Tropical Secondary Forest Management Influences Frugivorous Bat Composition,Abundance and Fruit Consumption in Chiapas,Mexico

Most studies on frugivorous bat assemblages in secondary forests have concentrated on differences among successional stages, and have disregarded the effect of forest management. Secondary forest management practices alter the vegetation structure and fruit availability, important factors associated with differences in frugivorous bat assemblage structure, and fruit consumption and can therefore modify forest succession. Our objective was to elucidate factors (forest structural variables and fruit availability) determining bat diversity, abundance, composition and species-specific abundance of bats in (i) secondary forests managed by Lacandon farmers dominated by Ochroma pyramidale, in (ii) secondary forests without management, and in (iii) mature rain forests in Chiapas, Southern Mexico. Frugivorous bat species diversity (Shannon H’) was similar between forest types. However, bat abundance was highest in rain forest and O. pyramidale forests. Bat species composition was different among forest types with more Carollia sowelli and Sturnira lilium captures in O. pyramidale forests. Overall, bat fruit consumption was dominated by early-successional shrubs, highest late-successional fruit consumption was found in rain forests and more bats consumed early-successional shrub fruits in O. pyramidale forests. Ochroma pyramidale forests presented a higher canopy openness, tree height, lower tree density and diversity of fruit than secondary forests. Tree density and canopy openness were negatively correlated with bat species diversity and bat abundance, but bat abundance increased with fruit abundance and tree height. Hence, secondary forest management alters forests’ structural characteristics and resource availability, and shapes the frugivorous bat community structure, and thereby the fruit consumption by bats.     

Bat assemblage structure in two dry forests of Colombia: Composition, species richness, and relative abundance

We studied the composition, species richness, and relative abundance of bat assemblages in the Colombian dry forests of Chicamocha and Patía. In Chicamocha, 11 bats of the family Phyllostomidae were captured with mist-nets, corresponding to 85–100% of the potential phyllostomids species in the area. Two bats of the family Vespertilionidae were also captured in Chicamocha. In Patía, 12 species were captured with mist-nets, all Phyllostomidae, representing 72–100% of the estimated total number of species in the zone. Minor differences in number of species and composition were detected among sampling periods in Chicamocha. The most common species in this dry forest were Glossophaga longirostris and Sturnira lilium. In Patía, notable differences in the number of species and composition were observed among sampling periods, and the most common species were Artibeus jamaicensis, Carollia perspicillata and Phyllostomus discolor. Arid-zone dwelling bats were absent in Patía and we suggest that this absence may be associated with the isolation of Patía from other northern dry zones of Colombia since Quaternary times. There was also low abundance of bats in Patía, which appears to be related to human disturbance. The most abundant phyllostomid bat species in the two dry forests studied are those that include fruit and/or nectar-pollen from columnar cacti as an important proportion of their diets.     

Three‐dimensional stratification pattern in an old‐growth lowland forest: How does height in canopy and season influence temperate bat activity?

The study of animal–habitat interactions is of primary importance for the formulation of conservation recommendations. Flying, gliding, and climbing animals have the ability to exploit their habitat in a three‐dimensional way, and the vertical canopy structure in forests plays an essential role for habitat suitability. Forest bats as flying mammals may seasonally shift their microhabitat use due to differing energy demands or changing prey availability, but the patterns are not well understood. We investigated three‐dimensional and seasonal habitat use by insectivorous bats in a temperate lowland old‐growth forest, the Belovezhskaya Pushcha in Belarus. We acoustically sampled broadleaved and mixed coniferous plots in the forest interior and in gaps in three heights during two reproductive periods (pregnancy/lactation vs. postlactation). In canopy gaps, vertical stratification in bat activity was less pronounced than in the forest interior. Vertical activity patterns differed among species. The upper canopy levels were important foraging habitats for the open‐space forager guild and for some edge‐space foragers like the Barbastelle bat Barbastella barbastellus and the soprano pipistrelle Pipistrellus pygmaeus. Myotis species had highest activity levels near the ground in forest gaps. Moreover, we found species‐dependent seasonal microhabitat shifts. Generally, all species and species groups considered except Myotis species showed higher activity levels during postlactation. Myotis species tended toward higher activity in the forest interior during postlactation. P. pygmaeus switched from high activity levels in the upper canopy during pregnancy and lactation to high activity levels near the ground during postlactation. We conclude that a full comprehension of forest bat habitat use is only possible when height in canopy and seasonal patterns are considered.     

