Building large-scale spatially explicit models to predict the distribution of suitable habitat patches for the Greater rhea (Rhea americana), a near-threatened species

We developed large-scale spatially explicit models to predict the distribution of suitable habitat patches for the Greater rhea (Rhea americana), a near-threatened species, in two areas of central Argentina with different land use: a grassland area (ca. 4943 km²) mainly devoted to cattle grazing and an agro-ecosystem area (ca. 4006 km²) mostly used for crop production. The models were developed using logistic regression and were based on current records of Greater rhea occurrence coupled with remote sensing data, including land cover and human presence variables. The habitat suitability maps generated were used to predict the suitable habitat patch structure for wild rhea populations in each area. Fifty-one percent of the total grassland area was suitable for the species, being represented by a single large patch that included 62% of the individual locations. In the agro-ecosystem, only 28% of the total area was suitable, which was distributed among four patches. Seventy percent of rhea observations were in suitable habitat, with all rheas grouped in the largest patch. Conservation efforts for preserving wild rhea populations should be focused on maintaining habitats similar to grasslands, which are less profitable for landowners at present. Consequently, the protection of the pampas grasslands, a key habitat for this species as well as for others with similar habitat requirements, will demand strong conservation actions through the reconciliation of interests between producers and conservationists, since the proportion of croplands is increasing.     

Plant communities and soil phytolith assemblages relationship in native grasslands from southeastern Buenos Aires province,Argentina

This study has compared the phytolith production and soil phytolith assemblages of two native grass communities typical in Argentina, namely the monospecific Paspalum quadrifarium community or “pajonal”, and the Stipa-Piptochaetium community or “flechillar”. Five sites – three of which had been unmanaged while other two had been submitted to fire and fire-grazing management activities – were selected for study. The results show that in these sites a) phytolith production of the plant communities differed due to their diverse floristic composition, b) it was possible to define the type of management or the type of community based on soil phytolith assemblages, c) there were differences between plant communities and soil phytolith assemblages. These results suggest that modern phytolith assemblages are composed not only of local and present vegetation but also of extralocal/regional and probably past vegetation. Hypotheses about the probable processes involved in phytolith incorporation to soil have emerged during the investigation.     

Habitat associations of the endangered damselfly, Coenagrion mercuriale, in a water meadow ditch system in southern England

The damselfly Coenagrion mercuriale (Charpentier) (Odonata: Coenagrionidae) is threatened throughout Europe. It is listed on the EC Habitats and Species Directive and is the only odonate currently given priority status in the UK Biodiversity Action Plan. Information about the habitat preferences of this species is required to guide conservation and management programmes. We obtained estimates of density of mature adult C. mercuriale during an intensive mark-release-recapture study over 7.65 km of a water meadow ditch network in the Itchen Valley, Hampshire. Detailed habitat information was also collected, including a variety of physical variables, and data about the in-channel and bankside vegetation. C. mercuriale density and movement were analysed in relation to habitat variables and local population size using Generalized Linear Models. Mean adjacent population density was the single most important factor determining density. However the species was also associated with a number of habitat features, the most important of which were: a channel substrate consisting primarily of silt, wide underwater ledges (berms), in-channel emergent dicots, and bankside monocots. The presence of trees was negatively associated with damselfly density. Mean net lifetime movement was greatest from sections with low density, with smaller than average berms, and with deeper water. The causes and consequences of these findings are discussed in relation to the conservation and management of this rare species.     

Habitat use and preliminary demographic evaluation of the critically endangered Mediterranean monk seal (Monachus monachus) in the Cilician Basin (Eastern Mediterranean)

Over the last few decades a dramatic decline in the number of Mediterranean monk seals (Monachus monachus) limited scientific studies on the biology, behaviour, and ecology of the species. This lack of information impaired the effectiveness of conservation strategies. Thus, any further information gathered on the monk seal has utmost value in the work to halt and reverse the plight of the species. A hitherto unknown Cilician Basin colony of the Mediterranean monk seal has been investigated during a 6-year period between 1995 and 2001. Results of direct in-cave surveys and land based seal-watch observations enabled the evaluation of habitat use and preliminary demographic information. A total of 39 caves were discovered among which only three were used for breeding. Scarcity of the breeding caves emphasised the importance of breeding habitat in the survival of the colony. Common features of the breeding caves are found as an entrance with a protective barrier against strong waves; a chamber with a wide beach; and a well-sheltered shallow pool. Supplementary data obtained from infrared monitors installed in three actively used caves were used to assess the number of individuals. Throughout the study period, 25 individuals were identified and 11 newborn pups were found. Evaluation of the results suggests the presence of sub-regions inhabited by an individually identified sub-group of monk seals. Recommendations for conservation guided by this information is a Marine Protected Area consisting of two zones; a restricted core zone prohibiting any human activity securing breeding habitat and a restricted fishery zone securing sustainability of the food source.     

