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Cai, L., McGuire, N.E., Hanlon, R.T., Mooney, T.A., Girdhar, Y. Semi-supervised visual tracking of marine animals using Autonomous Underwater Vehicles. 2023. International Journal of Computer Vision 131 (6): 1406-1427.

Moniz, H.A., Kruleski, S.M., Risbud, A.D., Louden, S.J.H., Hanlon, R.T., Stokes, A.N., Palmer, S.E., Feldman, C.R. 2023. Aposematic coloration of Pacific newts (Tracha) provides a qualitatively but not quantitatively honest signal to predators. Biol J Linnean Soc. 139 (1): 1-17.

Jones, I.T., Schumm, M., Stanley, J.A., Hanlon, R.T., and Mooney, T.A. 2023. Longfin squid reproductive behaviours and spawning withstand wind farm pile driving noise. ICES Journal of Marine Sciences 80, fsad117.


Kim, D., Buresch, K.C., Hanlon, R.T. and Kampff, A.R. 2022. An experimental method for evoking and characterizing dynamic color patterning of cuttlefish during prey capture. Journal of Biological Methods 9 (2): e161

Buresch, K.C., Sklar, K., Chen, J.Y., Madden, S.R., Mongil, A.S., Wise, G.V., Boal, J.G., Hanlon, R.T. 2022. Contact chemoreception in multi-modal sensing of prey by Octopus. J. Comparative Physiology A 208 (3): 435-442.

Drinkwater E., Allen, W.L., Endler, J.A., Hanlon, R.T. et al (21 authors). 2022. A synthesis of deimatic behaviour. Biological Reviews 97 (6): 2237-2267.

Barrett, C.J. …… Hanlon, R.T. …. Yen, D.T.H. (32 authors). 2022. Cuttlefish conservation: a global view of methods to ameliorate unwanted fishing mortality and other anthropogenic threats to sustainability. ICES Journal of Marine Science 79 (10): 2579-2596.


Senft, S.L., Kuzirian, A.M. and Hanlon, R.T. 2021. Networks of linked radial muscles could influence dynamic skin patterning of squid chromatophores. Journal of Morphology 282(1): 1245-1258.

Abbo, L.A., Himebaugh, N.E., Demelo, L.M., Hanlon, R.T., Crook, R.J. 2021. Anesthetic efficacy of magnesium chloride and ethyl alcohol in temperate octopus and cuttlefish species. J. American Association for Laboratory Animal Science 60 (5): 556-567.

Bennice, CO, Brooks, WR and Hanlon RT. 2021. Behavioral dynamics provide insight into resource exploitation and habitat coexistence of two octopus species in a shallow Florida lagoon. Journal of Experimental Marine Biology and Ecology. 542-543: 151592

Schnell, A.K., Clayton, N.S., Hanlon, R.T., Jozet-Alves, C. 2021. Episodic-like memory is preserved with age in cuttlefish. Proc. Roy. Soc. B. 288: 20211052.


Hanlon, R.T., and McManus, G. 2020. Flamboyant cuttlefish behavior: camouflage tactics and complex colorful reproductive behavior assessed during field studies at Lembeh Strait, Indonesia. J. Experimental Marine Biology and Ecology 529: 151397. DOI: 10.1016/j.jembe.2020.151397

Kennedy, E.B.L., Buresch, K.C., Boinapally, P., and Hanlon R.T. (2020). Octopus arms exhibit exceptional flexibility. Sci Rep 10:20872.


Marian, J.E.A.R., Apostólico, L.H., Chiao, C.C., Hanlon, R.T., Hirohashi, N., Iwata, Y., Mather, J.A., Sato, N., Shaw, P.W. (2019). Male alternative reproductive tactics and associated evolution of anatomical characteristics in Loliginid squid. Frontiers in Physiology, 10(1281). DOI: 10.3389/fphys.2019.01281

Bennice, C.O., Rayburn, A.P., Brooks, W.R. and Hanlon, R.T. (2019). Fine-scale habitat partitioning facilitates sympatry between two octopus species in a shallow Florida lagoon. Marine Ecology Progress Series 609: 151-161. DOI: 10.3354/meps12845

Iglesias, T.L., Boal, J.G., Frank, M.G., Zeil, J. and Hanlon, R.T. (2019). Cyclic nature of the REM-like state in the cuttlefish Sepia officinalis. Journal of Experimental Biology 222. DOI: 10.1242/jeb.174862

Williams, T.L., Senft, S.L., Yeo, J., Martin-Martinez, F.J., Kuzirian, A.M., Martin, C.A., DiBona, C.W., Chen, C.-T., Dineen, S.R., Nguyen, H.T., Gomes, C.M., Rosenthal, J.J.C., MacManes, M.D., Chu, F., Buehler, M.J., Hanlon, R.T., Deravi, L.F. (2019). Dynamic pigmentary and structural coloration within a cephalopod chromatophore organs. Nature Communications 10: 1004. DOI: 10.1038/s41467-019-08891-x

Schnell, A.K., Jozet-Alves, C., Hall, K.C., Radday, L. and Hanlon, R.T. (2019). Fighting and mating success in giant Australian cuttlefish is influenced by behavioural lateralization. Proceedings of the Royal Society B 286(1898):20182507. DOI:10.1098/rspb.2018.2507

Lutz, H., Ramirez-Puebla, T., Abbo, L., Durand, A., Schlundt, C., Gottel, N., Sjaarda, A., Hanlon, R.T., Gilbert, J.A., and J.L. Mark Welch. (2019). A simple microbiome in the European common cuttlefish, Sepia officinalis. mSystems DOI:10.1128/mSystems.00177-19


Gonzalez-Bellido, P.T., Scaros, A.T., Hanlon, R.T. and Wardill, T.J. (2018). Neural control of dynamic 3-dimensional skin papillae for cuttlefish camouflage. iScience 1: 24-34. DOI: 10.1016/j.isci.2018.01.001

Hanlon, R.T., Mäthger, L.M., Bell, G.R.R., Kuzirian, A.M & Senft, S.L. (2018). White reflection from cuttlefish skin leucophores. Bioinspiration and Biomimetics 13: 035002. DOI: 10.1088/1748-3190/aaa3a9

Chubb, C., Chiao, C.-C., Ulmer, K.M., Buresch, K.C., Birk, M. and Hanlon, R.T. (2018). Dark scene elements strongly influence cuttlefish camouflage responses in visually cluttered environments. Vision Research 149: 86-101. DOI: 10.1016/j.visres.2018.06.003


Pikul, J.H., Li, S., Bai, H., Hanlon, R.T., Cohen, I. and Shepherd, R.F. (2017). Stretchable surfaces with programmable 3-D texture morphing for synthetic camouflaging skins. Science 358: 210-214. DOI: 10.1126/science.aan5627

Panetta, D., Buresch, K.C., & Hanlon, R.T. (2017). Dynamic masquerade with morphing 3D skin in cuttlefish. Biology Letters 13 (3): 20170070 (featured in Nature Vol 544 Research Highlights). DOI: 10.1098/rsbl.2017.0070

Allen, J.J., Akkaynak, D., Schnell, A.K. & Hanlon, R.T. (2017). Dramatic fighting by male cuttlefish for a female mate. American Naturalist 190 (1): 144-151. DOI: 10.1086/692009

Panetta, D., Solomon, M., Buresch, K.C. & Hanlon, R.T. (2017). Small-scale rearing of cuttlefish (Sepia officinalis) for research purposes. Marine & Freshwater Behaviour and Physiology 50(2): 115-124. DOI: 10.1080/10236244.2017.1343631


Schnell, A.K., Smith, K-L., Hanlon, R.T., Hall, K.C. & Harcourt, R.T. (2016). Cuttlefish perform multiple agonistic displays to communicate a hierarchy of threats. Behav Ecol & Sociobiology 70 (10): 1643-1655. DOI: 10.1007/s00265-016-2170-7

