蝙蝠声音信号功能研究进展
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摘要
声音信号介导动物的资源竞争、配偶选择及反捕食等系列生活史事件,对维持动物种群稳定与群落平衡至关重要。蝙蝠占据夜空生态位,视觉退化,听觉发达,一直被视为声学研究的模式生物。本文围绕蝙蝠回声定位声波与交流声波的功能,综述当前研究现状,展望未来发展方向。纵观已有研究,蝙蝠回声定位声波具有多重功能,包括空间导航与猎物探测、水体与植被识别、协调觅食活动及传递交流信息。蝙蝠利用交流声波从事社群活动,包括社群联系、资源防御、繁殖活动及求救呼叫。绝大多数研究关注蝙蝠回声定位声波的功能,有关蝙蝠交流声波的功能近年才引起重视。未来工作有待进一步探究蝙蝠声音信号产生的影响机制。
Acoustic signals mediate a variety of life history events in animals,such as resource competition,mate selection,and antipredator defense. Therefore,it has been proposed that acoustic signals have profound effects on population stability and community dynamics. Echolocating bats occupy a nocturnal niche in the sky. They have comparatively poor vision but well-developed auditory sense,providing a good model for acoustic research. In this review,we summarize the current studies regarding the function of echolocation and social calls in bats. We provide some recommendations and directions for future work. We consider that the echolocation calls of bats serve multiple functions including spatial orientation,prey tracking,recognition of water and plant,foraging coordination,and social information transfer. By contrast,bats use social calls to engage in social activities,including social contact,resource competition,mate choice,and distress calling. Despite the large amount of previous studies which have focused on echolocation calls,study on the function of social calls in bats is emerging in recent years. Further work in this field is needed to explore the influence mechanisms of the production of acoustic signals among bats.
Acoustic signals mediate a variety of life history events in animals,such as resource competition,mate selection,and antipredator defense. Therefore,it has been proposed that acoustic signals have profound effects on population stability and community dynamics. Echolocating bats occupy a nocturnal niche in the sky. They have comparatively poor vision but well-developed auditory sense,providing a good model for acoustic research. In this review,we summarize the current studies regarding the function of echolocation and social calls in bats. We provide some recommendations and directions for future work. We consider that the echolocation calls of bats serve multiple functions including spatial orientation,prey tracking,recognition of water and plant,foraging coordination,and social information transfer. By contrast,bats use social calls to engage in social activities,including social contact,resource competition,mate choice,and distress calling. Despite the large amount of previous studies which have focused on echolocation calls,study on the function of social calls in bats is emerging in recent years. Further work in this field is needed to explore the influence mechanisms of the production of acoustic signals among bats.
引文
冯江.2001.蝙蝠回声定位行为生态研究[M].长春:吉林科学技术出版社.
吴飞健,陈其才.1997.蝙蝠听觉器的“视”功能[J].生物学通报,32(7):16-18.
张树义,冯江.1999.三种蝙蝠飞行状态下回声定位信号的比较[J].动物学报,45(4):385-389.
Arnold BD,Wilkinson GS.2011.Individual specific contact calls of pallid bats(Antrozous pallidus)attract conspecifics at roosting sites[J].Behavioral Ecology and Sociobiology,65(8):1581-1593.
August P.1979.Distress calls in Artibeus jamaicensis:ecology and evolutionary implications[M]//Eisenberg JF.Vertebrate ecology in the northern Neotropics.Washington:Smithsonian Institution Press:151-159.
Balcombe JP.1990.Vocal recognition of pups by mother Mexican free-tailed bats,Tadarida brasiliensis mexicana[J].Animal Behaviour,39(5):960-966.
Barclay RMR.1982.Interindividual use of echolocation calls:eavesdropping by bats[J].Behavioral Ecology and Sociobiology,10(4):271-275.
Barlow KE,Jones G.1997.Function of pipistrelle social calls:field data and a playback experiment[J].Animal Behaviour,53(5):991-999.
Bastian A,Schmidt S.2008.Affect cues in vocalizations of the bat,Megaderma lyra,during agonistic interactions[J].Journal of the Acoustical Society of America,124(1):598-608.
Behr O,Helversen O.2004.Bat serenades-complex courtship songs of the sac-winged bat(Saccopteryx bilineata)[J].Behavioral Ecology and Sociobiology,56(2):106-115.
Behr O,von Helversen O,Heckel G,et al.2006.Territorial songs indicate male quality in the sac-winged bat Saccopteryx bilineata(Chiroptera,Emballonuridae)[J].Behavioral Ecology,17(5):810-817.
Bohn KM,Schmidt-French B,Ma ST,et al.2008.Syllable acoustics,temporal patterns,and call composition vary with behavioral context in Mexican free-tailed bats[J].Journal of the Acoustical Society of America,124(3):1838-1848.
