Dr. Annette Denzinger
Prof. em. Dr. Hans-Ulrich Schnitzler

We investigate the echolocation behaviour of bats and dolphins and the acoustic communication behaviour of vertebrates with the main focus on echolocation in bats.

Bats have to solve a multitude of echolocation tasks depending on where they fly and forage, what they eat and how they acquire food. In the course of evolution bats evolved a repertoire of species-specific signal types and behaviours that have been adapted to specific echolocation tasks. In a tightly coupled action-perception loop bats continuously emit echolocation signals, evaluate the incoming echo information to form an inner representation of the outside world, and select the appropriate signal types and emission pattern to reach an intended behavioural goal.

We compare and analyze the echolocation signals and behaviours of various bat species in the laboratory and worldwide in the field to understand their adaptive value.

More group members

Graduate students

Simone Götze Dipl. Biol. -Niche partitioning and flight paths of foraging bat

Sonja Sändig, Dipl. Biol. -Variation of emission SPL in insect catching bats

Johanna Flick, Dipl. Biol. -Echolocation behaviour of gleaning bats

Melanie Hüpkes Dipl. Biol. -Species specific frequency channels in European pipistrelle bats

Diana Schöppler, Dipl. Biol. -Doppler shift compensation in rhinolophids and hipposiderids bats of Vietnam

Anna-Maria Seibert, Dipl. Biol. -Acoustic scanning behavior of free-ranging echolocating bats

Susanne Walther-Voolstra, Dipl. Biol. -Landmark orientation in the greater mouse-eared bat.

Diploma/Master students

Walter, Michael


Publications since 2008


Siebert AM, Koblitz J C, Denzinger A and Schnitzler H-U (2015)
Bidirectional echolocation in the bat B. barbastellus: different signals of low source level are emitted upward through the nose and downward through the mouth.
PLoS ONE 10(9): e0135590. doi:10.1371/journal.pone.0135590

Carter G, Schoeppler D, Manthey M, Knoernschild M and Dennzinger A (2015)
Distress calls of a fast-flying bat M.molossus provoke inspection flights but not cooperative mobbing.
PLoS ONE 10(9): e0136146. doi:10.1371/journal.pone.0136146


Denzinger A and Schnitzler HU (2013)
Batguilds,a concept to classify the highly diverse foraging and echolocation behaviors of microchiropteran bats
Frontiers in Physiol 4/164:1-15

Seibert AM, Koblitz J C, Denzinger A, Schnitzler HU.(2013)
Scanning Behavior in Echolocating Common PipistrelleBats (Pipistrellus pipistrellus)
PLOS ONE April 2013 ;Volume 8 ; Issue 4; e60752</span>


Stilz P, Schnitzler H-U. (2012)
Estimation of the acoustic range of bat ecolocation for extended targets
J. Acoust. Soc. Am 132/3:1765-1775

Siemers B, Kriner E, Kaipf I, Simon M and Greif S (2012)
Bats eavesdrop on the sound of copulating flies
Cur. Biol. 11(14)

Thong VD, Denzinger A, Dietz C, Bates PJJ, Furey NM, Racey PA, Schnitzler HU (2012)
Coelops frithii, a hipposiderid bat with a specialized echolocation system.
Proc. R. Soc. B

Thong, VD, Dietz C, Denzinger A, Bates PJJ, Puechmaille SJ, Callou C, Schnitzler HU (2012)
Resolving a mammal mystery: the identity of Paracoelops megalotis (Chiroptera: Hipposideridae)
Zootaxa 3505: 75-85

Thong VD, Puechmaille SJ, Denzinger A, Dietz C, Csorba G, Bates PJJ, Teeling EC, Schnitzler HU (2011)
A new species of Hipposideros (Chiroptera: Hipposideridae) from Vietnam.
Journal Mamm.93:1-11

