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.     

Beyond the wetland border: Estimating the impact of roads for two species of water snakes

We used models integrating road maps, traffic volume, and snake movements to examine the potential for roads to contribute to mortality in two species of water snakes that differ in their vagility, use of terrestrial habitats, and conservation status. Road networks and traffic volumes typical of three regions in Indiana, USA, may account for mortality of 14-21% of the population per year in the more vagile, terrestrial, and imperiled copperbelly water snake (Nerodia erythrogaster neglecta) but only 3-5% mortality in the more sedentary, aquatic, and common northern water snake (Nerodia sipedon). The majority (>91%) of road crossings and associated mortality are predicted to occur during overland migrations to other wetlands, suggesting roads bisecting travel routes between wetlands may function as mortality sinks. Our models highlight the proportionately greater risk of mortality for the more vagile and imperiled species, N. e. neglecta, and suggest current wetland conservation strategies that focus on the wetland alone are unlikely to adequately protect wetland biodiversity from certain types of anthropogenic habitat modification. What is needed is a landscape approach to wetland conservation that considers not only the quality of wetlands and nearby terrestrial habitats, but also ensures that terrestrial corridors between wetlands remain permeable and offer safe passage for wildlife.     

Pathogen exposure in endangered island fox (Urocyon littoralis) populations: Implications for conservation management

Island fox (Urocyon littoralis) populations on four California Channel Islands have declined severely since 1994. Canine distemper (CDV) was suspected to be responsible for the decline of the Santa Catalina Island fox, so knowledge of infectious disease exposure in the remaining island fox populations was urgently needed. This study reviewed previous pathogen exposure in island foxes and investigated the current threat by conducting a serologic survey of foxes on all islands and sympatric feral cats on three islands from 2001 to 2003 for antibodies against canid pathogens. Before the decline, foxes had evidence of exposure to CDV, canine adenovirus (CAV), canine parvovirus (CPV), and Toxoplasma, with exposure to these five pathogens differing greatly by island. Exposure to canine coronavirus (CCV), canine herpesvirus (CHV), and Leptospira was rare. In 2001-2003, wild-born foxes had evidence of exposure to CDV (5.2-32.8%) on 5 of 6 islands, CPV (28-100%) and CAV (4.7-100%) on five islands, and Toxoplasma gondii (2.3-15.4%) on four islands. Exposure to CCV, CHV and Leptospira was less common. Sharing of infectious agents between sympatric foxes and feral cats appeared minimal, but CDV exposure was detected in two cats on Santa Catalina Island. Domestic dogs have historically been present on the islands, but it is not known if canine diseases can be maintained in fox populations without the continual presence of dogs. Targeted vaccination programs against the most virulent pathogens and continued intensive disease surveillance may help protect the critically small remaining fox populations from disease outbreaks that could threaten the success of ongoing conservation efforts.     

Demographic consequences of anthropogenic influences: Florida black bears in north-central Florida

Anthropogenic habitat fragmentation poses a serious threat to conservation of large carnivores, due to their extensive movements and potential conflicts with humans. We studied the population ecology of Florida black bears (Ursus americanus floridanus) for 6 years in two study areas in north-central Florida: Ocala National Forest (ONF), a contiguous forested habitat, and an adjacent residential community of Lynne, a fragmented habitat with substantial human activities. We estimated age-specific survival and fecundity rates of bears using data from radio-collared bears, and parameterized and analyzed stage-structured matrix population models for the two study sites and also for data pooled from both sites. Annual survival rates of adult females were lower in Lynne (0.776 ± 0.074) than in ONF (0.966 ± 0.023). While cub survival rates were higher in Lynne (0.507 ± 0.135) than in ONF (0.282 ± 0.109), the rates at both sites were substantially lower than those reported for other black bear populations. Age-specific fecundities did not vary between sites. The asymptotic population growth rate for ONF was greater than one, whereas that for Lynne was less than one. Our results suggest that anthropogenic influences (primarily road density and vehicular traffic, through their effect on adult survival) can substantially affect the population dynamics of Florida black bears and other large carnivores with large home ranges. We recommend efforts such as constructing highway underpasses, which could reduce road-related mortalities, to ensure long-term persistence of Florida black bears facing threats from rapidly increasing human influences.     

