Sampling adequacy in an extreme environment: species richness patterns in Slovenian caves

Caves harbor a rich fauna unique to subterranean environments. Although extensive records of cave animals are available, only a small fraction of known caves in any region have been biologically assessed. We investigated the impact of incomplete sampling using one of the richest, best documented cave faunas in the world – that of the Dinaric karst of Slovenia. We utilized time snapshots (1940, 1970, and 2000) of the caves and cave fauna to analyze stability of hotspots, spatial pattern, and relationship between number of species and number of caves. Using data aggregated into 100km² hexagons, the location of hotspots, black–white joins, Moran's I, and spatial autocorrelation all remained constant, at least from 1970 on. The linear regression coefficient of the relationship between number of caves and number of species declined with time. Most hexagons had been sampled, but there was no indication that any hexagon had been sampled intensively enough for the accumulation curve of number of caves versus number of species within a hexagon to reach an asymptote. This appeared to be the result of a highly skewed distribution of species richness among caves. Number and position of hotspots can be predicted from information on these few high diversity caves.     

Orchids of the West Indies: predictability of diversity and endemism

Aim We examined phytogeographical patterns of West Indian orchids, and related island area and maximum elevation with orchid species richness and endemism. We expected strong species–area relationships, but that these would differ between low and montane island groups. In so far as maximum island elevation is a surrogate for habitat diversity, we anticipated a strong relationship with maximum elevation and both species richness and endemism for montane islands. Location The West Indies. Methods Our data included 49 islands and 728 species. Islands were classified as either montane (≥ 300 m elevation) or low (< 300 m). Linear and multivariate regression analyses were run to detect relationships between either area or maximum island elevation and species richness or the number of island endemic species. Results For all 49 islands, the species–area relationship was strong, producing a z‐value of 0.47 (slope of the regression line) and explaining 46% of the variation. For 18 relatively homogeneous, low islands we found a non‐significant slope of z = −0.01 that explained only 0.1% of the variation. The 31 montane islands had a highly significant species–area relationship, with z = 0.49 and accounting for 65% of the variation. Species numbers were also strongly related to maximum island elevation. For all islands < 750 km², we found a small‐island effect, which reduced the species–area relationship to a non‐significant z = 0.16, with only 5% of the variation explained by the model. Species–area relationships for montane islands of at least 750 km² were strong and significant, but maximum elevation was the best predictor of species richness and accounted for 79% of the variation. The frequency of single‐island endemics was high (42%) but nearly all occurred on just nine montane islands (300 species). The taxonomic distribution of endemics was also skewed, suggesting that seed dispersability, while remarkable in some taxa, is very limited in others. Montane island endemics showed strong species–area and species–elevation relationships. Main conclusions Area and elevation are good predictors of orchid species diversity and endemism in the West Indies, but these associations are driven by the extraordinarily strong relationships of large, montane islands. The species richness of low islands showed no significant relationship with either variable. A small‐island effect exists, but the montane islands had a significant relationship between species diversity and maximum elevation. Thus, patterns of Caribbean orchid diversity are dependent on an interplay between area and topographic diversity.     

Spatial correlations of Diceroprocta apache and its host plants: evidence for a negative impact from Tamarix invasion

Abstract 1. The hypothesis that the habitat‐scale spatial distribution of the Apache cicada Diceroprocta apache Davis is unaffected by the presence of the invasive exotic saltcedar Tamarix ramosissima was tested using data from 205 1‐m² quadrats placed within the flood‐plain of the Bill Williams River, Arizona, U.S.A. Spatial dependencies within and between cicada density and habitat variables were estimated using Moran's I and its bivariate analogue to discern patterns and associations at spatial scales from 1 to 30 m. 2. Apache cicadas were spatially aggregated in high‐density clusters averaging 3 m in diameter. A positive association between cicada density, estimated by exuvial density, and the per cent canopy cover of a native tree, Goodding's willow Salix gooddingii, was detected in a non‐spatial correlation analysis. No non‐spatial association between cicada density and saltcedar canopy cover was detected. 3. Tests for spatial cross‐correlation using the bivariate I_YZ indicated the presence of a broad‐scale negative association between cicada density and saltcedar canopy cover. This result suggests that large continuous stands of saltcedar are associated with reduced cicada density. In contrast, positive associations detected at spatial scales larger than individual quadrats suggested a spill‐over of high cicada density from areas featuring Goodding's willow canopy into surrounding saltcedar monoculture. 4. Taken together and considered in light of the Apache cicada's polyphagous habits, the observed spatial patterns suggest that broad‐scale factors such as canopy heterogeneity affect cicada habitat use more than host plant selection. This has implications for management of lower Colorado River riparian woodlands to promote cicada presence and density through maintenance or creation of stands of native trees as well as manipulation of the characteristically dense and homogeneous saltcedar canopies.     

