A preliminary account of patterns of endemism in Namibia's Sperrgebiet – the succulent karoo

Aim This study investigated spatial patterns of endemism in the flora of Namibia's succulent karoo in order to generate information for conservation planning. Location The study area, the Sperrgebiet, comprises the majority of Namibia's portion of the succulent karoo biome which is the south‐west corner of the country. This is an arid area that has been off limits to public access, farming and tourism for nearly a century due to restrictions imposed by the diamond industry. Methods Based on existing distribution records, areas of high concentrations of endemic plants were identified using numbers of endemics and weighted endemics according to area of occupancy. The resolution of the available data was quarter degree squares (15‐min intervals of latitude and longitude grids). Results At the scale of this study straight numbers of endemics generated similar results to the endemics weighted according to area of occupancy, which gives sparsely distributed species a higher weighting. Based on the current distribution records, 17.7% (184 species) of the Sperrgebiet's spermatophyte flora is endemic. The ‘hotspots of endemism’ comprised from north to south: Lüderitz‐Kowisberge, Klinghardt Mountains, Aurusberge‐Heioab, Witpütz, Skorpion and Obib‐Schakalsberge. Taking also areas into account that stand out because of their high proportion of local endemics, this adds Grillental and the central coastal area from Pomona to Baker's Bay to the areas of importance for plant endemism. Main conclusions The Sperrgebiet's endemic flora is special in taxonomic composition in that it does not present a subset of the total flora of this area, but shows a remarkably high representation of the families Mesembryanthemaceae and Liliaceae (sensu lato). Compared with other arid areas, the level of endemism in the Sperrgebiet is high, but not compared with the succulent karoo in general or other hotspots in the succulent karoo biome, such as the Richtersveld. The proportion of local endemics (13.5%) is high compared with some endemism hotspots in southern Africa. Hotspots of plant endemism provide an important tool to contribute to conservation planning studies. This study also highlighted the importance of centralized data bases without which these analyses would not have been possible. Further plant collecting is required to fill presently data‐deficient areas and studies at a finer spatial resolution taking habitat requirements into account are needed to elucidate some of the factors contributing to plant endemism in this area.     

ORIGINS AND CONSEQUENCES OF SERPENTINE ENDEMISM IN THE CALIFORNIA FLORA

Habitat specialization plays an important role in the creation and loss of biodiversity over ecological and evolutionary time scales. In California, serpentine soils have a distinctive flora, with 246 serpentine habitat specialists (i.e., endemics). Using molecular phylogenies for 23 genera containing 784 taxa and 51 endemics, we infer few transitions out of the endemic state, which is shown by an analysis of transition rates to simply reflect the low frequency of endemics (i.e., reversal rates were high). The finding of high reversal rates, but a low number of reversals, is consistent with the widely hypothesized trade‐off between serpentine tolerance and competitive ability, under which serpentine endemics are physiologically capable of growing in less‐stressful habitats but competitors lead to their extirpation. Endemism is also characterized by a decrease in speciation and extinction rates and a decrease in the overall diversification rate. We also find that tolerators (species with nonserpentine and serpentine populations) undergo speciation in serpentine habitats to give rise to new serpentine endemics but are several times more likely to lose serpentine populations to produce serpentine‐intolerant taxa. Finally, endemics were younger on average than nonendemics, but this alone does not explain their low diversification.     

Environmental heterogeneity,species diversity and co‐existence at different spatial scales

