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Journal of BirdLife Australia
RESEARCH ARTICLE

Vocal individuality of Little Spotted Kiwi (Apteryx owenii)

Andrew Digby A C , Ben D. Bell A and Paul D. Teal B
+ Author Affiliations
- Author Affiliations

A Centre for Biodiversity and Restoration Ecology, School of Biological Sciences, Victoria University of Wellington, Wellington, 6140, New Zealand.

B School of Engineering and Computer Science, Victoria University of Wellington, Wellington, 6140, New Zealand.

C Corresponding author. Email: andrewdigby@mac.com

Emu 114(4) 326-336 https://doi.org/10.1071/MU13114
Submitted: 26 August 2013  Accepted: 4 June 2014   Published: 8 September 2014

Abstract

The ability to identify individuals by call is particularly important for monitoring cryptic or nocturnal species. It provides key benefits in conservation management, such as allowing correction of biases in censuses and monitoring individual survival. Call surveys are a key tool in conservation of kiwi (Apterygidae), yet individuality of calls has previously been assessed in just one of the five species, the Brown Kiwi (formerly North Island Brown Kiwi, Apteryx mantelli). We have made the first test of whether Little Spotted Kiwi (Apteryx owenii; LSK) can be individually identified by call. Three classification methods (discriminant function analysis, support vector machines and statistical classifiers) were used to assign identity from either call or syllable variables. Using syllables, classification was significantly better than expected by chance for both males and females. Individual LSK can therefore be identified acoustically, suggesting that vocalisations may be used for conspecific identification. Classifier performance was worse using call variables, with no better than chance assignment for females and low accuracy for males. Significant differences in classification ability between the sexes support the hypothesis that the function of calls of male and female LSK differ. Furthermore, identification by call was much less reliable in LSK than in the more genetically diverse Brown Kiwi. This indicates that call diversity may be related to genetic diversity in kiwi, which, if confirmed, could provide a powerful conservation tool for rapid assessment of genetic differences among populations of these threatened species.

Additional keywords: acoustic communication, vocal discrimination.


References

Acevedo, M. A., Corrada-Bravo, C. J., Corrada-Bravo, H., Villanueva-Rivera, L. J., and Aide, T. M. (2009). Automated classification of bird and amphibian calls using machine learning: a comparison of methods. Ecological Informatics 4, 206–214.
Automated classification of bird and amphibian calls using machine learning: a comparison of methods.Crossref | GoogleScholarGoogle Scholar |

Appleby, B. M., and Redpath, S. M. (1997). Variation in the male territorial hoot of the Tawny Owl Strix aluco in three English populations. Ibis 139, 152–158.
Variation in the male territorial hoot of the Tawny Owl Strix aluco in three English populations.Crossref | GoogleScholarGoogle Scholar |

Araya-Ajoy, Y.-M., Chaves-Campos, J., Kalko, E. K. V., and DeWoody, J. A. (2009). High-pitched notes during vocal contests signal genetic diversity in Ocellated Antbirds. PLoS ONE 4, e8137.
High-pitched notes during vocal contests signal genetic diversity in Ocellated Antbirds.Crossref | GoogleScholarGoogle Scholar | 19956580PubMed |

Aubin, T., Jouventin, P., and Hildebrand, C. (2000). Penguins use the two-voice system to recognize each other. Proceedings of the Royal Society of London – B. Biological Sciences 267, 1081–1087.
Penguins use the two-voice system to recognize each other.Crossref | GoogleScholarGoogle Scholar | 1:STN:280:DC%2BD3M%2FhtFClsA%3D%3D&md5=37f00f9ecb3daad948e56e4964e30c04CAS |

Baldo, S., and Mennill, D. J. (2011). Vocal behavior of Great Curassows, a vulnerable Neotropical bird. Journal of Field Ornithology 82, 249–258.

Bard, S. C., Hau, M., Wikelski, M., and Wingfield, J. C. (2002). Vocal distinctiveness and response to conspecific playback in the Spotted Antbird, a Neotropical suboscine. Condor 104, 387–394.
Vocal distinctiveness and response to conspecific playback in the Spotted Antbird, a Neotropical suboscine.Crossref | GoogleScholarGoogle Scholar |

BirdLife International (2012). Apteryx owenii (Little Spotted Kiwi). In ‘The IUCN Red List of Threatened Species’, ver. 2014.1. Available at http://www.iucnredlist.org/details/22678129/0 [Verified 13 July 2014].