Population baseline data for monitoring genetic diversity loss for 2010: A case study for Brassica species in the UK

The Convention on Biological Diversity and the subsequent International Treaty on Plant Genetic Resources for Food and Agriculture have proved a watershed in plant genetic resources (PGR) conservation and consequent initiatives have set various PGR conservation targets to be met by 2010. The aim of our paper is to develop ideas and issues concerning the monitoring of natural genetic resources: particularly in terms of developing a baseline from which to measure levels of genetic diversity. Three species of Brassica (B. nigra, B. oleracea, B. rapa) found wild in the UK were assessed for levels of genetic diversity using AFLP. The relationship between genetic distribution and ecogeographic distribution was considered for each species to determine patterns that may be useful in formulating conservation strategies. Genetic distance between populations of B. nigra and B. rapa were correlated to geographic distance. Levels of genetic polymorphism in B. oleracea were correlated to soil pH while in B. rapa they were correlated to soil coarseness. In terms of PGR conservation these findings may suggest an emphasis toward in situ conservation of a selection of disparate populations would be appropriate where possible as such adaptations may be lost in ex situ collections.     

Modelling populations of long-lived birds of prey for conservation: A study of imperial eagles (Aquila heliaca) in Kazakhstan

Eagle populations worldwide are in decline and their demography is generally poorly understood. We use novel sensitivity analysis of stochastic simulation models to analyse the demography of the worlds highest-density and longest-studied population of eastern imperial eagles (Aquila heliaca), at the Naurzum National Nature Reserve in Kazakhstan. Single variable perturbation (a simple elasticity-type analysis) showed that population growth was most sensitive to changes in adult survival but provided no information on how interactions between parameters may influence population growth. Multiple-variable perturbations (a more comprehensive elasticity-type analysis) suggested that population growth is relatively more sensitive to adult survival than is indicated by single-variable perturbation but also that when adult survival is within a biologically reasonable range, other parameters are still highly consequential to model outputs. For Naurzum’s imperial eagles, and for other structured populations of vertebrates, effective conservation and management likely requires an approach that addresses the importance of simultaneous variation in multiple vital rates including both survivorship and reproductive output.     

Host status of 10 fungal isolates for two nematode species, Filenchus misellus and Aphelenchus avenae

The classification of nematodes in the family Tylenchidae into plant parasites, plant associates or fungal-feeders for community analyses, have been much discussed by nematode ecologists. For an appropriate classification, fungal-feeding habits in the family need to be studied. To evaluate the host status of 10 fungal isolates for Filenchus misellus (Tylenchidae) and Aphelenchus avenae (Aphelenchida, Aphelenchidae), population growth rates, body length and width and sex ratios of the nematodes were measured after 40-day culture on fungal colonies at 25 °C. For F. misellus, the fungi determined as good hosts were two Basidiomycota fungi (Agaricus bisporus, Coprinus cinereus), three Ascomycota fungi (Chaetomium cochlioides, Chaetomium funicola, Chaetomium globosum) and a plant-pathogenic fungus (Rhizoctonia solani) on the basis of nematode population growth rate and female body length. Interestingly Pleurotus ostreatus, known as a predaceous fungus for the other nematodes, was also a good host for F. misellus. While, for A. avenae, good hosts were four plant-pathogenic fungi (Fusarium oxysporum f. sp. conglutinans, F. oxysporum f. sp. cucumerinum, Pythium ultimum, R. solani) and A. bisporus. A. avenae was trapped and preyed upon by Pleurotus hyphae. In F. misellus, males were 7-21% of adults, but the ratio did not correlate significantly with the population growth rate. In A. avenae, no male occurred. Differences in habitat preference between Filenchus and Aphelenchus were explained on the basis of the host status and habitat preferences of the tested fungi.     