Bradbury J,Vehrencamp SL.1998.Principles of animal communication[M].Sunderland:Sinauer Associates.
Carter GG,Logsdon R,Arnold BD,et al.2012.Adult vampire bats produce contact calls when isolated:acoustic variation by species,population,colony,and individual[J].PLo S ONE,7(6):e38791.DOI:10.1371/journal.pone.0038791.
Carter G,Schoeppler D,Manthey M,et al.2015.Distress calls of a fast-flying bat(Molossus molossus)provoke inspection flights but not cooperative mobbing[J].PLo S ONE,10(9):e0136146.DOI:10.1371/journal.pone.0136146.
Chaverri G,Gillam EH,Vonhof MJ.2010.Social calls used by a leaf-roosting bat to signal location[J].Biology Letters,6(4):441-444.
Davidson SM,Wilkinson GS.2004.Function of male song in the greater white-lined bat,Saccopteryx bilineata[J].Animal Behaviour,67(5):883-891.
Dechmann DKN,Heucke SL,Giuggioli L,et al.2009.Experimental evidence for group hunting via eavesdropping in echolocating bats[J].Proceedings of the Royal Society B,276(1668):2721-2728.
Eckenweber M,Kn9rnschild M.2016.Responsiveness to conspecific distress calls is influenced by day-roost proximity in bats(Saccopteryx bilineata)[J].Royal Society Open Science,3(5):160151.DOI:10.1098/rsos.160151.
Fenton MB,Belwood JJ,Fullard JH,et al.1976.Responses of Myotis lucifugus(Chiroptera:Vespertilionidae)to calls of conspecifics and to other sounds[J].Canadian Journal of Zoology,54(9):1443-1448.
Fenton MB,Jacobs DS,Richardson EJ,et al.2004.Individual signatures in the frequency-modulated sweep calls of African large-eared,free-tailed bats Otomops martiensseni(Chiroptera:Molossidae)[J].Journal of Zoology,262(1):11-19.
Fenton MB.2003.Eavesdropping on the echolocation and social calls of bats[J].Mammal Review,33(3):193-204.
Fenton MB.2013.Questions,ideas and tools:lessons from bat echolocation[J].Animal Behaviour,85(5):869-879.
Fernandez AA,Fasel N,Kn9rnschild M,et al.2014.When bats are boxing:aggressive behaviour and communication in male Seba's short-tailed fruit bat[J].Animal Behaviour,98:149-156.
Furmankiewicz J,Ruczyński I,Urban R,et al.2011.Social calls provide tree-dwelling bats with information about the location of conspecifics at roosts[J].Ethology,117:480-489.
Ganesh A,Raghuram H,Nathan PT,et al.2010.Distress callinduced gene expression in the brain of the Indian short-nosed fruit bat,Cynopterus sphinx[J].Journal of Comparative Physiology A,196(2):155-164.
Geipel I,Jung K,Kalko EKV.2013.Perception of silent and motionless prey on vegetation by echolocation in the gleaning bat Micronycteris microtis[J].Proceedings of the Royal Society B,280(1754):20122830.DOI:10.1098/rspb.2012.2830.
Gillam E,Fenton MB.2016.Roles of acoustic social communication in the lives of bats[M]//Fenton BM,Grinnell DA,Popper NA,et al.Bat bioacoustics.New York:Springer:117-139.
Gillam EH.2007.Eavesdropping by bats on the feeding buzzes of conspecifics[J].Canadian Journal of Zoology,85(7):795-801.
Goerlitz HR,ter Hofstede HM,Zeale MR,et al.2010.An aerial-hawking bat uses stealth echolocation to counter moth hearing[J].Current Biology,20(17):1568-1572.
Gould E.1975.Neonatal vocalizations in bats of eight genera[J].Journal of Mammalogy,56(1):15-29.
Greif S,Siemers BM.2010.Innate recognition of water bodies in echolocating bats[J].Nature Communications,1(1):107.DOI:10.1038/ncomms1110.
Griffin DR.1944.Echolocation by blind men,bats and radar[J].Science,100(2609):589-590.
Hage SR,Jiang TL,Berquist SW,et al.2013.Ambient noise induces independent shifts in call frequency and amplitude within the Lombard effect in echolocating bats[J].Proceedings of the National Academy of Sciences of the United States of America,110(10):4063-4068.
Halfwerk W,Jones P,Taylor R,et al.2014.Risky ripples allow bats and frogs to eavesdrop on a multisensory sexual display[J].Science,343(2609):413-416.
Huang XB,Kanwal JS,Jiang TL,et al.2015.Situational and age-dependent decision making during life threatening distress in Myotis macrodactylus[J].PLo S ONE,10(7):e0132817.DOI:10.1371/journal.pone.0132817.