Blumstein, D. T., Mennill, D. J., Clemins, P., Girod, L., Yao, K., Patricelli, G., Deppe, J. L., Krakauer, A. H., Clark, C., Cortopassi, K. A., Hanser, S. F., McCowan, B., Ali, A. M., and Kirschel, A. N. G. (2011). Acoustic monitoring in terrestrial environments using microphone arrays: applications, technological considerations and prospectus. Journal of Applied Ecology 48, 758–767.
Acoustic monitoring in terrestrial environments using microphone arrays: applications, technological considerations and prospectus.Crossref | GoogleScholarGoogle Scholar |

Bradley, A. P. (1997). The use of the area under the ROC curve in the evaluation of machine learning algorithms. Pattern Recognition 30, 1145–1159.
The use of the area under the ROC curve in the evaluation of machine learning algorithms.Crossref | GoogleScholarGoogle Scholar |

Budka, M., and Osiejuk, T. S. (2013). Neighbour–stranger call discrimination in a nocturnal rail species, the Corncrake Crex crex. Journal of Ornithology 154, 685–694.
Neighbour–stranger call discrimination in a nocturnal rail species, the Corncrake Crex crex.Crossref | GoogleScholarGoogle Scholar |

Castro, I. (2011). ‘Kiwi. A Natural History.’ (New Holland: Auckland, New Zealand.)

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

Cheng, J., Xie, B., Lin, C., and Ji, L. (2012). A comparative study in birds: call-type- independent species and individual recognition using four machine-learning methods and two acoustic features. Bioacoustics 21, 157–171.
A comparative study in birds: call-type- independent species and individual recognition using four machine-learning methods and two acoustic features.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BC38XhtlagtLnF&md5=249ff29a1ae6b6be5f4e418d8f8ac174CAS |

Clark, J., Boersma, P., and Olmsted, D. (2006). Name that tune: call discrimination and individual recognition in Magellanic Penguins. Animal Behaviour 72, 1141–1148.
Name that tune: call discrimination and individual recognition in Magellanic Penguins.Crossref | GoogleScholarGoogle Scholar |

Cohen, J. (1960). A coefficient of agreement for nominal scales. Educational and Psychological Measurement 20, 37–46.
A coefficient of agreement for nominal scales.Crossref | GoogleScholarGoogle Scholar |

Colbourne, R. M. (2005). Kiwi (Apteryx spp.) on offshore New Zealand islands. Department of Conservation Research and Development Series 208, Wellington, New Zealand.

Colbourne, R. M. (2006). Kiwi call scheme. Investigation S5020/129. Department of Conservation, Wellington, New Zealand.

Corfield, J. (2005). Description, dueting, seasonal variations, and individual identification of the vocalisations of the Brown Kiwi (Apteryx mantelli). M.App.Sc. Thesis, Auckland University of Technology, Auckland, New Zealand.

Corfield, J., Gillman, L., and Parsons, S. (2008). Vocalisations of the North Island Brown Kiwi (Apteryx mantelli). Auk 125, 326–335.
Vocalisations of the North Island Brown Kiwi (Apteryx mantelli).Crossref | GoogleScholarGoogle Scholar |

Cristianini, N., and Shawe-Taylor, J. (2000). ‘An Introduction to Support Vector Machines and Other Kernel-based Learning Methods.’ (Cambridge University Press: Cambridge, UK.)

Digby, A. (2013). Whistling in the dark: an acoustic study of Little Spotted Kiwi. Ph.D. Thesis, Victoria University of Wellington, Wellington, New Zealand.

Digby, A., Bell, B. D., and Teal, P. D. (2013). Vocal cooperation between the sexes in Little Spotted Kiwi Apteryx owenii. Ibis 155, 229–245.
Vocal cooperation between the sexes in Little Spotted Kiwi Apteryx owenii.Crossref | GoogleScholarGoogle Scholar |

Digby, A., Bell, B. D., and Teal, P. D. (2014). Non-linear phenomena in Little Spotted Kiwi calls. Bioacoustics 23, 113–128.
Non-linear phenomena in Little Spotted Kiwi calls.Crossref | GoogleScholarGoogle Scholar |

Dundar, M., Krishnapuram, B., Bi, J., and Rao, R. B. (2007). Learning classifiers when the training data is not IID. In ‘Proceedings of the 20th International Joint Conference on Artificial Intelligence’, 6–12 January 2007 Hyderabad, India. pp. 756–761. (Morgan Kaufmann Publishers: San Francisco, CA.)

