Understanding narwhal diving behaviour using Hidden Markov Models with dependent state distributions and long range dependence

Diving behaviour of narwhals is still largely unknown. We use Hidden Markov models (HMMs) to describe the diving behaviour of a narwhal and fit the models to a three-dimensional response vector of maximum dive depth, duration of dives and post-dive surface time of 8,609 dives measured in East Greenland over 83 days, an extraordinarily long and rich data set. Narwhal diving patterns have not been analysed like this before, but in studies of other whale species, response variables have been assumed independent. We extend the existing models to allow for dependence between state distributions, and show that the dependence has an impact on the conclusions drawn about the diving behaviour. We try several HMMs with 2, 3 or 4 states, and with independent and dependent log-normal and gamma distributions, respectively, and different covariates to characterize dive patterns. In particular, diurnal patterns in diving behaviour is inferred, by using periodic B-splines with boundary knots in 0 and 24 hours.

[1]  Russel D. Andrews,et al.  First Long-Term Behavioral Records from Cuvier’s Beaked Whales (Ziphius cavirostris) Reveal Record-Breaking Dives , 2014, PloS one.

[2]  Roland Langrock,et al.  Estimation and simulation of foraging trips in land-based marine predators. , 2016, Ecology.

[3]  Toby A Patterson,et al.  Classifying movement behaviour in relation to environmental conditions using hidden Markov models. , 2009, The Journal of animal ecology.

[4]  Mads Peter Heide-Jørgensen,et al.  A note on the diet of narwhals (Monodon monoceros) in Inglefield Bredning (NW Greenland) , 1994 .

[5]  Mark P. Johnson,et al.  Cheetahs of the deep sea: deep foraging sprints in short-finned pilot whales off Tenerife (Canary Islands). , 2008, The Journal of animal ecology.

[6]  Jr. G. Forney,et al.  The viterbi algorithm , 1973 .

[7]  P. A. P. Moran,et al.  Statistical inference with bivariate gamma distributions , 1969 .

[8]  P.,et al.  Encyclopedia of Marine Mammals , 2017 .

[9]  Roland Langrock,et al.  Flexible and practical modeling of animal telemetry data: hidden Markov models and extensions. , 2012, Ecology.

[10]  Susanne Ditlevsen,et al.  Spatial and temporal patterns of sound production in East Greenland narwhals , 2018, PloS one.

[11]  Kristin L. Laidre,et al.  WINTER FEEDING INTENSITY OF NARWHALS (MONODON MONOCEROS) , 2005 .

[12]  John Calambokidis,et al.  A multivariate mixed hidden Markov model for blue whale behaviour and responses to sound exposure , 2017 .

[13]  William A. Watkins,et al.  SPERM WHALES TAGGED WITH TRANSPONDERS AND TRACKED UNDERWATER BY SONAR , 1993 .

[14]  Roland Langrock,et al.  Modeling the Diving Behavior of Whales: A Latent-Variable Approach with Feedback and Semi-Markovian Components , 2014 .

[15]  M. P. Heide-Jørgensen Narwhal: Monodon monoceros , 2009 .

[16]  Christian Lydersen,et al.  Distribution of endemic cetaceans in relation to hydrocarbon development and commercial shipping in a warming Arctic , 2014 .

[17]  Susanna B Blackwell,et al.  Stomach temperature of narwhals (Monodon monoceros) during feeding events , 2014, Animal Biotelemetry.

[18]  Jennifer Pohle,et al.  Selecting the Number of States in Hidden Markov Models: Pragmatic Solutions Illustrated Using Animal Movement , 2017 .

[19]  Peter L. Tyack,et al.  Formal Comment on Schorr GS, Falcone EA, Moretti DJ, Andrews RD (2014) First Long-Term Behavioral Records from Cuvier’s Beaked Whales (Ziphius cavirostris) Reveal Record-Breaking Dives. PLoS ONE 9(3): e92633. doi:10.1371/journal.pone.0092633 , 2015, PloS one.

[20]  Terrie M. Williams,et al.  Extreme physiological adaptations as predictors of climate‐change sensitivity in the narwhal, Monodon monoceros , 2011 .

[21]  David R. Anderson,et al.  Model Selection and Multimodel Inference , 2003 .

[22]  Andrew J. Read,et al.  Hidden Markov models reveal complexity in the diving behaviour of short-finned pilot whales , 2017, Scientific Reports.

[23]  Dirk Eddelbuettel,et al.  Rcpp: Seamless R and C++ Integration , 2011 .

[24]  W. Zucchini,et al.  Hidden Markov Models for Time Series: An Introduction Using R , 2009 .

[25]  Mads Peter Heide-Jørgensen,et al.  The predictable narwhal: satellite tracking shows behavioural similarities between isolated subpopulations , 2015 .