneity), irrigated area (increasing productivity), and other covarying factors (e.g. primary productivity) as predictors of species richness. We compared species richness–human density relationships among regions with different amounts of natural habitat, and investigated whether the full range of human densities decrease species richness in relation to primary productivity. Results Hypotheses 1, 2 and 3 were supported. Human densities and activities that increase habitat heterogeneity and productivity are important beneficial factors to common species, but not to rare species. The species richness–human density relationship persists only at low land transformation levels, and no significant relationship exists at higher levels. For common species, the relationship becomes non‐significant at lower land transformation levels than for rare species. Main conclusions The persistence of the species richness–human density relationship depends mostly on the amount of remaining natural habitat. In addition, certain human activities benefit especially common species. Common species seem to be more flexible than rare species in response to human activity and habitat loss.     

Density‐dependent effects of non‐native brown trout Salmo trutta on the species–area relationship in stream fish assemblages

The spatial scale and density‐dependent effects of non‐native brown trout Salmo trutta on species richness of fish assemblages were examined at 48 study sites in Mamachi Stream, a tributary of Chitose River, Hokkaido, Japan. The density of age ≥1 year S. trutta was high in the upstream side of the main stem of Mamachi Stream. Fish species richness increased with increasing area of study sites (habitat size), but the increasing magnitude of the species richness with area decreased with increasing age of ≥1 year S. trutta density. The relationships between age ≥1 year S. trutta, however, and presence–absence of each species seemed to be different among species. Species richness was also determined by location and physical environmental variables, i.e. it was high on the downstream side and in structurally complex environments.     

Speciation-rate dependence in species–area relationships

The general tendency for species number (S) to increase with sampled area (A) constitutes one of the most robust empirical laws of ecology, quantified by species–area relationships (SAR). In many ecosystems, SAR curves display a power-law dependence, S∝A^z. The exponent z is always less than one but shows significant variation in different ecosystems. We study the multitype voter model as one of the simplest models able to reproduce SAR similar to those observed in real ecosystems in terms of basic ecological processes such as birth, dispersal and speciation. Within the model, the species–area exponent z depends on the dimensionless speciation rate ν, even though the detailed dependence is still matter of controversy. We present extensive numerical simulations in a broad range of speciation rates from ν=10^-3 down to ν=10^-11, where the model reproduces values of the exponent observed in nature. In particular, we show that the inverse of the species–area exponent linearly depends on the logarithm of ν. Further, we compare the model outcomes with field data collected from previous studies, for which we separate the effect of the speciation rate from that of the different species lifespans. We find a good linear relationship between inverse exponents and logarithm of species lifespans. However, the slope sets bounds on the speciation rates that can hardly be justified on evolutionary basis, suggesting that additional effects should be taken into account to consistently interpret the observed exponents.     

Richness and composition of plants and birds on land‐bridge islands: effects of island attributes and differential responses of species groups

Aim To test relationships between the richness and composition of vascular plants and birds and attributes of habitat fragments using a model land‐bridge island system, and to investigate whether the effects of fragmentation differ depending on species natural history traits. Location Thousand Island Lake, China. Methods We compiled presence/absence data of vascular plant and bird species through exhaustive surveys of 41 islands. Plant species were assigned to two categories: shade‐intolerant and shade‐tolerant species; bird species were assigned to three categories: edge, interior, and generalist species. We analysed the relationships between island attributes (area, isolation, elevation, shape complexity, and perimeter to area ratio) and species richness using generalized linear models (GLMs). We also investigated patterns of composition in relation to island attributes using ordination (redundancy analysis). Results We found that island area explained a high degree of variation in the species richness of all species groups. The slope of the species–area relationship (z) was 0.16 for all plant species and 0.11 for all bird species. The lowest z‐value was for generalist birds (0.04). The species richness of the three plant species groups was associated with island area per se, while that of all, generalist, and interior birds was explained mainly by elevation, and that of edge bird species was associated primarily with island shape. Patterns of species composition were most strongly related to elevation, island shape complexity, and perimeter to area ratio rather than to island area per se. Species richness had no significant relationship with isolation, but species composition did. We also found differential responses among the species groups to changes in island attributes. Main conclusions Within the Thousand Island Lake system, the effects of fragmentation on both bird and plant species appear to be scale‐dependent and taxon‐specific. The number of plant species occurring on an island is strongly correlated with island area, and the richness of birds and the species composition of plants and birds are associated with variables related to habitat heterogeneity. We conclude that the effects of fragmentation on species diversity and composition depend not only on the degree of habitat loss but also on the specific patterns of habitat fragmentation.     

