Environmental determinants of woody and herbaceous riparian vegetation patterns in a semi-arid mediterranean basin

What environmental variables determine riparian vegetation patterns? Are there differences between woody and herbaceous species? To answer these questions, we first explored the composition and richness patterns of both riparian woody and herbaceous species in a semi-arid mediterranean basin. Then, we assessed the environmental factors (climate, geology, topography, hydrogeomorphology and land use) that best explain these patterns. We used the following methodological approaches: clustering analyses, distance-based linear models, generalised linear models and hierarchical partitioning procedures. Valley shape, drought duration, river habitat heterogeneity, water conductivity and agricultural land use were the most important variables explaining variation in species composition for both groups. Woody riparian richness was mainly influenced by flow conditions and valley shape, whereas herbaceous one was more dependent on substrate features. Thus, although some differences in the importance of individual variables were observed, we found a notable congruence in the composition and species richness of both groups and also in the main types of variables explaining these patterns (hydrogeomorphology and land use, especially agriculture). Our results show that both communities could be treated in a holistic way, since they respond similarly to the strong natural and anthropogenic environmental gradients present in mediterranean basins.     

Richness, diversity, and rate of primary succession over 20?year in tropical coastal dunes

The tropical coastal dunes in central Gulf of Mexico have been stabilizing over the last decades resulting in reduced substrate mobility, and promoting primary succession. We describe changes in species richness and diversity in dune vegetation during 20?years. Our questions: (a) Do species richness and diversity increase over time as predicted by models of ecological succession or do they show a hump-backed manner similar to the observations in temperate coastal dunes?, (b) What is the interaction between vegetation cover and diversity and species richness?, (c) Is there a relationship between species diversity and succession rate and does succession rate change over time?, and (d) How do plant functional types change during succession? In order to answer these questions, we set 140 4?×?4?m permanent plots in a mobile dune area and monitored vegetation cover and species richness from 1991 to 2011. In time, diversity increased in a logistic manner toward an asymptotic value once vegetation cover surpassed 60?%. Species richness increased in a humped-back shape, also reaching a maximum peak at 60?% vegetation cover. The succession rate of diversity was measured by the Euclidean distance, and showed a significant humped-back relation, meaning that it was slower in early and late successional stages. The study supports the intermediate disturbance theory. The conservation of coastal dunes vegetation should focus on all, species-poor and species-rich habitats that help to maintain the ecological integrity of these ecosystems. The understanding of community dynamics and diversity patterns becomes an essential component of coastal dune management and conservation.     

The role of landscape structure in determining palynological and floristic richness

The associations between floristic and palynological richness and landscape structure were studied based on modern pollen?Cvegetation data from a patchy cultural landscape in southern Estonia (northern temperate vegetation zone). Nine study sites (small lakes and their surrounding vegetation) represent land cover gradient from closed forest to semi-open vegetation. Floristic richness (number of species) and floristic richness of pollen types (number of pollen-equivalent taxa) were used to describe the vegetation within the radius of 250?m from the pollen sampling sites. Palynological richness was calculated to describe the modern pollen samples diversity. Landscape structure was estimated on the basis of landscape openness and three landscape diversity measures: richness of community patches, Simpson evenness of community patches and Simpson diversity of community patches. To study the effect of the spatial scale of landscapes on the vegetation?Clandscape and pollen?Clandscape associations, landscape structure was estimated within eight radii (250?C2,000?m) around each lake. The results showed that landscape openness was the most important determinant of both floristic richness and palynological richness in southern Estonia and that landscape diversity estimated by Simpson diversity index was also significantly associated with the richness estimates. Floristic and palynological richness were significantly positively correlated with landscape structure within the radii greater than 1,000?m from the pollen sampling sites, which is similar to the estimated Relevant Source Area of Pollen in southern Estonia. We conclude that within one floristic or climatic region, palynological richness gives reliable estimates about the variation in floristic richness and landscape structure; however, caution must be taken when comparing pollen-inferred vegetation diversities from different regions or when interpreting fossil pollen records from times with highly different vegetation associations.     

