The evolution of anterior coloration in carnivorans

  • Tim Caro
  • Hannah Walker
  • Sharlene E. Santana
  • Theodore Stankowich
Original Article


Some carnivorans have striking patches of fur on their faces (spots, bands, eye masks) and blazes on their chests that are primarily visible from a frontal view. We tested five hypotheses to explain the evolution of the complexity and contrast of these color patches. These were: signals of species identity to avoid hybridization, communication between conspecifics, signals used to warn of defensive anal secretions, signals of belligerence or pugnacity, and camouflage-related coloration used to break up the outline and facial features of the predator when approaching prey. Using phylogenetically controlled multifactorial analyses in six different families of carnivorans, examined separately, our analyses uncovered significant positive associations between measures of color pattern complexity and sociality across herpestid faces and canid chests, suggesting use in social communication. Mustelid facial color complexity was associated with ability to direct anal secretions accurately at predators, and facial markings were significantly or marginally associated with pugnacity in mustelids, viverrids, and herpestids. Facial complexity of viverrid and herpestid species was significantly or marginally related to a mammal-based diet. In ursids, facial contrast appeared less variable in species living in greater sympatry with other bears. Facial and chest coloration in Carnivora appears to have evolved under different selection pressures in different families.

Significance statement

The reasons that many carnivorans have colorful and memorable faces and chests are not yet understood. Here, we pit five different hypotheses against each other: species recognition, advertising either toxic anal defenses or pugnacity, recognizing group members, and trying to remain concealed when approaching mammalian prey. We find that measures of facial and chest complexity and contrast have evolved for different reasons depending on the carnivoran family. Anterior coloration appears to be involved with social communication in herpestids and canids; facial coloration is associated noxious secretions in mustelids, with pugnacity in mustelids, viverrids and herpestids; with reliance on a mammal-based diet in viverrids and herpestids; and with avoiding hybridization in bear species. There is no overriding evolutionary explanation for varied facial and chest pelage coloration across carnivorans.


Carnivores Chests Color complexity Contrast Faces 



We thank anonymous reviewers for their comments.

Compliance with ethical standards

Ethical statement

All sources of data were from the literature or the web and did not involve ethical approval. There was no funding for this project.

Conflict of interest

The authors declare that they have no conflict of interest.

Data availability

All data generated or analyzed during this study are included in the supplementary information files.

Supplementary material

265_2017_2402_MOESM1_ESM.docx (68 kb)
ESM 1 (DOCX 67 kb)
265_2017_2402_MOESM2_ESM.xlsx (23 kb)
ESM 2 (XLSX 22 kb)


