Roosting Behavior of Colonial and Solitary Flying Foxes in American Samoa (Chiroptera: Pteropodidae)1

We examined characteristics of roosting sites utilized by two flying fox species (Pteropus tonganus and P. samoensis) in American Samoa. The colonial roosting sites of P. tonganus were observed over a ten‐year period, including two years when severe hurricanes devastated bat populations and destroyed roost trees. Prior to the hurricanes, roosts were located on cliff faces above the ocean or steep mountainsides, locations that were either inaccessible to people or in protected areas where hunting was not allowed. In the years immediately following the hurricanes, P. tonganus colonies split into smaller groups that moved frequently to different locations. Four years after the second hurricane, colonies had coalesced and returned to many of the traditional roosting sites used before the hurricanes. Common tree species in upland and coastal forest were selected as roosts. The isolated locations selected for P. tonganus roosts were apparently the result of hunting pressure on the colonies. The solitary roosts of P. samoensis were observed during 29 months. Roosting bats were well concealed and hard to detect within the forest; even bats on exposed branches were cryptic. Mature primary forest was favored as roosting habitat. Individual bats used specific branches or trees as roosts and returned to them for up to 29 months. Unlike P. tonganus, people did not alarm roosting P. samoensis easily and some roosts were located near houses and along roads.     

Indirect Evidence that Flying Foxes Track Food Resources Among Islands in a Pacific Archipelago

Although flying foxes (fruit bats in the genus Pteropus ) in continental forests often fly between scattered resources, little is known about their ranging behavior among islands. The inhospitable water matrix that surrounds the food patches (islands) in archipelagos may prevent flying foxes from tracking resources as efficiently as their counterparts on larger landmasses do. Our aim in this study was to determine whether the abundance of foraging flying foxes ( Pteropus tonganus ) reflected food availability on islands in the Vava'u archipelago of Tonga, regardless of island size and isolation. Overall, food availability was the strongest determinant of flying fox abundance, and spatial aspects of the islands (land area within 10 km) had only a small influence. Food availability appears to regulate flying fox abundance only when food resources are low, but when food sources are plentiful, flying fox abundance may be high or low. These results provide indirect evidence that flying foxes are able to track food resources efficiently in an archipelago, and the water matrix that surrounds the food patches (islands) is not a strong deterrent for foraging animals.     

Use of Forest Edges by Bats in a Managed Pine Forest Landscape

ABSTRACT Forest edges often have increased species richness and abundance (edge effect) and affect spatial behaviors of species and dynamics of species interactions. Landscapes of intensively managed pine (Pinus spp.) stands are characterized by a mosaic of patches and linear forest edges. Managed pine forests are a primary landscape feature of the southeastern United States, but the effects of intensive management on bat communities are poorly understood. Insectivorous bats are important top predators in nocturnal forest food webs. We examined bat foraging behavior along forest edges and in 4 structurally distinct stand types (open-canopy pine, prethinned pine, thinned pine, and unmanaged forest) within a managed pine forest in the coastal plain of North Carolina, USA. During May-August, 2006 and 2007, we recorded echolocation calls using Pettersson D240X bat detectors linked to digital recorders at 156 sites. We also sampled nocturnal flying insects at each site using Malaise insect traps. We used negative binomial count regression models to describe bat foraging behavior relative to forest edges, stand types, and prey availability. Although some species showed affinities for certain stand types and prey items, bat activity patterns were most strongly related to forest edges. Edges were used extensively by 6 aerial-hunting bat species, but avoided by Myotis species. Forest edges function similarly to natural forest gaps, by providing foraging opportunities for aerial-hunting bat species. Therefore, the maintenance of forest edges in managed pine landscapes may enhance foraging habitat for aerial-hunting bat species.     

