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Inconsistent Patterns of Body Size Evolution in Co-Occurring Island Reptiles

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Inconsistent Patterns of Body Size Evolution in Co-Occurring Island Reptiles This is a repository copy of Inconsistent patterns of body size evolution in co-occurring island reptiles. White Rose Research Online URL for this paper: http://eprints.whiterose.ac.uk/158008/ Version: Accepted Version Article: Itescu, Y., Schwarz, R., Donihue, C.M. et al. (7 more authors) (2018) Inconsistent patterns of body size evolution in co-occurring island reptiles. Global Ecology and Biogeography, 27 (5). pp. 538-550. ISSN 1466-822X https://doi.org/10.1111/geb.12716 This is the peer reviewed version of the following article: Itescu, Y, Schwarz, R, Donihue, CM, et al. Inconsistent patterns of body size evolution in co‐ occurring island reptiles. Global Ecol Biogeogr. 2018; 27: 538– 550., which has been published in final form at https://doi.org/10.1111/geb.12716. This article may be used for non-commercial purposes in accordance with Wiley Terms and Conditions for Use of Self-Archived Versions. Reuse Items deposited in White Rose Research Online are protected by copyright, with all rights reserved unless indicated otherwise. They may be downloaded and/or printed for private study, or other acts as permitted by national copyright laws. The publisher or other rights holders may allow further reproduction and re-use of the full text version. This is indicated by the licence information on the White Rose Research Online record for the item. Takedown If you consider content in White Rose Research Online to be in breach of UK law, please notify us by emailing [email protected] including the URL of the record and the reason for the withdrawal request. [email protected] https://eprints.whiterose.ac.uk/ Global Ecology and Biogeography Inconsistent patterns of body size evolution in co-occurring island reptiles Journal: Global Ecology and Biogeography Manuscript ForID GEB-2017-0100.R2 Peer Review Manuscript Type: Research apers Date Submitted by the Author: n/a Complete List of Authors: Itescu, Yu,al- Tel A,i, .ni,ersity, /oolo0y Sch1ar2, Rachel- Tel A,i, .ni,ersity, Department of /oolo0y Donihue, Colin- 3ar,ard .ni,ersity Department of Or0anismic and E,olutionary Biolo0y, Department of Or0anismic and E,olutionary Biolo0y Sla,en5o, Ale6- Tel A,i, .ni,ersity, Department of /oolo0y Roussos, Stephanos- .ni,ersity of North Te6as, Department of Biolo0ical Sciences- Te6as Tech .ni,ersity, Department of Biolo0ical Sciences Sa0onas, 8ostas- 9ueen Mary .ni,ersity of London, School of Biolo0ical and Chemical Sciences :ala5os, Efstratios- National and 8apodistrian .ni,ersity of Athens Faculty of hysics, Department of Biolo0y Foufopoulos, Johannes- .ni,ersity of Michi0an, School of Natural Resources and En,ironment afilis, anayiotis- .ni,ersity of Athens, Biolo0y Meiri, Shai- Tel A,i, .ni,ersity, /oolo0y 8ey1ords: body si2e, competition, island area, isolation, li2ards, predation, sna5es Global Ecology and Biogeography Page 6 of 106 1 2 3 Research paper 1 4 5 6 Inconsistent patterns of body size evolution in co-occurring island reptiles. 2 7 8 1 1 9 Yuval Itescu ([email protected]), Rachel Schwarz 3 10 11 ([email protected]), Colin Donihue2 ([email protected]) , 4 12 13 Alex Slavenko1 ([email protected]), Stephanos A. Roussos3,4 5 14 15 ([email protected]), Kostas Sagonas5 ([email protected]), Efstratios 6 16 17 6 7 D. Valakos ([email protected]), Johannes Foufopoulos ([email protected]), 7 18 19 6 For Peer Review1 20 Panayiotis Pafilis ([email protected]), Shai Meiri ([email protected]) 8 21 22 1 School of Zoology, Tel Aviv University, Tel Aviv 6997801, Israel 23 9 24 2 25 epartment of Organismic and Evolutionary Biology, Harvard University, 10 26 27 Cambridge, MA 021,8, USA 11 28 29 3 30 epartment of Biological Sciences, University of North Te.as, enton, T/ 7620,, 12 31 32 USA 13 33 34 4 35 epartment of Biological Sciences, Te.as Tech University, 0ubboc1, T/ 79209, USA 14 36 37 5 38 School of Biological and Chemical Sciences, 3ueen Mary University of 0ondon, 15 39 40 0ondon E1 2NS, U4 16 41 42 43 6 epartment of Biology, National and 4apodistrian University of Athens, Athens 157- 17 44 45 84, 7reece 18 46 47 48 7School of Natural Resources and Environment, University of Michigan, Ann Arbor, 19 49 50 Ann Arbor, MI 28109, USA 20 51 52 53 54 Key words: body size, competition, island area, isolation, lizards, predation, 21 55 56 snakes. 