Conservation issues of temporary wetland Branchiopoda (Anostraca, Notostraca: Crustacea) in a semiarid agricultural landscape: What spatial scales are relevant?

Ecologists increasingly recognise the importance of spatial scale for conservation. This study focuses on threatened temporary wetland crustaceans, the fairy shrimp Branchinecta orientalis Sars (Anostraca) and the tadpole shrimp Triops cancriformis Bosc (Notostraca). Using redundancy analyses with a canonical variance partitioning approach, we studied how local habitat conditions and landscape features influence their densities at 4 spatial scales (100 m buffer strip around ponds, 1 km, 5 km and 10 km catchment scales). Branchinecta densities were negatively related with local conditions (trophic status) at all scales. Landscape effects (catchment:wetland size ratio) were only significant at the 10 km scale. However, trophic state conditions were influenced by local contamination rather than landscape conditions. Local degradation tended to be more pronounced in wetlands situated in catchments with a higher cover of natural vegetation compared to those in agricultural catchments. Triops was less influenced by local effects at all scales. The importance of landscape effects increased with landscape scale but effects were only significant at the 10 km scale, and were negatively explained by irrigated croplands. The importance of broad landscape scales and the difficulty to restore locally degraded sites challenges management. Because rationalisation of large-scale agricultural practises can conflict with socioeconomic demands, a first step to the conservation of actual Branchiopoda populations in this remnant wetland complex could benefit from the creation of vegetated buffer strips around the wetlands and/or hedgerows around agricultural fields to counteract atmosphere-mediated flux of particles and solutes from croplands to wetlands at broad landscape scales.     

The importance of spatial scale for the conservation of tidal flat macrobenthos: An example from New South Wales, Australia

Planners of Marine Protected Areas (MPAs) commonly use maps of habitat types when choosing areas to conserve. This assumes that habitats are homogeneous, and therefore, that any area of habitat will represent the full spectrum of ecological diversity within that habitat. Here, we report that macrobenthic assemblages in tidal flat habitats were spatially heterogeneous in terms of beta diversity (taxonomic turnover), abundance, taxonomic richness and Shannon-Wiener H′ Diversity. Importantly, the patterns of heterogeneity were scale dependent for the three spatial scales we examined; plots (20 m), sites (100s of m) and estuaries (<30 km). The three estuaries in the study were compositionally similar as they shared the same dominant taxa, although one estuary had significantly more taxa and a higher abundance of macrobenthos. Assemblages within tidal flats differed at scales of 100s of m for all ecological measures. Most notably, beta diversity was highest at this scale. Assemblages were relatively more homogeneous at the 20-m scale. These findings highlight the value of examining more than one ecological measure and estimating magnitude of effects across a variety of scales. This work presents two important considerations for MPAs. First, although tidal flats in different estuaries are compositionally similar for dominant taxa, rarer taxa and high heterogeneity in abundance should influence the choice and number of tidal flats in MPAs. Second, strong compositional heterogeneity within individual tidal flats implies that conservation of whole habitat, rather than sections of a tidal flat, is essential if this habitat type is to be used to represent taxonomic diversity.     

Do secondary forests act as refuges for old growth forest animals? Recovery of ant diversity in the Atlantic forest of Brazil

The extent to which secondary forests occupying degraded and abandoned lands provide suitable habitat for forest-adapted species is an important conservation issue in times of vanishing old growth forests. We used ants (Hymenoptera: Formicidae), a functionally important and diverse group of invertebrates, to investigate the recovery of soil taxa during secondary forest succession in the Atlantic Forest of Southern Brazil. We compared the resilience of epigeic vs. hypogeic ant assemblages. For this purpose we established 27 sites that encompassed a chronosequence from pastures to old growth forests on two contrasting soil types. Our results are based on a collection of 35 508 individuals in 40 genera.Richness and composition of ant assemblages in secondary forests have recovered slowly and have not approached conditions typical of old growth forests. The distribution of genera along the successional stages was arranged in a nested pattern where ant genera of younger successional stages were a subset of genera present in older stages. Edaphic conditions had no influence on the recovery process. Overall, richness of ants was lower at study sites with water-logged soils than at sites where soils did not exhibit hydromorphic properties. The hypogeic ant assemblage recovered more slowly than the epigeic assemblage.Our results show that secondary forests do not act as refuges for many forest-adapted animals which are currently restricted to discontinuous patches of old growth forest in the highly endangered Atlantic Forest of Brazil. Moreover, estimated recovery times of 50 to several hundred years suggest it would take much longer than previously presumed for complete recolonization.     

