Skip to main content
Intended for healthcare professionals
Restricted access
Research article
First published online September 1, 2009

Did Neanderthals and other early humans sing? Seeking the biological roots of music in the territorial advertisements of primates, lions, hyenas, and wolves

Abstract

Group defence of territories is found in many gregarious mammalian carnivores, including lions, canids, and hyenas. In these taxa, group members often mark territory boundaries and direct aggressive behaviour towards alien conspecifics found within the territory (Boydston et al., 2001). Middle Pleistocene hominids such as Neanderthals occupied an ecological niche similar to such large carnivores (Stiner, 2002), and so could be expected to share with them a suite of behavioural traits. Complex, coordinated vocalizations that function, at least in part, to advertise the group defence of a territory is one behavioural trait exhibited by several social carnivores, as well as many other gregarious animals, including primates. Hagen and Bryant (2003) proposed that the evolution of human music and dance was rooted in such coordinated auditory and visual territorial advertisements, an hypothesis we develop and expand upon here. Human proto-music, in essence, might have been functionally analogous to the howling of wolves.

Get full access to this article

View all access and purchase options for this article.

• References

Adams E. S. (2001). Approaches to the study of territory size and shape. Annual Review of Ecology and Systematics, 32, 277–303.
Alexander R. D. (1974). The evolution of social behaviour. Annual Review of Ecology and Systematics, 5, 325–83.
Arcadi A. C., Robert D., & Mugurusi F. (2004). A comparison of buttress drumming by male chimpanzees from two populations. Primates, 45, 135–39.
Baker R. R. (1983). Insect territoriality. Annual Review of Entomology, 28, 65–89.
Bermejo M., & Omedes A. (1999). Preliminary vocal repertoire and vocal communication of wild bonobos (Pan paniscus) at Lilungu (Democratic Republic of Congo). Folia Primatologica, 70, 328–57.
Bird R. B., & Smith E. A. (2005). Signalling theory, strategic interaction, and symbolic capital. Current Anthropology, 46, 221–48.
Bispham J. (2006). Rhythm in music: What is it? Who has it? And Why? Music Perception, 24, 125–34.
Bocherens H., Drucker D. G., Billiou D., Patou-Mathis M., & Vandermeersch B. (2005). Isotopic evidence for diet and subsistence pattern of the Saint-Cæsaire I Neanderthal: Review and use of a multi-source mixing model. Journal of Human Evolution, 49, 71–87.
Boydston E. E., Morelli T. L., & Holekamp K. E. (2001). Sex differences in territorial behaviour exhibited by the Spotted Hyena (Hyaenidae, Crocuta crocuta). Ethology, 107, 369–85.
Brantingham P. J. (1998). Hominid-carnivore coevolution and invasion of the predatory guild. Journal of Anthropological Archaeology, 176, 327–53.
Burrows E. G. (1945). Songs of Uvea and Futuna. Honolulu: Bernice P. Bishop Museum Bulletin No. 183.
Campbell R. D., Rosell F., Nolet B. A., & Dijkstra V. A. A. (2005). Territory and group sizes in Eurasian beavers (Castor fiber): Echoes of settlement and reproduction? Behavioral Ecology and Sociobiology, 58, 597–607.
Cashdan E. (1983). Territoriality among human foragers: Ecological models and an application to four Bushman groups. Current Anthropology, 24, 47–66.
Catchpole C. K., & Slater P. J. B. (1995). Bird song: Biological themes and variations. Cambridge, UK: Cambridge University Press.
Chagnon N. A. (1968). Yanomamö. New York: Holt, Rinehart and Winston.
Cheney D. L. (1987). Interactions and relationships between groups. In Smuts B. B., Cheney D. L., Seyfarth R. M., Wrangham R. W., & Struhsaker T. T. (eds), Primate societies (p. 267–81). Chicago: University of Chicago Press.
Colmenares F., Esteban M. M., & Zaragoza F. (2006). One-male units and clans in a colony of hamadryas baboons (Papio hamadryas hamadryas): Effect of male number and clan cohesion on feeding success. American Journal of Primatology, 68, 21–37.
Connor R. C. (2007). Dolphin social intelligence: complex alliance relationships in bottlenose dolphins and a consideration of selective environments for extreme brain size evolution in mammals. Philos Trans R Soc Lond B Biol Sci, 1–16.
