Ecology and Distribution of Melastomataceae in African Rain Forest Inselbergs1

The ecology and distribution of thirty‐seven Melastomataceae species found on 29 inselbergs in the rain forest of Equatorial Guinea, Gabon, and Cameroon are described. The relative contributions of climatic determinism, and stochastic and historical processes to the present distribution of the species amongst the study sites are investigated. Inselbergs are rock outcrops enduring very harsh edaphic and microclimatic conditions, forming “xeric islands” in the rain forest, where natural grasslands develop. For each Melastomataceae species, the habitats where they were collected on the inselbergs and in other biotopes were listed, and their phytogeographical distribution was established. Three species are restricted to rock outcrops. Most species are distributed inside the Guineo‐Congolian Region. Three species have disjunct distributions on rain forest inselbergs and have their main distribution area outside the forested zone. The distribution of the species amongst the inselbergs is correlated to the altitude and the annual rainfall of the sites, and to inselberg geographical location. This correlation of the species distribution with the geographical location of the inselberg is attributed to the spatial autocorrelation of environmental variables and to the combined effects of the history of past vegetation changes due to climatic fluctuations and dispersal limitation.     

Biodiversity and vegetation of small-sized inselbergs in a West African rain forest (Taı, Ivory Coast)

The vegetation of small granitic rock outcrops (geomorphologically small-sized inselbergs) which do not reach the canopy was studied in the Taı rain forest (southwestern Ivory Coast) under aspects of species diversity and phytogeographical affinities. Rock outcrops form edaphically arid (due to absent or very sparse soil cover) and microclimatologically xeric (i.e. low air humidity, temperature regularly exceeding 50°C) islands with cryptogamic crusts, succulents and poikilohydric vascular plants as characteristic elements of their vegetation which differs totally from the surrounding forest. Altogether sixty-six species of vascular plants out of twenty-nine families occur, the number of species correlates positively with inselberg size. Compared with large inselbergs the microclimatic attributes of small-sized rock outcrops are less pronounced. This is accompanied by a decrease of typical inselberg taxa (i.e. species mainly occurring on inselbergs). Low beta diversity between inselbergs indicates deterministic influences as important regulators of species composition. Annual Poaceae and Cyperaceae are richly represented. It can be hypothesized that inselbergs may represent natural growing sites of widely distributed tropical weeds today. Inselbergs might provide habitat resources for savanna elements in rain forest zones.     

The role of Namibian inselbergs in contributing to local and regional plant species richness

This study investigated four inselberg landscapes in Namibia's aridNama Karoo. Inselbergs of different size and geology were investigated (a) todetermine environmental variables influencing the number of inselberg restrictedplant species and (b) to investigate seed dispersal spectra as one parameterdescribing functional properties of inselberg specialists. With regard to therole of inselbergs in contributing to local and regional species richness, manyspecies restricted to inselberg habitats occurred (up to 79 species recorded inthis survey) and so contribute to local and regional species richness. As nospecies endemic to an inselberg were recorded in this study, inselbergspecialists may provide an indicator for conservation value of these mountains.Elevation rather than surface area provided a measure for predicting the numberof inselberg specialists. Besides elevation, underlying geology, and resultinglandforms and edaphic conditions, as well as distance to potential mainland alsoinfluenced the number of inselberg specialists. Functional relationships betweeninselbergs based on seed dispersal spectra of inselberg specialists were notconclusive.     

High species diversity and turnover in granite inselberg floras highlight the need for a conservation strategy protecting many outcrops

