Butterflies highlight the conservation value of hay meadows highly threatened by land-use changes in a protected Mediterranean area

Butterfly assemblages were used to characterize and evaluate the conservation value of the main habitat types in the Aiguamolls de l’Empordà Natural Park (north-eastern Spain), an important protected wetland area on the Mediterranean coast. Butterfly data were obtained from a number of transects walked as part of the Catalan Butterfly Monitoring Scheme, which uses a standardized methodology for monitoring butterflies. A Mantel test indicated a strong association between habitat types and the composition of butterfly assemblages and a principal component analysis ordinated individual butterfly species along a gradient from woodland to open areas, thereby indicating various degrees of shade tolerance. In addition, cluster analysis distinguished two main groups of habitats based on the similarities of their butterfly fauna: woodland and bramble clumps and a group of three different kinds of grasslands (traditionally hay meadows, pastures, and alfalfa fields). Hay meadows flooded in winter (the so-called closes) appeared always as the highest ranked habitat in terms of conservation evaluation: they have more butterflies and a slight tendency to harbor more and generally rarer species. This conclusion coincides with that of previous investigations that have indicated that the most diverse and rare plant communities in the whole Natural Park are present in the closes, and highlights the importance of traditionally managed hay meadows for wildlife. However, these hay meadows - one of the rarest habitats in the Mediterranean region - are in alarming decline and have become the most threatened habitat in this protected area: no longer profitable, we believe that the future of the closes will depend ultimately on the existence of agri-environmental schemes.     

Resource availability, matrix quality, microclimate, and spatial pattern as predictors of patch use by the Karner blue butterfly

Determination of which aspects of habitat quality and habitat spatial arrangement best account for variation in a species’ distribution can guide management for organisms such as the Karner blue butterfly (Lycaeides melissa samuelis), a federally endangered subspecies inhabiting savannas of Midwest and Eastern United States. We examined the extent to which three sets of predictors, (1) larval host plant (Lupinus perennis, wild lupine) availability, (2) characteristics of the matrix surrounding host plant patches, and (3) factors affecting a patch’s thermal environment, accounted for variation in lupine patch use by Karner blues at Indiana Dunes National Lakeshore, Indiana and Fort McCoy, Wisconsin, USA. Each predictor set accounted for 7-13% of variation in patch occupancy by Karner blues at both sites and in larval feeding activity among patches at Indiana Dunes. Patch area, an indicator of host plant availability, was an exception, accounting for 30% of variation in patch occupancy at Indiana Dunes. Spatially structured patterns of patch use across the landscape accounted for 9-16% of variation in patch use and explained more variation in larval feeding activity than did spatial autocorrelation between neighboring patches. Because of this broader spatial trend across sites, a given management action may be more effective in promoting patch use in some portions of the landscape than in others. Spatial trend, resource availability, matrix quality, and microclimate, in general, accounted for similar amounts of variation in patch use and each should be incorporated into habitat management planning for the Karner blue butterfly.     

An unseen foe in arthropod conservation efforts: The case of Wolbachia infections in the Karner blue butterfly

Infection by endosymbiotic bacteria is an underappreciated threat to endangered arthropods with serious implications for captive management programs. We examined the nature of Wolbachia infection in the North American endangered Karner blue butterfly, Lycaeides melissa samuelis, as a case study. Screening for Wolbachia across the range of the species confirmed widespread infection in the western half of the Karner blue’s range. Multilocus sequence typing using six genes confirmed that the infection in the western populations is attributable to a single strain of Wolbachia. This strain was also detected in the closely related Melissa blue butterfly, L. m. melissa, the presumed source of the infection. The infection in the Karner blue butterfly was perfectly correlated with the presence of a foreign mitochondrial DNA variant present in the Melissa blue butterfly, consistent with the hypothesis that the mitochondrial introgression was driven by the spread of Wolbachia. A single individual out of 71 screened from the eastern portion of the range of the Karner blue butterfly was also infected, however this infection was attributable to a different strain. Simulation models of the demographic effects of the spread of Wolbachia infection to uninfected populations and metapopulations suggest that such an infection might further reduce already small population sizes and substantially increase the probability of population extirpation. We discuss threats to other endangered arthropods in the light of this case study and make recommendations for minimizing the impact of endosymbiont infections in conservation plans, especially those including captive propagation and augmentation of endangered arthropod populations.     