Schnell, A.K., Hanlon, R.T., Benkada, A., & Jozet-Alves, C. (2016). Lateralization of eye use in cuttlefish: opposite direction for anti-predatory and predatory behavior. Frontiers in Physiology 7: Article 620. DOI: 10.3389/fphys.2016.00620

Oshima, M., di Pauli von Treuheim, T., Carroll, J., Hanlon, R.T., Walters, E.T., & Crook, R.J. (2016). Peripheral injury alters schooling behavior in squid, Doryteuthis pealeii. Behav Processes, 128: 89-95. DOI: 10.1016/j.beproc.2016.04.008


Chiao, C.C., Chubb, C., & 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

Buresch, K.C., Ulmer, K.M., Akkaynak, D., Allen, J.J., Mäthger, L.M., Nakamura, M., & 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

Tyrie, E.K., Hanlon, R.T., Siemann, L.A., & 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

Schnell, A.K., Smith, C.L., Hanlon, R.T., & Harcourt, R.T. (2015). Female receptivity, mating history, and familiarity influence the mating behavior of cuttlefish. Behavioral Ecology and Sociobiology, 69(2): 283–292. DOI: 10.1007/s00265-014-1841-5

Kingston, A.C.N., Kuzirian, A.M., Hanlon, R.T., & Cronin, T.W. (2015). Visual phototransduction components in cephalopod chromatophores suggest dermal photoreception. Journal of Experimental Biology, 218(10): 1596–1602. DOI: 10.1242/jeb.117945

Kingston, A.C., Wardill, T.J., Hanlon, R.T., & Cronin, T.W. (2015). An Unexpected Diversity of Photoreceptor Classes in the Longfin Squid, Doryteuthis pealeii. PLoS One, 10(9): e0135381. DOI: 10.1575/1912/7383

Buresch, K.C., Ulmer, K.M., Cramer, C., McAnulty, S., Davison, W., Mäthger, L.M., & 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. DOI: 10.13140/RG.2.1.3473.0965

Allen, J.J., Akkaynak, D., Sugden, A.U., & 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

Kripke, E., Senft, S.L., Mozzherin, D., & Hanlon, R.T. (2015). Visualizing Biological Complexity in Cephalopod Skin: A Synergy of Art and Science Technologies. Leonardo, 48(5): 486–487. DOI: 10.1162/LEON_a_01124

Schnell, A.K., Smith, C.L., Hanlon, R.T., & Harcourt, R.T. (2015). Giant Australian cuttlefish use mutual assessment to resolve male-male contests. Animal Behaviour, 107: 31–40. DOI: 10.1016/j.anbehav.2015.05.026


Samson, J.E., Mooney, T.A., Gussekloo, S.W.S., & Hanlon, R.T. (2014). Graded behavioral responses and habituation to sound in the common cuttlefish Sepia officinalis. Journal of Experimental Biology, 217(24): 4347–4355. DOI: 10.1242/jeb.113365

Yu, C., Li, Y., Zhang, X., Huang, X., Malyarchuk, V., Wang, S., Shi, Y., Gao, L., Su, Y., Zhang, Y., Xu, H., Hanlon, R.T., and Rogers, J. (2014). Adaptive optoelectronic camouflage systems with designs inspired by cephalopod skins. Proceedings of the National Academy of Sciences, 111 (36): 12998-13003. DOI:10.1073/pnas.1410494111

Watson, A.C., Siemann, L.A., & 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

Allen, J.J., Bell, G.R.R., Kuzirian, A.M., Velankar, S.S., & Hanlon, R.T. (2014). Comparative Morphology of Changeable Skin Papillae in Octopus and Cuttlefish. Journal of Morphology, 275(4): 371–390. DOI: 10.1002/jmor.20221

Deravi, L.F., Magyar, A.P., Sheehy, S.P., Bell, G.R.R., Mäthger, L.M., Senft, S.L., Wardill, T.J., Lane, W.S., Kuzirian, A.M., Hanlon, R.T., Hu, E.L. and Parker, K.K. (2014). The structure-function relationship of a natural nanoscale photonic device in cuttlefish chromatophores. J Royal Society Interface, 11(93): 20130942. DOI: 10.1098/rsif.2013.0942

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

Gonzalez-Bellido, P.T.*, Wardill, T.J.*, Ulmer, K.M., Buresch, K.C., Hanlon, R.T. (2014). Expression of squid iridescence depends on environmental luminance and peripheral ganglion control. J. Experimental Biology 217: 850-858. DOI: 10.1242/jeb.091884

Crook, R.J., Dickson, K., Hanlon, R.T., & Walters, E.T. (2014). Nociceptive Sensitization Reduces Predation Risk. Current Biology, 24(10): 1121–1125. DOI: 10.1016/j.cub.2014.03.043

Bell, G.R.R., Mäthger, L.M., Gao, M., Senft, S.L., Kuzirian, A.M., Kattawar, G.W., & Hanlon, R.T. (2014). Diffuse white structural coloration from multilayer reflectors in a squid. Advanced Materials, 26(25): 4352–4356. DOI: 10.1002/adma.201400383


Allen, J.J., Bell, G.R.R., Kuzirian, A.M., Hanlon, R.T. (2013). Cuttlefish skin papilla morphology suggests a muscular hydrostat function for rapid changeability. J Morphology 274 (6): 645-656. DOI: 10.1002/jmor.20121

Allen, J.J., Bell, G.R.R., Kuzirian, A.M, Velankar, S.S. and Hanlon, R.T. (2013). Comparative morphology of changeable skin papillae in octopus and cuttlefish. J. Morphology 275(4): 371-390. DOI: 10.1002/jmor.20221

Akkaynak, D., Allen, J.J., Mäthger, L.M., Chiao, C.-C., Hanlon, R.T. (2013). Quantification of cuttlefish (Sepia officinalis) camouflage in the eyes of fish predators: A study of color and luminance using in situ spectrometry. J. Comp. Physiol A. 199 (3): 211-225. DOI:10.1007/s00359-012-0785-3

Hanlon, R.T., Chiao, C.-C., Mäthger, L.M., 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

Bell, G.R.R., Kuzirian, A.M., Senft, S.L., Mäthger, L.M., Wardill, T.J., Hanlon, R.T. (2013). Chromatophore radial muscle fibers anchor in flexible squid skin. Invertebrate Biology 132: 120-132. DOI: 10.1111/ivb.12016

Chiao, C.C., Ulmer, K.M., Siemann, L.A., Buresch, K.C., Chubb, C., Hanlon, R.T. (2013). How visual edge features influence cuttlefish camouflage patterning. Vision Research 83: 40-47. DOI: 10.1016/j.visres.2013.03.001

Crook, R.J., Hanlon, R.T. and Walters, E.T. (2013). Squid have nociceptors that display widespread long-term sensitization and spontaneous activity after bodily injury. J. Neuroscience 33 (24): 10021-10026. DOI: 10.1523/JNEUROSCI.0646-13.2013

Kreit, E., Mäthger, L.M., Hanlon, R.T., Dennis, P.B., Naik, R.R., Forsythe, E. and Heikenfeld, J. (2013). Biological vs. electronic adaptive coloration: how can one inform the other? Journal of the Royal Society Interface 10 (78): article 20120601 DOI:10.1098/rsif.2012.0601

Wardill, T.J., Gonzalez-Bellido, P.T., Crook, R.J. and Hanlon, R.T. (2013). Neural control of tuneable skin iridescence in squid. Proc. R. Soc. B. 279: 4243-4252. DOI: 10.1098/rspb.2012.1374

Mäthger, L.M., Hanlon, R.T., Håkansson, J., Nilsson, D.-E. (2013). The W-shaped pupil in cuttlefish (Sepia officinalis): functions for improving horizontal vision. Vision Research 83: 19–24. DOI: 10.1016/j.visres.2013.02.016