Fitch, W., and Fritz, J. (2006). Rhesus Macaques spontaneously perceive formants in conspecific vocalizations. Journal of the Acoustical Society of America 120, 2132–2141.
Rhesus Macaques spontaneously perceive formants in conspecific vocalizations.Crossref | GoogleScholarGoogle Scholar | 17069311PubMed |

Fitch, W. T., Neubauer, J., and Herzel, H. (2002). Calls out of chaos: the adaptive significance of nonlinear phenomena in mammalian vocal production. Animal Behaviour 63, 407–418.
Calls out of chaos: the adaptive significance of nonlinear phenomena in mammalian vocal production.Crossref | GoogleScholarGoogle Scholar |

Fitzsimmons, L. P., Barker, N. K., and Mennill, D. J. (2008). Individual variation and lek-based vocal distinctiveness in songs of the Screaming Piha (Lipaugus vociferans), a suboscine songbird. Auk 125, 908–914.
Individual variation and lek-based vocal distinctiveness in songs of the Screaming Piha (Lipaugus vociferans), a suboscine songbird.Crossref | GoogleScholarGoogle Scholar |

Fox, E. J. S. (2008). A new perspective on acoustic individual recognition in animals with limited call sharing or changing repertoires. Animal Behaviour 75, 1187–1194.
A new perspective on acoustic individual recognition in animals with limited call sharing or changing repertoires.Crossref | GoogleScholarGoogle Scholar |

Galeotti, P., and Pavan, G. (1991). Individual recognition of male Tawny Owls (Strix aluco) using spectrograms of their territorial calls. Ethology Ecology and Evolution 3, 113–126.
Individual recognition of male Tawny Owls (Strix aluco) using spectrograms of their territorial calls.Crossref | GoogleScholarGoogle Scholar |

Galeotti, P., and Sacchi, R. (2001). Turnover of territorial Scops Owls Otus scops as estimated by spectrographic analyses of male hoots. Journal of Avian Biology 32, 256–262.
Turnover of territorial Scops Owls Otus scops as estimated by spectrographic analyses of male hoots.Crossref | GoogleScholarGoogle Scholar |

Geberzahn, N., Goymann, W., Muck, C., and ten Cate, C. (2009). Females alter their song when challenged in a sex-role reversed bird species. Behavioral Ecology and Sociobiology 64, 193–204.
Females alter their song when challenged in a sex-role reversed bird species.Crossref | GoogleScholarGoogle Scholar | 19946649PubMed |

Gilbert, G., McGregor, P. K., and Tyler, G. (1994). Vocal individuality as a census tool: practical considerations illustrated by a study of two rare species. Journal of Field Ornithology 65, 335–348.

Gilbert, G., Tyler, G., and Smith, K. (2002). Local annual survival of booming male Great Bittern Botaurus stellaris in Britain, in the period 1990–1999. Ibis 144, 51–61.
Local annual survival of booming male Great Bittern Botaurus stellaris in Britain, in the period 1990–1999.Crossref | GoogleScholarGoogle Scholar |

Greenewalt, C. H. (1968). ‘Bird Song: Acoustics and Physiology.’ (Smithsonian Institution Press: Washington, DC.)

Heather, B., and Robertson, H. (2005). ‘The Field Guide to the Birds of New Zealand’, 2nd edn. (Penguin Books: Auckland, New Zealand.)

Hojem, C. (2006). The real New Zealand national anthem: calling structure and calling rate of North Island Brown Kiwi (Apteryx mantelli) on Ponui Island and comparisons with Rarewarewa Reserve. M.Sc. Thesis, University of Auckland, Auckland, New Zealand.

Hutto, R. L., and Stutzman, R. J. (2009). Humans versus autonomous recording units: a comparison of point-count results. Journal of Field Ornithology 80, 387–398.
Humans versus autonomous recording units: a comparison of point-count results.Crossref | GoogleScholarGoogle Scholar |

Jiménez, I., Londoño, G. A., and Cadena, C. D. (2003). Efficiency, bias, and consistency of visual and aural surveys of curassows (Cracidae) in tropical forests. Journal of Field Ornithology 74, 210–216.