Threshold effect of eucalypt density on an aggressive avian competitor

General guidelines available to revegetation planners focus on the spatial context and dimensions of the revegetated site. However, site-specific habitat factors can have overarching importance for habitat value, especially where interactions with competitors or predators may play an important role. Current revegetation projects in Australia which aim to restore slow-growing buloke (Allocasuarina luehmannii) woodland, a threatened habitat important for bird conservation, usually include faster-growing eucalypts in plantings. This research aimed to identify whether eucalypt presence in buloke woodland facilitates invasion by the noisy miner (Manorina melanocephala), an aggressive competitor that is absent from pure buloke woodland. Birds were surveyed in buloke woodland remnants that contained eucalypts as a sub-dominant species at densities of 0-16 per ha. The probability of noisy miner presence in buloke woodland increased markedly where eucalypts were present at a density of approximately five per hectare. The presence of noisy miners resulted in a substantial difference in bird assemblage structure and composition. Small-bodied insectivorous birds which are experiencing population declines in southern Australia were recorded on average six times more often in transects without noisy miners (low-eucalypt density transects). Avian behaviour and habitat use was also altered, with birds flying more frequently in transects where noisy miners were present. A minor difference in habitat composition results in substantial degradation of the conservation value of non-eucalypt woodland in eastern Australia due to invasion by aggressive avian competitors. Revegetation and restoration practices should take into account the potential for such subtle floristic differences to result in substantial variation in conservation outcomes.     

Protecting more than the wetland: The importance of biased sex ratios and habitat segregation for conservation of the Hine’s emerald dragonfly, Somatochlora hineana Williamson

Within species habitat use may depend on age, season or sex of an individual. The distribution of males and females may vary both temporally and spatially due to differences in the costs of reproduction and the distribution of critical resources. Conservation of a species requires knowledge of the habitat use of both sexes in order to predict the population size and protect all habitats that a species requires. Adult dragonfly populations often have highly male-biased sex ratios at the breeding habitat. This bias has been attributed to females using alternative habitats to avoid male harassment, or to high female mortality. We monitored adult Hine’s emerald dragonfly (Somatochlora hineana Williamson) populations, in breeding and non-breeding habitats in Door County, Wisconsin and found significant differences in habitat use between males and females. Males primarily used wetland habitats, while females primarily used dry meadows and marginal breeding habitats, only coming into wetlands to lay-eggs or find mates. We assessed food resources in the different habitats and found that high quality insect prey (primarily adult Diptera) were more available in the wetland habitat, indicating that these areas were likely a more productive foraging area for adult dragonflies. The fact that females appear to avoid the wetland habitat is consistent with the hypothesis that male harassment alters female distribution patterns. Consideration of the patterns of habitat use by S. hineana indicates the need to develop a broader understanding of the importance of non-wetland areas in the conservation of wetland species.     

Estimating the potential effects of sudden oak death on oak-dependent birds

Sudden oak death (SOD), a disease induced by the fungus-like pathogen Phytophthora ramorum, threatens to seriously reduce or eliminate several oak species endemic to the west coast of North America. We investigated how the disappearance of one of these species, coast live oak (Quercus agrifolia), may affect populations of five resident oak-affiliated California birds - acorn woodpecker (Melanerpes formicivorus), Nuttall’s woodpecker (Picoides nuttallii), Hutton’s vireo (Vireo huttoni), western scrub-jay (Aphelocoma californica), and oak titmouse (Baeolophus inornatus) - using geocoded data from Audubon Christmas Bird Counts, North American Breeding Bird Surveys, and the California Gap Analysis. Capitalizing on observed relationships between the focal bird species and both oak species diversity and areal extent, we modeled relative bird abundance while assuming complete loss of Q. agrifolia and complete, partial, or no loss of oak habitat following a disease sweep. Post-SOD projections of bird populations occurring within the range of coast live oak were on average 25-68% smaller and 13-49% more variable relative to pre-SOD estimates. SOD effects were greatest for habitats with low initial oak species diversity. Climatic SOD models predicted that the disease stands to negatively impact populations of all five focal bird species throughout 20% of California’s coast live oak habitats. This study provides the first spatially explicit insights into the potential effects of SOD on avian distribution and abundance. Results may be used to help prioritize conservation plans aimed at minimizing overall community level disturbances resulting from the disease.     

