Ecological biogeography of Mexican bats: the relative contributions of habitat heterogeneity,beta diversity,and environmental gradients to species richness and composition patterns

Mexico has higher mammalian diversity than expected for its size and geographic position. High environmental hetero geneity throughout Mexico is hypothesized to promote high turnover rates (β‐diversity), thus contributing more to observed species richness and composition than within‐habitat (α) diversity. This is true if species are strongly associated with their environments, such that changes in environmental attributes will result in changes in species composition. Also, greater heterogeneity in an area will result in greater species richness. This hypothesis has been deemed false for bats, as their ability to fly would reduce opportunities for habitat specialization. If so, we would expect no significant relationships between 1) species composition and environmental variables, 2) species richness and environmental heterogeneity, 3) β‐diversity and environmental heterogeneity. We tested these predictions using 31 bat assemblages distributed across Mexico. Using variance partitioning we evaluated the relative contribution of vegetation, climate, elevation, horizontal heterogeneity (a variate including vegetation, climate, and elevational heterogeneity), spatial variation (lat‐long), and vertical hetero geneity (of vegetation strata) to variation in bat species composition and richness. Variation in vegetation explained 92% of the variation in species composition and was correlated with all other variables examined, indicating that bats respond directly to habitat composition and structure. Beta‐diversity and vegetational heterogeneity were significantly correlated. Bat species richness was significantly correlated with vertical, but not horizontal, heterogeneity. Nonetheless, neither horizontal nor vertical heterogeneity were random; both were related to latitude and to elevation. Variation in bat community composition and richness in Mexico were primarily explained by local landscape heterogeneity and environmental factors. Significant relationships between β‐diversity and environmental variation reveal differences in habitat specialization by bats, and explain their high diversity in Mexico. Understanding mechanisms acting along environmental or geographic gradients is as important for understanding spatial variation in community composition as studying mechanisms that operate at local scales.     

Bird species richness is associated with phylogenetic relatedness,plant species richness,and altitudinal range in Inner Mongolia

Bird species richness is mediated by local, regional, and historical factors, for example, competition, environmental heterogeneity, contemporary, and historical climate. Here, we related bird species richness with phylogenetic relatedness of bird assemblages, plant species richness, topography, contemporary climate, and glacial‐interglacial climate change to investigate the relative importance of these factors. This study was conducted in Inner Mongolia, an arid and semiarid region with diverse vegetation types and strong species richness gradients. The following associated variables were included as follows: phylogenetic relatedness of bird assemblages (Net Relatedness Index, NRI), plant species richness, altitudinal range, contemporary climate (mean annual temperature and precipitation, MAT and MAP), and contemporary‐Last Glacial Maximum (LGM) change in climate (change in MAT and change in MAP). Ordinary least squares linear, simultaneous autoregressive linear, and Random Forest models were used to assess the associations between these variables and bird species richness across this region. We found that bird species richness was correlated negatively with NRI and positively with plant species richness and altitudinal range, with no significant correlations with contemporary climate and glacial–interglacial climate change. The six best combinations of variables ranked by Random Forest models consistently included NRI, plant species richness, and contemporary‐LGM change in MAT. Our results suggest important roles of local ecological factors in shaping the distribution of bird species richness across this semiarid region. Our findings highlight the potential importance of these local ecological factors, for example, environmental heterogeneity, habitat filtering, and biotic interactions, in biodiversity maintenance.     

The importance of sward architectural complexity in structuring predatory and phytophagous invertebrate assemblages

Abstract.  1. Although the importance of plant community assemblages in structuring invertebrate assemblages is well known, the role that architectural complexity plays is less well understood. In particular, direct empirical data for a range of invertebrate taxa showing how functional groups respond to plant architecture is largely absent from the literature.
2. The significance of sward architectural complexity in determining the species richness of predatory and phytophagous functional groups of spiders, beetles, and true bugs, sampled from 135 field margin plots over 2 years was tested. The present study compares the relative importance of sward architectural complexity to that of plant community assemblage.
3. Sward architectural complexity was found to be a determinant of species richness for all phytophagous and predatory functional groups. When individual species responses were investigated, 62.5% of the spider and beetle species, and 50.0% of the true bugs responded to sward architectural complexity.
4. Interactions between sward architectural complexity and plant community assemblage indicate that the number of invertebrate species supported by the plant community alone could be increased by modification of sward architecture. Management practices could therefore play a key role in diversifying the architectural structure of existing floral assemblages for the benefit of invertebrate assemblages.
5. The contrasting effects of sward architecture on invertebrate functional groups characterised by either direct (phytophagous species) or indirect (predatory species) dependence on plant communities is discussed. It is suggested that for phytophagous taxa, plant community assemblage alone is likely to be insufficient to ensure successful species colonisation or persistence without appropriate development of sward architecture.     