Thong VD, Puechmaille SJ, Denzinger A, Bates PJJ, Dietz C, Csorba G, Soisook P, Teeling EC, Matsumura S, Furey NM, Schnitzler HU (2011)
Systematics of the Hipposideros turpis complex with a description of a new subspecies from Vietnam.
Mamm. Review 42:166-192


Koblitz J, Stilz P, Pflästerer W, Melcón ML, Schnitzler HU (2011) Source level reduction and sonar beam aiming in landing big brown bats (Eptesicus fuscus).
J Acoust Soc Am 130:3090-3099

Koblitz J, Wahlberg M, Stilz P, Madsen P, Beedholm K, Schnitzler HU (2011)
Asymmetry and dynamics of a narrow sonar beam in an echolocating harbour porpoise. Journal of the Acoustical Society of America (in press)

Koselj K, Schnitzler HU, Siemers BM.
Horseshoe bats make adaptive prey selection decisions, informed by echo cues.
Proceedings of the Royal Society B, doi:10.1098/rspb.2010.2793

Melcón ML, Yovel Y, Denzinger A, Schnitzler HU (2011)
How greater mouse-eared bats deal with ambiguous echoic scenes.J Comp Physiol A 197:505-514

Schnitzler HU, Denzinger A (2011)
Auditory fovea and Doppler shift compensation: adaptations for flutter detection in echolocating bats using CF-FM signals.J Comp Physiol A 197:541-559

Thong VD, Dietz C, Denzinger A, Bates PJJ, Furey NM, Csorba G, Hoye G, Thuy LD, Schnitzler HU (2011)
Further records of Murina tiensa from Vietnam with first information on its echolocation calls.
Hystrix It J Mamm 22:129-138

Yovel Y, Franz MO, Stilz P, Schnitzler HU (2011)
Complex echo classification by echo-locating bats.
J Comp Physiol A 197:475-490


Baumann-Pickering S, Wiggins SM, Hildebrand JA, Roch MA, Schnitzler HU (2010)
Discriminating features of echolocation clicks of melon-headed whales (Peponocephala electra), bottlenose dolphins (Tursiops truncatus), and Gray's spinner dolphins (Stenella longirostris longirostris).
J Acoust Soc Amer 128 (4):2212-2224

Baumann-Pickering S, Wiggins SM, Roth EH, Roch MA, Schnitzler HU, Hildebrand JA (2010)
Echolocation signals of a beaked whale at Palmyra Atoll.
J Acoust Soc Amer 127: 3790-3799

Koblitz JC, Stilz P, Schnitzler HU (2010)
Source levels of echolocation signals vary in correlation with wingbeat cycle in landing big brown bats (Eptesicus fuscus). J Experim Biol 213: 3263-3268


Fooks AR, Johnson N, Müller T, Vos A, Mansfield K, Hicks D, Nunez A, Freudling C, Kaipf I, Denzinger A, Franka R, Rupprecht CE (2009)
Detection of high levels of European bat lyssavirus Type-1 viral RNA in the thyroid gland of experimentally infected Eptesicus fuscus bats.
Zoonoses Public Health. 56: 270–277

Freuling C, Vos A, Johnson N, Kaipf I, Denzinger A, Neubert L, Mansfield K, Hicks D, Nuez A, Tordo N, Rupprecht C, Fooks AR, Müller T (2009)
Experimental infection of Serotine bats (Eptesicus serotinus) with European bat lyssavirus type 1a (EBLV-1a).
Journal of General Virology 90:2493-2502

Melcón ML, Schnitzler HU, Denzinger,A (2009)
Variability of the approach phase of landing echolocating Greater Mouse-eared bats.
J Comp Physiol A 195:69-77

Sümer S, Denzinger A, Schnitzler HU (2009)
Spatial unmasking in the echolocating Big Brown Bat, Eptesicus fuscus.
J Comp Physiol A 195:463-472