Connor R. C., Smolker R. A., & Bejder L. (2006). Synchrony, social behaviour and alliance affiliation in indian ocean bottlenose dolphins, Tursiops aduncus. Animal Behaviour, 72, 1371–78.
Connor R. C., Smolker R. A., & Richards A. F. (1992). Two levels of alliance formation among male bottlenose dolphins (Tursiops sp.). Proceedings of the National Academy of Sciences, 89, 987–90.
Crockford C., Herbinger I., Vigilant L., & Boesch C. (2004). Wild chimpanzees produce group-specific calls: A case for vocal learning? Ethology, 110, 221–43.
Darwin C. (1871). The descent of man and selection in relation to sex. London: John Murray.
Delgado R. A. (2006). Sexual selection in the loud calls of male primates: Signal content and function. International Journal of Primatology, 27, 5–25.
Dyson-Hudson R., & Smith E. A. (1978). Human territoriality: An ecological reassessment. American Anthropologist, 80, 21–41.
East M. L., & Hofer H. (1991). Loud calling in a female-dominated mammalian society. II: Behavioural contexts and functions of whooping of spotted hyaenas, Crocuta crocuta. Animal behaviour, 42, 651–69.
Ekman P. (1971). Universals and cultural differences in facial expressions of emotion. Nebraska: University of Nebraska Press.
Evans-Pritchard E. E. (1940). The Nuer: a description of the modes of livelihood and political institutions of a Nilotic people. Oxford: Oxford University Press.
Fitch W. T. (2006). The biology and evolution of music: A comparative perspective. Cognition, 100, 173–215.
Fox R. (1980). The red lamp of incest. Dutton Books.
Geissmann T. (2000). Gibbon songs and human music from an evolutionary perspective. In Wallin N. L., Merker B., & Brown S. (eds), The origins of music (p. 103–123). Cambridge, MA: MIT Press.
Geissmann T. (2002). Duet-splitting and the evolution of gibbon songs. Biological Reviews, 77, 57–76.
Geissmann T., & Hulftegger A. M. (1994). Olfactory communication in gibbons? In Roeder J. J., Thierry B., Anderson J. R., & Herrenschmidt N. (eds), Current primatology: Social development, learning and behavior (Vol. 2, p. 199–206). Strasbourg: Universitæ Louis Pasteur.
Getty T. (1987). Dear enemies and the prisoner's dilemma: Why should territorial neighbours form defensive coalitions? American Zoologist, 27, 327–36.
Gosling L. M., & Roberts S. C. (2001). Scent-marking by male mammals: Cheat-proof signals to competitors and mates. Adv Stud Behav, 30, 169–217.
Grinnell J., & McComb K. (1996). Maternal grouping as a defense against infanticide by males: Evidence from field playback experiments on African lions. Behavioral Ecology, 7, 55–9.
Grinnell J., Packer C., & Pusey A. E. (1995). Cooperation in male lions: Kinship, reciprocity, or mutualism. Animal Behaviour, 49, 95–105.
Hagen E. H., & Bryant G. A. (2003). Music and dance as a coalition signalling system. Human Nature, 14, 21–51.
Hall M. L. (2004). A review of hypotheses for the functions of avian duetting. Behavioral Ecology and Sociobiology, 55, 415–30.
Hall M. L., & Magrath R. D. (2007). Temporal coordination signals coalition quality. Current Biology, 17, R406–R407.
Hammerstein P. (1981). The role of asymmetries in animal contests. Animal Behaviour, 29, 193–205.
Hammerstein P. (ed). (2003). Genetic and cultural evolution of cooperation. Cambridge, MA: MIT Press.
Harrington F. H., & Mech L. D. (1979). Wolf howling and its role in territory maintenance. Behaviour, 68, 207–49.
Harrington F. H., & Mech L. D. (1983). Wolf pack spacing: Howling as a territory-independent spacing mechanism in a territorial population. Behavioral Ecology and Sociobiology, 12, 161–8.
Hayden B. (1995). Pathways to power: Principles for creating socioeconomic inequalities. In Price T., & Feinman G. (eds), Foundations of social inequality (p. 15–86). New York: Plenum Press.
Hediger H. (1949). Saugetier-Territorien und ihre Markierung. Bijdr. Dierkd, 28, 172–84.
Heinsohn R. (1997). Group territoriality in two populations of African lions. Animal Behaviour, 53, 1143–47.
Herbinger I., Boesch C., & Rothe H. (2001). Territory characteristics among three neighbouring chimpanzee communities in the Taï National Park, Côte d'Ivoire. International Journal of Primatology, 22, 143–67.