Hughes NK,Kelley JL,Banks PB.2012.Dangerous liaisons:the predation risks of receiving social signals[J].Ecology Letters,15(11):1326-1339.
Jakobsen L,Olsen MN,Surlykke A.2015.Dynamics of the echolocation beam during prey pursuit in aerial hawking bats[J].Proceedings of the National Academy of Sciences,112(26):8118-8123.
Jakobsen L,Ratcliffe JM,Surlykke A.2013.Convergent acoustic field of view in echolocating bats[J].Nature,493(7430):93-96.
Jiang TL,Metzner W,You YY,et al.2010.Variation in the resting frequency of Rhinolophus pusillus in mainland China:effect of climate and implications for conservation[J].Journal of the Acoustical Society of America,128(4):2204-2211.
Jin LR,Yang SL,Kimball RT,et al.2015.Do pups recognize maternal calls in pomona leaf-nosed bats,Hipposideros pomona?[J].Animal Behaviour,100:200-207.
Jones G,Siemers BM.2010.The communicative potential of bat echolocation pulses[J].Journal of Comparative Physiology A,197(5):447-457.
Jones G,Teeling E.2006.The evolution of echolocation in bats[J].Trends in Ecology&Evolution,21(3):149-156.
Jones G.1997.Acoustic signals and speciation:the roles of natural and sexual selection in the evolution of cryptic species[J].Advances in the Study of Behaviour,26(8):317-354.
Kalko EK,Schnitzler H.1998.How echolocating bats approach and acquire food[M]//Kunz TH,Racey PA.Bat biology and conservation.London:Smithsonian Institution Press:197-204.
Kanwal JS,Matsumura S,Ohlemiller K,et al.1994.Analysis of acoustic elements and syntax in communication sounds emitted by mustached bats[J].Journal of the Acoustical Society of America,96(3):1229-1254.
Kazial KA,Kenny TL,Burnett SC.2008.Little brown bats(Myotis lucifugus)recognize individual identity of conspecifics using sonar calls[J].Ethology,114(5):469-478.
Kazial KA,Masters WM.2004.Female big brown bats,Eptesicus fuscus,recognize sex from a caller's echolocation signals[J].Animal Behaviour,67(5):855-863.
Kn9rnschild M,Blüml S,Steidl P,et al.2017.Bat songs as acoustic beacons-male territorial songs attract dispersing females[J].Scientific Reports,7:13918.DOI:10.1038/s41598-017-14434-5.
Kn9rnschild M,Jung K,Nagy M,et al.2012.Bat echolocation calls facilitate social communication[J].Proceedings of the Royal Society B,279(1748):4827-4835.
Kn9rnschild M,Nagy M,Metz M,et al.2010.Complex vocal imitation during ontogeny in a bat[J].Biology Letters,6(2):156-159.
Kn9rnschild M,von Helversen O.2008.Nonmutual vocal mother-pup recognition in the greater sac-winged bat[J].Animal Behaviour,76(3):1001-1009.
Kober R,Schnitzler HU.1990.Information in sonar echoes of fluttering insects available for echolocating bats[J].Journal of the Acoustical Society of America,87(2):882-896.
Kunz TH,Fenton MB.2003.Bat ecology[M].Chicago:University of Chicago Press.
Lima SL,O'Keefe JM.2013.Do predators influence the behaviour of bats?[J].Biological Reviews,88(3):626-644.
Lin HJ,Kanwal JS,Jiang TL,et al.2015.Social and vocal behavior in adult greater tube-nosed bats(Murina leucogaster)[J].Zoology,118(3):192-202.
Liu Y,Feng J,Jiang YL,et al.2007.Vocalization development of greater horseshoe bat,Rhinolophus ferrumequinum(Rhinolophidae,Chiroptera)[J].Folia Zoologica,56(2):126-136.
Luo B,Huang XB,Li YY,et al.2017b.Social call divergence in bats:a comparative analysis[J].Behavioral Ecology,28(2):533-540.
Luo B,Jiang TL,Liu Y,et al.2013.Brevity is prevalent in bat short-range communication[J].Journal of Comparative Physiology A,199(4):325-333.
Luo B,Lu GJ,Chen K,et al.2017a.Social calls honestly signal female competitive ability in Asian particoloured bats[J].Animal Behaviour,127:101-108.
Luo JH,Goerlitz HR,Brumm H,et al.2015.Linking the sender to the receiver:vocal adjustments by bats to maintain signal detection in noise[J].Scientific Reports,5:18556.DOI:10.1038/srep18556.