Spatial variation in the climatic predictors of species compositional turnover and endemism

Previous research focusing on broad‐scale or geographically invariant species‐environment dependencies suggest that temperature‐related variables explain more of the variation in reptile distributions than precipitation. However, species–environment relationships may exhibit considerable spatial variation contingent upon the geographic nuances that vary between locations. Broad‐scale, geographically invariant analyses may mask this local variation and their findings may not generalize to different locations at local scales. We assess how reptile–climatic relationships change with varying spatial scale, location, and direction. Since the spatial distributions of diversity and endemism hotspots differ for other species groups, we also assess whether reptile species turnover and endemism hotspots are influenced differently by climatic predictors. Using New Zealand reptiles as an example, the variation in species turnover, endemism and turnover in climatic variables was measured using directional moving window analyses, rotated through 360°. Correlations between the species turnover, endemism and climatic turnover results generated by each rotation of the moving window were analysed using multivariate generalized linear models applied at national, regional, and local scales. At national‐scale, temperature turnover consistently exhibited the greatest influence on species turnover and endemism, but model predictive capacity was low (typically r² = 0.05, P < 0.001). At regional scales the relative influence of temperature and precipitation turnover varied between regions, although model predictive capacity was also generally low. Climatic turnover was considerably more predictive of species turnover and endemism at local scales (e.g., r² = 0.65, P < 0.001). While temperature turnover had the greatest effect in one locale (the northern North Island), there was substantial variation in the relative influence of temperature and precipitation predictors in the remaining four locales. Species turnover and endemism hotspots often occurred in different locations. Climatic predictors had a smaller influence on endemism. Our results caution against assuming that variability in temperature will always be most predictive of reptile biodiversity across different spatial scales, locations and directions. The influence of climatic turnover on the species turnover and endemism of other taxa may exhibit similar patterns of spatial variation. Such intricate variation might be discerned more readily if studies at broad scales are complemented by geographically variant, local‐scale analyses.     

Coastal forests of eastern Africa: status, endemism patterns and their potential causes

Eastern African coastal forests are located within the Swahili regional centre of endemism and Swahili-Maputaland regional transition zone in eastern Africa, between 1d? North and 25d? South, and 34—41d? East. Approximately 3167 km² coastal forest remains: 2 km² in Somalia, 660 km² in Kenya, 697 km² in Tanzania, 16 km² in Malawi, 3 km² in Zimbabwe and perhaps 1790 km² in Mozambique. Most forests are small (≤ 20 km²), and all but 19 are under 30 km² in area. Over 80% of coastal forest is located on government land, principally Forest Reserves; only 8.3 km² is found in National Parks (6.2 km² in Kenya (Arabuko-Sokoke), 2 km² in Tanzania (Mafia Island) and tiny patches in Zimbabwe). Coastal forests are an important and highly threatened centre of endemism for plants (c 550 endemic species), mammals (6 species), birds (9 species), reptiles (26 species), frogs (2 species), butterflies (79 species), snails (>86 species) and millipedes (>20 species). Endemic species are concentrated in the forests of the Tana River, between Malindi in Kenya to Tanga in northern Tanzania, and in southern Tanzania. Forests with highest numbers of endemics are: lower Tana River, Arabuko-Sokoke, Shimba Hills (Kenya); lowland East Usambara, Pugu Hills, Matumbi Hills, Rondo and Litipo and other plateaux near Lindi (Tanzania); the Tanzanian offshore island of Pemba; Bazaruto archipelago (Mozambique), and tiny forest remnants of southern Malawi, eastern Zimbabwe and Mozambique. Most coastal forest endemics have a narrow distributional range, often exhibiting single-site endemism or with scattered or disjunct distributional patterns. They are best interpreted as relicts and not the result of recent evolution. Relictualization probably started with the separation of the ancient Pan African rainforest into two parts during the Miocene. The coastal forests are interpreted as a ‘vanishing refuge’ with the endemic species gradually becoming more and more relict (and presumably extinct) due historically to climatic desiccation and more recently to human destruction.     