The positive relationship between spatial environmental heterogeneity and species diversity is a widely accepted concept, generally associated with niche limitation. However, niche limitation cannot account for negative heterogeneity–diversity relationships (HDR) revealed in several case studies. Here we explore how HDR varies at different spatial scales and provide novel theories for small‐scale species co‐existence that explain both positive and negative HDR. At large spatial scales of heterogeneity (e.g. landscape level), different communities co‐exist, promoting large regional species pool size and resulting in positive HDR. At smaller scales within communities, species co‐existence can be enhanced by increasing the number of different patches, as predicted by the niche limitation theory, or alternatively, restrained by heterogeneity. We conducted meta‐regressions for experimental and observational HDR studies, and found that negative HDRs are significantly more common at smaller spatial scales. We propose three theories to account for niche limitation at small spatial scales. (1) Microfragmentation theory: with increasing spatial heterogeneity, large homogeneous patches lose area and become isolated, which in turn restrains the establishment of new plant individuals and populations, thus reducing species richness. (2) Heterogeneity confounded by mean: when heterogeneity occurs at spatial scales smaller than the size of individual plants, which forage through the patches, species diversity can be either positively or negatively affected by a change in the mean of an environmental factor. (3) Heterogeneity as a separate niche axis: the ability of species to tolerate heterogeneity at spatial scales smaller than plant size varies, affecting HDR. We conclude that processes other than niche limitation can affect the relationship between heterogeneity and diversity.     

Community- Weighted Mean Plant Traits Predict Small Scale Distribution of Insect Root Herbivore Abundance

Small scale distribution of insect root herbivores may promote plant species diversity by creating patches of different herbivore pressure. However, determinants of small scale distribution of insect root herbivores, and impact of land use intensity on their small scale distribution are largely unknown. We sampled insect root herbivores and measured vegetation parameters and soil water content along transects in grasslands of different management intensity in three regions in Germany. We calculated community-weighted mean plant traits to test whether the functional plant community composition determines the small scale distribution of insect root herbivores. To analyze spatial patterns in plant species and trait composition and insect root herbivore abundance we computed Mantel correlograms. Insect root herbivores mainly comprised click beetle (Coleoptera, Elateridae) larvae (43%) in the investigated grasslands. Total insect root herbivore numbers were positively related to community-weighted mean traits indicating high plant growth rates and biomass (specific leaf area, reproductive- and vegetative plant height), and negatively related to plant traits indicating poor tissue quality (leaf C/N ratio). Generalist Elaterid larvae, when analyzed independently, were also positively related to high plant growth rates and furthermore to root dry mass, but were not related to tissue quality. Insect root herbivore numbers were not related to plant cover, plant species richness and soil water content. Plant species composition and to a lesser extent plant trait composition displayed spatial autocorrelation, which was not influenced by land use intensity. Insect root herbivore abundance was not spatially autocorrelated. We conclude that in semi-natural grasslands with a high share of generalist insect root herbivores, insect root herbivores affiliate with large, fast growing plants, presumably because of availability of high quantities of food. Affiliation of insect root herbivores with large, fast growing plants may counteract dominance of those species, thus promoting plant diversity.     

Multiscale Patterns of Riparian Plant Diversity and Implications for Restoration

Planning riparian restoration to resemble historic reference conditions requires an understanding of both local and regional patterns of plant species diversity. Thus, understanding species distributions at multiple spatial scales is essential to improve restoration planting success, to enhance long‐term ecosystem functioning, and to match restoration planting designs with historic biogeographic distributions. To inform restoration planning, we examined the biogeographic patterns of riparian plant diversity at local and regional scales within a major western U.S.A. drainage, California's Sacramento—San Joaquin Valley. We analyzed patterns of species richness and complementarity (β‐diversity) across two scales: the watershed scale and the floodplain scale. At the watershed scale, spatial patterns of native riparian richness were driven by herbaceous species, whereas woody species were largely cosmopolitan across the nearly 38,000 km² study area. At the floodplain scale, riparian floras reflected species richness and dissimilarity patterns related to hydrological and disturbance‐driven successional sequences. These findings reinforce the importance of concurrently evaluating both local and regional processes that promote species diversity and distribution of native riparian flora. Furthermore, as restoration activities become more prevalent across the landscape, strategies for restoration outcomes should emulate the patterns of species diversity and biogeographic distributions found at regional scales.     

