Re‐approaching the small island effect

Aim To propose a new approach to the small island effect (SIE) and a simple mathematical procedure for the estimation of its upper limit. The main feature of the SIE is that below an upper size threshold an increase of species number with increase of area in small islands is not observed. Location Species richness patterns from different taxa and insular systems are analysed. Methods Sixteen different data sets from 12 studies are analysed. Path analysis was used for the estimation of the upper limit of the SIE. We studied each data set in order to detect whether there was a certain island size under which the direct effects of area were eliminated. This detection was carried out through the sequential exclusion of islands from the largest to the smallest. For the cases where an SIE was detected, a log‐log plot of species number against area is presented. The relationships between habitat diversity, species number and area are studied within the limits of the SIE. In previous studies only area was used for the detection of the SIE, whereas we also encompass habitat diversity, a parameter with well documented influence on species richness, especially at small scales. Results An SIE was detected in six out of the 16 studied cases. The upper limit of the SIE varies, depending on the characteristics of the taxon and the archipelago under study. In general, the values of the upper limit of the SIE calculated according to the approach undertaken in our study differ from the values calculated in previous studies. Main conclusions Although the classical species–area models have been used to estimate the upper limit of the SIE, we propose that the detection of this phenomenon should be undertaken independently from the species–area relationship, so that the net effects of area are calculated excluding the surrogate action of area on other variables, such as environmental heterogeneity. The SIE appears when and where area ceases to influence species richness directly. There are two distinct SIE patterns: (1) the classical SIE where both the direct and indirect effects of area are eliminated and (2) the cryptic SIE where area affects species richness indirectly. Our approach offers the opportunity of studying the different factors influencing biodiversity on small scales more accurately. The SIE cannot be considered a general pattern with fixed behaviour that can be described by the same model for different island groups and taxa. The SIE should be recognized as a genuine but idiosyncratic phenomenon.     

Colonization and extinction of ant communities in a fragmented landscape

Abstract In this paper we tested the assumption that smaller and more isolated remnants receive fewer ant colonizers and lose more species. We also tested hypotheses to explain such a pattern. We sampled ants in Brazil for 3 years in 18 forest remnants and in 10 grasslands between them. We tested the influence of remnant area and isolation on colonization rate, as well as the effect of remnant area on extinction rate. We tested the correlation between remnant area and isolation to verify the landscape design. Colonization rate was not affected by remnant area or isolation. Extinction rate, however, was smaller in larger remnants. Remnant area and isolation were negatively correlated. We tested two hypotheses related to the decrease in ant species extinction rate with increased remnant area: (i) small remnants support smaller and more extinction‐prone populations; and (ii) small remnants are more often invaded by generalist species, which suffer higher extinction inside remnants. The density of ant populations significantly increased with area. Generalist species presented a lower colonization rate in larger remnants, contrary to the pattern observed in forest species. Generalist species suffered more extinction than expected inside remnants. The lack of response of colonization rate to remnant area can be explained by the differential colonization by generalist and forest species. The decrease of ant population density in smaller remnants could be related to loss of habitat quality or quantity. The higher colonization by generalist ant species in the smaller remnants could be related to landscape design, because smaller remnants are more similar to the matrix than larger ones. Our results have important implications for conservation strategies because small remnants seem to be more affected by secondary effects of fragmentation, losing more forest species and being invaded more often by generalist species. Studies that compare only species richness between remnants cannot detect such patterns in species composition.     

Spatio‐temporal scaling of biodiversity and the species–time relationship in a stream fish assemblage

1. The increase of species richness with the area of the habitat sampled, that is the species–area relationship, and its temporal analogue, the species–time relationship (STR), are among the few general laws in ecology with strong conservation implications. However, these two scale‐dependent phenomena have rarely been considered together in biodiversity assessment, especially in freshwater systems. 2. We examined how the spatial scale of sampling influences STRs for a Central‐European stream fish assemblage (second‐order Bernecei stream, Hungary) using field survey data in two simulation‐based experiments. 3. In experiment one, we examined how increasing the number of channel units, such as riffles and pools (13 altogether), and the number of field surveys involved in the analyses (12 sampling occasions during 3 years), influence species richness. Complete nested curves were constructed to quantify how many species one observes in the community on average for a given number of sampling occasions at a given spatial scale. 4. In experiment two, we examined STRs for the Bernecei fish assemblage from a landscape perspective. Here, we evaluated a 10‐year reach level data set (2000–09) for the Bernecei stream and its recipient watercourse (third‐order Kemence stream) to complement results on experiment one and to explore the mechanisms behind the observed patterns in more detail. 5. Experiment one indicated the strong influence of the spatial scale of sampling on the accumulation of species richness, although time clearly had an additional effect. The simulation methodology advocated here helped to estimate the number of species in a diverse combination of spatial and temporal scale and, therefore, to determine how different scale combinations influence sampling sufficiency. 6. Experiment two revealed differences in STRs between the upstream (Bernecei) and downstream (Kemence) sites, with steeper curves for the downstream site. Equations of STR curves were within the range observed in other studies, predominantly from terrestrial systems. Assemblage composition data suggested that extinction–colonisation dynamics of rare, non‐resident (i.e. satellite) species influenced patterns in STRs. 7. Our results highlight that the determination of species richness can benefit from the joint consideration of spatial and temporal scales in biodiversity inventory surveys. Additionally, we reveal how our randomisation‐based methodology may help to quantify the scale dependency of diversity components (α, β, γ) in both space and time, which have critical importance in the applied context.     

