University of Bristol - Explore Bristol Research. Peer reviewed version. Link to published version (if available): /jeb.

Transcription

1 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, Peer reviewed version Link to published version (if available): /jeb 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:

2 J Exp Biol Advance Online Articles. First posted online on 7 August 2014 as doi: /jeb Access the most recent version at 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 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 Published by The Company of Biologists Ltd

3 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 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 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

4 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 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 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

5 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 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 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

6 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 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 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

7 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 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 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

8 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 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 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

9 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 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 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

10 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 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 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

11 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 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/ ) 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 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

12 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 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 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

13 ˆ Š ˆ Š Ž ˆ ƒˆ ƒ ƒž Š ˆ ˆ Š ƒ ƒ ƒ Ž ˆ ƒ Š Ž 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 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 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

14 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 Polarization analysis of the maxilliped signals ƒ ˆ Š ƒš ŽŽ ˆ ƒ Ž ƒ 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 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

15 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 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 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

16 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 Statistical analysis ŽŽ ƒ ƒž ƒ ƒž ƒ ˆ ƒ ƒž ƒ Ž Š Š œ ƒžž ƒžž Žƒ œ 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 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 REFERENCES 15

17 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, Caldwell, R. and Dingle, H. (1975). Ecology and evolution of agonistic behavior in stomatopods. Naturwissenschaften 62, Candolin, U. (2003). The use of multiple cues in mate choice. Biol. Rev. 78, 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, 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 Biol. 18, 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,

18 Cronin, T. W., Marshall, N. J. and Land, M. F. (1991). Optokinesis in Gonodactyloid mantis shrimps (Crustacea, Stomatopoda, Gonodactylidae). J. Comp. Physiol. A. 168, Cronin, T. W., Caldwell, R. L. and Marshall, J. (2001). Sensory adaptation: tunable colour vision in a mantis shrimp. Nature 411, 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, 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, Cronin, T. W., Chiou, T. H., Caldwell, R. L., Roberts, N. and Marshall, J. (2009). Polarization signals in mantis shrimps. Proc. SPIE 7461, 74610C 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,

19 Dingle, H. and Caldwell, R. (1969). Aggressive and territorial behaviour of mantis shrimp Gonodactylus bredini Manning (Crustacea, Stomatopoda). Behaviour 33, Endler, J. and Basolo, A. (1998). Sensory ecology, receiver biases and sexual selection. Trends Ecol. Evol. 13, Guindon, S. and Gascuel, O. (2003). A simple, fast, and accurate algorithm to estimate large phylogenies by maximum likelihood. Syst. Biol. 52, Guilford, T. and Dawkins, M. (1991). Receiver psychology and the evolution of animal signals. Anim. Behav. 42, Hasson, O. (1991). Sexual displays as amplifiers: practical examples with an emphasis on feather decorations. Behav. Ecol. 2, Hebets, E. A. and Papaj, D. R. (2005). Complex signal function: developing a framework of testable hypotheses. Behav. Ecol. Sociobiol. 57, 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, Horváth, G. and Varjú, D. (2004). Polarized light in animal vision: polarization patterns in nature. Springer, U.K. 18

20 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, 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 Nucleic Acids Res. 30, 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, 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, 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 Microsc. Res. Tech. 47, 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,

21 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, Mäthger, L. M. and Hanlon, R. T. (2006). Anatomical basis for camouflaged polarized light communication in squid. Biol. Lett. 2, Mäthger, L. M., Shashar, N. and Hanlon, R. T. (2009). Do cephalopods communicate using polarized light reflections from their skin? J. Exp. Biol. 212, Miller, M. A., Holder, M. T., Vos, R., Midford, P. E., Liebowitz, T., Chan, L., Hoover, P. and Warnow, T. ( ). The CIPRES Portals. CIPRES. (Archived by WebCite at Š ƒ Ž ƒ). Accessed September Partan, S. R. and Marler, P. (2005). Issues in the classification of multimodal communication signals. Am. Nat. 166, 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, 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,

22 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, 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 wavelength region. Nat. Photonics, 3, 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, Rossel, S. and Wehner, R. (1986). Polarization vision in bees. Nature 323, Shashar, N., Rutledge, P. and Cronin, T. (1996). Polarization vision in cuttlefish - A concealed communication channel? J. Exp. Biol. 199, 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 biological functions. Phil. Trans. R. Soc. B 366, Stamatakis, A. (2006). RAxML-VI-HPC: Maximum likelihood-based phylogenetic analyses with thousands of taxa and mixed models. Bioinformatics 22,

23 Stamatakis, A., Hoover, P. and Rougemont, J. (2008). A rapid bootstrap algorithm for the RAxML Web servers. Syst. Biol. 57, 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-R 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, Waterman, T. H. (1954). Polarization patterns in submarine illumination. Science 120, Wehner, R. (1976). Polarized-light navigation by insects. Sci. Am. 235, Wehner, R. and Labhart, T. (2006). Polarisation vision. In Invertebrate Vision (eds. Warrant E., Nilsson D. -E.), Cambridge University Press, Cambridge, UK ƒ 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

24 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 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 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

25 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 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 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 ƒ Ž Š ƒ ˆ ŽŽ Š ˆ ƒ Žƒ Š Ž ƒ Š Ž ƒ ƒ ƒ ˆ ˆ 24

26 Figures

27 FIGURE 1 26

28 FIGURE 2 27

29 FIGURE 3

30 FIGURE 4 29

31 FIGURE 5 30

32 FIGURE 6 31

33 FIGURE 7 32