Mäthger, L.M., Senft, S.L., Gao, M., Karaveli, S., Bell, G.R.R., Zia, R., Kuzirian, A.M., Dennis, P.B., Crookes-Goodson, W.J., Naik, R.R., Kattawar, G.W., Hanlon, R.T. (2013). Bright white scattering from protein spheres in color changing, flexible cuttlefish skin. Advanced Functional Materials 23(32): 3980-3989. DOI: 10.1002/adfm.201203705

Nilsson, D.-E., Warrant, E.J., Johnsen, S., Hanlon, R.T., and Shashar, N. (2013). The giant eyes of giant squid are indeed unexpectedly large, but not if used for spotting sperm whales. BMC Evolutionary Biology 13: article 187. DOI: 10.1186/1471-2148-13-187

Shashar, N. and Hanlon, R.T. (2013). Spawning behavior dynamics at communal egg beds in the squid Doryteuthis (Loligo) pealeii (Mollusca: Cephalopoda). J. Experimental Marine Biology and Ecology 447:65-74. DOI: 10.1016/j.jembe.2013.02.011

Staudinger, M.D., Buresch, K.C., Mäthger, L.M., Fry, C., McAnulty, S., Ulmer, K.M., Hanlon, R.T. (2013). Defensive responses of cuttlefish to different teleost predators. Biological Bulletin 225 (3): 161-174. DOI: 10.1086/BBLv225n3p161

Ulmer, K.M., Buresch, K.C., Kossodo, M.M., Mäthger, L.M., Siemann, L.A., Hanlon, R.T. (2013). Visual background features in the vertical dimension have a strong influence on cuttlefish camouflage. Biological Bulletin 224(2): 110-118. DOI:10.1086/BBLv224n2p110



Wardill, T.J., Gonzalez-Bellido, P.T., Crook, R.J., Hanlon, R.T. (2012). Neural control of tuneable skin iridescence in squid. Proc. R. Soc. B. 279: 4243-52. DOI: 10.1098/rspb.2012.1374

Albertin, C.B., Bonnaud, L., Brown, T.C., Crookes-Goodsen, W.J., da Fonseca, R.R., Di Cristo, C., Dilkes, B.P., Edsinger-Gonzales, E., Freeman, R.M., Hanlon, R.T., & 16 others. (2012). Cephalopod genomics: a plan of strategies and organization. Standards in Genomic Sciences 7: 175-188. DOI: 10.4056/sigs.3136559

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

Gerlach, G., Buresch, K.C. and Hanlon, R.T. (2012). Population structure of the squid Doryteuthis (Loligo) pealeii on the eastern coast of the USA: comment on Shaw et al. (2010). Marine Ecology Progress Series 450: 281-283. DOI: 10.3354/meps09652

Mäthger, L.M., Bell, G.R.R., Kuzirian, A.M., Allen, J.J. and Hanlon, R.T. (2012). How does the blue-ringed octopus (Hapalochlaena lunulata) flash its blue rings? Journal of Experimental Biology. 215, 3752-3757. DOI: 10.1242/jeb.076869

Nilsson, D.-E., Warrant, E.J., Hanlon, R.T., Shashar, N., and Johnsen, S. (2012). A unique advantage for giant eyes in giant squid. Current Biology 22 (8): 683-688. DOI: 10.1016/j.cub.2012.02.031

Barbosa, A., Allen, J.J., Mäthger, L.M. and Hanlon, R.T. (2012). Cuttlefish use visual cues to determine arm postures for camouflage. Proc Roy Soc B 279:84-90. DOI: 10.1098/rspb.2011.0196


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

Chiao, C-C., Wickiser, J.K., Allen, J.J., Genter, B., & 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 108 (22): 9148-9153. DOI:10.1073/pnas.1019090108

Cummings, S.F., Boal, J.G., Buresch, K.C., Kuanpradit, C., Sobhon, P., Degnan, B.M., Nagle, G.T., & Hanlon, R.T. (2011). Extreme aggression in male squid induced by a β -MSP-like pheromone. Current Biology 21: 322-327. DOI: 10.1016/j.cub.2011.01.038

Shashar, N., Johnsen, S., Lerner, A., Sabbah, S., Chiao, C-C., Mäthger, L.M., & Hanlon, R.T. (2011). Underwater linear polarization – physical limitations to biological functions. Phil. Trans. Royal Society B 366: 649-654. DOI: 10.1098/rstb.2010.0190

Staudinger, M.D., Hanlon, R.T., & Juanes, F. (2011). Primary and secondary defenses 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

Zylinski, S., How, M.J., Osorio, D., Hanlon, R.T., & 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

Young, M.A., Kvitek, R.G., Iampietro, P.J., Garza, C.D., Malliet, R., & Hanlon, R.T. (2011). Seafloor mapping and landscape ecology analyses used to monitor variations in spawning site preference and benthic egg mop abundance for the California market squid (Doryteuthis opalescens). J Exp Marine Biology & Ecology 407: 226-233. DOI: 10.1016/j.jembe.2011.06.017

Crook, R.J., Lewis, T., Hanlon, R.T., & Walters, E.T. (2011). Peripheral injury induces long-term sensitization of defensive responses to visual and tactile stimuli in the squid, Loligo pealeii, Lesueur 1821. Journal of Experimental Biology 214: 3173-3185. DOI:10.1242/jeb.058131


Izumi, M., Sweeney, A.M., DeMartini, D., Weaver, J.C., Powers, M.L., Tao, A., Silvas, T.V., Kramer, R.M., Crookes-Goodson, W.J., Mäthger, L.M., Naik, R.R., Hanlon, R.T., & Morse, D. E. (2010). Changes in reflectin protein phosphorylation are associated with dynamic iridescence in squid. J. Royal Society Interface 7(44): 549-560. DOI: 10.1098/rsif.2009.0299

Chiao, C-C., Chubb, C., Buresch, K.C., Allen, J.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:10.1242/jeb.030247

Hanlon, R.T., Watson, A.C., & Barbosa, A. (2010). A “mimic octopus” in the Atlantic: flatfish mimicry and camouflage by Macrotritopus defilippi. Biological Bulletin 218 (1): 15-24. DOI: 10.1086/BBLv218n1p15 (Editor’s Choice Article and Cover Shot)

Allen, J.J., Mäthger, L.M., Barbosa, A., Buresch, K.C., Sogin, E., Schwartz, J., Chubb, C., & Hanlon, R.T. (2010). Cuttlefish dynamic camouflage: responses to substrate choice and integration of multiple visual cues. Proc. Roy. Soc. B 277(1684): 1031-1039. DOI:10.1098/rspb.2009.1694

Mooney, T.A., Lee, W-J., & Hanlon, R.T. (2010). Long-duration anesthetization of squid (Doryteuthis pealeii). Marine and Freshwater Behaviour and Physiology 43 (4): 297-303. DOI: 10.1080/10236244.2010.504334  PDF

Mäthger, L.M., Roberts, S.B., & Hanlon, R.T. (2010). Evidence for distributed light sensing in the skin of cuttlefish, Sepia officinalis. Biology Letters 6(5): 600-603. DOI: 10.1098/rsbl.2010.0223

Mooney, T.A., Hanlon, R.T., Christensen-Dalsgaard, J., Madsen, P.T., Ketten, D.R., & Nachtigall, P.E. (2010). Sound detection by the longfin squid (Loligo pealeii) studied with auditory evoked potentials: sensitivity to low-frequency particle motion and not pressure. Journal of Experimental Biology 213:3748-3759. Biology 213: 3953-3960. DOI: 10.1242/jeb.048348


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(1516): 429-437. DOI: 10.1098/rstb.2008.0270

Mäthger, L.M., Denton, E.J., Marshall, N.J., & Hanlon, R.T. (2009). Mechanisms and behavioral functions of structural coloration in cephalopods. Journal of the Royal Society Interface 6(Suppl 2): S149-S164. DOI: 10.1098/rsif.2008.0366.focus