Is landscape context important for riparian conservation? Birds in grassy woodland

An important challenge for riparian management is to determine the extent to which landscape context influences the faunal assemblages of riparian habitats. We examined this challenge in the variegated landscapes of southeastern Queensland, Australia where riparian vegetation is surrounded by both extensive grazing and intensive cropping. We investigated whether riparian habitats adjacent to different landuses support similar bird assemblages. Three types of riparian habitat condition were sampled (uncleared ungrazed; uncleared grazed; cleared grazed) in four different land-use contexts (ungrazed woodland; grazed woodland; native pasture; crop) although only six of the 12 possible treatment combinations were available. Eighty percent of bird species responded significantly to changes in both riparian habitat condition and landscape context, while fewer than 50% of species were significantly influenced by landscape context alone. The influence of landscape context on the bird assemblage increased as the surrounding land use became more intensive (e.g., woodland to native pasture to crop). Riparian zones have been shown to have consistently high biodiversity values relative to their extent. These findings suggest it is not enough to conserve riparian habitats alone, conservation and restoration plans must also take into consideration landscape context, particularly when that context is intensively used land.     

Differential habitat selection by Pipistrellus pipistrellus and Pipistrellus pygmaeus identifies distinct conservation needs for cryptic species of echolocating bats

Cryptic species are similar in morphology, and make interesting subjects for relating morphological differentiation to ecological resource partitioning. Can species that are morphologically almost identical occupy different ecological niches, and hence potentially need distinct conservation planning? The discovery that the most widespread bat in Europe - the pipistrelle - comprised two cryptic species (Pipistrellus pipistrellus and Pipistrellus pygmaeus) that emit echolocation calls at different frequencies provides a remarkable model system for investigating links between morphology, echolocation call design and resource partitioning. We investigated resource partitioning between the two cryptic species of sympatric pipistrelle bats by radio tracking breeding females. Habitat selection was investigated by using compositional analysis. P. pygmaeus selected riparian habitats over all other habitat types in its core foraging areas, whereas P. pipistrellus, although preferring deciduous woodland overall, was more of a generalist, spreading its foraging time in a wider range of habitats. Although morphologically very similar, the cryptic species show quite different patterns of habitat use. Our findings suggest that large-scale differences in habitat preferences can occur between sympatric bat species that are virtually identical in flight morphology; hence morphological differences may be a weak indication of ecological differences between taxa. Conservation planning needs to take account of these differences to meet policy and legal obligations associated with these protected cryptic species.     

A multi-scale analysis of species-environment relationships: rare moths in a pitch pine-scrub oak (Pinus rigida-Quercus ilicifolia) community

Community level analyses of species-environment relationships can provide critical information for conservation planners. A comprehensive analysis of the factors that influence species distributions should include variables measured at multiple scales because species-environment relationships are known to vary with the scale of observation. The pitch pine-scrub oak (Pinus rigida-Quercus ilicifolia) communities, or pine barrens, of the northeastern USA are severely threatened by development and fire suppression. They also provide critical habitat for many species of rare moths. We used partial canonical correspondence analysis to assess the relative effects of three levels of environmental variables (plot, patch, and landscape) on the distribution and abundance of 10 species of rare moths in a pine barrens community in southeastern Massachusetts, USA. We also used a set of spatial variables to quantify and partial out the effects of spatial autocorrelation of species composition among sampling locations. All three levels of environmental factors combined, independent of spatial factors, accounted for virtually half (48.4%) of the total variation in the moth community. Sequential partitioning of the variance explained by each level of environmental factors indicated that landscape level factors explained more than twice as much variance as plot and patch level factors. Another environmental model that included only landscape level variables explained 53% of the total variation in the moth community. Patch density and percentage of the landscape comprised of open and sparse canopy, scrub oak habitats were the most significant variables. These results suggest that the presence of scrub oak habitat within relatively large, heterogeneous landscape mosaics may be more important for the maintenance of many rare pine barrens associated moth populations than plot or patch level characteristics.     

