Additive diversity partitioning as a guide to regional montane reserve design in Asia: an example from Yunnan Province,China

Aim Data on spatial and temporal turnover in species composition within a region is essential to design regional protected areas. Montane systems are often recognized as biodiversity hotspots. The primary objective of this study is to identify patterns of montane bird diversity across multiple spatial and temporal scales using an additive diversity partitioning framework. Location The Ailao Mountains, central Yunnan Province, China. Methods We used point counts to sample bird communities in four elevational zones, on eastern and western slopes, during both the breeding and the non‐breeding seasons. Diversity (richness and Shannon) was partitioned across space (points, elevational zones and slopes) and time (seasons). We used permutation tests to compare observed values to values expected by random chance. A complementary cluster analysis was also used to evaluate beta diversity. Results Overall, the gamma diversity was attributed to significantly higher beta diversity (relative to that of randomization tests) among elevational zones and, to a lesser extent, between slopes. For Shannon–Wiener Index, beta diversity between seasons was significantly higher than expected and had a similar contribution to the gamma diversity as with beta diversity between slopes. Hierarchical cluster analysis supported the findings for Shannon–Wiener Index. The contribution of beta diversity among points to gamma diversity within each elevational zone generally lessened with increasing elevation. Main conclusions Our results show significantly high levels of beta diversity among elevational zones and between slopes, as well as between seasons for Shannon diversity, in a small area of the Ailao Mountain range. Thus, a regional montane reserve system should cover the entire elevational gradient and multiple slopes, rather than only the montane crest. Furthermore, higher pattern diversity in lower elevational zones suggests that larger areas should be preserved at lower elevational zones. Finally, the design of regional reserve systems require more studies conducted at multiple seasons at a regional scale.     

Animal species diversity driven by habitat heterogeneity/diversity: the importance of keystone structures

Aim In a selected literature survey we reviewed studies on the habitat heterogeneity–animal species diversity relationship and evaluated whether there are uncertainties and biases in its empirical support. Location World-wide. Methods We reviewed 85 publications for the period 1960–2003. We screened each publication for terms that were used to define habitat heterogeneity, the animal species group and ecosystem studied, the definition of the structural variable, the measurement of vegetation structure and the temporal and spatial scale of the study. Main conclusions The majority of studies found a positive correlation between habitat heterogeneity/diversity and animal species diversity. However, empirical support for this relationship is drastically biased towards studies of vertebrates and habitats under anthropogenic influence. In this paper, we show that ecological effects of habitat heterogeneity may vary considerably between species groups depending on whether structural attributes are perceived as heterogeneity or fragmentation. Possible effects may also vary relative to the structural variable measured. Based upon this, we introduce a classification framework that may be used for across-studies comparisons. Moreover, the effect of habitat heterogeneity for one species group may differ in relation to the spatial scale. In several studies, however, different species groups are closely linked to ‘keystone structures’ that determine animal species diversity by their presence. Detecting crucial keystone structures of the vegetation has profound implications for nature conservation and biodiversity management.     

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.     

Distinguishing area and habitat heterogeneity effects: a simulation test of the MacNally and Watson (1997) protocol

Distinguishing the roles that different factors, such as sampling effects and habitat heterogeneity, play in generating species‐area curves continues to be difficult in many communities. A recent response to this challenge is the proposal of a ‘zoom’ protocol in which species richness and habitat heterogeneity are sampled in successively larger units (transects or quadrats). The utility of this approach requires that there be justifiable, predictable and unambiguous relationships between richness and heterogeneity. Results of computer simulations that I have done to test the predicted relationships demonstrate, however, that the predicted patterns were not always observed and, on occasion, more complex relationships were observed in their place. While the development of such protocols may increase our understanding of species‐area curves, they are unlikely ever to pronounce unambiguously on their causes.     

Bird species richness patterns of northern Taiwan: primary productivity, human population density, and habitat heterogeneity

Energy, climate, habitat heterogeneity, and human activity are important correlates of spatial variation in species richness. We examined the correlation between species richness and these variables using the birds that breed in northern Taiwan. We conducted general linear models (GLMs) and spatial correlation models to examine the relationship between bird species richness (BSR) and environmental variables. We found that normalized difference vegetation index (NDVI) was the most important predictor of BSR. We suggest productivity is the primary process of BSR. Additionally, we hypothesized that scale dependency might exist in the relationship between BSR and NDVI in Taiwan. Human population density, the second most important factor, was inversely correlated with BSR. The factor and BSR did not have similar response to NDVI, which contradicted observations in most of the previous studies on human population vs. species richness. We proposed that the human population density had an effect on NDVI, which in turn had an effect on BSR. Moreover, we hypothesized that the contradiction between our study and the previous studies might arise from a higher level of human disturbance in Taiwan than in other areas. The necessity of conserving native species in intensively developed lowlands of Taiwan cannot be overemphasized. Number of land cover type was another significant predictor of BSR. Habitat heterogeneity may have an effect on BSR in Taiwan.     

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 richness and the proportion of predatory animal species in temporary freshwater pools: relationships with habitat size and permanence

The proportion of predatory animal species is often believed not to vary systematically across communities. However, we predict that larger temporary freshwater pools, and pools that are more permanent, will contain a higher proportion of predatory animal species. In 24 temporary rockpools in Northern Israel (supporting communities dominated by ostracods, copepods, cladocerans, flatworms, dipterans and amphibians), the mean proportion of macroscopic predatory species (averaged over a series of samples) increased with increasing pool area. For the highest possible proportion of predatory species (including microscopic species with uncertain diets), the relationship with pool area was not statistically significant. We did not find significant relationships between permanence and the proportion of either macroscopic or all possible predatory species. Larger pools and pools that were more permanent had more species. Species richness and the proportion of macroscopic predators were positively correlated. These patterns imply that species-poor ecosystems are likely to be functionally different from species-rich systems.     

Controls on pathogen species richness in plants' introduced and native ranges: roles of residence time, range size and host traits

Introduced species escape many pathogens and other enemies, raising three questions. How quickly do introduced hosts accumulate pathogen species? What factors control pathogen species richness? Are these factors the same in the hosts' native and introduced ranges? We analysed fungal and viral pathogen species richness on 124 plant species in both their native European range and introduced North American range. Hosts introduced 400 years ago supported six times more pathogens than those introduced 40 years ago. In hosts' native range, pathogen richness was greater on hosts occurring in more habitat types, with a history of agricultural use and adapted to greater resource supplies. In hosts' introduced range, pathogen richness was correlated with host geographic range size, agricultural use and time since introduction, but not any measured biological traits. Introduced species have accumulated pathogens at rates that are slow relative to most ecological processes, and contingent on geographic and historic circumstance.