Magellanic woodpecker ( Campephilus magellanicus) abundance and foraging in Tierra del Fuego, Chile

The Magellanic woodpecker (Campephilus magellanicus) is a vulnerable and poorly studied bird in the sub-antarctic deciduous and evergreen beech (Nothofagus) forests of South America. On Tierra del Fuego island (Chile), we compared Magellanic woodpecker abundance and its foraging habitat in two forest types: pure N. pumilio and mixed forests composed by N. pumilio and N. betuloides, including managed and non managed stands. At a regional scale, abundance of woodpeckers was greater in landscapes including both forest types than in pure N. pumilio landscapes. When both forest types occurred together, woodpecker abundance did not differ between them. The number of trees with foraging signs was correlated with Magellanic woodpecker abundance and was also associated with N. betuloides and snag densities, but was not affected by forest management. Occurrence of pecking on foraging trees was greater in mixed Nothofagus than pure N. pumilio stands. Woodpeckers foraged disproportionately more on larger diameter and more decayed trees. Moreover, trees used for foraging were positively correlated with canopy cover and snag density and were negatively correlated with distance to nearby peatlands and beaver ponds. Direct observation revealed that the flying distance between trees was negatively correlated with proportion of trees with foraging signs. Woodpeckers chose trees that were visited before, suggesting a pattern of tree recognition within foraging territories.Communicated by F. Bairlein     

Preference of an insular flying fox for seed figs enhances seed dispersal of a dioecious species

Interactions among multiple species form complex networks of interdependences and are considered primary factors in the generation and maintenance of biodiversity. Pteropodid bats are keystone species that provide important ecosystem services of pollination and seed dispersal in the tropics and subtropics. In this study, we investigated the utilization and preference of food resources by the insular frugivorous flying fox Pteropus dasymallus. We found that fig species constituted the major portion of the diet of the flying fox (94.6%). When foraging, the flying fox preferred seed figs from female trees over gall figs from male trees in functionally dioecious fig species. Germination experiments showed a significantly higher percentage of germination for fig seeds in feces than those from pellets and ripe figs (feces: 80.2%, pellets: 23.4%, ripe figs: 32.9%). Considering the active selection of seed figs and avoidance of gall figs by foraging flying foxes, we suggest that the abundance of seed figs accurately represents food availability for dioecy. This preference for seed figs or viable seeds can effectively promote the survival of pollinating wasps and might reinforce the evolution of dioecism in figs. In addition, the effects of gut passage on seed germination, in combination with the capacity of flying foxes to travel long distances, may substantially contribute to the efficiency of flying foxes as seed dispersers.     

Seasonal Shifts in Nocturnal Habitat Use by Coastal Bat Species

Sensitivity of bats to land use change depends on their foraging ecology, which varies among species based on ecomorphological traits. Additionally, because prey availability, vegetative clutter, and temperature change throughout the year, some species may display seasonal shifts in their nocturnal habitat use. In the Coastal Plain of South Carolina, USA, the northern long-eared bat (Myotis septentrionalis), southeastern myotis (Myotis austroriparius), tri-colored bat (Perimyotis subflavus), and northern yellow bat (Lasiurus intermedius) are species of conservation concern that are threatened by habitat loss. Our objective was to identify characteristics of habitat used by these species during their nightly active period and compare use between summer and winter. We conducted acoustic surveys at 125 sites during May–August and at 121 of the same 125 sites December–March 2018 and 2019 in upland forests, bottomland forests, fields, ponds, and salt marsh and used occupancy models to assess habitat use. The northern long-eared bat and southeastern myotis (i.e., myotis bats) used sites that were closer to hardwood stands, pine stands, and fresh water year-round. We did not identify any strong predictors of tri-colored bat habitat use in summer, but during winter they used bottomland forests, fields, and ponds more than salt marsh and upland forests. During summer and winter, northern yellow bats used sites close to fresh water and salt marsh. Additionally, during summer they used fields, ponds, and salt marsh more than upland and bottomland forests, but in winter they used bottomland forests, fields, and ponds more than upland forest and salt marsh. Our results highlight important land cover types for bats in this area (e.g., bottomland forests, ponds, and salt marsh), and that habitat use changes between seasons. Accounting for and understanding how habitat use changes throughout the year will inform managers about how critical habitat features may vary in their importance to bats throughout the year. © 2021 The Wildlife Society.     

