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Australian Journal of Zoology Australian Journal of Zoology Society
Evolutionary, molecular and comparative zoology
RESEARCH ARTICLE

Ghost bats exhibit informative daily and seasonal temporal patterns in the production of social vocalisations

Nicola Hanrahan https://orcid.org/0000-0001-8282-2590 A E , Christopher Turbill https://orcid.org/0000-0001-9810-7102 A , Kyle N. Armstrong https://orcid.org/0000-0002-3228-9182 B C , Anastasia H. Dalziell https://orcid.org/0000-0003-3602-0495 D and Justin A. Welbergen https://orcid.org/0000-0002-8085-5759 A
+ Author Affiliations
- Author Affiliations

A Hawkesbury Institute for the Environment, Western Sydney University, Richmond, NSW 2753, Australia.

B School of Biological Sciences, University of Adelaide, Adelaide, SA 5005, Australia.

C South Australian Museum, Adelaide, SA 5000, Australia.

D Centre for Sustainable Ecosystem Solutions, University of Wollongong, Wollongong, NSW 2522, Australia.

E Corresponding author. Email: n.hanrahan@westernsydney.edu.au

Australian Journal of Zoology 67(6) 305-315 https://doi.org/10.1071/ZO20055
Submitted: 19 June 2020  Accepted: 14 December 2020   Published: 5 January 2021

Abstract

The ghost bat (Macroderma gigas) is a colonial and highly vocal species that is impacted by human visitation of caves. The ability to document behaviours inside the roost by recording vocalisations could provide an important new tool for the management of this disturbance-prone species by removing the need for in-person confirmation of reproductive activity, and, in turn, identifying roosts of conservation importance. To assess whether vocalisations are indicators of daily and seasonal behavioural events, we aimed to determine whether total vocal activity significantly varied by time of day and time of year and, further, how the relative frequencies of occurrence of three common social vocalisations (‘Chirp-trill’, ‘Squabble’ and ‘Ultrasonic Social’) aligned with previously reported seasonal reproductive behaviour. We recorded sound inside the largest known maternity roost, extracted all vocal signals and classified them into types using semiautomated methods. Total vocal activity varied significantly by time of day and time of year, peaking around sunrise and sunset, and during the mating and nursing seasons. The relative frequencies of occurrence of vocalisation types varied significantly seasonally, with the Chirp-trill and Squabble produced most during the mating season and first flight periods, whereas the Ultrasonic Social peaked during parturition and weaning periods. This timing aligns with a previously suggested vocalisation function, providing further evidence that these signals are important in mating and maternity behaviours. Further, this suggests that peaks in the relative frequency of occurrence of distinct social vocalisations may act as indicators of in-roost reproductive and pup development behaviours and provides a low-disturbance, semiautomated method for using long-term acoustic recordings to study and monitor behaviour in this sensitive species.

Keywords: Chiroptera, Macroderma gigas, threatened species, social communication, in-roost behaviour, acoustic, reproduction, semiautomated classifier.


References

Adams, R. A., and Snode, E. R. (2015). Differences in the male mating calls of co-occurring epauletted fruit bat species (Chiroptera, Pteropodidae, Epomophorus wahlbergi and Epomophorus crypturus) in Kruger National Park, South Africa. Zoological Studies (Taipei, Taiwan) 54, 15.
Differences in the male mating calls of co-occurring epauletted fruit bat species (Chiroptera, Pteropodidae, Epomophorus wahlbergi and Epomophorus crypturus) in Kruger National Park, South Africa.Crossref | GoogleScholarGoogle Scholar |

Adams, A. M., Jantzen, M. K., Hamilton, R. M., and Fenton, M. B. (2012). Do you hear what I hear? Implications of detector selection for acoustic monitoring of bats. Methods in Ecology and Evolution 3, 992–998.
Do you hear what I hear? Implications of detector selection for acoustic monitoring of bats.Crossref | GoogleScholarGoogle Scholar |

Andrews, M. M., Hodnett, A. M., and Andrews, P. T. (2017). Social activity of lesser horseshoe bats (Rhinolophus hipposideros) at nursery roosts and a hibernaculum in north Wales, U.K. Acta Chiropterologica 19, 161–174.
Social activity of lesser horseshoe bats (Rhinolophus hipposideros) at nursery roosts and a hibernaculum in north Wales, U.K.Crossref | GoogleScholarGoogle Scholar |