Bromeliad architectural complexity and vertical distribution predict spider abundance and richness

Habitat complexity is a main predictor of the distribution of arthropods on vegetation. However, it remains poorly known whether plant architecture and fine‐scale spatial distribution affect the species richness and composition of associated arthropod guilds. In this study, we extensively sampled bromeliad species with a variety of rosette architectures in a megadiversity region. The aims were to investigate whether (i) possible differences in spider species composition among bromeliad species are related to the distinct architectures of the plants, and (ii) bromeliad architectural complexity (an intrinsic feature) and vertical distribution (an extrinsic feature) are good predictors of spider abundance and richness. Contrary to our expectations, spider species composition did not vary significantly among bromeliad species with different architectures. We found a positive effect of the mean number of leaves on spider abundance and species richness, but it occurred indirectly through spider abundance; factoring out the indirect effect revealed a negative effect of leaf number on species richness. Bromeliad species with wider vertical distributions harboured more spider species. Our results suggest that the dominance of a few spider species and reduced space for orb‐web spiders to attach their webs are the main explanations for lower spider richness on bromeliad species with higher architectural complexity. Our findings highlight the importance of both intrinsic and extrinsic plant features as co‐determinants of predator arthropod diversity.     

Disentangling the effects of area, energy and habitat heterogeneity on boreal forest bird species richness in protected areas

Aim  One of the few general laws in ecology is that species richness is a positive function of area. However, it has been proposed that area would merely be a proxy for energy. Additionally, habitat heterogeneity has been found to be an important factor determining species richness. Yet the relative importance of those relationships is little known, and it is still unclear how they are brought about. We aimed to dissect which factors drive the species richness of boreal forest birds, and to identify the most probable mechanisms.
Location  Forested protected areas in Finland.
Methods  Using bird line census data collected in 104 protected areas, we ran simultaneous autoregressive models to explain the species richness of forest birds. We explored the value of forest area, tree volume, tree growth, mean degree days and habitat heterogeneity as explanatory variables and used the species richness within different species groups, based on the predictions of hypothesized mechanisms, as a response variable.
Results  Energy, rather than area or habitat heterogeneity, seems to be the main driver of species richness in boreal forest birds. More specifically, productive energy was a better predictor of total species richness than solar energy. Among the tested hypothetical mechanisms, the sampling hypothesis received strong support. After accounting for sampling, solar energy had an effect on species richness.
Main conclusions  As productive energy, such as tree volume, is associated with species richness, high-energy areas should be prioritized in forest conservation planning. Reductions in productive energy may first lead to the disappearance of the rarest species due to the random sampling process. Climate change may result in increased species richness due to increasing amount of productive and solar energy in forests. However, the range shifts of bird species may not be fast enough to keep up with the temperature increases.     

The importance of canopy complexity in shaping seasonal spider and beetle assemblages in saltmarsh habitats

