Nutrition or detoxification: why bats visit mineral licks of the Amazonian rainforest

Many animals in the tropics of Africa, Asia and South America regularly visit so-called salt or mineral licks to consume clay or drink clay-saturated water. Whether this behavior is used to supplement diets with locally limited nutrients or to buffer the effects of toxic secondary plant compounds remains unclear. In the Amazonian rainforest, pregnant and lactating bats are frequently observed and captured at mineral licks. We measured the nitrogen isotope ratio in wing tissue of omnivorous short-tailed fruit bats, Carollia perspicillata, and in an obligate fruit-eating bat, Artibeus obscurus, captured at mineral licks and at control sites in the rainforest. Carollia perspicillata with a plant-dominated diet were more often captured at mineral licks than individuals with an insect-dominated diet, although insects were more mineral depleted than fruits. In contrast, nitrogen isotope ratios of A. obscurus did not differ between individuals captured at mineral lick versus control sites. We conclude that pregnant and lactating fruit-eating bats do not visit mineral licks principally for minerals, but instead to buffer the effects of secondary plant compounds that they ingest in large quantities during periods of high energy demand. These findings have potential implications for the role of mineral licks for mammals in general, including humans.     

Parrots Take it with a Grain of Salt: Available Sodium Content May Drive Collpa (Clay Lick) Selection in Southeastern Peru

Soils from 18 parrot collpas ('clay licks') in southeastern Peru averaged four times more available sodium than uneaten control soils. Collpa soils contained marginally more clay than control sites and clay content was uncorrelated with available sodium content. Parrots may select and ingest soils based on available sodium content.     

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.     

Geophagy and nutrient supplementation in the Ngorongoro Conservation Area, Tanzania, with particular reference to selenium, cobalt and molybdenum

Wildlife and livestock in the Ngorongoro Conservation Area (NCA), Tanzania, commonly practise geophagy. We investigated the nutrient content of nine earth licks and adjacent topsoils in the NCA, and for comparative purposes, two salt pans in north-eastern Tanzania, licks in central Tanzania and a lick near Puerto Maldonado, Peru. The licks had no consistent pattern of nutrient enrichment relative to unutilized topsoils, although the Na content of licks was greater than that of the adjacent topsoils at all but one site. The three largest licks in the NCA (Gibbs Farm, Seneto and Ascent Road) were enriched relative to topsoils and global averages in Se (maximum of 4.7 mg kg⁻¹), Co (maximum of 107 mg kg⁻¹) and/or Mo (maximum of 7.4 mg kg⁻¹). We suggest that licks do provide supplemental Na, but that Se, Co and/or Mo at the largest licks provide easily overlooked nutritional benefits and are perhaps the primary target for geophagy at these sites. Subsoil zones of clay deposition are likely to be enriched in various elements through illuviation or mineral precipitation from solution. Animals may use the taste of NaCl as a clue for locating such zones where they are likely to find a greater quantity of micronutrients relative to other soils. These findings have consequences for conservation and pastoralism in that these large licks may be key resources, providing micronutrients that are essential for maintaining the health and fecundity of animal as well as human populations in the region.     

Movement Patterns and Home Range Of The Bongo (Tragelaphus Eurycerus) In The Rain Forest Of The Dzanga National Park, Central African Republic

Bongos ( Tragelaphus eurycerus Ogilby) were studied for 8 months in the Dzanga National Park, Central African Republic. Tracks were followed and mapped with a compass and a pedometer to study movement patterns and home range. Natural licks were shown to be central points in the home range of the bongos: they visited the licks recurrently to consume soil, but also to forage on grass and herb species, and for social reasons. Forest areas far from licks were used much less than forest areas close to licks. When a lick was visited, distances between two resting sites were longer than in the forest without lick visits, caused by a direct and straight movement from the denser forest areas toward a lick. The study area of about 150 km² was presumably occupied by two groups of bongos. One of them seemed to split temporarily into two subgroups. Groups were not larger than 10–20 individuals. Estimated home ranges were at least 49 km² and 19 km² for the two groups, respectively. Estimated density in the Dzanga National Park was 0.25 bongos per km^2. This study shows the importance of natural licks for the largest social forest antelope, the bongo, and provides information which is important for its future conservation.     