Habitat use and spatial dynamics of takhi introduced to Hustai National Park, Mongolia

The successful introduction of captive bred takhi or Przewalski’s horse, Equus ferus przewalskii, into Mongolia in the 1990s is a good example of the benefits of ex situ conservation and one of the few examples of the recovery of an animal after it became extinct in the wild. This is also particularly interesting because virtually nothing was known about how takhi lived before they died out, and the introductions have enabled us to study how they have settled, and their ecology and behaviour within their former natural range. In this paper, we describe the movement, home range size and shape, and habitat use of takhi at one of the release areas, the 570 km² Hustai National Park in Mongolia. Harem home ranges varied between 129 and 2399 ha, with 80% core areas of between 61 and 1196 ha. There was no relationship between range size and harem size, or length of time since release. Initially, harems stayed near their release enclosures, but over time they established home ranges further away. There was little overlap between home ranges of different harems, but neither was there evidence of exclusive range use. The more nutritious vegetation at lower elevations was preferentially selected. Thus the present situation looks good, but, as the population continues to grow, we anticipate that there will be potential problems related to intraspecific competition for water and vegetation resources, and the potential for hybridisation with domestic horses belonging to the local people. We consider the time it may take for takhi to reach carrying capacity within Hustai National Park and emphasise that continual monitoring of the population is essential because interventional management is likely to be required in the future.     

Habitat of endangered white abalone, Haliotis sorenseni

Surveys with a submersible at offshore islands and banks in southern California found that white abalone were most abundant at depths between 43 and 60 m. This is deeper than estimates taken when white abalone were more abundant. Densities were highest at sites far from fishing ports. Controlling for depth and site found that white abalone were significantly more abundant in areas with Laminaria farlowii (an alga) but abalone were not associated with areas high in the cover of other algae (Pelagophycus porra or Eisenia arborea) or the amount of sand in the habitat (except that abalone always occurred on rock). Within an area with abalone, the particular rock they occurred on was significantly larger than unoccupied neighboring rocks. Occupied rocks were not significantly different in algal cover or in sea urchin density than unoccupied neighboring rocks. The position of abalone on a rock was nearer to the rock-sand interface than would be expected based on a random distribution. More white abalone were feeding when in association with red urchins, perhaps because both grazers capture drift algae to eat. These data may aid future efforts to locate white abalone brood stock and identify locations for outplanting.     

Habitat use and conservation of bumblebees (Bombus spp.) under different grassland management regimes

Declines in the natural populations of several bumblebee species across Britain and Europe are an increasing cause for concern. In this study the habitat use of bumblebees was investigated on Salisbury Plain Training Area, the largest remaining area of unimproved chalk grassland in north-west Europe. Habitat characteristics influencing the overall abundance, species richness and foraging activity of bumblebees included the diversity and abundance of flowering plant species (particularly of favoured forage plants such as Trifolium pratense), vegetation structure and height. It is suggested that different Bombus species respond to these habitat characteristics depending on their specific foraging and nesting requirements, the case of Bombus humilis being especially relevant. The effects of several grassland management practices were considered in terms of their suitability for the conservation of bumblebee habitats. Cattle grazing was shown to be preferable to both sheep grazing and the absence of any management, although the timing and intensity of such grazing was important. Small-scale disturbances caused by vehicle activity were also of value in producing locally abundant forage resources in less intensively managed grasslands.     