Spatial autocorrelation as a tool for identifying the geographical patterns of aphid annual abundance

Abstract 1 A spatial autocorrelation analysis was undertaken to investigate the spatial structure of annual abundance for the pest aphid Myzus persicae collected in suction traps distributed across north‐west Europe. 2 The analysis was applied at two different scales. The Moran index was used to estimate the degree of spatial autocorrelation at all sites within the study area (global level). The contributions of each site to the global index were identified by the use of a local indicator of spatial autocorrelation (LISA). A hierarchical cluster analysis was undertaken to highlight differences between groups of resulting correlograms. 3 Similarity between traps was shown to occur over large geographical distances, suggesting an impact of phenomena such as climatic gradients or land use types. 4 The presence of outliers and zones of similarity (hot‐spots) and of dissimilarity (cold‐spots) were identified indicating a strong impact of local effects. 5 Several groups of traps characterized by similarities in their local spatial structure (correlograms, value of Moran's I_i) also had similar values for land use variables (the area occupied by agricultural zones, forest and sea). 6 It is concluded that trap data can provide information about Myzus persicae that is representative of large geographical areas. Thus, trap data can be used to estimate the aerial abundance of this species, even if the suction traps are not regularly and densely distributed.     

HOW SEDENTARY ARE GREYWING FRANCOLINS (FRANCOLINUS AFRICANUS)?

Phasianids are considered to be sedentary birds with limited dispersal so that populations may be expected to show genetic isolation by distance. To test this, we examined genetic variability in 618 greywing francolins (Francolinus africanus) at 24 localities over a 1,500 km² area. We subdivided the samples to measure genetic population structure among localities separated by 6–60 km, and among coveys separated by 0.1–6 km. Thirteen of 30 (43%) allozyme loci were polymorphic, and heterozygosity ranged from 5.3 to 8.5% over 24 localities and averaged 7.0%, a value much larger than that found for other phasianids. Significant allele-frequency heterogeneity was detected among localities and among coveys at several localities for several loci. Mantel's test, however, showed that there was no correlation between geographical distance and the allele-frequency difference between localities for all but one allele. Although spatial autocorrelation was detected with Moran's I and Geary's c for two alleles, the geographical patterns of I in correlograms of 18 independent alleles showed a “crazy-quilt” pattern of allele-frequency patches. This shows that the isolation-by-distance model of subpopulation structure is inappropriate for these birds. Individuals, therefore, appear to disperse far beyond neighboring populations. “Private-allele” and F_ST estimates of migration under the island model were 8–9 individuals between localities of each generation. Allele-frequency heterogeneity, large amounts of gene flow, and the general lack of spatial autocorrelation imply that the small, socially-structured populations of greywing are subject to high rates of turnover, founder effects, and random drift.     

The small-scale spatial distribution of an invading moth

We studied the spread of a small leaf-mining moth [Phyllonorycter leucographella (Zeller), Gracillariidae] after its accidental introduction into the British Isles. At large geographical scales, previous work had shown the spread to be well described by a travelling weve of constant velocity. Here, we report the pattern of spread at scales of 1 km². By locating all bushes of the insect's foodplant (Pyracantha spp.) within 1-km² quadrats, the precise pattern of colonisation at finer spatial scales could be established. Where the 1-km² site was colonised by moths from the main advancing front, no spatial pattern in the order that bushes were infested was found. If the source of colonisation was a single or small group of infested plants within the site, there was some evidence that nearby plants were colonised first. We found no evidence of population turnover after colonisation. We interpret the results in terms of a two-stage model of invasion that produces different patterns at small and large geographical scales.     