Assessing and mitigating the effects of agricultural soil erosion on roadside ditches

Purpose

Roadside ditches line more than 6.3 million km of roadways in the USA, dissecting the natural topography and altering the flow of runoff from the catchments that drain into them. In agricultural regions, more than 30% of a watershed may directly drain into the roadside ditch system. Quantifying soil erosion and sediment export from agricultural watersheds is a crucial component when considering long-term soil sustainability.

Materials and methods

Our study evaluated the relation of catchment soil erosion and ditch sedimentation at six representative roadside ditches in Lime Creek watershed (eastern Iowa) and quantified the effectiveness of possible catchment conservation practices to reduce soil erosion and ditch sedimentation.

Results and discussion

Study results provide clear evidence linking roadside ditches to the agricultural catchments that drain into them. Among the six ditch sites, catchment erosion was found to be inversely related to sediment storage within the ditch due to erosive power of water entering the ditches from their basins. Of four catchment scenarios to reduce soil erosion, no-till with cover and graded terrace did not require land to be taken out of production and provided the most significant reductions in catchment erosion rates.

Conclusions

Results indicated that reducing nutrient and sediment loads to ditches by incorporating in-field conservation practices in ditch catchments may be more economical and environmentally sustainable than current management practices for both farmers and roadway managers because they trap detached soil sediments before they enter the ditch.

     

Conservation priorities for freshwater biodiversity: The Key Biodiversity Area approach refined and tested for continental Africa

Freshwater ecosystems represent one of the most threatened broad habitat types globally. Despite containing around a third of all vertebrates, area-based approaches to conservation planning rarely include freshwater species as an explicit target for conservation. Here we describe and apply a globally applicable methodology comparable to those for other groups (i.e. Important Bird Areas) to identify river and lake catchments that represent, or contain, freshwater Key Biodiversity Areas. We discuss the rationale behind the methodology and propose appropriate definitions and quantitative threshold values for the selection criteria. Thresholds are developed through spatial analysis of species information for four comprehensively assessed freshwater taxonomic groups in continental Africa, comprising 4203 species, as recently assessed for the IUCN Red List of Threatened Species?. To illustrate application of the methodology freshwater Key Biodiversity Areas are identified across continental Africa, and conservation planning software used to prioritise a network of catchments that captures 99% of the total species complement within catchments covering ca. 20% of the total land area. Within these prioritised catchments only 19% of river length falls within existing Protected Areas suggesting that, given the high connectivity within freshwater ecosystems and their dependence upon catchment management for effective conservation, modification or expansion of the protected area network is required to increase effective conservation of freshwater species. By applying this methodology, gaps in the coverage of freshwater species by existing Protected Areas can be identified and used to inform conservation policy and investment to ensure it is inclusive of, and effective for, freshwater biodiversity.     

Habitat-based conservation priorities for carabid beetles within the Picos de Europa National Park, northern Spain

Quick biodiversity studies on poorly studied taxa and areas are increasingly popular for setting conservation priorities over a wide range of spatial scales. However, the implementation of such studies is complicated by the variable extent to which the different criteria used in prioritisation are correlated to each other. Using methods of constrained ordination, we examined the species-habitat relationships of carabid beetles based on ground beetle assemblages from 22 sites in the Picos de Europa National Park, northern Spain. We found characteristic species assemblages for subalpine meadows, Genista shrublands, and pastures, whereas mown meadows, heathlands, beech and riparian woodlands were occupied by more habitat generalist species. Species associated with subalpine meadows and Genista shrublands tended to be mostly brachypterous and to have geographic ranges restricted to northern Spain. In contrast, we found no relationship between the degree of species' association with pastures and geographic range-wing size type. Although the species richness was higher in riparian woodlands and mown meadows, we suggest a higher conservation value for subalpine meadows and Genista shrublands across the landscape because they sustain characteristic assemblages dominated by species with restricted ranges and reduced powers of dispersal. Our study suggests that preserving areas in the landscape supporting higher biodiversity will not necessarily preserve those species potentially more susceptible to habitat loss and fragmentation. It also supports the feasibility of biodiversity studies based on multivariate techniques for setting conservation priorities over complex landscapes.     

