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How, M. J., Porter, M., Radford, A. N., Feller, K., Temple, S. E., R.L., C.,... Roberts, N. W. (2014). Out of the blue: the evolution of horizontally polarized signals in Haptosquilla (Crustacea, Stomatopoda, Protosquillidae). Journal of Experimental Biology, 217, 3425-3431. https://doi.org/10.1242/jeb.107581 Peer reviewed version Link to published version (if available): 10.1242/jeb.107581 Link to publication record in Explore Bristol Research PDF-document University of Bristol - Explore Bristol Research General rights This document is made available in accordance with publisher policies. Please cite only the published version using the reference above. Full terms of use are available: http://www.bristol.ac.uk/pure/about/ebr-terms

J Exp Biol Advance Online Articles. First posted online on 7 August 2014 as doi:10.1242/jeb.107581 Access the most recent version at http://jeb.biologists.org/lookup/doi/10.1242/jeb.107581 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 K d, ^ W Martin J How 1, Megan L Porter 2, Andrew N Radford 1, Kathryn D Feller 3, Shelby E Temple 1, Roy L Caldwell 4, N Justin Marshall 5, Thomas W Cronin 3 and Nicholas W Roberts 1* 1. School of Biological Sciences, University of Bristol, Tyndall Avenue, Bristol, BS8 1TQ, UK 2. Department of Biology, University of South Dakota, Vermillion, SD 57069, USA 3. Department of Biological Sciences, University of Maryland Baltimore County, 1000 Hilltop Circle, Baltimore, Maryland 21250, USA 4. Department of Integrative Biology, University of California, Berkeley, CA 94720, USA 5. Queensland Brain Institute, The University of Queensland, St Lucia, QLD 4072, Australia *Corresponding author: nicholas.roberts@bristol.ac.uk These authors contributed equally to this work. Running title: Evolution of polarization signals Keywords stomatopod, mantis shrimp, polarization vision, signal evolution, sensory bias, multi-modal signal. Š Žƒ œƒ ˆŽ Š ˆ ƒ Šƒ ƒ ƒ ƒž ˆ ƒ 36 37 38 39 40 41 ˆ ˆˆ ƒžž Šƒ ƒž ƒ Šƒ ƒ ƒ ƒ ƒ ŠƒŽ ŽŽ ƒ ƒž ƒž Šƒ ƒ Žƒ œ ˆ ƒ Šƒ ˆ ƒ ˆƒ Ž š Ž ƒž ƒž ƒž Š ƒ Š Ž ˆ Žƒ œ ƒž ˆƒ ŽŽƒ ƒ ƒ ˆ ƒ ƒ ŽŽƒ Žƒ ŽŽ ˆ ƒ 1 2013. Published by The Company of Biologists Ltd

42 43 44 45 46 47 48 š ƒ ƒ Š œ ƒžž Žƒ œ ƒž ƒ ƒ Šƒ Š ˆ Š Žƒ œƒ Š ƒ ƒž ƒ Š ƒž ƒ ˆ Š ƒž Š Ž Š Š ƒ ˆˆ ƒ Šƒ Žƒ œƒ ƒ Š ƒ ŠƒŽŽ ƒ ƒœ ˆ ƒ ˆ Š Ž ˆ Š Žƒ œ ˆ ˆ Š Ž š ƒž ƒ Š ˆˆ ƒ š ƒ ƒ Ž ƒ Ž 49 50 51 52 53 54 55 56 57 58 59 60 61 INTRODUCTION Žƒ œƒ ƒ ƒž ƒž œƒ Šƒ Šƒ Ž Š ƒž ƒ ƒ ƒ ƒ ƒž ƒ ƒ Žƒ Ž ƒž ƒ Š ƒ ƒ Šƒ Žƒ ƒ Š Ž ƒž Žƒ œƒ ƒ ˆ ƒ ƒ Š Žƒ Š ƒ Šƒ ƒ ƒ ƒž Š Ž Š ƒ ƒ ƒ ƒ ŠƒŽ ŽŽ Žƒ œƒ ˆ ƒ ƒ Ž ƒ ƒ ƒ ˆ ˆ ƒ Šƒ Šƒ ƒž ƒž ƒ Š ƒž Š ƒž ƒž Š ƒž Š š ˆ ƒ Žƒ œƒ ˆ ˆ 62 63 64 65 66 67 68 ƒž Š Š ƒž Š ƒ Š ƒ Š ƒž ƒ ƒž Š Žƒ œƒ ˆ ƒž ˆ Ž Š Š ƒ Ž ˆ Žƒ œƒ Š ƒ Š Š Š Ž ˆ Ž ˆ Š Ž Š ŽŽƒ Š Ž Š ˆ Žƒ œƒ ˆ Š š Š Š ƒ ŽŽƒ ƒ Š ƒ ƒ Ž ƒ ˆˆ ƒž Š ƒ Ž ˆ Žƒ œƒ Šƒ ƒžˆ ƒ ƒžƒ ƒ ƒ 2