Distribution of the endemic Balkan flora on serpentine I. – obligate serpentine endemics

Serpentine (ophiolithic) substrate covers large areas in the Balkans, more so than in any other part of Europe. These areas extend from north to south mainly in the mountainous central regions and represent specialized habitats for basiphilous-calcifugal plants. Biodiversity in the area is high, with a great number of interesting local and regional endemics. The high number of endemics indicates the importance of serpentine habitats as centres for floristic differentiation and speciation. The number of Balkan endemics growing on serpentine is c. 335 taxa (species and subspecies) of which 123 are obligate. Their distribution is presented in 50 × 50 km UTM squares as adopted in the Atlas Florae Europaeae project coordinated at Helsinki. The richest (in number of taxa) squares are situated in NW Greece (Epirus), the island of Evvia, N Albania together with SW Serbia, and N Greece (Vourinos). They indicate important centres of plant diversity in the Balkans, areas to be noted for conservation strategy. Features responsible for the distribution and abundance of these obligate serpentine endemics include: 1) edaphic isolation in relation to type of bedrock (lime, dolomite, marble, schist, etc.), 2) mountain island isolation (Smolikas, Vourinos, Ostrovica, etc.), 3) island isolation (Evvia) and 4) continuous long-term isolation without interruption or disturbance of speciation.     

Non‐interacting impacts of fertilization and habitat area on plant diversity via contrasting assembly mechanisms

Aim

The local‐ and regional‐based forms of anthropogenic change reducing grassland diversity are generally identified, but these scale‐dependent processes tend to co‐occur with unclear interactive effects. Here, we explicitly test how common local and regional perturbations simultaneously affect plant alpha and beta diversity in a multiyear community assembly experiment using fragments of grassland habitat of various sizes. We hypothesized that local disturbances and decreasing patch size would interact, suppressing local diversity while homogenizing composition among patches.

Location

North America.

Methods

We conducted a three‐year grassland assembly experiment, factorially manipulating local perturbation (nitrogen addition and mowing) and patch area for 36 patches over 13 ha. We quantified the individual and interactive effects of these local and regional factors on plant alpha and beta diversity within (quadrat scale) and among patches (patch scale). We also used a null model approach to disentangle between stochastic‐ and niche‐based assembly mechanisms.

Results

We detected a gradient of assembly outcomes driven by two non‐interacting factors—the effects of N fertilization on alpha (negative) and beta (positive) diversity regardless of spatial scale and the scale‐dependant effect of increasing patch size on alpha (positive) and beta (positive) diversity. These effects unfolded over time, with the constraints on richness and composition shifting from dispersal‐based during the first sampling year to perturbation‐and size‐based factors at year two and three. Fertilization effects were driven by a mixture of deterministic (i.e., selection at the species level) and stochastic (i.e., random extinctions) processes resulting in a decline in local richness but an increase in spatial heterogeneity in species composition. Area appeared to influence alpha diversity mainly via stochastic “sampling effect”—larger patches represented a larger sample of the regional pool. Niche‐based processes, however, led to convergence in beta diversity among smaller patches driving a positive overall effect of area on beta diversity.

Main conclusion

Our results illustrate how diversity regulation in contemporary grasslands can be simultaneously shaped by local and regional factors acting additively but via contrasting assembly mechanisms that operate at different spatial and temporal scales.

     

Facilitation allows plant coexistence in Cuban serpentine soils

Serpentine soils represent stressful habitats where plants have to cope with heavy metals, moisture limitation and low nutrient availability. We propose that facilitation is an important mechanism structuring plant communities under such stressful conditions. Facilitation has been shown to generate the spatial association of species, forming discrete vegetation patches of phylogenetically distant species. We measured these spatial and phylogenetic signatures left by facilitation in a serpentine plant community of central Cuba. Our results show that seedlings preferentially grow under plants of different species, and that adults are significantly aggregated into vegetation patches. In these patches, adults tend to co‐occur with distant relatives, ultimately forming phylogenetically diverse neighbourhoods. We discuss possible mechanisms explaining how species adapted to serpentine areas may be acting as nurses, reducing the stressful conditions for the establishment of other species.     