Triage for conserving populations of threatened species: The case of woodland caribou in Alberta

Prioritization of conservation efforts for threatened and endangered species has tended to focus on factors measuring the risk of extirpation rather than the probability of success and cost. Approaches such as triage are advisable when three main conditions are present: insufficient capacity exists to adequately treat all patients, patients are in a critical state and cannot wait until additional capacity becomes available, and patients differ in their likely outcome and/or the amount of treatment they require. The objective of our study was to document the status of woodland caribou (Rangifer tarandus) herds in Alberta, Canada, with respect to these three conditions and to determine whether a triage approach might be warranted. To do this we modeled three types of recovery effort - protection, habitat restoration, and wolf control - and estimated the opportunity cost of recovery for each herd. We also assessed herds with respect to a suite of factors linked to long-term viability. We found that all but three herds will decline to critical levels (<10 animals) within approximately 30 years if current population trends continue. The opportunity cost of protecting all ranges by excluding new development, in terms of the net present value of petroleum and forestry resources, was estimated to be in excess of 100 billion dollars (assuming no substitution of activity outside of the ranges). A habitat restoration program applied to all ranges would cost several hundred million dollars, and a provincial-scale wolf control program would cost tens of millions of dollars. Recovery costs among herds varied by an order of magnitude. Herds also varied substantially in terms of their potential viability. These findings suggest that woodland caribou in Alberta meet the conditions whereby triage should be considered as an appropriate conservation strategy.     

Conservation benefits of biological control: The recovery of a threatened plant subsequent to the introduction of a pathogen to contain an invasive tree species

Biological control using host-specific natural enemies is sometimes the only long-term and cost-effective option to contain dominant invasive alien species in natural ecosystems. To assess biocontrol success, most monitoring studies focus on the negative impacts of the released biocontrol agents on the target invasive species. Very few studies have shown the conservation benefits, i.e. their positive effects on the recovery of native species and habitats. A fungal pathogen (Colletotrichum gloeosporioides f. sp. miconiae) was successfully released on the island of Tahiti (South Pacific) to control the dominant invasive tree Miconia calvescens which forms dense monospecific stands in native rain- and cloud forests. The pathogen causes a leaf disease leading to partial (5–35%) defoliation of Miconia canopy trees. We studied the demographical parameters of the rare endemic sub-shrub Ophiorrhiza subumbellata, growing in the understory of montane cloud forests invaded by Miconia. We compared Ophiorrhiza populations in 13 plots set up in two sites located between 950 and 1000 m elevation, having the same Miconia invasion degree but different levels of leaf damage. Our results showed that, whereas the total population density of Ophiorrhiza is not affected, plants’ growth and fertility were significantly higher in more defoliated Miconia forests. A 3-year-long survey of Ophiorrhiza population at one study site indicated that the number of seedlings and juvenile plants increased, suggesting better recruitment and survival. This study demonstrates that biological control has contributed to the recovery of an endemic plant directly threatened by an invasive species, and may be considered as an important management tool for the partial restoration of native forests.     

Role of human-modified habitat in protecting specialist species: A case study in the threatened Florida Scrub-Jay

Habitat specialists are especially vulnerable to habitat alterations in vegetation structure and composition, making them difficult to protect and restore amidst agricultural landscapes. Protection strategies in such cases require information on species’ ability to survive and reproduce on marginal and modified habitat. We examined reproductive success, survival, and foraging efficiency of the Florida Scrub-Jay (Aphelocoma coerulescens), an extreme habitat specialist, in a human-modified habitat - regenerating pasture - along a pasture-native scrub interface. From 1985 to 2003, Florida Scrub-Jays were equally successful at producing young in regenerating pasture and native scrub. Production of eggs, nestlings, fledglings, independent young, and yearlings were not statistically different between territories containing pasture and those containing only scrub. Similar trends were observed for nest success and survival. When in pasture, individual jays were significantly more efficient at capturing small prey items, but significantly less efficient at capturing medium and large prey items. Availability of small prey items was significantly higher in pasture than in scrub; availability of medium and large prey items was not significantly different between habitat types. We conclude that regenerating pasture provides suitable supplemental habitat for Florida Scrub-Jays when in close proximity to native scrub, suggesting a novel conservation strategy for maintaining and potentially increasing local populations in fragmented agricultural settings. This case study illustrates how a broadened definition of ‘suitable habitat’ may augment existing conservation strategies for specialist species facing substantial and rapid modification of their native habitat.     