Mller R,Kuc R.2000.Foliage echoes:a probe into the ecological acoustics of bat echolocation[J].Journal of the A-coustical Society of America,108(2):836-845.
Morell V.2014.When the bat sings[J].Science,344(6190):1334-1337.
Moss CF,Zagaeski M.1994.Acoustic information available to bats using frequency-modulated sounds for the perception of insect prey[J].Journal of the Acoustical Society of America,95(5):2745-2756.
Nelson JE.1965.Behaviour of Australian pteropodidae(Megacheroptera)[J].Animal Behaviour,13(4):544-557.
Obrist MK.1995.Flexible bat echolocation:the influence of individual,habitat and conspecifics on sonar signal design[J].Behavioral Ecology and Sociobiology,36(3):207-219.
Racey PA,Swift SM.1985.Feeding ecology of Pipistrellus pipistrellus(Chiroptera:Vespertilionidae)during pregnancy and lactation.Ⅰ.Foraging behaviour[J].Journal of Animal E-cology,54(1):205-215.
Roverud RC,Nitsche V,Neuweiler G.1991.Discrimination of wingbeat motion by bats,correlated with echolocation sound pattern[J].Journal of Comparative Physiology A,168(2):259-263.
RuppL,Clment G,Herrel A,et al.2015.Environmental constraints drive the partitioning of the soundscape in fishes[J].Proceedings of the National Academy of Sciences of the United States of America,112(19):6092-6097.
Russ J,Racey P,Jones G.1998.Intraspecific responses to distress calls of the pipistrelle bat,Pipistrellus pipistrellus[J].Animal Behaviour,55(3):705-713.
Russ J,Jones G,Mackie I,et al.2004.Interspecific responses to distress calls in bats(Chiroptera:Vespertilionidae):a function for convergence in call design?[J].Animal Behaviour,67(6):1005-1014.
Russo D,Cistrone L,Jones G.2012.Sensory ecology of water detection by bats:a field experiment[J].PLo S ONE,7(10):e48144.DOI:10.1371/journal.pone.0048144.
Rydell J.1986.Feeding territoriality in female northern bats,Eptesicus nilssoni[J].Ethology,72(4):329-337.
Schnitzler HU,Moss CF,Denzinger A.2003.From spatial orientation to food acquisition in echolocating bats[J].Trends in Ecology&Evolution,18(8):386-394.
Schuchmann M,Puechmaille SJ,Siemers BM.2012.Horseshoe bats recognise the sex of conspecifics from their echolocation calls[J].Acta Chiropterologica,14(1):161-166.
Servick K.2014.Eavesdropping on ecosystems[J].Science,343(6173):834-837.
Siemers BM,Schnitzler HU.2004.Echolocation signals reflect niche differentiation in five sympatric congeneric bat species[J].Nature,429(6992):657-661.
Simmons JA,Ferragamo MJ,Moss CF.1998.Echo-delay resolution in sonar images of the big brown bat,Eptesicus fuscus[J].Proceedings of the National Academy of Sciences of the United States of America,95(21):12647-12652.
Simmons JA,Houser D,Kloepper L.2014.Localization and classification of targets by echolocating bats and dolphins[M]//Surlykke A,Nachtigall EP,Fay RR,et al.Biosonar.New York:Springer:169-193.
Simon R,Holderied MW,Koch CU,et al.2011.Floral acoustics:conspicuous echoes of a dish-shaped leaf attract bat pollinators[J].Science,333(6042):631-633.
Suga N,Niwa H,Taniguchi I,et al.1987.The personalized auditory cortex of the mustached bat:adaptation for echolocation[J].Journal of Neurophysiology,58(4):643-654.
Sun KP,Luo L,Kimball RT,et al.2013.Geographic variation in the acoustic traits of greater horseshoe bats:testing the importance of drift and ecological selection in evolutionary processes[J].PLo S ONE,8(8):e70368.DOI:10.1371/journal.pone.0070368.
Thiagavel J,Cechetto C,Santana SE,et al.2018.Auditory opportunity and visual constraint enabled the evolution of echolocation in bats[J].Nature Communications,9(1):98.DOI:10.1038/s41467-017-02532-x.
Ulanovsky N,Moss CF.2008.What the bat's voice tells the bat's brain[J].Proceedings of the National Academy of Sciences of the United States of America,105(205):8491-8498.
Vannoni E,McElligott AG.2008.Low frequency groans indicate larger and more dominant fallow deer(Dama dama)males[J].PLo S ONE,3(9):e3113.DOI:10.1371/journal.pone.0003113.
Vehrencamp SL.2000.Handicap,index,and conventional signal elements of bird song[M]//Espmark Y,Amundsen T,Rosenqvist G.Animal signals:signalling and signal design in animal communication.Norway:Tapir Academic Press:277-300.