Hofer H., & East M. L. (1993a). The commuting system of Serengeti spotted hyaenas: How a predator copes with migratory prey. I. Animal Behaviour, 46, 547–57.
Hofer H., & East M. L. (1993b). The commuting system of serengeti spotted hyaenas: How a predator copes with migratory prey. II: Intrusion pressure and commuters’ space use. Animal Behaviour, 46, 559–74.
Johnson D. D., Kays R., Blackwell P. G., & Macdonald D. W. (2002). Does the resource dispersion hypothesis explain group living? Trends in Ecology & Evolution, 17, 563–70.
Kappeler M. (1984). Vocal bouts and territorial maintenance in the moloch gibbon. In Preuschoft H. (ed), The lesser apes: Evolutionary and behavioral biology (p. 376–89). Edinburgh: Edinburgh University Press.
Kokko H., López-Sepulcre A., & Morrell L. J. (2006). From hawks and doves to self-consistent games of territorial behaviour. Am. Nat, 167, 901–12.
Kummer H. (1968). Social organization of hamadryas baboons: A field study. Basel: Karger.
Leiser J. K. (2003). When are neighbours “dear enemies” and when are they not? the responses of territorial male variegated pupfish, Cyprinodon variegatus, to neighbours, strangers and heterospecifics. Animal Behaviour, 65, 453–62.
Lerdahl F., & Jackendoff R. (1983). A generative theory of tonal music. Cambridge, MA: MIT Press.
Lowen C., & Dunbar R. I. M. (1994). Territory size and defendability in primates. Behavioral Ecology and Sociobiology, 35, 347–54.
Maher C. R., & Lott D. F. (1995). Definitions of territoriality used in the study of variation in vertebrate spacing systems. Animal behaviour, 49, 1581–97.
Maher C. R., & Lott D. F. (2000). A review of ecological determinants of territoriality within vertebrate species. The American Midland Naturalist, 143, 1–29.
Maynard Smith J., & Parker G. A. (1976). The logic of asymmetric contests. Animal Behaviour, 24, 159–75.
Merker B. (2000). Synchronous chorusing and human origins. In Wallin N. L., Merker B., & Brown S. (eds), The origins of music (p. 315–27). Cambridge, MA: MIT Press.
Merker B. (2005). The conformal motive in birdsong, music, and language: An introduction. Annals of the New York Academy of Sciences, 1060, 17–28.
Merker B. (2006). The uneven interface between culture and biology in human music. Music Perception, 24, 95–8.
Miller G. (2000). Evolution of human music through sexual selection. In Wallin N. L., Merker B., & Brown S. (eds), The origins of music (p. 271–300). Cambridge, MA: MIT Press.
Mills M. G. L. (1990). Kalahari hyaenas. London: Unwin Hyman.
Mitani J. C. (1990). Experimental field studies of Asian ape social systems. International Journal of Primatology, 11, 103–26.
Mitani J. C., & Rodman P. S. (1979). Territoriality: The relation of ranging pattern and home range size to defendability, with an analysis of territoriality among primate species. Behavioral Ecology and Sociobiology, 5, 241–51.
Mithen S. J. (2005). The singing Neanderthals: The origins of music, language, mind and body. London: Weidenfeld & Nicolson.
Müller C., & Manser M. (2007). “Nasty neighbours” rather than “dear enemies” in a social carnivore. Proceedings of the Royal Society B: Biological Sciences, 274, 959–65.
Nevison C. M., Armstrong S., Beynon R. J., Humphries R. E., & Hurst J. L. (2003). The ownership signature in mouse scent marks is involatile. Proceedings: Biological Sciences, 270, 1957–63.
O'Connell J. F., Hawkes K., Lupo K. D., & Blurton Jones N. G. (2002). Male strategies and Plio-Pleistocene archaeology. Journal of Human Evolution, 43, 831–72.
Packer C., Hilborn R., Mosser A., Kissui B., Borner M., Hopcraft G., (2005). Ecological change, group territoriality, and population dynamics in Serengeti lions. Science, 307, 390–3.
Palombit R. A. (1993). Lethal territorial aggression in a white-handed gibbon. American Journal of Primatology, 31, 311–18.
Pereira H. M., Bergman A., & Roughgarden J. (2003). Socially stable territories: The negotiation of space by interacting foragers. American Naturalist, 161, 143–52.
Pinker S. (1997). How the mind works. New York: W. W. Norton & Company.
Potter J. M. (2000). Pots, parties, and politics: Communal feasting in the American Southwest. American Antiquity, 65(3), 471–92.