Plant genetic diversity in the Canary Islands: a conservation perspective

The Canary Islands are an Atlantic volcanic archipelago with a rich flora of ～570 endemic species. The endemics represent ～40% of the native flora of the islands, and ～20% of the endemics are in the E (endangered) category of the International Union for Conservation of Nature. A review of allozyme variation in 69 endemic species belonging to 18 genera and eight families is presented. The average species-level genetic diversity (H(T)) at allozyme loci is 0.186, which is twice as high as the mean reported for endemics of Pacific archipelagos. Possible factors contributing to this higher diversity are discussed, but the reasons remain obscure. An average of 28% of the allozyme diversity within species resides among populations, indicating a high level of interpopulational differentiation. Studies of reproductive biology indicate that many of the endemic species are outcrossers. The high total diversity within species, the relatively high differentiation among populations, and the outcrossing breeding systems have implications for species conservation. Decreased population sizes in outcrossing species would promote biparental inbreeding and increase inbreeding depression. The relatively high proportion of allozyme diversity among populations indicates that the most effective strategy for preserving genetic variation in species is to conserve as many populations as possible. The genetic diversity in many Canary Island endemics is endangered by: (1) overgrazing by introduced animals, such as barbary sheep, goats, mouflons, rabbits, and sheep; (2) interspecific hybridization following habitat disturbance or planting of endemics along roadsides or in public gardens; (3) competition with alien plant species; and (4) decline of population size because of urban development and farming.     

Trade‐offs between savanna woody plant diversity and carbon storage in the Brazilian Cerrado

Incentivizing carbon storage can be a win‐win pathway to conserving biodiversity and mitigating climate change. In savannas, however, the situation is more complex. Promoting carbon storage through woody encroachment may reduce plant diversity of savanna endemics, even as the diversity of encroaching forest species increases. This trade‐off has important implications for the management of biodiversity and carbon in savanna habitats, but has rarely been evaluated empirically. We quantified the nature of carbon‐diversity relationships in the Brazilian Cerrado by analyzing how woody plant species richness changed with carbon storage in 206 sites across the 2.2 million km² region at two spatial scales. We show that total woody plant species diversity increases with carbon storage, as expected, but that the richness of endemic savanna woody plant species declines with carbon storage both at the local scale, as woody biomass accumulates within plots, and at the landscape scale, as forest replaces savanna. The sharpest trade‐offs between carbon storage and savanna diversity occurred at the early stages of carbon accumulation at the local scale but the final stages of forest encroachment at the landscape scale. Furthermore, the loss of savanna species quickens in the final stages of forest encroachment, and beyond a point, savanna species losses outpace forest species gains with increasing carbon accumulation. Our results suggest that although woody encroachment in savanna ecosystems may provide substantial carbon benefits, it comes at the rapidly accruing cost of woody plant species adapted to the open savanna environment. Moreover, the dependence of carbon‐diversity trade‐offs on the amount of savanna area remaining requires land managers to carefully consider local conditions. Widespread woody encroachment in both Australian and African savannas and grasslands may present similar threats to biodiversity.     

TRADE‐OFFS,SPATIAL HETEROGENEITY,AND THE MAINTENANCE OF MICROBIAL DIVERSITY

Specialization and concomitant trade‐offs are assumed to underlie the non‐neutral coexistence of lineages. Trade‐offs across heterogeneous environments can promote diversity by preventing competitive exclusion. However, the importance of trade‐offs in maintaining diversity in natural microbial assemblages is unclear, as trade‐offs are frequently not detected in artificial evolution experiments. Stressful conditions associated with patches of heavy‐metal enriched serpentine soils provide excellent opportunities for examining how heterogeneity may foster genetic diversity. Using a spatially replicated design, we demonstrate that rhizobium bacteria symbiotic with legumes inhabiting contrasting serpentine and nonserpentine soils exhibit a trade‐off between a genotype's nickel tolerance and its ability to replicate rapidly. Furthermore, we detected adaptive divergence in rhizobial assemblages across soil type heterogeneity at multiple sites, suggesting that this trade‐off may promote the coexistence of phenotypically distinct bacterial lineages. Trade‐offs and adaptive divergence may be important factors maintaining the tremendous diversity within natural assemblages of bacteria.     