Armstrong, K., and Anstee, S. D. (2000). The ghost bat in the Pilbara: 100 years on. Australian Mammalogy 22, 93–101.
The ghost bat in the Pilbara: 100 years on.Crossref | GoogleScholarGoogle Scholar |

Arnold, B. D., and Wilkinson, G. S. (2011). Individual specific contact calls of pallid bats (Antrozous pallidus) attract conspecifics at roosting sites. Behavioral Ecology and Sociobiology 65, 1581–1593.
Individual specific contact calls of pallid bats (Antrozous pallidus) attract conspecifics at roosting sites.Crossref | GoogleScholarGoogle Scholar |

Balcombe, J. P. (1990). Vocal recognition of pups by mother Mexican free-tailed bats, Tadarida brasiliensis mexicana. Animal Behaviour 39, 960–966.
Vocal recognition of pups by mother Mexican free-tailed bats, Tadarida brasiliensis mexicana.Crossref | GoogleScholarGoogle Scholar |

Barton, K. (2018). MuMIn: multi-model inference. R package version 1.42.1. Available at: https://CRAN.R-project.org/package=MuMIn [accessed 12 January 2019].

Bastian, A., and Schmidt, S. (2008). Affect cues in vocalizations of the bat, Megaderma lyra, during agonistic interactions. The Journal of the Acoustical Society of America 124, 598–608.
Affect cues in vocalizations of the bat, Megaderma lyra, during agonistic interactions.Crossref | GoogleScholarGoogle Scholar | 18647002PubMed |

Behr, O., Von Helversen, O., Heckel, G., Nagy, M., Voigt, C. C., and Mayer, F. (2006). Territorial songs indicate male quality in the sac-winged bat Saccopteryx bilineata (Chiroptera, Emballonuridae). Behavioral Ecology 17, 810–817.
Territorial songs indicate male quality in the sac-winged bat Saccopteryx bilineata (Chiroptera, Emballonuridae).Crossref | GoogleScholarGoogle Scholar |

Binder, C. M., and Hines, P. C. (2014). Automated aural classification used for inter-species discrimination of cetaceans. The Journal of the Acoustical Society of America 135, 2113–2125.
Automated aural classification used for inter-species discrimination of cetaceans.Crossref | GoogleScholarGoogle Scholar | 25235008PubMed |

Bioacoustics Research Program (2014). Raven Pro: interactive sound analysis software (version 1.5) [computer software]. The Cornell Lab of Ornithology. Available at: http://www.birds.cornell.edu/raven [accessed 10 December 2015].

Bittle, M., and Duncan, A. (2013). A review of current marine mammal detection and classification algorithms for use in automated passive acoustic monitoring. In ‘Annual Conference of the Australian Acoustical Society 2013. Acoustics 2013: Science, Technology and Amenity‘. pp. 208–215. Available at: https://www.scopus.com/inward/record.uri?eid=2-s2.0-84903825224&partnerID=40&md5=cefebf4f7a9b0b2d26be656b92d8af37 [accessed 23 June 2019].

Bohn, K. M., Wilkinson, G. S., and Moss, C. F. (2007). Discrimination of infant isolation calls by female greater spear-nosed bats, Phyllostomus hastatus. Animal Behaviour 73, 423–432.
Discrimination of infant isolation calls by female greater spear-nosed bats, Phyllostomus hastatus.Crossref | GoogleScholarGoogle Scholar | 18311319PubMed |

Bohn, K. M., Schmidt-French, B., Ma, S. T., and Pollak, G. D. (2008). Syllable acoustics, temporal patterns, and call composition vary with behavioral context in Mexican free-tailed bats. The Journal of the Acoustical Society of America 124, 1838–1848.
Syllable acoustics, temporal patterns, and call composition vary with behavioral context in Mexican free-tailed bats.Crossref | GoogleScholarGoogle Scholar | 19045674PubMed |

Bohn, K. M., Smarsh, G. C., and Smotherman, M. (2013). Social context evokes rapid changes in bat song syntax. Animal Behaviour 85, 1485–1491.
Social context evokes rapid changes in bat song syntax.Crossref | GoogleScholarGoogle Scholar |