Activity patterns of the Ryukyu flying fox on a subtropical island: responses to seasonal changes in night length

Almost all chiropteran species are nocturnal, but some species are occasionally active during the daytime. We conducted radio-tracking surveys and direct observations of the Ryukyu flying fox, Pteropus dasymallus, in two different habitats—urbanized and forested areas—on a subtropical island from April 2002 to January 2006. We recorded the departure time and return time from/to day roosts as well as behavioral time budgets during the night. The departure and return times shifted in correspondence with seasonal changes in sunset and sunrise times. The Ryukyu flying fox tended to depart earlier in summer when the night length was shorter, suggesting that it adjusts its active period by departing earlier. On the contrary, the amount of foraging performed by the bats in urbanized areas decreased in the summer when fruits of Ficus microcarpa were more abundant, suggesting that the bats adjust their behavioral time budgets in line with local food availability. Daytime activity was observed only in the forested area. In conclusion, the duration of Ryukyu flying fox activity was found to primarily depend on seasonal changes in the light–dark cycle, and this bat may adjust its behavioral time budget according to local food availability and the intensity of human activities.     

Foraging habitats of Myotis emarginatus in Central Europe

We radio-tracked Myotis emarginatus in Upper Bavaria, Germany to identify the key-foraging habitats and to enable an adequate habitat management for this endangered species. The studied females foraged at a distance of up to 8.1 km around their colony roost. The average distance of the foraging area was 3.7 km, where 70% of foraging areas were located within a distance of 5 km and 90% within 6 km of the nurseries. Moreover, these bats spent about 75% of their foraging time within 5 km and 85% within a 6-km radius. To reach the foraging areas, the bats usually used riparian woodlands, hedges and tree lines as flight paths. Specifically, 46.9% of the foraging areas were located in forests, 24.5% in cow sheds, 18.4% in riparian woodlands along streams and 10.2% in fields, villages, orchards, hedges and groves on open land. On average, the bats foraged in forests for 56.2% of the time, during which habitat allocation was possible. In cow sheds the percentage was 29.2%, in riparian deciduous woodland 11.5% and in the other habitats 3.1%. Within forests M. emarginatus avoided foraging in spruce monocultures. Pure stands of spruce covered 45% of the total forest area, but only 10% of the foraging areas were located in this forest type. Deciduous forests on the other hand were much more common as foraging sites (40% versus 11%). Therefore, the availability of native deciduous forest and of fly-infested stables within a radius of 6 km around the colony roosts should be the focus of conservation concepts for M. emarginatus.     

The effect of bark beetle infestation and salvage logging on bat activity in a national park

Salvage logging—the removal of dead trees in disturbed forest stands—has been controversially discussed. We investigated the impact of bark beetle attacks and subsequent salvage logging on insectivorous bats in a temperate mountain forest. We quantified bat activity (25,373?min counts; 32 plots) using batcorders during 221 all-night surveys in stands killed by bark beetles, with dead trees removed or not, and in vital, single- or multi-layered mature forest stands. We analysed the differences in activity of all bats in general and of bats of foraging guilds (open habitat, forest edge, closed habitat) in these habitats using a generalized linear Poisson mixed model, with plot and observation as random factors, and temperature and habitat as fixed factors. Only open-habitat foragers were slightly more active in salvage-logged stands than in bark-beetle-affected stands; they generally benefited from an open forest canopy, whereas closed-habitat foragers did not. Our results indicated that: (1) bats are less affected by salvage logging after a disturbance of a magnitude typical for European forests, probably because enough roosts are present in surrounding areas, (2) habitats for open foragers are improved by bark beetle infestation and (3) bats are poor bioindicators of negative impacts of salvage logging after natural disturbance in forests with a composition typical for Central Europe.     

Diversity of understory birds in old stands of native and Eucalyptus plantations

We compared the vegetation structure between old (>70 year) stands of planted diversified native forests and stands of Eucalyptus tereticornis embedded in a mosaic of Eucalyptus stands. We then tested for differences in the abundance, species richness, species composition, and ecological traits (forest dependence, sensitivity to forest fragmentation, and diet) of the understory bird assemblages inhabiting both kinds of stands. We expected differences in the structure of the bird assemblages because of the different origins and management strategies (contrary to native stands, Eucalyptus stands were selectively logged in the past). Three stands of each habitat (native and Eucalyptus) were sampled with mist nets during 11 months. Eucalyptus stands had a denser understory, whereas native plantations had a more developed vertical structure and a greater density of native trees. The abundance distribution of bird species was more homogeneous in Eucalyptus than in native stands. Eucalyptus had slightly higher species richness (36 species) than native stands (32 species). The composition of species and the occurrence of the diet, forest dependence, and sensitivity to forest fragmentation categories were similar between habitats. Some bird species (e.g. Turdus leucomelas), however, were more abundant in one habitat over the other. Old stands of Eucalyptus and planted native forest can harbor a diverse bird community similar in structure but not exactly equivalent for individual bird species. Planting native diversified forests and keeping set‐aside stands of the exotic tree should be viewed as complementary rather than alternative strategies for maintaining bird diversity within plantations.     