Trends associated with residential development in riparian breeding bird habitat along the Snake River in Jackson Hole, WY, USA: Implications for conservation planning

Throughout North America, bird population declines may be attributable to loss of habitat on the breeding grounds. Human land uses such as residential development, greatly impact landscapes and biota, particularly breeding birds. Riparian areas, which are essential to breeding birds, are particularly vulnerable to development pressures. In North America, most studies have focused on eastern birds and it is unclear how western bird assemblages respond to the effects of residential development. We sampled bird community parameters and habitat variables at three spatial scales (microhabitat, macrohabitat, and landscape) along a residential development gradient within the Snake River riparian corridor in Jackson Hole, WY, USA. Principal component analysis and multiple linear regression statistical tests were used to determine the effects of housing densities on avian community parameters, guilds, individual species distributions, and environmental variables. Landscape-level features were most affected by residential development and trends associated with increasing housing densities, such as anthropogenic habitat fragmentation primarily structured local bird communities. Overall species richness and diversity declined with increasing residential development. Neotropical migrant species were most negatively impacted and consistently declined in proportional representation on forested plots as residential development densities increased. Food generalists, ground gleaners, and avian nest predators all increased with increasing residential development. Brood parasites, on the other hand, did not increase with increasing housing densities and their distribution may reflect the availability of nest host species. These results suggest that residential development within riparian habitats may be exerting a strong negative influence on western bird communities and at high densities may lead to a depauperate avian biota. The results of our study indicate that preserving forest structure may benefit certain species but will not likely conserve as diverse and rich an avian assemblage as minimizing fragmentation associated with residential development.     

Large-scale habitat associations of birds in lowland Iceland: Implications for conservation

Iceland is responsible for many internationally important populations of breeding bird species, yet very little is currently known about how these species use the habitats available to them. Lowland areas of Iceland, in particular, have undergone significant landscape changes over the last century, such as widespread drainage of wetlands and conversion to agriculture, changes in grazing pressure and recently, extensive afforestation. The impact of these changes on breeding bird species will depend on the relative importance of different habitats for each species, and the threats facing those habitats. Here we report the results of a large-scale survey of the factors influencing patterns of habitat selection of eight populations of Charadriiform bird species throughout lowland Iceland; oystercatcher Haematopus ostralegus, golden plover Pluvialis apricaria, dunlin Calidris alpina, snipe Gallinago gallinago, whimbrel Numenius phaeopus, black-tailed godwit Limosa limosa, redshank Tringa totanus and arctic skua Stercorarius parasiticus. Ordination analyses and multiple logistic regression models are constructed to explore the components of habitats that influence the distribution of these species. Five of the eight species analysed showed significant preferences for lowland wetland habitats and four significantly selected areas containing wet features such as pools and high water tables. These results allow us to identify future conflicts in land use that are likely to result from government-supported large-scale afforestation of lowland areas and hydro-electric developments.     

Variation in wolf spider (Araneae: Lycosidae) distribution and abundance in response to the size and shape of woodland fragments

The wolf spider (Lycosidae) community of open paddocks and five different configurations of woodland habitat in the wheatbelt of New South Wales, Australia, was sampled by nocturnal spotlighting. A total of 16 species was detected amongst the 2769 individuals that were sufficiently mature to allow identification, and 80% of these individuals were accounted for by just three species. There were no significant differences in the spider community among any of the woodland configurations, but the community of all five configurations differed significantly from that of open paddocks. However, only three species differed significantly in abundance between paddocks and the woodland configurations. Two species, Venator spenceri and Sp. C appeared to be less common in paddocks than in the woodland habitats, while Sp. A was more abundant in the paddocks. The percentage cover of thistles, medium grass and high grass, as well as the total abundance of Callitris glaucophylla were the habitat variables that best explained variation in the wolf spider community between woodland sites. Of these, thistle cover had the strongest correlation suggesting that disturbance, rather than cover per se, might be an important determinant of wolf spider communities. This study indicates that habitat fragmentation, at the spatial scale associated with current agricultural practices, may not be presenting a threat to generalist ground predators such as wolf spiders.     