Potential distribution modelling, niche characterization and conservation status assessment using GIS tools: a case study of Iberian Copris species

Dung beetle populations, in decline, play a critical ecological role in extensive pasture ecosystems by recycling organic matter; thus the importance of their conservation status. Presence data available for Copris hispanus (L.) and Copris lunaris (L.) (Coleoptera, Scarabaeidae) in Comunidad de Madrid (CM), and BIOMAPPER, a GIS-based tool, was used to model their environmental niches. The so derived potential distributions of both species were used to exemplify the utility of this kind of methodologies in conservation assessment, as well as its capacity to describe the potential sympatry between two or more species. Both species, distributed along a Dry-Mediterranean to Wet-Alpine environmental conditions gradient, overlap in areas of moderate temperatures and mean annual precipitations in the north of CM. Copris are poorly conserved in the existing protected sites network, but protection provided by new sites included in the future Natura 2000 Network will improve the general conservation status of these species in CM.     

Evaluating alternative rodenticides for island conservation: roof rat eradication from the San Jorge Islands, Mexico

Introduced commensal rats (Rattus spp.) are a major contributor to the extinction and endangerment of island plants and animals. The use of the toxin brodifacoum to completely eradicate rats from islands is a powerful conservation tool. However, brodifacoum is toxic to animals other than rats and on some islands its use may not be feasible without prohibitively expensive mitigation. As part of a regional conservation program, we experimentally tested brodifacoum and two less toxic rodenticides, diphacinone and cholecalciferol, in eradicating Rattus rattus from three small islands in the northern Gulf of California, Mexico. All three rodenticides were successful in eradicating rats, suggesting that the less toxic diphacinone and cholecalciferol may be useful alternatives to brodifacoum for some island eradication programs. However, the choice of rodenticide must be balanced between efficacy and the risks to non-target species. Applied field research is needed on less toxic rodenticides, as well as improving palatability of baits. This may prove invaluable in preventing extinctions and in restoring larger and more diverse island ecosystems.     

Population sizes, site fidelity and residence patterns of Australian snubfin and Indo-Pacific humpback dolphins: Implications for conservation

Very little is known about the ecology of snubfin Orcaella heinsohni and Indo-Pacific humpback dolphins Sousa chinensis in Australian waters. We used photo-identification data collected between 1999 and 2002 in Cleveland Bay, northeast Queensland, to estimate abundance, site fidelity and residence patterns of these species in order to make recommendations for their effective conservation and management. Our abundance estimates indicate that less than a hundred individuals of each species inhabit this coastal area. Even with relatively unbiased and precise abundance estimates population trends will be extremely difficult to detect in less than three years unless changes in population size are very high (>20% p.a.). Though both species are not permanent residents in Cleveland Bay, they both used the area regularly from year to year following a model of emigration and reimmigration. Individuals of both species spend periods of days to a month or more in coastal waters of Cleveland Bay before leaving, and periods of over a month outside the study area before entering the bay again. Because of their small population sizes and movement patterns, snubfin and humpback dolphins are particularly vulnerable to local extinction. Our results illustrate that: (1) detection of population trends should not be a necessary criterion for enacting conservation measures of both species in this region, and (2) efforts to maintain viable populations of both species in Cleveland Bay must include management strategies that integrate anthropogenic activities in surrounding areas.     

The decline of Cantabrian capercaillie: How much does habitat configuration matter?

