Hanlon Lab – Camouflage & Adaptive Coloration

Nearly all animals have some sort of adaptive coloration or camouflage patterning that is often linked to a behavior to make it adaptive.

Some of the functions of adaptive coloration include:

  • defense against predators,
  • communication with conspecifics,
  • attracting or deceiving mates,
  • repelling or deceiving rivals,
  • signalling alarm to conspecifics and so forth,
  • protection from the environment (e.g., ultraviolet radiation, cold), and
  • approaching prey

iBioSeminar: Concepts and questions (28:00 min)


A primary defense against predators throughout the animal kingdom (and against the enemy during human warfare) is to avoid detection or recognition through camouflage. Achieving effective camouflage requires a suite of appropriate actions by an animal:

  1. sensing the local environment (including the animals in it),
  2. filtering the sensory input,
  3. using selected sensory input to make a behavior decision,
  4. directing the appropriate effectors (be they muscles/postures/color patterns, etc.) to achieve some form of camouflage, and
  5. implementing the appropriate behavior to render the camouflage effective.

How many kinds of camouflage are there?

There is still active debate on how to best "categorize" camouflage. Generally the tactics involve hindering or preventing detection or recognition. Some of the generally accepted mechanisms of camouflage include:

  • general background resemblance (or "background matching,"
  • deceptive resemblance (or "masquerade" including mimicry),
  • disruptive coloration,
  • countershading/concealment of the shadow.

Other mechanisms include self-shadow concealment, obliterative shading, distractive markings, flicker-fusion camouflage, motion dazzle and motion camouflage. Ongoing research worldwide is occurring on many of these features.

Cephalopods: ultimate adaptive coloration?

A distinguishing feature of cephalopods is that individual animals can change their appearance with a speed and diversity unparalleled in the animal kingdom: we term this “rapid, neurally controlled polymorphism.” Some squids, octopuses and cuttlefish can show 30-50 different appearances. In fact, these marine invertebrates manifest most aspects of their behavior through body patterning. An example of their versatility is that – unlike other animals that use one or a few mechanisms of camouflage – cephalopods use most of the mechanisms listed above.

Sensory/motor mechanisms of achieving adaptive coloration in cephalopods

Due to their sophisticated neural control of the skin, cephalopods can adapt to a wide range of backgrounds. What sensory cues do they use to achieve background matching? Vision is probably the main cue, but cephalopods do not seem to have color vision. We are currently investigating the mechanisms and functions of polarization sensitivity in cephalopods. We have also begun to look at visual features of the background that cuttlefish use to switch on disruptive coloration.

Tactile cues seem not to be used by cuttlefish for regulating their skin texture - vision seems to be used. Olfactory cues are used by cuttlefish females to choose mates, yet we do not know if/how this translates to certain body patterns for communication. Read more about skin ultrastructure and neurobiology.

How do you study camouflage?

Field work is mandatory to experience and understand the dynamic features of light throughout each daily, lunar, seasonal, and yearly cycle. Field work also enables observation and analysis of animal behavior under natural conditions. Laboratory experiments provide detailed testing of sensory cues, motor output and sequences of behavior. Computer simulations allow additional testing of hypotheses at multiple levels of analysis.

Publications on adaptive coloration and behavior:

Allen, J. J., Akkaynak, D., Sugden, A. U., and Hanlon, R. T. (2015). Adaptive body patterning, three-dimensional skin morphology and camouflage measures of the slender filefish Monacanthus tuckeri on a Caribbean coral reef. Biological Journal of the Linnean Society, 116(2), 377–396. doi: 10.1111/bij.12598

Buresch, K. C., Ulmer, K. M., Cramer, C., McAnulty, S., Davison, W., Mathger, L. M., and Hanlon, R. T. (2015). Tactical Decisions for Changeable Cuttlefish Camouflage: Visual Cues for Choosing Masquerade Are Relevant from a Greater Distance than Visual Cues Used for Background Matching. Biological Bulletin, 229(2), 160 – 166. Pubmed: 26504156

Chiao, C. C., Chubb, C., and Hanlon, R. T. (2015). A review of visual perception mechanisms that regulate rapid adaptive camouflage in cuttlefish. Journal of Comparative Physiology A-Sensory Neural and Behavioral Physiology, 201(9), 933–945. doi: 10.1007/s00359-015-0988-5

Tyrie, E. K., Hanlon, R. T., Siemann, L. A., and Uyarra, M. C. (2015). Coral reef flounders, Bothus lunatus , choose substrates on which they can achieve camouflage with their limited body pattern repertoire. Biological Journal of the Linnean Society, 114(3), 629–638. doi: 10.1111/bij.12442