69 70 71 72 73 74 75 Š ˆ ˆ ƒž ˆ ƒ ƒž ƒ ƒž ƒž Šƒ Šƒ ƒž ƒ ŠƒŽŽ Ž ƒ ƒ ƒ Š ƒ ƒž ˆ Š ŽŽ Ž Š ƒ ƒ ƒ ƒ ƒžž Šƒ ƒˆ ˆ Š ˆ ƒ ƒ ƒ ˆ ƒž ˆƒ Š ƒ ƒž Š Šƒ Š Žƒ œƒ ˆ Ž Š ƒ ƒ Šƒ Š ƒž Š ƒ ƒ ƒ Š Š ƒ Ž ƒ Ž 76 77 78 79 80 81 82 83 84 85 86 ˆ ˆ ƒ Šƒ Šƒ ƒž Š ƒž ƒ Š Š ˆ Ž Šƒ ˆ Š Š Ž ƒž Šƒ ˆ Žƒ œƒ ˆ ž Šƒ ƒ Œï ƒž Š ƒž Šƒ Š Š Žƒ œƒ ƒ Ž Žƒ œƒ Ž ƒž ƒ Š ƒž Š ƒ ƒ Ž Š ƒž Š ƒž ƒž ƒ Š Ž ƒ š ˆ Žƒ œƒ ƒž Žƒ Š ƒž ˆ ŽŽ Š ƒ ƒ ƒ ƒ ˆ Š ˆ Žƒ œƒ Ž Š Ž ƒž Šƒ ƒ Ž Š ƒ ƒž ƒ ƒ ˆ Žƒ œƒ Š ˆˆ Š Žƒ Š 87 88 89 90 91 92 93 Ž Ž Ž ƒ ƒ ŠƒŽŽ Š ƒž ƒž ƒ ƒ ŠƒŽŽ ƒž Š ƒž ƒž Ž ƒž ƒ Ž Žƒ Š ƒž ƒž Šƒ ƒ ƒž ˆ ƒ Š Ž Šƒ ˆ Žƒ œƒ ƒž Š Šƒ ƒž Š Šƒ ƒ ˆ ƒ ƒž ƒž Šƒ ƒ Š Ž ƒ Ž Žƒ Ž Žƒ œ Š ƒž Š ƒž ƒž Š ƒ ƒ ŽŽƒ 3

94 95 96 97 98 99 100 ˆƒ Ž ŽŽ ƒ ƒž ˆ Š ˆ ƒš ŽŽ ˆ Š š ƒžƒ ƒ ƒ Ž ƒ ƒž ŽŽ ƒž ŽŽ ƒ Ž Š ƒž ˆ ƒ ˆ ƒ ŽŽƒˆ ƒš ŽŽ Š ˆƒ Ž ƒž ˆŽ Š ƒž ƒž ŽŽ ƒ Š Ž ƒž ˆ ƒ ŽŽƒ ƒ Ž Ž ƒ ƒ ƒ ƒ ˆ Š ƒ Ž ƒƒ ƒ ƒ Š ˆŽ 101 102 103 104 105 106 107 108 109 110 111 ƒž Š œ ƒžž Žƒ œ Š ƒž ƒž š Ž Š ƒž Ž ƒ ƒ Š ƒ ˆ Žƒ œƒ ƒ ŽŽ ƒ š ƒ Š Š Š Šƒ ƒž ˆˆ ˆ ˆ Žƒ œƒ ƒž ƒ ˆ Š š Ž Š ƒ ƒž ƒ Šƒ ƒž Žƒ œ Ž Ž ˆ ƒ ˆ ƒ ŽŽ ƒ Ž Š ŽŽƒ ƒ Š š ƒ Š š ˆ Š ƒž Žƒ œƒ ƒ Š Š Š Ž ƒ Š Š ƒƒ Ž ƒ Ž ƒ ˆˆ ƒ Ž ˆ Žƒ œƒ ƒžž ƒ Š Ž ˆ ŽŽ Š 112 Ž ˆ Š Žƒ œƒ ˆ ƒš ŽŽ ƒž 113 114 115 116 117 118 RESULTS Responses to polarized stimuli ƒ Ž ƒ ƒžž Š ƒ ˆ ƒ Ž ƒ ƒ Ž ˆ Š Š œ ƒžž Žƒ œ Ž ƒ Šƒ ƒžž Žƒ œ Ž ƒ Ž š ˆ 4

119 120 121 122 123 124 Ž ˆ ˆ Š šƒž ƒ ƒ Š œ ƒžž Žƒ œ Ž Š Š ƒžž ƒ Ž œ Ž ƒ ƒž Š ƒ ˆ ƒ ˆˆ ƒ Ž ƒ Š Žƒ Š Ž ƒž ƒžž χ ˆ 125 126 127 128 129 130 131 132 133 134 135 136 137 Level of discrimination between two angles of linearly polarized light ƒ Š Ž Ž Ž Š Š ˆˆ Š Žƒ œƒ ƒ Ž ˆ Š Ž ƒ ƒ ƒ ƒ Ž ƒžƒ Ž ƒ Ž ˆ Ž Š ƒ ƒž ƒ Ž Š Žƒ œƒ Ž ƒ ƒž ˆ ƒ ƒ Š Žƒ ƒ ƒ ƒžž ˆ ƒ Š Ž Presence of polarized signals Š ƒš ŽŽ ˆŽ ˆ ƒ Ž ƒ Š šƒ Š ƒ Žƒ Š ƒ ƒ Ž Š Ž ˆŽ ˆ Š 138 139 140 141 142 143 ƒš ŽŽ ƒ Š ƒ ƒžž ƒ ˆŽ ˆ Š Š ŽŽƒ ˆ Š Ž ƒ ŽŽƒ ˆŽ ƒ ƒ Š œ ƒžž Žƒ œ Š ƒ Š ˆŽ ˆ Ž ƒ Žƒ œ ƒžƒ ƒž ˆ Š ˆƒ ŽŽƒ Š Šƒ Ž ƒ ƒ ƒ Ž Š ŽŽƒ ˆƒ ƒ ƒ Šƒ ˆ ƒ ƒžž Šƒ Ž ˆŽ 5

