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

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

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.     

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.     

Effects of landscape-scale broadleaved woodland configuration and extent on roost location for six bat species across the UK

Although forest fragmentation can greatly affect biodiversity, responses to landscape-scale measures of woodland configuration in Europe have been examined for only a limited range of taxa. Almost all European bat species utilise woodland, however little is known about how they are affected by the spatial arrangement of woodland patches. Here we quantify landscape structure surrounding 1129 roosts of six bat species and a corresponding number of control locations across the UK, to examine associations between roost location and landscape composition, woodland proximity and the size of the nearest broadleaved woodland patch. Analyses are performed at two spatial scales: within 1 km of the roost and within a radius equivalent to the colony home-range (3–7 km). For four species, models at the 1 km scale were better able to predict roost occurrence than those at the home-range scale, although this difference was only significant for Pipistrellus pipistrellus. For all species roost location was positively associated with either the extent or proximity of broadleaved woodland, with the greatest effect of increasing woodland extent seen between 0% and 20% woodland cover. P. pipistrellus, Pipistrellus pygmaeus, Rhinolophus hipposideros, Eptesicus serotinus and Myotis nattereri all selected roosts closer to broadleaved woodland than expected by chance, with 90% of roosts located within 440 m of broadleaved woodland. Roost location was not affected by the size of the nearest broadleaved patch (patches ranged from 0.06–2798 ha ± 126 SD). These findings suggest that the bat species assessed here will benefit from the creation of an extensive network of woodland patches, including small patches, in landscapes with little existing woodland cover.     

Modelling the influence of seasonal inter-habitat movements by an ecotone rodent

I assessed the long-term effects of seasonal inter-habitat movements for a common rodent, Microtus californicus, using matrix models tailored to demographic data obtained on populations living at the marsh-grassland ecotone. Three stochastic models were used to simulate the seasonal and spatial dynamics of this system, with the purpose of determining: (1) the growth rates of populations living in single versus a two-habitat landscape; and (2) if the amount, direction, and seasonality of rodent movements between the two habitats may impact these growth rates. The results indicated that grassland populations could be self-sustaining, but that isolated marsh populations would rapidly decline. Importantly, 98% of the two-habitat simulations yielded growth rates greater than those seen in the grassland model, with populations benefiting the most by movements from marsh into grassland at the end of the dry season. This study demonstrated that movements between marsh ‘sink’ and grassland ‘source’ habitats can potentially increase long-term population growth due to seasonal, inter-habitat differences in resource availability.     

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.     

Improving the biodiversity benefits of hedgerows: How physical characteristics and the proximity of foraging habitat affect the use of linear features by bats

Within agricultural landscapes, linear features such as hedgerows and tree-lines provide valuable habitat for many species. We use data from 315 transects, completed as part of a national acoustic survey of bat distribution, to examine the incidence of four bat species adjacent to linear features in rural areas. The use of linear features was assessed in relation to hedgerow width, tree density, the presence of water and woodland proximity. To examine the effect of tree density, linear features were classified as either hedgerows without trees, hedgerows with sparse trees (comprising <50% tree canopy) or tree-lines (>50% tree canopy). The use of linear features by Pipistrellus pipstrellus was not affected by tree density; linear features of all types were associated with a similar increase in P. pipistrellus incidence. The use of linear features by Pipistrellus pygmaeus was dependent on both tree density and the proximity of woodland; only linear features containing trees provided an increase in P. pygmaeus incidence regardless of woodland proximity. P. pipistrellus and P. pygmaeus incidence was not affected by hedgerow width or the presence of water. Incidence of Nyctalus noctula and Eptesicus serotinus was unaffected by the density of linear features of any type. Many agri-environment schemes offer financial incentives for the creation and management of hedgerows. Optimising the biodiversity gain provided by linear features will maximise the effectiveness of these schemes. Agri-environment measures that encourage the provision and retention of hedgerow trees will benefit bats in agricultural landscapes.     

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.     