Williams, S.B., Pizarro, O., How, M., Mercer, D., Powell, G., Marshall, N.J., & Hanlon, R.T. (2009). Surveying nocturnal cuttlefish camouflage behaviour using an AUV. IEEE International Conference on Robotics and Automation: 214-219. DOI:10.1109/ROBOT.2009.5152868

Mäthger, L.M., Shashar, N., & Hanlon, R.T. (2009). Do cephalopods communicate using polarized light reflections from their skin? Journal of Experimental Biology 212(Pt 14): 2133-2140. (Featured Commentary) DOI: 10.1242/jeb.020800

Allen, J.J., 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(6): 547-555. DOI: 10.1007/s00359-009-0430-y

Chiao, C-C., Chubb, C., Buresch, K.C., Siemann, L.A., & Hanlon, R.T. (2009). The scaling effects of substrate texture on camouflage patterning in cuttlefish. Vision Research 49 (13): 1647-1656. DOI: 10.1016/j.visres.2009.04.002

Buresch, K.C., Maxwell, M.R., Cox, M.R., & Hanlon, R.T. (2009). Temporal dynamics of mating and paternity in the squid Loligo pealeii. Marine Ecology Progress Series 387: 197-203. DOI: 10.3354/meps08052 

Crook, R.J., Hanlon, R.T., & Basil, J.A. (2009). Memory of visual and topographical features suggests spatial learning in nautilus (Nautilus pompilius L.). J. Comparative Psychology 123 (3): 264-274. DOI: 10.1037/a0015921

Miserez, A., Weaver, J.C., Pedersen, P.B., Schneeberk, T., Hanlon, R.T., Kisailus, D., & Birkedal, H. (2009). Microstructural and biochemical characterization of the nano-porous sucker rings from Dosidicus gigas. Advanced Materials 21(4): 1-6. DOI:10.1002/adma.200801197


Mäthger, L.M., Chiao, C-C., Barbosa, A., & Hanlon, R.T. (2008). Color matching on natural substrates in cuttlefish, Sepia officinalis. Journal of Comparative Physiology A, 194 (6): 577-585. DOI: 10.1007/s00359-008-0332-4

Sutherland, R.L., Mäthger, L.M., Hanlon, R.T., Urbas, A.M., & Stone, M.O. (2008). Cephalopod coloration model. I. Squid chromatophores and iridophores. Journal of the Optical Society of America A, 25 (3): 588-599. DOI:10.1364/JOSAA.25.000588

Sutherland, R.L., Mäthger, L.M., Hanlon, R.T., Urbas, A.M., & Stone, M.O. (2008). Cephalopod coloration model. II. Multiple layer skin effects. Journal of the Optical Society of America, 25 (8): 2044-2054. DOI: 10.1364/JOSAA.25.002044

Barbosa, A., Mäthger, L.M., Buresch, K.C., Kelly, J., Chubb, C., Chiao, C-C., & Hanlon, R.T. (2008). Cuttlefish camouflage: The effects of substrate contrast and size in evoking uniform, mottle or disruptive body patterns. Vision Research, 48(10): 1242-1253.DOI: 10.1016/j.visres.2008.02.011

Barbosa, A., Litman, L., and Hanlon, R.T. (2008). Changeable cuttlefish camouflage is influenced by horizontal and vertical aspects of the visual background. Journal of Comparative Physiology A, 194(4): 405-413. DOI: 10.1007/s00359-007-0311-1

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(1): 23-38. DOI: 10.1111/j.1095-8312.2007.00948.x

Hanlon, R.T. & Forsythe, J.W. (2008). Sexual cannibalism by Octopus cyanea on a Pacific coral reef. Marine and Freshwater Behaviour and Physiology. 41(1): 19-28. DOI:10.1080/10236240701661123       PDF


Wilson, M., Hanlon, R.T., Tyack, P.L., & Madsen, P.T. (2007). Intense ultrasonic clicks from echolocating toothed whales do not elicit anti-predator responses or debilitate the squid Loligo pealeii. Biology Letters 3(3): 225-227. DOI: 10.1098/rsbl.2007.0005

Madsen, P.T., Wilson, M., Johnson, M., Hanlon, R.T., Bocconcelli, A., Aguilar de Soto, N., & Tyack, P.L. (2007). Clicking for calamari: toothed whales can echolocate squid Loligo pealeii. Aquatic Biology 1: 141-150. (Feature Article) DOI: 10.3354/ab00014 

Mäthger, L.M., Chiao, C-C., Barbosa, A., Buresch, K.C., Kaye, S., & Hanlon, R.T. (2007). Disruptive coloration elicited on controlled natural substrates in cuttlefish, Sepia officinalis. J. Exp. Biol. 210(Pt 15): 2657-2666. DOI: 10.1242/jeb.004382

Chiou, T.H., Hanlon, R.T., Mäthger, L.M., & Cronin, T.W. (2007). Spectral and spatial properties of polarized light reflections from the arms of squid (Loligo pealeii) and cuttlefish (Sepia officinalis L.). Journal of Experimental Biology. 210(Pt 20): 3624-3635. DOI:10.1242/jeb.006932

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(1): 179-186. DOI: 10.1007/s00441-007-0384-8

Chiao, C-C., Chubb, C., & Hanlon, R.T. (2007). Interactive effects of size, contrast, intensity and configuration of background objects in evoking disruptive camouflage in cuttlefish. Vision Research 47(16):2223-2235. DOI: 10.1016/j.visres.2007.05.001

Hanlon, R.T. (2007). Cephalopod dynamic camouflage. Current Biology 17 (11): 400-404. DOI: 10.1016/j.cub.2007.03.034

Hanlon, R.T., Naud, M.-J., Forsythe, J.W., Hall, K.C., Watson, A.C., & McKechnie, J. (2007). Adaptable night camouflage by cuttlefish. American Naturalist 169 (4): 543-551. DOI: 10.1086/512106

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(Pt 7): 1139-1147. DOI: 10.1242/jeb.02741


Buresch, K.C., Gerlach, G., & Hanlon, R.T. (2006). Multiple genetic stocks of longfin squid Loligo pealeii in the NW Atlantic: stocks segregate inshore in summer, but aggregate offshore in winter. Mar Ecol Prog Ser, 310: 263–270. DOI:10.3354/meps310263

Foote, K.G., Hanlon, R.T., Lampietro, P.J., & Kvitek, R.G. (2006). Acoustic detection and quantification of benthic egg beds of the squid Loligo opalescens in Monterey Bay, California. Journal of the Acoustical Society of America, 119 (2) : 844-856. DOI:10.1121/1.2149840

Mäthger, L.M., Barbosa, A., Miner, S., & Hanlon, R.T. (2006). Color blindness and contrast perception in cuttlefish (Sepia officinalis) determined by a visual sensorimotor assay. Vision Research 46(11): 1746-1753. DOI: 10.1016/j.visres.2005.09.035

Mäthger, L.M., & Hanlon, R.T. (2006). Anatomical basis for camouflaged polarized light communication in squid. Biology Letters 2(4): 494-496. DOI: 10.1098/rsbl.2006.0542


Hanlon, R.T., Naud, M.-J., Shaw, P.W., & Havenhand, J.N. (2005). Behavioural ecology: transient sexual mimicry leads to fertilisation. Nature 433(7023): 212. DOI: 10.1038/433212a

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 (1): 7-11. DOI:10.2307/3593095

Naud, M.J., Shaw, P.W., Hanlon, R.T., & Havenhand, J.N. (2005). Evidence for biased use of sperm sources in wild female giant cuttlefish (Sepia apama). Proceedings of the Royal Society B, 272 (1567) 1047-1051. DOI: 10.1098/rspb.2004.3031

Mensinger, A.F., Hanlon, R.T., Walsh, P.J. (2005). Blood biochemistry of the oyster toadfish. Journal of Aquatic Animal Health 17(2): 170-176. DOI: 10.1577/H04-021.1