Buresch, K. C., Ulmer, K. M., Akkaynak, D., Allen, J. J., Maethger, L. M., Nakamura, M., and Hanlon, R. T. (2015). Cuttlefish adjust body pattern intensity with respect to substrate intensity to aid camouflage, but do not camouflage in extremely low light. Journal of Experimental Marine Biology and Ecology, 462, 121–126. doi: 10.1016/j.jembe.2014.10.017

Watson, A. C., Siemann, L. A., and Hanlon, R. T. (2014). Dynamic camouflage by Nassau groupers Epinephelus striatus on a Caribbean coral reef. Journal of Fish Biology, 85(5), 1634 – 1649. doi: 10.1111/jfb.12519

Akkaynak, D., Treibitz, T., Xiao, B., Gurkan, U.A., Allen, J.J., Demirci, U. and Hanlon, R.T. (2014). Use of commercial off-the-shelf digital cameras for scientific data acquisition and scene-specific color calibration. Journal of the Optical Society of America 31(2), 312-321. doi: 10.1364/JOSAA.31.000312

Chiao, C. C., Ulmer, K. M., Siemann, L. A., Buresch, K. C., Chubb, C., and Hanlon, R. T. (2013). How visual edge features influence cuttlefish camouflage patterning. Vision Research, 83, 40–7. doi: 10.1016/j.visres.2013.03.001

Staudinger, M. D., Buresch, K. C., Mäthger, L. M., Fry, C., McAnulty, S., Ulmer, K. M., and Hanlon, R. T. (2013). Defensive responses of cuttlefish to different teleost predators. The Biological Bulletin, 225(3), 161–74. Pubmed: 24445442

Ulmer, K. M., Buresch, K. C., Kossodo, M. M., Mäthger, L. M., Siemann, L. A., and Hanlon, R. T. (2013). Vertical visual features have a strong influence on cuttlefish camouflage. The Biological Bulletin, 224(2), 110–8. Pubmed: 23677976

Akkaynak, D., Allen, J. J., Mäthger, L. M., Chiao, C.-C., and Hanlon, R. T. (2013). Quantification of cuttlefish (Sepia officinalis) camouflage: a study of color and luminance using in situ spectrometry. Journal of Comparative Physiology. A Neuroethology, Sensory, Neural, and Behavioral Physiology, 199(3), 211–25. doi: 10.1007/s00359-012-0785-3

Hanlon, R. T., Chiao, C.-C., Mäthger, L. M., and Marshall, N. J. (2013). A fish-eye view of cuttlefish camouflage using in situ spectrometry. Biological Journal of the Linnean Society, 109(3), 535–551. doi: 10.1111/bij.12071

Mäthger, L. M., and Hanlon, R. T. (2012). Pigmentation or transparency? Camouflage tactics in deep-sea cephalopods. Pigment Cell & Melanoma Research, 25(3), 295–296. doi: 10.1111/j.1755-148X.2012.00985.x

Barbosa, A., Allen, J. J., Mäthger, L. M., and Hanlon, R. T. (2012). Cuttlefish use visual cues to determine arm postures for camouflage. Proceedings. Biological Sciences / The Royal Society, 279(1726), 84–90. doi: 10.1098/rspb.2011.0196

Buresch, K. C., Mäthger, L. M., Allen, J. J., Bennice, C., Smith, N., Schram, J., Chiao, C.C., and Hanlon, R. T. (2011). The use of background matching vs. masquerade for camouflage in cuttlefish Sepia officinalis. Vision Research, 51(23-24), 2362–8. doi: 10.1016/j.visres.2011.09.009

Zylinski, S., How, M.J., Osorio, D., Hanlon, R.T. and Marshall, N.J. (2011). To be seen or to hide: visual characteristics of body patterns for camouflage and communication in the Australian giant cuttlefish, Sepia apama. American Naturalist, 177(5), 681-690. doi: 10.1086/659626

Chiao, C.-C., Wickiser, J. K., Allen, J. J., Genter, B., and Hanlon, R. T. (2011). Hyperspectral imaging of cuttlefish camouflage indicates good color match in the eyes of fish predators. Proceedings of the National Academy of Sciences of the United States of America, 108(22), 9148–53. doi: 10.1073/pnas.1019090108

Staudinger, M. D., Hanlon, R. T., and Juanes, F. (2011). Primary and secondary defences of squid to cruising and ambush fish predators: variable tactics and their survival value. Animal Behaviour, 81(3), 585–594. doi: 10.1016/j.anbehav.2010.12.002