Multiple spatial scale logistic and autologistic habitat selection models for northern pygmy owls, along the eastern slopes of Alberta’s Rocky Mountains

Little is known about the distribution and habitat use of northern pygmy owls (Glaucidium gnoma), in Alberta or throughout their range. In Alberta they are ranked as ‘sensitive’, meaning they are not believed to be at immediate risk of extirpation or extinction but may require special attention or protection to prevent them from becoming at risk.Diurnal broadcast surveys were conducted to determine distribution and habitat selection throughout a 28,500 km² study area situated along the eastern slopes of the Alberta Rocky Mountains. Surveys lasted for eight weeks in 2001, during which time 1532 site visits were made. Forty-eight responses were recorded at 42 sites representing 40 individual northern pygmy owls. Predictive models of habitat selection were developed using stepwise logistic and autologistic regression. Autologistic models accounted for observed spatial dependencies and as a result, produced better fitting models that more accurately reflect the role of predictor variables in influencing species occurrence. All models considered biophysical variable selection at two spatial scales, the minimum (75 ha) and maximum (300 ha) home range size. Northern pygmy owls showed a preference for older, structurally diverse mixedwood habitats, with line-of-sight enhanced by increased edge and terrain roughness.The use of habitat selection models resulting from this project and Geographic Information Systems as a tool, will enable managers to identify key habitat features, focus future survey efforts, set habitat goals and evaluate the effects of management decisions on current and future habitat availability.     

Location of roosts of the lesser long-eared bat Nyctophilus geoffroyi and Gould's wattled bat Chalinolobus gouldii in a fragmented landscape in south-eastern Australia

The location of roosts of two species of vespertilionid bats, the lesser long-eared bat (Nyctophilus geoffroyi) and Gould’s wattled bat (Chalinolobus gouldii), was investigated in a remnant vegetation-farmland mosaic and adjacent floodplain forest in south-eastern Australia. A total of 45 individuals of N. geoffroyi and 27 C. gouldii were fitted with radio transmitters, which resulted in the location of 139 and 89 roosts respectively. Selection of roosting habitat showed both inter- and intra-specific differences. At the landscape level, locations of roosts used by male and female N. geoffroyi differed markedly. Most males roosted in the farmland mosaic within 3 km (mean 1.9±2.9 km) of where they were captured while foraging in remnant vegetation. In contrast, roost sites of females were predominantly in the floodplain forest, 6-12 km from their capture site in farmland (mean for all females, 6.7±2.9 km). All maternity roosts were in the extensive floodplain forest, 4-10 km from foraging areas. Distances moved by this species were greater than previously recorded and further than predicted by flight morphology. Most C. gouldii roosted in the floodplain forest, 4-10 km from their capture site (6.9±1.6 km). Within the floodplain forest, both species roosted in areas that had higher densities of hollow-bearing trees than generally available; dead hollow-bearing trees for N. geoffroyi and large, live trees for C. gouldii. The scale of movements undertaken by these species means that they can obtain resources from multiple landscape elements. Both species used different habitats for roosting and foraging despite the energetic costs of commuting relatively large distances. Conservation of bat populations in rural environments needs to be considered at the landscape scale, with particular attention to identifying landscape elements that provide key resources.     

Changes in agricultural management drive the diversity of Burkholderia species isolated from soil on PCAT medium

In order to assess the diversity of culturable Burkholderia populations in rhizosphere and bulk soil and to evaluate how different agricultural management regimes and land use history affect this diversity, four treatments were evaluated: permanent grassland; grassland converted into maize monoculture; arable land and arable land converted into grassland. Burkholderia isolates obtained on PCAT medium were grouped in 47 clusters using 16S ribosomal RNA gene based PCR-DGGE combined with BOX genomic fingerprinting (DGGE-BOX). The distribution of the isolates in the DGGE-BOX clusters was used to calculate the Shannon diversity index per treatment. Interestingly, we observed that the Burkholderia diversity was affected by changes in the agricultural management, since the highest diversity was observed in permanent grassland and in continuous arable land. In addition, the diversity tended to be higher in the rhizosphere than in the corresponding bulk soil. The use of species abundance models indicated that rhizosphere communities had more even distributions than communities collected from the bulk soil. Identification of isolates revealed that only 2% of these belonged to the B. cepacia complex and that the majority was assigned to either (1) new Burkholderia species or (2) Burkholderia species that had originally been isolated from soil. Isolates classified as B. hospita, B. caledonica and Burkholderia sp. ‘LMG 22934’ and ‘LMG 22936’ were found mainly in the arable land, while isolates belonging to Burkholderia sp. ‘LMG 22929’ and B. phytofirmans were associated with the grassland area. Another potentially new Burkholderia species, ‘LMG 22932’, was found in both areas, in close association with the maize rhizosphere.     

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.     

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.     

