Swarming of bats at underground sites in Britain—implications for conservation

We investigated the extent of visitation of underground sites during late summer and autumn (‘swarming’) by bats. Bats were captured at eight cave, mine and tunnel sites in southern England over 6 years (1995-2000). A total of 3077 bats representing 11 of Britain's 16 species was caught. Myotis bats predominated in autumn. At some sites these are rarely seen during winter hibernation counts. Myotis nattereri and M. daubentonii were most common. However, species composition changed seasonally. At one site, M. brandtii dominated early in the season and at all sites peak activity of M. daubentonii was earlier than in M. nattereri. Relatively high numbers of internationally vulnerable species such as M. bechsteinii and Barbastella barbastellus were recorded. There was a strong male bias in captures of swarming species. Swarming probably has an important reproductive function, most males are reproductively active at this time and such sites may be important for outbreeding. Due to this and the large number of individuals and species concerned, the conservation of ‘swarming sites’ is vitally important. Clearly swarming sites merit special consideration in conservation management strategies for cave-dwelling bats.     

Cave selection and use by swarming bat species

Caves are an important resource to a large proportion of temperate bat species, primarily as mating and hibernation sites. However, information on bat use exists for only a small fraction of caves in many parts of the temperate world, in part because current survey methods are inappropriate for many species. We surveyed the UK’s largest karst landscape, the Yorkshire Dales, by monitoring autumn swarming (mating) activity at 53 caves using automated echolocation call loggers, followed by trapping. Over 60% of caves surveyed were used by bats, but there was considerable variation in activity. Swarming activity was positively correlated with chamber development and negatively correlated with the amount of water the cave carried: together these two predictors explained 45% of the variance. Entrance orientation and shelter explained a further 10%. Activity was not correlated with entrance size, altitude or connectivity to the nearest summer habitat. All five resident swarming species were caught at most sites: Myotis brandtii, Myotis daubentonii, Myotis mystacinus, Myotis nattereri and Plecotus auritus. Behaviour, species composition and sex and age ratios were typical of swarming reported in other studies. Bat populations and catchment sizes were large and despite the proximity of other suitable caves, bats showed high fidelity to single sites. Our results show that the national importance of these caves has been overlooked. We suggest that survey for swarming activity is a quick and effective method of identifying underground bat sites and is likely to uncover many previously unknown and important sites throughout the temperate world.     

Conservation of bats in suburban landscapes: roost selection by Myotis yumanensis in a residential area in California

Protection of roosting habitat is essential to the conservation of bats in human-dominated landscapes. To help define bat roosting needs in suburban settings, we used radio telemetry to locate day roosts of a common North American species (Myotis yumanensis) within a residential area in California. Between June and August 2000, we tracked 16 bats to 20 roosts in two buildings and 18 trees. We used multiple logistic regression to assess roost selectivity at multiple spatial scales. Of 15 tree, plot, and site characteristics considered, only three helped distinguish roosts from random comparisons: tree diameter, distance to water, and forest cover. Myotis yumanensis preferred large trees (mean diameter 115 cm), and roosted only in the five species of largest mean diameter (Sequoia sempervirens, Pseudotsuga menziesii, Quercus agrifolia, Quercus lobata, and Acer macrophyllum). At the site level, these bats selected roosts that were close to water and had substantial forest cover in the surrounding 100-m radius. Unlike other North American bats, they often roosted in live trees (89% live). Relatively high roost fidelity (mean 4.8 days) and large travel distances between consecutive roosts (mean 1130 m) and between capture sites and roosts (mean 2007 m) may indicate a greater degree of roost limitation at this site versus other research sites. We recommend the preservation of large trees and forested parkland, particularly along stream corridors, to help maintain bat populations in urbanizing landscapes.     

