Hanlon Lab – List of Publications

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

Allen, J.J., Akkaynak, D., Schnell, A. & 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. & Hanlon, R.T. 2017. Small-scale rearing of cuttlefish (Sepia officinalis) for research purposes. Marine & Freshwater Behaviour and Physiology. doi: 10.1080/10236244.2017.1343631


Schnell, A., Smith, K-L., Hanlon, R.T., Hall, K. & Harcourt, R. 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. 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


Buresch, K. C., Ulmer, K. M., Akkaynak, D., Allen, J. J., Mathger, 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). doi: 10.1575/1912/7383

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., Cramer, C., McAnulty, S., Davison, W., Mathger, 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., 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. (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., Mathger, 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., Buresch, K., 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), 4352–6. 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.

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. epub ahead of print

Akkaynak, D., Allen, J. J., Mäthger, L. M., Chiao, C.-C., Hanlon, R. T. 2013. Quantification of cuttlefish (Sepiaofficinalis) 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.

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.

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

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 Research83: 40-47.

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.

Kreit, E., Mäthger, L., 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

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.

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

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 Materials23 (32): 3980-3989.

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.

Nordstrom, Karin and Gonzalez-Bellido, Paloma T. 2013. Invertebrate Vision: Peripheral Adaptation to Repeated Object Motion. Current Biology. Volume 23, Issue 15, 5 August 2013, Pages R655–R656

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 Ecology447:65-74.
Staudinger, M.D., Buresch, K.C., Mäthger, L.M., Fry, C., McAnulty, S., Ulmer, K.M., Hanlon, R.T. Defensive responses of cuttlefish to different teleost predators. 2013. Biological Bulletin 225 (3): 161-174.

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.


Gonzalez-Bellido PT, Wardill TJ. 2012. Labeling and confocal imaging of neurons in thick invertebrate tissue samples. Cold Spring Harb Protoc. doi: 10.1101/pdb.prot069625

Wardill TJ, Gonzalez-Bellido PT, Crook RJ, Hanlon RT. 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.

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.

Mäthger, L.M., Bell, G., Kuzirian, A.M., Allen, J.J. and Hanlon, R.T. 2012. How does the blue-ringed octopus 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.

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.

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.

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.

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.

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.

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

Young, M., Kvitek, R., Iampietro, P., 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.

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.


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: 549-560.

Chiao, C-C., Chubb, C., Buresch, K., 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.

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. (Editor’s Choice Article and Cover Shot)

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

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.

Mäthger, L.M., Roberts, S., & Hanlon, R.T. 2010. Evidence for distributed light sensing in the skin of cuttlefish, Sepia officinalis. Biology Letters 6: 600-603.

Mooney, T. A., Hanlon, R.T., Christensen-Dalsgaard, J., Madsen, P.T., Ketten, D.R., & Nachtigall, P.E. 2010. Hearing 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.


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.

Mäthger, L.M., Denton, E.J., Marshall, J., & Hanlon, R.T. 2009. Mechanisms and behavioral functions of structural coloration in cephalopods. Journal of the Royal Society Interface 6: S149-S164.

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)

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: 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.

Buresch, K.C., Maxwell, M.R., Cox, M.R., & Hanlon, R.T. 2009. Temporal dynamics of mating and paternity inthe squid Loligo pealeii, Marine Ecology Progress Series 387: 197-203.

Crook, R., Hanlon, R.T., & Basil, J. 2009. Memory of visual and topographical features suggests spatial learning in the ancient cephalopod Nautilus. J. Comparative psychology 123 (3): 264-274.

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 20: 1-6.


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.

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.

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, 25 (3): 588-599.

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.

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: 1242-1253.

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: 405-413.

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.

Hanlon, R.T.,& Forsythe, J.W. 2008. Sexual cannibalism by Octopus cyaneaon a Pacific coral reef. Marine and Freshwater Behaviour and Physiology. 41: 19-28.