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.     

Determinants of alpha and beta vascular plant diversity in Mediterranean island systems: the Ionian islands,Greece

The Ionian archipelago is the second largest Greek archipelago after the Aegean, but the factors driving plant species diversity in the Ionian islands are still barely known. We used stepwise multiple regressions to investigate the factors affecting plant species diversity in 17 Ionian islands. Generalized dissimilarity modelling was applied to examine variation in the magnitude and rate of species turnover along environmental gradients, as well as to assess the relative importance of geographical and climatic factors in explaining species turnover. The values of the residuals from the ISAR log₁₀‐transfomed models of native and endemic taxa were used as a measure of island floristic diversity. Area was confirmed to be the most powerful single explanatory predictor of all diversity metrics. Mean annual precipitation and temperature, as well as shortest distance to the nearest island are also significant predictors of vascular plant diversity. The island of Kalamos constitutes an important plant diversity hotspot in the Ionian archipelago. The recent formation of the islands, the close proximity to the mainland source and the relatively low dispersal filtering of the Ionian archipelago has resulted in islands with a flora principally comprising common species and a low proportion of endemics. Small islands keep a key role in conservation of plant priority sites.     

sars: an R package for fitting,evaluating and comparing species–area relationship models

The species–area relationship (SAR) constitutes one of the most general ecological patterns globally. A number of different SAR models have been proposed. Recent work has shown that no single model universally provides the best fit to empirical SAR datasets: multiple models may be of practical and theoretical interest. However, there are no software packages available that a) allow users to fit the full range of published SAR models, or b) provide functions to undertake a range of additional SAR‐related analyses. To address these needs, we have developed the R package ‘sars’ that provides a wide variety of SAR‐related functionality. The package provides functions to: a) fit 20 SAR models using non‐linear and linear regression, b) calculate multi‐model averaged curves using various information criteria, and c) generate confidence intervals using bootstrapping. Plotting functions allow users to depict and scrutinize the fits of individual models and multi‐model averaged curves. The package also provides additional SAR functionality, including functions to fit, plot and evaluate the random placement model using a species–sites abundance matrix, and to fit the general dynamic model of oceanic island biogeography. The ‘sars’ R package will aid future SAR research by providing a comprehensive set of simple to use tools that enable in‐depth exploration of SARs and SAR‐related patterns. The package has been designed to allow other researchers to add new functions and models in the future and thus the package represents a resource for future SAR work that can be built on and expanded by workers in the field.     

Threats of land use to the global diversity of vascular plants

Aim

Land use is a main driver of biodiversity loss worldwide. However, quantifying its effects on global plant diversity remains a challenge due to the limited availability of data on the distributions of vascular plant species and their responses to land use. Here, we estimated the global extinction threat of land use to vascular plant species based on a novel integration of an ecoregion-level species-area model and the relative endemism richness of the ecoregions.

Location

Global.

Methods

First, we assessed ecoregion-level extinction threats using a countryside species–area relationship model based on responses of local plant richness to land use types and intensities and a high-resolution global land use map. Next, we estimated global species extinction threat by multiplying the relative endemism richness of each ecoregion with the ecoregion-level extinction threats.

Results

Our results indicate that 11% of vascular plant species are threatened with global extinction. We found the largest extinction threats in the Neotropic and Palearctic realms, mainly due to cropland of minimal and high intensity, respectively.

Main Conclusions

Our novel integration of the countryside species–area relationship and the relative endemism richness allows for the identification of hotspots of global extinction threat, as well as the contribution of specific land use types and intensities to this threat. Our findings inform where the development of measures to protect or restore plant diversity globally are most needed.     