144 145 146 147 148 149 150 ƒš ŽŽ Ž Š ˆŽ ˆ Ž ƒ ƒ ƒ Ž Š ŽŽƒƒ ˆƒ ƒƒ Š œ ƒžž Žƒ œ Š Š ˆ Š ŽŽ ƒ ˆ ˆ Š Šƒ ƒ ƒž œ š ƒ ƒ ƒ Š š Š ŽŽƒ ƒ ŽŽƒˆ Ž Š Ž Ž ƒ Š œ ƒžž Žƒ œ ƒš ŽŽ ˆ Š ŽŽ ƒ š Š ƒ ˆ ƒ ƒ ˆ ˆƒ Ž Šƒ ˆ ƒš ŽŽ ƒž ˆ Š Ž Š 151 152 153 154 155 156 157 158 159 160 161 162 Ž ƒž ƒ Š Š Ž Š œ ƒžž Žƒ œ ƒž Phylogenetic analyses Š Ž ƒ ƒž ˆ ŽŽ Žƒ Š ƒ Žƒ ƒ ƒž Š ŽŽ ƒ ƒ ƒ ƒ Žƒ Š ƒ ŽŽƒ ƒ Š Ž Š Š ƒ ŽŽƒ Š Ž ˆ Šƒ Š ˆ ƒš ŽŽ ƒž Š Ž ƒ Žƒ œ Ž ƒ Žƒ œ DISCUSSION 163 164 165 166 167 168 Ž Š Šƒ ƒž ˆ ƒ Šƒ ƒ Š ƒ Šƒ ƒž ƒ Ž Ž ƒ Ž Žƒ œ Ž ƒžž Š ŽŽ Žƒ ƒ ƒ ƒ Ž ˆ Š œ ƒžž Žƒ œ Ž ƒ Š Š Šƒ ƒ ƒžž Žƒ œ Š ƒ ˆ Š ƒž Ž ƒ Žƒ œƒ ˆ Š ƒš ŽŽ ƒ Š Š 6

169 170 171 172 173 174 175 ƒ ƒ Š Ž ƒ ƒž ƒž Šƒ ˆ Š ŽŽ Ž Š ƒ ŽŽƒ Žƒ Š Ž ƒž Š Ž Š ˆ ƒ ŽŽƒ Ž ƒ Šƒ Š ƒ ƒž Žƒ œ ƒž Š Š Žƒ œƒ ˆ Š ƒž ƒž ƒ ƒ Š œ ƒžž Š ˆ Ž Šƒ Š Šƒ ƒž ƒ Š œ ƒžž Žƒ œ Ž ƒ Š ŽŽ Ž Šƒ ƒ Š Žƒ œƒ ˆ ƒš ŽŽ 176 177 178 179 180 181 182 183 184 185 186 ƒž Š œ ƒž Š ƒ Žƒ Žˆ ƒ ƒ Ž ƒ ƒ Ž ƒ Š ˆ ƒ Š Š ƒ š Š Ž ƒ Ž Žƒ Š ƒ ˆ ƒ Š œ ƒžƒ Ž ˆ Žƒ œƒ ƒ ˆ Š ˆƒ Šƒ Š ƒ Ž ƒž ˆŽ ˆ Œ ƒ ˆ ƒž ƒ Šƒ Ž Ž ƒ ƒ Ž ƒƒž Žƒ ƒ Š Š Ž ˆ ƒ Š Š ƒ Ž ˆ Žƒ œƒ Š ƒ ƒ Ž Ž ˆ ƒ Ž Š ƒž ƒ ˆ ƒ Ž ƒ Ž Žƒ œ ƒž ˆˆ Ž Ž ƒ ˆ Ž ƒ ƒ ƒ Š ƒ 187 188 189 190 191 192 Š Š Š Ž ƒ ˆ ƒ Š Š Šƒ ƒ Š ˆ Ž ƒ ƒ ƒž Ž ˆ Š ƒ Žƒ Ž ƒž ƒ ˆ ˆ Š Š ƒ ƒž ƒ Š Š Šƒ Š ƒ ƒ ƒ ŠƒŽ Žƒ œƒ ƒž ƒ Ž Š Š Ž Ž ˆ Žƒ œƒ ƒ Ž 7

193 194 195 196 197 198 199 Š Ž š ƒ ˆ ƒ Š Š ƒ ƒ ƒ ƒž œ ƒ ƒ Žƒ Ž ƒ Š ƒ ˆ ˆ ƒ Šƒ ƒ ˆ Š ƒž Šƒ Š ˆˆ Ž Š Š Šƒ Š Š Š ƒ ƒ Ž Ž ƒ ƒž ƒ ˆ Š ƒž Š Š Š ƒ Š ƒ ƒ Š Ž ƒž ƒ š ƒ Š Ž ƒ Žƒ œƒ ˆ ƒ ƒ ƒž Š 200 201 202 203 204 205 206 207 208 209 210 ˆ Š Ž Ž š Š ˆ Ž Ž Žƒ Š ƒ ƒž Š Š Š Š Ž ƒ ƒž ƒ ƒ Š ƒ ƒ ƒ Ž Š Ž ˆ ŽŽ ƒ ƒž ƒ ˆ Š ƒ Š Š ƒ ˆ ŽŽ Š ƒ ˆ ƒ ƒ Ž Š ˆ Šƒ Š ˆ Š ƒ Ž ƒž ˆ ƒ Ž ˆ ˆ ƒ ƒž ˆ Ž Š ƒž ˆ Š Žƒ œƒ ƒ Šƒ ƒ ˆ ƒžž ƒ ˆ Š ƒž Š ƒž ƒ ƒžš Š Š Šƒ ƒž š Š Ž ƒž Ž Šƒ Š ƒ ƒ Š Š Ž ˆ Š ƒ ƒ ˆ 211 212 213 214 215 216 217 Š ƒ ƒž Š ƒž ŽŽ ƒ Š Š ˆ Žƒ œƒ ƒˆˆ Šƒ ƒž Š Žƒ œƒ ƒž ƒžž ˆ ƒˆ ƒ ˆ ƒ Š ƒž Ž ƒ ƒ Š ƒ ˆ Š Žƒ œƒ ˆ Š ƒš ŽŽ ƒž ƒ ˆ Ž ƒ Ž ˆ ƒ Š Š Š Šƒ ƒ Šƒ ŽŽ ˆ ƒ ƒ Ž ƒ Šƒ ƒž Š 8