Verfuss UK, Miller LA, Pilz PKD, Schnitzler HU (2009)
Echolocation by two foraging harbour porpoises (Phocoena phocoena).
J Experim Biol 212:823-834

Verfuss UK, Miller LA, Pilz, PKD, Schnitzler HU (2009)
How porpoises track prey with echolocation.
J Experim Biol 212:I-II

Yovel Y, Melcón ML, Franz MO, Denzinger A, Schnitzler HU (2009).
The voice of bats: How greater mouse-eared bats recognize individuals based on their echolocation calls.
PLoS Comput Biol 5(6): e1000400.doi:10.1371/journal.pcbi.1000400

Yovel Y, Stilz P, Franz MO, Boonman A, Schnitzler HU (2009)
What does a plant sound like? The statistics of natural plant echoes as received by echolocating bats.
PLoS Comput Biol 5(7): e1000429.doi:10.1371/journal.pcbi.1000429


Yovel Y, Franz MO, Stilz P, Schnitzler HU (2008)
Plant classification from bat-like echolocation signals.
PLOS Comp Biol 4(3) e1000032. doi:10.1371/journal.pcbi.1000032

Johnson N, Vos A, Neubert L, Freudling C, Mansfield KL, Kaipf I, Denzinger A, Hicks D, Núnez A, Franka R, Rupprecht CE, Müller T, Fooks A (2008)
Experimental study of European bat lyssavirus type-2 infection in Daubenton’s bats (Myotis daubentonii).
J Gen Virol 89:2662-2672


Schnitzler HU, Denzinger A (2011)
„Ehrlichkeit“ in der akustischen Kommunikation bei Tieren.
In: Klosinski G (ed) Tarnen, Täuschen, Lügen. Narr Francke Attempto Verlag, Tübingen, pp 205-211

Schnitzler H-U, Denzinger A (2009)
Neuroethology of biosonar systems in bats.
In: Windhorst U, Binder M, Hirokawa N (eds)., Encyclopedia of Neuroscience, Part 14, Springer Verlag, Heidelberg, pp 2654-2661

Schnitzler HU, Denzinger A (2008)
Die Stimmen der Natur.
In: Geissner HK (ed) Das Phänomen Stimme: Natürliche Veranlagung und kulturelle Formung. Röhrig Universitätsverlag, St. Ingbert, pp 33 -41

Schnitzler HU (2006)
Das Singen der Vögel.
In: Das Phänomen Stimme: Imitation und Identität (ed Thomas Kopfermannn) Röhrig Universitätsverlag, St. Ingbert, pp 33-41

Denzinger A, Schnitzler HU (2004)
Perceptual tasks in echolocating bats.
In: Ilg UJ, Bülthoff HH, Mallot HA (eds) Dynamic Perception. Akademische Verlagsgesellschaft, Berlin, pp 33-38

Müller R, Schnitzler HU (2001)
Computational assessment of an acoustic flow hypothesis for cf-bats.
In: Greenberg M, Slaney M (eds) Computational models of auditory function. pp 141-150

Papers before 2008


Melcón ML, Denzinger A, Schnitzler HU (2007)
Aerial hawking and landing: approach behaviour in Natterer's bats, Myotis nattereri (Kuhl 1818).
J Exp Biol 210:4457-4464

Schaub A, Schnitzler HU (2007a)
Echolocation behavior of the bat Vespertilio murinus reveals the border between the habitat types "edge" and "open space".
Behav Ecol Sociobiol 61:513-523.

Schaub A, Schnitzler HU.
Flight and echolocation behaviour of three vespertilionid bat species while commuting on flyways (2007b)
J Comp Physiol A 193:1185-1194.

Vos A, Kaipf I, Denzinger A, Fooks T, Johnson N. Müller T (2007) European Bat Lyssaviruses – an ecological enigma.
Acta Chiropterologica 9(1):283-296


Suthers RA, Narins PM, Lin W, Schnitzler HU, Denzinger A, Xu C, Feng AS (2006)
Voices of the Dead: Complex Nonlinear Vocal Signals from the Larynx of an Ultrasonic Frog.
J Exp Biol 209:4984-4993.

