Worldwide, bats occupy unique ecological roles and provide valuable services to society. In addition to arthropod suppression, pollination and seed dispersal, bats contribute immensely to biodiversity (every fifth mammal’s a bat), technological advancements (sonar systems, biomedical ultrasound, autonomous aerial vehicles), cultural symbolism, and tourism (e.g., Carlsbad Caverns, Congress Avenue Bridge). In fact, bats fulfill every category of ‘ecosystem service’ – “defined as benefits to humankind derived from resources and processes supplied by natural ecosystems.” These categories include
This #BatWeek, we are celebrating bats – from A to Z. As you explore this interactive alphabet, you’ll discover the enormous contribution #batkind brings to our everyday lives and the world we live in. Worldwide, more than 1400 bat species enhance biodiversity and biomass, contribute to ecosystem functioning and disperse keystone resources important to wildlife communities. The alphabet highlights some of these extraordinary contributions.
So explore, learn, celebrate – or more fittingly, celeBrATe.
Rousettus aegyptiacus, Egyptian fruit bats, have an extensive geographic distribution, occurring throughout Africa and the Middle East. R. aegyptiacus, along with other species of Rousettus, are the only megachiropterids to employ echolocation, which they accomplish by emitting a series of sharp clicks with their tongues. They are frugivorous, and wild dates are a favorite, but they consume almost any soft, pulpy fruit.
DYK Chromatic disorders causing causing abnormal coloration, either from complete absence (albinism), partial deficiency (leucism) or excess (melanism) of melanin, are relatively rare within the order Chiroptera. Individuals with leucism - like this Rousettus aegyptiacus - display atypical white spots on the skin and/or pure white fur and normal colored eyes.
The genus Manilkara, of family Sapotaceae, includes approximately 80 species, three of particular importance - M. bidentata, balatá; M. chicle, chicle; and M. zapota, sapodilla. Pteropodids, including Pteropus giganetus, Indian flying fox, and phyllostomids, including Artibeus jamaicensis, common fruit bats and Stenoderma rufum, red fruit bats, contribute to pollination and seed dispersal. These three Manilkara species produce chicle - natural chewing gum.
Only a handful of companies use chicle to produce chewing gum including Glee Gum ™, Tree Hugger ®, and Simply Gum ™.
Through insect suppression and chirosurveillance, Eptesicus fuscus, big brown bats, provide valuable ecosystem services to the United States' agricultural industry.
E. fuscus consume numerous herbivorous arthropods, including Halyomorpha halys, brown marmorated stink bugs, a pentatomid native to East Asia. H. halys are capable of consuming more than 100 different species including apples, apricots, Asian pears, cherries, corn, grapes, lima beans, peaches, peppers, tomatoes, and soybeans.
Now common throughout eastern North America, H. halys nymphs and adults can decimate fruit orchards; in 2010, this invasive species cost mid Atlantic apple producers $37 million USD.
Furthermore, E. fuscus seasonally consume H. halys, detecting the insects 3-4 weeks earlier than existing monitoring tools. Chirosurveillance, "the use of native bats to detect invasive agricultural pests," may become a standard strategy for integrated pest management.
Leptonycteris species - Mexican long-nosed bats - pollinate several species of Agave, including Agave tequiliana, a source of agave syrup, a commercially available sweetener and substitute for sugar or honey.
Worldwide, insectivorous bats commonly consume deleterious arthropods such as Drosophila suzukii, spotted wing drosophila; Recurvaria leucatella, lesser budmoths; and Halyomorpha halys, brown marmorated stink bugs, which consume summer fruits including cherries, blueberries, raspberries, peaches, nectarines and apricots.
In Mediterranean agroecosystems, Rhinolophus hipposideros, lesser horseshoe bats, consume herbivorous arthropods that cause significant agricultural damage including Recurvaria nanella, lesser budmoths; and Hedya nubiferana, green budworm moths, which damage Prunus dulcis - almond trees. The edible nutlike seeds of these deciduous trees provide a nondairy alternative to traditional milk.
The botanical family Musaceae comprises approximately 91 species including Musa acuminata and M. balbisiana - the ancestors of modern, edible bananas.
Both opportunistic and specialized pteropodids pollinate Musa species; the 18-26 gram megabat, Macroglossus sobrinus, or long-tongued fruit bat, consumes nectar almost exclusively from these extraordinarily beautiful flowers.
In Yunnan, China, M. sabrinus pollinates M. itinerans - the Yunnan banana. This endangered banana species provides an important foodstuff for the endangered Asian elephant, the largest extant land mammal inhabiting Asia.
Beer, the third most popular beverage after water and tea, derives from brewing cereal grains - most commonly barley, wheat, maize, and rice.
Worldwide, molossids, particularly Tadarida brasiliensis, Mexican free-tailed bats, provide valuable ecosystem services by consuming arthropods, including those damaging staple crops such as maize, rice, wheat and barley.
In Australia and Indonesia, Ozimops planiceps, southern free-tailed bats, forage extensively on Nysius vinitor, rutherglen bugs, a serious agricultural pest of oil seeds including sunflower, canola, safflower and sorghum. Grain sorghum, an important cereal crop, provides a gluten-free grain for breads, cereals and beer. In fact, sorghum's the main ingredient in Anheuser Busch's 'Redbridge Gluten Free' beer.
Worldwide, insectivorous bats suppress populations of insects, including those damaging agricultural crops.
In North America, Myotis sodalis, Indiana bats, consume Mayetiola destructor, barley midges, a significant pest of cereal crops - the primary ingredient of bread.
In southwestern Europe, Rhinolophus spp., horseshoe bats, consume multiple insects including Mythimna unipuncta, true armyworm moths, which damage various Graminae, barley, corn, oats, rice, rye, sorghum and wheat.