Determining patterns of plant diversity on granite inselbergs is an important task for conservation biogeography due to mounting threats. However, beyond the tropics there are relatively few quantitative studies of floristic diversity, or consideration of these patterns and their environmental, biogeographic, and historical correlates for conservation. We sought to contribute broader understanding of global patterns of species diversity on granite inselbergs and inform biodiversity conservation in the globally significant Southwest Australian Floristic Region (SWAFR). We surveyed floristics from 16 inselbergs (478 plots) across the climate gradient of the SWAFR stratified into three major habitats on each outcrop. We recorded 1,060 species from 92 families. At the plot level, local soil and topographic variables affecting aridity were correlated with species richness in herbaceous (HO) and woody vegetation (WO) of soil‐filled depressions, but not in woody vegetation on deeper soils at the base of outcrops (WOB). At the outcrop level, bioclimatic variables affecting aridity were correlated with species richness in two habitats (WO and WOB) but, contrary to predictions from island biogeography, were not correlated with inselberg area and isolation in any of the three habitats. Species turnover in each of the three habitats was also influenced by aridity, being correlated with bioclimatic variables and with interplot geographic distance, and for HO and WO habitats with local site variables. At the outcrop level, species replacement was the dominant component of species turnover in each of the three habitats, consistent with expectations for long‐term stable landscapes. Our results therefore highlight high species diversity and turnover associated with granite outcrop flora. Hence, effective conservation strategies will need to focus on protecting multiple inselbergs across the entire climate gradient of the region.     

The inselberg flora of Atlantic Central Africa. I. Determinants of species assemblages

Aims To identify the relative contributions of environmental determinism, dispersal limitation and historical factors in the spatial structure of the floristic data of inselbergs at the local and regional scales, and to test if the extent of species spatial aggregation is related to dispersal abilities. Location Rain forest inselbergs of Equatorial Guinea, northern Gabon and southern Cameroon (western central Africa). Methods We use phytosociological relevés and herbarium collections obtained from 27 inselbergs using a stratified sampling scheme considering six plant formations. Data analysis focused on Rubiaceae, Orchidaceae, Melastomataceae, Poaceae, Commelinaceae, Acanthaceae, Begoniaceae and Pteridophytes. Data were investigated using ordination methods (detrended correspondence analysis, DCA; canonical correspondence analysis, CCA), Sørensen's coefficient of similarity and spatial autocorrelation statistics. Comparisons were made at the local and regional scales using ordinations of life‐form spectra and ordinations of species data. Results At the local scale, the forest‐inselberg ecotone is the main gradient structuring the floristic data. At the regional scale, this is still the main gradient in the ordination of life‐form spectra, but other factors become predominant in analyses of species assemblages. CCA identified three environmental variables explaining a significant part of the variation in floristic data. Spatial autocorrelation analyses showed that both the flora and the environmental factors are spatially autocorrelated: the similarity of species compositions within plant formations decreasing approximately linearly with the logarithm of the spatial distance. The extent of species distribution was correlated with their a priori dispersal abilities as assessed by their diaspore types. Main conclusions At a local scale, species composition is best explained by a continuous cline of edaphic conditions along the forest‐inselberg ecotone, generating a wide array of ecological niches. At a regional scale, these ecological niches are occupied by different species depending on the available local species pool. These subregional species pools probably result from varying environmental conditions, dispersal limitation and the history of past vegetation changes due to climatic fluctuations.     

Patterns of bird species richness and composition on islands off Arnhem Land,Northern Territory,Australia

The bird faunas of the adjacent Wessel and English Company island chains were sampled at two scales (0.25 ha quadrats and entire islands). Ninety‐six species were recorded from 226 quadrats, with the most frequently recorded species being mistletoebird Dicaeum hirundinaceum, brown honeyeater Lichmera indistincta, silver‐crowned friarbird Philemon argenticeps, bar‐shouldered dove Geopelia humeralis, northern fantail Rhipidura rufiventris and yellow white‐eye Zosterops lutea. At the quadrat scale, vegetation type was a major determinant of the abundance of individual species (and hence species composition), species richness and total bird abundance. Bird species composition and richness at the quadrat scale was also significantly affected by island isolation (particularly the amount of land within 20 km of the island perimeter). Island size had no effect on quadrat‐scale richness or total abundance. However, the abundance of 10 of the 38 most frequently recorded individual species was significantly related to island size, in most cases even when the comparison was restricted to similar habitats. The most striking cases were rufous fantail Rhipidura rufifrons, mangrove golden whistler Pachycephala melanura, brown honeyeater and yellow white‐eye, which were all significantly more abundant on smaller islands. One hundred and seventy‐one species were recorded from the 62 islands sampled. There was a very tight relationship between island size and the number of terrestrial species (73% of deviance explained) and of all species (84% of deviance explained). This relationship was improved (marginally) when isolation was included in the model. Ordination of islands by their terrestrial bird species composition was related to island size and isolation, and suggested an erratic species composition on small islands.     