Patterns of mineral lick visitation by spider monkeys and howler monkeys in Amazonia: are licks perceived as risky areas?

Mineral licks--also known as "salados," "saladeros," or "collpas"--are specific sites in tropical and temperate ecosystems where a large diversity of mammals and birds come regularly to feed on soil. Although the reasons for vertebrate geophagy are not completely understood, animals are argued to obtain a variety of nutritional and health benefits from the ingestion of soil at mineral licks. We studied the temporal patterns of mineral lick use by white-bellied spider monkey (Ateles belzebuth) and red howler monkey (Alouatta seniculus) in a lowland rain forest in Amazonian Ecuador. Using camera and video traps at four different mineral licks, combined with behavioral follows of one group of spider monkeys, we documented rates of mineral lick visitation by both primate species and the relative frequency and intensity of mineral lick use by spider monkeys. On the basis of 1,612 days and 888 nights of mineral lick monitoring, we found that A. belzebuth and A. seniculus both visit mineral licks frequently throughout the year (on average ～14% of days for both species), and mineral lick visitation was influenced by short-term environmental conditions (e.g. sunny and dry weather). For spider monkeys, the area surrounding the lick was also the most frequently and most intensively used region within the group's home range. The fact that spider monkeys spent long periods at the lick area before coming to the ground to obtain soil, and the fact that both species visited the lick preferentially during dry sunny conditions (when predator detectability is presumed to be relatively high) and visited simultaneously more often than expected by chance, together suggest that licks are indeed perceived as risky areas by these primates. We suggest that howler and spider monkeys employ behavioral strategies aimed at minimizing the probability of predation while visiting the forest floor at risky mineral lick sites.     

Scale‐dependent effects of forest restoration on Neotropical fruit bats

Neotropical fruit bats (family Phyllostomidae) facilitate forest regeneration on degraded lands by dispersing shrub and tree seeds. Accordingly, if fruit bats can be attracted to restoration sites, seed dispersal could be enhanced. We surveyed bat communities at 10 sites in southern Costa Rica to evaluate whether restoration treatments attracted more fruit bats if trees were planted on degraded farmlands in plantations or island configurations versus natural regeneration. We also compared the relative influence of tree cover at local and landscape spatial scales on bat abundances. We captured 68% more fruit bat individuals in tree plantations as in controls, whereas tree island plots were intermediate. Bat activity also responded to landscape tree cover within a 200‐m radius of restoration plots, with greater abundance but lower species richness in deforested landscapes. Fruit bat captures in controls and tree island plots declined with increasing landscape tree cover, but captures in plantations were relatively constant. Individual species responded differentially to tree cover measured at different spatial scales. We attribute restoration effects primarily to habitat structure rather than food resources because no planted trees produced fruits regularly eaten by bats. The magnitude of tree planting effects on fruit bats was less than previous studies have found for frugivorous birds, suggesting that bats may play a particularly important role in dispersing seeds in heavily deforested and naturally regenerating areas. Nonetheless, our results show that larger tree plantations in more intact landscapes are more likely to attract diverse fruit bats, potentially enhancing seed dispersal.     