Pteropodids, including Rousettus aegyptiacus, Egyptian fruit bats and Eidolon helvum, straw-coloured fruit bats, pollinate Adansonia digitata, the baobab, which possesses globose flower buds extending from long flower stalks. "The flowers of all other species of baobab grow on much shorter stalks, protruding either erectly or horizontally off the branches. Such orientation is ill-suited to bat pollination and the blooms rely on birds and insects to spread their pollen." Indigenous to the West African savannahs, the baobab blooms March to May, and at night, "fruit bats visit the trees for the soft floral parts and presumably feed on the stamens and nectar as well.”
'Africa's upside-down tree' has a "rather sharp and unpleasant" fragrance, resulting from the "substantial proportion of sulphur compounds in its floral scent."
Numerous phyllostomids pollinate Ochroma pyramidale - the balsa tree. “The big cuplike flowers are visited by Phyllostomus discolor, pale spear-nosed bats; P. hastatus, greater spear-nosed bats; a number of stenodermatine bats; Carollia perspicullata, Seba's short-tailed bats; and also by marsupials like Caluromys and monkeys. "While smaller stenodermatine bats “jump” into the Ochroma flowers with a head-first dive from the air, larger bats like P. hastatus and P. discolor hold the petals with their hind legs and dip their heads into the nectar, becoming completely covered with pollen and nectar on the head and back."
Theobroma cacao, commonly cacao tree or cocoa tree, produces seeds indispensable to chocolate liquor, cocoa solids, cocoa butter and chocolate. The fruit, or cacao pod, contains approximately 20-60 "beans," which are the primary ingredient of chocolate. The generic name 'Theobroma' derives from Greek 'theos,' meaning "god," and 'broma,' meaning "food" - hence, "food of the gods."
Anacardium occidentale, a tropical evergreen native to Central America, the Caribbean Islands and northern South America, produces cashew seeds, or simply cashews, and cashew apples.
Seed dispersing pteropodids, including Eidolon helvum, straw-coloured fruit bats; Epomophorus gambianus, Gambian epauletted fruit bats; and Micropteropus pusillus, Peters' dwarf epauletted fruit bats, visit “when the fleshy, swollen, edible pedicel, known as the cashew apple, is ripe and juicy. They carry off part of the pedicel and drop the hard nut when they are eating on the fleshy part."
In North America, aerial insectivores including Myotis sodalis, Indiana bats; Tadarida brasiliensis, Mexican free-tailed bats; and Nycticeius humeralis, evening bats, consume massive quantities of insects. Several of these species damage cotton crops including Acrosternum hilare, green stinkbugs; Diabrotica undecimpunctata, spotted cucumber beetles; Nysius raphanus, false chinch bugs; and especially Spodoptera frugiperda, fall armyworms.
S. frugiperda' scientific name derives from Latin frugiperda meaning 'lost fruit,' an indication of this species' destructive ability. T. brasiliensis' insect suppression services save cotton producers approximately $12.2 - $70.1 million USD annually. This soft, fluffy staple fiber remains the most common natural fiber textile available today, with global production averaging 25 million tons annually.
In Mexican coffee plantations, insectivorous bats reduce arthropods by a grande 84% - almost 30% more than birds.
In Baduy, Banten Indonesia, the Baduy people cultivate Coffea canephora, coffee robusta, for subsistence cultivation. The Baduy people obtain coffee seedlings from gardens, because mature seeds are "spread out by small bats from parent trees," i.e., "The small bats pick mature coffee fruit at night, eating it, but some of the beans fall on the ground and finally germinate into seedlings..."
C. canephora represents approximately 40% of global coffee production, with beans imparting "a distinctive earthy flavour."
In southcentral Texas and northern Mexico, Tadarida brasiliensis, Mexican free-tailed bats,
Scientists value the suppression of herbivory on corn alone may be > 1 billion USD globally, and bats further benefit farmers by indirectly limiting pest-associated fungus and mycotoxins. Globally, bats bolster the quality and quantity of corn via a trophic cascade, suppressing damage to economically valuable row crops during both reproductive and vegetative stages.
In which heartwarming Disney movie does someone say, "What am I looking at here? Why are you hanging from the earth by your feet like bats?"
Phoenix dactylifera, commonly date or date palm, produces economically valuable fruits originating from the Fertile Crescent region between Egypt and Mesopotamia. Pteropodids including Pteropus giganteus, Indian flying fox, and Rousettus aegyptiacus, Egyptian fruit bats, are valuable dispersal agents of these deliciously sweet fruits.
Fun Fact | Countries of the Middle East and North Africa produce nearly 8.5 million metric tons of dates annually.
Nocturnal nectivores, including Leptonycteris curasoae, southern long-nosed bats, and Choeronycteris mexicana, Mexican long-tongued bats, pollinate the hemiepiphytic cactus Hylocereus undatus. The cactus' edible fruits - dragonfruit or pitahaya - are popular both regionally and nationally.
Pteropodids disperse the seeds of Dracontomelon vitiense, dragon plum, an indigenous Oceanian species of fruit tree. "The Melasnesians appreciate the acid taste of the small fruits, which are eaten raw as a snack or cooked in coconut milk."
The genus Durio comprises 30 species; however, only one - Durio zibethinus - enjoys commercial cultivation and international recognition. This species of durian produces chiropterophilous blossoms - large, nocturnally blooming, odoriferous floral components attractive to nectarivorous pteropodids, particularly Eonycteris spelaea, cave nectar bats.
Species such as durian "rely on a highly specialized pollination system that depends entirely on the nocturnal pteropodids that the plant has coevolved with over millions of years."
Despite durian's odor, "best described as pig-shit, turpentine and onions, garnished with a gym sock," the fruit enjoys considerable popularity, including annual celebrations such as Thailand's 'World Durian Festival' and Phillipines' Kadayawan Festival.
Pteropodids are incredibly important pollinators of multiple Parkia species including Parkia speciosa, petai or 'stink beans;' P. timoriana, tree beans; P. biglobosa, African locust bean; and P. clappertoniana, whose seeds provide dawadawa, a traditional ingredient throughout West Africa, particularly Ghana.