High plant taxonomic beta diversity and functional and phylogenetic convergence between two Neotropical inselbergs

ABSTRACT

Background: Inselbergs (granitic and gneissic rock outcrops) are common elements in the Atlantic Forest and present large taxonomic (TD), functional (FD) and phylogenetic (PD) diversity.

Aims: We investigated how plant diversity changed across ecological and biogeographic scales by comparing TD, FD and PD of communities within and between two inselbergs. We expected converging FD and PD but distinct TD between outcrops, because of similar local environmental conditions in inselbergs and the long-term lineage isolation.

Methods: We calculated TD, PD and FD, and partitioned diversity into α (each inselberg), β (between inselbergs) and γ (whole sample) components. Phylogenetic signal was estimated for all traits. To link environmental predictors to functional traits a redundancy analysis was run. Variation in TD, FD and PD was analysed by general linear models with patch area and the two inselbergs as predictors.

Results: The inselbergs were taxonomically different, but showed convergence in their functional and phylogenetic diversity. The limited retention of phylogenetic signal suggests that different species may converge and respond similarly to environmental variables. Within inselbergs, larger patches displayed higher TD, FD and PD.

Conclusions: Seeking conservation strategies for inselbergs is challenging since, despite their functional and phylogenetic similarity, endemic species make individual rock outcrops unique.     

Inselbergs in a changing world — global trends

Abstract. Based on a review of recent literature, this paper puts forward hypotheses for global trends of inselbergs (isolated mountains) with regard to: (a) their bioclimatic position in relation to the surroundings; (b) their potential for providing habitat niches; and (c) human impacts that may influence ecological processes. This review takes a landscape-level perspective and highlights the challenges ahead in view of changing environmental conditions. Recognizing that inselbergs per se are composed of different microhabitats relative to their surroundings, inselbergs are hypothesized to be bioclimatic 'islands' of xeric conditions in a humid matrix in tropical and temperate regions, and 'islands' of mesic conditions in arid regions. The bioclimatic status of intermediate positions along this global axis (e.g. semiarid and subtropical savanna regions) is less clear. Here, other environmental variables may be of greater importance (microhabitat composition, size of inselberg, distance to other mountain habitats and biogeographical influences). Whether or not biotic communities match these hypothesized physical and bioclimatic trends warrants investigation and could contribute to explaining global species diversity patterns. Invasion of alien species in tropical and Mediterranean-climate regions, and altered fire regimes and resource use pose threats to inselberg communities. Their role as sources of native species to recolonize disturbed surroundings is important in degraded semiarid and arid regions. A generalized model is proposed hypothesizing possible processes between inselberg habitats and disturbed areas in their surroundings in different climatic regions. This model may help to direct further research towards substantiating these perceived trends.     

Contrasting patterns and mechanisms of spatial turnover for native and exotic freshwater fish in Europe

Aim We compare the distribution patterns of native and exotic freshwater fish in Europe, and test whether the same mechanisms (environmental filtering and/or dispersal limitation) govern patterns of decrease in similarity of native and exotic species composition over geographical distance (spatial species turnover). Locations Major river basins of Europe. Methods Data related to geography, habitat diversity, regional climate and species composition of native and exotic freshwater fish were collated for 26 major European river basins. We explored the degree of nestedness in native and exotic species composition, and quantified compositional similarity between river basins according to the beta‐sim (independent of richness gradient) and Jaccard (dependent of richness gradient) indices of similarity. Multiple regression on distance matrices and variation‐partitioning approaches were used to quantify the relative roles of environmental filtering and dispersal limitation in shaping patterns of decreasing compositional similarity over geographical distance. Results Native and exotic species exhibited significant nested patterns of species composition, indicating that differences in fish species composition between river basins are primarily the result of species loss, rather than species replacement. Both native and exotic compositional similarity decreased significantly with increasing geographical distance between river basins. However, gradual changes in species composition with geographical distance were found only for exotic species. In addition, exotic species displayed a higher rate of similarity decay (higher species turnover rate) with geographical distance, compared with native species. Lastly, the majority of explained variation in exotic compositional similarity was uniquely related to geography, whereas native compositional similarity was either uniquely explained by geography or jointly explained by environment and geography. Main conclusions Our study suggests that large‐scale patterns of spatial turnover for exotic freshwater fish in Europe are generated by human‐mediated dispersal limitation, whereas patterns of spatial turnover for native fish result from both dispersal limitation relative to historical events (isolation by mountain ranges, glacial history) and environmental filtering.     