Linking the spatial scale of environmental variation and the evolution of phenotypic plasticity: selection favors adaptive plasticity in fine-grained environments

Adaptive phenotypic plasticity and adaptive genetic differentiation enable plant lineages to maximize their fitness in response to environmental heterogeneity. The spatial scale of environmental variation relative to the average dispersal distance of a species determines whether selection will favor plasticity, local adaptation, or an intermediate strategy. Habitats where the spatial scale of environmental variation is less than the dispersal distance of a species are fine grained and should favor the expression of adaptive plasticity, while coarse-grained habitats, where environmental variation occurs on spatial scales greater than dispersal, should favor adaptive genetic differentiation. However, there is relatively little information available characterizing the link between the spatial scale of environmental variation and patterns of selection on plasticity measured in the field. I examined patterns of spatial environmental variation within a serpentine mosaic grassland and selection on an annual plant (Erodium cicutarium) within that landscape. Results indicate that serpentine soil patches are a significantly finer-grained habitat than non-serpentine patches. Additionally, selection generally favored increased plasticity on serpentine soils and diminished plasticity on non-serpentine soils. This is the first empirical example of differential selection for phenotypic plasticity in the field as a result of strong differences in the grain of environmental heterogeneity within habitats.     

Habitat diversity associated to island size and environmental filtering control the species richness of rock‐savanna plants in neotropical inselbergs

Disentangling the multiple factors controlling species diversity is a major challenge in ecology. Island biogeography and environmental filtering are two influential theories emphasizing respectively island size and isolation, and the abiotic environment, as key drivers of species richness. However, few attempts have been made to quantify their relative importance and investigate their mechanistic basis. Here, we applied structural equation modelling, a powerful method allowing test of complex hypotheses involving multiple and indirect effects, on an island‐like system of 22 French Guianan neotropical inselbergs covered with rock‐savanna. We separated the effects of size (rock‐savanna area), isolation (density of surrounding inselbergs), environmental filtering (rainfall, altitude) and dispersal filtering (forest‐matrix openness) on the species richness of all plants and of various ecological groups (terrestrial versus epiphytic, small‐scale versus large‐scale dispersal species). We showed that the species richness of all plants and terrestrial species was mainly explained by the size of rock‐savanna vegetation patches, with increasing richness associated with higher rock‐savanna area, while inselberg isolation and forest‐matrix openness had no measurable effect. This size effect was mediated by an increase in terrestrial‐habitat diversity, even after accounting for increased sampling effort. The richness of epiphytic species was mainly explained by environmental filtering, with a positive effect of rainfall and altitude, but also by a positive size effect mediated by enhanced woody‐plant species richness. Inselberg size and environmental filtering both explained the richness of small‐scale and large‐scale dispersal species, but these ecological groups responded in opposite directions to altitude and rainfall, that is positively for large‐scale and negatively for small‐scale dispersal species. Our study revealed both habitat diversity associated with island size and environmental filtering as major drivers of neotropical inselberg plant diversity and showed the importance of plant species growth form and dispersal ability to explain the relative importance of each driver.     

Patterns of resource tracking by avian frugivores at multiple spatial scales: two case studies on discordance among scales

Scaling is relevant for the analysis of plant‐frugivore interaction, since the ecological and evolutionary outcomes of seed dispersal depend on the spatial and temporal scale at which frugivory patterns emerge. We analyse the relationship between fruit abundance and frugivore activity at local and landscape spatial scales in two different systems composed, respectively, by the bird‐dispersed woody plants Juniperus communis and Bursera fagaroides, and their frugivore assemblages. We use a hierarchical approach of nested patchiness of fruit‐resource, where patches are defined by individual plants within site, at the local scale, and by sites within region, at the landscape scale. The structure of patches is also described in terms of contrast (differences in fruit availability among patches) and aggregation (spatial distribution of patches). For J. communis, frugivore activity was positively related to fruit availability at the landscape scale, this pattern seldom emerging at the local scale; conversely, B. fagaroides showed a general trend of positive local pattern that disappeared at the landscape scale. These particular trends might be partially explained by differences in contrast and aggregation. The strong contrast among plants within site together with a high aggregation among sites would promote the B. fagaroides pattern to be only local, whereas in J. communis, low aggregation among sites within region would favour a sharp landscape‐scale pattern. Both systems showed discordant patterns of fruit‐resource tracking among consecutive spatial scales, but the sense of discordance differed among systems. These results, and the available multi‐scale frugivory data, suggest that discordance among successive scales allows to link directly frugivory patterns to resource‐tracking mechanisms acting at particular scales, resulting, thus, more informative than concordance observational data, in which landscape patterns might result from accumulated effect of local mechanisms. In this context, we propose new methodological approaches for a better understanding of the hierarchical behavioural mechanisms underpinning the multi‐scale resource tracking by frugivores.     