The chiropterophilous inflorescence of P. clappertoniana "hangs clear of the foliage on a flexible but stout peduncle. It produces a large reservoir of nectar at night in a position where it can be taken by large flying animals which, at the same time, receive or transfer pollen which adheres to the fur of their adpressed breasts."
Pteropus poliocephalis, gray-headed flying fox, consume several species of eucalypt, particularly Corymbia [Eucalyptus] gummifera, red bloodwood; E. muelleriana, yellow stringybark; E. globoidea, white stringybark; and E. botryoides, bangalay, while Pteropus medius, Indian flying fox, disperse the seeds of Eucalyptus citriodora, lemon eucalyptus.
The essential oil of E. citriodora consists predominately of citronellal, providing a distinctive lemony fragrance to perfumes and insect repellents.
Phyllanthus emblica, commonly emblic, Indian gooseberry or amla, are "arguably one of the most important plants" relative to Ayervedic medicine. These deciduous trees possess multiple therapeutic properties, including antiinflammatory, antioxidant, and antiaging benefits. Frugivorous animals, particularly bats and birds, disperse emblic fruits, which are "sour, bitter and astringent," and "possibly one of the richest natural sources of vitamin C."
In Chinese culture, bats are auspicious creatures representing good fortune, and this association has linguistic origins - the pronunciation of “bat” in Chinese (蝠), is “fú” — the same as that of “blessing” (福). For centuries, bats have been ubiquitous motifs within Chinese art; for example, 'famille rose' porcelains portray five red bats alongside graceful branches of fruiting peaches. The five bats represent five blessings - longevity, health, prosperity, virtue and natural death; the color red symbolizes abundance - red bats were especially emphatic harbingers of fortune.
Numerous frugivorous bats disperse the seeds of multiple Ficus spp., which produce figs. In tropical forest ecosystems, Ficus are keystone species - their fruits are an important resource for many wildlife including capuchin monkeys, langurs, hornbills and parrots.
Frugivorous species such as Epomophorus gambianus, Gambian epauletted fruit bats, and Pteropus medius, Indian flying fox, disperse the seeds of Psidium guajava - guavas.
"Apart from large plantations, guavas are found in the back yards of houses in towns and villages in West Africa. When the fruits are ripe, fruit bats ... feed on them. When a guava tree is near the roof of a house that is covered with corrugated iron sheets, the bats create continuous commotion by plucking the guavas with their mouths and dropping them on the roof while feeding." Some pteropodids are capable of carrying entire fruits while flying and thus, effectively distribute seeds.
In the United States, Nycticeius humeralis, evening bats, and Perimyotis subflavus, tri-colored bats consume multiple species capable of damaging grapes including Drosophila suzukii, spotted wing drosophila; Spodoptera exigua, beet armyworms; and Harmonia axyridis, harlequin ladybeetles.
In the Gironde region of southwest France, bats and vineyards are the perfect pairing - 19 of 22 native bat species consume grapevine and cochylis moths - the bane of French winegrowers.
Insectivorous species such as Eptesicus fuscus, big brown bats, consume Halyomorpha halys, brown marmorated stink bugs - one of the most harmful, invasive insect pests inhabiting North America. The host range of H. halys encompasses more than 120 different plants including many economically important fruit and vegetable crops such as peaches, nectarines, apples, asian pears, soybeans, okra, and hazelnuts.
Galleria mellonella, greater wax moths or honeycomb moths, possess sensitive tympanic "ears," an adaption resulting from selective pressures from insectivorous bats. Ultrasounds at frequencies simulating bats' echolocation calls elicit evasive maneuvers including dropping, looping and freezing. Worldwide, G. mellonella are ubiquitous parasites of honeybees, causing substantial economic losses both regionally and globally. In the southern United States, losses due to G. mellonella infestation approximate $3-4 million annually.
More than 200,000 species of bats, birds, small mammals, flies, beetles, wasps, ants, butterflies, moths and bees, pollinate more than 300,000 angiosperms - or flowering plants. Pollinators are essential to our daily lives, providing 1 of every 3 bites of food we eat. North America’s approximately 4,000 wild bee species pollinate agricultural crops of value to humans and the seeds, nuts, and fruits important to wild animals, from songbirds to grizzlies.
There are two bees hovering around the letter 'H.' Can you find 8 more bees hidden throughout the alphabet?
Inga edulis, native to South America, produces leguminous pods known as ice cream beans, a name originating from the sweet, succulent pulp surrounding the seeds, which resembles vanilla ice cream.
In Columbia, frugivores such as Artibeus planirostris, flat-faced fruit-eating bats and Dermanura phaeotis, pygmy fruit-eating bats, disperse the seeds of multiple pioneer species, particularly Bidens pilosa, Spanish needle; Piper crassinervium, pepper; and Physalis peruviana, - Inca berries.
Inca berries are economically important exotic fruits. In the United States, these delicious fruits are marketed as goldenberries or Pichuberries.
In Samoa, approximately 30% of rainforest canopy trees rely, at least partially, on two Pteropus species - P. samoensis, Samoan flying fox; and P. tonganus, Pacific flying fox, for pollination and seed dispersal. In various islands of the South Pacific, Pteropus tonganus consumes the fruits of multiple Artocarpus species including A. altilis, breadfruit and A. heterophylla, jackfruit.
Jackfruit, native to southeastern India and Sri Lanka, weighs up to 36 kg, with an average seed size and weight of 1.5 cm and 14 g, respectively. Fruit size and hardness limits use by phyllostomids - New World leaf-nosed bats- to small and medium size fruits. However,
"during a mammalian diversity research developed at Ilha Grande, Rio de Janeiro State, Brazil, it was observed an unusual feeding behaviour by Phyllostomus hastatus (one of the largest bats in Brazil, weighing more than 100g. In September 08th 2006, at least five bats approached a mature jackfruit (5 m height)and chewed the fruit shell, until they were able to enter the fruit. More than one bat was observed removing the shell at the same time, and pieces of the shell could be found fallen near the tree. The bats entered the hole, one by one, and ate the soft parts. Intense activity was observed, suggesting a group feeding behavior... Being the second species in size in neotropical region, P. hastatus, could be the sole bat species able to open the jackfruit and therefore eat it when the fruit is still hanging on the trunk. Although seed carrying was not observed, the disperser role of P. hastatus on A. heterophyllus can not be neglected."