Habitat associations of bats in Northern Ireland: implications for conservation

Agricultural intensification in Northern Ireland has brought about large-scale changes to the landscape with a detrimental effect on biodiversity. Between 1996 and 1998, we surveyed a stratified random sample of 1 km squares for bats using a spot-sample technique and time expansion bat detector to establish linear and area habitat associations. Bats strongly selected water bodies with bankside vegetation, treelines, and deciduous and mixed woodland edge, avoiding open areas such as upland/unimproved grassland and improved grassland. Of three sympatric pipistrelle species (Pipistrellus pipistrellus, P. pygmaeus and P. nathusii), two were shown to forage in different habitats; P. pipistrellus being more of a ‘generalist’, foraging in a wider variety of habitats than P. pygmaeus, a ‘specialist’ which foraged in a smaller range of habitats. Nyctalus leisleri selected parkland/amenity grassland, deciduous woodland edge and rivers/canals and avoided improved grassland, hedgerows, whilst Myotis spp. selected deciduous woodland centre, lake/reservoir margins and rivers/canals. Reduction in area and quality of inland water, deciduous woodlands and field boundaries in Northern Ireland may impact on bat populations. Habitat management should focus on improving those habitats which are selected by bats, including the maintenance and enhancement of connecting linear habitats.     

Habitat selection by the pale-headed brush-finch (Atlapetes pallidiceps) in southern Ecuador: implications for conservation

The pale-headed brush-finch (Atlapetes pallidiceps) is threatened with extinction due to habitat loss, but very little is known about its ecological requirements. We used multiple logistic regression to study habitat selection of this species at landscape, territory, and nest site scales in order to make recommendations about effective management. Habitat selection by the sympatric stripe-headed brush-finch (Buarremon torquatus) was examined with the same methods in order to analyse interspecific resource partitioning and potential competition. The pale-headed brush-finch selected semi-open habitat types with intermediate scrub heights, and avoided forests. Nest sites depended on the presence of vines or bamboo. By contrast, the stripe-headed brush-finch chose dense habitat with low ground cover under tall vegetation and avoided semi-open habitat. The two species had overlapping territories but differed significantly in microhabitat use and the use of vegetation strata. We found no convincing evidence that the stripe-headed brush-finch displaces the pale-headed brush-finch from optimal habitat. The preservation of semi-open scrubland maintained by low-intensity grazing is suggested for future conservation of the pale-headed brush-finch.     

Status and conservation of the grey seal, Halichœrus grypus, in France

We investigated the status of the grey seal in the Molène archipelago, western Brittany, where a Marine National Park is planned. We combined regular censuses, photo-identification and satellite telemetry to investigate trends in grey seal abundance, site fidelity, individual movements and haulout site use. Individual movements and seasonal variations in abundance suggest that the seals used the Molène archipelago alternatively with other haulout sites of the South-west British Isles during their annual cycle. Nine out of 16 seals fitted with Satellite Relay Data Loggers (SRDLs) moved to the Isles of Scilly, English Cornwall, Wales or the Channel islands. On average the seals spent 85% of their time Near Haulout, 56.8% of which in the Molène archipelago. Overall the seals spent 66% of their tracking time within the boundaries of the Marine National Park. From 1997 to 2000 in the Molène archipelago, we observed a peak in number during the moulting season (mean = 49.8 seals) and a lower abundance during the breeding season (mean = 38.3 seals). The sex ratio was strongly skewed towards males during the moult (between 3.1 and 5.3-1), and was close to 1:1 during the summer. Inter-annual site fidelity outside the breeding season was very high, varying from 70% to 95% of identified seals, suggesting that the observed movements constitute rotations rather than random dispersion. From 1991/1994 to 1997/2000, the mean number of seals counted at haulout sites increased by 7%, close to the rates of increase observed in the core population of the British Isles at the same period. The Marine National Park may be too restricted for the overall conservation of the population but management plans should be aimed at preserving the characteristics of the habitat for the grey seal.     

Habitat selection and conservation of Sharpe's longclaw (Macronyx sharpei), a threatened Kenyan grassland endemic

Montane grassland is a severely threatened habitat in Kenya. Despite a high level of faunal endemism, it has received very little conservation attention. We investigated habitat selection in a threatened grassland endemic bird species, Sharpe's longclaw (Macronyx sharpei) (family Motacillidae), in order to understand its likely response to land-use changes. Between November 1995 and May 1996, we studied 41 territories of this species on the Kinangop Plateau in central Kenya. With an overall density of 0.4 birds ha⁻¹, longclaws lived in permanent groups of two to seven individuals. They were sedentary and territorial, with a mean home range size of 0.5 ha. The species avoided non-grassland areas entirely, and within grassland showed a strong preference for short grass with tussocks. Territory sizes and foraging ranges were smaller, and rates of pecking for food higher, in this grassland type than in open short grass or long grass. Land use changes, in particular conversion to cultivation or woodlots and ploughing up of grassland to remove tussocks, pose a serious and immediate threat to this species. Dairy farming is potentially compatible with grassland conservation, and conservation-friendly livestock rearing needs to be promoted through economic incentives, awareness-raising and technical advice.     