Multi‐scale patterns of moss and lichen richness on the Antarctic Peninsula

Mosses and lichens are the dominant macrophytes of the Antarctic terrestrial ecosystem. Using occurrence data from existing databases and additional published records, we analyzed patterns of moss and lichen species diversity on the Antarctic Peninsula at both a regional scale (1°latitudinal bands) and a local scale (52 and 56 individual snow‐ and ice‐free coastal areas for mosses and lichens, respectively) to test hypothesized relationships between species diversity and environmental factors, and to identify locations whose diversity may be particularly poorly represented by existing collections and online databases. We found significant heterogeneity in sampling frequency, number of records collected, and number of species found among analysis units at the two spatial scales, and estimated species richness using projected species accumulation curves to account for potential biases stemming from sample heterogeneity. Our estimates of moss and lichen richness for the entire Antarctic Peninsula region were within 20% of the total number of known species. Area, latitude, spatial isolation, mean summer temperature, and penguin colony size were considered as potential covariates of estimated species richness. Moss richness was correlated with isolation and latitude at the local scale, while lichen richness was correlated with summer mean temperature and, for 17 sites where penguins where present with <20 000 breeding pairs, penguin colony size. At the regional scale, moss richness was correlated with temperature and latitude. Lichen richness, by contrast, was not significantly correlated with any of the variables considered at the regional scale. With the exception of temperature, which explained 91% of the variation in regional moss diversity, explained variance was very low. Our results show that patterns of moss and lichen biodiversity are highly scale‐dependent and largely unexplained by the biogeographic variables found important in other systems.     

Influence of Habitat and Climate Variables on Arbuscular Mycorrhizal Fungus Community Distribution,as Revealed by a Case Study of Facultative Plant Epiphytism under Semiarid Conditions

In semiarid Mediterranean ecosystems, epiphytic plant species are practically absent, and only some species of palm trees can support epiphytes growing in their lower crown area, such as Phoenix dactylifera L. (date palm). In this study, we focused on Sonchus tenerrimus L. plants growing as facultative epiphytes in P. dactylifera and its terrestrial forms growing in adjacent soils. Our aim was to determine the possible presence of arbuscular mycorrhizal fungi (AMF) in these peculiar habitats and to relate AMF communities with climatic variations. We investigated the AMF community composition of epiphytic and terrestrial S. tenerrimus plants along a temperature and precipitation gradient across 12 localities. Epiphytic roots were colonized by AMF, as determined by microscopic observation; all of the epiphytic and terrestrial samples analyzed showed AMF sequences from taxa belonging to the phylum Glomeromycota, which were grouped in 30 AMF operational taxonomic units. The AMF community composition was clearly different between epiphytic and terrestrial root samples, and this could be attributable to dispersal constraints and/or the contrasting environmental and ecophysiological conditions prevailing in each habitat. Across sites, the richness and diversity of terrestrial AMF communities was positively correlated with rainfall amount during the most recent growing season. In contrast, there was no significant correlation between climate variables and AMF richness and diversity for epiphytic AMF communities, which suggests that the composition of AMF communities in epiphytic habitats appears to be largely determined by the availability and dispersion of fungal propagules from adjacent terrestrial habitats.     

A Multi-Clade Test Supports the Intermediate Dispersal Model of Biogeography

Background

Biogeography models typically focus on explaining patterns through island properties, such as size, complexity, age, and isolation. Such models explain variation in the richness of island biotas. Properties of the organisms themselves, such as their size, age, and dispersal abilities, in turn may explain which organisms come to occupy, and diversify across island archipelagos. Here, we restate and test the intermediate dispersal model (IDM) predicting peak diversity in clades of relatively intermediate dispersers.