In South India, Hipposideros ater, dusky leaf-nosed bats, control multiple lepidopteran and coleopteran pests such as Hippotion celerio, pest of grapevine; Othreis meterna and O. fullonica, pests of citrus and grapevine; Melanitis leda ismene, pest of rice, and Nausinoe geometralis, pest of jasmine.
The beautifully fragrant flowers of Jasminum species - jasmines - are both culturally and economically important; e.g., perfumes, religious ceremonies and festivals, jasmine-flower tea.
Bats are the primary pollinators of Ceiba pentandra, whose seed pods provide a beautifully fluffy, lightweight ecofriendly fibre known as kapok or silk cotton.
In West Africa, “when the flowers of Ceiba pentandra open, visits to them are made by Epomophorus gambianus, [Gambian epauletted fruit bats]; Nanonycteris veldkampi, [Veldkamp's dwarf epauletted fruit bats] and Eidolon helvum [straw-coloured fruit bats] … of these 3 bat species, N. veldkampi is somewhat the most important visitor, since these bats stay on the trees much longer in search of nectar."
Artibeus jamaicensis, Jamaican fruit bats, inhabiting the Yucatán Peninsula, Mexico, function as pollen vectors for Ceiba, which "play an important ecological role in the Yucatán landscape, supplying shade, nectar and fruit for wildlife."
Bombax ceiba - cotton trees - produce capsules containing cotton-esque fibers, similar to Ceiba pentandra. Both C. pentandra and B. ceiba share the name 'kapok' and 'silk cotton.'
B. ceiba's beautifully vibrant flowers bloom between February and April, attracting "insects and birds during day time and fruit bats during night." These "cotton-tree flowers" are the official flower of Guangzhou, the capital of Guangdong Province, southern China, and an essential ingredient of nam ngiao spicy noodle soup of Shan State and northern Thailand.
The evergreen Garcinia indica, or commonly kokum, produces reddish purple berries with fleshy endocarps. Numerous mammalian frugivores, including fruit bats, disperse kokum seeds, which represent 20-23% of the fruits' weight. G. indica has multiple culinary and pharmaceutical uses including kokum butter, a chocolate and sugar confectionary ingredient, and kokum sharbat, a refreshing juice containing kokum fruits, sugar, and bhuna jeera.
In South Africa, insectivorous species such as Scotophilus dinganii, African yellow bats; Neoromicia nana, banana bats; and Nycteris thebaica, Egyptian slit-faced bats consume Cryptophlebia peltastica, litchi moths, the primary pest of Litchi chinensis - lychees.
Lychees are native to the Guangdong and Fujian provinces of southeastern China - the main producers of these fleshy fruits.
From India to Madagascar, insectivorous bats are suppressors of multiple agricultural pests, particularly Ericeia inangulata, sober tabby; Eudocima materna, dot-underwing moths; and E. phalonia, common fruit-piercing moths, which damage citrus fruits, including lemons and limes.
In eastern Mediterranean habitats, Rousettus aegyptiacus, Egyptian fruit bats, naturally consume fruits of Ceratonia siliqua, carob; Ficus carica, figs; Morus nigra, mulberry; and Eriobotrya japonica, loquat; thereby functioning as important dispersal agents for > 4 fruit-bearing plant species.
In Italy, loquat seeds provide Nespolino liqueur, whose bitterness resembles other Italian seed-based liqueurs such as amaretto and nocino.
Several frugivorous species, including Pteropus tonganus, insular flying fox, consume Mangifera indica - the delicious mango. Mangos are the favorite foodstuff of Pteropus voeltzkowi, Pemba flying fox - an endearing species endemic to Pemba, an island approximately 31 miles off the Tanzanian coast. On the island, P. voeltzkowi may be one of the only species capable of dispersing large seeds. Furthermore, germination rates of bat‐ingested seeds are higher than those of ripe fruit, which appears to be linked to selective ingestion of viable seeds by bats.
In eastern Mediterranean habitats, Rousettus aegyptiacus, Egyptian fruit bats, disperse the seeds of several fruit-bearing plant species, including Morus nigra - mulberry.
In Costa Rica, Carollia perspicillata, Seba's short-tailed bats disperse the seeds of Chlorophora tinctoria, dyer's mulberry, which produces fustic, a bright yellow dye primarily known for coloring khaki fabric for US military apparel during World War I.
In South African macadamia orchards, insectivorous bat species opportunistically consume numerous insect pests, including Bathycoelia distincta, twin-spot stinkbugs; Nezara viridula, green vegetable bugs; and macadamia nut borers. These insect species can damage the macadamia kernel directly, thereby decreasing nut quality, and indirectly by causing nuts to drop immaturely, germinate or mold.
Flamboyant, pendulous vegetarian sausages and fragrant, magenta flowers are spectacular characteristics of Kigelia africana - the sausage tree. Micropteropus pussilus, the dwarf epaulet fruit bat, “lands on the lower tip of the flower and thrusts its head inside the flower to lap nectar ... On leaving the flower, the bat is dusted with pollen [which is] deposited on the stigma of the same or the next flower and thus effects pollination.”
The large, sausage-esque fruits enhance flavoring and facilitate fermentation of muratina - a staple of Central Kenya’s Kikuyu community. This artisan beer "is crisp and moderately sour with a mild, residual sweetness and smooth aftertaste."
In addition to muratina, Kigelia africana exhibits antiinflammatory, analgesic, antimicrobial and anticancer properties, valuable to cosmetics and topical pharmaceutical applications.