218 219 220 221 222 223 224 ƒ ƒ ƒ Ž Š š Š Ž ƒž Ž ƒ Š ˆˆ ƒ ƒ Žˆ ƒ ƒ ƒ ƒ ƒœ Š Žƒ œƒ ƒ Ž ƒ Ž ˆ ˆ ƒž ˆ ƒ ƒ Žƒ Ž ƒžž ƒ ƒ Ž Ž Š Šƒ Š Šƒ Š ˆ ƒ Šƒ Š ƒ ƒž ˆˆ ƒ Š Ž ˆ Š š ƒ ƒž Š ˆ ˆƒ Ž Ž ƒ Š Šƒ ƒž ƒ ƒ 225 226 227 228 229 230 231 232 233 234 235 Š œ ƒž Žƒ œ Ž Š ƒ ˆ Š š Žƒ ƒ ˆ Š Š Žƒ œ ˆ Š ƒž Šƒ Ž Š œ ƒžž Žƒ œ Š Š ƒ ˆ Žƒ œƒ Š ƒž ƒ ƒ ˆ Š ƒ Š Š š ƒ ƒ Ž ƒ Ž ˆ Š Ž ˆ Š Ž š ƒž ˆ Š ˆ ƒ ƒ ƒ Žƒ Š Žƒ Žƒ œƒ ƒ ˆ Š ƒž ƒ Š ƒ ˆŽ Š ƒ Ž Š ƒž ƒ Žƒ œƒ ƒ Ž ˆ ˆ ˆ ƒ ƒ Š ƒˆ ƒž ƒ ˆ šƒ Ž ƒ Š ƒ ƒ ˆ ƒ ƒ ƒ Ž ˆ Š Š ˆ Ž ƒž ƒ ƒ Ž 236 237 238 239 240 241 ƒ ƒ ƒœ Šƒ ˆ ˆ Žƒ œƒ ƒ Ž ƒž Š ƒ ƒž Š Ž ƒž ƒ ˆ ŽŽ Š Š ƒž ƒ Š ƒž Š ƒžž Š Š Ž ƒž ƒ ƒ Ž š ƒž ƒ Š ˆ ˆ ƒž ƒ ƒ ƒœ Š Ž ƒ Žƒ œƒ ƒž 9

242 243 244 245 246 247 248 ƒ ƒ š ŽŽ Šƒ ƒž ƒ Š ˆ ƒ Ž ˆ ƒž Ž š MATERIALS AND METHODS Animals ƒ Š Š ƒ Ž ˆ ƒ ƒ Šƒ ƒž 249 250 251 252 253 254 255 256 257 258 259 Žƒ œ Ž ŽŽ ƒž ˆ ƒ ƒž ˆ Š Ž ƒ ƒ ˆ ƒž ˆ Š š ˆ ˆˆ Š ˆ ƒ œƒ Žƒ ƒ ƒ ˆ ƒž ƒ (Queensland GBRMPA permit G12/35042.1) ƒž ƒ ƒ ˆ ƒ ƒ ƒž ƒ ƒ ˆŽ Š Š ƒ ƒ ƒ ˆƒ Ž ƒ Š œƒ Žƒ ƒ Š ƒ ƒ ƒž ƒ Ž Š ŽŽ ƒ ƒ ˆ ˆˆ œ Š ŽŽ ƒ Š ƒž Š ˆ Š ˆ Žƒ Relationship between behavioural responses and polarization stimulus 260 261 262 263 264 265 266 content ƒž ƒ Žƒ ƒ š š ƒ ƒ Ž ƒž ƒ Š ƒ ƒ Š ƒž ƒ Žƒ ƒ ƒ Ž ƒ ƒ ƒ ƒ ƒžžƒ ˆˆ Š Ž ƒ ƒž ƒž Š ƒ ƒž ƒ Š Šƒ Š ˆ ƒ ˆ Š ˆ ˆ Š Ž ƒ Š ƒ ƒž ƒ ƒ ƒ ƒ ƒ ƒ Šƒ Š ƒ ˆ Š Ž Š 10

267 268 269 270 271 272 273 ˆ Š ƒ ƒ ˆ ˆ Š ƒ ƒž ƒ ƒ Ž š ƒ ƒž Ž ƒ œ Š ƒ ƒ ƒ ˆ ƒ š ƒ Ž ˆ Š Š ˆ Žƒ œ ˆ Š ƒ ƒ Š Šƒ ƒ ƒž ƒž ˆ Š Žƒ Š Š Ž ƒž Žƒ œƒ Ž ŽŽ ƒ ƒž Ž Š œ ƒž Ž Ž ƒ ŽŽ ƒž Š Ž š ƒ ˆ Š ƒž ˆ Ž ƒ Ž ƒ 274 275 276 277 278 279 280 281 282 283 284 ƒ ˆ ƒ ƒ Š ƒ Žƒ ƒ ƒ Š Ž Ž Ž ˆ Š Žƒ œƒ ˆ Š Ž Š Ž ƒ Ž ˆ Š Žƒ œƒ ƒ Ž ƒ Š œ Ž ƒ ƒ Š ƒ ƒ Š ƒ ƒ Š Ž Ž Š Š Žƒ œƒ ˆ Š ƒ ƒ ƒ ƒ ƒ ƒ ƒ ƒ Žƒ Š Žƒ œ ƒ ƒ ƒ Ž Š Š Š Š ƒ ˆ Š Žƒ œƒ ŽŽ ˆ ƒ Š ˆ Š Ž ˆ ƒ ƒ Ž Š ŽŽƒ ƒž ƒ ƒžƒ ƒ œ ƒ ˆ Ž ƒ Ž ƒ ƒ ƒ Žƒ Ž Ž ƒ Ž Žƒ œ 285 286 287 288 289 290 291 ƒ Šƒ Š ƒ ˆ Š ƒ Ž ƒ ƒ Ž ƒ Š Ž ˆ œ ƒ ˆˆ ˆ Šƒ ƒ Š Š Š ƒ ƒž Š Ž Š ˆ Š ˆ ƒžƒ ƒž ˆ ƒ Š Ž ƒ ƒ ƒ ƒ Š Š Š ƒž ƒ Ž Š Š ˆ ƒ šƒ Ž Š ƒ ƒ ƒ 11