Carter, G., Logsdon, R., Arnold, B. D., Menchaca, A., and Medellin, R. A. (2012). Adult vampire bats produce contact calls when isolated: acoustic variation by species, population, colony, and individual. PLoS One 7, e38791.
Adult vampire bats produce contact calls when isolated: acoustic variation by species, population, colony, and individual.Crossref | GoogleScholarGoogle Scholar | 22719947PubMed |

Carter, G., Schoeppler, D., Manthey, M., Knörnschild, M., and Denzinger, A. (2015). Distress calls of a fast-flying bat (Molossus molossus) provoke inspection flights but not cooperative mobbing. PLoS One 10, e0136146.
Distress calls of a fast-flying bat (Molossus molossus) provoke inspection flights but not cooperative mobbing.Crossref | GoogleScholarGoogle Scholar | 26445502PubMed |

Charif, R. A., Waack, A. M., and Strickman, L. M. (2010). ‘Raven Pro 1.4 User’s Manual.’ (Cornell Lab of Ornithology: Ithaca, NY.)

Chua, M. A. H., and Aziz, S. A. (2019). Into the light: atypical diurnal foraging activity of Blyth’s horseshoe bat, Rhinolophus lepidus (Chiroptera: Rhinolophidae) on Tioman Island, Malaysia. Mammalia 83, 78–83.
Into the light: atypical diurnal foraging activity of Blyth’s horseshoe bat, Rhinolophus lepidus (Chiroptera: Rhinolophidae) on Tioman Island, Malaysia.Crossref | GoogleScholarGoogle Scholar |

Churchill, S. (2008). ‘Australian Bats.’ (Allen & Unwin.)

Clement, M. J., and Kanwal, J. S. (2012). Simple syllabic calls accompany discrete behavior patterns in captive Pteronotus parnellii: an illustration of the motivation–structure hypothesis. The Scientific World Journal 2012, 12869510.1100/2012/128695.10.1100/2012/128695

Cook, G. D., and Heerdegen, R. G. (2001). Spatial variation in the duration of the rainy season in monsoonal Australia. International Journal of Climatology 21, 1723–1732.
Spatial variation in the duration of the rainy season in monsoonal Australia.Crossref | GoogleScholarGoogle Scholar |

Douglas, A. D. A. (1967). The natural history of the ghost bat, Macroderma gigas (Microchiroptera, Megadermatidae) in Western Australia. West Australian Naturalist 10, 125–138.

Eckenweber, M., and Knörnschild, M. (2016). Responsiveness to conspecific distress calls is influenced by day-roost proximity in bats (Saccopteryx bilineata). Royal Society Open Science 3, 160151.
Responsiveness to conspecific distress calls is influenced by day-roost proximity in bats (Saccopteryx bilineata).Crossref | GoogleScholarGoogle Scholar | 27293797PubMed |

Engler, S., Rose, A., and Knörnschild, M. (2017). Isolation call ontogeny in bat pups (Glossophaga soricina). Behaviour 154, 267–286.
Isolation call ontogeny in bat pups (Glossophaga soricina).Crossref | GoogleScholarGoogle Scholar |

Erkert, H. G. (1978). Sunset-related timing of flight activity in neotropical bats. Oecologia 37, 59–67.
Sunset-related timing of flight activity in neotropical bats.Crossref | GoogleScholarGoogle Scholar | 28309285PubMed |

Esser, K.-H., and Schmidt, U. (1989). Mother–infant communication in the lesser spear-nosed bat Phyllostomus discolor (Chiroptera, Phyllostomidae) – evidence for acoustic learning. Ethology 82, 156–168.
Mother–infant communication in the lesser spear-nosed bat Phyllostomus discolor (Chiroptera, Phyllostomidae) – evidence for acoustic learning.Crossref | GoogleScholarGoogle Scholar |

Fernandez, A. A., and Knörnschild, M. (2017). Isolation calls of the bat Saccopteryx bilineata encode multiple messages. Animal Behavior and Cognition 4, 169–186.
Isolation calls of the bat Saccopteryx bilineata encode multiple messages.Crossref | GoogleScholarGoogle Scholar |

Fox, J., Weisberg, S., Adler, D., Bates, D., Baud-Bovy, G., Ellison, S., Firth, D., Friendly, M., Gorjanc, G., and Graves, S. (2012). Package ‘car’. R Foundation for Statistical Computing, Vienna.