Responses of waterbirds to flooding in an arid region of Australia and implications for conservation

Floods are a frequent but irregular feature of Australia's dryland river catchments. We investigated changes in abundances of waterbirds in north western New South Wales with changes in wetland distribution at local, catchment and broad scales. The abundance of most functional groups of waterbirds changed in response to broad scale changes in wetland distribution, while local abundance remained highly variable. Patterns of abundance varied among functional groups of waterbirds, with some immediately responding to changes in wetland distribution and area flooded, and others apparently responding to sequences of wetting and drying. In Australia, the main conservation issue for waterbirds is water and its use across the landscape and not the spatial arrangement of any fixed array of reserves established to protect them.     

Spatial genetic variation of a riparian wolf spider Pardosa agricola (Thorell, 1856) on lowland river banks: The importance of functional connectivity in linear spatial systems

Habitat patches along river systems are often highly isolated and characterised by a high degree of heterogeneity at different spatio-temporal scales. The connectivity between river bank arthropod populations directly adjoining the river channel is often greatly disturbed due to river regulation. While flight-active arthropods easily disperse upstream, less mobile species are expected to show predominant downstream dispersal unless specific upward movements are prevalent. In linear river ecosystems, downstream drift of organisms may therefore prevail with subsequently strong asymmetrical gene flow. We analysed patterns of genetic variation within and among nine spatially structured populations of the highly stenotopic riparian wolf spider (Lycosidae) Pardosa agricola (Thorell, 1856) using Amplified Fragment Length Polymorphism (AFLP) markers. No evidence was found for downstream accumulation of genetic diversity. The high genetic diversity is hypothesised to be the result of historical drift processes. Nearby populations on the same river shore were less genetically differentiated compared with populations further away and/or on the opposite shore. This indicates that short-distance dispersal still occurs on river banks during low water flow levels, but at the same time that the river channel constitutes a physical barrier for species exchange between opposite shores. The rehabilitation of the riparian corridor will increase the amount of suitable habitat, the functional connectivity during periods of low flow-discharges and eventually riparian spider population persistence.     

Methods for conservation outside of formal reserve systems: The case of ants in the seasonally dry tropics of Veracruz, Mexico

Like most ecosystems of the world, tropical dry forests of the central coast of the Gulf of Mexico are inadequately preserved. Given that reserve expansion is unlikely, it is imperative that the conservation capacity of the matrix surrounding reserves is enhanced. Here, we examine the habitat value of isolated pasture trees and patches of secondary regrowth in terms of their terrestrial and arboreal ant assemblages in both a wet and dry season. These simplified wooded systems increase species densities within the agricultural matrix and provide habitat for some forest ant species. Estimated species richness of arboreal ants was particularly low on isolated trees, highlighting an important limitation. This was not the case for terrestrial ants, which were particularly species rich under isolated trees. We also found that the inter-site variations in species densities and similarity to the forest ant assemblage for terrestrial and arboreal strata were not correlated, suggesting that responses to restoration may not be as uniform as often thought. This has important implications for the use of indicator taxa in suggesting the response of other taxa. In terms of species composition, neither secondary regrowth nor isolated trees were appropriate replacements for forest fragments, even though the studied forest fragments were small (13-32 ha). The ant assemblages did not exhibit a seasonal change in composition. However, season influenced the contrast between habitats, with isolated trees being more distinct from pasture, and regrowth more closely resembling forests, during the wet season. Microclimatic variables indicate that the forests were least affected by the tropical dry season, and this may contribute to their characteristic fauna. We conclude that even small forest patches make a unique contribution to landscape conservation and that, where reserves are limited, conscientious management of the landscape matrix may provide some species with sufficient new habitat to survive outside of reserve systems. These conclusions are influenced by both season and strata studied.     

Trends and controls of Holocene floodplain sedimentation in the Rhine catchment

Holocene floodplain sedimentation in the Rhine catchment is controlled by human and climate impacts. Intricate river behaviour modifies the fluvial response to the external impacts making cause–effect analysis difficult, especially on large spatial scales. To better understand the relative importance and interdependencies of external and internal controls, temporally resolved floodplain sedimentation rates are established using three different methods: i) floodplain storage studies on the trunk stream, ii) depth/age-analysis of overbank deposits from different parts of the catchment and iii) cumulative frequency distributions of ¹⁴C-ages from floodplain deposits from various parts of the catchment. The applied methodology strongly differs with the available temporal resolution and the size of the corresponding catchment. All three methods show a strong increase in sedimentation rate for more recent periods that can be linked to increasing human impact. Evidences for climate impacts and intricate river behaviour are less clear and hindered by insufficient temporal resolution of the currently available data.     

