Foraging strategies of generalist and specialist Old World nectar bats in response to temporally variable floral resources

Foraging theory predicts that generalist foragers should switch resources more readily, while specialist foragers should remain constant to preferred food resources. Plant‐pollinator interactions provide a convenient system to test such predictions because floral resources are often temporally patchy, thus requiring long‐lived pollinators to switch resources seasonally. Furthermore, flowering phenologies range from ‘steady‐state’ (low‐rewarding but highly reliable) to ‘big‐bang’ (high‐rewarding but ephemeral) plant species. We assessed how nectarivorous Old World bats respond to this temporally variable floral environment by examining their diets throughout the year. Over 15 months of fieldwork in southern Thailand, we simultaneously: (1) recorded the flowering phenologies of six bat‐pollinated plant taxa; and (2) assessed the diets of seven common flower‐visiting bat species. As predicted, the generalist nectarivore (Eonycteris spelaea) frequently switched diets and utilized both big‐bang and steady‐state resources, while the specialist nectarivores (Macroglossus minimus and M. sobrinus) foraged on one or two steady‐state plant species year‐round. Our results suggest that larger and faster bat species are able to fly longer distances in search of big‐bang resources, while smaller bat species rely on highly predictable food resources. This study supports the theory that generalist foragers have flexible diets, while specialist species restrict foraging to preferred floral resources even when other floral resources are more abundant. Moreover, these findings demonstrate how plant flowering phenology and pollinator diet breadth can shape the frequency and constancy of pollinator visits; we further discuss how such interactions can influence the potential extent of gene flow within a patchy floral environment.     

Old World fruit bats can be long-distance seed dispersers through extended retention of viable seeds in the gut

Seed dispersal and pollination by animals play a crucial role in the maintenance of forest ecosystems worldwide. Frugivorous bats are important pollen and seed dispersers in both the Palaeo- and Neotropics, and at least 300 plant species are known to rely on Old World fruit bats (Megachiroptera, Pteropodidae) for their propagation. However, rapid food transit times (generally less than 30 minutes) in frugivorous bats have been thought to limit their ability to disperse seeds to just a few tens of kilometres. Here we demonstrate regular daytime (greater than 12 hours) retention of food and viable fig seeds (Ficus, Moraceae) in the gut of the Old World fruit bat Cynopterus sphinx: a behaviour not previously reported for any frugivorous bat. Field observations indicate that this behaviour also occurs in other genera. Old World fruit bats are highly mobile and many species undertake considerable foraging and migration flights. Our findings indicate that Old World fruit bats have the potential to disperse small seeds hundreds of kilometres. This necessitates a reappraisal of their importance in transporting zoochorous seeds to remote areas and facilitating gene flow between isolated populations of plants, both within mainlands and across ocean barriers.     

Seasonal variation in food availability and relative importance of dietary items in the Gambian epauletted fruit bat (Epomophorus gambianus)

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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.     

Ecological constraints on fruit initiation by Calyptrogyne ghiesbreghtiana (Arecaceae): FLORAL HERBIVORY,POLLEN AVAILABILITY,AND VISITATION BY POLLINATING BATS

Failure of a flower to initiate fruit can be attributed to an insufficiency of resources, genetic incompatibilities, or ecological constraints on pollination. Flower visitors can increase fruit initiation by pollinating flowers, or decrease fruit initiation by damaging them. If, however, resources limit fruit initiation, variation in visitation may be relatively unimportant. Visitors to Calyptrogyne ghiesbreghtiana, a rain forest understory palm, leave a record of their activity by feeding on floral tissue. I recorded variation in inflorescence visitation, floral display, and fruit initiation during 7 mo over two field seasons. On average, bats visited 50-60% of flowers; katydids and scarab beetles visited and damaged the remaining 40-50%. Four species of bat (Phyllostomidae) captured near inflorescences were found to be carrying C. ghiesbreghtiana pollen. Approximately 18% of flowers visited by bats and not damaged by insects initiated fruit, whereas insect-damaged flowers set a negligible number. Some variation in fruit initiation was explained by nightly variation in the proportion of inflorescences at each site that were potential pollen donors (male-phase inflorescences). Visitors responded to variation in floral display; katydids damaged more flowers on taller inflorescences, and in one season bats visited a greater proportion of flowers on inflorescences with many flowers. However, floral display only explained a small amount of variation in visitation. A similar amount of variation in flower visitation was explained by the degree to which inflorescences were obstructed by surrounding vegetation. I used path analysis to summarize the overall effect of floral display on visitation by both pollinators and florivores and the effect of visitation and site sex ratio on fruit initiation. Because almost all flowers were visited by bats or damaged by katydids, the amount of bat visitation was strongly negatively correlated with katydid damage. In spite of the low fruit set from bat-visited flowers, variation in bat visitation explained 27-37% of the variation in fruit initiation.     