Autumn swarming behaviour of Natterer’s bats in the UK: Population size, catchment area and dispersal

Many species of temperate bats visit underground sites in autumn, prior to hibernation, and chase each other in and around the entrances in an event known as autumn swarming. It has been suggested that swarming is a mating event that facilitates gene flow between bats from otherwise isolated summer colonies. We describe swarming behaviour at a cluster of four caves in the North of England and provide direct evidence for seasonal migration of Natterer’s bats (Myotis nattereri ) between these caves and many summer nursery roosts in a catchment area of at least 60 km radius. Mark-recapture analysis suggests that an estimated 2000-6000 Natterer’s bats use the three best-studied caves, together with smaller numbers of four other species: M. daubentonii, M. brandtii, M. mystacinus and Plecotus auritus. Recaptures also show that there is a small interchange of bats between these three caves, which are 0.6-1.5 km apart. Capture rates, automatic logging and video recordings at one cave suggest that an average 310-400 bats visit the cave each night. Mating at swarming sites would be particularly important for gene flow if a species showed no other dispersal pattern. We provide evidence for natal philopatry, with little or no dispersal, in both sexes of Natterer’s bats. The protection of swarming sites is discussed in light of our findings.     

Scale-independent significance of river and riparian zones on three sympatric Myotis species in an agricultural landscape

Conserving biodiversity in agricultural landscapes is an urgent issue. The effective conservation of biodiversity requires plans based on species’ habitat preferences at multiple spatial scales. We examined how the foraging habitat selection of bats varied with grain size (50, 150, 250, and 350 m) and how habitat selection in the home range differed from that in the foraging habitat in an agricultural landscape. Focusing on three sympatric Myotis species (Myotis petax, Myotis gracilis, and Myotis frater), we radio-tracked 10 individuals of each species for 121 nights in Hokkaido, northern Japan. Rivers and broadleaved riparian forests were commonly preferred at multiple scales, although the preferred land-cover type was dependent on both extent and grain size for most species. The best grain sizes for predicting the foraging-habitat use of M. petax and M. gracilis were 50 and 150 m. By contrast, M. frater showed no tendency across the grain sizes. Our results indicate that it is necessary to consider both extent and grain size to understand the habitat selection of bats. Our findings also suggest that focusing primarily on preferred land-cover types at multiple scales is effective for conservation planning, given the limited resources in terms of time, manpower, and finances. Although arable did not negatively affect the selection of foraging habitats and home ranges in these bat species, preventing the loss of rivers and forests should be prioritized over preventing arable land expansion.     

Bat activity on riparian zones and upper slopes in Australian timber production forests and the effectiveness of riparian buffers

Unlogged buffers are used to ameliorate impacts to riparian areas in timber production forests. One function of these buffers is to protect the biodiversity of riparian areas. We measured bat activity in buffered streams with ultrasonic detectors across four different stream orders in logged, regrowth and mature forests (60 sites). Bat activity, foraging rates and species richness were similar in buffered streams surrounded by logged, regrowth and mature forests, suggesting that riparian areas effectively provide habitat for foraging and commuting bats in selectively logged forests. Vespadelus pumilus was the only species that responded to logging history, with decreased activity in mature forests. We found higher activity on larger rather than smaller order streams, a pattern also not affected by logging history. Bat activity along paired forest trail flyways on upper slopes (60 sites) was measured simultaneously with riparian flyway activity (for a total of 120 sites) to determine the importance of riparian areas relative to other available flyways. Activity was higher on upper slopes than on small streams, but similar to levels on larger streams. Total foraging activity was similar between riparian zones and upper slopes. Upper slopes contained higher species diversity, with Chalinolobus gouldii, Miniopterus schreibersii, Mormopterus norfolkensis, Scotorepens sp., Vespadelus pumilus and, to a lesser extent, Vespadelus darlingtoni detected more often than along streams. Other species (Rhinolophus megaphyllus, Nyctophilus spp. and Miniopterus australis) were not affected by topography. Estimates of total vegetation cover and, in particular, rainforest cover, were negatively associated with bat activity, highlighting the need for management of forest ‘clutter’ in regrowth forests for a suite of bat species. Streams and forest trails provide areas of lower clutter, which assist in maintaining high species diversity in regrowth forests. Our results support the use of riparian buffers, and point to the need for greater recognition of tracks on upper slopes as important habitat.     

Monitoring trends in bat populations through roost surveys: methods and data from Rhinolophus hipposideros

Volunteers counted lesser horseshoe bats (Rhinolophus hipposideros) emerging from 79 roosts throughout Wales prior to parturition over a 5-year period. Analysis of the count data and environmental variables revealed that there was no statistically significant change in the number of bats counted during the monitoring period. A maximum count of 5118 lesser horseshoe bats for one of the years of the study, suggests a Welsh population of >10,000. Within years, date had a strong influence on roost count with higher counts occurring later in the year. Analyses of roosts for which data was available in most years failed to find any evidence of changes in number of emerging bats over time either between roosts or across geographical regions. Over the time period studied, the population of R. hipposideros was stable. Power analysis suggested that the present monitoring programme should be able to detect a 5% increase or decrease over a 5-year period and could detect smaller trends over longer periods of time even when some data were missing. Roost surveys can therefore be a valuable tool to assess the status of a bat species, alert conservationists to population declines and provide feedback regarding the success of conservation management for bats.     