Reduced-impact Logging has Little Effect on Temporal Activity of Frugivorous Bats (Chiroptera) in Lowland Amazonia

Reduced-impact logging (RIL) represents a viable option for sustainable use of Neotropical lowland forests while minimizing negative effects on local biodiversity. Many Neotropical bats of the family Phyllostomidae provide ecosystem services associated with pollination and seed dispersal that promote the regeneration of disturbed areas; therefore, effects of RIL on these species is of particular concern. We determined patterns of temporal activity, degree of temporal overlap of activity, and dispersion in peaks of activity for seven abundant species of frugivorous bat in Tapajós National Forest, Pará, Brazil. In addition, we evaluated the effects of RIL at a harvest level of 18.7 m³/ha and habitat physiognomy on temporal patterns of activity for these species. Bats were surveyed for four nights at each of 96 sites for a total sampling effort of 64,512 net-m-h. Sites were distributed among four experimental blocks, two blocks of unlogged forest and two blocks of forest subjected to RIL. Half of the sites in each management type were in forest gaps and half were in closed-canopy forest. In general, species exhibited similar patterns of activity, and greater than expected temporal overlap in activity among species. RIL and forest physiognomy had little effect on activity patterns of species. RIL in Amazonia removes fewer trees than do naturally occurring treefalls and such changes in habitat structure do not alter activity patterns of frugivorous bats. Evidence suggests that RIL does not have an appreciable adverse effect on frugivorous bats in Amazonia.     

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.     

Use of Fruit Essential Oils to Assist Forest Regeneration by Bats

Frugivorous bats can be attracted with essential oils from ripe chiropterochoric fruit. We evaluated the efficiency of these oils to attract bats in degraded areas within the Atlantic Rain Forest, particularly pasture and agricultural land. We hypothesized that induction units (IUs), each containing a rubber septum impregnated with oil, would have more bat activity than their respective control units (CUs; without the oil). To test this hypothesis we monitored bat flight activity with night‐vision infrared visors in eight IU and CU from August 2006 to July 2007. We also verified the probability of arrival of chiropterochoric seeds by analyzing the diet of bats captured in a neighboring forest area. Our initial hypothesis that units with odor would lead to greater bat activity was confirmed. Results indicated a rich community of fruit‐eating bats, and dietary analysis revealed a huge potential for dispersion of a vast amount of seeds from different plant species at the IU. Although our study does not reveal with certainty which bat species are attracted to the oil, the flying patterns coincide with those described for the foraging behavior of fruit‐eating phyllostomids. Furthermore, the fact that the bats spend more time flying around the odor source compared to flying time around CU suggest an increase in seed rain. Taken together, these results suggest that the use of essential oils from chiropterochoric fruits induces a qualitative and quantitative increase in seed dispersal in areas that otherwise would not be frequently visited by frugivorous bats.     

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.     

Feeding ecology of a large social antelope in the rainforest

We studied the feeding ecology of the bongo (Tragelaphus eurycerus) in the Dzanga National Park in the rainforest of the Central African Republic to understand better the trade-off between food selectivity, ranging behaviour and social organization of a large, forest-dwelling, social antelope. Food plants and vegetation types were registered along a 311-km route travelled by bongos. Food availability was determined by identifying and counting the plants in 19 randomly chosen forest plots. Bongos showed pronounced selectivity for 26 out of 100 woody forest species. They predominantly consumed younger leaves, which suggests that high protein and low fibre content influence plant choice. In addition to leaves, bongos also ate fruits of two and flowers of one species. Furthermore, the diet was supplemented by grasses and herbs consumed on large natural licks. Such licks were regularly visited by the bongos. According to Jarman's ecological classification of antelopes, selective browsers are relatively small and live alone or in pairs to avoid competition over food. The bongo's large size and gregariousness should not allow it to survive in the rainforest as a pure selective browser. Our data suggest that the bongo relies on the opportunity to graze in bulk which it finds on the natural licks. We hypothesize that such licks either limit the distribution of bongos in other rainforest areas or allow larger group sizes than in areas without licks. Received: 8 June 1998 / Accepted: 30 November 1998     

Drinking and Flying: Does Alcohol Consumption Affect the Flight and Echolocation Performance of Phyllostomid Bats?