Voigt-Heucke SL,Taborsky M,Dechmann DKN.2010.A dual function of echolocation:bats use echolocation calls to identify familiar and unfamiliar individuals[J].Animal Behaviour,80(1):59-67.
von der Emde G,Schnitzler HU.1990.Classification of insects by echolocating greater horseshoe bats[J].Journal of Comparative Physiology A,167(3):423-430.
von Helversen D,von Helversen O.1999.Acoustic guide in bat-pollinated flower[J].Nature,398:759-760.
Wilkinson GS,Boughman JW.1998.Social calls coordinate foraging in greater spear-nosed bats[J].Animal Behaviour,55(2):337-350.
Yovel Y,Franz MO,Stilz P,et al.2008.Plant classification from bat-like echolocation signals[J].PLo S Computational Biology,4(3):e1000032.DOI.10.1371/journal.pcbi.1000032.
Yovel Y,Melcon ML,Franz MO,et al.2009b.The voice of bats:how greater mouse-eared bats recognize individuals based on their echolocation calls[J].PLo S Computational Biology,5(6):e1000400.DOI:10.1371/journal.pcbi.1000400.
Yovel Y,Stilz P,Franz MO,et al.2009a.What a plant sounds like:the statistics of vegetation echoes as received by echolocating bats[J].PLo S Computational Biology,5(7):e1000429.DOI:10.1371/journal.pcbi.1000429.
吴飞健,陈其才.1997.蝙蝠听觉器的“视”功能[J].生物学通报,32(7):16-18.
张树义,冯江.1999.三种蝙蝠飞行状态下回声定位信号的比较[J].动物学报,45(4):385-389.
Arnold BD,Wilkinson GS.2011.Individual specific contact calls of pallid bats(Antrozous pallidus)attract conspecifics at roosting sites[J].Behavioral Ecology and Sociobiology,65(8):1581-1593.
August P.1979.Distress calls in Artibeus jamaicensis:ecology and evolutionary implications[M]//Eisenberg JF.Vertebrate ecology in the northern Neotropics.Washington:Smithsonian Institution Press:151-159.
Balcombe JP.1990.Vocal recognition of pups by mother Mexican free-tailed bats,Tadarida brasiliensis mexicana[J].Animal Behaviour,39(5):960-966.
Barclay RMR.1982.Interindividual use of echolocation calls:eavesdropping by bats[J].Behavioral Ecology and Sociobiology,10(4):271-275.
Barlow KE,Jones G.1997.Function of pipistrelle social calls:field data and a playback experiment[J].Animal Behaviour,53(5):991-999.
Bastian A,Schmidt S.2008.Affect cues in vocalizations of the bat,Megaderma lyra,during agonistic interactions[J].Journal of the Acoustical Society of America,124(1):598-608.
Behr O,Helversen O.2004.Bat serenades-complex courtship songs of the sac-winged bat(Saccopteryx bilineata)[J].Behavioral Ecology and Sociobiology,56(2):106-115.
Behr O,von Helversen O,Heckel G,et al.2006.Territorial songs indicate male quality in the sac-winged bat Saccopteryx bilineata(Chiroptera,Emballonuridae)[J].Behavioral Ecology,17(5):810-817.
Bohn KM,Schmidt-French B,Ma ST,et al.2008.Syllable acoustics,temporal patterns,and call composition vary with behavioral context in Mexican free-tailed bats[J].Journal of the Acoustical Society of America,124(3):1838-1848.
Bradbury J,Vehrencamp SL.1998.Principles of animal communication[M].Sunderland:Sinauer Associates.
Carter GG,Logsdon R,Arnold BD,et al.2012.Adult vampire bats produce contact calls when isolated:acoustic variation by species,population,colony,and individual[J].PLo S ONE,7(6):e38791.DOI:10.1371/journal.pone.0038791.
Carter G,Schoeppler D,Manthey M,et al.2015.Distress calls of a fast-flying bat(Molossus molossus)provoke inspection flights but not cooperative mobbing[J].PLo S ONE,10(9):e0136146.DOI:10.1371/journal.pone.0136146.
Chaverri G,Gillam EH,Vonhof MJ.2010.Social calls used by a leaf-roosting bat to signal location[J].Biology Letters,6(4):441-444.
Davidson SM,Wilkinson GS.2004.Function of male song in the greater white-lined bat,Saccopteryx bilineata[J].Animal Behaviour,67(5):883-891.
Dechmann DKN,Heucke SL,Giuggioli L,et al.2009.Experimental evidence for group hunting via eavesdropping in echolocating bats[J].Proceedings of the Royal Society B,276(1668):2721-2728.