The importance of cenotes in conserving bat assemblages in the Yucatan, Mexico

In the Yucatan, cenotes or water sinkholes formed by the dissolution of limestone and surrounded with a characteristic dense layer of heterogeneous vegetation, are the main sources of water for plant and animal assemblages. We investigated their importance to bats by comparing the assemblage structure and diversity between tropical forest and pastureland, with and without cenotes. We set ground level mist nets, sub-canopy nets and harp traps for 96 nights and caught 2819 bats of 26 species and six families. Forest had a greater bat species diversity and abundance than pastureland, as well as rare and threatened species. In forest, sites with and without cenotes had a similar bat assemblage structure and diversity, whereas in pastureland there was a greater species diversity and abundance and number of rare and threatened species at cenotes than sites without cenotes. Chrotopterus auritus and Micronycteris schmidtorum, both threatened in Mexico, and Eptesicus furinalis, were exclusively captured at cenotes. We suggest that it may be the vegetation around cenotes rather than the water bodies per se that attracts many bat species in pastureland. Cenotes are threatened by nitrate contamination from untreated domestic and animal waste and fertilizers and the removal of vegetation to allow livestock access to water. In Yucatan’s agricultural landscapes, protection of cenotes and their characteristic vegetation is key not only for bat conservation but also because they represent potential seed sources for the regeneration of the lowland forest, in which bats play key roles as pollinators and seed dispersers.     

Species-specific barrier effects of a motorway on the habitat use of two threatened forest-living bat species

It is generally recognized that roads can adversely affect local animal populations but little is known how roads affect bats. In particular, no study compared the response of bats that differ in foraging ecology to motorways that cut through the breeding habitat. As bats are key species in conservation, such data are urgently needed for designing management plans. Using radio-telemetry, mist netting, and mark-recapture data we investigated the effects of a motorway with heavy traffic on the habitat use of two threatened forest-living bats. We compared barbastelle bats (Barbastella barbastellus), which forage in open space, to Bechstein’s bats (Myotis bechsteinii), which glean prey from the vegetation. Five of six radio-tracked barbastelle bats crossed the motorway during foraging and roost switching, flying through underpasses and directly over the motorway. In contrast, only three of 34 radio-tracked Bechstein’s bats crossed the motorway during foraging, all three using an underpass. Bechstein’s bats, unlike barbastelle bats, never crossed the motorway during roost switching. Moreover, only in Bechstein’s bats individuals foraging close to the motorway had smaller foraging areas than individuals foraging further away, whereas other forest edges had no such effect. Our data show that motorways can restrict habitat accessibility for bats but the effect seems to depend on the species’ foraging ecology and wing morphology. We suggest that motorways have stronger barrier effects on bats that forage close to surfaces than on bats that forage in open space, and discuss the implications of our findings for bat conservation during road construction.     

Intra-specific effects of forest loss on the distribution of the forest-dependent northern long-eared bat (Myotis septentrionalis)

Forest fragmentation by the expansion of human activities is acknowledged as an important factor driving declines of forest species worldwide and quantitatively characterizing this trend is essential to understanding the response of populations of forest-dependent species. To examine the impacts of forest cover loss and changes in composition on forest-dependent bats, we examined the effects of these factors on the distribution of the northern long-eared bat (Myotis septentrionalis) in a case study. We surveyed forest fragments in the forest-agricultural landscape of Prince Edward Island, Canada, to assess the occupancy of bats in fragments. Logistic regression and multimodel inference using Akaike’s information criteria were used to identify potentially important predictor variables influencing the distribution of the northern long-eared bat at the fragment and landscape level and quantified their effects. A compositional variable, area of deciduous stands, was found to be the best predictor of the presence of M. septentrionalis. We found the effects of fragmentation were greatest at the fragment level for females and at the landscape level for males. This case study emphasizes the importance of examining intra-specific resource selection in how it affects the response of a forest-dependent species to fragmentation.     