Contrasting alien and native plant species–area relationships: the importance of spatial grain and extent

Aim The proportion of alien plant species in floras is increasingly being used to indicate the threat of invasions to native species and/or the homogenization of biodiversity. However, this indicator is only valuable if it is independent of the spatial extent and grain of observation. This study tested the equivalence of native and alien species–area relationships (SARs) in order to assess the support for scale invariance in the proportion of alien species in floras. Location England, UK. Methods Nested SARs were generated by assessing the richness of native and alien plant species drawn from the New atlas of the British and Irish flora for six areas comprising 100, 400, 900, 1600, 2500 and 3600 km² with each larger area containing all smaller areas. Five replicate sets of nested areas encompassing northern, southern, eastern, western and central regions were chosen. For each set of nested areas, the log‐transformed species richness was regressed on log‐transformed area to fit a power function to the SAR. Results Native and alien plant SARs reveal consistent differences in slope, highlighting that the proportion of alien species is a function of spatial grain. Aliens are more rare than natives and have higher spatial turnover leading to faster accumulation of species as area increases. However, equivalent samples drawn from a larger spatial extent reveal similar alien and native SARs. Main conclusions The significant differential scale dependence in native and alien species richness observed in this study reflects dissimilar influences of regional drivers such as habitat, but potentially also propagule pressure and introduction history, that leads to the relative rarity and high spatial turnover of alien species. Maps of invasion hotspots that identify areas where the proportion of the alien flora is particularly high should therefore be treated with considerable caution since patterns across most grains used for species monitoring will be scale dependent.     

Species diversity and endemism of five major Malesian islands: diversity–area relationships

Aim A comparison of biodiversity patterns within Malesia in relation to surface area. Location Analysis of the patterns in species richness and endemism of vascular plants in the five major Malesian islands, i.e. Java, Sulawesi, Sumatra, Borneo and New Guinea. Methods Available data on species richness and species ranges in correlation with the surface area of the respective islands were examined in this work. Estimations of total species numbers for these islands are presented based on extrapolation of all available published Flora Malesiana information and recent checklists, all in all comprising 12,000 different species. The regression analysis of overall species richness and endemism were studied for all species together as well as for different plant families to compare the fit with the Arrhenius species–area model. Results The five islands form a series of independent areas of increasing size suited for an analysis of the species–area relationships at the regional scale. All species taken together and those of families with even distribution throughout Malesia show significant species–area relationships. Non‐significant relationships were detected in families with western or eastern‐centred Malesian distribution patterns. Relationships between number of endemic species and surface area are significant for all species and for the majority of the families with significant species–area relationships. Main conclusions Species–area relationships of families appear to be dependent on species number. Families with high numbers of species usually have a significant species–area relationship whereas small families have not. For the families that display an eastern or western Malesian centred pattern, a historical biogeographical explanation should be invoked. Island surface area appears to be a predictor for island percent endemism in Malesian vascular plants. None of the islands appears to be a hotspot of endemism nor of species diversity, as no significant departure from the Arrhenius model was noted for any of them.     