Seed Rain Produced by Bats and Birds in Forest Islands in a Neotropical Savanna1

The high degree of isolation of forest “islands” relative to “continental” forested areas creates a naturally fragmented landscape in the savanna ecosystem. Because fragmentation can affect the intensity and quality of biological interactions (e.g., seed dispersal) we examined the abundance and species richness of seed rain produced by birds and bats in three different parts of forest islands (center, edge, and exterior) located at the Estación Biológica del Beni, Bolivia. Despite the fact that we found higher species density of seeds in the seed rain at the center of forest islands, when comparing species richness corrected for observed differences in density, species richness was higher at the edge of islands. The three parts of the islands did not differ in total number of seeds. Three genera (Byrsonima, Ficus, and Piper) contributed the most seeds to the seed rain. We found differences in the abundance of dispersed seeds probably because of the variation related with the disturbance line, where the savanna matrix interacts with the forest islands. Carollia perspicillata, Carollia brevicauda, and Sturnira lilium were the bats that contributed most to seed dispersal within forest islands, and Schistochlamys melanopis and Tyranneutes stolzmanni were the most important birds. The movement of seeds produced by bats and birds within forest islands of the savanna is crucial to assure the continuity of ecological process and dynamics of these forest islands.     

广州地区犬蝠的食性

2004年5月-2006年4月采用拾遗法、粪便内容物分析法及实地观察对广州地区常见的食果蝙蝠-犬蝠(Cynopterus sphinx)进行了食性研究。对26份食物残留物和粪便样品的分析结果表明:犬蝠的食物包含13科21种的植物果实,3科3种的植物叶片,如:蒲桃(Syzygium jambos)、蒲葵(Livistona subglobosa)及龙眼(Dimocarous longan)的果实。其食性随果实的成熟季节而出现明显的季节性变化,夏秋两季大量食用各种水果,而在食物欠缺的春冬两季则主要食用棕榈科蒲葵的种子。广州地区犬蝠的繁殖期在每年的5-10月。     

Do Bats Roost and Forage in Shade Coffee Plantations? A Perspective from the Frugivorous Bat Sturnira hondurensis

Shade coffee plantations are considered important habitats for frugivorous bats. However, it is not known if bats use this agricultural habitat for shelter, food resources, or both. This study addresses these questions using the highland yellow‐shouldered bat (Sturnira hondurensis) as an example. Twenty‐six adult individuals of S. hondurensis were captured, 50 percent in tropical montane cloud forest (TMCF) and 50 percent in shade coffee plantations (SCP) in Veracruz, Mexico, and each was fitted with a radio transmitter for locating roosts and feeding areas. Data were obtained from 24 of them. The fieldwork was conducted between October 2010 and October 2011 covering all seasons. Twenty‐two day roosts were located in the cavities of twelve different species of tree. Roosts located in TMCF differed significantly from those in SCP, having a smaller crown area and a greater species richness and density of plants around the roost. In SCP, both the average home range and the average core use area were smaller than in TMCF, but the differences were not statistically significant. Distances travelled by bats were generally longer and more variable in the SCP; the distance between capture site and foraging site was significantly greater in SCP than in TMCF. In SCP, there were fewer understory chiropterochorous plants, which are the main item in the diet of this bat and many other sympatric species of frugivorous bats. Although S. hondurensis does use roosts and foraging sites in the SCP, it is important to note that this species and others with similar requirements primarily depend on the preservation of intact forest adjacent to modified landscapes, where roosts and fruit are constantly available in abundance. Management practices should guarantee a greater density and diverse of trees and the preservation of understory plants with fruits in the coffee plantations that allow a long‐term survival of frugivorous bats populations.     

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.     

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.     