1. Habitat structure, including vegetation structural complexity, largely determines invertebrate assemblages in semi‐natural grasslands. The importance of structural complexity to the saltmarsh invertebrate community, where the interplay between vegetation characteristics and tidal inundation is key, is less well known. 2. It was hypothesised that canopy complexity would be a more important predictor of spider and beetle assemblages than simple vegetation attributes (e.g. height, community type) and environmental variables (e.g. elevation) alone, measured in two saltmarsh regions, south‐east (Essex) and north‐west (Morecambe Bay) U.K. Canopy complexity (number of non‐vegetated ‘gaps’ in canopy ≥ 1 mm wide) was assessed using side‐on photography. Over 1500 spiders and beetles were sampled via suction sampling, winter and summer combined. 3. In summer, saltmarshes with abundant spider and beetle populations were characterised by high scores for canopy complexity often associated with tussocky grass or shrub cover. Simple vegetation attributes (plant cover, height) accounted for 26% of variation in spider abundance and 14% in spider diversity, rising to 46% and 41%, respectively, with the addition of canopy complexity score. Overwintering spider assemblages were associated with elevation and vegetation biomass. Summer beetle abundance, in particular the predatory and zoophagous group, and diversity were best explained by elevation and plant species richness. 4. Summer canopy complexity was identified as a positive habitat feature for saltmarsh spider communities (ground‐running hunters and sheet weavers) with significant ‘added value’ over more commonly measured attributes of vegetation structure.     

How does habitat diversity affect the species–area relationship?

Aim   To examine the way in which 'area' and 'habitat diversity' interact in shaping species richness and to find a simple and valid way to express this interaction.
Location   The Natura 2000 network of terrestrial protected areas in Greece, covering approximately 16% of the national territory.
Methods   We used the Natura 2000 framework, which provides a classification scheme for natural habitat types, to quantify habitat heterogeneity. We analysed data for the plant species composition in 16,143 quadrats in which 5044 species and subspecies of higher plants were recorded. We built a simple mathematical model that incorporates the effect of habitat diversity on the species–area relationship (SAR).
Results   Our analysis showed that habitat diversity was correlated with area. However, keeping habitat diversity constant, species richness was related to area; while keeping area constant, species richness was related to habitat diversity. Comparing the SAR of the 237 sites we found that the slope of the species–area curve was related to habitat diversity.
Main conclusions   Discussion of the causes of the SAR has often focused on the primacy of area per se versus habitat heterogeneity, even though the two mechanisms are not mutually exclusive and should be considered jointly. We find that increasing habitat diversity affects the SAR in different ways, but the dominant effect is to increase the slope of the SAR. While a full model fit typically includes a variety of terms involving both area and habitat richness, we find that the effect of habitat diversity can be reduced to a linear perturbation of the slope of the species accumulation curve.     

How does forest landscape structure explain tree species richness in a Mediterranean context?

Although the strong relationship between vegetation and climatic factors is widely accepted, other landscape composition and configuration characteristics could be significantly related with vegetation diversity patterns at different scales. Variation partitioning was conducted in order to analyse to what degree forest landscape structure, compared to other spatial and environmental factors, explained forest tree species richness in 278 UTM 10 × 10 km cells in the Mediterranean region of Catalonia (NE Spain). Tree species richness variation was decomposed through linear regression into three groups of explanatory variables: forest landscape (composition and configuration), environmental (topography and climate) and spatial variables. Additionally, the forest landscape characteristics which significantly contributed to explain richness variation were identified through a multiple regression model. About 60% of tree species richness variation was explained by the whole set of variables, while their joint effects explained nearly 28%. Forest landscape variables were those with a greater pure explanatory power for tree species richness (about 15% of total variation), much larger than the pure effect of environmental or spatial variables (about 2% each). Forest canopy cover, forest area and land cover diversity were the most significant composition variables in the regression model. Landscape configuration metrics had a minor effect on forest tree species richness, with the exception of some shape complexity indices, as indicators of land use intensity and edge effects. Our results highlight the importance of considering the forest landscape structure in order to understand the distribution of vegetation diversity in strongly human-modified regions like the Mediterranean.     

Vascular plant species richness on wetland remnants is determined by both area and habitat heterogeneity

