Ecosystem services are the benefits obtained from the environment that increase human well-being. Economic valuation is conducted by measuring the human welfare gains or losses that result from changes in the provision of ecosystem services. Bats have long been postulated to play important roles in arthropod suppression, seed dispersal, and pollination; however, only recently have these ecosystem services begun to be thoroughly evaluated. Here, we review the available literature on the ecological and economic impact of ecosystem services provided by bats. We describe dietary preferences, foraging behaviors, adaptations, and phylogenetic histories of insectivorous, frugivorous, and nectarivorous bats worldwide in the context of their respective ecosystem services. For each trophic ensemble, we discuss the consequences of these ecological interactions on both natural and agricultural systems. Throughout this review, we highlight the research needed to fully determine the ecosystem services in question. Finally, we provide a comprehensive overview of economic valuation of ecosystem services. Unfortunately, few studies estimating the economic value of ecosystem services provided by bats have been conducted to date; however, we outline a framework that could be used in future studies to more fully address this question. Consumptive goods provided by bats, such as food and guano, are often exchanged in markets where the market price indicates an economic value. Nonmarket valuation methods can be used to estimate the economic value of nonconsumptive services, including inputs to agricultural production and recreational activities. Information on the ecological and economic value of ecosystem services provided by bats can be used to inform decisions regarding where and when to protect or restore bat populations and associated habitats, as well as to improve public perception of bats.

Three bat communities were studied for 1 year at each of two localities in the Panama Canal Zone and one locality in western Costa Rica. Removal sampling and banding techniques using Japanese mist nets were employed to document community structure, food habits. reproductive cycles, and movement patterns of these bats.Results indicated that 27-31 species occur at or near ground level at each locality. Species diversity, as measured by H', was highest in the Costan Rican community; each community contained 3-4 common species and many uncommon species. Based on body size and general food habits, niche overlap appears to be greatest among small to intermediate-sized insectivores and frugivores, many of which, however, are apparently uncommon. Four basic reproductive patterns are found among the species. Most frugivores are seasonally polyestrous whereas some insectivores are monestrous and at least one is polyestrous. It is postulated that in both insectivores and frugivores birth peaks coincide with maximum food levels. Recapture patterns of several species suggest that home range size may be positively correlated with body size; omnivorous species may have larger home ranges than similarly sized species with more restricted diets.

JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact support@jstor.org.. Ecological Society of America is collaborating with JSTOR to digitize, preserve and extend access to Ecology.Abstract. Many tropical bats eat fruit, nectar, and pollen, and many tropical plants are pollinated or dispersed by bats. These groups make up a coevolved system that illustrates the effects plants can have on animal populations, and conversely, the effects animals can have on plant populations. This paper reports a 2-yr intensive study of one bat-plant community in a seasonal Costa Rican forest.Seven common phyllostomatid bat species visited flowers or fruits. These were Glossophaga soricina, Phyllostomus discolor, Artibeus jamaicensis, Artibeus lituratus, Artibeus phaeotis, Sturnira lilium, and Carollia perspicillata. The incidence of nectar ivory was high among these species, even among the species previously considered to be frugivores. Floral resources were seasonally abundant and competition for these resources appeared minimal during the period of peak abundance, but only G. soricina was nectarivorous during the period of low flower abundance. Fruit resources were more evenly available through the year and were more finely divided among the species of bats. There was a correlation between size of bat and the average size of fruits eaten. To the extent that competition for food has influenced the structure of this bat community, competition for fruits was more important than competition for nectar and pollen in determining species diversity. Resource distributions through space and time also influenced bat foraging and reproductive behaviors. Small species of bats feed on resources of high abundance, whereas large species utilize resources that are patchy in time and space. The latter species probably forage in groups to maximize foraging efficiency. Reproductive behavior is seasonal and correlated with resource abundance.The flowering periods of plant species tend to be displaced, suggesting competition among plants for bat-pollinator services. A combination of the timing of plant-reproductive behavior and bat foraging characteristics promotes long-distance outcrossing and seed dispersal. These are critical factors in determining the low-density populations characteristic of many tropical plant species. E. RAYMOND HEITHAUS ET AL.Ecology, Vol. 56, No. 4 tion to plants that are potentially competing for agents of seed dispersal or services of pollinators affects reproductive success and, ultimately, population structure. Nightly foraging patterns can affect the probability of cross-pollination among plants or the dispersal distance of seeds. These factors partially determine the spatial distributions and effective breeding size of plant populations. On the other hand,...