The endangered Cantabrian capercaillie (Tetrao urogallus cantabricus) has undergone a severe decline during recent decades. Capercaillies require large tracts of mature forest, and are thus sensitive to landscape-level habitat alteration. The high degree of fragmentation and anthropogenic disturbances in the habitat of Cantabrian capercaillie suggested that habitat patterns may be related to the population decline. To evaluate this we developed predictive habitat suitability models. We used a geographic information system (GIS) to integrate environmental variables and spatial context into two logistic models, comparing (a) presence vs. pseudo-absence habitat units (General model) and (b) presence vs. abandoned habitat units (Decline model). We obtained low overall habitat suitability and poor connectivity between high-suitability areas. We found evidence that habitat suitability is indirectly related to the process of population decline, and that the relationship between capercaillie presence and habitat configuration goes beyond the local spatial scale and the average size of forest fragments. We suggest that the combination of these two modelling procedures provides a deeper insight into the decline process, differentiating overall optimal and suboptimal areas (General model) and predicting the direction of eventual local extinctions (Decline model).     

Relationship between population size and genetic diversity in endangered populations of the European bullhead (Cottus gobio): implications for conservation

This study investigated the relationship between the current size of endangered bullhead (Cottus gobio) populations and microsatellite genetic variability. Additionally, the microsatellite data were used to evaluate whether a genetic test for population bottlenecks was able to provide evidence of recent severe population declines. Finally, our results were used to develop conservation priorities and measures. Population size appears to be a crucial parameter in determining the amount of genetic diversity that can be preserved in bullheads, since a significant positive correlation was observed between both variables. Furthermore, in some populations we were able to detect genetic signatures of the documented decline in population size. We suggest that the most immediate goal for bullhead conservation should be to increase the size and the range of the populations, and in doing so minimise or even reverse further genetic erosion. Potential management actions like habitat quality improvement, reduction of river fragmentation and supplementation programmes (translocation, supportive breeding) are discussed.     

Microsatellite analysis of genetic population structure of the endangered cyprinid Anaecypris hispanica in Portugal: implications for conservation

The endangered fish species Anaecypris hispanica is restricted to eight disjunct populations in the Portuguese Guadiana drainage. The genetic structure of these populations was studied in order to determine levels of genetic variation within and among populations and suggest implications for conservation of the species. Based on five microsatellite loci, the null hypothesis of population homogeneity was tested. Tests for genetic differentiation revealed highly significant differences for pairwise comparisons between all populations, and substantial overall population subdivision (F_ST=0.112). All sampled populations contained unique alleles. Our findings indicate marked genetic structuring and emphasise limited dispersal ability. The high levels of genetic diversity detected within and among A. hispanica populations suggest, however, that the observed fragmentation and reduction in population size of some populations during the last two decades, has impacted little on levels of genetic variability. Data imply that most A. hispanica populations should be managed as distinct units and that each has a high conservation value containing unique genetic variation. It is argued that geographic patterns of genetic structuring indicate the existence of eight management units.     

Use of the Biological Flora framework in the United Kingdom Overseas Territories: Euphorbia origanoides L.

The United Kingdom Overseas Territories (UKOTs) are globally important for a high diversity of endemic and threatened plant species but are poorly represented in plant ecological literature. This lack of ecological research is compounded by a lack of funding and skills. Cost effective approaches of compiling conservation relevant information are required. Here we present the first examination of a species from the UKOTs presented within the standard framework of a Biological Flora. This framework allows a convenient way to compile ecological information and assess missing data. The account reviews all available information on Euphorbia origanoides L. (Ascension spurge) from Ascension Island (South Atlantic Ocean) relevant to understanding its ecology and conservation, including soil chemistry, climate and plant community data. E. origanoides is an endemic perennial, found in dry, lava plains of Ascension Island with soils comprised of weathered volcanic scoria. E. origanoides has suffered habitat loss through the introduction of invasive species and survival in the wild is currently under threat. We relate the information gathered for this Biological Flora to the conservation of the species in the wild and propose the framework should be used as one way of compiling information relevant for conservation managers. The framework is beneficial as it allows an evidence-based approach to conservation but also permits the prioritisation of research and can help conservation managers to meet targets for the Convention on Biological Diversity and the Global Strategy for Plant Conservation.     