Low-Intensity Agricultural Landscapes in Transylvania Support High Butterfly Diversity: Implications for Conservation

European farmland biodiversity is declining due to land use changes towards agricultural intensification or abandonment. Some Eastern European farming systems have sustained traditional forms of use, resulting in high levels of biodiversity. However, global markets and international policies now imply rapid and major changes to these systems. To effectively protect farmland biodiversity, understanding landscape features which underpin species diversity is crucial. Focusing on butterflies, we addressed this question for a cultural-historic landscape in Southern Transylvania, Romania. Following a natural experiment, we randomly selected 120 survey sites in farmland, 60 each in grassland and arable land. We surveyed butterfly species richness and abundance by walking transects with four repeats in summer 2012. We analysed species composition using Detrended Correspondence Analysis. We modelled species richness, richness of functional groups, and abundance of selected species in response to topography, woody vegetation cover and heterogeneity at three spatial scales, using generalised linear mixed effects models. Species composition widely overlapped in grassland and arable land. Composition changed along gradients of heterogeneity at local and context scales, and of woody vegetation cover at context and landscape scales. The effect of local heterogeneity on species richness was positive in arable land, but negative in grassland. Plant species richness, and structural and topographic conditions at multiple scales explained species richness, richness of functional groups and species abundances. Our study revealed high conservation value of both grassland and arable land in low-intensity Eastern European farmland. Besides grassland, also heterogeneous arable land provides important habitat for butterflies. While butterfly diversity in arable land benefits from heterogeneity by small-scale structures, grasslands should be protected from fragmentation to provide sufficiently large areas for butterflies. These findings have important implications for EU agricultural and conservation policy. Most importantly, conservation management needs to consider entire landscapes, and implement appropriate measures at multiple spatial scales.     

A floristic-structural gradient related to land forms in the southern Chihuahuan Desert

Abstract. The purposes of this study were to elucidate the floristic and structural characteristics of the vegetationin the Mapimi subdivision of the Chihuahuan Desert, Mexico, and to relate them to environmental variation. The main question addressed was: How does floristic composition, total species richness and life-form species richness vary in relation to environmental change? 154 sites, randomly selected and stratified over seven landscape units, were analyzed. Results showed the existence of a land form gradient along which vegetation types were ordered. Species richness varied along the gradient, the richest land form was bajada, indicating that the maximum species richness did not occur at one extreme of the water availability gradient but in a moderately poor situation. The lowest species richness was found in the playa land form. Cover-based life form spectra varied significantly with land forms, while presence-absence based life form spectra did not. It is suggested that this may be a consequence of the relatively young age of this desert.     

Species and functional responses of ants to inter-row tillage and vegetation in organic Mediterranean vineyards

Ants are arthropods providing crucial ecosystem services such as soil structuring, nutrient cycling, seed dispersal and pest predation. Thus, their abundance and diversity need to be considered in approaches to improve sustainability of land use such as Mediterranean viticulture. In our study, we tested whether (1) inter-row vegetation and the absence of tillage increase the species richness and/or functional diversity of ants in vineyards and (2) ground cover vegetation drives ant species composition. We included 23 Mediterranean organic vineyards in our analyses and distinguished three types of inter-row management: all inter-rows tilled, half of the inter-rows tilled, and all inter-rows are untilled and covered by vegetation. The occurrence of ant species was analysed in six pitfall traps per vineyard. Around each trap, the floristic composition of inter-row vegetation was analysed in 2 × 2 m² plots. We found that inter-row tillage significantly affected ant species richness, being higher in partially than in fully tilled vineyards whereas untilled vineyards were not different from the other tillage types. Grass cover and the perennial/annual rate were positively correlated with ant species richness. Ant functional diversity and the frequency of most predatory ants were not correlated neither with plant functional groups nor with tillage type. In conclusion, ant communities benefit from inter-row vegetation and/or absence of soil disturbance but partial inter-row tillage of vineyards may be tolerated and even benefit several species. In particular, grasses and perennial plant species favour ants in our system and need to be considered in inter-row sowing.     

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.     