Methodology

We test the model through a review of terrestrial and freshwater organisms in the western Indian Ocean examining the correlation among species richness and three potential explanatory variables: dispersal ability quantified as the number of estimated dispersal events, average body size for animals, and clade age.

Conclusions

Our study supports the IDM with dispersal ability being the best predictor of regional diversity among the explored variables. We find a weaker relationship between diversity and clade age, but not body size. Principally, we find that richness strongly and positively correlates with dispersal ability in poor to good dispersers while a prior study found a strong decrease in richness with increased dispersal ability among excellent dispersers. Both studies therefore support the intermediate dispersal model, especially when considered together. We note that many additional variables not here considered are at play. For example, some taxa may lose dispersal ability subsequent to island colonization and some poor dispersers have reached high diversity through within island radiations. Nevertheless, our findings highlight the fundamental importance of dispersal ability in explaining patterns of biodiversity generation across islands.     

Rotifer species richness along an altitudinal gradient in the Alps

Aim Biodiversity patterns along altitudinal gradients are less studied in aquatic than terrestrial systems, even though aquatic sites provide a more homogeneous environment independent of moisture constraints. We studied the altitudinal species richness pattern for planktonic rotifers in freshwater lakes and identified the environmental predictors for which altitude is a proxy. Location Two hundred and eighteen lakes of Trentino–South Tyrol (Italy) in the eastern Alps; lakes covered 98% (range 65–2960 m above sea level) of the altitudinal gradient in the Alps. Methods We performed: (1) linear regression between species richness and altitude to evaluate the general pattern, (2) multiple linear regression between species richness and environmental predictors excluding altitude to identify the most important predictors, and (3) linear regression between the residuals of the best model of step (2) and altitude to investigate any additional explanatory power of altitude. Selection of environmental predictors was based on limnological importance and non‐parametric Spearman correlations. We applied ordinary least squares regression, generalized linear, and generalized least squares modelling to select the most statistically appropriate model. Results Rotifer species richness showed a monotonic decrease with altitude independent of scale effects. Species richness could be explained (R²= 51%) by lake area as a proxy for habitat diversity, reactive silica and total phosphorus as proxies for productivity, water temperature as a proxy for energy, nitrate as a proxy for human influence and north–south and east–west directions as covariates. These predictors completely accounted for the species richness–altitude pattern, and altitude had no additional effect on species richness. Main conclusions The linear decrease of species richness along the altitudinal gradient was related to the interplay of habitat diversity, productivity, heat content and human influence. These factors are the same in terrestrial and aquatic habitats, but the greater environmental stability of aquatic systems seems to favour a linear pattern.     

Within-population spatial genetic structure in four naturally fragmented species of a neotropical inselberg radiation, Alcantarea imperialis, A. geniculata, A. glaziouana and A. regina (Bromeliaceae)

Studies of organisms on 'terrestrial islands' can improve our understanding of two unresolved issues in evolutionary genetics: the likely long-term effects of habitat fragmentation and the genetic underpinnings of continental species radiations in island-like terrestrial habitats. We have addressed both issues for four closely related plant species of the adaptive radiation Bromeliaceae, Alcantarea imperialis, A. geniculata, A. regina and A. glaziouana. All four are adapted to ancient, isolated inselberg rock outcrops in the Brazilian Atlantic rainforest and are thus long-term fragmented by nature. We used eight nuclear microsatellites to study within-population spatial genetic structure (SGS) and historical gene dispersal in nine populations of these species. Within-population SGS reflected known between-species differences in mating systems. The strongest SGS observed in A. glaziouana (Sp=0.947) was stronger than literature estimates available for plants. Analysis of short- and long-distance components of SGS identified biparental inbreeding, selfing and restricted seed dispersal as main determinants of SGS, with restricted pollen dispersal by bats contributing in some localities. The ability of Alcantarea spp. to colonize isolated inselbergs probably stems from their flexible mating systems and an ability to tolerate inbreeding. Short-ranging gene dispersal (average sigma=7-27 m) is consistent with a loss of dispersal power in terrestrial island habitats. Population subdivision associated with sympatric colour morphs in A. imperialis is accompanied by between-morph differences in pollen and seed dispersal. Our results indicate a high potential for divergence with gene flow in inselberg bromeliads and they provide base-line data about the long-term effects of fragmentation in plants.     