Pollination by the locally endangered island flying fox (Pteropus hypomelanus) enhances fruit production of the economically important durian (Durio zibethinus)

Fruit bats provide valuable pollination services to humans through a unique coevolutionary relationship with chiropterophilous plants. However, chiropterophily in the Old World and the pollination roles of large bats, such as flying foxes (Pteropus spp., Acerodon spp., Desmalopex spp.), are still poorly understood and require further elucidation. Efforts to protect these bats have been hampered by a lack of basic quantitative information on their role as ecosystem service providers. Here, we investigate the role of the locally endangered island flying fox Pteropus hypomelanus in the pollination ecology of durian (Durio zibethinus), an economically important crop in Southeast Asia. On Tioman Island, Peninsular Malaysia, we deployed 19 stations of paired infrared camera and video traps across varying heights at four individual flowering trees in a durian orchard. We detected at least nine species of animal visitors, but only bats had mutualistic interactions with durian flowers. There was a clear vertical stratification in the feeding niches of flying foxes and nectar bats, with flying foxes feeding at greater heights in the trees. Flying foxes had a positive effect on mature fruit set and therefore serve as important pollinators for durian trees. As such, semi‐wild durian trees—particularly tall ones—may be dependent on flying foxes for enhancing reproductive success. Our study is the first to quantify the role of flying foxes in durian pollination, demonstrating that these giant fruit bats may have far more important ecological, evolutionary, and economic roles than previously thought. This has important implications and can aid efforts to promote flying fox conservation, especially in Southeast Asian countries.     

Abundance responses of frugivorous bats (Stenodermatinae) to coffee cultivation and selective logging practices in mountainous central Veracruz,Mexico

Shifting and permanent cultivation, selective logging, cattle production and coffee plantations are among the most important factors in montane cloud forest conversion and disturbance. Although shaded-coffee plantations can contribute to the preservation of local species richness, abundance of organisms could be determined by habitat resource availability in agricultural landscapes. We compared abundance of Sturnira and Artibeus bats (Phyllostomidae, Stenodermatinae), in shade coffee plantations and disturbed cloud forest fragments, which represent habitats with different chiropterochorous plant density. We also investigated the relationship between bat species abundance and food plant richness, abundance and diversity. We captured 956 bats, 76% in cloud forest fragments and 24% in shaded coffee plantations. Abundance of Sturnira spp. (small bats) was greater in cloud forest than in coffee plantations, but Artibeus spp. (large bats) abundance was similar in both habitats. Chiropterochorous plant abundance was positively related with bat abundance for Sturnira spp., while chiropterochorous plant richness and diversity were negatively related for Artibeus spp. This suggests that frugivorous bats with different morphological and ecological characteristics respond differentially to anthropogenic activities. For landscape management purposes, the maintenance and augmentation of diverse food resources, for frugivorous bats with different foraging requirements in coffee plantations, will benefit the resilience of bats to modification of their natural habitat.     

Diurnal Habitat Relationships of Canada Lynx in an Intensively Managed Private Forest Landscape in Northern Maine

Abstract In March 2000, Canada lynx (Lynx canadensis) were listed as a federally threatened species in 14 states at the southern periphery of their range, where lynx habitat is disjunct and snowshoe hare (Lepus americanus) densities are low. Forest conditions vary across lynx range; thus, region-specific data on the habitat requirements of lynx are needed. We studied lynx in northern Maine, USA, from 1999 to 2004 to assess quality and potential for forests in Maine to sustain lynx populations. We trapped and radiocollared 43 lynx (21 M, 22 F) during this period and evaluated diurnal habitat selection by 16 resident adult lynx (9 M, 7 F) monitored in 2002. We evaluated lynx selection of 8 habitats at multiple spatial scales, and related lynx habitat selection to snowshoe hare abundance. Lynx preferred conifer-dominated sapling stands, which supported the highest hare densities on our study site (x̄ = 2.4 hares/ha), over all other habitats. The habitats where lynx placed their home ranges did not differ by sex. However, within their home ranges, males not only preferred conifer-dominated sapling stands, but also preferred mature conifer, whereas females singularly preferred conifer-dominated sapling stands. Approximately one-third of Maine's spruce-fir forest and nearly 50% of our study area was regenerating conifer or mixed-sapling forest, resulting from a disease event and intensive forest management (e.g., large clear-cuts). Our findings suggest that current habitat conditions in Maine are better than western montane regions and approach conditions in boreal forests during periods of hare abundance. We recommend that forest landowners maintain a mosaic of different-aged conifer stands to ensure a component of regenerating conifer-dominated forest on the landscape.     