Saidel, W.M., Shashar, N., Schmolesky, M.T., Hanlon, R.T. (2005). Discriminative responses of squid (Loligo pealeii) photoreceptors to polarized light. Comparative Biochemistry & Physiology A: 142(3): 340-346. DOI: 10.1016/j.cbpa.2005.08.003

Walker, I.D., Dawson, D.M., Flash, T., Grasso, F.W., Hanlon, R.T., Hochner, B., Kier, W.M., Pagano, C.C., Rahn, C.D., & Zhang, Q.M. (2005). Continuum Robot Arms Inspired by Cephalopods. Proceedings of SPIE. 5804: 303-314. [37]. DOI: 10.1117/12.606201   PDF


Naud, M.-J., Hanlon, R.T., Hall, K.C., Shaw, P.W., & Havenhand, J.N. (2004). Behavioral and genetic assessment of reproductive success in a spawning aggregation of the Australian giant cuttlefish, Sepia apama. Animal Behaviour 67(6): 1043-1050. DOI:10.1016/j.anbehav.2003.10.005

Hanlon, R.T., Kangas, N., & Forsythe, J.W. (2004). Egg capsule deposition and how behavioral interactions influence spawning rate in the squid Loligo opalescens in Monterey Bay, California. Marine Biology 145(5):923-930. DOI: 10.1007/s00227-004-1383-x

Forsythe, J.W., Kangas, N., & Hanlon, R.T. (2004). Does the California Market Squid, Loligo opalescens, spawn naturally during the day or at night? A note on the successful use of ROVs to obtain basic fisheries biology data. Fish. Bull. 102(2): 389-392. PDF

Buresch, K.C., Boal, J.G., Nagle, G.T., Knowles, J., Nobuhara, R., Sweeney, K., & Hanlon, R.T. (2004). Experimental evidence that ovary and oviducal gland extracts influence male agonistic behavior in squids. Bio. Bull. 206(1): 1-3. DOI: 10.2307/1543192

Boal, J.G., Shashar, N., Grable, M.M., Vaughan, K.H., Loew, E.R., & Hanlon, R.T. (2004). Behavioral evidence for intraspecific signaling with achromatic and polarized light by cuttlefish (Mollusca: Cephalopoda). Behaviour 141(7): 837-861. DOI:10.1163/1568539042265662     PDF


Hanlon, R.T., & Shashar, N. (2003). Aspects of the sensory ecology of cephalopods. In: Collin, S.P., Marshall, N.J. (eds.) Sensory Processing in Aquatic Environments. Springer-Verlag, New York, NY, pp. 266-282. DOI: 10.1007/978-0-387-22628-6_14

King, A.J., Adamo, S.A., & Hanlon, R.T. (2003). Squid egg mops provide sensory cues for increased agonistic behavior between male squid. Anim. Behav. 66(1): 49-58. DOI: 10.1006/anbe.2003.2197

Mensinger, A.F., Price, N.N., Richmond, H.E., Forsythe, J.W., & Hanlon, R.T. (2003). Mariculture of the oyster toadfish: juvenile growth and survival. North American Journal of Aquaculture 65(4):289-299. DOI: 10.1577/C02-056

Karson, M.A., Boal, J.G, & Hanlon, R.T. (2003). Experimental evidence for spatial learning in cuttlefish (Sepia officinalis). J. Comp. Psychol. Vol. 117(2): 149-155. DOI:10.1037/0735-7036.117.2.149  PDF

Buresch, K.C., Boal, J.G., Knowles, J., DeBose, J., Nichols, A., Erwin, A., Painter, S.D., Nagle, G.T., & Hanlon, R.T. (2003). Contact chemosensory cues in egg bundles elicit male-male agonistic conflicts in the squid Loligo pealeii. J. Chem. Ecol. 29(3):547-560. DOI:10.1023/A:1022846603591     PDF

Anderson, J.C., Baddeley, R.J., Osorio, D., Shashar, N., Tyler, C.W., Ramachandran, V.S., Crook, A.C., & Hanlon, R.T. (2003). Modular organization of adaptive colouration in flounder and cuttlefish revealed by independent component analysis. Comput. Neural Syst. 14(2): 321-333. DOI: 10.1088/0954-898X_14_2_308       PDF


Shashar, N., Milbury, C.A., & Hanlon, R.T. (2002). Polarization vision in cephalopods: neuroanatomical and behavioral features that illustrate aspects of form and function. Mar. Freshwat. Behav. Physiol. 35(1-2):57- 68. DOI: 10.1080/10236240290025617

Basil, J.A., Lazenby, G.B., Nakanuku, L., & Hanlon, R.T. (2002). Female Nautilus are attracted to male conspecific odor. Bull. Mar. Sci. 70(1):217-225. PDF

Hall, K.C., & Hanlon, R.T. (2002). Principal features of the mating system of a large spawning aggregation of the giant Australian cuttlefish Sepia apama(Mollusca: Cephalopoda). Mar. Biol. 140(3):533-545. DOI: 10.1007/s00227-001-0718-0 

Hanlon, R.T., Smale, M.J., & Sauer, W.H.H. (2002). The mating system of the squid Loligo vulgaris reynaudii (Cephalopoda, Mollusca) off South Africa: Fighting, guarding, sneaking, mating and egg laying behavior. Bull. Mar. Sci. 71(1): 331-345. PDF

O’Dor, R.K., Adamo, S.A., Aitken, J.P., Andrade, Y., Finn, J., Hanlon, R.T., & Jackson, G.D. (2002). Currents as environmental constraints on the behavior, energetics and distribution of squid and cuttlefish. Bull. Mar. Sci., 71(2): 601-617. PDF

Grable, M.M., Shashar, N., Gilles, N.L., Chiao, C-C., & Hanlon, R.T. (2002). Cuttlefish body patterns as a behavioral assay to determine polarization perception. Biol. Bull., 203(2): 232-234. DOI: 10.2307/1543414


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(12): 2119-2125. LINK

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(2):269-270. DOI: 10.2307/1543359

Buresch, K.M., Hanlon, R.T., Maxwell, M.R., & Ring, S. (2001). Microsatellite DNA markers indicate a high frequency of multiple paternity within individual field-collected egg capsules of the squid Loligo pealeii. Mar. Ecol. Prog. Ser. 210: 161-165. DOI:10.3354/meps210161

Hatfield, E.M.C., Hanlon, R.T., Forsythe, J.W., & Grist, E.P.M. (2001). Laboratory testing of a growth hypothesis for juvenile squid Loligo pealeii (Cephalopoda: Loliginidae). Can. J. Fish. Aquat. Sci. 58(5):845-857. DOI:10.1139/f01-030

Mensinger, A.F., Stephenson, K.A., Pollema, S.L., Richmond, H.E., Price, N., & Hanlon, R.T. (2001). Mariculture of the toadfish, Opsanus tau. Biol. Bull. 201(2):282-283. DOI: 10.2307/1543367

Shashar, N., Borst, D.T., Ament, S.A., Saidel, W.M., Smolowitz, R.M., & Hanlon, R.T. (2001). Polarization reflecting iridophores in the arms of the squid Loligo pealeii. Biol. Bull. 201(2):267-268. DOI: 10.2307/1543358


Basil, J.A., Hanlon, R.T., Sheikh, S.I., & Atema, J. (2000). Three-dimensional odor tracking by Nautilus pompilius. J. Exp. Biol. 203(9): 1409-1414. LINK

Boal, J.G., Dunham, A.W., Williams, K.T., & Hanlon, R.T. (2000). Experimental evidence for spatial learning in octopuses (Octopus bimaculoides). J. Comp. Psychol. 114(3): 246-252. DOI:10.1037/0735-7036.114.3.246

Boal, J.G., Wittenberg, K.M., & Hanlon, R.T. (2000). Observational learning does not explain improvement in predation tactics by cuttlefish (Mollusca: Cephalopoda). Behav. Processes 52(2-3):141-153. DOI:10.1016/S0376-6357(00)00137-6