Genetic connectivity and diversity in inselberg populations of Acacia woodmaniorum,a rare endemic of the Yilgarn Craton banded iron formations

Historically rare plant species with disjunct population distributions and small population sizes might be expected to show significant genetic structure and low levels of genetic diversity because of the effects of inbreeding and genetic drift. Across the globe, terrestrial inselbergs are habitat for rich, often rare and endemic flora and are valuable systems for investigating evolutionary processes that shape patterns of genetic structure and levels of genetic diversity at the landscape scale. We assessed genetic structure and levels of genetic diversity across the range of the historically rare inselberg endemic Acacia woodmaniorum. Phylogeographic and genetic structure indicates that connectivity is not sufficient to produce a panmictic population across the limited geographic range of the species. However, historical levels of gene flow are sufficient to maintain a high degree of adaptive connectivity across the landscape. Genetic diversity indicates gene flow is sufficient to largely counteract any negative genetic effects of inbreeding and random genetic drift in even the most disjunct or smallest populations. Phylogeographic and genetic structure, a signal of isolation by distance and a lack of evidence of recent genetic bottlenecks suggest long-term stability of contemporary population distributions and population sizes. There is some evidence that genetic connectivity among disjunct outcrops may be facilitated by the occasional long distance dispersal of Acacia polyads carried by insect pollinators moved by prevailing winds.     

Diversity and ecology of saxicolous vegetation mats on inselbergs in the Brazilian Atlantic rainforest

Inselbergs occur as mostly dome-shaped rock outcrops in all climatic and vegetational zones of the tropics. Consisting of Precambrian granites and gneisses, they form ancient and stable landscape elements. Due to harsh edaphic and microclimatic conditions, the vegetation of inselbergs differs markedly from those of the surroundings. Monocotyledonous mats form one of the most characteristic communities of this ecosystem. The floristic composition of this community was studied on six inselbergs located in the Brazilian Atlantic rainforest. Dominating are Bromeliaceae ( Alcantarea , Encholirium ), Velloziaceae ( Vellozia ), Cactaceae ( Coleocephalocereus ) and the Cyperaceae Trilepis . The alpha diversity of the mats was relatively uniform among the six outcrops. However, beta diversity varied considerably between the different sites. Beta diversity was highest at the most speciose locality indicating a high degree of stochasticity in colonization. In contrast to the low diversity mats on West African inselbergs, the Brazilian rock outcrops bear a floristically more diverse mat community rich in endemics. Thus the edaphically controlled inselberg vegetation reflects the outstanding diversity of the Mata Atlaântica. Possibly the high species richness of mats on East Brazilian inselbergs is a consequence of a large species-pool. The processes that regulate regional and local diversity in the Mata Atlaântica are not fully clear. It is assumed that historical (i.e. long-term stability) and biotic (i.e. evolutionary interchange of taxa between canopies and rock outcrops followed by differentiation of local populations) conditions have promoted high rates of speciation and their coexistence in isolated habitats. A danger to the unique vegetation of East Brazilian inselbergs is the establishment of invasive weeds.     

A global model of island biogeography

Aim The goal of our study was to build a global model of island biogeography explaining bird species richness that combines MacArthur and Wilson's area–isolation theory with the species–energy theory. Location Global. Methods We assembled a global data set of 346 marine islands representing all types of climate, topography and degree of isolation on our planet, ranging in size from 10 ha to 800,000 km². We built a multiple regression model with the number of non‐marine breeding bird species as the dependent variable. Results We found that about 85–90% of the global variance in insular bird species richness can be explained by simple, contemporary abiotic factors. On a global scale, the three major predictors — area, average annual temperature and the distance separating the islands from the nearest continent — all have constraining (i.e. triangular rather than linear) relationships with insular bird species richness. We found that the slope of the species–area curve depends on both average annual temperature and total annual precipitation, but not on isolation. Insular isolation depends not only on the distance of an island from the continent, but also on the presence or absence of other neighbouring islands. Range in elevation — a surrogate for diversity of habitats — showed a positive correlation with bird diversity in warmer regions of the world, while its effect was negative in colder regions. We also propose a global statistical model to quantify the isolation‐reducing effect of neighbouring islands. Main conclusions The variation in avian richness among islands worldwide can be statistically explained by contemporary environmental variables. The equilibrium theory of island biogeography of MacArthur and Wilson and the species–energy theory are both only partly correct. Global variation in richness depends about equally upon area, climate (temperature and precipitation) and isolation. The slope of the species richness–area curve depends upon climate, but not on isolation, in contrast to MacArthur and Wilson's theory.     