Habitat selection of endangered and endemic large flying-foxes in Subic Bay, Philippines

Large flying-foxes in insular Southeast Asia are the most threatened of the Old World fruit bats due to high levels of deforestation and hunting and effectively little local conservation commitment. The forest at Subic Bay, Philippines, supports a rare, large colony of vulnerable Philippine giant fruit bats (Pteropus vampyrus lanensis) and endangered and endemic golden-crowned flying-foxes (Acerodon jubatus). These large flying-foxes are optimal for conservation focus, because in addition to being keystone, flagship, and umbrella species, the bats are important to Subic Bay’s economy and its indigenous cultures. Habitat selection information streamlines management’s efforts to protect and conserve these popular but threatened animals. We used radio telemetry to describe the bats’ nighttime use of habitat on two ecological scales: vegetation and microhabitat. The fruit bats used the entire 14,000 ha study area, including all of Subic Bay Watershed Reserve, as well as neighboring forests just outside the protected area boundaries. Their recorded foraging locations ranged between 0.4 and 12 km from the roost. We compared the bats’ use to the availability of vegetative habitat types, riparian areas, and bat trees. The fruit bats’ locations showed a preference for undisturbed forest types and selection against disturbed and agricultural areas. Bat locations also showed selection for particular fruiting/flowering bat trees. The bats showed strong preference for riparian areas; locations were in riparian areas over four times more than expected. From these results we recommend that management focus flying-fox conservation efforts on undisturbed forest and riparian areas.     

Habitat selection of the Eurasian woodcock in winter in relation to earthworms availability

The Eurasian woodcock (Scolopax rusticola) is a game species experiencing high hunting pressure, long-term modifications of its habitats, and with questions regarding its current conservation status. Winter is a season of highest concentration of birds and hunting pressure but woodcock precise habitat requirements are poorly known. It is crucial to assess threats and to develop sustainable management options for the conservation of woodcock populations. During three consecutive winters, we monitored 65 individual woodcocks fitted with radio-tags in Brittany, France. Habitat selection was analysed using GIS and compositional analysis, in relation to vegetation types, soil variables (humus types) and the abundance of their main prey (earthworms). Woodcocks used different habitats diurnally and nocturnally, generally preferring areas with high earthworm biomass. Diurnal habitat selection in forests was associated with humus type (preference for mulls, rich in earthworms) and dense shrub strata (better protection). Hedges with a high density of trees and shrub were also important habitat. At night, grazed meadows were the preferred habitat, containing five times higher biomass of earthworms compared to cultivated fields. Sustainable management of populations requires protection and management of habitats that incorporates food and cover. Forestry practices should preserve rich humus types and coppices by choosing tree species that ameliorate the soil and soil tilling. Changes in landscapes and intensive agricultural practices are current threats to woodcock populations: destruction of hedges, decrease of permanent grazed meadows, impoverishment of soils fauna biomasses from ploughing and chemical applications. However, woodcocks may benefit from the recent development of set-asides, grass field-borders and simplified farm practices (no-tillage and direct sowing).     

Assessing habitat use of the endangered marine mollusc Patella ferruginea (Gastropoda, Patellidae) in northern Africa: preliminary results and implications for conservation

The limpet Patella ferruginea Gmelin, 1791, endemic to the Mediterranean, is the most endangered marine species on the list of the European Council Directive 92/43/EEC on the Conservation of Natural Habitat of Wild Fauna and Flora (1992). Its Mediterranean range has progressively contracted to a few restricted areas and the species is now threatened with extinction. Seventeen stations were sampled along the littoral zone on the coast of Ceuta, North Africa, for quantifying environmental factors (water movement, siltation and suspended solids), and 70 transects were selected for P. ferruginea sampling. The study revealed the presence of a well-established population of P. ferruginea with values of density and size of 0.67 individuals m⁻¹ ±0.96 and 48.94 mm±11.61 respectively (mean±standard error of mean). The biggest densities of P. ferruginea were found inside the harbour of Ceuta. The limpets were more abundant on artificial harbour stones than on natural rocky shores, and the areas under the highest human pressure were characterised by the lowest densities and the smallest specimens probably due mainly to the predation for food and fishing. Human pressure is probably the main contributing factor to the currently endangered status of P. ferruginea. Programmes of environmental education to avoid its collection for fishing, food or for fun as decorative objects should be conducted, and further experimental studies dealing with the reproductive biology of this species are needed to properly assess the future programmes of conservation.     