Maxwell, M.R., Buresch, K.M., & Hanlon, R.T. (2000). Pattern of inheritance of microsatellite loci in the squid Loligo pealeii (Mollusca: Cephalopoda). Mar. Biotechnol. 2(6): 517-521. DOI: 10.1007/s101260000036

Maxwell, M.R., & Hanlon, R.T. (2000). Female reproductive output in the squid Loligo pealeii: multiple egg clutches and implications for a spawning strategy. Mar. Ecol. Prog. Ser. 199: 159-170. DOI: 10.3354/meps199159

Shashar, N., Hagan, R., Boal, J.G., & Hanlon, R.T. (2000). Cuttlefish use polarization sensitivity in predation on silvery fish. Vision Res. 40(1): 71-75. DOI: 10.1016/S0042-6989(99)00158-3


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): 1-22. DOI:   10.1111/j.1095-8312.1999.tb01914.x

Boal, J.G., Hylton, R.A., Gonzalez, S.A., & Hanlon R.T. (1999). Effects of crowding on the social behavior of cuttlefish (Sepia officinalis). Contemp. Topics Lab. Anim. Sci. 38(1): 49-55. PDF

Gabr, H.R., Hanlon, R.T., El-Etreby, S.G., & Hanafy, M.H. (1999). Reproductive versus somatic tissue growth during the life cycle of the cuttlefish Sepia pharaonis Ehrenberg, 1831. Fish. Bull. 97(4): 802-811. PDF

Gabr, H.R., Hanlon, R.T., Hanafy, M.H., & El-Etreby, S.G. (1999). Reproductive versus somatic tissue allocation in the cuttlefish Sepia dollfusi Adam (1941). Bull. Mar. Sci. 65(1): 159-173. PDF

Hanley, J.S., Shashar, N., Smolowitz, R., Mebane, W.N., & Hanlon, R.T. (1999). Soft- sided tanks improve long-term health of cultured cuttlefish. Biol. Bull. 197(2): 237-238. DOI: 10.2307/1542623

Hanlon, R.T., Ament, S.A., & Gabr, H.R. (1999). Behavioral aspects of sperm competition in cuttlefish, Sepia officinalis (Sepioidea: Cephalopoda). Mar. Biol. 134(4): 719-728. DOI: 10.1007/s002270050588

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. DOI:10.2307/1542996

King, A.J., Adamo, S.A., & Hanlon, R.T. (1999). Contact with squid egg capsules increases agonistic behavior in male squid (Loligo pealei). Biol. Bull. 197(2): 256. DOI: 10.2307/1542636

Tang, K.Q., Price, N.N., O’Neill, M.D., Mensinger, A.F., & Hanlon, R.T. (1999). Temperature effects on first-year growth of cultured oyster toadfish, Opsanus tau. Biol. Bull. 197(2): 247-248. DOI: 10.2307/1542630


Shashar, N., Hanlon, R.T., & Petz, A.deM. (1998). Polarization vision helps detect transparent prey. Nature 393(6682): 222-223. DOI: 10.1038/30380

Hanlon, R.T. (1998). Mating systems and sexual selection in the squid Loligo: How might commercial fishing on spawning squids affect them? Calcofi Rep. 39: 92-100. PDF

Gabr, H.R., Hanlon, R.T., Hanafy, M.H., & El-Etreby, S.G. (1998). Maturation, fecundity and seasonality of reproduction of two commercially valuable cuttlefish, Sepia pharaonis and S. dollfusi, in the Suez Canal. Fish. Res. 36(2-3): 99-115. DOI:10.1016/S0165-7836(98)00107-6

Hanley, J.S., Shashar, N., Smolowitz, R., Bullis, R.A., Mebane, W.N., Gabr, H.R., & Hanlon, R.T. (1998). Modified laboratory culture techniques for the European cuttlefish Sepia officinalis. Biol. Bull. 195(2): 223-225. DOI: 10.2307/1542850

Maxwell, M.R., Macy, W.K., Odate, S., & Hanlon, R.T. (1998). Evidence for multiple spawning by squids (Loligo pealei) in captivity. Biol. Bull. 195(2): 225- 226. DOI: 10.2307/1542851

O’Neill, M.D., Wesp, H.M., Mensinger, A.F., & Hanlon, R.T. (1998). Initial baseline blood chemistry of the oyster toadfish, Opsanus tau. Biol. Bull. 195(2): 228-229. DOI: 10.2307/1542853

Shashar, N., Harosi, F.I., Banaszak, A.T., & Hanlon, R.T. (1998). UV radiation blocking compounds in the eye of the cuttlefish Sepia officinalis. Biol. Bull. 195(2): 187-188. DOI: 10.2307/1542830


Sauer, W.H.H., Roberts, M.J., Lipinski, M.R., Smale, M.J., Hanlon, R.T., Webber, D.M., & O’Dor, R.K. (1997). Choreography of the squid’s “nuptial dance.” Biol. Bull. 192(2): 203-207. DOI: 10.2307/1542714

Ament, S.A., Bullis, R.A., Hanlon, R.T., & Mensinger, A.F. (1997). Righting response and escape response in Opsanus tau are temperature dependent. Biol. Bull. 193(2): 265-266. DOI: 10.1086/BBLv193n2p265

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(4): 1243-1246. DOI: 10.1017/S0025315400038789

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. DOI: 10.1111/j.1439-0310.1997.tb00010.x

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(1-2): 15-31. DOI: 10.1016/S0022-0981(96)00057-3

Hanlon, R.T., Claes, M.F., Ashcraft, S.E., & Dunlap, P.V. (1997). Laboratory culture of the sepiolid squid Euprymna scolopes: A model system for bacteria- animal symbiosis. Biol. Bull. 192(3): 364-374. DOI: 10.2307/1542746

Hanlon, R.T., Maxwell, M.R., & Shashar, N. (1997). Behavioral dynamics that would lead to multiple paternity within egg capsules of the squid Loligo pealei. Biol. Bull. 193(2): 212-214. DOI: 10.1086/BBLv193n2p212

Jackson, G.D., Forsythe, J.W., Hixon, R.F., & Hanlon, R.T. (1997). Age, growth, and maturation of Lolliguncula brevis (Cephalopoda: Loliginidae) in the northwestern Gulf of Mexico with a comparison of length-frequency versus statolith age analysis. Can. J. Fish. Aquat. Sci. 54(12): 2907-2919. DOI: 10.1139/f97-192

Oestmann, D.J., Scimeca, J.M., Forsythe, J.W., Hanlon, R.T., & Lee, P.G. (1997). Special considerations for keeping cephalopods in laboratory facilities. Contemp. Top. Lab. Anim. Sci. 36(2): 89-93. PDF

Rosenberg, G., Tillier, S., Tillier, A., Kuncio, G.S., Hanlon, R.T., Masselot, M., & Williams, C.J. (1997). Proceedings of an international meeting: Ribosomal RNA phylogeny of selected major clades in the Mollusca. J. Molluscan Stud. 63(3): 301-309. DOI:10.1093/mollus/63.3.301

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. DOI: 10.1086/BBLv193n2p207


Adamo, S.A., & Hanlon, R.T. (1996). Do cuttlefish (Cephalopoda) signal their intentions to conspecifics during agonistic encounters? Anim. Behav. 52(1): 73-81. DOI: 10.1006/anbe.1996.0153

Hanlon, R.T. (1996). Evolutionary games that squids play: Fighting, courting, sneaking, and mating behaviors used for sexual selection in Loligo pealei. Biol. Bull. 191(2): 309-310. DOI: 10.1086/BBLv191n2p309

Lee, K.W., Baker, R., Galione, A., Gilland, E.H., Hanlon, R.T., & Miller, A.L. (1996). Ionophore-induced calcium waves activate unfertilized zebrafish (Danio rerio) eggs. Biol. Bull. 191(2): 265-267. DOI: 10.1086/BBLv191n2p265