Rhinolophus euryale, Mediterranean horseshoe bats, predate Peribatodes rhomboidari, willow beauty moths whose herbivorous larval stage consumes "tea tree" leaves. Matcha, the exquisitely green, finely ground powder of shade-grown green tea leaves, provides flavoring and coloring for numerous foodstuff including confections and matcha lattes.
Insectivorous bats are integral components of terrestrial ecosystems, consuming insects worldwide. In agroecosystems, bats - like this Plecotus auritus, brown long-eared bat - provide valuable services by suppressing crop pests. In the United States alone, these services average $28 billion annually.
Azadirachta indica, the neem or Margosa tree, was introduced to West Africa from India "to supplement the indigenous species with exotic species that might be useful in the economic activities of the West African countries." Epomophorus gambianus, Gambian epauletted fruit bats, “are perhaps attracted to the Neem tree more than any other West African fruit bat. At dusk these bats leave their roosting places and converge in the forest reserve as well as the several acres of neem trees planted as “amentity trees in towns and villages throughout the country. The bats feed for long periods; sometimes a flock will stay on a tree for at least one hour before flying away. During their departure, hoewver, many of them carry neem fruits in their mouths and drop the seeds during flight before or after they have completed feeding on the fruit.”
Pteropus giganteus, Indian flying fox, structure forest communities through long-distance dispersal and pollination. In the Palaeotropics, bat-dispersed fruits are morphologically variable, exhibiting a variety of colors. Additionally, some fruits have a pungent fragrance, such as Morinda citrifolia, which entices bats with a "strong, vomit-like odor." The fruits of Morinda citrifolia, commonly noni, compose various beverages, powders, and cosmetics.
Pteropodids, Old World fruit bats, are phytophagous - consuming fruit, floral resources (nectar and pollen), leaves, and occasionally, insects. Phytophagous bats are neither true generalists nor specialists, but rather 'sequential specialists,' favoring at any given time one or a few plant species amongst the group of potential food sources available.
Pteropodids provide pollination and/or seed dispersal services to > 300 plants species of ~ 200 genera. These mammals are extraordinarily important long-distance dispersal agents, and perhaps the most important vertebrate dispersers on many oceanic islands, where they are the only animals capable of carrying large-seeded fruits.
In France, insectivorous Pipistrellus kuhlii, Kuhl's pipistrelles, consume Bactrocera oleae, olive fruit flies, a serious pest of olives. This species' range parallels those of genus Olea and remains the preeminent pest of olives, significantly affecting the quality and quantity of production within most olive-growing regions.
In southeastern Europe, Rhinolophus hipposideros, lesser horseshoe bats, predate 55 different pest species affecting a diversity of crops including grapevines, vegetables, and olive groves (e.g., Prays oleae, olive moths)
In North America, Tadarida brasiliensis, Mexican free-tailed bats, colonies consume enormous quantities of insects including several lepidopterans whose larvae are significant agricultural pests, such as Spodoptera frugiperda, fall armyworms; Trichoplusia ni, cabbage loopers; Heliothis virescens, tobacco budworms; and Helicoverpa zea, corn earworms. S. frugiperda, whose scientific name frugiperda, means lost fruit, decimates a multitude of crops including cotton, tobacco, sweet corn, rice, peanuts, and fruits such as apples, and oranges.
Worldwide, insectivorous bats such as Eptesicus fuscus, big brown bats; and Rhinolophus hipposideros, lesser horseshoe bats, consume numerous arthropods, including those damaging economically important rosaceous species such as almonds, cherries and raspberries.
Originally a combination of almonds and barley, orgeat syrups are an irreplaceable ingredient of Mai Tais and other classic libations. Today's orgeat syrups contain almonds, sugar and orange blossom water, adding a subtle sweetness and nuttiness to favorite cocktails.
Bats pollinate and disperse Pandanus spp., 'the vanilla of southeast Asia.' Prominent in Asian curries and Indian rice dishes, pandan leaf adds a delicately sweet floral fragrance to desserts.
In various islands of the South Pacific, Pteropus tonganus, Pacific flying fox, consume the fruits of Pandanus tectorius, or commonly, hala, a large fruit resembling a pineapple. Although an important resource for inhabitants of Micronesia, P. tectorius supports the beautifully bluish-green Megacrania batesii, peppermint stick insect, which lives and feeds only on this species.
Millions of Tadarida brasiliensis, Mexican free-tailed bats, ascend to high altitudes to exploit the seasonal migrations of billions of moths including Helicoverpa zea, corn earworms, and Spodoptera frugiperda, fall armyworms, from crops throughout the lower Rio Grande Valley of northern Mexico and southern Texas. Nearly 40% of the crop species that armyworms target are economically important including sweetcorn, oranges, rice, peanuts, papayas, strawberries, apples, cotton, grapes and peaches.
Pteropodids including Pteropus tonganus, Pacific flying fox, and Pteropus mariannus, Mariana fruit bats, disperse numerous fruits including Carica papaya - the papaya or pawpaw.
The largely neotropical genus Passiflora - the passion flowers or passion vines - comprises nearly 560 species, many of which require biotic pollination. Pollinators of Passiflora include hymenopterans (bees, wasps), trochilids (hummingbirds) and of course, nectarivorous chiropterans including Glossophaga commissarisi, Commissaris's long-tongued bats; Glossophaga soricina, Pallas's long-tongued bats; Anoura fistulata, tube-lipped nectar bats; Carollia perspicillata, Seba's short-tailed bats; and Monophyllus redmani, Leach's single leaf bats.
Additionally, frugivorous species consume the fruits of several Passiflora species thereby providing seed dispersal services.
In California, activity and foraging behavior of multiple bat species including Myotis californicus, California myotis; M. yumanensis, Yuma myotis; Lasiurus blossevillii, red bats; Tadarida brasiliensis, Mexican free-tailed bats; and Antrozous pallidus, pallid bats, "were detected in every Fairfield pear orchard that was surveyed and occurred during periods when the codling moth," one of the main pests of pears, "was expected to be active."