Predictive Occurrence Models for Bat Species in California

Abstract: The ability to accurately predict the potential occurrence of species of management concern is useful for wildlife managers, particularly for those whose management activities involve large areas where sampling is difficult due to logistical or financial constraints. During the summers of 2002 and 2003, we used mist nets to capture bats (Myotis yumanensis, M. californicus, M. evotis, M. thysanodes, Eptesicus fuscus, Lasionycteris noctivagans, Tadarida brasiliensis, Antrozous pallidus, Lasiurus borealis, and Lasiurus cinereus) in Whiskeytown National Recreation Area in north-central California, USA. We used landscape-scale variables, logistic regression, and Akaike's Information Criterion (AIC_c) to model species distributions and produce spatially discerning predictive occurrence maps. We developed a priori models that we used to determine which landscape-scale variables best discriminated between capture sites and non-capture sites. The odds of capturing a bat were 3.3 greater when total edge increased by 10,000 m, whereas for Yuma myotis (Myotis yumanensis), the odds of predicting presence were 0.2 greater when distance to lakes and ponds decreased by 2,000 m. Elevation was important in predicting the distribution of silver-haired bats (Lasionycteris noctivagans) and big brown bats (Eptesicus fuscus). Increasing elevation by 400 m decreased the odds of capturing a silver-haired bat by 0.1 and a big brown bat by 0.4. Classification accuracy for our models ranged from 80.9% for all bat species combined to 72.3% for Yuma myotis and silver-haired bats. Predictive occurrence models can be valuable to bat conservation efforts because they provide spatial data important for evaluating the effects of management activities on species distributions.     

Diets of Frugivorous Bats in Montane Rain Forest and Coffee Plantations in Southeastern Chiapas,Mexico

Frugivorous are one of the main diaspore dispersers in tropical ecosystems, particularly in open areas and sites in the early stages of ecological succession. Frugivorous bat species respond differentially to habitat modification, and in the context of their diaspore dispersal functions it is important to understand species' ecological requirements. We compared the diversity of diaspores, obtained from fecal samples and from fruits carried by frugivorous bats, among five shaded coffee plantations under different management regimes and a montane rain forest in southeastern Chiapas, Mexico. At each site, bats were captured every 2 mo from March 2004 to July 2005, using six mist‐nets, during two consecutive nights. We captured 2589 individuals from 18 frugivorous species, from which we collected 969 fecal samples, containing 42 diaspore species associated with early and late successional plants. Although, we captured more frugivorous bat species in montane rain forest, the number of diaspore species in this site (N=14) was not significantly different from the coffee plantations under different management regimes (16–24). In montane rain forest, Sturnira ludovici fed mainly on Piper auritum, but in coffee plantations ate Peperomia sp., Saurauia madrensis, Solanum chrysotrichum and Solanum diphyllum. Artibeus jamaicensis and Artibeus intermedius feed mostly Cecropia obtusifolia and Ficus cookii in all coffee plantations. We suggest that the presence of frugivorous bats in shaded coffee plantations is favored by trees and shrubs associated with secondary and introduced vegetation that farmers have allowed to grow within or around the plantations.     

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.     

Comparison of frugivory by howling monkeys (Alouatta palliata) and bats (Artibeus jamaicensis) in the tropical rain forest of Los Tuxtlas,Mexico

Information on the fruit diets of howling monkeys and fruit-eating bats in the tropical rain forest of Los Tuxtlas, Mexico was collected for a year to compare the plant species used. Howling monkeys used 19 plant species whereas bats used 32 plant species as fruit sources. Eleven species were common in the diet of both mammals. A rank analysis at the plant species level showed that the fruit diets of Alouatta and Artibeus were very different. In contrast to bats, howling monkeys displayed a seasonal pattern in fruit consumption. Diet overlap between the two mammals was highest during the monkeys' fruit-eating season. Measures of fruit production in eight trees (four species) indicated marked variations in fruit biomass produced and in length of fruiting from tree to tree and species to species. Peaks in fruit production were typical both at the species and the individual tree levels, demonstrating the very patchy nature of the fruit available to the monkeys and the bats.     

Nectarivory by Endemic Malagasy Fruit Bats During the Dry Season1

Madagascar has a distinctive fruit bat community consisting of Pteropus rufus, Eidolon dupreanum, and Rousettus madagascariensis. In this study, we observed fruit bat visits to flowering baobabs (Adansonia suarezensis and Adansonia grandidieri) and kapok trees (Ceiba pentandra) during the austral winter. Eidolon dupreanum was recorded feeding on the nectar of baobabs and kapok, P. rufus was observed feeding on kapok only and no R. madagascariensis were seen. Three mammals species, two small lemurs (Phaner furcifer and Mirza coquereli) and E. dupreanum, made nondestructive visits to flowering A. grandidieri and are therefore all potential pollinators of this endangered baobab. This is the first evidence to show that A. grandidieri is bat‐pollinated and further demonstrates the close link between fruit bats and some of Madagascar's endemic plants. Eidolon dupreanum was the only mammal species recorded visiting A. suarezensis and visits peaked at the reported times of maximum nectar concentration. Pteropus rufus visited kapok mostly before midnight when most nectar was available, but E. dupreanum visited later in the night. These differences in timing of foraging on kapok can be explained either by differing distances from the roost sites of each species or by resource partitioning. We advocate increased levels of protection, education awareness, and applied research on both mammal‐pollinated baobab species and fruit bats, and suggest that both baobabs and bats are candidate “flagship species” for the threatened dry forests of Madagascar.     