Eckenweber M,Kn9rnschild M.2016.Responsiveness to conspecific distress calls is influenced by day-roost proximity in bats(Saccopteryx bilineata)[J].Royal Society Open Science,3(5):160151.DOI:10.1098/rsos.160151.
Fenton MB,Belwood JJ,Fullard JH,et al.1976.Responses of Myotis lucifugus(Chiroptera:Vespertilionidae)to calls of conspecifics and to other sounds[J].Canadian Journal of Zoology,54(9):1443-1448.
Fenton MB,Jacobs DS,Richardson EJ,et al.2004.Individual signatures in the frequency-modulated sweep calls of African large-eared,free-tailed bats Otomops martiensseni(Chiroptera:Molossidae)[J].Journal of Zoology,262(1):11-19.
Fenton MB.2003.Eavesdropping on the echolocation and social calls of bats[J].Mammal Review,33(3):193-204.
Fenton MB.2013.Questions,ideas and tools:lessons from bat echolocation[J].Animal Behaviour,85(5):869-879.
Fernandez AA,Fasel N,Kn9rnschild M,et al.2014.When bats are boxing:aggressive behaviour and communication in male Seba's short-tailed fruit bat[J].Animal Behaviour,98:149-156.
Furmankiewicz J,Ruczyński I,Urban R,et al.2011.Social calls provide tree-dwelling bats with information about the location of conspecifics at roosts[J].Ethology,117:480-489.
Ganesh A,Raghuram H,Nathan PT,et al.2010.Distress callinduced gene expression in the brain of the Indian short-nosed fruit bat,Cynopterus sphinx[J].Journal of Comparative Physiology A,196(2):155-164.
Geipel I,Jung K,Kalko EKV.2013.Perception of silent and motionless prey on vegetation by echolocation in the gleaning bat Micronycteris microtis[J].Proceedings of the Royal Society B,280(1754):20122830.DOI:10.1098/rspb.2012.2830.
Gillam E,Fenton MB.2016.Roles of acoustic social communication in the lives of bats[M]//Fenton BM,Grinnell DA,Popper NA,et al.Bat bioacoustics.New York:Springer:117-139.
Gillam EH.2007.Eavesdropping by bats on the feeding buzzes of conspecifics[J].Canadian Journal of Zoology,85(7):795-801.
Goerlitz HR,ter Hofstede HM,Zeale MR,et al.2010.An aerial-hawking bat uses stealth echolocation to counter moth hearing[J].Current Biology,20(17):1568-1572.
Gould E.1975.Neonatal vocalizations in bats of eight genera[J].Journal of Mammalogy,56(1):15-29.
Greif S,Siemers BM.2010.Innate recognition of water bodies in echolocating bats[J].Nature Communications,1(1):107.DOI:10.1038/ncomms1110.
Griffin DR.1944.Echolocation by blind men,bats and radar[J].Science,100(2609):589-590.
Hage SR,Jiang TL,Berquist SW,et al.2013.Ambient noise induces independent shifts in call frequency and amplitude within the Lombard effect in echolocating bats[J].Proceedings of the National Academy of Sciences of the United States of America,110(10):4063-4068.
Halfwerk W,Jones P,Taylor R,et al.2014.Risky ripples allow bats and frogs to eavesdrop on a multisensory sexual display[J].Science,343(2609):413-416.
Huang XB,Kanwal JS,Jiang TL,et al.2015.Situational and age-dependent decision making during life threatening distress in Myotis macrodactylus[J].PLo S ONE,10(7):e0132817.DOI:10.1371/journal.pone.0132817.
Hughes NK,Kelley JL,Banks PB.2012.Dangerous liaisons:the predation risks of receiving social signals[J].Ecology Letters,15(11):1326-1339.
Jakobsen L,Olsen MN,Surlykke A.2015.Dynamics of the echolocation beam during prey pursuit in aerial hawking bats[J].Proceedings of the National Academy of Sciences,112(26):8118-8123.
Jakobsen L,Ratcliffe JM,Surlykke A.2013.Convergent acoustic field of view in echolocating bats[J].Nature,493(7430):93-96.
Jiang TL,Metzner W,You YY,et al.2010.Variation in the resting frequency of Rhinolophus pusillus in mainland China:effect of climate and implications for conservation[J].Journal of the Acoustical Society of America,128(4):2204-2211.
Jin LR,Yang SL,Kimball RT,et al.2015.Do pups recognize maternal calls in pomona leaf-nosed bats,Hipposideros pomona?[J].Animal Behaviour,100:200-207.
Jones G,Siemers BM.2010.The communicative potential of bat echolocation pulses[J].Journal of Comparative Physiology A,197(5):447-457.
Jones G,Teeling E.2006.The evolution of echolocation in bats[J].Trends in Ecology&Evolution,21(3):149-156.
