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Diel and Seasonal Patterns in Acoustic Presence and Foraging Behaviour of Free-Ranging Harbour Porpoises

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Diel and Seasonal Patterns in Acoustic Presence and Foraging Behaviour of Free-Ranging Harbour Porpoises Vol. 547: 257–272, 2016 MARINE ECOLOGY PROGRESS SERIES Published April 7 doi: 10.3354/meps11627 Mar Ecol Prog Ser OPENPEN ACCESSCCESS Diel and seasonal patterns in acoustic presence and foraging behaviour of free-ranging harbour porpoises Tobias Schaffeld1,2,3,*, Stefan Bräger1,4, Anja Gallus1, Michael Dähne1,3, Kathrin Krügel1, Anne Herrmann1, Martin Jabbusch1, Thomas Ruf5, Ursula K. Verfuß1,6, Harald Benke1, Jens C. Koblitz1,7 1German Oceanographic Museum, Katharinenberg, 18439 Stralsund, Germany 2University of Natural Resources and Life Sciences, 1180 Vienna, Austria 3Institute for Terrestrial and Aquatic Wildlife Research, University of Veterinary Medicine Hannover, Foundation, 25761 Büsum, Germany 4International Seabed Authority, Kingston, Jamaica 5Research Institute of Wildlife Ecology, University of Veterinary Medicine Vienna, 1160 Vienna, Austria 6SMRU Consulting, Scottish Oceans Institute, St. Andrews KY16 9SR, UK 7BioAcoustics Network, 18439 Stralsund, Germany ABSTRACT: A temporal and geographical analysis of echolocation activity can give insights into the behaviour of free-ranging harbour porpoises Phocoena phocoena. Seasonal and diel patterns in the presence and foraging activity of harbour porpoises were investigated based on a year-long passive acoustic monitoring data set recorded at 5 sites in the western Baltic Sea. Diel patterns in detection rates were found at 4 sites. A year-round rhythm in presence, however, was found at only 1 station, whereas the other 3 stations showed diel rhythms for 2 to 3 seasons. Three of the sites showed diel patterns in foraging sequences on a seasonal level, but no station showed such patterns for the complete year of investigation. Both diurnal and nocturnal patterns in harbour porpoise detections were observed, indicating that diel rhythmic behaviour is more complex than previously reported. In contrast, foraging behaviour showed only nocturnal rhythms. Owing to the limitations in passive acoustic monitoring, all categorized foraging sequences are a minimum esti- mate. Therefore, classified foraging sequences are most likely pelagic foraging, while bottom grubbing could have been missed. Differences in the occurrence of foraging sequences between station, season and time of day lead to the assumption that the long-term echolocation diel pat- terns of porpoises strongly depend on the temporal changes in food availability and composition within a certain habitat. Echolocation behaviour of foraging porpoises is strongly influenced by seasonally available prey resources, which require adaptive foraging strategies. Therefore, owing to seasonal variations, analyses of diel patterns need to be conducted over sufficiently long time periods and large geographic scales to allow generalized interpretation of the findings. Conse- quently, no general conclusion regarding diel rhythms in harbour porpoise echolocation was found. We hypothesize that porpoises in the study area alternate between foraging on benthic prey in shallow waters at daytime and in the pelagic in deeper waters at night. KEY WORDS: Cosinor analysis · Diel rhythms · Predation · Phocoena phocoena INTRODUCTION ronment. To do so they emit series of pulsed, high- frequency clicks to receive information on their Like all toothed whales (Odontoceti), harbour surroundings, including possible prey items, from the porpoises have developed an active acoustic sense, returning echo (Au 1993). Harbour porpoises are echolocation, to acquire information about their envi- known to echolocate almost constantly, emitting © The authors 2016. Open Access under Creative Commons by *Corresponding author: [email protected] Attribution Licence. Use, distribution and reproduction are un - restricted. Authors and original publication must be credited. Publisher: Inter-Research · www.int-res.com 258 Mar Ecol Prog Ser 547: 257–272, 2016 echolocation clicks on average every 12.3 s (Aka- al. 2014). The other population, inhabiting in the matsu et al. 2007). Maximum periods with no clicks, Baltic Proper, has been classified by the International or clicks with amplitudes below recording threshold, Union for Conservation of Nature (IUCN) as Critically ranging from 99 to 1300 s were found for 3 free-rang- Endangered (Hammond et al. 2008). P. phocoena ing porpoises equip ped with acoustic data loggers have high energy demands due to their small body (Linnenschmidt et al. 2013). Although diel variations size and thermoregulatory costs caused by the tem- and rare periods of silence occurred, Linnenschmidt perate waters they inhabit (Koopman 1998). Their et al. (2013) concluded that harbour porpoises echo - small size does not allow for the storage of energy locate almost continuously. Verfuß et al. (2009) reserves, thus leading to a high dependence on the showed that harbour porpoises use echolocation for year-round availability of food resources (Santos et al. orientation and foraging during daylight in relatively 2004). Therefore, seasonal shifts in porpoise distribu- clear water conditions, independent of their ability to tion (Verfuß et al. 2007, Benke et al. 2014) may be ex- use eyesight or not (e.g. when blindfolded). Porpoises plained best by changes in prey densities (Sveegaard thus appear to use their echolocation independently et al. 2012) and availa bility. For example, spring of light conditions. They emit specific click patterns migrations of harbour porpoises into the Baltic Sea for orientation and prey capture (Verfuß et al. 2005, are known to track migrating herring to the spawning DeRuiter et al. 2009, Verfuß et al. 2009) and also for grounds (Koschinski 2002 and citations therein), and intraspecific communication (reviewed in Koschinski harbour porpoise occurrence is positively correlated et al. 2008, Clausen et al. 2011). Click patterns thus with herring densities (Sveegaard et al. 2012). Diet allow the discrimination of different behavioural cat- regularly consists of the most abundant prey items egories such as communication (Clausen et al. 2011), (Christensen & Richardson 2008), which is typical foraging (Koschinski et al. 2008, DeRuiter et al. 2009, for a generalist and opportunistic feeder. According Verfuß et al. 2009, Linnenschmidt et al. 2013), or the to stomach content analysis, the most abundant fish use of local objects to lock on and approach during species in the local harbour porpoise diet are herring the navigation from one point to another (Verfuß et al. Clupea harengus, cod Gadus morhua and gobies 2005). According to Verfuß et al. (2009) foraging se- (Gobiidae) (Lick 1991, Sveegaard et al. 2012). In the quences can be separated into different phases by period between April and October, cod is the most changes in echolocation behaviour, leading to altered important prey item, whereas in the period from inter-click intervals (ICI). The pattern in ICI during November to March, cod and herring are of equal the search phase seems to be driven by a range lock- importance for harbour porpoises (Sveegaard et al ing to objects until a potential target can be detected. 2012). These species vary in size, caloric content, life In captivity, the initial part of the approach phase is cycle, water depth preference and migratory poten- characterised by relatively stable ICI of around 50 ms tial, and the availability of preferred fish species (Verfuß et al. 2005). The terminal part of the approach varies throughout the year (Sveegaard et al. 2012). A phase is marked by a sudden and rapid shortening of daily consumption of 3.5 to 5.5 kg has been reported ICI to levels of below 10 ms (DeRuiter et al. 2009, Ver- for porpoises in Danish waters (Lockyer 2007). Fur- fuß et al. 2009). The ICI can reach a minimum of thermore, there appear to be differences in the diet around 1.5 ms and remains constant at a distance of composition between harbour porpoises of different less than 1 m to the prey (Verfuß et al. 2009). In accor- ages. For instance, gobies and shrimps are more dance with patterns in echolocation sequences of in- commonly preyed upon by sub-adults (Lick 1991, sectivorous bats during the final stage of a foraging Santos et al. 2004). These dietary changes could pos- attempt, click sequences with a short and stable ICI of sibly be a result of different foraging strategies, de- below 10 ms are called ‘buzzes’ (Miller et al. 1995, pending on available prey items throughout the year, DeRuiter et al. 2009, Verfuß et al. 2009, Madsen & and depending on life stage to maximise energy Surlykke 2013). A rapid increase in ICI marks the end intake while minimising energy expenditure during of the terminal part of the approach phase. At least 2 foraging. genetically and morphologically distinct populations Just as Mysticetes (baleen whales) show rhythms of harbour porpoise Phocoena phocoena have been in their vocal behaviour (e.g. Risch et al. 2013), diel identified in the waters of the Baltic Sea (Wiemann et and seasonal changes in acoustic (and foraging) al. 2010, Galatius et al. 2012). One population inhab- behaviour are known also for various odontocete iting the southern part of the Kattegat, the Belt Seas, (toothed whale) species. Pacific white-sided dolphins the Sound and the western Baltic has an abundance Lagenorhynchus obliquidens change between 2 of 40 475 (CI 25 614 to 65 041) individuals (Viquerat et different click bout patterns from day to night, which Schaffeld et al.: Behavioural patterns of free-ranging harbour porpoises 259 is assumed to be related to prey species (Soldevilla For the results presented here,
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