292 293 294 ˆ Š ˆ Š Ž ˆ ƒˆ ƒ ƒž Š ˆ ˆ Š ƒ ƒ ƒ Ž ˆ ƒ Š Ž 295 296 297 298 Discrimination threshold between two angles of linearly polarized light Žƒ Š ƒ ƒ Š Žƒ œƒ ƒ Žƒ ƒ ˆ ƒ ƒž ƒ ƒ ƒž Š ƒ š 299 300 301 302 303 304 305 306 307 308 309 310 š ƒ ƒ ƒ ƒ š ƒ Ž ˆ Š ˆ ƒžž ˆˆ Žƒ œƒ ƒž ˆ ƒž ƒ ƒ Ž Šƒ ƒ Š Žƒ ƒ ƒ ƒ Š ˆ ƒžž Ž Ž Ž š ƒ ˆ Š ƒž ˆ Ž ƒ Ž ƒ ƒ ˆ ƒ ƒ Š ƒ Žƒ ƒ ƒ Š ƒž ƒž ƒ Š Žƒ ˆ Š ƒ ƒ ƒ ƒ ˆ Š Ž Ž ƒ Ž Šƒ ƒ Š ƒ Ž ˆ Žƒ œƒ ƒ ƒ ƒš œ ƒžž Žƒ œ ƒ Š Šƒ Š Ž Š ƒ Š Ž ƒ ƒž ƒ ƒ ƒ ƒ Š ˆ Š ˆ ƒ ƒ ƒžž Ž 311 312 313 314 315 316 ƒ ƒ ƒ ƒžž ƒ Š ƒ ƒ Š Š Ž š ƒ Š š Š œ Š Ž ƒ ˆƒ ƒ ƒž ƒ ƒ Ž Ž ˆ Š Š Š ˆ Š ƒ ƒ ƒ ˆ ƒ ƒ ƒ ƒ Š ˆ ƒ ƒˆ Š Ž ƒ ƒ 12

317 318 319 320 ƒ ƒž ˆ Ž ƒ Ž ˆ ƒ ˆ Ž Š Ž ƒ ˆ ŽŽ ƒ œ ƒ Š ƒž Ž ƒ ƒ œ ƒ 321 322 323 Polarization analysis of the maxilliped signals ƒ ˆ Š ƒš ŽŽ ˆ ƒ Ž ƒ 324 325 326 327 328 329 330 331 332 333 334 335 Š š ƒ Š Š ƒ œ ƒ œž ƒ ƒ š Œ ƒ ƒ ƒž ƒ ƒ ƒ Ž ŽŽ ƒ Š š Š Žƒ Š ƒ ƒž ˆŽ ƒ ƒ ˆ Š ƒ ˆ ƒ ƒ ƒ ŠƒŽ Ž Š ƒ ƒ Š ƒ ˆ ƒž Žƒ ˆ Š ƒ ŽŽ ƒ Š ƒš ŽŽ ƒžž Š ˆŽ Ž Š ƒ ŽŽ ƒ Š ƒ ˆ ƒž Žƒ ˆ Š ƒ ƒ ˆ ƒ ƒ ƒ Š œ ƒž ƒ ƒž Žƒ œƒ ˆ Ž Žƒ Š ƒ Š ˆ Š ˆŽ Ž Š Š ŽŽ ƒ Š Žƒ œ ˆŽ ƒ ƒž ƒ Š Ž ƒ Žƒ œ ƒ ˆ Š ƒš ŽŽ ˆ Š ƒ 336 337 338 339 340 341 ˆ ƒ ƒ Š ˆƒ Ž ƒ Ž ƒ Šƒ ƒ ƒžž Š ƒš ŽŽ Š Š ƒ ƒ ƒ Ž Ž ƒ Žƒ œ Phylogenetic analyses 13

342 343 344 345 346 347 348 ƒ Š ƒž Ž ƒ ƒ Š ƒ ˆ Ž ƒ Žƒ œƒ ƒž Š Š ƒ ŽŽƒ ˆ Š Ž ƒ ƒ Š ƒž ˆ ƒžžƒ ƒ Žƒ Ž Š ƒ ˆ ƒ ƒ ƒ ƒ ˆ ŽŽ Š Š ˆ ƒž Ž ƒž ƒ Ž ƒž ƒ ƒ ˆ Š Š ƒ ˆƒ Ž ŽŽ ƒ ƒ ˆƒ Ž ƒ Ž ƒ Ž 349 350 351 352 353 354 355 356 357 358 359 ƒ Ž ƒ ƒ Ž ƒ Š Š Š Ž ƒ ƒ ƒ ƒ ƒ ƒ š ˆ Ž ˆ Š Š š ƒ ƒ Š ƒž ƒ Š ƒ Ž ƒ ƒž Š Š Š ˆ ƒ ƒ Žƒ Ž ƒ ƒ Ž ƒžƒ Ž ˆ ˆ ŽŽ ˆ ƒ ƒ ƒ ƒ Ž ˆ Š ƒ ƒž Š ƒ Š ƒˆˆ Œ ƒž ƒ Š ƒž ƒ Š ƒž Š ˆ ˆ Ž Š ƒ Š ƒ ƒžž ƒž Š ƒ Žƒ ƒ ƒ Š ˆ 360 361 362 363 364 365 366 Š ƒ ƒ ƒ Š ƒ ƒ ƒ ƒ Š Ž ƒ œ š Ž ƒ ƒš Ž Š š Š ƒ ƒ ƒ Ž Š ˆ ƒ ˆ Š Ž ƒ Š ƒž ƒ ƒ ƒ ƒ ƒ ƒ ƒž ŽŽ ƒž Š ƒ ƒ ˆ Š šƒ ƒž ƒ ƒ ƒžž ˆ Š ƒž ƒ ŽŽ 14