Numerical and Functional Responses of Forest Bats to a Major Insect Pest in Pine Plantations

Global change is expected to modify the frequency and magnitude of defoliating insect outbreaks in forest ecosystems. Bats are increasingly acknowledged as effective biocontrol agents for pest insect populations. However, a better understanding is required of whether and how bat communities contribute to the resilience of forests to man- and climate-driven biotic disturbances. We studied the responses of forest insectivorous bats to a major pine defoliator, the pine processionary moth pityocampa, which is currently expanding its range in response to global warming. We used pheromone traps and ultrasound bat recorders to estimate the abundance and activity of moths and predatory bats along the edge of infested pine stands. We used synthetic pheromone to evaluate the effects of experimentally increased moth availability on bat foraging activity. We also evaluated the top-down regulation of moth population by estimating T. pityocampa larval colonies abundance on the same edges the following winter. We observed a close spatio-temporal matching between emergent moths and foraging bats, with bat activity significantly increasing with moth abundance. The foraging activity of some bat species was significantly higher near pheromone lures, i.e. in areas of expected increased prey availability. Furthermore moth reproductive success significantly decreased with increasing bat activity during the flight period of adult moths. These findings suggest that bats, at least in condition of low prey density, exhibit numerical and functional responses to a specific and abundant prey, which may ultimately result in an effective top-down regulation of the population of the prey. These observations are consistent with bats being useful agents for the biocontrol of insect pest populations in plantation forests.     

A habitat suitability analysis at multi-spatial scale of two sympatric flying fox species reveals the urgent need for conservation action

In this study we used a multi-spatial scale approach to investigate habitat suitability, roosting characteristics, and ecological niche in two flying fox species on the Comoros Islands—Pteropus livingstonii and Pteropus seychellensis comorensis. At a broad scale, we assessed the ecological niche and habitat suitability for both species using the Species Distribution Modeling method based on the recent ensembles of small models (ESM) approach. At a fine scale, Ecological Niche Factor Analysis (ENFA) was applied to assess habitat selection by each species. Direct observation was used at each roost to estimate the total number of individuals and to identify the roost characteristics. At both broad and fine scales, the analyses highlighted clear niche partitioning by the two species. We found that P. livingstonii has a very limited distribution, restricted to steep, high-elevation slopes of the islands’ remaining natural forests, and the patterns were the same for roosting, foraging sites and the entire habitat. By contrast, P. s. comorensis has a relatively large geographic range that extends over low-elevation farmlands and villages and it was negatively correlated to natural forest across the entire area and all roosting sites, but its foraging areas were positively correlated to natural forest and high elevation areas. Both species selected large, tall trees for roosting. The total number of individuals in the studied area was estimated to be 1243 P. livingstonii and 11,898 P. s. comorensis. The results of our study demonstrated that these two species use different habitat types and ensure different ecosystem services in pollination and seed dispersion and thus are both critical for maintaining overall ecosystem dynamics. However, the currently high level of hunting pressure and roost disturbance makes them vulnerable to extinction. To ensure the viability of both species, conservation measures need to be taken by the Comoros government.     