Raaum R. L., Sterner K. N., Noviello C. M., Stewart C. B., & Disotell T. R. (2005). Catarrhine primate divergence dates estimated from complete mitochondrial genomes: concordance with fossil and nuclear DNA evidence. Journal of Human Evolution, 48, 237–57.
Ralls K. (1971). Mammalian scent marking. Science, 171, 443–9.
Randall J. A. (1984). Territorial defense and advertisement by footdrumming in bannertail kangaroo rats (Dipodomys spectabilis) at high and low population densities. Behavioral Ecology and Sociobiology, 16, 11–20.
Randall J. A. (2001). Evolution and function of drumming as communication in mammals. American Zoologist, 41, 1143–56.
Reichard U. H. (2003). Social monogamy in gibbons: The male perspective. In Reichard U. H., & Boesch C. (eds), Monogamy: Mating strategies and partnerships in birds, humans and other mammals (p. 190–213). New York: Cambridge University Press.
Richards M. P., Pettitt P. B., Trinkaus E., Smith F. H., Paunovic M., & Karavanic I. (2000). Neanderthal diet at Vindija and Neanderthal predation: The evidence from stable isotopes. Proc Natl Acad Sci USA, 97, 7663–6.
Rodseth L., Wrangham R. W., Harrigan A. M., & Smuts B. B. (1991). The human community as a primate society. Current Anthropology, 32, 221–54.
Roebroeks W. (2001). Hominid behaviour and the earliest occupation of Europe: An exploration. Journal of Human Evolution, 41, 437–61.
Rohland N., Pollack J. L., Nagel D., Beauval C., Airvaux J., Pääbo S., (2005). The population history of extant and extinct hyenas. Molecular Biology and Evolution, 22, 2435–43.
Rosell F., & Nolet B. A. (1997). Factors affecting scent-marking behaviour in Eurasian beaver (Castor fiber). Journal of Chemical Ecology, 23, 673–89.
Schmidt P. A., & Mech L. D. (1997). Wolf pack size and food acquisition. The American Naturalist, 150, 513–17.
Slater P., & Mann N. (2004). Why do the females of many bird species sing in the tropics? Journal of Avian Biology, 35, 289–94.
Smith D. R., & Dworkin M. (1994). Territorial interactions between two Myxococcus species. Journal of Bacteriology, 176, 1201–5.
Spence M. (1973). Job market signalling. The Quarterly Journal of Economics, 87, 355–74.
Stanford C. B. (1998). The social behaviour of chimpanzees and bonobos: Empirical evidence and shifting assumptions. Current Anthropology, 39, 399–420.
Stiner M. C. (1992). Overlapping species “choice” by Italian Upper Pleistocene predators. Current Anthropology, 33, 433–51.
Stiner M. C. (2002). Carnivory, coevolution, and the geographic spread of the genus Homo. Journal of Archaeological Research, 10, 1–63.
Stringer C. (2003). Human evolution: Out of Ethiopia. Nature, 423, 747–52.
Számadó S., & Szathmáry E. (2006). Selective scenarios for the emergence of natural language. Trends in Ecology & Evolution, 21, 555–61.
Thorpe W. H. (1961). Bird-song: The biology of vocal communication and expression in birds. Cambridge, UK: Cambridge University Press.
Trainer J. M., McDonald D. B., & Learn W. A. (2002). The development of coordinated singing in cooperatively displaying long-tailed manakins. Behavioral Ecology, 13, 65–9.
Ujhelyi M. (1996). Is there any intermediate stage between animal communication and language? Journal of Theoretical Biology, 180, 71–6.
Ujhelyi M. (2000). Territorial song and facial gesture: A language precursor in apes. Behavioral and Brain Sciences, 22, 572–3.
Vila C., Amorim I. R., Leonard J. A., Posada D., Castroviejo J., Petrucci-Fonseca F., (1999). Mitochondrial DNA phylogeography and population history of the grey wolf Canis lupus. Molecular Ecology, 8, 2089–103.
Wich S. A., & Nunn C. L. (2002). Do male “long-distance calls” function in mate defense? a comparative study of long-distance calls in primates. Behavioral Ecology and Sociobiology, 52, 474–84.
Wiessner P., & Schiefenhovel W. (eds). (1996). Food and the status quest: An interdisciplinary perspective. Providence: Berghahn Books.
Wilson M. L., Hauser M. D., & Wrangham R. W. (2001). Does participation in intergroup conflict depend on numerical assessment, range location, or rank for wild chimpanzees? Animal Behaviour, 61, 1203–16.
Yamaguchi N., Cooper A., Werdelin L., & Macdonald D. W. (2004). Evolution of the mane and group-living in the lion (Panthera leo): A review. Journal of Zoology, 263, 329–42.
Zahavi A. (1975). Mate selection-a selection for a handicap. Journal of Theoretical Biology, 53, 205–14.