Habitat use, predicted distribution and conservation of green peafowl (Pavo muticus) in Dak Lak Province, Vietnam

In the last few decades, green peafowl numbers have declined drastically across their south-east Asian range, primarily, it is believed, due to human persecution and loss of habitat. They were formerly common across Vietnam, but are now restricted to a few central provinces. In this study green peafowl were surveyed in Dak Lak province, Vietnam, using a point-count methodology to assess their status and distribution, and to investigate their habitat use. Results showed that the most used habitat was dry deciduous forest with access to permanent water and free from human disturbance. Green peafowl density was lower in mixed and evergreen forest and lower when water was distant and/or human settlement was nearby. A map of predicted distribution was created which predicted that eastern and north-eastern Dak Lak are important areas containing much of the remaining habitat and populations of green peafowl. Several key river systems within this area were predicted to be of particular value. Deciduous forest < 2 km from water and > 2 km from human settlement was estimated to occupy only 3% of the province but to hold 37% of green peafowl. Conservation of green peafowl in Dak Lak, and elsewhere, should concentrate on maintaining undisturbed access to permanent water within large tracts of dry deciduous forest. The spread of human settlement presents the greatest threat; both directly due to hunting pressure and habitat loss and indirectly by preventing access to water in otherwise suitable areas.     

Landscape occupancy and local population size depends on host plant distribution in the butterfly Cupido minimus

We tested whether the landscape occupancy and local population size of the monophagous butterfly Cupido minimus can be predicted by patch size and isolation of its host plant or by other habitat characteristics. C. minimus and its larval food plant Anthyllis vulneraria are classified as rare and endangered in northern Germany. Adults of C. minimus are ranked as the most sedentary butterfly species in northern Europe.Around the city of Göttingen (Germany), we checked all known locations of A. vulneraria (n=70) in June 2002 for butterfly eggs (in blooming flowerheads) and adult butterflies (within 20-min transects).We found eggs of C. minimus or a high number of adults (>7) in all habitats with A. vulneraria (which are calcareous grasslands) even when isolated up to 2-4 km. In multiple regression analyses, local population size of adult butterflies was positively related to the cover of its larval food plant A. vulneraria explaining 65% of variance. Cover of A. vulneraria increased with increasing habitat area and increasing cover of plant species in flower and decreased with increasing cover of shrub layer. Habitat isolation and further factors describing habitat quality were not related to C. minimus population size or cover of its larval food plant.The results suggest that dispersal ability of C. minimus is greater than expected and that management should focus to increase A. vulneraria patches. For conservation, low impact grazing once a year and removing of excessive shrubs in winter seems to be the most appropriate strategies.     

Bighorn sheep habitat use and selection near an urban environment

To further understand the impact of urban development on wildlife populations, we examined habitat use and selection by female bighorn sheep Ovis canadensis in two endangered subpopulations near a metropolitan area in southern California. One subpopulation, which had previously been found to have low reproductive success, selected urban environments while the other did not use urban areas. In the subpopulation that used urban areas, females had smaller core activity areas and selected lower elevations and gentler slopes. These females used urban sources of water but a clear relationship between levels of urban use and periods of increased water need was not evident. Diet quality was higher among females that selected urban areas, and this increase was correlated with the level of urban use. Thus, optimal foraging behavior may have contributed to the selection of urban areas. Urban use was lowest during peak months of parturition, suggesting that reproductive strategies may also have influenced temporal patterns of urban use. Although urban areas provided bighorn sheep with forage and water resources, the use of these areas may have substantial costs. For example, females using urban areas had internal parasites that were not found elsewhere in the Peninsular Ranges.     