Jolly, J. (1989). A field study of the breeding biology of the Little Spotted Kiwi (Apteryx owenii) with emphasis on the causes of nest failures. Journal of the Royal Society of New Zealand 19, 433–448.
A field study of the breeding biology of the Little Spotted Kiwi (Apteryx owenii) with emphasis on the causes of nest failures.Crossref | GoogleScholarGoogle Scholar |

Jones, D., and Smith, G. (1997). Vocalisations of the Marbled Frogmouth II: An assessment of vocal individuality as a potential census technique. Emu 97, 296–304.
Vocalisations of the Marbled Frogmouth II: An assessment of vocal individuality as a potential census technique.Crossref | GoogleScholarGoogle Scholar |

Karatzoglou, A., Smola, A., Hornik, K., and Zeileis, A. (2004). Kernlab – an S4 package for kernel methods in R. Journal of Statistical Software 11, 1–20.

Kirschel, A. N. G., Earl, D. A., Yao, Y., Escobar, I. A., Viches, E., Vallejo, E. E., and Taylor, C. E. (2009). Using songs to identify individual Mexican Antthrush Formicarius moniliger: comparison of four classification methods. Bioacoustics 19, 1–20.
Using songs to identify individual Mexican Antthrush Formicarius moniliger: comparison of four classification methods.Crossref | GoogleScholarGoogle Scholar |

Kirschel, A. N. G., Cody, M. L., Harlow, Z. T., Promponas, V. J., Vallejo, E. E., and Taylor, C. E. (2011). Territorial dynamics of Mexican Ant-thrushes Formicarius moniliger revealed by individual recognition of their songs. Ibis 153, 255–268.
Territorial dynamics of Mexican Ant-thrushes Formicarius moniliger revealed by individual recognition of their songs.Crossref | GoogleScholarGoogle Scholar |

Kraemer, H. C., Periyakoil, V. S., and Noda, A. (2004). ‘Kappa Coefficients in Medical Research.’ (Wiley: Hoboken, NJ.)

Kroodsma, D. (2005). ‘The Singing Life of Birds.’ (Houghton Mifflin: Boston, MA.)

Kuhn, M. (2008). Building predictive models in R using the caret package. Journal of Statistical Software 28, 1–26.

Kuhn, M., Weston, S., and Coulter, N. (2013). C50: C5.0 decision trees and rule-based models. Available at http://CRAN.R-project.org/package=C50 [Verified 1 December 2013].

Langmore, N. (1998). Functions of duet and solo songs of female birds. Trends in Ecology & Evolution 13, 136–140.
Functions of duet and solo songs of female birds.Crossref | GoogleScholarGoogle Scholar | 1:STN:280:DC%2BC3M7itFymtQ%3D%3D&md5=4b023b81a10c070c79c147b1ddf204bdCAS |

Loesche, P., Stoddard, P., Higgins, B. J., and Beecher, M. D. (1991). Signature versus perceptual adaptations for individual vocal recognition in swallows. Behaviour 118, 15–25.
Signature versus perceptual adaptations for individual vocal recognition in swallows.Crossref | GoogleScholarGoogle Scholar |

Marshall, R. C., Buchanan, K. L., and Catchpole, C. K. (2003). Sexual selection and individual genetic diversity in a songbird. Biology Letters 270, S248–S250.

May, L. (1998). Individually distinctive Corncrake Crex crex calls: a further study. Bioacoustics 9, 135–148.
Individually distinctive Corncrake Crex crex calls: a further study.Crossref | GoogleScholarGoogle Scholar |

McDonald, P. G., and Wright, J. (2011). Bell Miner provisioning calls are more similar among relatives and are used by helpers at the nest to bias their effort towards kin. Proceedings of the Royal Society of London – B. Biological Sciences 278, 3403–3411.
Bell Miner provisioning calls are more similar among relatives and are used by helpers at the nest to bias their effort towards kin.Crossref | GoogleScholarGoogle Scholar |

McKown, M. W. (2008). Acoustic communication in colonial seabirds: individual, sexual, and species-specific variation in acoustic signals of Pterodroma petrels. Ph. D. Thesis, University of North Carolina.