Input data, analytical methods and biogeography of Elegia (Restionaceae)

Aim The aim of this paper is to determine the optimal methods for delimiting areas of endemism for Elegia L. (Restionaceae), an endemic genus of the Cape Floristic Region. We assess two methods of scoring the data (presence–absence in regular grids, or in irregular eco‐geographical regions) and three methods for locating biogeographical centres or areas of endemism, and evaluate one method for locating biotic elements. Location The Cape Floristic Region (CFR), South Africa. Methods The distribution of all 48 species of Elegia was mapped as presence–absence data on a quarter‐degree grid and on broad habitat units (eco‐geographical areas). Three methods to delimit areas of endemism were applied: parsimony analysis of endemism (PAE), phenetic cluster analysis, and NDM (‘end em ism’). In addition, we used presence–absence clustering (‘Prabclus’) to delimit biotic elements. The performances of these methods in elucidating the geographical patterns in Elegia were compared, for both types of input data, by evaluating their efficacy in maximizing the proportion of endemics and the number of areas of endemism. Results Eco‐geographical areas perform better than quarter‐degree grids. The eco‐geographical areas are potentially more likely to track the distribution of species. The phenetic approach performed best in terms of its ability to delimit areas of endemism in the study area. The species richness and the richness of range‐restricted species are each highest in the south‐western part of the CFR, decreasing to the north and east. The phytogeographical centres identified in the present study are the northern mountains, the southern mountains (inclusive of the Riviersonderend Mountains and the Cape Peninsula), the Langeberg range, the south coast, the Cape flats, and the west coast. Main conclusions This study demonstrates that (1) eco‐geographical areas should be preferred over a grid overlay in the study of biogeographical patterns, (2) phenetic clustering is the most suitable analytical method for finding areas of endemism, and (3) delimiting biotic elements does not contribute to an understanding of the biogeographical pattern in Elegia. The areas of endemism in Elegia are largely similar to those described in other studies, but there are many detailed differences.     

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.     

Potential migration routes and barriers for vascular plants of the Neotropical Guyana Highlands during the Quaternary

Aim To reconstruct in detail the potential migration routes and barriers for vascular plants from the summits of the Guyana mountains during the Quaternary, in order to test the possibility of migration among them during the glaciations. These changes in connectivity are predicted based on the altitudinal migration of plant communities associated with glacial cooling. To examine the effects of these cycles, the extent of the potential biotic interchange and its influence on patterns of endemism was modelled. Location The summits of the tepuis or table mountains of the Neotropical Guyana Highlands, which constitute the peculiar and discontinuous Pantepui phytogeographical province (total surface 5000 km², altitudinal range 1500–3014 m a.s.l.), and is characterized by a unique and diverse flora with a high degree of endemism. Methods GIS‐based palaeotopographical reconstruction using a high‐precision digital elevation model, combined with phytogeographical analysis by means of a data base built up from the Flora of the Venezuelan Guyana, which includes the geographical and altitudinal ranges for each Pantepui species. Results During the Last Glacial Maximum, which serves as a representative of a standard Quaternary glaciation, most migration pathways among tepuis were open for species with lower altitudinal levels (LAL) ≤ 1500 m (1678 species or c. 69% of the total Pantepui species), and closed for species with LAL ≥ 2300 m altitude (c. 3%). The species in between these altitudes have intermediate migratory possibilities, depending on the district and the tepui considered. If these local factors are considered, the number of species with no possibility of glacial interchange increases to 202 or c. 8% of the total. The strongest topographical barrier separated eastern locations above 1600–1700 m elevation from all others. The highest possibility of interchange was among the tepuis of the eastern sector, in which internal topographical barriers were only effective for species with LALs at or above 1900 m. Main conclusions The Quaternary evolution of the vascular flora from the Guyana Highlands took place in a predominantly migration‐prone, glacial‐era landscape, in which more than 70% of the flora (maximum estimate) was able to move from one tepuian district to another, thanks to the downward bioclimatic shift caused by cooling. Interglacials were too short to drive significant evolutionary diversification. A number of present high‐altitude local endemics are species that were unable to migrate, even during glaciations. However, some endemic species do appear to have been able to migrate among regions, suggesting that topographical isolation alone is not enough to explain patterns of endemism. Other factors such as tepui summit area, habitat heterogeneity or pre‐Quaternary evolution are considered. These studies should be complemented with palaeoecological and phylogeographical surveys.     