The influence of livestock grazing and weed invasion on habitat use by birds in grassy woodland remnants

Remnants of native vegetation in regions dominated by agriculture are subject to degradation, especially by livestock grazing and weed invasion. Ground-foraging birds are amongst the most threatened bird groups in Australia, and these agents of degradation might be contributing to their decline by causing a reduction in food availability. We studied the foraging behaviour and microhabitat use of seven species of ground-foraging insectivores in south-eastern Australian buloke woodland remnants with native, grazed and weedy ground-layers. If birds must resort to using more energetically expensive prey-attack manoeuvres, or selectively use substrates and microhabitats that are less available in degraded habitats, then such degradation is likely to be negatively impacting on these species. We found evidence of a negative impact of one or both of these types of degradation on five of the seven bird species. Three species that employ a range of foraging manoeuvres to attack prey used potentially more energetically expensive aerial manoeuvres significantly more frequently in weedy remnants than in remnants with a native or grazed ground layer. Red-capped robins Petroica goodenovii and brown treecreepers Climacteris picumnus both selectively foraged near trees in grazed sites, and hooded robins Melanodryas cucullata, red-capped robins and willie wagtails Rhipidura leucophrys avoided foraging in microhabitats with a high percentage cover of exotic grasses in weedy sites. Brown treecreepers were also less likely to be present in weedy sites that had been protected from grazing than in either grazed or native sites. These results suggest that although grazing appears to have a detrimental impact on foraging habitat of ground-foraging birds, the exclusion of livestock grazing from previously disturbed buloke remnants alone is not adequate to restore habitat values for ground-foraging birds. A conservation strategy for this habitat type should consider the exclusion of heavy grazing from sites with an intact cryptogamic crust and the management of weeds in disturbed remnants, potentially through the use of carefully controlled light grazing.     

Habitat structure and proximity to forest edge affect the abundance and distribution of forest-dependent birds in tropical coastal forests of southeastern Madagascar

Despite the fact that Madagascar is classified a biological `hotspot' due to having both high levels of species endemism and high forest loss, there has been no published research on how Madagascan bird species respond to the creation of a forest edge or to degradation of their habitat. In this study, we examined how forest bird communities and different foraging guilds were affected by patch habitat quality and landscape context (forest core, forest edge and matrix habitat) in the threatened littoral forests of coastal southeastern Madagascar. We quantified habitat use and community composition of birds by conducting 20 point counts in each landscape contextual element in October and November 2002. We found that littoral forest core habitats had significantly (p<0.01) more bird species than forest edge and matrix habitats. Thirty-one (68%) forest dependent species were found to be edge-sensitive. Forest edge sites had fewer species, and a higher representation of common species than forest interior sites. Twenty-nine species were found in the matrix habitat, and the majority of matrix-tolerant forest species had their greatest abundance within littoral forest edge habitats. Guild composition also changed with landscape context. Unlike other tropical studies with which we are familiar, we found that frugivorous species were edge-sensitive while sallying insectivores were edge-preferring. The majority of canopy insectivores (n=15, 88%), including all six endemic vanga species, were edge-sensitive. When habitat quality was assessed, the distributions of nine edge-sensitive species were significantly (p<0.01) affected by changes in habitat complexity and vegetation vertical structure in core or edge point counts. Therefore, we believe that changes in vegetation structure at the edge of littoral forest remnants may be a key indicator of mechanisms involved in edge sensitivity of forest dependent species in these forests. Our findings indicate that habitat fragmentation and degradation affect Madagascan bird communities and that these processes threaten many species. With continued deforestation and habitat degradation in Madagascar, we predict the further decline of many bird species.     