Metapopulation dynamics and conservation of the marsh fritillary butterfly: Population viability analysis and management options for a critically endangered species in Western Europe

This study investigates the dynamics and viability of a marsh fritillary butterfly Euphydryas aurinia metapopulation in a Belgian successional landscape. Based on capture-mark-recapture and winter nest census data, we first estimated demography (survival and recruitment rates, population size, density dependence) and dispersal parameters (emigration rate, effect of patch connectivity on dispersal, mortality during dispersal). Then using RAMAS/GIS platform, we parameterised a population viability analysis (PVA) model with these parameters to simulate the future of this metapopulation under different scenarios.The metapopulation does not seem viable even if natural reforestation is controlled by adequate management. In its present state, the patch system is not able to sustain enough individuals: due to the large temporal fluctuations in demographic parameters, a carrying capacity far higher than currently would be necessary to limit extinction risk to 1%, suggesting the existence of an extinction debt for the species in Belgium. The situation of E. aurinia appears much worse compared to two other fritillary species threatened in Belgium, for which similar PVA are available. It is therefore urgent to increase the carrying capacity of the patch system. How and where it is achieved are of secondary importance for the gain in viability: improvement of habitat quality through restoration, or increase of habitat quantity via enlargement of existing patches and/or creation of new habitat in the matrix. A regime of management based on regular re-opening and maintenance of habitat patches may be the only guarantee of long-term persistence for this critically endangered species in Belgium.     

Factors influencing survival and long-term population viability of New Zealand long-tailed bats (Chalinolobus tuberculatus): Implications for conservation

A population viability analysis is important for the management of endangered populations and requires the estimation of survival parameters. The long-tailed bat (Chalinolobus tuberculatus) is one of only two native terrestrial mammals currently found in New Zealand and is classed as vulnerable. Its viability in temperate beech (Nothofagus) forest, Eglinton Valley, Fiordland, New Zealand was estimated using mark-recapture data collected between 1993 and 2003 using the Program MARK. Survival was estimated based on a total of 5286 captures representing 1026 individuals. Overall annual survival varied between 0.34 and 0.83 but varied significantly among three sub-populations and with sex and age. Females generally had a higher survival rate compared to males; and adults had higher survival relative to juveniles. Survival of all bats was lower in years when the number of introduced mammalian predators was high and when the winter temperature was warmer than average. High numbers of introduced predators occurred during three of the 10 years in the study. Climate change may mean that the conditions that promote high predator numbers may occur more frequently. A preliminary population viability analysis using a projection matrix on the overall adult female population showed an average 5% decline per year (λ = 0.95). Increased predator control targeting a range of predators is required in years when their numbers are high in order to halt the decline of this population of long-tailed bats. Population estimates using minimum number alive estimates supported the population estimates derived from Program MARK and a population viability analysis using matrices.     

Impact of grassland farming intensification on the breeding ecology of an indicator insectivorous passerine, the Whinchat Saxicola rubetra: Lessons for overall Alpine meadowland management

The decline of insectivorous farmland passerines has been attributed mostly to global decrease in arthropod availability, as a result of intensification of agricultural practices. The diminution of the Alpine Whinchat, once a widespread insectivorous passerine, has been ascribed to nest losses due to earlier and more frequent mowings. However, potential conjugated effects of deteriorated arthropod food availability had yet to be investigated. We compared food supply and nestling diet in intensively vs. traditionally managed grassland. Abundance and diversity of arthropods were much lower in intensive areas, where small-sized invertebrates, which do not enter nestling diet, were also predominant. Parents breeding in intensive habitats fed less biomass to nestlings than adults from traditional habitats. Nestling diet was less diverse and dominated by less profitable prey items in intensive than in traditional habitats. Feeding rate did not differ between the two habitats, but foraging distances from nest tended to be greater in intensive farmland. There were no significant differences in clutch sizes and hatching success with respect to management intensity, but fledging success was higher in traditional habitats. The recent intensification of farming practices has led to a decrease in the availability of grassland invertebrates, and of important Whinchat nestling food in particular, affecting parents’ foraging efficiency and reproductive success. Conservation actions must not only reduce nest losses by postponing mowing, but should also promote grassland farming that is less detrimental to invertebrates. Dominant at the study site, organic grassland farming does seemingly not provide sufficient conditions for Alpine Whinchats.