Background

In the wild, frugivorous and nectarivorous bats often eat fermenting fruits and nectar, and thus may consume levels of ethanol that could induce inebriation. To understand if consumption of ethanol by bats alters their access to food and general survival requires examination of behavioural responses to its ingestion, as well as assessment of interspecific variation in those responses. We predicted that bats fed ethanol would show impaired flight and echolocation behaviour compared to bats fed control sugar water, and that there would be behavioural differences among species.

Methodology/Principal Findings

We fed wild caught Artibeus jamaicensis, A. lituratus, A. phaeotis, Carollia sowelli, Glossophaga soricina, and Sturnira lilium (Chiroptera, Phyllostomidae) sugar water (44 g of table sugar in 500 ml of water) or sugar water with ethanol before challenging them to fly through an obstacle course while we simultaneously recorded their echolocation calls. We used bat saliva, a non-invasive proxy, to measure blood ethanol concentrations ranging from 0 to >0.3% immediately before flight trials. Flight performance and echolocation behaviour were not significantly affected by consumption of ethanol, but species differed in their blood alcohol concentrations after consuming it.

Conclusions/Significance

The bats we studied display a tolerance for ethanol that could have ramifications for the adaptive radiation of frugivorous and nectarivorous bats by allowing them to use ephemeral food resources over a wide span of time. By sampling across phyllostomid genera, we show that patterns of apparent ethanol tolerance in New World bats are broad, and thus may have been an important early step in the evolution of frugivory and nectarivory in these animals.     

Temporal patterns of visitation of birds and mammals at mineral licks in the Peruvian Amazon

Mineral licks are key ecological resources for many species of birds and mammals in Amazonia, providing essential dietary nutrients and clays, yet little is known about which species visit and their behaviors at the mineral licks. Studying visitation and behavior at mineral licks can provide insight into the lives of otherwise secretive and elusive species. We assessed which species visited mineral licks, when they visited, and whether visits and the probability of recording groups at mineral licks were seasonal or related to the lunar cycle. We camera trapped at 52 mineral licks in the northeastern Peruvian Amazon and detected 20 mammal and 13 bird species over 6,255 camera nights. Generalized linear models assessed visitation patterns and records of groups in association with seasonality and the lunar cycle. We report nocturnal curassows (Nothocrax urumutum) visiting mineral licks for the first time. We found seasonal trends in visitation for the black agouti (Dasyprocta fuliginosa), red howler monkey (Alouatta seniculus), blue‐throated piping guan (Pipile cumanensis), red brocket deer (Mazama americana), collared peccary (Pecari tajacu), and tapir (Tapirus terrestris). Lunar trends in visitation occurred for the paca (Cuniculus paca), Brazilian porcupine (Coendou prehensilis), and red brocket deer. The probability of recording groups (>1 individual) at mineral licks was seasonal and related to lunar brightness for tapir. Overall, our results provide important context for how elusive species of birds and mammals interact with these key ecological resources on a landscape scale. The ecological importance of mineral licks for these species can provide context to seasonal changes in species occupancy and movement.

Many species of animals in the Amazon visit mineral licks, consuming soil to supplement their diet. Using camera trap data, we show that visits to mineral licks are seasonal for many species. These trends in visitation are likely due to species‐specific factors such as reproduction, predator avoidance, seasonal diet shifts, and resource use.     

The Role of Fruits and Insects in the Nutrition of Frugivorous Bats: Evaluating the Use of Stable Isotope Models1