Optical trait indicators for remote sensing of plant species composition: Predictive power and seasonal variability

Most plant species feature similar biochemical compositions and thus similar spectral signals. Still, empirical evidence suggests that the spectral discrimination of species and plant assemblages is possible. Success depends on the presence or absence of faint but detectable differences in biochemical (e.g., pigments, leaf water and dry matter content) and structural properties (e.g., leaf area, angle, and leaf structure), i.e., optical traits. A systematic analysis of the contributions and spatio-temporal variability of optical traits for the remote sensing of organismic vegetation patterns has not yet been conducted. We thus use time series of optical trait values retrieved from the reflectance signal using physical models (optical trait indicators, OTIs) to answer the following questions: How are optical traits related among patterns of floristic composition and reflectance? How variable are these relations in space and time? Are OTIs suitable predictors of plant species composition?We conducted a case study of three temperate open study sites with semi-natural vegetation. The canopy reflectance of permanent vegetation plots was measured on multiple dates over the vegetation period using a field spectrometer. We recorded the cover fractions of all plant species found in the vegetation plots and extracted gradients of species composition from these data. The physical PROSAIL leaf and canopy optical properties model was inverted with random forest regression models to retrieve time series of OTIs for each plot from the reflectance spectra. We analyzed these data sets using correlation analyses. This approach allowed us to assess the distribution of optical traits across gradients of species composition. The predictive performance of OTIs was tested in relation to canopy reflectance using random forest models.OTIs showed pronounced relationships with floristic patterns in all three study sites. These relationships were subject to considerable temporal variability. Such variability was driven by short-term vegetation dynamics introduced by local resource stress. In 72% of all cases OTIs out-performed the original canopy reflectance spectra as indicators of plant species composition. OTIs are also easier to interpret in an ecological sense than spectral bands or features. We thus conclude that optical traits retrieved from reflectance data have a high indicative value for ecological research and applications.     

Floristics and soil relations of upland swamp vegetation near Sydney

Abstract We examined relations between vegetation and soils, using multivariate methods, in hitherto poorly-known upland swamps on the Woronora Plateau, south of Sydney. A major trend in floristic composition was related to the height and cover of the herbaceous stratum and reflected a gradient in soil moisture and nutrients. A second trend in floristic composition was related to the height and cover of the shrub stratum, and may reflect the influence of recurring fires on certain dominant shrub species. Five plant communities were recognized on the basis of floristic composition and were distinguishable by their different soil habitats and/or structural characteristics. We report some of the highest species-richness values in the world (at scales of 1–15 m₂) for shrub/sedge-dominated vegetation, with up to 70 vascular plant species in 15 m2. Variation in species richness is inversely related to the resource gradient and positively related to the penetration of light through the vegetation canopy. This pattern is consistent with resource-competition models and warrants further investigation.     

Habitat structure components are effective predictors of trap-nesting Hymenoptera diversity

Habitat heterogeneity can be the major factor affecting species diversity in a community and measuring bee and wasp community habitat preferences in natural systems may provide insights for biodiversity management and conservation. In the present study, we investigate the effects of habitat structure components on solitary bee and wasp species richness and abundance. The research was conducted in an urban forest remnant in southeast Brazil. Our main questions were: (1) is similarity in habitat structure mirrored by similarity in Aculeate assemblage composition? and (2) what are the vegetation features that could be used as predictors of solitary bee and wasp richness and abundance? Aculeate bees and wasps were sampled using trap nests from February to November 2004. Trap nests were placed in sampling units located in 6 ha of secondary mesophytic forest. One hundred and thirty-seven trap nests were occupied by four species of wasps and seven species of bees. Altogether, our sampling units had a mean capture rate (relative to expected richness) of 72% during all the study period. The more similar sampling units were in terms of vegetation structure, the more similar they were in solitary bee and wasp species composition. The variance of tree abundance, shrub height and the abundance of wood logs were good predictors of solitary bee and wasp species richness and abundance in the study area. We demonstrate that even in a small scale it is possible to detect significant influences of habitat features on alpha diversity and that some of them are effective as predictors of trap-nesting Hymenoptera richness and abundance.     

Afforestation causes changes in post‐fire regeneration in native shrubland communities of northwestern Patagonia,Argentina