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.     

Phyllostomid Bat Assemblage Structure in Amazonian Flooded and Unflooded Forests

In Amazonia, the assemblages of several taxa differ significantly between upland terra firme and white‐water flooded várzea forests, but little is known about the diversity and distribution of bats in these two forest types. We compare the spatio‐temporal patterns of bat assemblage composition and structure in adjacent terra firme and várzea forests in the lower Purus River region of central Brazilian Amazonia. Bats were sampled using mist nets at five sites in each forest type during 40 nights (2400 net‐hours). We captured 1069 bats representing 42 species and Phyllostomidae bats comprised 99.3 percent of all captures. The bat assemblages in várzea and terra firme forests were significantly different, mainly due to a marked dissimilarity in species composition and in the number of captures during high‐water season. In addition, bat assemblages within forest types differed significantly between seasons for both terra firme and várzea. Frugivores dominated the bat assemblages in both forest types. Overall guild structure did not change between várzea and terra firme or between seasons, but frugivore and animalivore abundance increased significantly in várzea forest during the inundation. The difference in assemblage structure observed in the high‐water season is probably caused by the annual várzea flooding, which provides an effective barrier to the persistence of many understory bats. We also hypothesize that some bat species may undertake seasonal movements between forest types in response to fruit abundance, and our results further underline the importance of floodplain habitats for the conservation of species in the Amazon.     

Vertical stratification of an avian community in New Guinean tropical rainforest

Vertical stratification of avian communities has been studied in both temperate and tropical forests; however, the majority of studies used ground-based methods. In this study we used ground-to-canopy mist nets to collect detailed data on vertical bird distribution in primary rain forest in Wanang Conservation Area in Papua New Guinea (Madang Province). In total 850 birds from 86 species were caught. Bird abundance was highest in the canopy followed by the understory and lowest in the midstory. Overall bird diversity increased towards the canopy zone. Insectivorous birds represented the most abundant and species-rich trophic guild and their abundances decreased from the ground to canopy. The highest diversity of frugivorous and omnivorous birds was confined to higher vertical strata. Insectivorous birds did not show any pattern of diversity along the vertical gradient. Further, insectivores preferred strata with thick vegetation, while abundance and diversity of frugivores increased with decreasing foliage density. Our ground-to-canopy (0–27 m) mist netting, when compared to standard ground mist netting (0–3 m), greatly improved bird diversity assessment and revealed interesting patterns of avian community stratification along vertical forest strata.     

Bat Diversity and Movement in an Agricultural Landscape in Matiguás, Nicaragua

Although agriculture dominates much of Central America, little is known about the bat assemblages that occur within agricultural landscapes and how bats use different types of tree cover within these landscapes. Using mist-nets and a mark-recapture protocol, we compared bat diversity and movement across six types of tree cover within an agricultural landscape in central Nicaragua. The tree cover types surveyed included secondary forests, riparian forests, forest fallows, live fences, pastures with high tree cover and pastures with low tree cover. We captured a total of 3084 bats of 39 species, including two new species records for the country ( Lonchorhina aurita and Molossops greenhalli ). Of these, 2970 bats and 27 species were in the Phyllostomidae family. There were significant differences in mean species density, abundance and evenness of phyllostomid bats across the different types of tree cover, but not in bat diversity. Riparian forests had the highest mean species density and bat abundance per plot. In contrast, mean bat abundance and species density were lowest in pastures with low tree cover. Of the 1947 phyllostomid bats marked, a total of 64 bats of eight species were recaptured. The average linear distance between extra-site recaptures was 2227 m (± 228 SE) and the maximum distance was 10.6 km. Bats were recorded moving between almost all types of tree cover, and especially to and from riparian forests. Our study suggests that agricultural landscapes retaining a heterogeneous tree cover may maintain a diverse bat assemblage, and that bats visit and use a variety of tree cover types within the agricultural matrix.     