Discussion of successional change has traditionally focused on plants. The role of animals in producing and responding to successional change has received far less attention. Dispersal of plant propagules by animals is a fundamental part of successional change in the tropics. Here we review the role played by frugivorous bats in successional change in tropical forests. We explore the similarities and differences of this ecological service provided by New and Old World seed-dispersing bats and conclude with a discussion of their current economic and conservation implications. Our review suggests that frugivorous New World phyllostomid bats play a more important role in early plant succession than their Old World pteropodid counterparts. We propose that phyllostomid bats have shared a long evolutionary history with small-seeded early successional shrubs and treelets while pteropodid bats are principally dispersers of the seeds of later successional canopy fruits. When species of figs (Ficus) are involved in the early stages of primary succession (e.g. in the river meander system in Amazonia and on Krakatau, Indonesia), both groups of bats are important contributors of propagules. Because they disperse and sometimes pollinate canopy trees, pteropodid bats have a considerable impact on the economic value of Old World tropical forests; phyllostomid bats appear to make a more modest direct contribution to the economic value of New World tropical forests. Nonetheless, because they critically influence forest regeneration, phyllostomid bats make an important indirect contribution to the economic value of these forests. Overall, fruit-eating bats play important roles in forest regeneration throughout the tropics, making their conservation highly desirable.

Aim We review several aspects of the structure of regional and local assemblages of nectar-feeding birds and bats and their relationships with food plants to determine the extent to which evolutionary convergence has or has not occurred in the New and Old World tropics.Location Our review is pantropical in extent and also includes the subtropics of South Africa and eastern Australia. Within the tropics, it deals mostly with lowland forest habitats.Methods An extensive literature review was conducted to compile data bases on the regional and local species richness of nectar-feeding birds and bats, pollinator sizes, morphology, and diets. Coefficients of variation (CVs) were used to quantify the morphospace occupied by the various families of pollinators. The extent to which plants have become evolutionarily specialized for vertebrate pollination was explored using several criteria: number and diversity of growth forms of plant families providing food for all the considered pollinator families; the most common flower morphologies visited by all the considered pollinator families; and the number of plant families that contain genera with both bird-and bat-specialized species.Results Vertebrate pollinator assemblages in the New World tropics differ from those in the Old World in terms of their greater species richness, the greater morphological diversity of their most specialized taxa, and the greater degree of taxonomic and ecological diversity and morphological specialization of their food plants. Within the Old World tropics, Africa contains more specialized nectarfeeding birds than Asia and Australasia; Old World nectar-feeding bats are everywhere less specialized than their New World counterparts.Main conclusions We propose that two factors -phylogenetic history and spatio-temporal predictability (STP) of flower resources -largely account for hemispheric and regional differences in the structure of vertebrate pollinator assemblages. Greater resource diversity and resource STP in the New World have favoured the radiation of small, hovering nectar-feeding birds and bats into a variety of relatively specialized feeding niches. In contrast, reduced resource diversity and STP in aseasonal parts of Asia as well as in Australasia have favoured the evolution of larger, non-hovering birds and bats with relatively generalized feeding niches. Tropical Africa more closely resembles the Neotropics than Southeast Asia and Australasia in terms of resource STP and in the niche structure of its nectar-feeding birds but not its flower-visiting bats.

Abstract. I discuss the evolution of feeding strategies in frugivorous bats by addressing two basic questions: (1) To what extent is diet choice opportunistic or selective and (2) What ecological factors influence diet choice and foraging behavior? I review the dietary diversity and feeding selectivity of bats of several common genera of phyllostomid bats and conclude that the evolution of food choice has involved: (1) specialization on a core plant taxon whose fruit are available throughout the year and (2) the opportunistic addition of other fruits to this core diet. Specialization on a core diet involves a tradeoff between fruit quality and fruit quantity. Bats eating high quality fruit (e.g. Piper and Solanum) tend to have broader diets then those eating low quality but high density fruits (e.g. Ficus). Specialization on different plant taxa as core dietary items has influenced the geographic (elevational) distributions, local abundance, foraging and food handling behavior, and feeding anatomy of frugivorous bats.

