Scaling Marine Fish Movement Behavior from Individuals to Populations

This is a repository copy of Scaling marine fish movement behavior from individuals to populations. White Rose Research Online URL for this paper: http://eprints.whiterose.ac.uk/131598/ Version: Published Version Article: Griffiths, C.A., Patterson, T.A., Blanchard, J. et al. (4 more authors) (2018) Scaling marine fish movement behavior from individuals to populations. Ecology and Evolution, 8 (14). pp. 7031-7043. ISSN 2045-7758 https://doi.org/10.1002/ece3.4223 Reuse This article is distributed under the terms of the Creative Commons Attribution (CC BY) licence. This licence allows you to distribute, remix, tweak, and build upon the work, even commercially, as long as you credit the authors for the original work. More information and the full terms of the licence here: https://creativecommons.org/licenses/ 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/ Received November | Revised February | Accepted March DOI: 10.1002/ece3.4223 ORIGINAL RESEARCH Scaling marine fish movement behavior from individuals to populations Christopher A. Griffiths1,2,3 |Toby A Patterson4|Juia L Banchard2 | David A. Righton3 |Serena R Wright3 |Jon W Pitchford5|Pau G Backwe1 1Schoo of Mathematics and Statistics University of Sheffied Sheffied Abstract UK Understanding how where and when animas move is a centra probem in marine 2 Institute for Marine and Antarctic ecoogy and conservation Key to improving our knowedge about what drives anima Studies University of Tasmania Hobart TAS Austraia movement is the rising depoyment of teemetry devices on a range of free roaming 3Centre for Environment Fisheries and species An increasingy popuar way of gaining meaningfu inference from an ani Aquacuture Science Lowestoft Laboratory mas recorded movements is the appication of hidden Markov modes HMMs Lowestoft UK 4CSIRO Marine and Atmospheric Research which aow for the identification of atent behaviora states in the movement paths Hobart TAS Austraia of individuas However the use of HMMs to expore the popuation eve conse 5 Department of Bioogy University of York quences of movement is often imited by mode compexity and insufficient sampe York UK sizes Here we introduce an aternative approach to current practices and provide Correspondence evidence of how the incusion of prior information in mode structure can simpify the Christopher A Griffiths Schoo of Mathematics and Statistics University of appication of HMMs to mutipe anima movement paths with two cear benefits a Sheffied Sheffied UK consistent state aocation and b increases in effective sampe size To demonstrate Emai cagriffithssheffiedacuk the utiity of our approach we appy HMMs and adapted HMMs to over muti Funding information variate movement paths consisting of conditionay dependent daiy horizonta and Natura Environment Research Counci GrantAward Number NEL vertica movements in two species of demersa fish Atantic cod Gadus morhua; n and European paice Pleuronectes platessa; n We identify atent states corresponding to two main underying behaviors resident and migrating As our ana ysis considers a reativey arge sampe size and states are aocated consistenty we use coective mode output to investigate state dependent spatiotempora trends at the individua and popuation eves In particuar we show how both species shift their movement behaviors on a seasona basis and demonstrate popuation space use patterns that are consistent with previous individua eve studies Tagging studies are increasingy being used to inform stock assessment modes spatia management strategies and monitoring of marine fish popuations Our approach provides a prom ising way of adding vaue to tagging studies because inferences about movement behavior can be gained from a arger proportion of datasets making tagging studies more reevant to management and more cost effective This is an open access artice under the terms of the Creative Commons Attribution License which permits use distribution and reproduction in any medium provided the origina work is propery cited The Authors Ecology and Evolution pubished by John Wiey Sons Ltd Ecology and Evolution. 