367 368 369 ƒ ƒ ƒž œ Š Š ƒ ƒ Ž ƒ Š ƒ Š ˆ Žƒ ƒ Žƒ Ž š ƒ Š Ž ˆ Œ Ž ƒ ƒ Ž ƒ ƒ 370 371 372 373 Statistical analysis ŽŽ ƒ ƒž ƒ ƒž ƒ ˆ ƒ ƒž ƒ Ž Š Š œ ƒžž ƒžž Žƒ œ 374 375 376 377 378 379 380 381 382 383 384 Ž Ž ƒ ƒž Ž š ƒ ƒ ˆˆ ƒž Žƒ ƒ ƒž ƒžž ƒ Š ƒž ƒ ƒ ˆ ƒ ˆˆ ƒ Žƒ ƒ ƒ ƒž ƒ ƒ ACKNOWLEDGEMENTS ƒ ƒ ˆ Ž Š œƒ Žƒ ƒ Š ƒ ƒˆˆ ˆ ƒžž Š ƒ ƒ Š ŽŽ ˆ Š Ž Š Š ƒ ƒž ƒ ƒž ƒ ƒ Ž ƒ ˆ ƒ ƒ FUNDING 385 386 387 388 Š Ž ƒ Ž ƒž ƒ Š Ž ƒ ƒ Š ˆˆ ˆ ˆ ƒ Š ƒ Š ƒ ƒ ˆƒ Ž Ž ƒ ƒ œƒ Žƒ ƒž ŽŽ Š 389 390 REFERENCES 15

391 392 393 394 Barber, P. and Boyce, S. L. (2006). Estimating diversity of Indo-Pacific coral reef stomatopods through DNA barcoding of stomatopod larvae. Proc. R. Soc. B. 273, 2053 2061. 395 396 397 Caldwell, R. and Dingle, H. (1975). Ecology and evolution of agonistic behavior in stomatopods. Naturwissenschaften 62, 214-222. 398 399 400 401 402 403 404 405 406 407 408 409 Candolin, U. (2003). The use of multiple cues in mate choice. Biol. Rev. 78, 575-595. Chiou, T. H., Cronin, T.W., Caldwell R.L. and Marshall, J. (2005). Biological polarized light reflectors in stomatopod crustaceans. Proc. SPIE 5888, 58881B. Chiou, T. H., Mäthger L. M., Hanlon, R. T. and Cronin, T. W. (2007). Spectral and spatial properties of polarized light reflections from the arms of squid (Loligo pealeii) and cuttlefish (Sepia officinalis L.). J. Exp. Biol. 210, 3624-3635. Chiou, T. H., Kleinlogel, S., Cronin, T., Caldwell, R., Loeffler, B., Siddiqi, A. and Marshall, J. (2008). Circular polarization vision in a stomatopod crustacean. Curr. 410 411 412 413 414 415 Biol. 18, 429-434. Chiou, T. H., Marshall, N. J., Caldwell, R. L. and Cronin, T. W. (2011). Changes in light-reflecting properties of signalling appendages alter mate choice behaviour in a stomatopod crustacean Haptosquilla trispinosa. Mar. Freshw. Behav. Physiol. 44, 1-11. 16

416 417 418 419 Cronin, T. W., Marshall, N. J. and Land, M. F. (1991). Optokinesis in Gonodactyloid mantis shrimps (Crustacea, Stomatopoda, Gonodactylidae). J. Comp. Physiol. A. 168, 233-240. 420 421 422 Cronin, T. W., Caldwell, R. L. and Marshall, J. (2001). Sensory adaptation: tunable colour vision in a mantis shrimp. Nature 411, 547-548. 423 424 425 426 427 428 429 430 431 432 433 434 Cronin, T. W., Shashar, N., Caldwell, R. L., Marshall, J., Cheroske, A. G. and Chiou, T. H. 2003a). Polarization signals in the marine environment. Proc. SPIE 5158, 85-92. Cronin, T. W., Shashar, N., Caldwell, R. L., Marshall, J., Cheroske, A. G. and Chiou, T. H. 2003b). Polarization vision and its role in biological signaling. Integr. Comp. Biol. 43, 549-558. Cronin, T. W., Chiou, T. H., Caldwell, R. L., Roberts, N. and Marshall, J. (2009). Polarization signals in mantis shrimps. Proc. SPIE 7461, 74610C. 435 436 437 438 439 440 Cronin, T. W., Johnsen, S., Marshall, J. and Warrant, E. J. (2014). Visual Ecology. Princeton New Jersey, Princeton University Press. Dacke, M., Nilsson, D-E., Scholtz, C. H., Byrne, M. and Warrant, E. J. (2003). Animal behaviour: Insect orientation to polarized moonlight. Nature 424, 33. 17

