Visualizing sound: counting wolves by using a spectral view of the chorus howling

IntroductionMonitoring large carnivores is a central issue in conservation biology. The wolf (Canis lupus) is the most studied large carnivore in the world. After a massive decline and several local extinctions, mostly due to direct persecutions, wolves are now recolonizing many areas of their historical natural range. One of the main monitoring techniques is the howling survey, which is based on the wolves’ tendency to use vocalisations to mark territory ownership in response to howls of unknown individuals. In most cases wolf howling sessions are useful for the localisation of the pack, but they provide only an aural estimation of the chorus size.We tested and present a new bioacoustic approach to estimate chorus size by recording wolves’ replies and visualising choruses through spectrograms and spectral envelopes. To test the methodology, we compared: a) the values detected by visual inspections with the true chorus size to test for accuracy; b) the bioacoustic estimations of a sample of free-ranging wolves’ replies developed by different operators to test for precision of the method; c) the aural field estimation of chorus size of a sample of free-ranging wolves’ replies with the sonogram analysis of the same recordings to test for difference between methods.ResultsVisual inspection of the chorus by spectrogram and spectrum proved to be useful in determining the number of concurrent voices in a wolf chorus. Estimations of chorus size were highly correlated with the number of wolves counted in a pack, and 92 % of 29 known chorus sizes were recognized by means of bioacoustic analysis. On the basis of spectrographic evidence, it was also possible to identify up to seven concurrent vocalisations in a chorus of nine wolves. Spectral analysis of 37 free ranging wolves’ replies showed a high correlation between the chorus size estimations of the different operators (92.8 %), but a low correlation with the aural estimation (59.2 %).ConclusionsWolf howling monitoring technique could be improved by recording wolves’ replies and by using bioacoustic tools such as spectrograms and spectral envelopes to determine the size of the wolf chorus. Compared with other monitoring techniques (i.e., genetic analysis), bioacoustic analysis requires widely available informatic tools (i.e., sound recording set of devices and sound analysis software) and a low budget. Information obtained by means of chorus analysis can also be combined with that provided by other techniques.Moreover, howls can be recorded and stored in audio file format with a good resolution (i.e. in “Wave” format), thus representing a useful tool for future listening and investigations, which can be countlessly employed without risks of time deterioration.

[1]  L. Mech,et al.  An analysis of howling response parameters useful for wolf pack censusing , 1982 .

[2]  M. Hebblewhite,et al.  Status and Ecological Effects of the World’s Largest Carnivores , 2014, Science.

[3]  L. Mech,et al.  WOLF HOWLING AND ITS ROLE IN TERRITORY MAINTENANCE , 1979 .

[4]  E. Font,et al.  Iberian Wolf Howls: Acoustic Structure, Individual Variation, and a Comparison with North American Populations , 2007 .

[5]  F. Dessì-Fulgheri,et al.  Group specific vocal signature in free-ranging wolf packs , 2012 .

[6]  L. Waits,et al.  Estimating gray wolf pack size and family relationships using noninvasive genetic sampling at rendezvous sites , 2011 .

[7]  W Seidner,et al.  Nonlinear phenomena in the natural howling of a dog-wolf mix. , 2000, The Journal of the Acoustical Society of America.

[8]  O. Liberg,et al.  Recovery of large carnivores in Europe’s modern human-dominated landscapes , 2014, Science.

[9]  K. Berger Carnivore‐Livestock Conflicts: Effects of Subsidized Predator Control and Economic Correlates on the Sheep Industry , 2006, Conservation biology : the journal of the Society for Conservation Biology.

[10]  F. Harrington Aggressive howling in wolves , 1987, Animal Behaviour.

[11]  R. H. Wiley,et al.  Reverberations and Amplitude Fluctuations in the Propagation of Sound in a Forest: Implications for Animal Communication , 1980, The American Naturalist.

[12]  Michael Schoeffler,et al.  Human ability of counting the number of instruments in polyphonic music , 2013 .

[13]  David Huron Voice Denumerability in Polyphonic Music of Homogeneous Timbres , 1989 .

[14]  N. Selva,et al.  KILL RATES AND PREDATION BY WOLVES ON UNGULATE POPULATIONS IN BIAŁOWIEŻA PRIMEVAL FOREST (POLAND) , 2002 .

[15]  M. Apollonio,et al.  Wolves in the Casentinesi Forests: insights for wolf conservation in Italy from a protected area with a rich wild prey community , 2004 .

[16]  M. Walpole,et al.  Tourism and flagship species in conservation , 2002, Biodiversity & Conservation.

[17]  Julián Velasco Valdés,et al.  Implementation of a diffusive differential reassignment method for signal enhancement: An application to wolf population counting , 2007, Appl. Math. Comput..