Question: What are the effects of fire in native shrubland communities and in pine plantations established in these shrublands? Location: Northern Patagonia, Argentina. Methods: We surveyed four sites in Chall‐Huaco valley, located in northwest Patagonia. Each site was a vegetation mosaic composed of an unburned Pinus ponderosa plantation, a plantation burned in 1996, and an unburned matorral and a matorral burned by the same fire. We recorded the cover of all vascular plant species. We also analysed species richness, total cover, proportion of exotic species, abundance of woody species and herb species, cover of exotic species, abundance of woody and herb species and differences in composition of species. For both shrubs and tree species we recorded the main strategy of regeneration (by resprouting or by seed). Results: We found that fire had different effects on native matorral and pine plantations. Five years after fire, plantations came to be dominated by herbs and exotic species, showing differences in floristic composition. In contrast, matorral communities remained very similar to unburned matorral in terms of species richness, proportion of woody species, and herb species and proportion of exotics. Also, pine plantations were primarily colonized by seedlings, while matorrals were primarily colonized by resprouting. Conclusions: Matorrals are highly fire resilient communities, and the practice of establishing plantations on matorrals produces a strong reduction in the capacity of matorral to return to its original state. The elimination of shrubs owing to the effect of plantations can hinder regeneration of native ecosystems. Burned plantations may slowly develop into ecosystems similar to the native ones, or they may produce a new ecosystem dominated by exotic herbs. This study shows that plantations of exotic conifers affect native vegetation even after they have been removed, as in this case by fire.     

Changes in alpine snowbed‐wetland vegetation over three decades in northern Norway

We have quantified floristic changes in alpine snowbeds and wetland vegetation during three decades and analyzed to what extent these changes are related to initial variations in snow cover duration and distance to groundwater level. Vascular plant species richness and total plant cover were estimated along three transects in northern Norway. Three different vegetation zones were identified along the original transects: relatively dry snowbeds, wet snowbeds and wetlands. The resampling shows major changes in species richness and plant cover. In general, there was a net immigration of species and 13 new species were found. Five rare species with initial low cover were lost. In the dry and wet snowbeds, species richness and total plant cover increased, mostly because of invasion by shrubs, graminoids and herbs. A general trend was that species indicating high soil moisture were strongly reduced. In the wetland zones there were no significant floristic changes but hygrophilous species had decreased and were replaced by graminoids and shrub species with lower water requirements. These floristic changes were significantly related to snow and soil moisture conditions, important factors for rate and direction of change. Contrasting vegetation responses within very short distances demonstrate the importance of detailed knowledge of the actual microhabitats when effects of climate change in alpine habitats are considered.     

Water–energy,land‐cover and heterogeneity drivers of the distribution of plant species richness in a mountain region of the European Alps

Aim To evaluate the relative importance of water–energy, land‐cover, environmental heterogeneity and spatial variables on the regional distribution of Red‐Listed and common vascular plant species richness. Location Trento Province (c. 6200 km²) on the southern border of the European Alps (Italy), subdivided regularly into 228 3′ × 5′ quadrants. Methods Data from a floristic inventory were separated into two subsets, representing Red‐Listed and common (i.e. all except Red‐Listed) plant species richness. Both subsets were separately related to water–energy, land‐cover and environmental heterogeneity variables. We simultaneously applied ordinary least squares regression with variation partitioning and hierarchical partitioning, attempting to identify the most important factors controlling species richness. We combined the analysis of environmental variables with a trend surface analysis and a spatial autocorrelation analysis. Results At the regional scale, plant species richness of both Red‐Listed and common species was primarily related to energy availability and land cover, whereas environmental heterogeneity had a lesser effect. The greatest number of species of both subsets was found in quadrants with the largest energy availability and the greatest degree of urbanization. These findings suggest that the elevation range within our study region imposes an energy‐driven control on the distribution of species richness, which resembles that of the broader latitude gradient. Overall, the two species subsets had similar trends concerning the relative importance of water–energy, land cover and environmental heterogeneity, showing a few differences regarding the selection of some predictors of secondary importance. The incorporation of spatial variables did not improve the explanatory power of the environmental models and the high original spatial autocorrelation in the response variables was reduced drastically by including the selected environmental variables. Main conclusions Water–energy and land cover showed significant pure effects in explaining plant species richness, indicating that climate and land cover should both be included as explanatory variables in modelling species richness in human‐affected landscapes. However, the high degree of shared variation between the two groups made the relative effects difficult to separate. The relatively low range of variation in the environmental heterogeneity variables within our sampling domain might have caused the low importance of this complex factor.     