Gadziola, M. A., Grimsley, J. M. S., Faure, P. A., and Wenstrup, J. J. (2012). Social vocalizations of big brown bats vary with behavioral context. PLoS One 7, e44550.
Social vocalizations of big brown bats vary with behavioral context.Crossref | GoogleScholarGoogle Scholar | 22970247PubMed |

Gillam, E. H., and Chaverri, G. (2012). Strong individual signatures and weaker group signatures in contact calls of Spix’s disc-winged bat, Thyroptera tricolor. Animal Behaviour 83, 269–276.
Strong individual signatures and weaker group signatures in contact calls of Spix’s disc-winged bat, Thyroptera tricolor.Crossref | GoogleScholarGoogle Scholar |

Goymann, W., Leippert, D., and Hofer, H. (1999). Parturition, parental behaviour, and pup development in Indian false vampire bats, Megaderma lyra. Zeitschrift für Säugetierkunde 64, 321–331.

Griffin, D. R., Webster, F. A., and Michael, C. R. (1960). The echolocation of flying insects by bats. Animal Behaviour 8, 141–154.
The echolocation of flying insects by bats.Crossref | GoogleScholarGoogle Scholar |

Griffiths, R. W., and Griffiths, R. W. (2007). Activity patterns of long‐tailed bats (Chalinolobus tuberculatus) in a rural landscape, south Canterbury, New Zealand. New Zealand Journal of Zoology 34, 247–258.
Activity patterns of long‐tailed bats (Chalinolobus tuberculatus) in a rural landscape, south Canterbury, New Zealand.Crossref | GoogleScholarGoogle Scholar |

Guppy, A., Coles, R., and Pettigrew, J. (1985). Echolocation and acoustic communication in the Australian ghost bat, Macroderma gigas (Microchiroptera: Megadermatidae). Australian Mammalogy 8, 299–308.

Hanrahan, N. (2020). The acoustic ecology of the ghost bat (Macroderma gigas): form, function and applied uses of vocalisations. Ph.D. Thesis, Western Sydney University, Sydney.

Hayes, J. P. (1997). Temporal variation in activity of bats and the design of echolocation-monitoring studies. Journal of Mammalogy 78, 514–524.
Temporal variation in activity of bats and the design of echolocation-monitoring studies.Crossref | GoogleScholarGoogle Scholar |

Hengjan, Y., Iida, K., Doysabas, K. C. C., Phichitrasilp, T., Ohmori, Y., and Hondo, E. (2017). Diurnal behavior and activity budget of the golden-crowned flying fox (Acerodon jubatus) in the Subic Bay forest reserve area, the Philippines. The Journal of Veterinary Medical Science 79, 1667–1674.
Diurnal behavior and activity budget of the golden-crowned flying fox (Acerodon jubatus) in the Subic Bay forest reserve area, the Philippines.Crossref | GoogleScholarGoogle Scholar | 28804092PubMed |

Holland, R. A., Meyer, C. F. J., Kalko, E. K. V., Kays, R., and Wikelski, M. (2011). Emergence time and foraging activity in Pallas’ mastiff bat, Molossus molossus (Chiroptera: Molossidae) in relation to sunset/sunrise and phase of the moon. Acta Chiropterologica 13, 399–404.
Emergence time and foraging activity in Pallas’ mastiff bat, Molossus molossus (Chiroptera: Molossidae) in relation to sunset/sunrise and phase of the moon.Crossref | GoogleScholarGoogle Scholar |

Hope, P. R., and Jones, G. (2013). An entrained circadian cycle of peak activity in a population of hibernating bats. Journal of Mammalogy 94, 497–505.
An entrained circadian cycle of peak activity in a population of hibernating bats.Crossref | GoogleScholarGoogle Scholar |

Huang, X., Metzner, W., Zhang, K., Wang, Y., Luo, B., Sun, C., Jiang, T., and Feng, J. (2018). Acoustic similarity elicits responses to heterospecific distress calls in bats (Mammalia: Chiroptera). Animal Behaviour 146, 143–154.
Acoustic similarity elicits responses to heterospecific distress calls in bats (Mammalia: Chiroptera).Crossref | GoogleScholarGoogle Scholar |

Hughes, P. M., Rayner, J. M. V., and Jones, G. (1995). Ontogeny of ‘true’flight and other aspects of growth in the bat Pipistrellus pipistrellus. Journal of Zoology 236, 291–318.
Ontogeny of ‘true’flight and other aspects of growth in the bat Pipistrellus pipistrellus.Crossref | GoogleScholarGoogle Scholar |