Island–matrix relationships in Nama Karoo inselberg landscapes. Part I: Do inselbergs provide a refuge for matrix species?

In a first step to investigate the potential source-sink functionof isolated mountains (i.e. inselbergs) in an arid landscape,affinities of inselberg floras with their surrounding were investigated in foursets of inselbergs in Namibia's Nama Karoo. The questions focussed on(a) the potential of inselbergs to serve as sources for degradedrangelands, and (b) the role of species attributes in the relationshipbetween inselberg and matrix. The study showed that inselbergs can serve assource for degraded rangeland, but at regional and landscape level employed inthis analysis, the species attributes growth form and seed dispersal did notprovide any conclusive trends.     

Plants on small islands revisited: the effects of spatial scale and habitat quality on the species–area relationship

Understanding how species diversity is related to sampling area and spatial scale is central to ecology and biogeography. Small islands and small sampling units support fewer species than larger ones. However, the factors influencing species richness may not be consistent across scales. Richness at local scales is primarily affected by small‐scale environmental factors, stochasticity and the richness at the island scale. Richness at whole‐island scale, however, is usually strongly related to island area, isolation and habitat diversity. Despite these contrasting drivers at local and island scales, island species–area relationships (SARs) are often constructed based on richness sampled at the local scale. Whether local scale samples adequately predict richness at the island scale and how local scale samples influence the island SAR remains poorly understood. We investigated the effects of different sampling scales on the SAR of trees on 60 small islands in the Raja Ampat archipelago (Indonesia) using standardised transects and a hierarchically nested sampling design. We compared species richness at different grain sizes ranging from single (sub)transects to whole islands and tested whether the shape of the SAR changed with sampling scale. We then determined the importance of island area, isolation, shape and habitat quality at each scale on species richness. We found strong support for scale dependency of the SAR. The SAR changed from exponential shape at local sampling scales to sigmoidal shape at the island scale indicating variation of species richness independent of area for small islands and hence the presence of a small‐island effect. Island area was the most important variable explaining species richness at all scales, but habitat quality was also important at local scales. We conclude that the SAR and drivers of species richness are influenced by sampling scale, and that the sampling design for assessing the island SARs therefore requires careful consideration.     

Species richness patterns and metapopulation processes – evidence from epiphyte communities in boreo-nemoral forests

For several epiphyte species, dispersal limitation and metapopulation dynamics have been suggested. We studied the relative importance of local environmental conditions and spatial aggregation of species richness of facultative and obligate epiphytic bryophytes and lichens within two old‐growth forests in eastern Sweden. The effect of the local environment was analyzed using generalized linear models (GLM). We tested whether species richness was spatially structured by fitting variogram models to the residuals of the GLM. In addition, we analyzed the species‐area relationship (area=tree diameter). Different environmental variables explained the richness of different species groups (bryophytes vs lichens, specialists vs generalists, sexual vs asexual dispersal). In most groups, the total variation explained by environmental variables was higher than the variation explained by the spatial model. Spatial aggregation was more pronounced in asexually than in sexually dispersed species. Bryophyte species richness was only poorly predicted by area, and lichen species richness was not explained by area at all. Spatial aggregation may indicate effects of dispersal limitation and metapopulation dynamics on community species richness. Our results suggest that species groups differ in habitat requirements and dispersal abilities; there were indications that presence of species with different dispersal strategies is linked to the age of the host tree. Separate analyses of the species richness of species groups that differ in the degree of habitat specialization and dispersal ability give insights into the processes determining community species richness. The poor species‐area relationship, especially in lichens, may indicate species turnover rather than accumulation during the lifetime of the host tree. Epiphyte species extinctions may be mainly caused by deterministic processes, e.g. changes in habitat conditions as the host tree grows, ages and dies, rather than by stochastic population processes.