  1. Allen WL, Cuthill IC, Scott-Samuel N, Baddeley R (2010) Why the leopard got its spots: relating pattern development to ecology in felids. Proc R Soc B (2011) 278:1373–1380.
  2. Allen WL, Stevens M, Higham JP (2014) Character displacement of Cercopithecini primate visual signals. Nat Commun 5:4266CrossRefPubMedPubMedCentralGoogle Scholar
  3. Arnold C, Matthews LJ, Nunn CL (2010) The 10kTrees website: a new online resource for primate phylogeny. Evol Anthropol 19:114–118CrossRefGoogle Scholar
  4. Caro T (2009) Contrasting colouration in terrestrial mammals. Philos T Roy Soc B 364:537–548CrossRefGoogle Scholar
  5. Caro T (2013) The colours of extant mammals. Sem cell Devel Biol 24:542–552CrossRefGoogle Scholar
  6. Caro T, Allen WL (2017) Interspecific visual signalling in animals and plants: a functional classification. Philos T Roy Soc B 372:20160344CrossRefGoogle Scholar
  7. Caro T, Walker H, Rossman Z, Hendrix M, Stankowich T (2017) Why is the giant panda black and white? Behav Ecol 28:657–667CrossRefGoogle Scholar
  8. Cott HB (1940) Adaptive coloration in animals. Methuen & Co. Ltd, LondonGoogle Scholar
  9. Davis AK, Woodall N, Moskowitz JP, Castleberry N, Freeman BJ (2013) Temporal change in fur color in museum specimens of mammals: reddish-brown species get redder with storage time. Int J Zool 2013:876347CrossRefGoogle Scholar
  10. Francis CM (2008) A guide to the mammals of Southeast Asia. Princeton University Press, PrincetonGoogle Scholar
  11. Gorman ML, Trowbridge BJ (1989) The role of odor in the social lives of carnivores. In: Gittleman JL (ed) Carnivore behavior, ecology, and evolution. Chapman & Hall, London, pp 57–88CrossRefGoogle Scholar
  12. Hill GE (2015) Sexiness, individual condition, and species identity: the information signaled by ornaments and assessed by choosing females. Evol Biol 42:251–259CrossRefGoogle Scholar
  13. Hunter L (2011) Carnivores of the world. Princeton University Press, Princeton, NJGoogle Scholar
  14. Kelly BP, Whiteley A, Tallmon D (2010) The Arctic melting pot. Nature 468:891–891CrossRefPubMedGoogle Scholar
  15. Kingdon J (1977) East African mammals. 3, part a (carnivores). Academic Press, LondonGoogle Scholar
  16. Lehman N, Eisenhawer A, Hansen K, Mech LD, Peterson RO, Gogan PJ, Wayne RK (1991) Introgression of coyote mitochondrial DNA into sympatric North American gray wolf populations. Evolution 45:104–119CrossRefPubMedGoogle Scholar
  17. Myers P, Espinosa R, Parr CS, Jones T, Hammond GS, Dewey TA (2013) The Animal Diversity Web (online),
  18. Newman C, Buesching CD, Wolff JO (2005) The function of facial masks in “midguild” carnivores. Oikos 108:623–633CrossRefGoogle Scholar
  19. Nowak RM (1999) Walker’s mammals of the world, 6th edn. Johns Hopkins University Press, BaltimoreGoogle Scholar
  20. Orme D, Freckleton R, Thomas G, Petzoldt T, Fritz S, Isaac N, Pearse W (2012) Caper: comparative analysis of phylogenetics and evolution in R. R package version 0.5,
  21. Ortolani A (1999) Spots, stripes, tail tips and dark eyes: predicting the function of carnivore colour patterns using the comparative method. Biol J Linn Soc 67:433–476CrossRefGoogle Scholar
  22. Ortolani A, Caro T (1996) The adaptive significance of color patterns in carnivores: phylogenetic tests of classic hypotheses. In: Gittleman JL (ed) Carnivore behavior, ecology and evolution, vol II. Cornell University press, Ithaca, New York, pp 132–188Google Scholar
  23. Osorio D, Vorobyev M (1996) Colour vision as an adaptation to frugivory in primates. Proc R Soc Lond B 263:593–599CrossRefGoogle Scholar
  24. R Development Core Team (2008) R: a language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria Google Scholar
  25. Santana SE, Alfaro JL, Alfaro ME (2012) Adaptive evolution of facial colour patterns in Neotropical primates. Proc R Soc Lond B 279:2204–2211CrossRefGoogle Scholar
  26. Santana SE, Alfaro JL, Noonan A, Alfaro ME (2013) Adaptive response to sociality and ecology drives the diversification of facial colour patterns in catarrhines. Nat Commun 4:2765CrossRefPubMedGoogle Scholar
  27. Stankowich T, Caro T, Cox M (2011) Bold coloration and the evolution of aposematism in terrestrial carnivores. Evolution 65:3090–3099CrossRefPubMedGoogle Scholar
  28. Stankowich T, Havercamp P, Caro T (2014) Ecological drivers of antipredator defenses in carnivores. Evolution 68:1415–1425CrossRefPubMedGoogle Scholar
  29. Stoner CJ, Caro TM, Graham CM (2003) Ecological and behavioral correlates of coloration in artiodactyls: systematic analyses of conventional hypotheses. Behav Ecol 14:823–840CrossRefGoogle Scholar
  30. Tibbetts EA, Mullen SP, Dale J (2017) Signal function drives phenotypic and genetic diversity: the effects of signaling individual identity, quality or behavioural strategy. Philos T Roy Soc B 372:20160347CrossRefGoogle Scholar
  31. Van Dyck S (2006) In: Strahan R (ed) The mammals of Australia. New Holland Publishing, SydneyGoogle Scholar
  32. Wilson DE, Mittermeier RA (2009) Handbook of the mammals of the world, vol 1. Carnivores. Lynx Edicions, BarcelonaGoogle Scholar
  33. Wilson DE, Reeder DM (eds) (2005) Mammal species of the world: a taxonomic and geographic reference. John Hopkins University Press, BaltimoreGoogle Scholar

Copyright information

© Springer-Verlag GmbH Germany, part of Springer Nature 2017

Authors and Affiliations

  • Tim Caro
    • 1
  • Hannah Walker
    • 2
  • Sharlene E. Santana
    • 3
  • Theodore Stankowich
    • 2
  1. 1.Department of Wildlife, Fish and Conservation BiologyUniversity of CaliforniaDavisUSA
  2. 2.Department of Biological SciencesCalifornia State UniversityLong BeachUSA
  3. 3.Department of Biology and Burke Museum of Natural Hosry and CultureUniversity of WashingtonSeattleUSA

Personalised recommendations