Patterns in the use of rainforest vertical space by Neotropical aerial insectivorous bats: all the action is up in the canopy

In tropical rainforests environmental conditions vary dramatically from the ground to the canopy, resulting in a marked stratification in the way vertical space is used by organisms, but research work is often limited to the understorey layer. Aerial insectivorous bats are a highly diverse group that plays key roles in the ecology of rainforests, but their use of vertical space remains elusive. Using automatic ultrasound recording stations placed in the canopy, subcanopy and understorey we tested if bat activity and species diversity are vertically stratified, both in the forest interior and near the edges of water bodies. These patterns were tested separately for individual species, and for two functional groups – open space and edge space bats. Insectivorous bat activity increased by roughly seven fold, and species diversity doubled, from the understorey to the canopy. Both edge space and open space bats were more active in the upper strata, but this tendency was much more accentuated in the latter. Myotis riparius was the only species with greater activity near the understorey. These patterns were altered at the edges of water bodies, where vertical stratification was much less marked. The observed patterns are parsimoniously explained by constraints imposed by vegetation clutter that change across strata, which affect bat species differently. Only bats better adapted to closed spaces are usually capable of foraging within the understorey, whereas the majority of species can exploit the free spaces immediately below the canopy; open space bats seem to concentrate their activity above the canopy. This importance of the inter strata open spaces for bat foraging highlights the need to preserve pristine stratified rainforests, as even selective logging usually disrupts vertical stratification. Moreover, the concentration of insectivorous bats at the upper strata of rainforests underlines the need to include canopy level sampling in ecological studies.     

On the structure of rodent communities in tropical forests of southern Vietnam

The abundance, species diversity, and spatial distribution of small mammals were studied in two localities in southern Vietnam: the Cat Tien National Park and the Binh Chau Nature Reserve. Rodents were trapped on the ground and in the tropical forest canopy (2–12 m above the ground), marked, and released on the spot. Species diversity of small mammals in undisturbed tropical forest (9 species) was higher than in disturbed tropical forest (14 species). The rat Maxomys surifer and the squirrel Callosciurus flavimanus were dominant. In undisturbed forest, the frequencies of rodent occurrence on the ground and in the canopy were of the same order; in disturbed forest, rodents occurred in the canopy an order of magnitude less frequently than on the ground.     

Two new species of yellow-shouldered bats,genus Sturnira Gray, 1842 (Chiroptera,Phyllostomidae) from Costa Rica,Panama and western Ecuador

Two new species of yellow-shouldered bats Sturnira Gray, 1842 (Chiroptera, Phyllostomidae) from Central America and western South America are described using molecular and morphological data. The two new species, which occur in Costa Rica and Panama and in western Ecuador, were previously confused with S. ludovici, and S. lilium and S. luisi, respectively. Sturnira now includes 22 described species, making it the most speciose genus in the Neotropical family Phyllostomidae.     

Effects of Agricultural Intensification on the Assemblage of Leaf-Nosed Bats (Phyllostomidae) in a Coffee Landscape in Chiapas,Mexico

The agricultural matrix surrounding forested areas serves critical functions as dispersal corridors and alternate habitat for wildlife. Agricultural intensification, however, can reduce the conservation value of these areas. To evaluate the effects of agroecosystem management on bat assemblages, we studied the abundance and diversity of leaf-nosed bats (family: Phyllostomidae) in southwestern Chiapas, Mexico, a landscape dominated by shade coffee agroforestry. During 2104 mist-net hour (MNH), we captured 3167 bats of 27 phyllostomid species. Total species richness in each land-use type varied from 24 species in forest fragments to 22 species in commercial shade polycultures. Although the cumulative observed species richness showed little change in response to management intensity, the number of bats captured per MNH declined significantly in the more intensively managed (i.e., low-shade monocultures) plantations. Intensively managed coffee plantations had lower phyllostomid diversity and species similarity, and had lower proportions of nectarivorous and animalivorous bats. Among frugivores, the proportion of large (>25 g) frugivores captured increased with management intensity. Recapture frequency was significantly higher than expected in forest fragments, and lower than expected in more intensively managed coffee. Our results suggest that less intensively managed coffee agroforests can serve as valuable feeding and commuting areas for most leaf-nosed bats, and that maintaining forest fragments in agricultural landscapes contributes to bat diversity. Declines in populations of gleaning insectivores, however, could compromise natural suppression of insect pests in these agricultural areas.     