Nectarivorous bats, including Eonycteris spelaea, cave nectar bats, are principal pollinators of economically significant crops including Durio spp., durians; and Parkia spp., including Parkia speciosa - petai.
Petai, or 'stink beans,' are popular culinary ingredients of Indonesia, Malaysia, Singapore, Laos, southern Thailand, Burma, and northeastern India, despite their offensive, lingering aroma. In southern Thailand alone, the annual economic contribution of pollinating bats, relative to durian and petai, equals more than $137 million USD.
Melicoccus bijugatus, quenepa, of the soapberry family Sapindaceae, are native to northern South America and cultivated and naturalized widely throughout the tropics. In Trinidad and Tobago, bats and birds are responsible for seed dispersal. Quenepa fruits are delicious, sweet-tart 'drupes' with a flavor reminiscent of green seedless grapes. The fruit is used in preparing fruit juice, alcoholic beverages, and jellies.
In Ponce, Puerto Rico, the city celebrates quenepa during the annual 'Festival Nacional de la Quenepa' (National Quenepa Fruit Festival).
Cydonia oblonga, quince, are commercially grown fruits native to western Asia, Armenia, Turkey, Georgia, northern Iran and Afghanistan. In Europe, quinces are commonly grown in central and southern areas where the summers are sufficiently hot for the fruit to ripen.
In southwestern Europe, Rhinolophus hipposideros, lesser horseshoe bats, consume nearly 60 different pest arthropods including Cydia pomonella, codling moths. These lepidopterans damage several economically important fruits of the Rosaceae family including apples, pears, apricots, plums, cherries, and quinces.
Multiple frugivorous species, including Pteropus giganteus, Indian flying fox; P. tonganus, Pacific flying fox; Hypsignathus monstrosus, hammer-headed bats; and Cynopterus brachyotis, lesser short-nosed fruit bats consume and disperse the seeds of numerous Syzygium spp., including S. aqueum, S. jambos, and S. malaccense, whose beautiful fruits are called rose apples.
Worldwide, insectivorous bats contribute to sustainable Oryza production through effective biocontrol of major rice pests.
In Thailand, Chaerephon plicatus, wrinkle-lipped bats, consume large quantities of fulgoroids, or planthoppers, which are significant pests of Asian rice fields.
In southwestern Europe, Mythimna unipuncta, rice armyworms, are dietary constituents of both Rhinolophus euryale, Mediterranean horseshoe bats, and Rhinolophus hipposideros, lesser horseshoe bats. In addition to rice, this nocturnal lepidopteran damages various Graminae and other crops including barley, maize, oats, rye, sorghum, sugarcane, wheat, alfalfa, artichoke, bean, cabbage, carrot, celery, cucumber, lettuce, onion, parsley, pepper, and sweet potato.
In South India, Hipposideros ater, dusky leaf-nosed bats, control populations of multiple agricultural pests including Melanitis leda ismene, whose larvae, rice green-horned caterpillars, and adults, rice butterflies, are serious pests of rice.
In Swaziland, two African molossids - Chaerephon pumilus, little free-tailed bats, and Mops condylurus, Angolan free-tailed bats - preferentially forage over sugarcane plantations, thereby functioning as pest-controlling agents.
Sugarcane are perennial grasses of the genus Saccharum - essentially "reeds that produce honey without bees." Sugarcane agriculture accounts for 79% of global sugar production - the remaining percentage derives from Beta vulgaris, sugar beets. In addition to sugar, sugarcane provides falernum, molasses, rum, cachaça, bagasse, and ethanol.
Female Tadarida teniotis, European free-tailed bats, exhibit a dietary bias towards large migratory moths, including Autographa gamma, silver Y, and Agrotis ipsilon, ipsilon darts - two lepidopteran pests affecting crops such as tomatoes, potatoes and sugar beets.
Beta vulgaris, sugar beets, contain a high concentration of sucrose, and commercial cultivation of these plants provides approximately 55% of the United States' sugar production.
Numerous frugivorous species, including Artibeus lituratus, great fruit-eating bats; Epomophorus gambianus, Gambian epauletted fruit bats; and Cynopterus brachyotis, lesser short-nosed fruit bats consume and disperse Annona fruits such as A. muricata, soursop; A. squamosa, sweetsop; and A. montana, mountain soursop.
In North America, Nycticeius humeralis, evening bats; and Myotis lucifugus, little brown bats consume Harpalus pensylvanicus, Pennsylvania ground beetles; Paria fragariae, strawberry rootworms; and Korscheltellus lupulina, common swifts. These arthropods can damage Fragaria species including strawberries.
Vitellaria paradoxa, shea trees, common to West African savannahs, produce white flowers “that are clustered at the ends of the shoots and one-seeded fruits about 4-5 cm long with fleshy pulp. The bats feed on this fleshy pulp, releasing the seeds that fall to the ground where, after the fleshy pulp decomposes, the seeds are gathered by man.”
Three pteropodids native to western Africa - Epomophorus gambianus, Gambian epauletted fruit bats; Nanonycteris veldkampii, Veldkamp's dwarf epauletted fruit bats; and Micropteropus pusillus, Peter's dwarf epauletted fruit bats - are fundamental to long distance dispersion of V. paradoxa seeds.
Shea trees are incredibly valuable, both locally, nationally and internationally, producing edible oil, nutritious fruits, and shea butter.
In North America, insectivores including Tadarida brasiliensis, Mexican free-tailed bats; Eptesicus fuscus, big brown bats; Nycticeius humeralis, evening bats; and Myotis sodalis, Indiana bats; consume multiple economically relevant agricultural pests, such as Chinavia hilaris, green stinkbugs; Halyomorpha halys, brown marmorated stinkbugs; Helicoverpa zea, corn earworms; and Diabrotica undecimpunctata, spotted cucumber beetles. These species can decimate many agricultural crops, including Glycine max - soybeans.
Although native to Asia, soybeans are globally important, providing numerous foodstuff including soy milk, soy vegetable oil, tofu, soy sauce, fermented bean paste, nattō, and tempeh.