Coppice management effects on experimentally established populations of three herbaceous layer woodland species

Traditional coppice management creates a temporal release of resources. We determined how this affected three herbaceous species (Cardamine pratensis, Primula elatior and Geum urbanum) and if it was suitable for their conservation within woodland given their differing phenologies and habitat affinities for woodland.Reproductive adults were transplanted and their fate, i.e. survival and counts of leafs and flowers, plus the fate of their progeny, were monitored for three years following cutting of coppice shoots (three light levels) or yearly autumn mowing.Cardamine pratensis and P. elatior produced more flowers with increasing light availability. Mowing increased flower and leaf production with time for P. elatior. Seedling numbers followed a similar trend. Geum urbanum initially produced more flowers with increasing light and when mown, but the effect disappeared and did not increase seedling numbers. Its basal leaves showed the opposite pattern. Population growth rates (λ), calculated for P. elatior and G. urbanum, confirmed the strong treatment effects on the former and the absence of effects on the latter. Yet, decomposition of treatment effects, showed considerable flexibility in life history of G. urbanum, except for contributions of fecundity. The latter, however, contributed most to positive effects on λ for P. elatior.Early flowering species with an affinity for open habitats (C. pratensis > P. elatior) benefited more from temporal resource release than the later flowering, typical woodland species. Coppice management thereby maintains both typical forest herbs and herbs with affinity for more open habitats. This is an important conservation tool especially in intensively managed landscapes.     

Multi-scale factors affecting bird use of isolated remnant oak trees in agro-ecosystems

With recent emphasis on sustainable agriculture, conservation of native biota within agricultural systems has become a priority. Remnant trees have been hypothesized to increase biological diversity in agro-ecosystems. We investigated how remnant Oregon white oak (Quercus garryana) trees contribute to conserving bird diversity in the agro-ecosystem of the Willamette Valley, Oregon, USA. We compared bird use of isolated oak trees in three landscape contexts - croplands, pastures, and oak savanna reserves - and ranked the relative importance of four factors thought to influence bird use of individual trees: (i) tree architecture; (ii) tree isolation; (iii) tree cover in the surrounding landscape; and (iv) landscape context, defined as the surrounding land use. We evaluated species-specific responses and four community-level responses: (i) total species richness; (ii) richness of oak savanna-associates; (iii) tree forager richness; and (iv) aerial and ground forager richness. We documented 47 species using remnant oaks, including 16 species typically occurring in oak savanna. Surprisingly, landscape context was unimportant in predicting frequency of use of individual trees. Tree architecture, in particular tree size, and tree cover in the surrounding landscape were the best predictors of bird use of remnant trees. Our findings demonstrate that individual remnant trees contribute to landscape-level conservation of bird diversity, acting as keystone habitat structures by providing critical resources for species that could not persist in otherwise treeless agricultural fields. Because remnant trees are rarely retained in contemporary agricultural landscapes in the United States, retention of existing trees and recruitment of replacement trees will contribute to regional conservation goals.     

Do regional gradients in land-use influence richness, composition and turnover of bird assemblages in small woods?

Small patches of natural or semi-natural habitat have an important role in the conservation of biodiversity in human-dominated environments. The values of such areas are determined by attributes of the patch as well as its context in the surrounding land mosaic. There is a need for better understanding of the ways in which assemblages are influenced by patch context and the scale over which this occurs. Here we examine the influence of regional environmental gradients on the richness, annual turnover and composition of breeding bird species in small woods in south-eastern England. Regional gradients were defined independently of woods by an ordination of attributes for 5 km × 5 km landscape units across a 2100 km² region. Patch-level attributes, particularly area, were the most important predictors for most bird variables. For woodland migrants and woodland-dependent species, variables representing the context of each wood, either at a local or regional scale, explained significant additional variance in species richness after accounting for wood area, but did not do so for species turnover. Significant context effects for woodland-dependent species related to the extent of hedges and woodland cover in the local vicinity (<1 km radius), whereas for woodland seasonal migrants the best predictors of richness after patch area were two regional environmental gradients. The initial cue to settlement for migrants may be at a coarse regional scale, with selection for suitable landscapes that have a greater extent of woodland cover. Edge species showed different responses: they were influenced by the diversity of structural features in woods, and were a more-dominant component of the avifauna in isolated woods in open fenland environments of the region. Significant relationships between coarse regional gradients (25 km² units) and bird assemblages in small woods (0.5-30 ha) suggest that population and community processes in the avifauna operate across a broader scale than local patch neighbourhoods. They also highlight the importance of adopting a landscape or regional perspective on potential changes to land-use in rural environments, and on the conservation management of small reserves.