Spatial patterns of tree recruitment in a relict population of Pinus uncinata: forest expansion through stratified diffusion

Aim To infer future changes in the distribution of isolated relict tree populations at the limit of a species’ geographical range, a deep understanding of the regeneration niche and the spatial pattern of tree recruitment is needed. Location A relict Pinus uncinata population located at the south‐western limit of distribution of the species in the Iberian System of north‐eastern Spain. Methods Pinus uncinata individuals were mapped within a 50 × 40‐m plot, and their size, age and reproductive status were estimated. Data on seed dispersal were obtained from a seed‐release experiment. The regeneration niche of the species was assessed based on the associations of seedling density with substrate and understorey cover. The spatial pattern of seedlings was described using point‐pattern (Ripley's K) and surface‐pattern (correlograms, Moran's I) analyses. Statistical and inverse modelling were used to characterize seedling clustering. Results Pine seedlings appeared aggregated in 6‐m patches. Inverse modelling estimated a longer mean dispersal distance (27 m), which corresponded to the size of a large cluster along the north to north‐eastward direction paralleled by an eastward trend of increasing seedling age. The two spatial scales of recruitment were related to two dispersal processes. The small‐scale clustering of seedlings was due to local seed dispersal in open areas near the edge of Calluna vulgaris mats: the regeneration niche. The long‐range expansion might be caused by less frequent medium‐distance dispersal events due to the dominant north‐westerly winds. Main conclusions To understand future range shifts of marginal tree populations, data on seed dispersal, regeneration niche and spatial pattern of recruitment at local scales should be obtained. The monitoring of understorey communities should be a priority in order to predict correctly shifts in tree species range in response to global warming.     

Long‐term population monitoring of the threatened and endemic black‐headed squirrel monkey (Saimiri vanzolinii) shows the importance of protected areas for primate conservation in Amazonia

Population monitoring of endangered species is essential to the improvement of their management and conservation plans. The black‐headed squirrel monkey (Saimiri vanzolinii) is a vulnerable species on the IUCN Red List and has extreme geographical endemism, exhibiting the smallest known distribution among Neotropical primates (ca. 870 km²), over 90% of which occurs in white‐water flooded forests within the Mamirauá Sustainable Development Reserve (MSDR), Brazilian Amazonia. To assess the effectiveness of this protected area in conserving the species, we conducted population monitoring of black‐headed squirrel monkeys across five consecutive years (2009–2013) on nine trails 2 km each. Each year samples included both low and high river water periods. We used the distance sampling method, recording the distance to each observed social group as well as counting component individuals. We also calculated annual encounter rates based on the number of individuals sighted every 10 km traveled. Densities ranged from 256 individuals/km² (2011) to 453 individuals/km² (2013), and no seasonal differences were detected. Population size was estimated to be 147,848 mature individuals. Encounter rates ranged from 100 individuals/10 km (2010) to 179 individuals/10 km (2013); no significant difference among years was found. We found that S. vanzolinii populations remained stable throughout the years, which indicates that the MSDR has been playing an essential role on protecting this species. Due to difficulties of fulfilling assumptions of the distance sampling method, we consider the encounter rate analysis to be more effective for monitoring this and other Saimiri species. Given the critical endemism and worrying conservation status of S. vanzolinii, we suggest that monitoring of the species population should be carried out regularly.     

Morphology,ultrastructure and phylogenetic affinities of the single-island endemic Anthoceros cristatus Steph. (Ascension Island)

Oceanic islands, due to their geographical isolation, number, precisely defined boundaries and their geomorphological and climatic diversity, have provided enormous insights into speciation, dispersal, adaptive radiations and macroecological processes. One of the key components of these island studies is the role of single-island endemics (SIEs) as, in many instances, island biogeography models use the proportion of SIEs to infer evolutionary processes. It is, therefore, imperative to undertake critical taxonomic revisions to evaluate SIEs because changes in the number of SIEs have a key impact on downstream biogeographic analyses. We revise the special case of a putative SIE Anthoceros cristatus on Ascension Island using light and electron microscopy, as well phylogenomic tools. A. cristatus lies within the A. agrestis/A. punctatus complex but differs from the sister species A. agrestis and A. punctatus in spore morphology and gametophytic lamellae fringed with caducous marginal cells. The present confirmation, from both molecules and morphology, of the SIE status of Anthoceros cristatus and its restricted distribution on the Island makes the preservation of its habitat a conservation priority. Ascension Island is the tip of an undersea volcano that is thought to have emerged from the ocean 1 million years ago with an area of approximately 91?km², with Green Mountain as the highest elevation (～859?m a.s.l.). Ascension has a relative low bryophyte species diversity of 87 spp but this includes 12 endemics (～14%); a much higher level of endemism than on the far more speciose Macaronesian Islands.     