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.

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

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.

Chiou, T.H., Hanlon, R.T., 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.

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 47:2223-2235.

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.


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.

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

Mäthger, L.M., Barbosa, A., Miner, S., & Hanlon, R.T. 2006. Color blindness and contrast perception in cuttlefish (Sepia officinalis). Vision Research 46: 1746-1753.

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


Hanlon, R.T., Naud, M.-J., Shaw, P.W., & Havenhand, J.N. 2005. Transient sexual mimicry leads to fertilisation. Nature 430: 212.

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.

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

Mensinger, A.F., Hanlon, R.T.,Walsh, P.J. 2005. Blood biochemistry of the toadfish Opsanus tau. Journal of Aquatic Animal Health 17: 170-176.

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.

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.


Naud, M.-J., Hanlon, R.T., Hall, K.C., Shaw, P.W., & Havenhand, J.N. 2004. Behavioral and genetic assessment of mating success in a natural spawning aggregation of the giant cuttlefish (Sepia apama) in southern Australia. Animal Behaviour 67: 1043-1050.

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 145:923-930.

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: 389-392.

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: 1-3.

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


Hanlon, R.T., & Shashar, N. 2003. Aspects of the sensory ecology of cephalopods. In: Sensory Processing in the Aquatic Environment. Collin, S.P., Marshall, N.J. (eds.). Springer-Verlag, New York, pp. 266-282.

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: 49-58.

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 65:289-299.

Karson, M.A., Boal, J.G, Usook, S., & Hanlon, R.T. 2003. Experimental evidence for spatial learning in the cuttlefish, Sepia officinalis. J. Comp. Psychol. Vol. 117, No. 2, 149-155.

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 (Mollusca: Cephalopoda). J. Chem. Ecol. 29(3):547-560.

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 coloration in flounder and cuttlefish revealed by independent component analysis. Comput. Neural Syst. 14: 321-333.


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.

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.

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.

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.

O’Dor, R.K., Adamo, S., 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.

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: 232-234.


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.

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.

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.

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. 201:282-283.

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. 201:267-268.


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.

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.

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 52:141-153.

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: 517-521.

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.

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.


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.

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.

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: 802-811.

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.

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

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

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.

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: 256.

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.


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

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.

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: 99-115.

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.

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.

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.

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.


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.

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

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.

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.

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: 364-374.

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.

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.

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

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

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.


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. 1996. Evolutionary games that squids play: Fighting, courting, sneaking, and mating behaviors used for sexual selection in Loligo pealei. Biol. Bull. 191: 309-310.

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: 265-267.


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.


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.

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.

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


Jackson, G.D., Arkhipkin, A.I., Bizikov, V.A., & Hanlon, R.T. 1993. Laboratory and field corroboration of age and growth from statoliths and gladii of the loliginid squid Sepioteuthis lessoniana (Mollusca: Cephalopoda). Recent Advances in Cephalopod Fisheries Biology. Okutani, T., O’Dor, R.K., Kubodera, T. (eds.) Tokai University Press, Tokyo, pp. 189-199.


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: 431-442.

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

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.


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.

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.

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.

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.


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.

Adams, P., 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.

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- 14.

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.

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: 15- 34.

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.


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.

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.

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.

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.

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.


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

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.

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: 451-455.

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: 1-25.

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.

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.

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.

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.


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.

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.

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.

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.

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.

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.

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.

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.

Marthy, H.-J., & Hanlon, R.T. 1983. Effects of laser microirradiation in chromatophore organs of embryonic and juvenile cephalopods. Mikroscopie (Wien) 40: 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: 77-88.

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.


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.

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.

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.

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.

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.


Hanlon, R.T. 1977. Laboratory rearing of the Atlantic reef octopus, Octopus briareus Robson, and its potential for mariculture. Proc. World Maricult. Soc. 8: 471-482.

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