2018;8:7031–7043. www.ecolevol.org | 7031 | GRIFFITHS ET AL. KEYWORDS Atantic cod data storage tags European paice hidden Markov modeing movement behavior popuationeve patterns priors |INTRODUCTION Kranstauber et a it woud appear that the most viabe route toward robust popuation eve inferences is approaches that The spatia management of the marine word requires in depth in make the most of the tagging data we aready have formation about how animas move when they move and where Among the many methodoogica deveopments that utiize they move to Key to increasing our understanding of species space movement data to answer ecoogica questions hidden Markov use movement patterns and how individuas interact with the en modes HMMs and hidden semiMarkov modes have taken cen vironment they inhabit is the rising depoyment of sma and rei ter stage eg DeRuiter et a McKear Langrock Waters abe data oggers and transmitters on free roaming marine animas Keser Micheot Langrock Patterson Towner et Costa Breed Robinson Hays et a Hussey et a a Favored because they match our initiative understanding Capabe of recording a range of movement metrics incuding that movement is governed by switches in an animas motivation horizonta and vertica movement aongside basic environmenta in Patterson et a HMMs provide a computationay efficient formation such as water temperature sainity and ambient dayight means of objectivey cassifying movement into discrete states with these devices have revoutionized our understanding of fundamen different statistica properties indicating differences in underying ta ecoogy Hussey et a documented oceanwide dispersa behavior Langrock et a events Bock et a highighted areas that are essentia for HMMs have been fitted to mutipe individua pathways simu species surviva Raymond et a and even aowed us to test taneousy in both the frequentist Langrock et a McKear the effectiveness of current conservation poicies Pittman et a et a and Bayesian statistica paradigms Jonsen Scott et a McCintock et a However these approaches are typicay One of the main motivations for anima borne teemetry studies impemented by speciaist statisticians and require the couping of is that by understanding individua movement behavior we might HMM and hierarchica structures producing a hierarchica Hidden infer the popuation species and community eve consequences Markov mode HHMM The aternative is the use of HMMs or other of movement Bock et a Hinde et a Raymond et a state space approaches that fit on an individua by individua basis Wakefied et a This is especiay true in marine sys Jonsen Myers James Micheot et a This atter tems as individua observations provide our ony insight into the more frequenty used approach has its advantages the most notabe otherwise unobservabe Achieving this scaing of inference from being an ease of use for statisticians and bioogists aike Fitting per individua movement patterns to popuation dynamics requires two individua aso has its disadvantages The first is that it requires indi important components The first is an adequate sampe size num vidua movement paths that are suitaby data rich to achieve mode ber of individuas to address the ecoogica question of interest convergence imposing even stricter restrictions on sampe size The Hebbewhite Haydon and second a statistica means by second is a distinct ack of any forma process by which state one in which we gain meaningfu inference at the individua and popua anima A is ensured consistency with state one in anima B This ack tion eve from a finite sampe of individuas Jonsen Langrock of consistency means that estimated parameters can readiy inform et a McCintock Russe Matthiopouos King individua eve movement studies but wi resut in tricky interspe The issue of sampe size has been extensivey discussed espe cific and intraspecific comparisons imiting a researchers abiity to ciay when considering how movement studies can inform marine ask post hoc popuation eve questions of their data conservation and spatia management Hebbewhite Haydon Our objective is to introduce an aternative framework that uses McGowan et a Nguyen et a Ogburn et a HMMs to overcome the described imitations of individuay fitted Tags can be expensive McGowan et a are iabe to HMMs whie maintaining their heraded ease of use advantages Our occasiona faiure or oss and often resut in individua pathways approach combines an N state HMM and severa hierarchica struc that are data poor or have a ow number of observations As a re tures but bypasses the need to integrate over the random effects sut meeting the minimum sampe size of individuas when as in HHMMs Langrock et a by using information we gain making simpe statistica comparisons between popuations is un from our data rich pathways as a priori approximations of each states common Hebbewhite Haydon with even greater num movement parameters Doing so not ony aows us to achieve co bers needed when testing for the effects of age sex and species herent individua and popuation eve state cassification but aso identity Lindberg Waker In the absence of a coabora ensures that we maximize our sampe size by gaining meaningfu in tive effort across mutipe institutions Bock

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