Dietrich S, Zaum DP, Kiefer A, Schnitzler H-U, Denzinger A (2006) Echolocation signals by the plecotine bat, Plecotus macrobullaris.
Acta Chiropt 8:465-475.

Götz T, Verfuss U K, Schnitzler H U (2006)
‘Eavesdropping’ in wild rough-toothed dolphins (Steno bredanensis)?
Biology Letters 2, 5-7

Thies W, Kalko E K V, Schnitzler H U (2006)
Influence of environment and resource availability on activity of Carollia castanea (Phyllostomidae) in Panama.
Journal of Mammalogy 87, 331-338


Boonman A, Schnitzler HU (2005)
Frequency modulation patterns in the echolocation signals of two vespertilionid bats.
J Comp Physiol A 191:13-21

Siemers BM, Baur E, Schnitzler H-U (2005)
Acoustical mirror effect increases prey detection distance in trawling bats.
Naturwissenschaften 92: 272-276

Verfuss UK, Miller LA, Schnitzler H-U (2005)
Spatial orientiation in echolocating harbour porpoises (Phocoena phocoena)
J Experim Biol 208: 3385-3394


Denzinger A, Kalko EKV, Jones G (2004)
Ecological and Evolutionary Aspects of Echolocation in Bats.
In: Thomas J, Moss C, Vater M (eds) Echolocation in bats and dolphins. University of Chicago Press, Chicago pp. 311-326

Narins PM, Feng AS, Lin WY, Schnitzler HU, Denzinger A, Suthers RA, Xu CH (2004)
Old World frog and bird, vocalizations contain prominent ultrasonic harmonics.
J Acoust Soc Am 115:910-913

Schnitzler HU, Kalko EKV, Denzinger A (2004)
Evolution of echolocation and foraging behavior in bats.
In: Thomas J, Moss CF, Vater M (eds) Echolocation in bats and dolphins. University of Chicago Press, Chicago, pp 331-339

Siemers BM, Schnitzler HU (2004)
Echolocation signals reflect niche differentiation in five sympatric congeneric bat species.
Nature 429:657-661


Schnitzler HU, Moss CF, Denzinger A (2003)
From spatial orientation to food acquisition in echolocating bats.
Trends Ecol Evol 18:386-394


Denzinger A, Siemers BM, Schaub A, Schnitzler HU (2001) Echolocation by the barbastelle bat, Barbastella barbastellus.
J Comp Physiol,A 187:521-528

Schnitzler HU, Kalko EKV (2001)
Echolocation by insect-eating bats.
Bioscience 51:557-569

Siemers BM, Dietz C, Nill D, Schnitzler HU (2001)
Myotis daubentonii is able to catch small fish.
Acta Chiropterologica 3:71-75

Siemers BM, Kalko EKV, Schnitzler HU (2001)
Echolocation behavior and signal plasticity in the Neotropical bat Myotis nigricans (Schinz, 1821) (Vespertilionidae): a convergent case with European species of Pipistrellus?
Behav Ecol Sociobiol 50:317-328

Siemers BM, Stilz P, Schnitzler HU (2001)
The acoustic advantage of hunting at low heights above water: behavioural experiments on the European 'trawling' bats Myotis capaccinii, M. dasycneme and M. daubentonii.
J Exp Biol 204:3843-3854


Müller R, Schnitzler HU (2000)
Acoustic flow perception in cf-bats: extraction of parameters.
J Acoust Soc Am 108:1298-1307

Siemers BM, Schnitzler HU (2000)
Natterer's bat (Myotis nattereri, Kuhl, 1818) hawks for prey close to vegetation using echolocation signals of very broad bandwidth.
Behav Ecol Sociobiol 47:400-412