Natural plant species inventory of hotspot areas in Arabian Peninsula: Southwest Al-Baha region,Saudi Arabia

Plant hotspot areas are the areas that are very rich in plant species diversity. These areas have a priority for conservation. To highlight the plant diversity for nature conservation purposes a case study in Al Baha region, Saudi Arabia is presented, in which the importance of the natural vegetation and flora of one of the hotspot areas of Saudi Arabia is evaluated through the explanation of its natural plant species. A survey study has been conducted in an area of 167.6 km2, a 97 sample each with 20X20 m were laid out covering the whole ecological zones of the study site. Data of flora, vegetation cover and topography were gathered from each sample site. The study revealed about 319 plant species belonging to 228 genera and 75 families. Two species were found endemic to Saudi Arabia, 14 were endemics to Arabian Peninsula, and five were regional endemics that are only found in East Africa and Arabian Peninsula, while 39 species are rare and endangered.     

Local variation in plant quality influences large‐scale population dynamics

Spatial variation in ecological systems can arise both as a consequence of variation in the quality and availability of resources and as an emergent property of spatially structured interactions. We used a spatially explicit model to simulate populations of herbivore hosts and their parasitoids in landscapes with different levels of variance in plant patch quality and different spatial arrangements of high‐ and low‐quality plant patches. We found that even small variation in patch quality at a fine spatial scale decreased overall herbivore populations, as parasitoid populations on low‐quality plant patches were subsidized by those from high‐quality neighbors. On landscapes with large, homogeneous regions of high‐ and low‐quality plant patches, herbivore populations increased with variation in patch quality. Overall, our results demonstrate that local variation in resource quality profoundly influences global population dynamics. In particular, fine‐scale variation in plant patch quality enhanced biological control of herbivores by parasitoids, suggesting that adding back plant genetic variation into perennial production systems may enhance the biological control of herbivores by their natural enemies.     

云南地区种子植物多样性分布格局和多样性分化特点研究

利用大尺度的植物分布信息结合地理信息系统和数理统计分析,探讨了云南种子植物多样性的地理分布格局和多样性分化特点。结果表明,从科到属和种,植物多样性和区系分化强度的数据集离散程度均呈递增趋势。相对属和种而言,科的多样性的空间分异和高值区域并不明显,这可能与科、属、种所代表的不同地质历史时期有关;南部地区属和种的多样性明显高于除滇西北以外的大部分地区。种的多样性分布中心主要集中在滇西北地区、云南南部地区、哀牢山、无量山地区和昆明地区,这可能与生境异质性和热量条件有关。相对科而言,属的区系分化强度的空间分异更为明显,并与物种多样性的分布格局比较接近,暗示着植物多样性与区系分化之间的密切联系。相对科而言,属的区系分化中心高值区域更为明显。总体上,云南地区植物多样性的分布格局及其分化可能与其生境、热量条件和地质历史有关。     

Effect of habitat area and isolation on plant trait distribution in European forests and grasslands