Cite article

Cite article

Cite article

OR

Download to reference manager

If you have citation software installed, you can download article citation data to the citation manager of your choice

Share options

Share

Share this article

Share with email
EMAIL ARTICLE LINK
Share on social media

Share access to this article

Sharing links are not relevant where the article is open access and not available if you do not have a subscription.

For more information view the Sage Journals article sharing page.

Information, rights and permissions

Information

Published In

Article first published online: September 1, 2009
Issue published: September 2009

Keywords

  1. biomusicology
  2. territorial signaling
  3. coalitions
  4. alliances
  5. feasting.

Rights and permissions

© 2009 European Society for the Cognitive Sciences of Music.

Authors

Affiliations

Edward H. Hagen
Department of Anthropology, Washington State University
Peter Hammerstein
Institute for Theoretical Biology, Humboldt University

Notes

Department of Anthropology Washington State University 14204 NE Salmon Creek Avenue, Vancouver, WA 98686–9600, USA [email protected]

Metrics and citations

Metrics

Journals metrics

This article was published in Musicae Scientiae.

VIEW ALL JOURNAL METRICS

Article usage*

Total views and downloads: 642

*Article usage tracking started in December 2016


Altmetric

See the impact this article is making through the number of times it’s been read, and the Altmetric Score.
Learn more about the Altmetric Scores



