Modelling movements of Saimaa ringed seals using an individual-based approach

Movement is a fundamental element of animal behaviour, and it is the primary way through which animals respond to environmental changes. Therefore, understanding the drivers of individual movement is essential for species conservation. The endangered Saimaa ringed seal (Phoca hispida saimensis) lives land-locked in Lake Saimaa and is affected by various anthropogenic factors. Telemetry studies provide critical information but are insufficient to identify the mechanisms responsible for particular movement patterns. To better understand these mechanisms and to predict how changed movement patterns could influence the subspecies’ spatial ecology, we developed an individual-based movement model. We divided the seals’ daily routines into foraging and resting and explored how well the model captured observed home ranges and other movement metrics. Here we present the model, its predictions of home ranges and its sensitivity to model assumptions and parameter uncertainty. We used movement data from one individual to calibrate the model, but this resulted in poor predictions of home range sizes of five seals used for validation. This suggests that differences in movement paths not only reflect different landscape configurations but also differences among the individuals’ state and personalities. Therefore, we separately re-calibrated the model to data from five individuals, reproducing their home ranges, habitat use and movement paths more accurately. Although ignoring many aspects of seal behaviour, the model can be applied as a tool to guide further data collection and analysis, study seal ecology, and evaluate the efficacy of various conservation measures.

[1]  T. Alerstam,et al.  Satellite tracking of Swedish Ospreys Pandion haliaetus: autumn migration routes and orientation , 2001 .

[2]  Mervi Kunnasranta Behavioural biology of two ringed seal (Phoca hispida) subspecies in the large European lakes Saimaa and Ladoga , 2001 .

[3]  T. Sipilä,et al.  Long-term effects of land use on perinatal mortality in the Endangered Saimaa ringed seal population , 2017 .

[4]  A. Valtonen,et al.  Haulout patterns of Saimaa ringed seals and their response to boat traffic during the moulting season , 2013 .

[5]  W. Richardson,et al.  Factors influencing local abundance and haulout behaviour of ringed seals (Phoca hispida) on landfast ice of the Alaskan Beaufort Sea , 2002 .

[6]  Andreas Focks,et al.  Towards better modelling and decision support: Documenting model development, testing, and analysis using TRACE , 2014 .

[7]  Uta Berger,et al.  Pattern-Oriented Modeling of Agent-Based Complex Systems: Lessons from Ecology , 2005, Science.

[8]  Steven F Railsback,et al.  Pattern-oriented modelling: a ‘multi-scope’ for predictive systems ecology , 2012, Philosophical Transactions of the Royal Society B: Biological Sciences.

[9]  Spatial and temporal variation in genetic diversity of an endangered freshwater seal , 2012, Conservation Genetics.

[10]  Daniel Fortin,et al.  Adaptive models for large herbivore movements in heterogeneous landscapes , 2005, Landscape Ecology.

[11]  Steven F. Railsback,et al.  Agent-Based and Individual-Based Modeling: A Practical Introduction , 2011 .

[12]  M. Niemi Behavioural ecology of the Saimaa ringed seal : implications for conservation , 2013 .

[13]  Norman A. Slade,et al.  Relating Body Size to the Rate of Home Range Use in Mammals , 1988 .

[14]  Dirk Eisinger,et al.  Spatial pattern formation facilitates eradication of infectious diseases , 2008, The Journal of applied ecology.

[15]  C. F. Estades,et al.  Individual-based modeling as a decision tool for the conservation of the endangered huemul deer (Hippocamelus bisulcus) in southern Chile , 2012 .

[16]  David A. Bennett,et al.  Modelling adaptive, spatially aware, and mobile agents: Elk migration in Yellowstone , 2006, Int. J. Geogr. Inf. Sci..

[17]  Estimating the demographic effective population size of the Saimaa ringed seal (Phoca hispida saimensis Nordq.) , 1998 .

[18]  Peter Leimgruber,et al.  Non‐Markovian maximum likelihood estimation of autocorrelated movement processes , 2014 .

[19]  R. Gramacy,et al.  Categorical Inputs, Sensitivity Analysis, Optimization and Importance Tempering with tgp Version 2, an R Package for Treed Gaussian Process Models , 2010 .

[20]  J. P. Ball,et al.  Partial migration by large ungulates: characteristics of seasonal moose Alces alces ranges in northern Sweden , 2001, Wildlife Biology.

[21]  Miina Auttila The endangered Saimaa ringed seal in a changing climate : challenges for conservation and monitoring , 2015 .

[22]  O. Liberg,et al.  Multistage, Long-Range Natal Dispersal by a Global Positioning System–Collared Scandinavian Wolf , 2007 .

[23]  J. Teilmann,et al.  How a simple adaptive foraging strategy can lead to emergent home ranges and increased food intake , 2013 .

[24]  J. Palo,et al.  Demographic histories and genetic diversities of Fennoscandian marine and landlocked ringed seal subspecies , 2014, Ecology and evolution.