Stable isotope canopy effects for sympatric monkeys at Ta? Forest,C?te d'Ivoire

This study tests the hypothesis that vertical habitat preferences of different monkey species inhabiting closed canopy rainforest are reflected in oxygen isotopes. We sampled bone from seven sympatric cercopithecid species in the Taï forest, Côte d''Ivoire, where long-term study has established taxon-specific patterns of habitat use and diet. Modern rib samples (n = 34) were examined for oxygen (δ¹⁸O_ap) and carbon (δ¹³C_ap) from bone apatite (‘bioapatite’), and carbon (δ¹³C_co) and nitrogen (δ¹⁵N_co) from bone collagen. Results are consistent for C₃ feeders in a closed canopy habitat. Low irradiance and evapotranspiration, coupled with high relative humidity and recycled CO₂ in forest understory, contribute to observed isotopic variability. Both δ¹³C_co and δ¹³C_ap results reflect diet; however, δ¹³C values are not correlated with species preference for canopy height. By contrast, δ¹⁸O_ap results are correlated with mean observed height and show significant vertical partitioning between taxa feeding at ground, lower and upper canopy levels. This oxygen isotope canopy effect has important palaeobiological implications for reconstructing vertical partitioning among sympatric primates and other species in tropical forests.     

EFFECT OF HABITAT AND FORAGING HEIGHT ON BAT ACTIVITY IN THE COASTAL PLAIN OF SOUTH CAROLINA

Abstract: We compared bat activity levels in the Coastal Plain of South Carolina among 5 habitat types: forested riparian areas, clearcuts, young pine plantations, mature pine plantations, and pine savannas. We used time-expansion radio-microphones and integrated detectors to simultaneously monitor bat activity at 3 heights (30, 10, 2 m) in each habitat type. Variation in vegetative clutter among sampling heights and among habitat types allowed us to examine the differential effect of forest vegetation on the spatial activity patterns of clutter-adapted and open-adapted bat species. Moreover, monitoring activity at 30, 10, and 2 m permitted us to also compare bat activity above and below the forest canopy. We detected calls of 5 species or species groups: eastern red/Seminole bats (Lasiurus borealis/L. seminolus), eastern pipistrelles (Pipistrellus subflavus), evening bats (Nycticeius humeralis), big brown bats (Eptesicus fuscus), and hoary bats (Lasiurus cinerius). At 2 and 10 m, bat activity was concentrated in riparian areas, whereas we detected relatively low levels of bat activity in upland habitats at those heights. Activity was more evenly distributed across the landscape at 30 m. Bat activity levels above the forest canopy were almost 3 times greater than within or below the canopy. We detected significantly greater activity levels of 2 open-adapted species (hoary and big brown bats) above rather than within or below the forest canopy. However, activity levels of 2 clutter-adapted species (eastern red/Seminole bats and eastern pipistrelles) did not differ above, within, or below the forest canopy. Despite classification as a clutter-adapted species, evening bat activity was greater above rather than within or below the forest canopy. We believe our results highlight the importance of riparian areas as foraging habitat for bats in pine-dominated landscapes in the southeastern United States. Although acoustical surveys conducted below forest canopies can provide useful information about species composition and relative activity levels of bats that forage in cluttered environments, our results showing activity above canopy suggest that such data may not accurately reflect relative activity of bats adapted to forage in more open conditions, and therefore may provide an inaccurate picture of bat community assemblage and foraging habitat use.     