Jones G.1997.Acoustic signals and speciation:the roles of natural and sexual selection in the evolution of cryptic species[J].Advances in the Study of Behaviour,26(8):317-354.
Kalko EK,Schnitzler H.1998.How echolocating bats approach and acquire food[M]//Kunz TH,Racey PA.Bat biology and conservation.London:Smithsonian Institution Press:197-204.
Kanwal JS,Matsumura S,Ohlemiller K,et al.1994.Analysis of acoustic elements and syntax in communication sounds emitted by mustached bats[J].Journal of the Acoustical Society of America,96(3):1229-1254.
Kazial KA,Kenny TL,Burnett SC.2008.Little brown bats(Myotis lucifugus)recognize individual identity of conspecifics using sonar calls[J].Ethology,114(5):469-478.
Kazial KA,Masters WM.2004.Female big brown bats,Eptesicus fuscus,recognize sex from a caller's echolocation signals[J].Animal Behaviour,67(5):855-863.
Kn9rnschild M,Blüml S,Steidl P,et al.2017.Bat songs as acoustic beacons-male territorial songs attract dispersing females[J].Scientific Reports,7:13918.DOI:10.1038/s41598-017-14434-5.
Kn9rnschild M,Jung K,Nagy M,et al.2012.Bat echolocation calls facilitate social communication[J].Proceedings of the Royal Society B,279(1748):4827-4835.
Kn9rnschild M,Nagy M,Metz M,et al.2010.Complex vocal imitation during ontogeny in a bat[J].Biology Letters,6(2):156-159.
Kn9rnschild M,von Helversen O.2008.Nonmutual vocal mother-pup recognition in the greater sac-winged bat[J].Animal Behaviour,76(3):1001-1009.
Kober R,Schnitzler HU.1990.Information in sonar echoes of fluttering insects available for echolocating bats[J].Journal of the Acoustical Society of America,87(2):882-896.
Kunz TH,Fenton MB.2003.Bat ecology[M].Chicago:University of Chicago Press.
Lima SL,O'Keefe JM.2013.Do predators influence the behaviour of bats?[J].Biological Reviews,88(3):626-644.
Lin HJ,Kanwal JS,Jiang TL,et al.2015.Social and vocal behavior in adult greater tube-nosed bats(Murina leucogaster)[J].Zoology,118(3):192-202.
Liu Y,Feng J,Jiang YL,et al.2007.Vocalization development of greater horseshoe bat,Rhinolophus ferrumequinum(Rhinolophidae,Chiroptera)[J].Folia Zoologica,56(2):126-136.
Luo B,Huang XB,Li YY,et al.2017b.Social call divergence in bats:a comparative analysis[J].Behavioral Ecology,28(2):533-540.
Luo B,Jiang TL,Liu Y,et al.2013.Brevity is prevalent in bat short-range communication[J].Journal of Comparative Physiology A,199(4):325-333.
Luo B,Lu GJ,Chen K,et al.2017a.Social calls honestly signal female competitive ability in Asian particoloured bats[J].Animal Behaviour,127:101-108.
Luo JH,Goerlitz HR,Brumm H,et al.2015.Linking the sender to the receiver:vocal adjustments by bats to maintain signal detection in noise[J].Scientific Reports,5:18556.DOI:10.1038/srep18556.
Mller R,Kuc R.2000.Foliage echoes:a probe into the ecological acoustics of bat echolocation[J].Journal of the A-coustical Society of America,108(2):836-845.
Morell V.2014.When the bat sings[J].Science,344(6190):1334-1337.
Moss CF,Zagaeski M.1994.Acoustic information available to bats using frequency-modulated sounds for the perception of insect prey[J].Journal of the Acoustical Society of America,95(5):2745-2756.
Nelson JE.1965.Behaviour of Australian pteropodidae(Megacheroptera)[J].Animal Behaviour,13(4):544-557.
Obrist MK.1995.Flexible bat echolocation:the influence of individual,habitat and conspecifics on sonar signal design[J].Behavioral Ecology and Sociobiology,36(3):207-219.
Racey PA,Swift SM.1985.Feeding ecology of Pipistrellus pipistrellus(Chiroptera:Vespertilionidae)during pregnancy and lactation.Ⅰ.Foraging behaviour[J].Journal of Animal E-cology,54(1):205-215.
Roverud RC,Nitsche V,Neuweiler G.1991.Discrimination of wingbeat motion by bats,correlated with echolocation sound pattern[J].Journal of Comparative Physiology A,168(2):259-263.
RuppL,Clment G,Herrel A,et al.2015.Environmental constraints drive the partitioning of the soundscape in fishes[J].Proceedings of the National Academy of Sciences of the United States of America,112(19):6092-6097.