Habitat, human pressure, and social behavior: Partialling out factors affecting large-scale territory extinction in an endangered vulture

Extinctions are often the result of multiple factors that are difficult to disentangle and so methods for identifying simple and combined types of extinctions are valuable for both basic and applied ecology. We applied a modification of variance partitioning and a hierarchical partitioning analysis to test several hypotheses that attempt to explain the recent large-scale disappearance of Egyptian Vulture breeding territories in Spain. Our aim was to identify and then separate the simple (or pure) from the combined effects of habitat features, human pressure, and the social behavior of the species on the risk of extinction from a territory while controlling for spatial autocorrelation. Deviance partitioning showed that a complex mix of factors is significantly related to the disappearance of more than 400 territories throughout Spain. Abandoned territories were located in areas that are isolated from other conspecific territories and far from communal roost sites. In addition, these territories were found in places where there is a lack of natural habitats, high habitat fragmentation and reduced habitat diversity, and where food availability seems to be low and the illegal use of poison to control predators is a common practice. Deviance partitioning also showed an important spatial component in the probability of extinction. Abandoned territories were not randomly distributed; rather, they were aggregated in extinction ‘hotspots’, mainly related to food availability and human pressure. Deviance partitioning turned out to be an useful tool for identifying the relative contribution of a variety of factors - and their combined effects - associated with an extinction process. The deviance explained by each factor must be interpreted, however, in the context of a good knowledge of the life history of the species. Hierarchical partitioning can help rank conservation priorities and, by using as an objective criterion the relative weight of each independent variable that could be effectively managed for conservation, may provide wildlife managers with a means of saving funds and optimizing action plans.     

The habitat requirements of the rufous treecreeper (Climacteris rufa). 1. Preferential habitat use demonstrated at multiple spatial scales

Determining the habitat requirements of a species is fundamental to effective conservation, particularly if the species is declining in areas where its habitat is being modified. Multi-scaled investigations of habitat use are essential because different selection processes may operate at different scales. I examined the habitat use of a declining woodland passerine, the rufous treecreeper (Climacteris rufa), at three spatial scales (landscape, woodland and territory) in the wheatbelt of Western Australia. Preferential habitat use was exhibited at all scales. At the landscape scale, wandoo (Eucalyptus wandoo) woodland was used at a significantly greater rate than three other common vegetation types. Territory use within woodlands was positively related to the density of hollow-bearing logs, the density of nest sites, and tree age. Within an individual territory, nest sites (hollows) were favoured if they had a spout angle of ?50° to the horizontal and an entrance size of between 5 and 10 cm. The rufous treecreeper preferentially used habitat with traits characteristic of old-growth wandoo woodland. Degradation of wandoo through habitat modification (e.g. grazing, logging, fire and removal of deadwood) represents a significant threat to the persistence of treecreepers.     

Quantification of Wautersia [Ralstonia] basilensis in the mycorrhizosphere of Pinus thunbergii Parl. and its effect on mycorrhizal formation

The bacterium Wautersia [Ralstonia] basilensis has been shown to enhance the mycorrhizal symbiosis between Suillus granulatus and Pinus thunbergii (Japanese black pine). However, no information is available about this bacterium under field conditions. The objectives of this study were to detect W. basilensis in bulk and mycorhizosphere soils in a Japanese pine plantation in the Tottori Sand Dunes, determine the density of W. basilensis in soil, and determine the optimal cell density of W. basilensis for mycorrhizal formation in pine seedlings. We designed and validated 16S rRNA gene-targeted specific primers for detection and quantification of W. basilensis. SYBR Green I real-time PCR assay was used. A standard curve relating cultured W. basilensis cell density (10³-10⁸ cells ml⁻¹) to amplification of DNA showed a strong linear relationship (R = 0.9968). The specificity of the reaction was confirmed by analyzing DNA melting curves and sequencing of the amplicon. The average cell density of W. basilensis was >4.8 × 10⁷ cells g⁻¹ of soil in the mycorrhizosphere and 7.0 × 10⁶ cells g⁻¹ in the bulk soil. We evaluated the W. basilensis cell density required for mycorrhizal formation using an in vitro microcosm with various inoculum densities ranging from 10² to 10⁷ cells g⁻¹ soil (10⁴-10⁹ cells ml⁻¹). Cell densities of W. basilensis of >10⁶ cells g⁻¹ of soil were required to stimulate mycorrhizal formation. In vivo and in vitro experiments showed that W. basilensis was sufficiently abundant to enhance mycorrhizal formation in the mycorrhizosphere of Japanese black pine sampled from the Tottori Sand Dunes.