The more‐individuals hypothesis revisited: the role of community abundance in species richness regulation and the productivity–diversity relationship

Species richness increases with energy availability, yet there is little consensus as to the exact processes driving this species–energy relationship. The most straightforward explanation is the more‐individuals hypothesis (MIH). It states that higher energy availability promotes a higher total number of individuals in a community, which consequently increases species richness by allowing for a greater number of species with viable populations. Empirical support for the MIH is mixed, partially due to the lack of proper formalisation of the MIH and consequent confusion as to its exact predictions. Here, we review the evidence of the MIH and evaluate the reliability of various predictions that have been tested. There is only limited evidence that spatial variation in species richness is driven by variation in the total number of individuals. There are also problems with measures of energy availability, with scale‐dependence, and with the direction of causality, as the total number of individuals may sometimes itself be driven by the number of species. However, even in such a case the total number of individuals may be involved in diversity regulation. We propose a formal theory that encompasses these processes, clarifying how the different factors affecting diversity dynamics can be disentangled.     

Island biogeography is not a single‐variable discipline: the small island effect debate

In some island systems, an ‘anomalous’ feature of species richness on smaller islands, in comparison with larger ones, has been observed. This has been described as the small island effect (SIE). The precise meaning of the term remains unresolved, as does the explanation for the phenomenon and even whether it exists. Dengler (2010 ; Diversity Distrib, 16 , 256–266.) addresses a number of conceptual and methodological issues concerning the nature and the detection of the SIE but fails to settle conclusively most of the issues he raises. We contend that his approach is theoretically flawed, especially in its treatment of habitat diversity. We offer a few suggestions of what is needed to advance understanding of the SIE.     

The environmental determinants of the insular butterfly faunas of the British Isles

A multiple regression analysis was performed upon selected environmental variables for a series of islands in the British Isles, to establish their effects upon the size of the butterfly fauna, measured as he number of species regularly breeding, S_B. So that the data be normally distributed, the regression analyses were performed upon log₁₀ transformed data only, with the data for outliers, mainland Britain and Ireland, the two largest islands, excluded. Most highly correlated with the number of butterfly species breeding upon an island is the number breeding within a 25 km radius of the nearest point of the mainland, r²=0.5941, followed by the correlations with the latitude of the mid-point of the island, r²=0.5541, the number of plant species comprising the island Hora, r²=0.5225, and the distance separating the island from the mainland, r²=0.4514. A partial correlation analysis confirms the importance of the parameters distance separating the island from the mainland, D₁, and the size of the faunal source S_F, and rejects the importance of the size of the flora and the latitude of the island. This is further confirmed by the results of a step-wise regression analysis, the two variables D₁ and S_F accounting for 66% of the variation of the butterfly fauna. If an alternative measure of isolation, D₂, which allows for the geographical clumping of islands, is combined with the variable S_F, then 69% of the variation of the butterfly fauna is accounted for.     

The environmental determinants of the insular butterfly faunas of the British Isles

A multiple regression analysis was performed upon selected environmental variables for a series of islands in the British Isles, to establish their effects upon the size of the butterfly fauna, measured as he number of species regularly breeding, S_B .
So that the data be normally distributed, the regression analyses were performed upon log₁₀ transformed data only, with the data for outliers, mainland Britain and Ireland, the two largest islands, excluded.
Most highly correlated with the number of butterfly species breeding upon an island is the number breeding within a 25 km radius of the nearest point of the mainland, r ²=0.5941, followed by the correlations with the latitude of the mid-point of the island, r ²=0.5541, the number of plant species comprising the island Hora, r ²=0.5225, and the distance separating the island from the mainland, r ²=0.4514.
A partial correlation analysis confirms the importance of the parameters distance separating the island from the mainland, D ₁, and the size of the faunal source S _F , and rejects the importance of the size of the flora and the latitude of the island. This is further confirmed by the results of a step-wise regression analysis, the two variables D ₁ and S_F accounting for 66% of the variation of the butterfly fauna.
If an alternative measure of isolation, D ₂, which allows for the geographical clumping of islands, is combined with the variable S_F , then 69% of the variation of the butterfly fauna is accounted for.     

Climate and food diversity as drivers of mammal diversity in Inner Mongolia

Traditionally, geographical distribution of biodiversity is assumed to be codetermined by multiple factors, for example, temperature, precipitation, environmental heterogeneity, and biotic interactions. However, few studies have simultaneously compared the relative roles of these factors in shaping the mammal diversity patterns for different feeding groups, that is, herbivores, insectivores, and carnivores. In this study, we assessed the relations between mammal diversity and current climate (mean annual temperature and precipitation), altitudinal range as well as mammal's food diversity in Inner Mongolia. Our results showed that the species richness for the three feeding guilds of mammals consistently increased with their food diversity, that is, species richness of plants, insects, and rodents. Mammal diversity also significantly decreased with mean annual temperature and precipitation. Random Forest models indicated that climate and food diversity were always included in the combinations of variables most associated with mammal diversity. Our findings suggest that while climate is an important predictor of large scale distribution of mammal diversity, biotic interactions, that is, food diversity, could also play important roles.     

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.