Scale‐dependent responses to forest cover displayed by frugivore bats

Despite vast evidence of species turnover displayed by Neotropical bat communities in response to forest fragmentation, the exact shape of the relationship between fragment area and abundance for individual bat species is still unclear. Bats’ ample variation in diet, morphology, and movement behaviour can potentially influence species’ perception of the landscape. Thus, studies describing fragment area at a single spatial scale may fail to capture the amount of forest available from the perspective of individual bat species. In the present paper, we study the influence of forest cover on bats inhabiting a fragmented forest in Mexico, focusing on some of the most common frugivore species: Artibeus jamaicensis, Carollia spp. (C. brevicauda/C. perspicillata) and Sturnira spp. (S. lilium/S. ludovici). We quantified forest cover at scales ranging from 50 to 2000 m, and measured the influence of forest cover on bat capture success, a surrogate for abundance. The three species displayed positive and significant scale‐dependent associations with forest cover. Abundance of A. jamaicensis increased with forest cover measured at scales ranging between 500 and 2000 m, while Carollia spp. responded more strongly to variation in forest cover measured at scales 100–500 m. For Sturnira spp., abundance was a function of presence of creeks near mist‐netting sites, and amount of secondary forest present at a 200 m scale. The observed variation in responses to forest cover can be explained in light of interspecific differences in diet, home range, and body size. Our results illustrate a method for measuring the effect of forest fragmentation on mobile species and suggest that changes in abundance in fragmented landscapes emerge from the interaction between species’ traits and landscape structure.     

Morphology,Diet and Flower‐visiting by Phyllostomid Bats in Cuba

Pollinator morphology can play an important role in structuring plant–pollinator relationships and a pollinator's morphology may be associated with aspects of its diet. We examined the relationship between morphology and the partitioning of flower‐based food resources for five species of flower‐visiting Cuban bats: Artibeus jamaicensis, Brachyphylla nana, Erophylla sezekorni, Monophyllus redmani and Phyllonycteris poeyi. We analyzed cranial traits and body size to assess differences among species with respect to morphological specializations. We also collected dietary data from guano and used acoustic monitoring to assess bat activity at flowers. We found evidence that bats partition floral resources, but we found no direct evidence that plants were limiting resources for the bats. Morphological similarity among bat species did not predict dietary overlap. Rather it was associated with phylogenic relationships among some species. Species with different morphological specialization for flower‐visiting consumed resources and visited food plants at different frequencies.     

Trophic structure of a neotropical frugivore community: is there competition between birds and bats?

Summary Dietary overlap and competition between frugivorous birds and bats in the Neotropics have been presumed to be low, but comparative data have been lacking. We determined the diets of volant frugivores in an early successional patch of Costa Rican wet forest over a one month period. Ordination of the diet matrix by Reciprocal Averaging revealed that birds and bats tend to feed on different sets of fruits and that diets differed more among bat species than among bird species. However, there was overlap between Scarlet-rumped Tanagers and three Carollia bat species on fruits of several Piper species which comprised most of the diet of these bats. Day/night exclosure experiments on P. friedrichsthalli treetlets provided evidence that birds deplete the amount of ripe fruit available to bats. These results indicate that distantly related taxa may overlap in diet and compete for fruit, despite the apparent adaptation of animal-dispersed plant species for dispersal by particular animal taxa.     

Columnar cacti as sources of energy and protein for frugivorous bats in a semi‐arid ecosystem

Columnar cacti constitute the dominant elements in the vegetation structure of arid and semi‐arid New World ecosystems representing a plethora of food resources for vertebrate consumers. Previous stable isotope analysis in Central Mexico showed that columnar cacti are of low importance to build tissue for frugivorous bats. We used carbon stable isotope analysis of whole blood and breath samples collected from four species of frugivorous bats (Sturnira parvidens, Sturnira ludovici, Artibeus jamaicensis, and Artibeus intermedius) to reconstruct the importance of cactus plants in their diet. Breath samples were collected within 10 min (B₁₀) of bat capture and ~12 h after capture (B₇₂₀), representing the oxidation of recently ingested food and of body reserves, respectively. We expected that bats relied primarily on non‐cactus food to construct tissues and fuel oxidative metabolism. Non‐cactus food strongly predominated for tissue building, whereas oxidative metabolism was supported by a moderate preponderance of non‐cactus food for B₁₀ samples, and a moderate preponderance of cactus food or an equal contribution of both sources for B₇₂₀ samples. Artibeus and Surnira species appear to cover a narrow part of the diet with cactus food, confirming that the incorporation of nutrients derived from these plants is not generalized among vertebrate consumers.