Evaluation of floristic diversity in urban areas as a basis for habitat management

Questions: How can floristic diversity be evaluated in conser‐vation plans to identify sites of highest interest for biodiversity? What are the mechanisms influencing the distribution of species in human‐dominated environments? What are the best criteria to identify sites where active urban management is most likely to enhance floristic diversity? Location: The Hauts‐de‐Seine district bordering Paris, France. Methods: We described the floristic diversity in one of the most urbanized French districts through the inventory of ca. 1000 sites located in 23 habitats. We built a new index of floristic interest (IFI), integrating information on richness, indigeneity, typicality and rarity of species, to identify sites and habitats of highest interest for conservation. Finally, we explored the relationship between site IFI and land use patterns (LUP). Results: We observed a total of 626 vascular plant species. Habitats with highest IFI were typically situated in seminatural environments or environments with moderate human impact. We also showed that neighbouring (urban) structures had a significant influence on the floristic interest of sites: for example, the presence of collective dwellings around a site had a strong negative impact on IFI. Conclusions: Our approach can be used to optimize management in urban zones; we illustrate such possibilities by defining a ‘Site Potential Value’, which was then compared with the observed IFI, to identify areas (e.g. river banks) where better management could improve the district's biodiversity.     

Relationship between topographic heterogeneity and vegetation patterns in a Californian salt marsh

Questions: Are species richness and species abundances higher in the presence of tidal creeks? Do species richness and species abundances vary with plot size? Location: Intertidal plain of Volcano Marsh, Bahia de San Quintin, Mexico. Methods: We analysed vegetation patterns in large areas (cells) with tidal creeks (+creek) and without (‐creek). We surveyed vegetation cover, microtopography, habitat type, and distance to creeks in nested plots of five sizes, 0.1, 0.25, 1, 2.5, and 10 m². Results: Species richness, frequency, cover, and assemblages differed between ±creek cells. Richness tended to be higher in +creek cells, and cover and frequency of individual species differed significantly between ±creek cells. We found consistent patterns in vegetation structure across plot sizes. We encountered 13 species that occurred in 188 unique assemblages. The most common assemblage had six species: Batis maritima, Frankenia salina, Salicornia bigelovii, S. virginica, Salicornia spec. and Triglochin concinna. This assemblage occurred in ±creek cells and at all spatial scales. Of the most common assemblages all but one were composed of multiple species (3–9 species/plot). Conclusions: The persistence of vegetation patterns across a 100‐fold range in spatial scale suggests that similar environmental factors operate broadly to determine species establishment and persistence. Differences in assemblage composition result from variation of frequency and cover of marsh plain species, particularly Suaeda esteroa and Monanthochloe littoralis. The recommendation for restoration of Californian salt marshes is to target (and plant) multi‐species assemblages, not monocultures.     

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.     

Seed bank,seed dispersal and vegetation cover: Colonization along a newly‐created river channel

Question : What is the relative importance of the initial seed bank and subsequent seed dispersal for floristic composition of bank vegetation two years after creation of a newly‐cut reach of a river channel? Location : River Cole, West Midlands, United Kingdom. Methods : We took bank and bed sediment samples from a 0.5‐km reach of a new river channel cut into intact flood‐plain. After river diversion, seed samples deposited on artificial turf mats placed on the river banks and flood‐plain edge were taken in summer and winter 2002 and 2003. Seed rain samples from funnel traps were taken during summer 2002 and 2003. We undertook greenhouse germination trials to assess viable seed species within these samples. In summer 2004, we surveyed river bank vegetation. Agglomerative cluster analysis was used to investigate floristic similarity between seed bank, seed rain, seed deposition samples and final bank vegetation cover. DCA was used to explore contrasts between the samples and to assess whether these reflected interpretable environmental gradients. Results : Seed rain samples contained a small subset of species in the summer depositional samples. 38 species were found within the final vegetation, the seed bank, and at least one of the four sets of depositional samples; a further 30 species not present in the seed‐bank samples were present in at least one of the four sets of depositional samples and the final vegetation. Floristic composition of the vegetation was most similar to the depositional samples from winter 2002 and 2003 and summer 2003. DCA axis 1 reflected a time sequence from seed‐bank samples through depositional samples to the final vegetation. Conclusions : Newly cut river banks were colonized rapidly. Seed remobilization and hydrochorous transport from the upstream catchment are important for colonization. Species richness was highest in samples deposited during winter when high river flows can remobilize and transport viable seeds from upstream. This process would also have enhanced the species richness of seed production along the banks during the second summer (2003).     

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.