A number of studies show contrasting results in how plant species with specific life‐history strategies respond to fragmentation, but a general analysis on whether traits affect plant species occurrences in relation to habitat area and isolation has not been performed. We used published data from forests and grasslands in north‐central Europe to analyse if there are general patterns of sensitivity to isolation and dependency of area for species using three traits: life‐span, clonality, and seed weight. We show that a larger share of all forest species was affected by habitat isolation and area as compared to grassland species. Persistence‐related traits, life‐span and clonality, were associated to habitat area and the dispersal and recruitment related trait, seed weight, to isolation in both forest and grassland patches. Occurrence of clonal plant species decreased with habitat area, opposite to non‐clonal plant species, and long‐lived plant species decreased with grassland area. The directions of these responses partly challenge some earlier views, suggesting that further decrease in habitat area will lead to a change in plant species community composition, towards relatively fewer clonal and long‐lived plants with large seeds in small forest patches and fewer clonal plants with small seeds in small grassland patches. It is likely that this altered community has been reached in many fragmented European landscapes consisting of small and isolated natural and semi‐natural patches, where many non‐clonal and short‐lived species have already disappeared. Our study based on a large‐scale dataset reveals general and useful insights concerning area and isolation effects on plant species composition that can improve the outcome of conservation and restoration efforts of plant communities in rural landscapes.     

Relationships between spatial configuration of tropical forest patches and woody plant diversity in northeastern Puerto Rico

The destruction and fragmentation of tropical forests are major sources of global biodiversity loss. A better understanding of anthropogenically altered landscapes and their relationships with species diversity and composition is needed in order to protect biodiversity in these environments. The spatial patterns of a landscape may control the ecological processes that shape species diversity and composition. However, there is little information about how plant diversity varies with the spatial configuration of forest patches especially in fragmented tropical habitats. The northeastern part of Puerto Rico provides the opportunity to study the relationships between species richness and composition of woody plants (shrubs and trees) and spatial variables [i.e., patch area and shape, patch isolation, connectivity, and distance to the Luquillo Experimental Forest (LEF)] in tropical forest patches that have regenerated from pasturelands. The spatial data were obtained from aerial color photographs from year 2000. Each photo interpretation was digitized into a GIS package, and 12 forest patches (24–34 years old) were selected within a study area of 28 km². The woody plant species composition of the patches was determined by a systematic floristic survey. The species diversity (Shannon index) and species richness of woody plants correlated positively with the area and the shape of the forest patch. Larger patches, and patches with more habitat edge or convolution, provided conditions for a higher diversity of woody plants. Moreover, the distance of the forest patches to the LEF, which is a source of propagules, correlated negatively with species richness. Plant species composition was also related to patch size and shape and distance to the LEF. These results indicate that there is a link between landscape structure and species diversity and composition and that patches that have similar area, shape, and distance to the LEF provide similar conditions for the existence of a particular plant community. In addition, forest patches that were closer together had more similarity in woody plant species composition than patches that were farther apart, suggesting that seed dispersal for some species is limited at the scale of 10 km.     

Reduced vertebrate diversity independent of spatial scale following feral swine invasions

Biological invasions often have contrasting consequences with reports of invasions decreasing diversity at small scales and facilitating diversity at large scales. Thus, previous literature has concluded that invasions have a fundamental spatial scale‐dependent relationship with diversity. Whether the scale‐dependent effects apply to vertebrate invaders is questionable because studies consistently report that vertebrate invasions produce different outcomes than plant or invertebrate invasions. Namely, vertebrate invasions generally have a larger effect size on species richness and vertebrate invaders commonly cause extinction, whereas extinctions are rare following invertebrate or plant invasions. In an agroecosystem invaded by a non‐native ungulate (i.e., feral swine, Sus scrofa), we monitored species richness of native vertebrates in forest fragments ranging across four orders of magnitude in area. We tested three predictions of the scale‐dependence hypothesis: (a) Vertebrate species richness would positively increase with area, (b) the species richness y‐intercept would be lower when invaded, and (c) the rate of native species accumulation with area would be steeper when invaded. Indeed, native vertebrate richness increased with area and the species richness was 26% lower than should be expected when the invasive ungulate was present. However, there was no evidence that the relationship was scale dependent. Our data indicate the scale‐dependent effect of biological invasions may not apply to vertebrate invasions.