Research Article: Time and tide: Seasonal, diel and tidal rhythms in Wadden Sea Harbour porpoises (Phocoena phocoena)

Date Published: March 20, 2019

Publisher: Public Library of Science

Author(s): Beate Zein, Benno Woelfing, Michael Dähne, Tobias Schaffeld, Stefan Ludwig, Jacob Hansen Rye, Johannes Baltzer, Andreas Ruser, Ursula Siebert, Songhai Li.


Odontocetes have evolved a rich diversity of prey- and habitat-specific foraging strategies, which allows them to feed opportunistically on locally and temporally abundant prey. While habitat-specific foraging strategies have been documented for some odontocete species, this is less known for the harbour porpoise (Phocoena phocoena). We collected multiple years of acoustic data using echolocation click loggers to analyse porpoise occurrence and buzzing behaviour, indicating feeding, in the German Wadden Sea (North Sea). Seasonal, diel and tidal effects were studied using Generalised Estimating Equations (GEE-GAMs). Locally season, time of day and tidal time significantly influenced the probability of porpoise detections and detection of foraging sequences (buzzes). Hunting strategies, and therefore frequency of buzzes, were likely affected by prey distribution and large differences between POD locations indicated that porpoises used highly specific behaviour adapted to tide and time of day to efficiently feed on the available prey. Strong seasonal and spatial variation in diel and tidal effects underline the importance of long-term observations. Studies on porpoise behaviour are often based on short-term observations and might rather reflect a seasonal than a general pattern. The results of this study show clearly that significant changes in porpoise behaviour can be found in short and long-term observations. Here some features are based on short term determinants and others are stable over years and care should be taken about drawing general conclusions based on local patterns. Highly variable spatio-temporal patterns indicate a high flexibility of porpoises in a highly variable environment and address a challenge for complex conservation management plans.

Partial Text

In a fast-changing environment with variable prey abundance, odontocetes have evolved highly adaptive foraging strategies. For an optimal foraging success, predators need to locate prey, which might be in localised patches due to biotic or abiotic factors [1,2]. While habitat-specific foraging strategies have been documented for some odontocete species, this is less known for the harbour porpoise (Phocoena phocoena). The dependency of predators’ distribution to prey availability should be especially high for harbour porpoises because they have a high energy demand due to their small size and their life in temperate waters [3–5].

At all POD stations echolocation clicks of harbour porpoises were regularly recorded and recording effort was 9888 days from 17 different positions. At four POD sites T11, T13, T15 and T18 recorded data was less than 3 months and they were therefore excluded from further analyses. Results of the GEE-GAM models indicate highly significant effects of day of the year, tide and time of the day on harbour porpoise detections at most POD sites (Table 1).

We analysed long term acoustic data of harbour porpoises in the German Wadden Sea and found strong and statistically significant seasonal, tidal and daylight effects on detections and buzzing behaviour at the different deployment positions over the time period from 2003 to 2016.




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