Conservation of Pteropus livingstonii based on roost site habitat characteristics on Anjouan and Moheli, Comoros islands

Livingstone's fruit bats, Pteropus livingstonii, (Pteropodidae: Chiroptera) are endemic to only two islands, Moheli and Anjouan of the Comoros Islands in the West Indian Ocean. With an estimated population of less than 2000 individuals, P. livingstonii is listed as Critically Endangered on the International Union for the Conservation of Nature's Red List. Yet the habitat characteristics of these giant fruit bats, or flying foxes, are poorly known. I conducted field research during 5 months in 1998 focused on finding new roost sites, measuring characteristics of both roost sites and roost trees at new as well as previously identified roost sites, and recording the same characteristics for a set of absence sites. I identified six new roost sites allowing me to measure the characteristics of 15 known roost sites and seven absence sites. The study spanned both dry and rainy seasons and I found a significant increase in the number of individuals at roost sites during the rainy season. Bat roost sites were associated with presence of water, southeast aspect, steep slopes, and presence of several native tree species. Roost habitat was characterized based on biotic, abiotic and human factors. This research on habitat characteristics is contributing to the development of a conservation action plan for the Livingstone's fruit bats. However, further research on dry season roost sites and distribution as well as feeding sites is necessary in order to better understand their seasonal habitat requirements and therefore ensure the conservation of P. livingstonii.     

Patterns of roost use by bats in a neotropical savanna: implications for conservation

Neotropic bats comprise a highly species-rich group and are considered important regulators of complex ecological processes. Resource partitioning of roosts and food is considered a key element in shaping bat communities. In this paper we examine the roosting patterns of 10 bat species belonging to a neotropical bat community in a Bolivian savanna. Roosts were more common in open woodlands than in forest islands and human settlements. In open woodlands, Tabebuia heptaphylla comprised the largest proportion of roosting trees, whereas in forest islands, Gallesia integrifolia was the most important roosting tree for phyllostomid bats. By comparing active roosts with control holes in Tabebuia heptaphylla, we found that roost site selection by Noctilio albiventris was mainly based on tree diameter and inner width of the cavities. The earlier results are discussed in view of resource partitioning within savanna bat communities and their practical conservation.     

European bat lyssaviruses: Distribution, prevalence and implications for conservation

Worldwide, there are more than 1100 species of the Order Chiroptera, 45 of which are present in Europe, and 16 in the UK. Bats are reservoirs of, or can be infected by, several viral diseases, including rabies virus strains (in the Lyssavirus genus). Within this genus are bat variants that have been recorded in Europe; European bat lyssavirus 1 (EBLV-1), European bat lyssavirus 2 (EBLV-2) and, four currently unclassified isolates. Since 1977, 783 cases of EBLVs (by isolation of viral RNA) have been recorded in Europe. EBLV-1 or EBLV-2 has been identified in 12 bat species, with over 95% of EBLV-1 infections identified in Eptesicus serotinus. EBLV-2 is associated with Myotis species (Myotis daubentonii and Myotis dasycneme). A programme of passive surveillance in the United Kingdom between 1987 and 2004 tested 4871 bats for lyssaviruses. Of these, four M. daubentonii (3.57% of submitted M. daubentonii) were positive for EBLV-2. Potential bias in the passive surveillance includes possible over-representation of synanthropic species and regional biases caused by varying bat submission numbers from different parts of the UK. In 2003, active surveillance in the UK began, and has detected an antibody prevalence level of 1-5% of EBLV-2 in M. daubentonii (n = 350), and one bat with antibodies to EBLV-1 in E. serotinus (n = 52). No cases of live lyssavirus infection or lyssavirus viral RNA have been detected through active surveillance. Further research and monitoring regarding prevalence, transmission, pathogenesis and immunity is required to ensure that integrated bat conservation continues throughout Europe, whilst enabling informed policy decision regarding both human and wildlife health issues.     