[25]  J. Palo,et al.  Causes and consequences of fine-scale population structure in a critically endangered freshwater seal , 2014, BMC Ecology.

[26]  Marc Deconchat,et al.  Modelling the overwintering strategy of a beneficial insect in a heterogeneous landscape using a multi-agent system , 2007 .

[27]  E. Hämäläinen,et al.  DIVING BEHAVIOR OF THE SAIMAA RINGED SEAL (PHOCA HISPIDA SAIMENSIS NORDQ.) , 1995 .

[28]  B. McConnell,et al.  Grey and harbour seals in France : distribution at sea, connectivity and trends in abundance at haulout sites , 2017 .

[29]  A. Harestad,et al.  Home Range and Body Weight‐‐A Reevaluation , 1979 .

[30]  T. Sipilä Conservation biology of Saimaa ringed seal (Phoca hispida saimensis) with reference to other European seal populations , 2003 .

[31]  H. Thulke,et al.  The strength of 70%: revision of a standard threshold of rabies control. , 2008, Developments in biologicals.

[32]  Hauke Reuter,et al.  Emerging properties on the individual level: modelling the reproduction phase of the European robin , 1999 .

[33]  J. Gareth Polhill,et al.  The ODD protocol: A review and first update , 2010, Ecological Modelling.

[34]  H. Hyvärinen,et al.  The diet of the Saimaa ringed seal Phoca hispida saimensis , 1999 .

[35]  T. Sipilä,et al.  Morphometrics, body condition, and growth of the ringed seal (Pusa hispida saimensis) in Lake Saimaa: Implications for conservation , 2016 .

[36]  C. Linard,et al.  A multi-agent simulation to assess the risk of malaria re-emergence in southern France , 2008 .

[37]  P. Boveng,et al.  Seasonal home ranges and fidelity to breeding sites among ringed seals , 2010, Polar Biology.

[38]  Kristen M. Hart,et al.  Satellite telemetry of marine megavertebrates: the coming of age of an experimental science , 2009 .

[39]  Lucas N Joppa,et al.  Understanding movement data and movement processes: current and emerging directions. , 2008, Ecology letters.

[40]  Juan M. Morales,et al.  EXTRACTING MORE OUT OF RELOCATION DATA: BUILDING MOVEMENT MODELS AS MIXTURES OF RANDOM WALKS , 2004 .

[41]  Pernille Thorbek,et al.  Linking pesticide exposure and spatial dynamics: An individual-based model of wood mouse (Apodemus sylvaticus) populations in agricultural landscapes , 2013 .

[42]  Patrice Boily,et al.  Theoretical heat flux in water and habitat selection of phocid seals and beluga whales during the annual molt , 1995 .

[43]  Paola Annoni,et al.  Variance based sensitivity analysis of model output. Design and estimator for the total sensitivity index , 2010, Comput. Phys. Commun..

[44]  K. Kovacs,et al.  Spring haul-out behavior of ringed seals (Pusa hispida) in Kongsfjorden, Svalbard , 2006 .

[45]  Deborah Austin,et al.  Intraspecific variation in movement patterns: modeling individual behaviour in a large marine predator , 2004 .

[46]  Broder Breckling,et al.  A spatiotemporal individual-based fish model to investigate emergent properties at the organismal and the population level , 2005 .

[47]  Jonas Hedin,et al.  Using radio telemetry to study dispersal of the beetle Osmoderma eremita, an inhabitant of tree hollows , 2002 .

[48]  W. Bowen,et al.  Sex differences in diving at multiple temporal scales in a size‐dimorphic capital breeder , 2003 .

[49]  Darcy R. Visscher,et al.  Memory keeps you at home: a mechanistic model for home range emergence , 2009 .

[50]  Miina Auttila,et al.  Estimating and Mitigating Perinatal Mortality in the Endangered Saimaa Ringed Seal (Phoca hispida saimensis) in a Changing Climate , 2014 .

[51]  Uta Berger,et al.  Structural realism, emergence, and predictions in next-generation ecological modelling: Synthesis from a special issue , 2016 .

[52]  E. Revilla,et al.  Trends and missing parts in the study of movement ecology , 2008, Proceedings of the National Academy of Sciences.

[53]  T. Selhorst,et al.  Emergency vaccination of rabies under limited resources – combating or containing? , 2005, BMC infectious diseases.

[54]  Volker Grimm,et al.  Ecological models supporting environmental decision making: a strategy for the future. , 2010, Trends in ecology & evolution.

[55]  A. Valtonen,et al.  The Effects of Sex and Season on Home Range in European Hedgehogs at the Northern Edge of the Species Range , 2013 .

[56]  B. Nolet,et al.  Foraging behaviour and fuel accumulation of capital breeders during spring migration as derived from a combination of satellite- and ground-based observations. , 2016 .