Mennill, D. J. (2011). Individual distinctiveness in avian vocalizations and the spatial monitoring of behaviour. Ibis 153, 235–238.
Individual distinctiveness in avian vocalizations and the spatial monitoring of behaviour.Crossref | GoogleScholarGoogle Scholar |

Mennill, D., and Vehrencamp, S. (2008). Context-dependent functions of avian duets revealed by microphone-array recordings and multispeaker playback. Current Biology 18, 1314–1319.
Context-dependent functions of avian duets revealed by microphone-array recordings and multispeaker playback.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD1cXhtVykurvN&md5=a9a525ad2013db811568e736fafb5fa3CAS | 18771924PubMed |

Mikkelsen, G., Dale, S., Holtskog, T., Budka, M., and Osiejuk, T. S. (2013). Can individually characteristic calls be used to identify long-distance movements of Corncrakes Crex crex? Journal of Ornithology 154, 751–760.
Can individually characteristic calls be used to identify long-distance movements of Corncrakes Crex crex? Crossref | GoogleScholarGoogle Scholar |

Miller, P., and Pierce, R. (1995). Distribution and decline of the North Island Brown Kiwi (Apteryx australis mantelli) in Northland. Notornis 42, 203–211.

Miller, H. C. H., Bowker-Wright, G. G., Kharkrang, M. M., and Ramstad, K. K. (2011). Characterisation of class II B MHC genes from a ratite bird, the Little Spotted Kiwi (Apteryx owenii). Immunogenetics 63, 223–233.
Characterisation of class II B MHC genes from a ratite bird, the Little Spotted Kiwi (Apteryx owenii).Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BC3MXivFyitrY%3D&md5=e84440de375ac0dca03e6b9b6b52a384CAS |

Mitchell, B. R., Makagon, M. M., Jaeger, M. M., and Barrett, R. H. (2006). Information content of Coyote barks and howls. Bioacoustics 15, 289–314.
Information content of Coyote barks and howls.Crossref | GoogleScholarGoogle Scholar |

Mundry, R., and Sommer, C. (2007). Discriminant function analysis with nonindependent data: consequences and an alternative. Animal Behaviour 74, 965–976.
Discriminant function analysis with nonindependent data: consequences and an alternative.Crossref | GoogleScholarGoogle Scholar |

Parker, T. A. (1991). On the use of tape recorders in avifaunal surveys. Auk 108, 443–444.

Peake, T., McGregor, P. K., Smith, K., and Tyler, G. (1998). Individuality in Corncrake (Crex crex) vocalizations. Ibis 140, 120–127.
Individuality in Corncrake (Crex crex) vocalizations.Crossref | GoogleScholarGoogle Scholar |

Petrusková, T., Osiejuk, T., Linhart, P., and Petrusek, A. (2008). Structure and complexity of perched and flight songs of the Tree Pipit (Anthus trivialis). Annales Zoologici Fennici 45, 135–148.
Structure and complexity of perched and flight songs of the Tree Pipit (Anthus trivialis).Crossref | GoogleScholarGoogle Scholar |

Pfaff, J. A., Zanette, L., MacDougall-Shackleton, S. A., and MacDougall-Shackleton, E. A. (2007). Song repertoire size varies with HVC volume and is indicative of male quality in Song Sparrows (Melospiza melodia). Proceedings of the Royal Society of London – B. Biological Sciences 274, 2035–2040.
Song repertoire size varies with HVC volume and is indicative of male quality in Song Sparrows (Melospiza melodia).Crossref | GoogleScholarGoogle Scholar |

Pierce, R., and Westbrooke, I. (2003). Call count responses of North Island Brown Kiwi to different levels of predator control in Northland, New Zealand. Biological Conservation 109, 175–180.
Call count responses of North Island Brown Kiwi to different levels of predator control in Northland, New Zealand.Crossref | GoogleScholarGoogle Scholar |

Pierotti, R. (1991). Infanticide versus adoption: an intergenerational conflict. American Naturalist 138, 1140–1158.
Infanticide versus adoption: an intergenerational conflict.Crossref | GoogleScholarGoogle Scholar |

Pinheiro, J., Bates, D., DebRoy, S., and Sarkar, D., and R Development Core Team (2013). nlme: linear and nonlinear mixed effects models, ver. 3.1–111. Available at http://cran.r-project.org/web/packages/nlme/index.html [Verified 23 November 2013]

R Development Core Team (2013). R: a language and environment for statistical computing, ver. 2.15. (R Foundation for Statistical Computing: Vienna, Austria.) Available at http://www.R-project.org [Verified 23 November 2013].