We estimated the relative contribution of fruits and insects as sources of dietary protein in two species of Neotropical frugivorous bats (Artibeus jamaicensis and Sturnira lilium) using stable carbon and nitrogen isotope analyses. An insectivorous species (Pteronotus parnellii) was also included for comparison. We found constant patterns in stable carbon and nitrogen isotope composition in blood that separated the two species of frugivorous bats from the insectivorous bat. When we used these isotopic values (combined with those of dietary fruits and insects) to estimate the percent contribution of fruits and insects to the diet of the bats, we obtained different results, depending on assumptions and model adopted. We tested models using both 8“N and 8′³C results simultaneously and separately and further used diet‐tissue fractionation factors of 3%o for nitrogen and 1 and 3.5%o for carbon. We found that a carbon‐based model with a diet‐blood enrichment factor of 3.5%o produced the most parsimonious results. The model estimated that A. jamaicensis and S. lilium obtained most of their protein requirements from fruits, whereas P. parnellii fed mostly on insects. No sexual or seasonal variations in the diet of the two frugivorous species were detected. We found no evidence that the diet of sexually active females differed from that of nonsexually active females in the two species of frugivorous bats. We suggest that future studies better define isotopic fractionation between diet and tissues of bats using captive rearing and controlled diets.     

Asymmetric Contribution of Isotopically Contrasting Food Sources to Vertebrate Consumers in a Subtropical Semi‐arid Ecosystem

Habitat heterogeneity is a primary ecological factor that is particularly pronounced in arid ecosystems. The Tehuacán valley is a subtropical semi‐arid ecosystem in which several species of columnar cacti and agave (i.e., CAM plants) constitute the dominant elements accompanied by patches of trees and shrubs (i.e., C₃ plants). Vegetation in Tehuacán is isotopically heterogenous because CAM plants have less depleted δ¹³C values than C₃ plants. Fruits and flowers of cactus and agaves offer abundant food to vertebrates, but their leaves might be less attractive to insects than the leaves of C₃ plants. Therefore, we use carbon and nitrogen stable isotope analysis to test the hypothesis that C₃ and CAM food would contribute asymmetrically to different guilds of birds and bats. We predict that granivorous and frugivorous birds and nectarivorous and frugivorous bats will consume a CAM diet, whereas insectivorous birds and bats will consume a C₃ diet. Due to omnivory of bird and bat consumers, we predict that the importance of CAM food will decrease as the trophic level of the animal increases. Our results showed that CAM food predominated in plant‐eating birds and in some flower‐visiting bats, whereas C₃ food predominated in insect‐eating bats and birds and frugivorous bats. Habitat heterogeneity in Tehuacán is important for conservation due to the asymmetric role of CAM and C₃ food in the nutrition of different feeding guilds of vertebrates. Our study provides basic information to evaluate the potential impact of habitat loss on functional groups of consumers in a semi‐arid ecosystem.     

Parrot behavior at a Rio Manu (Peru) clay lick: temporal patterns,associations, and antipredator responses

Although eating clay at licks (a form of geophagy) has been described, there are few behavioral data on temporal patterns, social interactions, species associations, or reactions to potential predators. We examined the behavior of nine species of macaws, parrots, and parakeets at the Machiguenga Ccolpa, a clay lick on the Rio Manu, Peru in the dry season. Three distinct mixed-species groups used the licks: in the early morning (parrots and small macaws), in mid-morning (large macaws), and in the early afternoon (parakeets), although the latter two groups used the licks at other times of day as well. The first parrots to begin eating at the lick in the early morning were yellow-crowned parrots (Amazona ochrocephala) and dusky-headed parakeets (Aratinga weddellii), followed by blue-headed parrots Pionus sordidus, and then by mealy (Amazona farinosa) and orange-cheeked (Pionopsitta barrabandi) parrots, and chestnut-fronted macaws (Ara severa). Although blue-headed parrots fed in dense groups of over 50, the others rarely exceeded 20 individuals. Scarlet macaws (A. macao) sometimes fed alone or joined the early morning groups, but most associated with a large group of red and green macaws (A. chloroptera) that arrived, often scaring off the smaller birds. On average, about 100 macaws and parrots fed in the early morning, macaw feeding groups averaging just over 40, and the parakeets averaged over 70. Average time at the lick ranged from 28 min for yellow-crowned parrots to 47 min for tui parakeets. Of the early morning group, blue-headed and mealy parrots were the most aggressive and orange-cheeked parrots were the least aggressive. Red and green macaws were more aggressive than scarlet macaws; the parakeets were equally aggressive. All species had more aggressive interactions with conspecifics than with other species. Responses to intruders and predators varied by species of parrot/macaw and type of intruder. In response to intruders or loud calls, responses could be partial (some individuals flew away, circled, and returned), temporary (all individuals flew away but returned within a few minutes), or total (all flew away and abandoned feeding for at least a half hour). The large macaws showed the lowest rate of total abandonment and the parakeets showed the highest. People passing up or down river in boats scared birds from the lick. The local residents (Machiguenga tribespeople in boats) elicited a much greater response than did the researchers. In the recent past, macaws and parrots were hunted for food, feathers, and the pet trade, and the birds response, as well as the presence of parrot and macaw feathers in local villages we visited, suggests some continued exploitation, or a long-term memory in the birds.Communicated by R.F. Oliveira     