[18]  L. Mech,et al.  Patterns of homesite attendance in two Minnesota wolf packs , 1982 .

[19]  Thierry Aubin,et al.  SEEWAVE, A FREE MODULAR TOOL FOR SOUND ANALYSIS AND SYNTHESIS , 2008 .

[20]  E. Randi,et al.  Monitoring wolves (Canis lupus) by non-invasive genetics and camera trapping: a small-scale pilot study , 2012, European Journal of Wildlife Research.

[21]  R Core Team,et al.  R: A language and environment for statistical computing. , 2014 .

[22]  J. C. Fentress,et al.  Individually distinct vocalizations in timber wolves, Canis lupus , 1990, Animal Behaviour.

[23]  Robert L. Beschta,et al.  Trophic cascades in Yellowstone: The first 15 years after wolf reintroduction , 2012 .

[24]  Christopher S. Evans,et al.  Animal Acoustic Communication: Sound Analysis and Research Methods , 2011 .

[25]  U. Breitenmoser Large predators in the Alps: The fall and rise of man's competitors , 1998 .

[26]  P. Joslin,et al.  Movements and Home Sites of Timber Wolves in Alǵonquin Park , 1967 .

[27]  F. Harrington CHORUS HOWLING BY WOLVES: ACOUSTIC STRUCTURE, PACK SIZE AND THE BEAU GESTE EFFECT , 1989 .

[28]  F. Harris On the use of windows for harmonic analysis with the discrete Fourier transform , 1978, Proceedings of the IEEE.

[29]  T. Fuller,et al.  Evaluation of a Simulated Howling Survey for Wolves , 1988 .

[30]  Roger Pradel,et al.  Capture–recapture population growth rate as a robust tool against detection heterogeneity for population management , 2011 .

[31]  D. Macdonald,et al.  A review of financial instruments to pay for predator conservation and encourage human–carnivore coexistence , 2011, Proceedings of the National Academy of Sciences.

[32]  R. Primack,et al.  Essentials of Conservation Biology , 1994 .

[33]  S. H. Fritts,et al.  The relationship of wolf recovery to habitat conservation and biodiversity in the northwestern United States , 1994 .

[34]  J. Lanszki,et al.  Expansion range of the golden jackal in Hungary between 1997 and 2006 , 2009 .

[35]  Julián Velasco Valdés,et al.  On a chirplet transform-based method applied to separating and counting wolf howls , 2008, Signal Process..

[36]  J. Kleiven,et al.  Factors influencing the social acceptability of large carnivore behaviours , 2004, Biodiversity & Conservation.

[37]  H. Herzel,et al.  SUBHARMONICS, BIPHONATION, AND DETERMINISTIC CHAOS IN MAMMAL VOCALIZATION , 1998 .

[38]  W. Tecumseh Fitch,et al.  Visualization of system dynamics using phasegrams , 2013, Journal of The Royal Society Interface.

[39]  K. Schmidt,et al.  Howling activity of free-ranging wolves (Canis lupus) in the Białowieża Primeval Forest and the Western Beskidy Mountains (Poland) , 2007, Journal of Ethology.

[40]  Paul Boersma,et al.  Praat, a system for doing phonetics by computer , 2002 .

[41]  Paul Boersma,et al.  Praat: doing phonetics by computer , 2003 .

[42]  S. Sgardelis,et al.  Wolf pack rendezvous site selection in Greece is mainly affected by anthropogenic landscape features , 2014, European Journal of Wildlife Research.

[43]  C. L. Elliott,et al.  Comparison of Remotely-triggered Cameras vs. Howling Surveys for Estimating Coyote (Canis latrans) Abundance in Central Kentucky , 2011 .

[44]  L. Mech,et al.  The Wolf: The Ecology and Behavior of an Endangered Species , 1970 .

[45]  M. Apollonio,et al.  Temporal changes of howling in south European wolf packs , 2002 .

[46]  L. Mech,et al.  Wolf Pack Size And Food Acquisition , 1997, The American Naturalist.

[47]  Benjamín Dugnol Álvarez,et al.  Wolf population counting by spectrogram image processing , 2007, Appl. Math. Comput..

[48]  R. Ruff,et al.  Howling by coyotes (Canis latrans) : variation among social classes, seasons, and pack sizes , 1998 .

[49]  Mech Ld,et al.  Wolf pack size and food acquisition. , 1997 .

[50]  L. Adams,et al.  Evaluation of wolf density estimation from radiotelemetry data , 2005 .

[51]  David J. Mladenoff,et al.  Predicting gray wolf landscape recolonization: logistic regression models vs. new field data , 1999 .

[52]  J. Blanco Surveying wolves without snow: a critical review of the methods used in Spain , 2011 .