Ramstad, K. M., Pfunder, M., Robertson, H. A., Colbourne, R. M., Allendorf, F. W., and Daugherty, C. H. (2010). Fourteen microsatellite loci cross-amplify in all five kiwi species (Apteryx spp.) and reveal extremely low genetic variation in Little Spotted Kiwi (A. owenii). Conservation Genetics Resources 2, 333–336.
Fourteen microsatellite loci cross-amplify in all five kiwi species (Apteryx spp.) and reveal extremely low genetic variation in Little Spotted Kiwi (A. owenii).Crossref | GoogleScholarGoogle Scholar |

Ramstad, K. M., Colbourne, R. M., Robertson, H. A., Allendorf, F. W., and Daugherty, C. H. (2013). Genetic consequences of a century of protection: serial founder events and survival of the Little Spotted Kiwi (Apteryx owenii). Proceedings of the Royal Society of London – B. Biological Sciences 280, 20130576.
Genetic consequences of a century of protection: serial founder events and survival of the Little Spotted Kiwi (Apteryx owenii).Crossref | GoogleScholarGoogle Scholar |

Raudys, S. (2001). ‘Statistical and Neural Classifiers.’ (Springer: London.)

Robertson, H. A., and Colbourne, R. M. (2004). Survival of Little Spotted Kiwi (Apteryx owenii) on Kapiti Island. Notornis 51, 161–163.

Robertson, H. A., and Fraser, J. R. (2009). Use of trained dogs to determine the age structure and conservation status of kiwi Apteryx spp. populations. Bird Conservation International 19, 121–129.
Use of trained dogs to determine the age structure and conservation status of kiwi Apteryx spp. populations.Crossref | GoogleScholarGoogle Scholar |

Robin, V. V., Katti, M., Purushotham, C., Sancheti, A., and Sinha, A. (2011). Singing in the sky: song variation in an endemic bird on the sky islands of southern India. Animal Behaviour 82, 513–520.
Singing in the sky: song variation in an endemic bird on the sky islands of southern India.Crossref | GoogleScholarGoogle Scholar |

Saunders, D., and Wooller, R. (1988). Consistent individuality of voice in birds as a management tool. Emu 88, 25–32.
Consistent individuality of voice in birds as a management tool.Crossref | GoogleScholarGoogle Scholar |

Seddon, N., Amos, W., Mulder, R. A., and Tobias, J. A. (2004). Male heterozygosity predicts territory size, song structure and reproductive success in a cooperatively breeding bird. Proceedings of the Royal Society of London – B. Biological Sciences 271, 1823–1829.
Male heterozygosity predicts territory size, song structure and reproductive success in a cooperatively breeding bird.Crossref | GoogleScholarGoogle Scholar |

Suthers, R. (1994). Variable asymmetry and resonance in the avian vocal tract: a structural basis for individually distinct vocalizations. Journal of Comparative Physiology – A. Neuroethology, Sensory, Neural, and Behavioral Physiology 175, 457–466.
Variable asymmetry and resonance in the avian vocal tract: a structural basis for individually distinct vocalizations.Crossref | GoogleScholarGoogle Scholar | 1:STN:280:DyaK2M%2FlsFejug%3D%3D&md5=b87d9140f9d4c6e0d1586251be3c5042CAS |

Terry, A., Peake, T., and McGregor, P. K. (2005). The role of vocal individuality in conservation. Frontiers in Zoology 2, 10.
The role of vocal individuality in conservation.Crossref | GoogleScholarGoogle Scholar | 15960848PubMed |

Tooze, Z. J., Harrington, F. H., and Fentress, J. C. (1990). Individually distinct vocalizations in Timber Wolves, Canis lupus. Animal Behaviour 40, 723–730.
Individually distinct vocalizations in Timber Wolves, Canis lupus.Crossref | GoogleScholarGoogle Scholar |

Venables, W. N., and Ripley, B. D. (2002). ‘Modern Applied Statistics with S.’ (Springer Verlag: New York.)

Wilden, I., Herzel, H., Peters, G., and Tembrock, G. (1998). Subharmonics, biphonation, and deterministic chaos in mammal vocalization. Bioacoustics 8, 1–30.

Ziesemann, B. (2011). The social organisation and mating system of the Brown Kiwi (Apteryx mantelli). Ph.D. Thesis, Massey University, Albany, New Zealand.

Zuur, A. F., Ieno, E. N., and Smith, G. M. (2007). ‘Analysing Ecological Data’, 1st edn. (Springer: New York.)