Smale, M.J., Sauer, W.H.H., & Hanlon, R.T. (1995). Attempted ambush predation on spawning squids Loligo vulgaris reynaudii by benthic pyjama sharks, Poroderma africanum, off South Africa. J. Mar. Biol. Assoc. U.K. 75(3): 739-742. DOI:10.1017/S002531540003914X


Forsythe, J.W., DeRusha, R.H., & Hanlon, R.T. (1994). Growth, reproduction and life span of Sepia officinalis (Cephalopoda: Mollusca) cultured through seven consecutive generations. J. Zool. Lond. 233(2): 175-192. DOI:10.1111/j.1469-7998.1994.tb08582.x

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. DOI: 10.2307/1542293

Lee, P.G., Turk, P.E., Yang, W.T., & Hanlon, R.T. (1994). Biological characteristics and biomedical applications of the squid Sepioteuthis lessoniana cultured through multiple generations. Biol. Bull. 186(3): 328-341. DOI: 10.2307/1542279


Forsythe, J.W., Hanlon, R.T., Bullis, R.A., & Noga, E.J. (1991). Octopus bimaculoides (Pickford & McConnaughey, 1949): A marine invertebrate host for ectoparasitic protozoans. J. Fish. Dis. 14(4): 431-442. DOI: 10.1111/j.1365-2761.1991.tb00595.x

Hanlon, R.T. (1991). Integrative Neurobiology and Behavior of Mollusks Symposium: Introduction, perspectives, and round-table discussion. Biol. Bull. 180(2): 197-199. DOI: 10.2307/1542388

Hanlon, R.T., Turk, P.E., & Lee, P.G. (1991). Squid and cuttlefish mariculture: An updated perspective. J. Cephalopod Biol. 2(1): 31-40.

Lee, P.G., Forsythe, J.W., DiMarco, F.P., DeRusha, R.H., & Hanlon, R.T. (1991). Initial palatability and growth trials on pelleted diets for cephalopods. Bull. Mar. Sci. 49(1-2): 362-372. LINK PDF


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(1): 3-14. DOI: 10.1007/BF00571424     PDF

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(1): 15-24. DOI: 10.1007/BF00571425  PDF

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. DOI: 10.1016/0006-8993(90)90094-r   PDF

Porteiro, F.M., Martins, H.R., Hanlon, R.T. (1990). Some observations on the behaviour of adult squids, Loligo forbesi, in captivity. J. Mar. Biol. Assoc. U.K. 70(2): 459-472. DOI: 10.1017/S0025315400035530


Long, T.M., Hanlon, R.T., ter Maat, A., & Pinsker, H.M. (1989). Non-associative learning in the squid Lolliguncula brevis (Mollusca, Cephalopoda). Mar. Behav. Physiol. 16(1) :1-9. DOI:10.1080/10236248909378736

Adams, P.M., Hanlon, R.T., & Forsythe, J.W. (1989). Toxic exposure to ethylene dibromide and mercuric chloride: Effects on laboratory-reared octopuses. Neurotoxicol. Teratol. 10(6): 519-523. DOI: 10.1016/0892-0362(88)90087-6

DeRusha, R.H., Forsythe, J.W., DiMarco, F.P., & Hanlon, R.T. (1989). Alternative diets for maintaining and rearing cephalopods in captivity. Lab. Anim. Sci. 39(4): 306-312.

Forsythe, J.W., & Hanlon, R.T. (1989). Growth of the Eastern Atlantic squid, Loligo forbesi Steenstrup (Mollusca: Cephalopoda). Aquacul. Fish. Manag. 20(1): 1- 14. DOI: 10.1111/j.1365-2109.1989.tb00437.x 

Hanlon, R.T., Bidwell, J.P., & Tait, R. (1989). Strontium is required for statolith development and thus normal swimming behaviour of hatchling cephalopods. J. Exp. Biol. 141: 187-195. LINK

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., Yang, W.T., Turk, P.E., Lee, P.G., & Hixon, R.F. (1989). Laboratory culture and estimated life span of the Eastern Atlantic squid Loligo forbesi Steenstrup, 1856 (Mollusca: Cephalopoda). Aquacul. Fish. Manag. 20(1): 15- 34. DOI:10.1111/j.1365-2109.1989.tb00438.x

Yang, W.T., Hanlon, R.T., Lee, P.G., & Turk, P.E. (1989). Design and function of closed seawater systems for culturing loliginid squids. Aquacult. Eng. 8(1): 47-65. DOI:10.1016/0144-8609(89)90020-4


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(1200): 437-487. DOI:10.1098/rstb.1988.0087

Forsythe, J.W., & Hanlon, R.T. (1988). Behavior, body patterning and reproductive biology of Octopus bimaculoides from California. Malacologia 29(1): 41- 55.

Forsythe, J.W., & Hanlon, R.T. (1988). Effect of temperature on laboratory growth, reproduction and life span of Octopus bimaculoides. Mar. Biol. 98(3): 369-379. DOI: 10.1007/BF00391113

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

Segawa, S., & Hanlon, R.T. (1988). Oxygen consumption and ammonia excretion rates in Octopus maya, Loligo forbesi and Lolliguncula brevis (Mollusca: Cephalopoda). Mar. Behav. Physiol. 13(4): 389-400. DOI: 10.1080/10236248809378687

Segawa, S., Yang, W.T., Marthy, H.-J., & Hanlon, R.T. (1988). Illustrated embryonic stages of the eastern Atlantic squid Loligo forbesi. Veliger 30(3): 230-243. PDF

Wells, M.J., Hanlon, R.T., Lee, P.G., & DiMarco, F.P. (1988). Respiratory and cardiac performance in Lolliguncula brevis (Cephalopoda, Myopsida): The effects of activity, temperature and hypoxia. J. Exp. Biol. 138: 17-36. PDF   LINK


Hanlon, R.T., & Budelmann, B.-U. (1987). Why cephalopods are probably not “deaf.” Am. Nat. 129(2): 312-317. DOI:10.1086/284637

DeRusha, R.H., Forsythe, J.W., & Hanlon, R.T. (1987). Laboratory growth, reproduction and life span of the Pacific pygmy octopus, Octopus digueti. Pac. Sci. 41(1-4): 104-121. PDF LINK

Hanlon, R.T., Turk, P.E., Lee, P.G., & Yang, W.T. (1987). Laboratory rearing of the squid Loligo pealei to the juvenile stage: Growth comparisons with fishery data. Fish. Bull. 85(1): 163-167.


Cooper, K.M., & Hanlon, R.T. (1986). Correlation of iridescence with changes in iridophore platelet ultrastructure in the squid Lolliguncula brevis. J. Exp. Biol. 121(1): 451-455. LINK

Dubas, F., Hanlon, R.T., Ferguson, G.P., & Pinsker, H.M. (1986). Localization and stimulation of chromatophore motoneurones in the brain of the squid, Lolliguncula brevis. J. Exp. Biol. 121(1): 1-25. LINK

Dubas, F., Leonard, R.B., & Hanlon, R.T. (1986). Chromatophore motoneurons in the brain of the squid, Lolliguncula brevis: An HRP study. Brain Res. 374(1): 21-29. DOI: 10.1016/0006-8993(86)90390-2

Ford, L.A., Alexander, S.K., Cooper, K.M., & Hanlon, R.T. (1986). Bacterial populations of normal and ulcerated mantle tissue of the squid, Lolliguncula brevis. J. Invertebr. Pathol. 48(1): 13-26. DOI: 10.1016/0022-2011(86)90138-2

Hanlon, R.T., & Hixon, R.F. (1986). Behavioral associations of coral reef fishes with the sea anemone Condylactis gigantea in the Dry Tortugas, Florida. Bull. Mar. Sci. 39(1): 130-134. LINK

Turk, P.E., Hanlon, R.T., Bradford, L.A., & Yang, W.T. (1986). Aspects of feeding, growth and survival of the European squid, Loligo vulgaris Lamarck, 1799, reared through the early growth stages. Vie Milieu 36(1): 9-13.