Allen, J.J., Mäthger,L.M., Buresch, K.C., Fetchko, T. Gardner, M. and Hanlon, R.T. (2010) Night vision by cuttlefish enables changeable camouflage. J. Experimental Biology 213: 3953-3960. doi:

Allen, J. J., Mäthger, L. M., Barbosa, A., Buresch, K. C., Sogin, E., Schwartz, J., and Hanlon, R. T. (2010). Cuttlefish dynamic camouflage: responses to substrate choice and integration of multiple visual cues. Proceedings. Biological Sciences / The Royal Society, 277 (1684), 1031–9. doi:

Hanlon, R.T., Watson, A.C., and Barbosa, A. (2010). A “mimic octopus” in the Atlantic: flatfish mimicry and camouflage by Macrotritopus defilippi. Biological Bulletin 218 (1): 15-24. (Editor’s Choice Article and Cover Shot) Pubmed: 20203250

Chiao, C.C., Chubb, C., Buresch, K., Allen, J., Barbosa, A., Mäthger, L.M., Hanlon, R.T. (2010). Mottle camouflage patterns in cuttlefish: quantitative characterization and visual stimuli that evoke them. Journal of Experimental Biology 213: 187-199. doi:

Hanlon, R.T., & Messenger, J.B. 1996. Cephalopod Behaviour. Cambridge University Press.

Hanlon R.T., Chiao C-C., Mäthger, L.M., Barbosa A., Buresch K.C., & Chubb, C. 2009. Cephalopod dynamic camouflage: bridging the continuum between background matching and disruptive coloration. Philosophical Transactions of the Royal Society B 364: 429-437.

Williams, S.B., Pizarro, O., How, M., Mercer, D., Powell, G., Marshall, J., & Hanlon, R. 2009. Surveying nocturnal cuttlefish camouflage behaviour using an AUV. IEEE International Conference on Robotics and Automation: 214-219.

Mäthger, L.M., Shashar, N., & Hanlon, R.T. 2009. Do cephalopods communicate using polarized light reflections from their skin? Journal of Experimental Biology 212: 2133-2140. (Featured Commentary)

Mäthger, L.M., Barbosa, A., & Hanlon, R.T. 2009. Cuttlefish use visual cues to control 3-dimensional skin papillae for camouflage. Journal of Comparative Physiology A 195: 547-555.

Chiao, C-C., Chubb, C., Buresch, K., Siemann, L., & Hanlon, R.T. 2009. The scaling effects of substrate texture on camouflage patterning in cuttlefish. Vision Research 49 (13): 1647-1656.

Mäthger, L.M., Denton, E.J., Marshall, J., & Hanlon, R.T. 2008. Mechanisms and behavioral functions of structural coloration in cephalopods. Journal of the Royal Society Interface 6: S149-S164.

Hanlon, R.T., Conroy, L.-A., & Forsythe, J.W. 2008. Mimicry and foraging behaviour of two tropical sand-flat octopus species off North Sulawesi, Indonesia. Biological Journal of the Linnean Society, 93: 23-38.

Mäthger, L.M., Chiao, C-C., Barbosa, A, Buresch, K., Kaye, S., & Hanlon, R.T. 2007. Disruptive coloration elicited on controlled natural substrates in cuttlefish, Sepia officinalis J. Exp. Biol. 210: 2657-2666.

Mäthger, L.M., & Cronin, T.W. 2007. Spectral and spatial properties of the polarized light reflections on the arms of squid (Loligo pealeii) and cuttlefish (Sepia officinalis L.). Journal of Experimental Biology. 210: 3624-3635.

Mäthger, L.M., & Hanlon, R.T. 2007. Malleable skin coloration in cephalopods: selective reflectance, transmission and absorbance of light by chromatophores and iridophores. Cell and Tissue Research 329: 179-186.

Hanlon, R.T. 2007. Cephalopod dynamic camouflage. Current Biology 17 (11): 400-404.

Hanlon, R.T., Naud, M.-J., Forsythe, J.W., Hall, K., Watson, A.C., & McKechnie, J. 2007. Adaptable night camouflage by cuttlefish. American Naturalist 169 (4): 543-551.

Barbosa, A., Mäthger, L.M., Chubb, C., Florio, C., Chiao, C-C., & Hanlon, R.T. 2007. Disruptive coloration in cuttlefish: a visual perception mechanism that regulates ontogenetic adjustment of skin patterning. Journal of Experimental Biology 210: 1139-1147.

Mäthger, L.M., & Hanlon, R.T. 2006. Anatomical basis for camouflaged polarized light communication in squid. Biology Letters 2(4): 494-496.