Evening Bat Summer Roost-Site Selection on a Managed Pine Landscape

ABSTRACT Creation and maintenance of forested corridors to increase landscape heterogeneity has been practiced for decades but is a new concept in intensively managed southern pine (Pinus spp.) forests. Additionally, more information is needed on bat ecology within such forest systems. Therefore, we examined summer roost-site selection by evening bats (Nycticeius humeralis) in an intensively managed landscape with forested corridors in southeastern South Carolina, USA, 2003–2006. We radiotracked 53 (26 M, 27 F) adult evening bats to 75 (31 M, 44 F) diurnal roosts. We modeled landscape-level roost-site selection with logistic regression and evaluated models using Akaike's Information Criterion for small samples. Model selection results indicated that mature (≥40 yr) mixed pine-hardwood stands were important roost sites for male and lactating female evening bats. Upland forested corridors, comprised of mature pine or mixed pine-hardwoods, were important roosting habitats for males and, to a lesser extent, lactating females. Male roosts were farther from open stands and lactating female roosts were farther from mid-rotation stands than randomly selected structures. Our results suggest roost structures (i.e., large trees and snags) in mature forests are important habitat components for evening bats. We recommend maintaining older (>40 yr old) stand conditions in the form of forest stands or corridors across managed landscapes to provide roosting habitat. Furthermore, our results suggest that an understanding of sex-specific roost-site selection is critical for developing comprehensive guidelines for creating and maintaining habitat features beneficial to forest bats.     

Poor effectiveness of Natura 2000 beech forests in protecting forest-dwelling bats

With the establishment of the Natura 2000 (N2000) network, the European Union intends to develop strategies to conserve Europe's threatened habitats and species, including bats. Forest-dwelling bats are highly reliant on forest structures, such as snags and hollow trees, which the bats need as roosts. The decrease in such forest microhabitats significantly affects the habitat use and, therefore, the activity in forests. To determine whether N2000 beech forests under active timber production offer better habitats for bats compared to commercially used non-N2000 forests, we measured the bat activity and assessed the potential roosts in trees and snags in eleven pairs of stands. All survey stands represented mesotrophic beech forests (Fagus sylvatica L.) of the N2000 habitat type 9130 (Asperulo-Fagetum) in three European Biogeographic Regions. The activity of all bat species, the activity of priority N2000 species, the species number, the number of trees with roosts and the snag volume did not differ significantly between the N2000 and non-N2000 stands. We conclude that the current management of the N2000 beech forests is almost identical to that of non-N2000 commercial forests, and thus, the N2000 status has not led to an increase of bat-relevant habitat variables yet. Consequently, additional efforts beyond the administrative assignment of N2000 areas are required to build and ensure an ecologically effective and sustainable network of beech forests in Europe, including increasing important forest requirements for bats, such as roosts and snags.     

The role of farm structure on bird assemblages around a Kenyan tropical rainforest

Although it is clear that the farmlands neighbouring fragmented forests are utilized by some forest birds, it is not clear how birds in general respond to farmland habitat mosaic. An effort was made to determine how bird density and foraging assemblages were influenced by farm structural characteristics and distance from forest edge. Thirty farms up to a distance of 12 km around Kakamega forest in western Kenya were studied. Farm structure entailed size, hedge volume, habitat heterogeneity, woody plant density, plant diversity and crop cover. Birds were surveyed using line transects and DISTANCE analyses and classified into six feeding guilds and three habitat associations. Size of farms increased away from the forest, as woody plant density, plant diversity, indigenous trees and subsistence crop cover declined. The most important farm structure variable was hedge volume, which enhanced bird species richness, richness of shrub‐land bird species and insectivorous bird density (R = 0.58, P < 0.01). Bird density increased with tree density while indigenous trees were suitable for insectivores and nectarivores. There were very few forest bird encounters. Agricultural practices incorporating maintenance of hedges and sound selection of agroforestry trees can enhance conservation of birds on farmland, though, not significantly for forest species.     

Isolation and characterisation of polymorphic microsatellite loci in the vulnerable spectacled flying fox, Pteropus conspicillatus

The spectacled flying fox, Pteropus conspicillatus, is listed as vulnerable in Australia and is under threat from numerous impacts. Primers to amplify eight co-dominant microsatellite loci were designed for Pteropus conspicillatus, based on an enriched genomic library. Four loci were monomorphic in this species while the remaining four loci were highly polymorphic with 16–23 alleles. Two of the four monomorphic loci were found to be polymorphic in Pteropus alecto, a closely related congener. All but one of the six polymorphic loci were in Hardy Weinberg equilibrium. Additionally, six microsatellite loci isolated for Pteropus rodricensis were tested against individuals of P. conspicillatus with all loci amplifying reliably. These loci will be used to investigate population genetic structure in the vulnerable spectacled flying fox.