Ensemble composition and activity levels of insectivorous bats in response to management intensification in coffee agroforestry systems

Shade coffee plantations have received attention for their role in biodiversity conservation. Bats are among the most diverse mammalian taxa in these systems; however, previous studies of bats in coffee plantations have focused on the largely herbivorous leaf-nosed bats (Phyllostomidae). In contrast, we have virtually no information on how ensembles of aerial insectivorous bats--nearly half the Neotropical bat species--change in response to habitat modification. To evaluate the effects of agroecosystem management on insectivorous bats, we studied their diversity and activity in southern Chiapas, Mexico, a landscape dominated by coffee agroforestry. We used acoustic monitoring and live captures to characterize the insectivorous bat ensemble in forest fragments and coffee plantations differing in the structural and taxonomic complexity of shade trees. We captured bats of 12 non-phyllostomid species; acoustic monitoring revealed the presence of at least 12 more species of aerial insectivores. Richness of forest bats was the same across all land-use types; in contrast, species richness of open-space bats increased in low shade, intensively managed coffee plantations. Conversely, only forest bats demonstrated significant differences in ensemble structure (as measured by similarity indices) across land-use types. Both overall activity and feeding activity of forest bats declined significantly with increasing management intensity, while the overall activity, but not feeding activity, of open-space bats increased. We conclude that diverse shade coffee plantations in our study area serve as valuable foraging and commuting habitat for aerial insectivorous bats, and several species also commute through or forage in low shade coffee monocultures.     

Vertical stratification of the termite assemblage in a neotropical rainforest

The importance of termites as decomposers in tropical forests has long been recognized. Studies on the richness and diversity of termite species and their ecological function have flourished in more recent times, but these have been mostly conducted in a thin stratum within a standing man’s reach. Our aims were to evaluate the specific richness and composition of the termite assemblage in the canopy of a tropical rainforest and to determine its originality with respect to the sympatric ground-level fauna. We conducted systematic searches for canopy termites, together with conventional sampling of the sympatric ground-level fauna, in the San Lorenzo forest, Panama. We hypothesized that (1) the canopy accommodates two categories of wood-feeding termites (long-distance foragers and small-colony “one-piece” species) and possibly soil-feeders in suspended soil-like habitats; (2) due to the abundance of soil-feeders, the overall diversity of the ground fauna is higher than that of the canopy; (3) differences in microclimate and resource accessibility favour vertical stratification among wood-feeders. Sixty-three canopy samples yielded ten species of termites, all wood-feeders. Five of these were not found at ground level, although a total of 243 ground samples were collected, representing 29 species. In addition to long-distance foragers (Microcerotermes and Nasutitermes spp.) and small-colony termites (mostly Kalotermitidae), the canopy fauna included Termes hispaniolae, a wood-feeding Termitidae from an allegedly soil-feeding genus, living in large dead branches. Soil-feeders were absent from the canopy, probably because large epiphytes were scarce. As predicted, the ground fauna was much richer than that of the canopy, but the species richness of both habitats was similar when only wood-feeders were considered. Vertical stratification was strongly marked among wood-feeders, as all common species, apart from the arboreal-nesting Microcerotermes arboreus, could unequivocally be assigned to either a ground or a canopy group. The canopy, therefore, contributes significantly to the total species richness of the termite assemblage, and the diversity, abundance and ecological importance of canopy termites in tropical rainforests may be higher than previously recognized.Electronic Supplementary Material Supplementary material is available to authorized users in the online version of this article at .     

The effects of rainforest canopy loss on arboreal dung beetles in Borneo: implications for the measurement of biodiversity in derived tropical ecosystems