Colonisation of newly established habitats by soil decomposer organisms: the effect of habitat corridors in relation to colonisation distance and habitat size

The aim of the present 2.5-year-long field experiment was to explore the ability of various members of the detrital food web to colonise newly established habitat patches in field conditions, either in the presence or absence of habitat corridors. Patch size and distance to the “mainland” (colonisation source) were manipulated to explore the scale dependency of the corridor effects. Sterilised humus patches, embedded in mineral soil regarded as uninhabitable (or non-preferred) matrix for the soil organisms, functioned as newly established habitats. Intact forest soil served as the source of colonisers. Three kinds of patches were established: large ones situated at relatively long distance from the colonisation source, and small ones situated either at long or short distance from the source. Corridors, when present, connected the patches to the intact forest soil. The presence of corridors consistently increased the species richness of soil fungi in the patches. Also bacterial species richness, number of microarthropod taxa and abundance of enchytraeid worms were sometimes increased by the presence of corridors. The response of bacterial species was assumed to be indirect, caused by the response of enchytraeid worms functioning as dispersal agents for the bacteria. Colonisation of the patches by soil organisms was virtually independent of patch size and colonisation distance, both in the presence and absence of corridors. This can be related to the resource-poor or otherwise sub-optimal conditions in the initially sterile humus soil, restricting the number of species able to establish themselves in the patches and/or utilise the corridors. Nevertheless, the results indicate that habitat corridors can facilitate the colonisation of unpopulated habitats by organisms with greatly different life history traits.     

A multi-taxa assessment of nestedness patterns across a multiple-use Amazonian forest landscape

Understanding how biodiversity is partitioned among alternative land-uses is an important first step for developing effective conservation plans in multiple-use landscapes. Here, we analysed nestedness patterns of species composition for nine different taxonomic groups [dung beetles, fruit-feeding butterflies, orchid bees, scavenger flies, leaf-litter amphibians, lizards, bats, birds and woody plants (trees and lianas)] in a multiple-use forestry landscape in the Brazilian Amazon containing primary, secondary and Eucalyptus plantation forests. A formal nestedness analysis was performed to investigate whether species-poor land-uses were comprised of a subset of species from more diverse forests, and the extent to which this pattern varied among taxa. At the landscape-scale the species-by-sites matrices were significantly nested for all nine taxonomic groups when both sites and species were sorted to maximally pack the species/occurrence matrix and, except for orchid bees when sorted by land-use intensity (primary forest to Eucalyptus plantation). Different patterns emerged when we conducted pairwise analyses of nestedness between the three forest types: (a) most of the taxonomic groups were nested in accordance with increased land-use intensity; (b) neither orchid bees nor leaf-litter amphibians from secondary forest made up a significant nested subset of primary forest species, although species found in Eucalyptus plantation sites were nested within secondary forest communities; and (c) lizards from Eucalyptus plantations were not a nested subset of either primary or secondary forest. Our findings emphasize the complex nature of patterns of species occupancy in tropical multiple-use forestry landscapes, and illustrate that there may be no easy solutions to questions regarding the conservation value of secondary and exotic plantation forests.     

Process-based modelling of erosion,sediment transport and reservoir siltation in mesoscale semi-arid catchments

Purpose

To support scientifically sound water management in dryland environments a modelling system has been developed for the quantitative assessment of water and sediment fluxes in catchments, transport in the river system, and retention in reservoirs. The spatial scale of interest is the mesoscale because this is the scale most relevant for management of water and land resources.

Materials and methods

This modelling system comprises process-oriented hydrological components tailored for dryland characteristics coupled with components comprising hillslope erosion, sediment transport and reservoir deposition processes. The spatial discretization is hierarchically designed according to a multi-scale concept to account for particular relevant process scales. The non-linear and partly intermittent run-off generation and sediment dynamics are dealt with by accounting for connectivity phenomena at the intersections of landscape compartments. The modelling system has been developed by means of data from nested research catchments in NE-Spain and in NE-Brazil.