In South Asia, multiple pteropdids including Pteropus giganteus, Indian flying fox, and Cynopterus sphinx, greater short-nosed fruit bats, disperse the seeds of Madhuca longifolia, an Indian tropical tree commonly known as mahua or butter tree.
In India, the leaves of Madhuca longifolia are eaten by the moth Antheraea paphia, South Indian small tussore, which produces tussar silk, a commercially important wild silk.
In the southwestern United States, Mexico, and northern South America, three Leptonycteris species pollinate numerous economically and ecologically important members of Agavaceae, agave family, and Cactaceae, cactus family. The bat-pollinated Agave tequilana provides commercial tequila, a multimillion dollar industry; other Agave species produce similar alcoholic beverages including pulque, mescal, and bacanora.
Artibeus lituratus, great fruit-eating bats, consume seeds of secondary forest species Cecropia obtusa, and also fruits of primary forest species such as Caryocar glabrum, souari trees; Swartzia panacoco, Brazilian ebony; and Dipteryx odorata - coumarou - whose seeds are known as tonka beans.
Tonka seeds contain coumarin, a chemical responsible for the beans' intoxicating fragrance - aromas of vanilla, licorice, caramel, cherry and clove. The aromatic seeds "impart a flavour so transcendent, tonka has been dubbed the most delicious ingredient you’ve never heard of."
In Southeast Asia, Cynopterus brachyotis, lesser dog-faced fruit bats, and Pteropus marianus, Mariana fruit bats, disperse the fruits of Calophyllum inophyllum, the source of culturally importanttamanu oil. One West Malaysia study indicates C. brachyotis consumes fruits from 38 different plant species, including C. inophyllum and Ficus fistulosa, whose fruits are available throughout the year. Consumption of such fruits may be critical to maintaining stable populations of C. brachyotis.
Borassus flabellifer, tala palm or toddy palm, of the palm family Arecaceae, are native to the Indian subcontinent and Southeast Asia; including Nepal, India, Bangladesh, Sri Lanka, Cambodia, Laos, Burma, Thailand, Vietnam, Malaysia, Indonesia and the Philippines. Frugivorous mammals, particularly Pteropus giganteus, Indian flying fox, consume fleshy fruits - Palmyra fruits - which contain deliciously "sweet jelly seed sockets."
A sugary sap, or toddy, derives from the young inflorescence, and was Thailand's predominant source of sugar before sugarcane.
In Columbia, after coffee, flower crops represent an economically significant agricultural commodity, with exports exceeding $800 million USD annually. The US Department of Agriculture, Animal and Plant Health Inspection Service frequently intercepts Copitarsia spp. on fresh commodities arriving from Mexico, Central America and South America. "Whenever the genus is detected in imported goods, shipments must be disinfested, destroyed, or returned to the country of origin," thereby precluding entry of exotic arthropods, which threaten species diversity and ecosystem processes. Simulating the fundamental parameters that characterize the echolocation signals of Tadarida brasiliensis, Mexican free-tailed bats, initiates an evasive response by Copitarsia decolora. This ecologically friendly management option "simulates the presence from the natural predator of the plague, the bat, to prevent the entrance of Copitarsia decolora to places of flower packaging, post-harvest rooms."
Capitarsia larvae cause both direct and indirect damage, affecting both quality and yield of numerous crop species, including Ullucus tuberosus - ulluco, one of the most economically important root crops of South America.
To overcome the hazards of chemical fertilizers and pesticides, modern India's agriculturalists are returning to traditional practices, employing organic techniques, fertilizers and pesticides. As natural manure, bat guano contains "all the macro and micronutrients that plants require in a natural form and hence ably serve as plant fertilizer, soil builder, soil cleanser, fungicide, nematocide and compost activator."
Small quantities of this biofertilizer have 'plant growth promoting properties,' including microflora and requisite nutrients e.g., nitrogen and phosphorus. The application of guano to Vigna mungo, urad beans; and Vigna radiata, mung beans, achieve "better growth, biomass and nitrogen content."
Cerrado, a biodiversity hotspot covering nearly 10% of South America, harbors more than 4800 endemic species. An incredible 44% of Cerrado's mammalian species are bats, more than half of which are phyllostomids, New World leaf-nosed bats. Members of this extraordinarily diverse family are critical seed dispersers and pollinators of > 549 and 360 species of Neotropical plants, respectively. In addition to hummingbirds, Glossophaga soricina, Pallas' long-tongued bats, and Artibeus planirostris, flat-faced fruit-eating bats, pollinate Vanilla chamissonis, or commonly Chamisso's vanilla - one of ~ 110 species of vanilla orchid. The genus Vanilla exhibits considerable morphological diversity, suggesting different animals can be effective pollinators.
The natural pollinator(s) of Vanilla planifolia - source of commercial vanilla - remains unclear, "and for a long time it has been said that bees (Melipona beechii), hummingbirds (Cynniris spp.) and bats pollinate vanilla." However, a preponderance of evidence favors pollination by Euglossa viridissima, shiny green orchid bees.
DYK | Mexico is one of the few countries where it is possible to obtain vanilla beans through natural pollination, although it happens rarely, accounting for only about 1% of all fruits.
Mucuna spp. are leguminous lianas producing 'buoyant sea-beans' with a characteristic 3-layer appearance, inspiring common names such as deer-eye beans, donkey-eye beans, ox-eye beans or hamburger beans. Mucuna species are generally bat-pollinated, with some species exhibiting extraordinary morphologies indicative of an evolutionary adaptation to echolocating glossophagine bats. Mucuna pruriens - velvet beans - are an important source of L-dopa, a common component of nootropics, or "smart drugs."
Worldwide, insectivorous bats suppress arthropod populations, providing natural biocontrol services for vineyards. In Europe's vineyard agroecosystems, Rhinolophus hipposideros, lesser horseshoe bats, consumes multiple significant grapevine pests including Ephestia parasitella, false cacao moths; Lobesia botrana, European grapevine moths; Peridroma saucia, variegated cutworms; Sparganothis pilleriana, long-palped tortrix; Spodoptera exigua, beet armyworms; and Drosophila suzukii, spotted wing drosophila.