Articles citing this one

Receive email alerts when this article is cited

Web of Science: 57 view articles Opens in new tab

Crossref: 55

  1. The origins of musicality in the motion of primates
    Go to citation Crossref Google Scholar
  2. Wie ist Rhythmus an Interaktion beteiligt? Sportsoziologische Potenzia...
    Go to citation Crossref Google Scholar
  3. How music-induced emotions affect sexual attraction: evolutionary impl...
    Go to citation Crossref Google Scholar
  4. Elephant on the Dance Floor: Revealing the Significance of Dancers’ Se...
    Go to citation Crossref Google Scholar
  5. Biyokültürel bakış açısıyla popüler müzikte üretilen ve seçilen benzer...
    Go to citation Crossref Google Scholar
  6. Pace setting as an adaptive precursor of rhythmic musicality
    Go to citation Crossref Google Scholar
  7. Music as aposematic signal: predator defense strategies in early human...
    Go to citation Crossref Google Scholar
  8. Imprinting-like effects of early adolescent music
    Go to citation Crossref Google Scholar
  9. Psychedelia: The interplay of music and psychedelics
    Go to citation Crossref Google Scholar
  10. Sex-related communicative functions of voice spectral energy in human ...
    Go to citation Crossref Google Scholar
  11. Nocturnal selective pressures on the evolution of human musicality as ...
    Go to citation Crossref Google Scholar
  12. Universal interpretations of vocal music
    Go to citation Crossref Google Scholar
  13. Free rider recognition—A missing link in the Baldwinian model of music...
    Go to citation Crossref Google Scholar
  14. Conceptual and Theoretical Exploration of Music Tourism
    Go to citation Crossref Google Scholar
  15. The Biological Roots of Music and Dance
    Go to citation Crossref Google Scholar
  16. Darwin’s sexual selection hypothesis revisited: Musicality increases s...
    Go to citation Crossref Google Scholar
  17. Development of synchrony‐dominant expectations in observers
    Go to citation Crossref Google Scholar
  18. Pitch syntax as part of an ancient protolanguage
    Go to citation Crossref Google Scholar
  19. Musicality in human vocal communication: an evolutionary perspective
    Go to citation Crossref Google Scholar
  20. Group dancing as the evolutionary origin of rhythmic entrainment in hu...
    Go to citation Crossref Google Scholar
  21. Signals and cues of social groups
    Go to citation Crossref Google Scholar
  22. Music Listening and Homeostatic Regulation: Surviving and Flourishing ...
    Go to citation Crossref Google Scholar
  23. On reappearance and complexity in musical calling
    Go to citation Crossref Google Scholar
  24. The Role of Canalization and Plasticity in the Evolution of Musical Cr...
    Go to citation Crossref Google Scholar
  25. From Animal Signals to Art: Manipulative Animal Signaling and the Evol...
    Go to citation Crossref Google Scholar
  26. Edward Hagen
    Go to citation Crossref Google Scholar
  27. Art Production, Appreciation, and Fitness
    Go to citation Crossref Google Scholar
  28. Origins of music in credible signaling
    Go to citation Crossref Google Scholar
  29. Toward a productive evolutionary understanding of music
    Go to citation Crossref Google Scholar
  30. Recognizing affiliation in colaughter and cospeech
    Go to citation Crossref Google Scholar
  31. Changes in behavioural synchrony during dog‐assisted therapy for child...
    Go to citation Crossref Google Scholar
  32. Music and the Evolution of Embodied Cognition
    Go to citation Crossref Google Scholar
  33. Function of private singing in instrumental music learning: A multiple...
    Go to citation Crossref Google Scholar
  34. Music and Language in Social Interaction: Synchrony, Antiphony, and Fu...
    Go to citation Crossref Google Scholar
  35. Singing to the self: Children’s private speech, private song, and exec...
    Go to citation Crossref Google Scholar
  36. Art Production, Appreciation and Fitness
    Go to citation Crossref Google Scholar
  37. Which Melodic Universals Emerge from Repeated Signaling Games? A Note ...
    Go to citation Crossref Google Scholar
  38. Edward Hagen
    Go to citation Crossref Google Scholar
  39. Plio-Pleistocene Foundations of Hominin Musicality: Coevolution of Cog...
    Go to citation Crossref Google Scholar
  40. The Role of the Baldwin Effect in the Evolution of Human Musicality
    Go to citation Crossref Google Scholar
  41. Simultaneous Cooperation and Competition in the Evolution of Musical B...
    Go to citation Crossref Google Scholar
  42. Harmonious Contact: Stories About Intergroup Musical Collaboration Imp...
    Go to citation Crossref Google Scholar
  43. Synchronized behavior increases assessments of the formidability and c...
    Go to citation Crossref Google Scholar
  44. “It Don’t Mean a Thing if It Ain’t Got that Swing”– an Alternative Con...
    Go to citation Crossref Google Scholar
  45. What Pinnipeds Have to Say about Human Speech, Music, and the Evolutio...
    Go to citation Crossref Google Scholar
  46. Evolving perceptual biases for antisynchrony: a form of temporal coord...
    Go to citation Crossref Google Scholar
  47. Four principles of bio-musicology
    Go to citation Crossref Google Scholar
  48. Körperliche Rhythmisierung und rituelle Interaktion
    Go to citation Crossref Google Scholar
  49. Traces across the body: the influence of music-dance synchrony on the ...
    Go to citation Crossref Google Scholar
  50. The evolution of coordinated vocalizations before language
    Go to citation Crossref Google Scholar
  51. Vocal learning, prosody, and basal ganglia: Don't underestimate their ...
    Go to citation Crossref Google Scholar
  52. The Evolutionary Origins of Rhythm: A Top-down/bottom-up Approach to T...
    Go to citation Crossref Google Scholar
  53. Music marketing: A history and landscape
    Go to citation Crossref Google Scholar
  54. Strategic Aspects of Communication
    Go to citation Crossref Google Scholar
  55. Gestures of Despair and Hope: A View on Deliberate Self-harm From Econ...
    Go to citation Crossref Google Scholar

Figures and tables

Figures & Media

Tables

View Options

Get access

Access options

If you have access to journal content via a personal subscription, university, library, employer or society, select from the options below:


Alternatively, view purchase options below:

Purchase 24 hour online access to view and download content.

Access journal content via a DeepDyve subscription or find out more about this option.

View options

PDF/ePub

View PDF/ePub