Spatial ordination of vegetation data using a generalization of Wartenberg's multivariate spatial correlation

Question: Are there spatial structures in the composition of plant communities? Methods: Identification and measurement of spatial structures is a topic of great interest in plant ecology. Univariate measurements of spatial autocorrelation such as Moran's I and Geary's c are widely used, but extensions to the multivariate case (i.e. multi‐species) are rare. Here, we propose a multivariate spatial analysis based on Moran's I (MULTISPATI) by introducing a row‐sum standardized spatial weight matrix in the statistical triplet notation. This analysis, which is a generalization of Wartenberg's approach to multivariate spatial correlation, would imply a compromise between the relations among many variables (multivariate analysis) and their spatial structure (autocorrelation). MULTISPATI approach is very flexible and can handle various kinds of data (quantitative and/or qualitative data, contingency tables). A study is presented to illustrate the method using a spatial version of Correspondence Analysis. Location: Territoire d'Etude et d'Expérimentation de Trois‐Fontaines (eastern France). Results: Ordination of vegetation plots by this spatial analysis is quite robust with reference to rare species and highlights spatial patterns related to soil properties.     

The scale of demographic heterogeneity in a population of Peromyscus leucopus

Summary The patterns of spatial heterogeneity in density and demography were studied in a population of Peromyscus leucopus inhabiting a deciduous forest in west-central Indiana. A series of 9 live-trapping grids sampled densities from 3 spatial scales: 3 ha, 80 ha and 350 km². We found high levels of variation within all three spatial scales. There was as much variation within a single, large grid as within the entire Sugar Creek Valley (350 km²). The patterns of density variation were not temporally stable and thus represent variation in demographic processes rather than simply differences in habitat quality. We suggest that the Sugar Creek Valley represents a metapopulation of Peromyscus leucopus composed of a number of temporally unstable subpopulations. This structure provides considerable demographic variation that dispersing animals can exploit.     

Habitat distribution models for intertidal seaweeds: responses to climatic and non‐climatic drivers

Aim Because intertidal organisms often live close to their physiological tolerance limits, they are potentially sensitive indicators of climate‐driven changes in the environment. The goals of this study were to assess the effect of climatic and non‐climatic factors on the geographical distribution of intertidal macroalgae, and to predict future distributions under different climate‐warming scenarios. Location North‐western Iberian Peninsula, southern Europe. Methods We developed distribution models for six ecologically important intertidal seaweed species. Occurrence and microhabitat data were sampled at 1‐km² resolution and analysed with climate variables measured at larger spatial scales. We used generalized linear models and applied the deviance and Bayesian information criterion to model the relationship between environmental variables and the distribution of each target species. We also used hierarchical partitioning (HP) to identify predictor variables with higher independent explanatory power. Results The distributions of Himanthalia elongata and Bifurcaria bifurcata were correlated with measures of terrestrial and marine climate, although in opposite directions. Model projections under two warming scenarios indicated the extinction of the former at a faster rate in the Cantabrian Sea (northern Spain) than in the Atlantic (west). In contrast, these models predicted an increase in the occurrence of B. bifurcata in both areas. The occurrences of Ascophyllum nodosum and Pelvetia canaliculata, species showing rather static historical distributions, were related to specific non‐climatic environmental conditions and locations, such as the location of sheltered sites. At the southernmost distributional limit, these habitats may present favourable microclimatic conditions or provide refuges from competitors or natural enemies. Model performances for Fucus vesiculosus and F. serratus were similar and poor, but several climatic variables influenced the occurrence of the latter in the HP analyses. Main conclusions The correlation between species distributions and climate was evident for two species, whereas the distributions of the others were associated with non‐climatic predictors. We hypothesize that the distribution of F. serratus responds to diverse combinations of factors in different sections of the north‐west Iberian Peninsula. Our study shows how the response of species distributions to climatic and non‐climatic variables may be complex and vary geographically. Our analyses also highlight the difficulty of making predictions based solely on variation in climatic factors measured at coarse spatial scales.     