A nonalcoholic byproduct of wine production, verjus bestows a "titillating combination of acidity and sweetness" upon salad dressings, marinades, sauces and cocktails. Verjus are flavorful juices made from pressing unripe, unfermented grapes, and were once a staple of French provincial cooking.
The genus Piper includes > 1000 species and exhibits pantropical distribution, with the greatest diversity occurring within the American tropics (700 spp.) and Southern Asia (300 spp.), where the economically important species Piper nigrum, black pepper; and Piper betle, betel leaf, originate. Piper species are commonly dominant elements within tropical forests; "not surprisingly, Piper species are of great ecological importance and have been considered "key" species on the basis of their association with frugivorous bats." In fact, "all of the known fruit dispersal in Piper is carried out by bats primarily and birds secondarily."
Thus, eastern Madagascar's three frugivorous species - Pteropus rufus, Madagascan flying fox; Eidolon dupreanum, Madagascan fruit bats; and Rousettus madagascariensis, Madagascan rousettes may similarly disperse Piper borbonense, Madagascan wild pepper, whose peppercorns provide the deliciously aromatic spice, voatsiperifery.
Cydia pomonella, codling moths, are significant pests of pome fruits and walnuts worldwide. In California walnut orchards, insectivorous predators including Tadarida brasiliensis, Mexican free-tailed bats, consume these destructive lepidopterans.
In southcentral Texas and northern Mexico, enormous colonies of Tadarida brasiliensis,Mexican free-tailed bats, consume multiple noctuids whose larvae damage agricultural crops, such as Spodoptera frugiperda, fall armyworms; Trichoplusia ni, cabbage loopers; Heliothis virescens, tobacco budworms; and Helicoverpa zea, corn earworms, whose larvae can decimate a variety of plants, including watermelon.
In Ghana, 30 plant species belonging to > 16 families are foodstuff for Epomophorus gambianus, Gambian epauletted fruit bats. These dietary sources include the edible fruits of Melothria spp. - Mexican miniature watermelons, or more commonly mouse melons. By consuming such fruits, E. gambianus contribute to the successful dispersion of seeds.
In central Thailand, Chaerephon plicatus, wrinkle-lipped bats, consume massive quantities of fulgoroids, or leafhoppers - which can devastate Oryza fields. One species, Oryza sativa - Asian rice - "has supported a greater number of people for a longer period of time than any other crop since it was domesticated between 8,000 to 10,000 years ago." Rice remains the staple foodstuff of 2.7 billion people - over half the world's population.
Oryza sativa var. glutinosa - sticky rice or glutinous rice - provides xôi, a popular Vietnamese fare made from glutinous rice and other ingredients, such as xôi sầu riêng - sticky rice and durian.
Approximately a quarter of Cactaceae's 27 genera exhibit chiropterophily, a pollination syndrome where angiosperms exhibit "bat loving" floral characteristics such as nocturnal anthesis and pale coloration. These include Pilosocereus spp., tree cactus; Stenocereus spp., Cereus repandus, Peruvian apple cactus; Hylocereus spp.; Pachycereus pringlei, cardón cactus; and Carnegiea gigantea, saguaro cactus.
Bats also contribute to Opuntia spp., a genus of cacti comprising the prickly pears. Opuntia jonocostle and Opuntia matudae produce sour prickly pears, or xoconostles - popular ingredients of Mexican cuisine. Fertilization influences plant production and the genus Opuntia especially, responds rapidly to organic fertilizers. The application of organic fertilizer - bat guano - increases the yield, weight, and onset of fructification - producing fruits one year earlier than plants without fertilizer.
Uroderma species, tent-making bats,
are phyllostomids - commonly "leaf-nosed bats" - of Central and South America. These primarily frugivorous mammals are 1 of only 22 bat species that modify natural structures (e.g., leaves) to construct suitable roosts.
On Tutuila and Aunu'u Island, American Samoa, Pteropus samoensis, Samoan flying fox, preferentially consume, and disperse, numerous native fruits, including Ficus spp., figs; Inocarpus fagifer, Tahitian chestnut; Terminalia catappa, sea almond, and Cananga odorata fragrant cananga, whose beautiful flowers yield the essential oil, ylang-ylang, a popular aromatherapy and perfumery ingredient.
Worldwide, frugivorous and nectarivorous bats contribute to the persistence of valuable timber species, including Pachira quinata, pochote; Kigelia africana, sausage tree; Daniellia oliveri, West African copal tree; Castanospermum australe, blackbean tree; Talipariti elatum, blue mahoe tree; and Caryocar spp., souari trees.
Artibeus lituratus are "capable of transporting fruits and seeds almost as large as itself," including those of Bocoa prouacensis, which produces "one of the hardest and densest [timbers] in the world."
The contiguous Maya forest of Belize, Guatemala, and Mexico boasts an extraordinary and distinctive biodiversity, comprising an abundance of economically important flora, particularly for Maya descendants. "Over 98% of tropical forest plants are pollinated by fauna, that is insects, birds, and bats; the Maya forest is no exception."
Dominant bat-pollinated plants include Musa paradisiacal, banana; Manilkara zapota, chicle; and Pouteria sapota, zac-xa-nal, or more commonly, mamey sapote.
In Indiana, Eptesicus fuscus, big brown bats, preferentially consume agricultural pest species including Scarabaeidae, scarab beetles; pentatomids, stinkbugs; cicadellids, leafhoppers; and Diabrotica undecimpunctata, spotted cucumber beetles - a significant pest of cucurbits such as cucumbers, watermelons, pumpkins, and zucchini. Larvae of genus Diabrotica, commonly corn rootworms, are "probably the most important agricultural pest in the United States," costing American farmers approximately 1 billion USD annually.