Results and discussion

In the semi-arid NE of Brazil sediment retention along the topography is the main process for sediment retention at all scales, i.e. the sediment delivery is transport limited. This kind of deposition retains roughly 50 to 60 % of eroded sediment, maintaining a similar deposition proportion in all spatial scales investigated. On the other hand, the sediment retained in reservoirs is clearly related to the scale, increasing with catchment area. With increasing area, there are more reservoirs, increasing the possibility of deposition. Furthermore, the area increase also promotes an increase in flow volume, favouring the construction of larger reservoirs, which generally overflow less frequently and retain higher sediment fractions. The second example comprises a highly dynamic Mediterranean catchment in NE-Spain with nested sub-catchments and reveals the full dynamics of hydrological, erosion and deposition features. The run-off modelling performed well with only some overestimation during low-flow periods due to the neglect of water losses along the river. The simulated peaks in sediment flux are reproduced well, while low-flow sediment transport is less well captured, due to the disregard of sediment remobilization in the riverbed during low flow.

Conclusions

This combined observation and modelling study deepened the understanding of hydro-sedimentological systems characterized by flashy run-off generation and by erosion and sediment transport pulses through the different landscape compartments. The connectivity between the different landscape compartments plays a very relevant role, regarding both the total mass of water and sediment transport and the transport time through the catchment.     

Network thinking in riverscape conservation – A graph-based approach

Graph theoretic approaches have received increased interest recently in landscape planning and conservation in the terrestrial realm, because these approaches facilitate the effective modelling of connectivity among habitats. We examined whether basic principles of graph theory can be extended to other ecosystems. Specifically, we demonstrate how a network-based context can be used for enhancing the more effective conservation of riverine systems. We first show how to use graph theoretic techniques to model riverscapes at the segment level. Then we use a real stream network (Zagyva river basin, Hungary) to examine the topological importance of segments in maintaining riverscape connectivity, using betweenness centrality, a commonly used network measure. Using the undirected graph model of this riverscape, we then prioritize segments for conservation purpose. We examine the value of each of the 93 segments present in the Zagyva river basin by considering the conservation value of local fish assemblages, connectivity and the size of the habitat patches. For this purpose we use the ‘integral index of connectivity’, a recently advocated habitat availability index. Based on the results the selection of the most valuable habitat segments can be optimized depending on conservation resources. Because of their inherent advantage in the consideration of connectivity relationships, we suggest that network analyses offer a simple, yet effective tool for searching for key segments (or junctions) in riverscapes for conservation and environmental management. Further, although the joint consideration of aquatic and terrestrial networks is challenging, the extension of network analyses to freshwater systems may facilitate the more effective selection of priority areas for conservation in continental areas.     

Functional conservation units for the endangered Alcon Blue butterfly Maculinea alcon in Belgium (Lepidoptera: Lycaenidae)

To organize and prioritise species-specific conservation efforts, we delineate `functional conservation units' for the threatened Alcon Blue butterfly Maculinea alcon in Belgium. We used detailed distribution data on the butterfly, its host plant and its habitat, present-day population sizes and its mobility and colonization capacity to determine functional conservation units (FCUs) on different spatial scales: FCU-1, i.e., the 12 presently occupied habitat patches plus the area within a range of 500 m surrounding them (the maximum local movement distance, based on mark-release-recapture data), FCU-2, i.e., the areas within a range of 2 km around the occupied habitat patches (the maximum observed colonization capacity) and FCU-3, i.e., potential re-introduction sites (sites where M. alcon went extinct recently). We suggest different management and planning measures for each type of functional conservation unit and discuss translocation and re-introduction as `intensive care' conservation measures for this threatened and sedentary species.     

Reserve selection using Red Listed taxa in three global biodiversity hotspots: Dragonflies in South Africa