There is an ongoing ecological debate on whether area per se or habitat heterogeneity is the main driver for species richness. The wetland remnants in the Sanjiang Plain, NE China harbor a high biodiversity and play an important role for local ecosystems. Fifty-one wetland remnants were sampled to examine the effect of area and habitat heterogeneity on vascular plant species richness. Number of community types, elevation, water heterogeneity and soil resource heterogeneity were employed as habitat heterogeneity variables, but only water heterogeneity was identified as the proper surrogate for habitat heterogeneity. Compared with the classic species-area model, the choros model achieved better fitness when water heterogeneity and elevation were employed as habitat heterogeneity variables. Nevertheless, elevation was poorly correlated with species richness. It suggests, without a comprehensive analysis of habitat heterogeneity variables, the choros model might result in a misleading result. In this study, species richness was significantly influenced by water heterogeneity, area and number of community types. Water heterogeneity and area both controlled the number of community types, and they were the two main determinants of species richness. As area was significantly and positively correlated with water heterogeneity, the variance in species richness was mainly related to the mutual effect of area and water heterogeneity. The results of this study confirmed that the relationship between the area per se hypothesis and the habitat heterogeneity hypothesis was conjunct rather than mutually exclusive. In addition, it is critical that both area and water heterogeneity should be taken into account for biodiversity conservation and management in wetland remnants.     

Spatial heterogeneity and habitat configuration overcome habitat composition influences on alpha and beta mammal diversity

The effects of habitat fragmentation on different taxa and ecosystems are subject to intense debate, and disentangling them is of utmost importance to support conservation and management strategies. We evaluated the importance of landscape composition and configuration, and spatial heterogeneity to explain α- and β-diversity of mammals across a gradient of percent woody cover and land use diversity. We expected species richness to be positively related to all predictive variables, with the strongest relationship with landscape composition and configuration, and spatial heterogeneity respectively. We also expected landscape to influence β-diversity in the same order of importance expected for species richness, with a stronger influence on nestedness due to deterministic loss of species more sensitive to habitat disturbance. We analyzed landscape structure using: (a) landscape metrics based on thematic maps and (b) image texture of a vegetation index. We compared a set of univariate explanatory models of species richness using AIC, and evaluated how dissimilarities in landscape composition and configuration and spatial heterogeneity affect β-diversity components using a Multiple Regression on distance Matrix. Contrary with our expectations, landscape configuration was the main driver of species richness, followed by spatial heterogeneity and last by landscape composition. Nestedness was explained, in order of importance, by spatial heterogeneity, landscape configuration, and landscape composition. Although conservation policies tend to focus mainly on habitat amount, we advocate that landscape management must include strategies to preserve and improve habitat quality and complexity in natural patches and the surrounding matrix, enabling landscapes to harbor high species diversity.     

Relative influence of habitat heterogeneity, climate, human disturbance, and spatial structure on vertebrate species richness in Spain

Many factors affect the distribution of species richness. This study examines the relative influence of habitat heterogeneity, climate, human disturbance, and spatial structure on the species-richness distribution of terrestrial vertebrates (amphibians, reptiles, birds and mammals) in mainland Spain. The results indicate that spatial structure and environment exert similar influences on species richness. For all four taxa, species richness increases southward and northward, being lower in the center of the country, when controlled for other variables. This may be the result of a peninsular effect, as found in other studies, and reflect the importance of historical events on species richness in the Iberian Peninsula. Climate is more important than habitat heterogeneity in determining species richness. Temperature is positively correlated with amphibian, reptile, and bird species richness, while mammalian species richness is highest at intermediate temperatures. This effect is stronger in ectotherms than among endotherms, perhaps reflecting physiological differences. Precipitation positively correlates with bird and mammalian species richness, but has no effect on ectotherm species richness. Amphibian species richness increases with altitudinal range, and bird species richness with habitat diversity. Human population density is positively correlated with bird and mammalian species richness, but does not affect ectotherm species richness, while amphibian and bird species richness is highest at moderate levels of human land alteration (farmland). However, unexplained variance remains, and we discuss that the effects of environmental variables on species richness may vary geographically, causing different effects to be obscured on a national scale, diminishing the explanatory power of environmental variables.     