Yang, W.T., Hixon, R.F., Turk, P.E., Krejci, M.E., Hulet, W.H., & Hanlon, R.T. (1986). Growth, behavior, and sexual maturation of the market squid, Loligo opalescens, cultured through the life cycle. Fish. Bull. 84(4): 771-798. LINK


Florey, E., Dubas, F., & Hanlon, R.T. (1985). Evidence for L-glutamate as a transmitter substance of motoneurons innervating squid chromatophore muscles. Comp. Biochem. Physiol. C. 82(2): 259-268. DOI: 10.1016/0742-8413(85)90160-4

Forsythe, J.W., & Hanlon, R.T. (1985). Aspects of egg development, post-hatching behavior, growth and reproductive biology of Octopus burryi Voss, 1950 (Mollusca: Cephalopoda). Vie Milieu 35(3-4): 273-282. LINK

Hanlon, R.T. (1985). Cephalopods of the northwestern Gulf of Mexico. Texas Conchol. 21(3): 90-95.

Hanlon, R.T., & Forsythe, J.W. (1985). Advances in the laboratory culture of octopuses for biomedical research. Lab. Anim. Sci. 35(1): 33-40. LINK

Hanlon, R.T., Forsythe, J.W., & Boletzky, S.V. (1985). Field and laboratory behavior of “macrotritopus larvae” reared to Octopus defilippi Verany, 1851 (Mollusca: Cephalopoda). Vie Milieu 35(3-4): 237-242.


Colmers, W.F., Hixon, R.F., Hanlon, R.T., Forsythe, J.W., Ackerson, M.V., Wiederhold, M.L., & Hulet, W.H. (1984). “Spinner” cephalopods: Defects of statocyst suprastructures in an invertebrate analogue of the vestibular apparatus. Cell Tissue Res. 236(3): 505-515. DOI: 10.1007/BF00217217

Hanlon, R.T., Forsythe, J.W., Cooper, K.M., DiNuzzo, A.R., Folse, D.S., & Kelly, M.T. (1984). Fatal penetrating skin ulcers in laboratory-reared octopuses. J. Invertebr. Pathol. 44(1): 67-83. DOI:10.1016/0022-2011(84)90047-8

Hanlon, R.T., Forsythe, J.W., Hixon, R.F., & Yang, W.T. (1984). Recent advances in octopus mariculture. Aquabiology 6(6): 420-426. [In Japanese, with English abstract]


Hanlon, R.T., Hixon, R.F., & Hulet, W.H. (1983). Survival, growth, and behavior of the loliginid squids Loligo plei, Loligo pealei, and Lolliguncula brevis (Mollusca: Cephalopoda) in closed sea water systems. Biol. Bull. 165(3): 637-685. DOI: 10.2307/1541470

Boletzky, S.V., & Hanlon, R.T. (1983). A review of the laboratory maintenance, rearing and culture of cephalopod molluscs. Mem. Natl. Mus. Victoria 44: 147- 187. DOI: 10.24199/j.mmv.1983.44.11

Hanlon, R.T., Hixon, R.F., & Smith, D.G. (1983). Behavioral associations of seven West Indian reef fishes with sea anemones at Bonaire, Netherlands Antilles. Bull. Mar. Sci. 33(4): 928-934. LINK

Marthy, H.-J., & Hanlon, R.T. (1983). Effects of laser microirradiation in chromatophore organs of embryonic and juvenile cephalopods. Mikroscopie (Wien) 40(1-2): 35-40.

Yang, W.T., Hanlon, R.T., Krejci, M.E., Hixon, R.F., & Hulet, W.H. (1983). Laboratory rearing of Loligo opalescens, the market squid of California. Aquaculture 31(1): 77-88. DOI:10.1016/0044-8486(83)90259-4

Yang, W.T., Hixon, R.F., Turk, P.E., Krejci, M.E., Hanlon, R.T., & Hulet, W.H. (1983). Culture of California market squid from hatching - completion of the rearing cycle to second generation hatchlings. Aquabiology 5(5): 328-339. [In Japanese, with English abstract]


Hanlon, R.T. (1982). The functional organization of chromatophores and iridescent cells in the body patterning of Loligo plei (Cephalopoda: Myopsida). Malacologia 23(1): 89-119. LINK


Forsythe, J.W., & Hanlon, R.T. (1981). First rearing of Octopus joubini Robson, 1929 on mysidacean and caridean shrimps. Bull. Amer. Malac. Union 1980: 42- 45.

Hixon, R.F., Hanlon, R.T., & Hulet, W.H. (1981). Growth and maximal size of the long- finned squid Loligo pealei in the northwestern Gulf of Mexico. J. Shellfish Res. 1(2): 181-185.


Hanlon, R.T., & Hixon, R.F. (1980). Body patterning and field observations of Octopus burryi Voss, 1950. Bull. Mar. Sci. 30(4): 749-755. PDF

Forsythe, J.W., & Hanlon, R.T. (1980). A closed marine culture system for rearing Octopus joubini and other large-egged benthic octopods. Lab. Anim. 14(2): 137-142. DOI:10.1258/002367780780942737

Hanlon, R.T., Hixon, R.F., Forsythe, J.W., & Hendrix, J.P., Jr. (1980). Cephalopods attracted to experimental night lights during a saturation dive at St. Croix, U.S. Virgin Islands. Bull. Amer. Malac. Union 1979: 53-58.

Hixon, R.F., Hanlon, R.T., Gillespie, S.M., & Griffin, W.L. (1980). Squid fishery in Texas: Biological, economic, and market considerations. Mar. Fish. Rev. 42(7-8): 44-50. LINK

Hulet, W.H., Hanlon, R.T., & Hixon, R.F. (1980). Lolliguncula brevis - a new squid species for the neuroscience laboratory. TINS 3(4): 1-2.

McConathy, D.A., Hanlon, R.T., & Hixon, R.F. (1980). Chromatophore arrangements of hatchling loliginid squids (Cephalopoda, Myopsida). Malacologia 19(2): 279-288. LINK

Yang, W.T., Hanlon, R.T., Hixon, R.F., & Hulet, W.H. (1980). First success in rearing hatchlings of the squid Loligo pealei Lesueur 1821. Malacol. Rev. 13(1- 2): 79-80.

Yang, W.T., Hanlon, R.T., Krejci, M.E., Hixon, R.F., & Hulet, W.H. (1980). Culture of California market squid from hatching - first rearing of Loligo to subadult stage. Aquabiology 2(6): 412-418. [In Japanese, with English abstract]


Hanlon, R.T., Hixon, R.F., Hulet, W.H., & Yang, W.T. (1979). Rearing experiments on the California market squid Loligo opalescens Berry, 1911. Veliger 21(4): 428-431.

Hulet, W.H., Villoch, M.R., Hixon, R.F., & Hanlon, R.T. (1979). Fin damage in captured and reared squids. Lab. Anim. Sci. 29(4): 528-533.

Rathjen, W.F., Hixon, R.F., & Hanlon, R.T. (1979). Squid fishery resources and development in the Northwest Atlantic and Gulf of Mexico. Proc. Gulf Carib. Fish. Inst. 31: 145-157. LINK


Hanlon, R.T. (1977). Laboratory rearing of the Atlantic reef octopus, Octopus briareus Robson, and its potential for mariculture. Proc. World Maricult. Soc. 8(1-4): 471-482. DOI: 10.1111/j.1749-7345.1977.tb00137.x

Rathjen, W.F., Hanlon, R.T., & Hixon, R.F. (1977). Is there a squid in your future? Proc. Gulf Carib. Fish. Inst. 29: 14-25. LINK


Hanlon, R.T., Kaufman, L. (1976). Associations of seven West Indian reef fishes with sea anemones. Bull. Mar. Sci. 26(2): 225-232.LINK