We examine the effect of selective timber extraction, and corresponding forest canopy loss, on arboreal dung beetles in the tropical rainforests of Sabah, Malaysian Borneo. Changes in vertical distribution of beetles are measured through differences in the abundance of beetles in ground-based pitfall traps in primary, logged and plantation forest. Previous research has demonstrated that arboreal dung beetles are not generally collected in pitfall traps in primary forest, but are present in large numbers above the ground in canopy vegetation: the presence of arboreal beetles in pitfalls in areas of reduced or modified canopy cover may therefore reflect a response to the absence or modification of their usual habitat, and the proliferation of these beetles nearer to the ground. In this paper, statistically significant differences are found in the abundance of beetles in ground pitfall traps from logged forest compared to primary forest. Results show that virtually no arboreal dung beetles are recorded in primary forest traps, with an increased abundance of arboreal dung beetles in traps from logged and plantation forest, with 1.72% of the total number of arboreal beetles recorded from primary forest, 22.32% from logged forest, and 75.96% from plantation forest. The presence of arboreal dung beetles in plantations demonstrates that arboreal dung beetles can survive outside their normal habitat, and we relate these observations to adaptations to upper rainforest canopy conditions, and proliferation of these microclimatic conditions in man-made habitats. Results are also discussed in terms of their relevance to the measurement of species richness and diversity in logged and other derived ecosystems, where mixing of the ground-based and arboreal faunas occurs.     

Leishmania (L.) mexicana Infected Bats in Mexico: Novel Potential Reservoirs

Leishmania (Leishmania) mexicana causes cutaneous leishmaniasis, an endemic zoonosis affecting a growing number of patients in the southeastern states of Mexico. Some foci are found in shade-grown cocoa and coffee plantations, or near perennial forests that provide rich breeding grounds for the sand fly vectors, but also harbor a variety of bat species that live off the abundant fruits provided by these shade-giving trees. The close proximity between sand flies and bats makes their interaction feasible, yet bats infected with Leishmania (L.) mexicana have not been reported. Here we analyzed 420 bats from six states of Mexico that had reported patients with leishmaniasis. Tissues of bats, including skin, heart, liver and/or spleen were screened by PCR for Leishmania (L.) mexicana DNA. We found that 41 bats (9.77%), belonging to 13 species, showed positive PCR results in various tissues. The infected tissues showed no evidence of macroscopic lesions. Of the infected bats, 12 species were frugivorous, insectivorous or nectarivorous, and only one species was sanguivorous (Desmodus rotundus), and most of them belonged to the family Phyllostomidae. The eco-region where most of the infected bats were caught is the Gulf Coastal Plain of Chiapas and Tabasco. Through experimental infections of two Tadarida brasiliensis bats in captivity, we show that this species can harbor viable, infective Leishmania (L.) mexicana parasites that are capable of infecting BALB/c mice. We conclude that various species of bats belonging to the family Phyllostomidae are possible reservoir hosts for Leishmania (L.) mexicana, if it can be shown that such bats are infective for the sand fly vector. Further studies are needed to determine how these bats become infected, how long the parasite remains viable inside these potential hosts and whether they are infective to sand flies to fully evaluate their impact on disease epidemiology.     

Horizontal distribution affects the vertical distribution of native and invasive container‐inhabiting Aedes mosquitoes within an urban landscape

The vertical dimension constitutes an important niche axis along which mosquitoes may adjust their distribution. Here, we evaluated whether the vertical distribution of container‐inhabiting Aedes mosquitoes differs along a gradient of anthropogenic land‐use intensity within an urban landscape. Using a pulley system, we hung oviposition cups at three heights (ground level, 4.5, and 9 m) and in three habitats: forest, park, and a built environment. We hypothesized that mosquito abundance and diversity would be highest in the least disturbed forest habitat, decrease in the park, and be lowest at the UNC‐Greensboro campus. We also expected Aedes albopictus (Skuse) and Ae. triseriatus (Say) to mainly oviposit at ground level and Ae. hendersoni (Cockerell) at canopy height. Aedes albopictus was the most common species (68.8%) collected in all three habitat types and was the only species found in the built environment. In that habitat, Ae. albopictus exhibited a bimodal distribution with the lowest activity at the intermediate height (4.5 m). Aedes triseriatus (28.9%) did not differ in egg abundance between the forest and park habitats but did exhibit diverse vertical habitat use while avoiding the canopy in the park habitat. Aedes hendersoni (2.3%) was the most sylvatic species and oviposited only at ground level. Our results indicate that the vertical distribution of mosquitoes is affected by the type of habitat in which they occur, and that this variation could be driven via local‐scale modification of microclimatic factors.