Environmental drivers of spider community composition at multiple scales along an urban gradient

Broad-scale modification of natural ecosystems associated with urbanisation often leads to localised extinctions and reduced species richness. Despite this, habitats within the urban matrix are still capable of supporting biodiversity to varying degrees. As species have different responses to anthropogenic habitat modification, the species composition of urban areas can depend greatly on the habitat characteristics of the local and surrounding areas. The aim of this study was to compare the community composition of spiders in private gardens, urban parks, patches of remnant vegetation and continuous bushland sites, so as to identify habitat variables associated with variation in spider populations along and within the urban gradient and matrix. Overall spider abundances and richness were highest in remnant vegetation patches and were associated with increased vegetation cover at microhabitat and landscape-scales. While gardens were not as diverse as remnant patches, they did support a surprisingly high diversity of spiders. We also found that species composition differed significantly between gardens and other urban green spaces. Higher richness within gardens was also associated with greater vegetation cover, indicating the importance of private management decisions on local biodiversity. Differences in community composition between land-use types were driven by a small number of urban-tolerant species, and spider guilds showed different responses to habitat traits such as vegetation cover and human population densities. This study demonstrates that urban land-uses support unique spider communities and that maintaining vegetation cover within the urban matrix is essential in order to support diverse spider communities in cities.     

Environmental features influencing Carabid beetle (Coleoptera) assemblages along a recently deglaciated area in the Alpine region

Abstract.  1. The spatio-temporal approach was used to evaluate the environmental features influencing carabid beetle assemblages along a chronosequence of an Italian Alpine glacier foreland. The influence of environmental variables on species richness, morphology (wing and body length), and distribution along the chronosequence was tested.
2. Species richness was found to be a poor indicator of habitat due to weak influences by environmental variables. It seems that the neighbouring habitats of a glacier foreland are not able to determine significant changes in carabid species richness.
3. Instead it appears that history (age since deglaciation) and habitat architecture of a glacier foreland are strongly correlated to species adaptive morphological traits, such as wing morphology and body length. Assemblages characterised by species with reduced wing size are linked to the older stages of the chronosequence, where habitat is more structured. Assemblages characterised by the largest species are linked to the younger sites near the glacier. These morphological differentiations are explained in detail.
4. Habitat age can therefore be considered the main force determining assemblage composition. On the basis of the relationship between morphological traits and environmental variables, it seems likely that age since deglaciation is the main variable influencing habitat structure (primary effect) on the Forni foreland. The strong relationship between carabid assemblages and habitat type indicates that site age has but a secondary effect on carabid assemblages. This may be utilised to interpret potential changes in assemblages linked to future glacier retreat.     

Conservation of arthropod diversity in montane wetlands: effect of altitude, habitat quality and habitat fragmentation on butterflies and grasshoppers

1. The patterns of arthropod diversity were investigated in 24 montane wetlands in Switzerland. These differed in altitude, management regime (cattle-grazing vs. mowing), vegetation structure (index combining vegetation height and density) and degree of habitat fragmentation.
2. The general arthropod diversity was determined by net sampling at 10 sampling points per site. The diversity of grasshoppers and butterflies was measured by counting species richness at the site and species density (species richness per unit area) on transects. The species richness of grasshoppers and butterflies was found to be more sensitive to the geographical attributes of the site whereas species density was more affected by the habitat quality.
3. Grasshopper diversity decreased within the observed altitudinal range (800–1400 m) and was higher at grazed sites, whereas butterfly diversity was higher at mown sites. Arthropod diversity but not abundance of arthropods was positively related to the vegetation structure.
4. The species richness of butterflies was negatively influenced by the degree of habitat fragmentation: both the size of habitat as well as the area of wetland habitats within 4 km were related positively to the number of specialist wetland butterflies.
5. Late mowing as well as low-density cattle-grazing are appropriate management actions to maintain arthropod diversity in montane wetlands. In order to establish site-specific management plans, the biology of the present target species as well as the historical context should be considered.
6. We suggest that the best protection for the species examined in this study would be a network of wetland sites managed using a variety of traditional, non-intensive methods. This can only be achieved by coordinated planning of conservation measures among sites.     

Refuge from cannibalism in complex-structured habitats: implications for the accumulation of invertebrate predators