Relating bat species presence to simple habitat measures in a central Appalachian forest

We actively sampled the bat community at 63 sites using detection and non-detection metrics on the Fernow Experimental Forest (FEF) in the central Appalachians of West Virginia using Anabat acoustical equipment May-June 2001-2003 to relate species presence to simple habitat measures such as proximity to riparian areas, forest canopy cover, forest canopy gap width, and forest canopy height. We acoustically detected eight species on the FEF, including the endangered Myotis sodalis. The presence of Lasiurus cinereus, M. lucifugus, M. sodalis, and Pipistrellus subflavus was associated more with riparian areas than upland areas. Both univariate comparisons and multiple logistic regression modeling showed that the probability that clutter-adapted foraging species such as M. septentrionalis and M. sodalis would be detected was greater as forest canopy cover increased or forest canopy gap size decreased, whereas the opposite was true for open-adapted foraging species such as Eptesicus fuscus and L. cinereus. The overall proportion of unidentifiable bat echolocation sequences to those identified to species was related to upland sites with increasing forest canopy cover indicating some sampling bias between cluttered and uncluttered habitats. However, given sufficient sample points, bat community surveys using acoustical detection show the ability to quickly develop generalized habitat associations for rugged areas such as the central Appalachians where traditional mist-net survey efforts often are logistically difficult and are lacking in scope. Moreover, these acoustical surveys also could lend themselves to species-specific predictive mapping of foraging habitat as well as allowing researchers to formulate testable hypotheses about detailed bat habitat relationships to be definitively tested with radio-telemetry techniques.     

Sex-specific roost movements and population dynamics of the vulnerable long-fingered bat, Myotis capaccinii

The roosts of many IUCN-listed cave-roosting bat species are under threat from tourist development in SE Europe and other regions of the world. Much-needed conservation strategies require, among other information, an understanding of their roost movements and population dynamics, which can now be obtained relatively quickly using advanced models. We have studied the long-fingered bat, Myotis capaccinii, an obligate cave-dweller, in Dadia National Park, Greece. The species formed colonies of up to a few thousand individuals and was highly mobile, frequently switching summer roosts up to 39 km apart, even during late pregnancy. The bats migrated to distant hibernacula including a cave in Bulgaria 140 km NW of the Park. Adult recapture probabilities varied with season and sex: low female recapture rates in autumn, relative to spring and summer, indicated non-random temporary emigration following nursery colony dispersal. The opposite pattern was seen in males: increasing recapture rates in the autumn suggest that males gather in these roosts to mate with females in transit. Adult survival (0.86-0.94) was similar in females and males, similar in winter and summer, and comparable to recent estimates for other bats based on similar modelling techniques. Sex-based differences in juvenile recapture suggest female philopatry and male-biased dispersal. Our work shows that protection of M. capaccinii roosts must extend beyond the Park’s and indeed the country’s boundaries: its conservation requires large-scale, trans-national integrated conservation plans. Our results will apply to many other warm-temperate species with similar life history cycles.     

Changes of isopod assemblages along an urban–suburban–rural gradient in Hungary

Responses of isopod assemblages to urbanisation were studied along an urban–suburban–rural gradient representing a decrease in the intensity of human disturbance. Pitfall trapping collected six species (Armadillidium vulgare, Porcellio scaber, Porcellium collicola, Trachelipus ratzeburgii, Cylisticus convexus, and Trachelipus rathkii). A. vulgare occurred abundantly in all sites reflecting the broad tolerance and invasive nature of this species. Indicator species analysis demonstrated that P. scaber and T. rathkii were significant quantitative character species for the urban site, while T. ratzeburgii was characteristic for the natural habitats (suburban and rural sites). CANOCO revealed that ground and air temperature show positive correlation with the distribution of P. scaber and T. rathkii, and negative correlation with T. ratzeburgii. Nested ANOVA on trap level showed that there were no significant differences between the number of isopod species and individuals, and the diversity of isopod assemblages in the three studied areas. Significant differences were observed at site level. The results did not support the hypothesis that diversity should decrease in response to habitat disturbance. They also contradicted the intermediate disturbance hypothesis; species richness was not the highest in the moderately disturbed suburban area. Multivariate methods detected that the isopod assemblages of the rural and suburban areas were relatively similar, while that of the urban area was relatively separated.     