Jiang, T., Long, Z., Ran, X., Zhao, X., Xu, F., Qiu, F., Kanwal, J. S., and Feng, J. (2016). Using sounds for making decisions: greater tube-nosed bats prefer antagonistic calls over non-communicative sounds when feeding. Biology Open 5, 1864–1868.
Using sounds for making decisions: greater tube-nosed bats prefer antagonistic calls over non-communicative sounds when feeding.Crossref | GoogleScholarGoogle Scholar | 27815241PubMed |

Kaňuch, P. (2007). Evening and morning activity schedules of the noctule bat (Nyctalus noctula) in western Carpathians. Mammalia 71, 126–130.
Evening and morning activity schedules of the noctule bat (Nyctalus noctula) in western Carpathians.Crossref | GoogleScholarGoogle Scholar |

Kawakita, S., and Ichikawa, K. (2019). Automated classification of bees and hornet using acoustic analysis of their flight sounds. Apidologie 50, 71–79.
Automated classification of bees and hornet using acoustic analysis of their flight sounds.Crossref | GoogleScholarGoogle Scholar |

Klose, S. M., Welbergen, J. A., Goldizen, A. W., and Kalko, E. K. V. (2009). Spatio-temporal vigilance architecture of an Australian flying-fox colony. Behavioral Ecology and Sociobiology 63, 371–380.
Spatio-temporal vigilance architecture of an Australian flying-fox colony.Crossref | GoogleScholarGoogle Scholar |

Knörnschild, M., and von Helversen, O. (2008). Nonmutual vocal mother–pup recognition in the greater sac-winged bat. Animal Behaviour 76, 1001–1009.
Nonmutual vocal mother–pup recognition in the greater sac-winged bat.Crossref | GoogleScholarGoogle Scholar |

Knörnschild, M., Behr, O., and von Helversen, O. (2006). Babbling behavior in the sac-winged bat (Saccopteryx bilineata). Naturwissenschaften 93, 451–454.
Babbling behavior in the sac-winged bat (Saccopteryx bilineata).Crossref | GoogleScholarGoogle Scholar | 16736178PubMed |

Knörnschild, M., Nagy, M., Metz, M., Mayer, F., and von Helversen, O. (2012). Learned vocal group signatures in the polygynous bat Saccopteryx bilineata. Animal Behaviour 84, 761–769.
Learned vocal group signatures in the polygynous bat Saccopteryx bilineata.Crossref | GoogleScholarGoogle Scholar |

Knörnschild, M., Blüml, S., Steidl, P., Eckenweber, M., and Nagy, M. (2017). Bat songs as acoustic beacons – male territorial songs attract dispersing females. Scientific Reports 7, 13918.
Bat songs as acoustic beacons – male territorial songs attract dispersing females.Crossref | GoogleScholarGoogle Scholar | 29066803PubMed |

Kulzer, E., Nelson, J. E., McKean, J. L., and Moehres, F. P. (1984). Prey-catching behavior and echolocation in the Australian ghost bat Macroderma gigas Microchiroptera Megadermatidae. Australian Mammalogy 7, 37–50.

MacSwiney, G., Clarke, F. M., and Racey, P. A. (2008). What you see is not what you get: the role of ultrasonic detectors in increasing inventory completeness in Neotropical bat assemblages. Journal of Applied Ecology 45, 1364–1371.
What you see is not what you get: the role of ultrasonic detectors in increasing inventory completeness in Neotropical bat assemblages.Crossref | GoogleScholarGoogle Scholar |

Markus, N., and Blackshaw, J. K. (2002). Behaviour of the black flying fox Pteropus alecto: 1. An ethogram of behaviour, and preliminary characterisation of mother–infant interactions. Acta Chiropterologica 4, 137–152.
Behaviour of the black flying fox Pteropus alecto: 1. An ethogram of behaviour, and preliminary characterisation of mother–infant interactions.Crossref | GoogleScholarGoogle Scholar |

Milne, D. J. (2002). Key to the bat calls of the Top End of the Northern Territory. Parks and Wildlife Commission of the Northern Territory.