Abstract.  1. Like many invertebrate predators, the wolf spider Pardosa littoralis Banks (Araneae: Lycosidae) accumulates in complex-structured habitats replete with leaf litter (thatch). Here we test the hypothesis that P. littoralis accumulates in complex habitats to gain refuge from cannibalism.
2. A laboratory experiment examined the effects of habitat complexity (thatch present or absent) and size-class pairing of conspecific spiders (large vs. small, small vs. small, and large vs. large) on the incidence of cannibalism. Spider survival was significantly higher (22%) in complex-structured habitats with thatch than in simple-structured habitats lacking thatch. Furthermore, cannibalism occurred more frequently in P. littoralis when the size of conspecifics was asymmetric (large vs. small spiders) than when spiders were of equal size. There was no interactive effect of habitat complexity and size-class pairing on spider survival.
3. A field experiment examined the effects of habitat complexity, conspecific density, and access to alternative prey on the prevalence of cannibalism in P. littoralis . Access to alternative prey significantly increased the number of spiders recovered from field enclosures, as did the presence of leaf litter thatch. That fewer spiders were recovered when thatch and alternative prey were absent suggests that cannibalism was most prevalent under these conditions.
4. Overall, results suggest that habitat complexity reduces agonistic interactions and cannibalism among wolf spiders, providing encouragement to pest managers that the structure of agricultural habitats can be managed to maximise densities of generalist predators for enhanced pest suppression.     

Habitat structure determines spider diversity in highland ponds

Wetlands (e.g. ponds, meadows) can be found in many landscapes, playing an important role in maintaining regional biodiversity and supporting heterogeneous communities. Spiders are diversified predators that are highly influenced by changes in plant community structure, heterogeneous habitats sustain high spider diversity and abundance. We investigated the characteristics of spider biodiversity in ponds with different habitat structures, by examining patterns across habitats of ponds with different vegetation levels. Sampling took place in four occasions over a year. We compared spider abundance, species richness and composition among ponds including distinct vegetation variables, related to life form, type of leaves, coverage and height. Overall 1174 individuals (194 adults) of 11 families and 37 morphospecies were sampled. We found mostly expected differences in the manner that communities were structured between different habitats. Thus, higher variability of abundance was explained for higher habitat structure of ponds. We also found differences in species composition between ponds with low emergent vegetation and higher habitat structures. Additionaly, spiders were consistently structured more by turnover than nestedness components, with a greater beta diversity of web-builders. Our results suggest varying levels of habitat structures and species substitution shape pond spider communities, depending on habitat heterogeneity and spider guild. Those findings demonstrate the clear role of spatial habitat structure, with more spider species preferring to build webs or actively hunt at vegetated environments on ponds.     

Plant species richness–environment relationships across the Subantarctic–Patagonian transition zone

Aim To evaluate the relative importance of climate, productivity, environmental heterogeneity, biotic associations and habitat use by cattle to account for the species richness of trees, shrubs and herbs across the Subantarctic–Patagonian transition. Location An area of c. 150 × 150 km, within the transition zone between the Subantarctic and Patagonian subregions on the eastern slope of the Andes (c. 39–42° S, 70–72° W). Methods All vascular plants found at each one of 50 (10 × 10 m) sampling plots were counted to estimate the local tree, shrub and herb species richness. Path analysis was used to evaluate the relationship between the richness of the three life‐forms and plant cover, dried litter biomass, mean annual temperature, annual precipitation, daily temperature range, substrate heterogeneity and number of faecal pats. Principal coordinates of neighbour matrices was used to model the spatial autocorrelation of the data. Results Total plant species richness showed a unimodal pattern of spatial variation across the transition. Richness responded positively to indirect effects of precipitation mediated through plant cover, but there was a negative overall effect of precipitation on richness towards the west of the transition, most strongly for trees. An increase in substrate heterogeneity promoted a local increase in herb and shrub richness; the richness of trees increased in sites with steeper slopes. Canopy closure had a direct negative impact on herb richness; it also increased the local accumulation of litter, which negatively affected shrub and herb richness. The impact of habitat use by cattle negatively affected herb richness in areas to the east of the biogeographical transition. Main conclusions We suggest that the importance of indirect climatic effects mediated by vegetation cover can account for species richness patterns across this transition, most strongly for woody species, which supports the productivity hypothesis. The southern temperate forests towards the west may represent a deviation from the predictions of the water–energy dynamics hypothesis. Dissimilar spatial patterns of variation in the richness of woody and herbaceous species, and their different responses to climatic and heterogeneity variables across the transition, suggest that plant life‐form influences the plant species richness–environment relationships.     