Bat activity affected by sewage effluent in Irish rivers

Bat activity and benthic macroinvertebrates were sampled at nine paired sites, upstream and downstream from sewage effluent discharges into Irish rivers. Bat activity was measured using broadband acoustic detectors and macroinvertebrates by three 30-s standard benthic kick samples per site. Biological indices of water quality were significantly lower downstream from sewage outfalls, relative to upstream. The soprano pipistrelle (Pipistrellus pygmaeus) was significantly more active (as measured by bat passes per unit time) at downstream sites, while Daubenton’s bat (Myotis daubentonii) was less active. These results contrast with those of a similar study in England, where P. pygmaeus were less active, and Myotis spp. were more active downstream from sewage outfalls. We suggest that P. pygmaeus were more active downstream in our study because of a preference for preying on small orthocladiinid Chironomidae (non-biting midges), which were significantly more abundant downstream. M. daubentonii may prefer Trichoptera (caddis fly), which were significantly more abundant upstream. Organic pollution may then affect bats, but its effect appears to be more complex than previously implied. Implications of changing nutrient levels in freshwaters for populations of M. daubentonii and P. pygmaeus may be different from what is currently suggested in the literature.     

Bat diversity in montane rainforest and shaded coffee under different management regimes in southeastern Chiapas, Mexico

Differences in alpha and beta bat diversity among montane rainforest and five shaded coffee plantations under different management regimes, as well as some environmental factors and vegetation parameters influencing bat richness, were evaluated for the first time in southeastern Chiapas, Mexico. In each site, bats were captured every 2 months from March 2004 to July 2005, with six mist-nets, during two nights, using the capture-recapture method. We captured 2970 individuals of 43 bat species. Montane rainforest had the greatest alpha diversity (H′ = 2.681; n = 37), whereas alpha diversity was similar among coffee plantations (H′ = 2.229-2.364; n = 23-26). The number of frugivorous and nectarivorous species was similar among the sites; the greatest exchange in species composition (beta diversity) occurred for insectivorous bats, which reduce their number in coffee plantations as pesticides are incorporated. Bat richness species was significantly related to the number of vegetation strata, height, and cover of trees. We suggest that coffee plantations could act as corridors, facilitating connection among different elements of the landscape in the Sierra Madre de Chiapas for some frugivorous and nectarivorous bats.     

Bats in continuous forest, forest fragments and in an agricultural mosaic habitat-island at Los Tuxtlas, Mexico

In order to document the responses of bats to destruction and fragmentation of their natural habitat and the value of different types of man-made vegetation for bat conservation in the Neotropics, bats were sampled with mist nets to compare species richness and species composition in a tract of continuous forest, forest fragments and a habitat-island consisting of a mosaic of forest and arboreal crops in Los Tuxtlas, southern Mexico. We captured 3835 bats representing 39 species: 76% were captured in continuous forest, 74% in the mosaic habitat and 87% in forest fragments. In the mosaic habitat we captured 43% of the total number of bats, 33% in the forest fragments and 24% in continuous forest. On average the habitats studied had 64% species in common. Evidence of continuous breeding activity was determined for a high number of species at the three habitats (> 70% lactating and 65% with embryos). A few bat species (Carollia brevicauda, Pteronotus parnelli, Sturnira lilium, Artibeus jamaicensis, Dermanura phaeotis, Vampyrodes caraccioli, Glossophaga soricina, Dermanuta toltecus, Cheoroniscus godmani, Platyrrhinus helleri) dominated the sample, but their relative dominance varied among habitats. Recapture of bats provided evidence for inter-habitat movement. The co-occurrence of the three habitats helps conserve a diverse assemblage of bat species in the local landscape.     

The impact of clonality on an endangered tree (Elaeocarpus williamsianus) in a fragmented rainforest

The occurrence and distribution of clonality in the endangered rainforest tree Elaeocarpus williamsianus (Elaeocarpaceae) was investigated using SSR and RAPD analyses for 170 apparent individual trees found across seven sites. The results obtained with the two molecular techniques were in complete agreement in showing that single clones are present in most of the E. williamsianus sites with two genets occurring at the largest and most adequately protected site. In addition, seed production, viability and germinability were determined for four of these populations. Fruit were produced in all four populations tested although sterile fruit were very common. Only two E. williamsianus trees representing different genets within the same site produced viable seed. The overall genetic diversity within E. williamsianus is much lower than expected and thus the potential for sexual reproduction has been significantly diminished. It is concluded that habitat fragmentation removed an existing balance between vegetative and sexual reproduction in this species. Such findings have added urgency to the management of this species which could include a reintroduction program incorporating the collation of all clones within selected sites.