441 442 443 Dingle, H. and Caldwell, R. (1969). Aggressive and territorial behaviour of mantis shrimp Gonodactylus bredini Manning (Crustacea, Stomatopoda). Behaviour 33, 115-136. 444 445 446 Endler, J. and Basolo, A. (1998). Sensory ecology, receiver biases and sexual selection. Trends Ecol. Evol. 13, 415-420. 447 448 449 450 451 452 453 454 455 456 457 458 459 Guindon, S. and Gascuel, O. (2003). A simple, fast, and accurate algorithm to estimate large phylogenies by maximum likelihood. Syst. Biol. 52, 696-704. Guilford, T. and Dawkins, M. (1991). Receiver psychology and the evolution of animal signals. Anim. Behav. 42, 1-14. Hasson, O. (1991). Sexual displays as amplifiers: practical examples with an emphasis on feather decorations. Behav. Ecol. 2, 189-197. Hebets, E. A. and Papaj, D. R. (2005). Complex signal function: developing a framework of testable hypotheses. Behav. Ecol. Sociobiol. 57, 197-214. 460 461 462 463 464 465 How, M. J., Pignatelli, V., Temple, S. E., Marshall, N. J. and Hemmi, J. M. (2012). High e-vector acuity in the polarisation vision system of the fiddler crab Uca vomeris. J. Exp. Biol. 215, 2128-2134. Horváth, G. and Varjú, D. (2004). Polarized light in animal vision: polarization patterns in nature. Springer, U.K. 18

466 467 468 469 Johnsen, S., Marshall, N. J. and Widder, E. A. (2011). Polarization sensitivity as a contrast enhancer in pelagic predators: lessons from in situ polarization imaging of transparent zooplankton. Phil. Trans. R. Soc. B 366, 655-670. 470 471 472 Katoh, K., Misawa, K., Kuma, K. I. and Miyata, T. (2002). MAFFT: a novel method for rapid multiple sequence alignment based on fast Fourier transform. 473 474 475 476 477 478 479 480 481 482 483 484 Nucleic Acids Res. 30, 3059-3066. Katoh, K., Kuma, K. I., Toh, H. and Miyata, T. (2005). MAFFT version 5: improvement in accuracy of multiple sequence alignment. Nucleic Acids Res. 33, 511-518. Kleinlogel, S. and Marshall, N. J. (2006). Electrophysiological evidence for linear polarization sensitivity in the compound eyes of the stomatopod crustacean Gonodactylus chiragra. J. Exp. Biol. 209, 4262-4272. Labhart, T. and Meyer, E. P. (1999). Detectors for polarized skylight in insects: a survey of ommatidial specializations in the dorsal rim area of the compound eye. 485 486 487 488 489 490 Microsc. Res. Tech. 47, 368-379. Land, M. F., Marshall, J. N., Brownless, D. and Cronin, T.W. (1990). The eyemovements of the mantis shrimp Odontodactylus scyllarus (Crustacea, Stomatopoda). J. Comp. Physiol. A. 167, 155-166. 19

491 492 493 Marshall, N. J., Land, M. F., King, C. A. and Cronin T. W. (1991). The compound eyes of mantis shrimps (Crustacea, Hoplocarida, Stomatopoda). I. Compound eye structure: the detection of polarized light. Phil. Trans. R. Soc. B 334, 33-56. 494 495 496 Mäthger, L. M. and Hanlon, R. T. (2006). Anatomical basis for camouflaged polarized light communication in squid. Biol. Lett. 2, 494-496. 497 498 499 500 501 502 503 504 505 506 507 508 509 Mäthger, L. M., Shashar, N. and Hanlon, R. T. (2009). Do cephalopods communicate using polarized light reflections from their skin? J. Exp. Biol. 212, 2133-2140. Miller, M. A., Holder, M. T., Vos, R., Midford, P. E., Liebowitz, T., Chan, L., Hoover, P. and Warnow, T. (2009-08-04). The CIPRES Portals. CIPRES. (Archived by WebCite at Š ƒ Ž ƒ). Accessed September 29 2010. Partan, S. R. and Marler, P. (2005). Issues in the classification of multimodal communication signals. Am. Nat. 166, 231-245. 510 511 512 513 514 515 Pignatelli, V., Temple, S. E., Chiou, T. H., Roberts, N. W., Collin, S. P. and Marshall, N. J. (2011). Behavioural relevance of polarization sensitivity as a target detection mechanism in cephalopods and fishes. Phil. Trans. R. Soc. B 366, 734-741. Porter, M. L., Bok, M. J., Robinson, P. R. and Cronin, T. W. (2009). Molecular diversity of visual pigments in Stomatopoda (Crustacea). Vis. Neurosci. 26, 255-265. 20

516 517 518 519 Porter, M. L., Zhang, Y., Desai, S., Caldwell, R. L. and Cronin, T. W. (2010). Evolution of anatomical and physiological specialization in the compound eyes of stomatopod crustaceans. J. Exp. Biol. 213, 3473-3486. 520 521 522 Roberts, N. W., Chiou, T. H., Marshall, N. J. and Cronin, T. W. (2009). A biological quarter-wave retarder with excellent achromaticity in the visible 523 524 525 526 527 528 529 530 531 532 533 534 wavelength region. Nat. Photonics, 3, 641-644. Roberts, N. W., Porter, M. L. and Cronin, T. W. (2011). The molecular basis of mechanisms underlying polarization vision. Phil. Trans. R. Soc. B. 366, 627-637. Rossel, S. and Wehner, R. (1986). Polarization vision in bees. Nature 323, 128-131. Shashar, N., Rutledge, P. and Cronin, T. (1996). Polarization vision in cuttlefish - A concealed communication channel? J. Exp. Biol. 199, 2077-2084. Shashar, N., Johnsen, S., Lerner, A., Sabbah, S., Chiao, C. C., Mäthger, L. M. and Hanlon, R. T. (2011). Underwater linear polarization: physical limitations to 535 536 537 538 539 biological functions. Phil. Trans. R. Soc. B 366, 649-654. Stamatakis, A. (2006). RAxML-VI-HPC: Maximum likelihood-based phylogenetic analyses with thousands of taxa and mixed models. Bioinformatics 22, 2688-2690. 21