Disentangling the Role of Climate,Topography and Vegetation in Species Richness Gradients

Environmental gradients (EG) related to climate, topography and vegetation are among the most important drivers of broad scale patterns of species richness. However, these different EG do not necessarily drive species richness in similar ways, potentially presenting synergistic associations when driving species richness. Understanding the synergism among EG allows us to address key questions arising from the effects of global climate and land use changes on biodiversity. Herein, we use variation partitioning (also know as commonality analysis) to disentangle unique and shared contributions of different EG in explaining species richness of Neotropical vertebrates. We use three broad sets of predictors to represent the environmental variability in (i) climate (annual mean temperature, temperature annual range, annual precipitation and precipitation range), (ii) topography (mean elevation, range and coefficient of variation of elevation), and (iii) vegetation (land cover diversity, standard deviation and range of forest canopy height). The shared contribution between two types of EG is used to quantify synergistic processes operating among EG, offering new perspectives on the causal relationships driving species richness. To account for spatially structured processes, we use Spatial EigenVector Mapping models. We perform analyses across groups with distinct dispersal abilities (amphibians, non-volant mammals, bats and birds) and discuss the influence of vagility on the partitioning results. Our findings indicate that broad scale patterns of vertebrate richness are mainly affected by the synergism between climate and vegetation, followed by the unique contribution of climate. Climatic factors were relatively more important in explaining species richness of good dispersers. Most of the variation in vegetation that explains vertebrate richness is climatically structured, supporting the productivity hypothesis. Further, the weak synergism between topography and vegetation urges caution when using topographic complexity as a surrogate of habitat (vegetation) heterogeneity.     

Effects of habitat loss, habitat fragmentation, and isolation on the density, species richness, and distribution of ladybeetles in manipulated alfalfa landscapes

Abstract.  1. Habitat loss and fragmentation are the main causes of changes in the distribution and abundance of organisms, and are usually considered to negatively affect the abundance and species richness of organisms in a landscape. Nevertheless, habitat loss and fragmentation have often been confused, and the reported negative effects may only be the result of habitat loss alone, with habitat fragmentation having nil or even positive effects on abundance and species richness.
2. Manipulated alfalfa micro-landscapes and coccinellids (Coleoptera: Coccinellidae) are used to test the effects habitat loss (0% or 84%), fragmentation (4 or 16 fragments), and isolation (2 or 6 m between fragments) on the density, species richness, and distribution of native and exotic species of coccinellids.
3. Generally, when considering only the individuals in the remaining fragments, habitat loss had variable effects while habitat fragmentation had a positive effect on the density of two species of coccinellids and on species richness, but did not affect two other species. Isolation usually had no effect. When individuals in the whole landscape were considered, negative effects of habitat loss became apparent for most species, but the positive effects of fragmentation remained only for one species.
4. Native and exotic species of coccinellids did not segregate in the different landscapes, and strong positive associations were found most often in landscapes with higher fragmentation and isolation.
5. The opposing effects of habitat loss and fragmentation may result in a nil global effect; therefore it is important to separate their effects when studying populations in fragmented landscapes.     

Habitat Complexity and Invasive Species: The Impacts of Gamba Grass (Andropogon gayanus) on Invertebrates in an Australian Tropical Savanna

Habitat complexity is an important factor structuring local faunal assemblages. Many invasive plant species alter vegetation complexity because they are both functionally and structurally different from native flora. This study investigated the effects of an invasive grass (Andropogon gayanus Kunth.), which is invading undisturbed savannas in northern Australia. Although A. gayanus is a similar lifeform to that dominant in the native understory (i.e., a C4 grass), it is structurally different and significantly alters habitat complexity. We, therefore, hypothesized that there would be a substantial effect of the invasive grass on ant, spider, and other invertebrate assemblages. Contrary to our hypothesis, there was no effect of A. gayanus on ant species richness, abundance, or composition, nor were spiders or other invertebrates affected. Instead the change in weather conditions with season was more important in structuring the local invertebrate assemblages. Change in habitat complexity was most pronounced vertically, rather than horizontally at ground level, thus although there was a clear difference in the vertical structure between invaded and noninvaded habitats, the limited invertebrate response to different A. gayanus densities suggests some invertebrates are less sensitive to vertical changes in vegetation structure.