Chiao, C-C., Kelman, E.J., & Hanlon, R.T. 2005. Disruptive body patterning of cuttlefish (Sepia officinalis) requires visual information regarding edges and contrast of objects in natural substrate backgrounds. Biological Bulletin. 208: 7-11.

Anderson, J.C., Baddeley, R.J., Osorio, D., Shashar, N., Tyler, C.W., Ramachandran, V.S., Cook A.C., & Hanlon, R.T. 2003. Modular organization of adaptive colouration in flounder and cuttlefish revealed by independent component analysis. Network: Comput. Neural Syst. 14: 321-333

Chiao, C-C., & Hanlon, R.T. 2001. Cuttlefish camouflage: visual perception of size, contrast and number of white squares on artificial substrata initiates disruptive coloration. J. Exp. Biol. 204: 2119-2125.

Chiao, C-C., & Hanlon, R.T. 2001. Cuttlefish cue visually on area—not shape or aspect ratio—of light objects in the substrate to produce disruptive body patterns for camouflage. Biol. Bull. 201:269-270.

Hanlon, R.T., Forsythe, J.W., & Joneschild, D.E. 1999. Crypsis, conspicuousness, mimicry and polyphenism as antipredator defences of foraging octopuses on Indo-Pacific coral reefs, with a method of quantifying crypsis from video tapes. Biol. J. Linn. Soc. 66: 1-22.

Hanlon, R.T., Maxwell, M.R., Shashar, N., Loew, E.R., & Boyle, K.-L. 1999. An ethogram of body patterning behavior in the biomedically and commercially valuable squid Loligo pealei off Cape Cod, Massachusetts. Biol. Bull. 197(1): 49-62.

Shashar, N., & Hanlon, R.T. 1997. Squids (Loligo pealei and Euprymna scolopes) can exhibit polarized light patterns produced by their skin. Biol. Bull. 193(2): 207-208.

Forsythe, J.W., & Hanlon, R.T. 1997. Foraging and associated behavior by Octopus cyanea Gray, 1849 on a coral atoll, French Polynesia. J. Exp. Mar. Biol. Ecol. 209: 15-31.

Cornwell, C.J., Messenger, J.B., & Hanlon, R.T. 1997. Chromatophores and body patterning in the squid Alloteuthis subulata. J. mar. biol. Assoc. U.K. 77: 1243-1246.

DiMarco, F.P. ,& Hanlon, R.T. 1997. Agonistic behavior in the squid Loligo plei (Loliginidae, Teuthoidea): Fighting tactics and the effects of size and resource value. Ethology 103(2): 89-108.

Hanlon, R.T., & Messenger, J.B. 1996. Cephalopod Behaviour. Cambridge University Press.

Adamo, S.A., & Hanlon, R.T. 1996. Do cuttlefish (Cephalopoda) signal their intentions to conspecifics during agonistic encounters? Anim. Behav. 52: 73-81.

Hanlon, R.T., Smale, M.J., & Sauer, W.H.H. 1994. An ethogram of body patterning behavior in the squid Loligo vulgaris reynaudii on spawning grounds in South Africa. Biol. Bull. 187(3): 363-372.

Cooper, K.M., Hanlon, R.T., & Budelmann, B.U. 1990. Physiological color change in squid iridophores II. Ultrastructural mechanisms in Lolliguncula brevis. Cell Tissue Res. 259: 15-24.

Hanlon, R.T., Cooper, K.M., Budelmann, B.U., & Pappas, T.C. 1990. Physiological color change in squid iridophores I. Behavior, morphology and pharmacology in Lolliguncula brevis. Cell Tissue Res. 259: 3-14.

Cooper, K.M., Hanlon, R.T., & Budelmann, B.U. 1990. Physiological color change in squid iridophores II. Ultrastructural mechanisms in Lolliguncula brevis. Cell Tissue Res. 259: 15-24.

Novicki, A., Budelmann, B.U., & Hanlon, R.T. 1990. Brain pathways of the chromatophore system in the squid Lolliguncula brevis. Brain Res. 519(1-2): 315-323.

Hanlon, R.T., & Wolterding, M.R. 1989. Behavior, body patterning, growth and life history of Octopus briareus cultured in the laboratory. Am. Malacol. Bull. 7(1): 21-45.

Hanlon, R.T. 1988. Behavioral and body patterning characters useful in taxonomy and field identification of cephalopods. Malacologia 29(1): 247-264.

Hanlon, R.T., Messenger, J.B. 1988. Adaptive coloration in young cuttlefish (Sepia officinalis L.): The morphology and development of body patterns and their relation to behaviour. Phil. Trans. R. Soc. Lond. B 320: 437-487.