Frugivory and the effects of ingestion by bats on the seed germination of three pioneering plants

The dispersion and seedling establishment of pioneering plants can be favoured by the presence of frugivorous bats because the bats usually improve seed germination after ingestion. Although seed germinability is known to vary greatly after ingestion by different bats, the relative contribution of each bat species to seed germination within plant communities is poorly understood. In this study, we first determined the fauna of frugivorous bats in a semideciduous seasonal forest remnant in southern Brazil and subsequently identified the plant species of the seeds passed through their guts. Second, the germination performance (i.e., germination percentage and speed) of the seeds of three pioneering plants (Piper aduncum, Piper hispidinervum and Solanum granuloso-leprosum) ingested by the most abundant bats was compared with that of the non-ingested seeds (seeds collected from fruits). Additionally, the effects on seed germination of different bat species were compared. During one year, five species of frugivorous bats were caught, and the seeds of eleven identifiable plant species (not counting those of undetermined species) were found in their faeces. We found that the germination performance of the seeds of Piper species was significantly enhanced after ingestion by bats, whereas S. granuloso-leprosum seeds had neutral or reduced germinability when seeds in faeces were compared with pulp-removed seeds. Our results revealed that the bat species that were captured exerted different effects upon seed germination; such a disparity is expected to result in different rates of early establishment of these pioneer plants in tropical forests, most likely affecting forest composition and structure, particularly during the initial stages of succession.     

Sources of assimilated protein in five species of New World frugivorous bats

Fruits are N-poor items and their availability in the tropics varies throughout the year. Field and experimental studies debate whether frugivorous bats have to switch to N-rich sources of food during part of the year or if they are able to subsist on a fruit-only diet. Different strategies to meet their N requirements may influence the way in which frugivorous bats partition food resources allowing the coexistence of numerous species in tropical communities. We examined the extent to which five species of frugivorous bats relied on plant and insect sources of assimilated protein using stable-N isotope analysis. We assumed that bats only had access to fruits and insects in our analysis but we also collected fecal samples to examine the presence of other food items. We conducted the study during at least 1 full year depending on the species of bat in a tropical rain forest in southern Mexico. In the five species of bats examined, plant sources (i.e. fruits) provided most of the protein assimilated during the year, although there was a general trend for all species to show a decrease in relative plant contribution at the end of the rainy season and beginning of the dry season. In Artibeus jamaicensis, Uroderma bilobatum and Dermanura phaeotis, plants were still a major source of protein during this period, but in some individuals of Sturnira lilium and Carollia brevicauda insects represented an important contribution to their diet. Fecal samples of most bats presented fruit remains, and insects and pollen were found in small proportions. Bat reproductive activity was detected at the end of the dry season and in the middle of the rainy season, and plants were the major source of protein during this period with the exception of pregnant S. lilum and one pregnant D. phaeotis during the dry season. Our findings showed that frugivorous bats might differ in their strategies to satisfy their N demands with some species relying almost completely on fruits during most of the year and some species switching to insects when fruits were less abundant.