A cryptic genetic boundary in remnant populations of a long‐lived,bird‐pollinated shrub Banksia sphaerocarpa var. caesia (Proteaceae)

Plant species often exhibit genetic structure at multiple spatial scales. Detection of this structure may depend on the sampling strategy used. We intensively sampled a common, naturally patchy Banksia species within a 200 km² region, in order to assess patterns of genetic diversity and structure at multiple spatial scales. In total, 1321 adult shrubs from 37 geographical populations were genotyped using eight highly polymorphic microsatellite markers developed for the species. Genetic structure was detected at three spatial scales. First, we identified a stark and unexpected division of the landscape into two genetic subregions, one to the north‐east and one to the south‐west of the sampling grid. This differentiation was based on sudden, highly structured differences in common allele frequencies, the cause of which is unknown but that may relate to physical and reproductive barriers to gene flow, localised selection, and/or historical processes. Second, we observed genetic differentiation of populations within these subregions, reflecting previously described patterns of restricted pollen flow in this species. Finally, fine‐scale genetic structure, although weak, was observed within some of the populations (mean S_P = 0.01837). When feasible, intensive sampling may uncover cryptic patterns of genetic structure that would otherwise be overlooked when sampling at broader spatial scales. Further studies using a similar sampling strategy may reveal this type of discontinuity to be a feature of other south‐western Australian taxa and has implications for our understanding of evolution in this landscape as well as conservation into the future. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 115 , 241–255.     

Climatic and spatial patterns of diversity in the serpentine plants of California

Abstract . The insular distribution of distinctive substrates, such as outcrops of serpentine rock, may either promote plant diversity by enhancing opportunities for speciation or reduce diversity by increasing rates of extinction. To examine the relationship between diversity and the spatial structure of habitats, we studied large‐scale patterns of diversity in the flora of serpentine in California. We used multiple linear regressions on geographical information system (GIS)‐derived data for 85 subregions of the state to analyse the climatic and spatial correlates of plant species richness. The diversity of plants endemic to serpentine declined from north to south and from the coast inland, in association with decreasing rainfall; the same trends were seen in the total flora, but the trends were stronger in serpentine endemics. Diversity of serpentine endemics increased with the area of serpentine and decreased with the mean isolation of serpentine patches in a subregion. The diversity of endemics was not correlated with the number of serpentine patches or their mean perimeter‐to‐area ratio. We conclude that patchiness in this terrestrial habitat does not appear to promote diversity, even at the large spatial scale associated with speciation and endemism.     

Response of native ungulates to drought in semi‐arid Kenyan rangeland

The distribution and abundance of native ungulates were measured on commercially managed, semi‐arid rangeland in central Kenya over a 3‐year period that encompassed severe drought and above‐average rainfall. Native ungulate biomass density averaged 5282 kg km^?2 over the study and was dominated by elephant (Loxodonta africana), impala (Aepyceros melampus) and dik‐dik (Madoqua kirkii). Biomass density of domestic cattle (Bos taurus) averaged 2280 kg km^?2 during the study. Responses of native ungulates to severe drought were variable. Impala densities were similar to or greater than densities for similar habitat in protected areas, and varied from 12 to 16 km^?2 during and following the drought to 24–29 km^?2 following above‐average rainfall. Dik‐dik densities were also greater than densities reported for protected areas and were surprisingly stable throughout the study despite the wide annual fluctuations in rainfall. Elephant migrated out of the region during drought but were present at high densities (2.9–5.2 km^?2) during wet seasons, consistent with telemetry studies emphasizing the importance of Acacia bushland habitat on commercial rangelands for the migratory portion of the Laikipia–Samburu elephant population. Results show that substantial densities of native browsing and mixed‐feeding ungulates can occur on rangeland managed for commercial beef production and suggest that the capacity for ungulates to move over large spatial scales (>100 km²) and to shift distributions in response to locally variable thunderstorms may be important for sustaining these populations.