The Red List can be used a gauging tool by conservationists to assess which species require focused conservation attention. Mapping the relative distributions of species, and identification of centers of richness, endemism and threat are a first step towards site-oriented conservation action. We use here a specially developed biodiversity index, based on three weighted sub-components assigned to each species: geographical distribution, Red List status, and sensitivity to habitat change. We test this approach using what is called here the Dragonfly Biotic Index (DBI) to prioritize sites for conservation action, with special emphasis on species occurrence in three global hotspots in southern Africa. Using a selected set of the 23 top prioritized sites, we compare the DBI’s performance to that of a rarity-complementarity algorithm. As with several other taxa, local endemism levels are highest in the Cape Floristic Region (CFR), while richness is highest in the north east, particularly in the stream systems of the Maputaland-Pondoland-Albany (MPA) hotspot. Red Listed Odonata species are also concentrated in the CFR, while richness is highest in the MPA hotspot. Site prioritization using the DBI reveals that CFR sites protect Red Listed taxa rather well, despite the fact that catchments are only partially protected. The DBI demonstrates high levels of redundancy in representing Red Listed species, in other words, the same species are represented in several catchments. The value in the DBI thus lies in maximizing redundancy (i.e. representation) of globally Red Listed species. The rarity-complementarity algorithm represents all species, but without greater emphasis on the rare and threatened (i.e. Red Listed) species. We conclude that the DBI is of great value in selecting biodiversity hotspots, while the algorithm is useful for selecting complementarity hotspots. We identify protection gaps and thus recommend continued searches in centers of endemism and existing reserves, as well as gap areas. These searches will hone Red List assessments and identify priority sites, as well as monitor already-identified sites for changes in quality of habitat.     

Long-term chemical and biological trends in Scottish streams and lochs

Between 1975 and 1985 several catchments in Scotland were established as reference sites to determine short-term (hours, days, weeks) and long-term (years, decades) changes in major physical, chemical and biological variables. The initial selection criteria included catchment size, land-use and deposition loading. Changes in fish status of lochs were determined by systematic netting while quantitative estimates of fish populations were made at stream sites. Egg survival and year class distribution assisted interpretation. Historical changes in pH based on stratigraphical changes in diatom assemblages are compared with recent measured changes which have occurred during the past 20 years when SO₂ emissions declined. Significant chemical and biological trends due to changes in pollutant loading are best detected in sensitive lochs which are unmanaged and with low organic content and acid neutralising capacity (ANC) close to zero. Significant trends are less apparent in lochs with managed (forested) catchments. Many stream sites also reveal long-term improvements in pH, sulphate (SO₄) Alkalinity (Alk) etc, but the significance of these trends is much less because of the greater inherent chemical variability. Stream sites close to ANC=0 provide the most wide-ranging evidence of ecological change, ie, increase in Alk, decrease in SO₄ and increase in numbers of juvenile salmonids. Measured chemical changes are compared with predicted estimates, based on critical load calculations and problems in interpretation are discussed.     

大通江、平洛河水保措施减水减沙分析

嘉陵江流域水土流失严重,“长治”工程首批治理的四大片区中,嘉陵江上游的陕南陇南地区和中下游土石山区两大片区名列其中。自20世纪80年代末开始实施水土保持治理以来,对控制流域内水土流失现象起到了较为明显的积极作用。选择了分别位于两大片区的大通江和平洛河两条典型河流,以实测水沙资料为依据,采用水文学方法,着重分析研究两个典型流域水保措施实施以来的水沙变化。     

Fragmented but not isolated: Contribution of single trees, small patches and long-distance pollen flow to genetic connectivity for Gomortega keule, an endangered Chilean tree

Habitat fragmentation is a major threat to species survival worldwide due to genetic isolation, inbreeding depression, genetic drift and loss of adaptive potential. However the data on how gene-flow changes following habitat fragmentation is contradictory. If there is significant gene-flow between spatially isolated populations then limited conservation resources could be directed away from projects to ‘establish genetic connectivity’ and used to address other consequences of habitat fragmentation.This research focused on an endangered tree species Gomortega keule (Gomortegaceae) in a fragmented landscape in the Central Chile Biodiversity Hotspot and addressed three questions: (1) How far does pollen move between pollen donors and seed trees and what is the shape of the dispersal curve? (2) Do insect pollinators travel outside of forest patches? (3) Do small populations and single trees contribute to genetic connectivity across the landscape?Paternity analysis results show that G. keule’s insect pollinators travel outside of forest patches, over distances of 6 km, beyond the scale of population fragmentation or genetic structure. Pollen moved from small sites and single trees into large sites, as well as in the other direction, indicating these sites play a key role as functioning elements of the wider population and as stepping stones between sites. Fragmentation at the scale investigated has not led to genetic isolation, thus genetic connectivity per se is not a conservation priority. Other consequences of land-use change, specifically continuing habitat loss and population reduction, still threaten the survival of the species.