Möhres, F. P., and Kulzer, E. (1957). Megaderma – ein konvergenter Zwischentyp der Ultraschallpeilung bei Fledermäusen. Naturwissenschaften 44, 21–22.
Megaderma – ein konvergenter Zwischentyp der Ultraschallpeilung bei Fledermäusen.Crossref | GoogleScholarGoogle Scholar |

Montero, K., and Gillam, E. H. (2015). Geographic variation in contact calls emitted by a leaf-roosting bat suggest distinct modes of vocal transmission. The Journal of the Acoustical Society of America 138, 1932.
Geographic variation in contact calls emitted by a leaf-roosting bat suggest distinct modes of vocal transmission.Crossref | GoogleScholarGoogle Scholar |

Muñoz-Romo, M. (2006). Ethogram and diurnal activities of a colony of Artibeus lituratus (Phyllostomidae: Stenodermatinae). Acta Chiropterologica 8, 231–238.
Ethogram and diurnal activities of a colony of Artibeus lituratus (Phyllostomidae: Stenodermatinae).Crossref | GoogleScholarGoogle Scholar |

Neuweiler, G. (1989). Foraging ecology and audition in echolocating bats. Trends in Ecology & Evolution 4, 160–166.
Foraging ecology and audition in echolocating bats.Crossref | GoogleScholarGoogle Scholar |

Park, K. J., Jones, G., and Ransome, R. D. (1999). Winter activity of a population of greater horseshoe bats (Rhinolophus ferrumequinum). Journal of Zoology 248, 419–427.
Winter activity of a population of greater horseshoe bats (Rhinolophus ferrumequinum).Crossref | GoogleScholarGoogle Scholar |

Petrželková, K. J., Downs, N. C., Zukal, J., and Racey, P. A. (2006). A comparison between emergence and return activity in pipistrelle bats Pipistrellus pipistrellus and P. pygmaeus. Acta Chiropterologica 8, 381–390.
A comparison between emergence and return activity in pipistrelle bats Pipistrellus pipistrellus and P. pygmaeus.Crossref | GoogleScholarGoogle Scholar |

Prat, Y., Taub, M., and Yovel, Y. (2016). Everyday bat vocalizations contain information about emitter, addressee, context, and behavior. Scientific Reports 6, 39419.
Everyday bat vocalizations contain information about emitter, addressee, context, and behavior.Crossref | GoogleScholarGoogle Scholar | 28005079PubMed |

Prat, Y., Taub, M., Pratt, E., and Yovel, Y. (2017). An annotated dataset of Egyptian fruit bat vocalizations across varying contexts and during vocal ontogeny. Scientific Data 4, 170143.
An annotated dataset of Egyptian fruit bat vocalizations across varying contexts and during vocal ontogeny.Crossref | GoogleScholarGoogle Scholar | 28972574PubMed |

R Core Team (2017). ‘R: A Language and Environment for Statistical Computing.’ (R Foundation for Statistical Computing: Vienna, Austria.)

Reynolds, D. S., Shoemaker, K., Von Oettingen, S., and Najjar, S. (2017). High rates of winter activity and arousals in two New England bat species: implications for a reduced White-nose Syndrome impact? Northeastern Naturalist 24, B188–B208.
High rates of winter activity and arousals in two New England bat species: implications for a reduced White-nose Syndrome impact?Crossref | GoogleScholarGoogle Scholar |

Ripley, B., Venables, W., and Ripley, M. B. (2016). Package ‘nnet’. R Package version 7, 3–12.

Russ, J. M., Jones, G., Mackie, I. J., and Racey, P. A. (2004). Interspecific responses to distress calls in bats (Chiroptera: Vespertilionidae): a function for convergence in call design? Animal Behaviour 67, 1005–1014.
Interspecific responses to distress calls in bats (Chiroptera: Vespertilionidae): a function for convergence in call design?Crossref | GoogleScholarGoogle Scholar |

Russo, D., Cistrone, L., Garonna, A. P., and Jones, G. (2011a). The early bat catches the fly: daylight foraging in soprano pipistrelles. Mammalian Biology 76, 87–89.
The early bat catches the fly: daylight foraging in soprano pipistrelles.Crossref | GoogleScholarGoogle Scholar |

Russo, D., Maglio, G., Rainho, A., Meyer, C. F. J., and Palmeirim, J. M. (2011b). Out of the dark: diurnal activity in the bat Hipposideros ruber on São Tomé Island (West Africa). Mammalian Biology 76, 701–708.
Out of the dark: diurnal activity in the bat Hipposideros ruber on São Tomé Island (West Africa).Crossref | GoogleScholarGoogle Scholar |

Schulz, M., and Menkhorst, K. (1986). Roost preferences of cave-dwelling bats at Pine Creek, Northern Territory. Macroderma 2, 2–7.