22 57 58 1 59 60 Page 7 of 106 Global Ecology and Biogeography 1 2 3 Short running title: Body size evolution in insular reptiles 23 4 5 6 Corresponding Author: Yuval Itescu, Department of Zoology, Tel Aviv University, 24 7 8 Tel Aviv 6997801, Israel; phone: +972 640 9811, fax: +972 640 9403; 25 9 10 11 [email protected] 26 12 13 14 Nu ber of words in the abstract: 300 27 15 16 17 Nu ber of words in the ain text: 5515 28 18 19 For Peer Review 20 Nu ber of references: 73 29 21 22 23 30 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 2 59 60 Global Ecology and Biogeography Page 8 of 106 1 2 3 ABSTRACT 31 4 5 6 Ai Animal body sizes are often remarkably variable across islands, but despite 32 7 8 much research we still poorly understand both patterns and drivers of bodyAsize 33 9 10 11 evolution. Theory predicts that interspecific competition and predation pressures are 34 12 13 relaxed on small, remote islands, and that these conditions promote 0odyAsize 35 14 15 evolution. Be studied 0ody size variation across multiple insular populations of 16 36 16 17 reptile species coAoccurring in the same archipelago and tested which island 37 18 19 characteristics primarilyFor drive Peer 0odyAsize evolution Review, what the common patterns are, 38 20 21 and whether coAoccurring species respond similarly to insular conditions. 39 22 23 24 40 25 26 Location Aegean Sea islands. 41 27 28 42 29 30 Ti e period 1984A2016. 43 31 32 44 33 34 Major taxa studied Reptiles. 45 35 36 37 46 38 39 Methods Be com0ined field work, museum measurements, and a comprehensive 47 40 41 literature survey to collect data on nearly 10,000 individuals representing eight lizard 48 42 43 and eight snake species across 233 islands. Be also ,uantified a large array of 49 44 45 predictors to directly assess the effects of island area, isolation (0oth spatial and 50 46 47 temporal), predation and interAspecific competition on 0ody size evolution. Be used 51 48 49 50 linear models and metaAanalyses to determine which predictors are informative for all 52 51 52 reptiles, for lizards and snakes separately, and for each single species. 53 53 54 54 55 56 57 58 3 59 60 Page 9 of 106 Global Ecology and Biogeography 1 2 3 Results 7ody size varies with different predictors across the species we studied, and 55 4 5 patterns differ within families and 0etween lizards and snakes. Bhile each predictor 56 6 7 influenced 0ody size in at least one species, no general trend was recovered. As a 57 8 9 group, lizards are hardly affected 0y any of the predictors we tested, whereas snake 58 10 11 size generally increases with area, competitor and predator richness, and decreases 12 59 13 14 with isolation. 60 15 16 61 17 18 Main conclusions Co factor emerges as a predominant driver of Aegean reptile sizes. 62 19 For Peer Review 20 This contradicts theories of general 0odyAsize evolutionary tra4ectories on islands. Be 63 21 22 conclude that overarching generalizations overAsimplify patterns and processes of 64 23 24 reptile 0odyAsize evolution on islands. Instead, speciesD autecology and island 65 25 26 27 particularities interact to drive the course of size evolution. 66 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 4 59 60 Global Ecology and Biogeography Page 10 of 106 1 2 3 INTRO./CTION 67 4 5 6 Identifying general pathways in the evolution of 0ody size variation among insular 68 7 8 populations has received decades of attention, and multiple patterns and drivers have 69 9 10 11 0een proposed and contradicted. Trait evolution on islands is often perceived as 70 12 13 strong, predicta0le and consistent across taxa (.an .alen, 1933; Eomolino, 2005; 71 14 15 KFhler et al., 2008). The most de0ated pattern is the Gisland ruleH, suggesting insular 72 16 17 animals tend to evolve a medium 0ody size (.an .alen, 1933; Eomolino, 2005; 73 18 19 2aubry I Svenning,For 2016 cf. Peer 6eiri, 2003; ReviewItescu et al., 2014; Eeisler I Binkler, 74 20 21 2015Insular faunas are generally depauperate, 0ecoming speciesApoor as islands 75 22 23 24 0ecome smaller and more isolated (e.g., arlington, 1953; 6acArthur I Bilson, 76 25 26 1963). Therefore, insular animals are thought to experience relaxed interspecific 77 27 28 competition and predation pressures, which, in turn, promote higher population 78 29 30 densities and conse,uently stronger intraspecific competition (6elton, 1982). 79 31 32 Together with resource limitation, these ecological processes are commonly thought 80 33 34 to drive 0ody size evolution on islands (Case, 1938; 6elton, 1982; Eomolino, 2005). 81 35 36 37 Heaney (1938) suggested that the effect of each of these factors changes with the size 82 38 39 of the focal island and animal.
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