[57]  Volker Grimm,et al.  Merging validation and evaluation of ecological models to ‘evaludation’: A review of terminology and a practical approach , 2014 .

[58]  W T Sobol,et al.  Analysis of variance for 'component stripping' decomposition of multiexponential curves. , 1993, Computer methods and programs in biomedicine.

[59]  Jacqueline L. Frair,et al.  Building the bridge between animal movement and population dynamics , 2010, Philosophical Transactions of the Royal Society B: Biological Sciences.

[60]  Meeri Koivuniemi,et al.  Photo-ID as a tool for studying and monitoring the endangered Saimaa ringed seal , 2016 .

[61]  Steven F. Railsback,et al.  Movement rules for individual-based models of stream fish , 1999 .

[62]  M. Niemi,et al.  Home range, survival, and dispersal of endangered Saimaa ringed seal pups: Implications for conservation , 2013 .

[63]  Valery E. Forbes,et al.  The Effects of Landscape Modifications on the Long-Term Persistence of Animal Populations , 2010, PloS one.

[64]  M. D. McKay,et al.  A comparison of three methods for selecting values of input variables in the analysis of output from a computer code , 2000 .

[65]  Birgit Müller,et al.  A standard protocol for describing individual-based and agent-based models , 2006 .

[66]  T. Sipilä,et al.  Comparative use of vendace by humans and Saimaa ringed seal in Lake Pihlajavesi, Finland , 2005 .

[67]  M. Niemi,et al.  Diet composition and seasonal feeding patterns of a freshwater ringed seal (Pusa hispida saimensis) , 2015 .

[68]  E. Revilla,et al.  A movement ecology paradigm for unifying organismal movement research , 2008, Proceedings of the National Academy of Sciences.

[69]  W. Richardson,et al.  Effects of an offshore oil development on local abundance and distribution of ringed seals (Phoca hispida) of the Alaskan Beaufort Sea , 2005 .

[70]  L. Sundström,et al.  Movement and home range in relation to dominance; a telemetry study on brown trout Salmo trutta , 2007 .

[71]  Carol E. Sparling,et al.  Seals like it hot: changes in surface temperature of harbour seals (Phoca vitulina) from late pregnancy to moult , 2012 .

[72]  Steven W. Buskirk,et al.  HOME RANGE, TIME, AND BODY SIZE IN MAMMALS' , 1986 .

[73]  Robert B. Gramacy,et al.  tgp: An R Package for Bayesian Nonstationary, Semiparametric Nonlinear Regression and Design by Treed Gaussian Process Models , 2007 .

[74]  H. Hyvärinen,et al.  Dive types and circadian behaviour patterns of Saimaa ringed sealsPhoca hispida saimensis during the open-water season , 2002, Acta Theriologica.

[75]  Donald L. DeAngelis,et al.  Spatially Explicit Modeling in Ecology: A Review , 2016, Ecosystems.

[76]  M. Niemi,et al.  Movement data and their application for assessing the current distribution and conservation needs of the endangered Saimaa ringed seal , 2012 .

[77]  J. Calabrese,et al.  The Correlated Random Walk and the Rise of Movement Ecology , 2014 .

[78]  R Core Team,et al.  R: A language and environment for statistical computing. , 2014 .

[79]  Patrick A. Zollner,et al.  SEARCH: Spatially Explicit Animal Response to Composition of Habitat , 2013, PloS one.

[80]  E. Hämäläinen,et al.  Movements and use of haul-out sites of radio-tagged Saimaa ringed seal (Phoca hispida saimensis Nordq.) during the open-water season , 2002 .

[81]  Simon J. Goodman,et al.  Global threats to pinnipeds , 2012 .

[82]  R. Alford,et al.  Nomadic movement in tropical toads , 2002 .

[83]  M. Jansen Analysis of variance designs for model output , 1999 .

[84]  D. DeAngelis,et al.  Individual-based models in ecology after four decades , 2014, F1000prime reports.

[85]  H. Kokko,et al.  BACKCASTING POPULATION SIZES OF RINGED AND GREY SEALS IN THE BALTIC AND LAKE SAIMAA DURING THE 20TH CENTURY , 1999 .

[86]  C H Fleming,et al.  Rigorous home range estimation with movement data: a new autocorrelated kernel density estimator. , 2015, Ecology.

[87]  Volker Grimm,et al.  Using pattern-oriented modeling for revealing hidden information: a key for reconciling ecological theory and application , 2003 .

[88]  Thorsten Wiegand,et al.  Dealing with Uncertainty in Spatially Explicit Population Models , 2004, Biodiversity & Conservation.

[89]  J. Teilmann,et al.  Habitat Use of Ringed Seals ( Phoca hispida ) in the North Water Area (North Baffin Bay) , 2004 .

[90]  Marja Niemi,et al.  Identifying foraging habitats of Baltic ringed seals using movement data , 2015, Movement ecology.