Microbial communities of a native perennial bunchgrass do not respond consistently across a gradient of land-use intensification

To test if native perennial bunchgrasses cultivate the same microbial community composition across a gradient in land-use intensification, soils were sampled in fall, winter and spring in areas under bunchgrasses (‘plant’) and in bare soils (‘removal’) in which plots were cleared of living plants adjacent to native perennial bunchgrasses (Nassella pulchra). The gradient in land-use intensification was represented by a relict perennial grassland, a restored perennial grassland, and a perennial grass agriculture site on the same soil type. An exotic annual grassland site was also included because perennial bunchgrasses often exist within a matrix of annual grasses in California. Differences in soil resource pools between ‘plant’ and ‘removal’ soils were observed mainly in the relict perennial grassland and perennial grass agriculture site. Seasonal responses occurred in all sites. Microbial biomass carbon (C) and dissolved organic C were greater under perennial bunchgrasses in the relict perennial grassland and perennial grass agriculture site when comparing treatment means of ‘plant’ vs. ‘removal’ soil. In general, soil moisture, microbial respiration, and nitrate decreased from fall to spring in ‘plant’ and ‘removal’ soils, while soil ammonium and net mineralizable nitrogen (N) increased only in ‘plant’ soils. A canonical correspondence analysis (CCA) of phospholipid fatty acid (PLFA) profiles from all sites showed that land-use history limits the similarity of microbial community composition as do soil C and N dynamics among sites. When PLFA profiles from individual sites were analyzed by CCA, different microbial PLFA markers were associated with N. pulchra in each site, indicating that the same plant species does not retain a unique microbial fingerprint across the gradient of land-use intensification.     

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

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

Gap-crossing by gliding marsupials: thresholds for use of isolated woodland patches in an agricultural landscape

Single trees and small patches of trees in farmland are conspicuous components of agricultural landscapes around the world, but their value for the conservation of biodiversity is not well known. In this study, arboreal mammals were censused by using hair-sampling tubes in small patches of woodland (single trees to patches <1.0 ha) in cleared farmland adjacent to a linear network of woodland known to support resident populations of arboreal mammals. Ninety-one small isolates were stratified by size (single trees or small patches) and distance from the linear network to test the capacity of animals to cross habitat ‘gaps’. The genus Petaurus (small gliding marsupials), the most commonly detected taxon, was recorded in 31% of hair-tubes (98 of 316). It occurred in 21% of sites in isolated trees and patches, and in all linear strips. A logistic regression model demonstrated that Petaurus sp. was most likely to occur in isolates in close proximity to linear strips and other patches of woodland. Ninety-five per cent of sites at which this taxon occurred were within 75 m of the linear network. This threshold corresponds with the maximum distance that animals can glide in a single movement between trees. The size of isolates did not influence utilisation rates. Such isolates are smaller than a single home range and were probably used to supplement home ranges centred on the linear network, by providing additional foraging habitat and den sites. Protection and restoration of isolated trees and small woodland clumps in cleared landscapes contributes to mammal conservation and this study provides quantitative data that can assist landscape design and habitat restoration in rural environments.     

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.     

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.     

Effects of Medicago sativa, Taraxacum officinale and Bromus sterilis on the density and diversity of Collembola in grassy arable fallows of different ages

The density and diversity of Collembola of nine grassy arable fallows of different ages were investigated in a factorial design with the factors ‘plant species’ (legume: Medicago sativa, herb: Taraxacum officinale, grass: Bromus sterilis) and ‘age class’ (2-3, 6-8 and 12-15 years) including the random effect ‘site’ (1-9). In May 2008, four plots were selected randomly at each fallow. Within each plot five M. sativa, T. officinale and B. sterilis plants were extracted with their associated soil using steel cylinders. The material from each plant species was used for extraction of Collembola and for determination of environmental parameters. Thus, the new aspect of the present study compared to other field studies investigating the relationships between plant and Collembolan communities is the focus on the “micro-scale”, investigating the Collembolan communities of the soil associated with single plants.We found that species richness and density of total and euedaphic Collembola were significantly higher in B. sterilis than in T. officinale samples with the M. sativa samples being intermediate. Fine-root feeding euedaphic Collembola particularly benefited from the higher amount of fine roots in B. sterilis samples. We also discovered that the age of the fallows had no significant influence on the number of Collembolan species and the density of the Collembolan groups. Notably, however, species of the epedaphic genera Lepidocyrtus and Sminthurinus were associated with the 12-15 year-old fallows and presumably benefited from the high number of plant species in the old fallows. Finally, canonical correspondence analysis (CCA) indicated that the soil organic matter content and the microbial biomass, both potential food sources for many Collembolan species, were important structuring forces for the Collembolan communities.