Smarsh, G. C. (2015). Singing away from home: song is used to create and defend foraging territories in the African megadermatid bat, Cardioderma cor. The Journal of the Acoustical Society of America 138, 1932.
Singing away from home: song is used to create and defend foraging territories in the African megadermatid bat, Cardioderma cor.Crossref | GoogleScholarGoogle Scholar |

Thomson, S. C., Brooke, A. P., and Speakman, J. R. (1998). Diurnal activity in the Samoan flying fox, Pteropus samoensis. Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences 353, 1595–1606.
Diurnal activity in the Samoan flying fox, Pteropus samoensis.Crossref | GoogleScholarGoogle Scholar |

Tidemann, C. R., Priddel, D. M., Nelson, J. E., and Pettigrew, J. D. (1985). Foraging behaviour of the Australian ghost bat, Macroderma gigas (Microchiroptera: Megadermatidae). Australian Journal of Zoology 33, 705–713.
Foraging behaviour of the Australian ghost bat, Macroderma gigas (Microchiroptera: Megadermatidae).Crossref | GoogleScholarGoogle Scholar |

Toop, J. (1985). Habitat requirements, survival strategies and ecology of the ghost bat, Macroderma gigas Dobson (Microchiroptera, Megadermatidae) in central coastal Queensland. Macroderma 1, 37–41.

Turbill, C., Law, B. S., and Geiser, F. (2003). Summer torpor in a free-ranging bat from subtropical Australia. Journal of Thermal Biology 28, 223–226.
Summer torpor in a free-ranging bat from subtropical Australia.Crossref | GoogleScholarGoogle Scholar |

Walter, M. H., and Schnitzler, H.-U. (2019). Spectral call features provide information about the aggression level of greater mouse-eared bats (Myotis myotis) during agonistic interactions. Bioacoustics 28, 1–25.
Spectral call features provide information about the aggression level of greater mouse-eared bats (Myotis myotis) during agonistic interactions.Crossref | GoogleScholarGoogle Scholar |

Welbergen, J. A. (2006). Timing of the evening emergence from day roosts of the grey-headed flying fox, Pteropus poliocephalus: the effects of predation risk, foraging needs, and social context. Behavioral Ecology and Sociobiology 60, 311.
Timing of the evening emergence from day roosts of the grey-headed flying fox, Pteropus poliocephalus: the effects of predation risk, foraging needs, and social context.Crossref | GoogleScholarGoogle Scholar |

Welbergen, J. A. (2008). Variation in twilight predicts the duration of the evening emergence of fruit bats from a mixed-species roost. Animal Behaviour 75, 1543–1550.
Variation in twilight predicts the duration of the evening emergence of fruit bats from a mixed-species roost.Crossref | GoogleScholarGoogle Scholar |

Woinarski, J. C. Z., Burbidge, A. A., and Harrison, P. L. (2014). ‘The Action Plan For Australian Mammals 2012.’ (CSIRO Publishing: Melbourne.)

Wu, X., Pang, Y., Luo, B., Wang, M., and Feng, J. (2019). Function of distress calls in least horseshoe bats: a field study using playback experiments. Acta Chiropterologica 20, 455–464.
Function of distress calls in least horseshoe bats: a field study using playback experiments.Crossref | GoogleScholarGoogle Scholar |

Yates, M. D., and Muzika, R. M. (2006). Effect of forest structure and fragmentation on site occupancy of bat species in Missouri Ozark forests. The Journal of Wildlife Management 70, 1238–1248.
Effect of forest structure and fragmentation on site occupancy of bat species in Missouri Ozark forests.Crossref | GoogleScholarGoogle Scholar |

Zhao, Z., Zhang, S.-H., Xu, Z.-Y., Bellisario, K., Dai, N.-H., Omrani, H., and Pijanowski, B. C. (2017). Automated bird acoustic event detection and robust species classification. Ecological Informatics 39, 99–108.
Automated bird acoustic event detection and robust species classification.Crossref | GoogleScholarGoogle Scholar |