Testing species–stone area and species–bryophyte cover relationships in riverine macroinvertebrates at small scales

1. The species–area relationship is considered amongst the few genuine laws in ecology. Although positive species richness–stone area relationships have been found previously in stream systems, very few studies have simultaneously examined species–individuals, individuals–area, species–bryophyte biomass and individuals–bryophyte biomass relationships. We examined these relationships based on temporally replicated assessments of macroinvertebrates on stones at two river sites. 2. We found only one significant species–area relationship out of six relationship tested, and two significant individuals–area relationships. Even these significant relationships were weak, however. By contrast, we detected significant and rather strong relationships between species richness and the number of individuals at both river sites on all three sampling dates. We also found significant relationships of both species richness and the number of individuals with bryophyte biomass at both river sites on all sampling occasions. One of the river sites was disturbed by a bulldozer, and the species–bryophyte biomass relationships were somewhat stronger after the disturbance event. 3. Our findings are quite surprising, given that there were very weak species–area relationships on stream stones. By contrast, our results suggest a pivotal role for bryophyte biomass in determining the species richness and the number of individuals of stream macroinvertebrates at this small scale. The most probably origin of these relationships begins with bryophyte cover, which determines the number of individuals, and subsequently passively affects species richness. Thus, there is not necessarily a direct mechanism that determines the variability of species richness on stream stones. 4. Experimental studies are needed to disentangle the various mechanisms (e.g. passive sampling, provision of more food, more niche space, flood disturbance refugia) by which bryophyte biomass affects stream macroinvertebrates.     

Plants on small islands revisited: the effects of spatial scale and habitat quality on the species–area relationship

Understanding how species diversity is related to sampling area and spatial scale is central to ecology and biogeography. Small islands and small sampling units support fewer species than larger ones. However, the factors influencing species richness may not be consistent across scales. Richness at local scales is primarily affected by small‐scale environmental factors, stochasticity and the richness at the island scale. Richness at whole‐island scale, however, is usually strongly related to island area, isolation and habitat diversity. Despite these contrasting drivers at local and island scales, island species–area relationships (SARs) are often constructed based on richness sampled at the local scale. Whether local scale samples adequately predict richness at the island scale and how local scale samples influence the island SAR remains poorly understood. We investigated the effects of different sampling scales on the SAR of trees on 60 small islands in the Raja Ampat archipelago (Indonesia) using standardised transects and a hierarchically nested sampling design. We compared species richness at different grain sizes ranging from single (sub)transects to whole islands and tested whether the shape of the SAR changed with sampling scale. We then determined the importance of island area, isolation, shape and habitat quality at each scale on species richness. We found strong support for scale dependency of the SAR. The SAR changed from exponential shape at local sampling scales to sigmoidal shape at the island scale indicating variation of species richness independent of area for small islands and hence the presence of a small‐island effect. Island area was the most important variable explaining species richness at all scales, but habitat quality was also important at local scales. We conclude that the SAR and drivers of species richness are influenced by sampling scale, and that the sampling design for assessing the island SARs therefore requires careful consideration.     

Species‐time‐area and phylogenetic‐time‐area relationships in tropical tree communities

The species‐area relationship (SAR) has proven to be one of the few strong generalities in ecology. The temporal analog of the SAR, the species‐time relationship (STR), has received considerably less attention. Recent work primarily from the temperate zone has aimed to merge the SAR and the STR into a synthetic and unified species‐time‐area relationship (STAR) as originally envisioned by Preston (1960). Here we test this framework using two tropical tree communities and extend it by deriving a phylogenetic‐time‐area relationship (PTAR). The work finds some support for Preston's prediction that diversity‐time relationships, both species and phylogenetic, are sensitive to the spatial scale of the sampling. Contrary to the Preston's predictions we find a decoupling of diversity‐area and diversity‐time relationships in both forests as the time period used to quantify the diversity‐area relationship changes. In particular, diversity‐area and diversity‐time relationships are positively correlated using the initial census to quantify the diversity‐area relationship, but weakly or even negatively correlated when using the most recent census. Thus, diversity‐area relationships could forecast the temporal accumulation of biodiversity of the forests, but they failed to “back‐cast” the temporal accumulation of biodiversity suggesting a decoupling of space and time.