Russ J,Racey P,Jones G.1998.Intraspecific responses to distress calls of the pipistrelle bat,Pipistrellus pipistrellus[J].Animal Behaviour,55(3):705-713.
Russ J,Jones G,Mackie I,et al.2004.Interspecific responses to distress calls in bats(Chiroptera:Vespertilionidae):a function for convergence in call design?[J].Animal Behaviour,67(6):1005-1014.
Russo D,Cistrone L,Jones G.2012.Sensory ecology of water detection by bats:a field experiment[J].PLo S ONE,7(10):e48144.DOI:10.1371/journal.pone.0048144.
Rydell J.1986.Feeding territoriality in female northern bats,Eptesicus nilssoni[J].Ethology,72(4):329-337.
Schnitzler HU,Moss CF,Denzinger A.2003.From spatial orientation to food acquisition in echolocating bats[J].Trends in Ecology&Evolution,18(8):386-394.
Schuchmann M,Puechmaille SJ,Siemers BM.2012.Horseshoe bats recognise the sex of conspecifics from their echolocation calls[J].Acta Chiropterologica,14(1):161-166.
Servick K.2014.Eavesdropping on ecosystems[J].Science,343(6173):834-837.
Siemers BM,Schnitzler HU.2004.Echolocation signals reflect niche differentiation in five sympatric congeneric bat species[J].Nature,429(6992):657-661.
Simmons JA,Ferragamo MJ,Moss CF.1998.Echo-delay resolution in sonar images of the big brown bat,Eptesicus fuscus[J].Proceedings of the National Academy of Sciences of the United States of America,95(21):12647-12652.
Simmons JA,Houser D,Kloepper L.2014.Localization and classification of targets by echolocating bats and dolphins[M]//Surlykke A,Nachtigall EP,Fay RR,et al.Biosonar.New York:Springer:169-193.
Simon R,Holderied MW,Koch CU,et al.2011.Floral acoustics:conspicuous echoes of a dish-shaped leaf attract bat pollinators[J].Science,333(6042):631-633.
Suga N,Niwa H,Taniguchi I,et al.1987.The personalized auditory cortex of the mustached bat:adaptation for echolocation[J].Journal of Neurophysiology,58(4):643-654.
Sun KP,Luo L,Kimball RT,et al.2013.Geographic variation in the acoustic traits of greater horseshoe bats:testing the importance of drift and ecological selection in evolutionary processes[J].PLo S ONE,8(8):e70368.DOI:10.1371/journal.pone.0070368.
Thiagavel J,Cechetto C,Santana SE,et al.2018.Auditory opportunity and visual constraint enabled the evolution of echolocation in bats[J].Nature Communications,9(1):98.DOI:10.1038/s41467-017-02532-x.
Ulanovsky N,Moss CF.2008.What the bat's voice tells the bat's brain[J].Proceedings of the National Academy of Sciences of the United States of America,105(205):8491-8498.
Vannoni E,McElligott AG.2008.Low frequency groans indicate larger and more dominant fallow deer(Dama dama)males[J].PLo S ONE,3(9):e3113.DOI:10.1371/journal.pone.0003113.
Vehrencamp SL.2000.Handicap,index,and conventional signal elements of bird song[M]//Espmark Y,Amundsen T,Rosenqvist G.Animal signals:signalling and signal design in animal communication.Norway:Tapir Academic Press:277-300.
Voigt-Heucke SL,Taborsky M,Dechmann DKN.2010.A dual function of echolocation:bats use echolocation calls to identify familiar and unfamiliar individuals[J].Animal Behaviour,80(1):59-67.
von der Emde G,Schnitzler HU.1990.Classification of insects by echolocating greater horseshoe bats[J].Journal of Comparative Physiology A,167(3):423-430.
von Helversen D,von Helversen O.1999.Acoustic guide in bat-pollinated flower[J].Nature,398:759-760.
Wilkinson GS,Boughman JW.1998.Social calls coordinate foraging in greater spear-nosed bats[J].Animal Behaviour,55(2):337-350.
Yovel Y,Franz MO,Stilz P,et al.2008.Plant classification from bat-like echolocation signals[J].PLo S Computational Biology,4(3):e1000032.DOI.10.1371/journal.pcbi.1000032.
Yovel Y,Melcon ML,Franz MO,et al.2009b.The voice of bats:how greater mouse-eared bats recognize individuals based on their echolocation calls[J].PLo S Computational Biology,5(6):e1000400.DOI:10.1371/journal.pcbi.1000400.
Yovel Y,Stilz P,Franz MO,et al.2009a.What a plant sounds like:the statistics of vegetation echoes as received by echolocating bats[J].PLo S Computational Biology,5(7):e1000429.DOI:10.1371/journal.pcbi.1000429.