540 541 Stamatakis, A., Hoover, P. and Rougemont, J. (2008). A rapid bootstrap algorithm for the RAxML Web servers. Syst. Biol. 57, 758-771. 542 543 544 545 Temple, S. E., Pignatelli, V., Cook, T., How, M. J., Chiou, T. H., Roberts, N. W. and Marshall, N. J. (2012). High-resolution polarisation vision in a cuttlefish. Curr. Biol. 22, R121-R122. 546 547 548 549 550 551 552 553 554 555 556 557 558 Thoen, H. H., How, M. J., Chiou, T. H. and Marshall, N. J. (2014). A new form of colour vision in Mantis shrimps. Science 343, 411-413. Waterman, T. H. (1954). Polarization patterns in submarine illumination. Science 120, 927-932. Wehner, R. (1976). Polarized-light navigation by insects. Sci. Am. 235, 106-115. Wehner, R. and Labhart, T. (2006). Polarisation vision. In Invertebrate Vision (eds. Warrant E., Nilsson D. -E.), Cambridge University Press, Cambridge, UK. 291-348. ƒ 559 560 561 562 563 564 ŽŽ ƒ šƒ Ž Š ƒ ˆ Š ƒš ŽŽ ƒž ƒ Ž Ž ƒƒ ƒ ƒ ƒ Ž ˆ Š ƒ Ž ˆ ˆ ƒ Š ƒž Š ƒžž ƒ Š œ ƒžž Žƒ œ Ž ˆ 22

565 566 567 568 569 570 571 Ž ƒ ƒ Š ƒ Ž Š ˆ ƒž Šƒ Š Šƒ ƒ Ž ƒ Ž ƒ Š š Responses of H. trispinosa (black dots) to differences between the angles of polarization of the stimulus and the background (x-axis). The response data are fitted with a hyperbolic tangent (dashed line). The background level of false positive 572 573 574 575 576 577 578 579 580 581 582 583 responses are represented for each stimulus type (white dots) and as an overall mean (dotted line). McNemar s test was used to determine which response values differed from the level of false positives (* = p < 0.05). ƒ ˆ Š ƒš ŽŽ ƒ Ž ƒ Š š ƒ ƒ Š Ž ƒ Š ˆŽ ƒˆ Š ƒ ƒ Š Ž ƒ Š ƒ Š ƒž Ž Ž Š Š œ ƒžž Žƒ œ ˆŽ ƒ ƒ Ž Š ƒžž Žƒ œ ˆŽ ƒ Š ƒžƒ Š œ ƒž Ž Žƒ Š ƒš ˆ Š ƒš ŽŽ 584 585 586 587 588 589 ƒš Ž Ž Š Š Ž ˆ ŽŽ Žƒ Š ƒ ˆ Š ƒ Ž ƒ ƒ Š ƒž ƒ ƒ Š ƒ ŽŽƒ ƒ Š ˆƒ Ž ŽŽ ƒ ƒ ƒ ƒ Ž Š Š ƒ ˆ Ž ƒž ƒ Žƒ œ ƒž Š ƒš ŽŽŽ Šƒ ƒ 23

590 591 592 Š Š Ž ƒ Ž ƒ Š ƒ ƒž ƒ Š š ƒžž ˆ Š ŠŠƒ ƒ ƒ Š œ ƒžž Žƒ œ Ž ŽŽ ƒ Š ƒ Š ƒ ƒ ˆ Š ŽŽ ƒ 593 594 595 596 Š ƒ ƒ ƒ ˆ Š š ƒž ƒ ƒ ƒ Š ƒ ˆ ˆ Š šƒ Ž ƒ ˆ Š ƒž œ ƒ ƒ ˆ Š Š œ ƒžž Žƒ œ Ž ƒ ƒˆ ˆ 597 598 599 600 601 602 603 604 605 606 607 608 ƒ Ž Š šƒ Ž ˆ Š ƒžƒ Š œ ƒž Žƒ œƒ ŽŽ ƒ ƒ ˆ Š Šƒ ƒž ƒ ƒ ˆ Š ƒ ˆ ƒ ˆ ƒ ŽŽ ƒ Š ƒž ƒ ƒ ƒ ƒ Ž Žƒ œ ƒ Ž Š œ ƒžž Žƒ œ ƒ ƒžž Žƒ œ ƒ ƒ Š ƒ ƒ Ž ˆ ƒ ƒ š ƒ Ž ƒ ƒ Š ˆ ƒ Š Š ˆ Š Ž Š ƒ Š Š ƒˆ Š Ž ƒ Š ˆ ƒž ƒ Š Š ƒ š ƒˆ Š Ž Š ƒ Šƒ Š ƒ Žƒ ƒ ƒ ˆ Š ƒž ƒ ƒˆ ˆ Š ˆ Š Ž Žƒ œ 609 610 611 612 613 614 ƒ Ž Š ƒ ˆ ŽŽ Š ˆ ƒ Žƒ Š Ž ƒ Š Ž ƒ ƒ ƒ ˆ ˆ 24

615 616 617 618 619 620 621 622 623 624 625 626 627 628 629 630 Figures 631 25

632 633 634 635 636 637 638 639 640 641 642 643 644 645 646 647 648 649 650 651 652 653 654 655 656 FIGURE 1 26

657 658 659 660 FIGURE 2 27

661 662 663 664 665 666 667 668 669 670 671 672 FIGURE 3

673 674 675 FIGURE 4 29

676 677 678 679 680 681 682 683 684 685 686 687 688 689 690 FIGURE 5 30

691 692 693 694 695 696 697 698 699 700 701 702 703 704 705 706 707 708 709 710 711 712 713 714 FIGURE 6 31

715 716 717 718 719 720 721 FIGURE 7 32