We report a new obligate pollination mutualism involving the senita cactus, Lophocereus schottii (Cactaceae, Pachyceereae), and the senita moth, Upiga virescens (Pyralidae, Glaphyriinae) in the Sonoran Desert and discuss the evolution of specialized pollination mutualisms. L. schottii is a night-blooming, self-incompatible columnar cactus. Beginning at sunset, its¯owers are visited by U. virescens females, which collect pollen on specialized abdominal scales, actively deposit pollen on¯ower stigmas, and oviposit a single egg on a¯ower petal. Larvae spend 6 days eating ovules before exiting the fruit and pupating in a cactus branch. Hand-pollination and pollinator exclusion experiments at our study site near Bahia Kino, Sonora, Mexico, revealed that fruit set in L. schottii is likely to be resource limited. About 50% of hand-outcrossed and open-pollinated senita¯owers abort by day 6 after¯ower opening. Results of exclusion experiments indicated that senita moths accounted for 75% of open-pollinated fruit set in 1995 with two species of halictid bees accounting for the remaining fruit set. In 1996,¯owers usually closed before sunrise, and senita moths accounted for at least 90% of open-pollinated fruit set. The net outcome of the senita/senita moth interaction is mutualistic, with senita larvae destroying about 30% of the seeds resulting from pollination by senita moths. Comparison of the senita system with the yucca/yucca moth mutualism reveals many similarities, including reduced nectar production, active pollination, and limited seed destruction. The independent evolution of many of the same features in the two systems suggests that a common pathway exists for the evolution of these highly specialized pollination mutualisms. Nocturnal¯ower opening, self-incompatible breeding systems, and resource-limited fruit production appear to be important during this evolution.

We studied variation in flowering phenology, fruit and seed set, and the abundance of the pollinators of four species of night-blooming Sonoran Desert columnar cacti for up to eight years at one site in Mexico and one year at one site in Arizona. We determined how spatiotemporal variation in plant-pollinator interactions affects the evolution of generalized pollination systems. We conducted pollinator exclusion and hand pollination experiments to document annual variability in pollinator reliability and to determine whether pollination systems were redundant (different species are partially or totally substitutable) or complementary (different species have an additive effect on fruit set). The cacti we studied included three species with generalized pollination systems involving bats, birds, and bees (cardon, Pachycereus pringlei; saguaro, Carnegiea gigantea; and organ pipe, Stenocereus thurberi) and one specialized moth-pollinated species (senita, Lophocereus schottii). We predicted that the migratory lesser long-nosed bat, Leptonycteris curasoae, is a less reliable pollinator than birds and bees, and that cacti with generalized pollination systems have more variable flowering phenologies than the specialized species.Annual time of peak flowering and mean size of flower crops were relatively invariant in saguaro and organ pipe. Time of peak flowering in cardon varied by as much as six weeks, and mean flower crop size varied threefold over six years. In senita, peak flowering varied by as much as 5-8 wk among years. Peak numbers of the nectar bat L. curasoae varied among years, and bat density (0.9/ha) was an order of magnitude lower than that of cactus-visiting birds at both study sites. The abundance of migratory hummingbirds was also highly variable among years.Pollinator exclusion experiments indicated that bats were major pollinators of cardon, whereas diurnal visitors accounted for most fruit set in saguaro (except in 1995 when bats were most important) and organ pipe at our Mexican site; honeybees accounted for 64-87% of diurnal fruit set in these species. Annual variation in the contribution to fruit set by bats was substantially higher than that of diurnal pollinators in saguaro and organ pipe, but not in cardon. There was little geographic variation in the relative importance of nocturnal vs. diurnal pollinators in saguaro and senita, but bats were much more important for fruit set in organ pipe in Arizona than in Mexico. We generally detected no effect of different pollinators on number of seeds per fruit in any species.Annual variation in fruit set was lowest in saguaro, the species with the most diurnal pollination system, and highest in organ pipe, the species with the most generalized pollination system. Fruit set was strongly pollen limited only in females of cardon (a trioecious species) and in organ pipe (at both sites). The ''missing'' pollinators in both species are likely Leptonycteris bats. The pollination systems of saguaro and cardon were partially redundant, whereas that of organ pipe was...