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Acoustic Determination of Activity and Flipper Stroke Rate in Foraging Northern Fur Seal Females

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Acoustic Determination of Activity and Flipper Stroke Rate in Foraging Northern Fur Seal Females Vol. 4: 147–155, 2008 ENDANGERED SPECIES RESEARCH Printed January 2008 doi: 10.3354/esr00050 Endang Species Res Published online October 19, 2007 OPENPEN ACCESSCCESS THEME SECTION Acoustic determination of activity and flipper stroke rate in foraging northern fur seal females S. J. Insley1, 3,*, B. W. Robson2, 4, T. Yack3, R. R. Ream2, W.C. Burgess4 1University of California Santa Cruz, Institute of Marine Sciences, Long Marine Laboratory, 100 Shaffer Rd., Santa Cruz, California 95060, USA 2National Marine Mammal Laboratory, National Marine Fisheries Service, 7600 Sand Point Way, Seattle, Washington 98115, USA 3Hubbs-SeaWorld Research Institute, 2595 Ingraham Street, San Diego, California 92109, USA 4Greeneridge Sciences, 1411 Firestone Rd., Santa Barbara, California 93117, USA ABSTRACT: Foraging effort for lactating female otariid pinnipeds is largely a function of the energy expended swimming to a site and diving in search of prey. This is especially true for northern fur seal Callorhinus ursinus females, which predictably punctuate their suckling with 7 to 12 d foraging trips at sea, with swimming distances often exceeding 400 km. In the present study we tested a unique approach (flow noise from onboard acoustic dataloggers) to empirically measure swim effort in free ranging female northern fur seals, the first such field measurements on an otariid pinniped. We first measured behavioural activity budgets of seals from a combination of satellite telemetry, pressure (depth), and onboard acoustic data. From these data we were able to quantify the time spent in each of 4 mutually exclusive forms of behaviour: locomoting, diving, resting, and surface activity. Second, flipper stroke rates and stroke rate patterns were measured from the acoustic data for each seal dur- ing 3 dive types (i.e. locomoting, shallow and mid/deep dives) and during 3 dive parts (descent, bot- tom time and ascent). Although stroke rates during each of the 3 dive types were similar (ca. 0.5 Hz), they were distinct during the different parts of a dive. In each case, variation among individuals was significant. Stroke rate patterns were distinct for the different dive types and dive parts. Overall, in addition to applying a unique technique to measure foraging effort in a declining population, the results emphasize the importance of accounting for individual variation to obtain accurate estimates of foraging cost. KEY WORDS: Flipper stroke rate ⋅ Effort ⋅ Activity budget ⋅ Diving behaviour ⋅ Foraging effort ⋅ Marine mammal ⋅ Pinniped ⋅ Otariid ⋅ Fur seal Resale or republication not permitted without written consent of the publisher INTRODUCTION 1986). While gravity is the primary force behind the energetic cost of locomotion in terrestrial mammals While caring for their young, female pinnipeds in the (Cavagna et al 1977, Taylor et al 1980), in marine family Otariidae (i.e. sea lions and fur seals) are central mammals it is a combination of hydrostatic pressure, place foragers, alternating between suckling on land body drag, and buoyancy, all of which vary with depth and foraging at sea (Orians & Pearson 1979, Bowen et (Williams 2001). In the present paper we investigate a al. 2002). Foraging success during the period of lacta- novel method (flow noise combined with pressure and tion is therefore a key factor determining offspring sur- geo-location data) to measure the swimming effort of vival and the female’s reproductive success. Foraging lactating female northern fur seals Callorhinus ursinus effort is largely a function of the amount of energy during foraging trips at sea. expended while swimming, specifically locomoting to Reliable measurements of swimming effort in aqua- a site and diving in search of prey (Gentry & Kooyman tic mammals are difficult to obtain in nature (Boyd et *Email: [email protected] © Inter-Research 2008 · www.int-res.com 148 Endang Species Res 4: 147–155, 2008 al. 1995). Two essential components are (1) the amount al. 2002, Towel et al. 2006). The cause of the decline is of time spent in each aquatic activity and (2) estimates currently unknown; however, decreased food avail- of the energy expenditure for each. The first require- ability, whether due to anthropogenic or heterospecific ment can be satisfied with an accurate behavioural competition or climate change, is a principal suspect. If activity budget (Martin & Bateson 1993). Although food availability has decreased, the effort or cost to activity budgets are basic to an understanding of an find sufficient food may be increasing. Consequently, animal’s behavioural ecology, they are logistically dif- empirical measurement of foraging costs, particularly ficult and consequently rare for pelagic animals. for lactating females, is needed. Despite considerable recent advances in remote The goal of this study was to determine whether we instrumentation techniques on free ranging animals, could combine flow noise from onboard acoustic data- studies of pinniped pelagic activity are mostly re- loggers with pressure and geo-location data to empiri- stricted to location (using satellite transmitters, also cally measure swim effort in free ranging female referred to as platform terminal transmitters, PTTs) northern fur seals. To do so, we first produced behav- and diving behaviour (using time-depth recorders, ioural activity budgets from the combined dataset to TDRs; but see Bailleul et al. 2005 and Guinet et al. determine the time spent in each activity while at sea. 2005). We then measured flipper stroke rates and stroke rate A number of different techniques including swim patterns as a proxy for effort for each of the activities speed, heart rate and swimming stroke rate have been that involved swimming. The result is an empirical used to satisfy the second requirement, accurate esti- measurement of swim effort during different behav- mates of swimming effort (i.e. cost) by free ranging ioural states combined with the amount of time spent animals (Costa & Williams 1999). Of these methods, in each state, the first such field measurements on an stroke rate appears to be the best predictor of swim- otariid pinniped. ming effort (Williams et al. 2004, Wilson et al. 2006). Stroke rate frequencies and patterns have been suc- cessfully measured in a number of free ranging marine MATERIALS AND METHODS animals, including phocid pinnipeds (Williams et al. 2000, 2004), cetaceans (Skrovan et al. 1999, Williams et Subjects, location and equipment. Four lactating al. 2000, Nowacek et al. 2001) and diving seabirds (van females located on St. Paul Island (area known as Reef Dam et al. 2002, Watanuki et al. 2003, Wilson et al. Rookery, 57° 06.466’ N, 170° 17.815’ W), Alaska, were 2006). More importantly, stroke rate appears to have captured and instrumented during late August 2002. A a simple and predictable relationship to dive cost set of 3 instruments was attached to the pelage on the (Williams et al. 2004). Stroke rate, as mostly measured dorsal surface of each female with 5-minute epoxy: an by video and acceleration data, is thus proving to be a acoustic datalogger (Bioacoustic Probe, BP), a PTT, and standard metric for estimating energetic expenditure a VHF radio transmitter. Each BP was mounted along in many free ranging animals (Wilson et al. 2006). the central axis of the dorsal surface of the seal, posi- We focused our attention on the foraging trips of lac- tioned to minimize any body contact during the spinal tating northern fur seal females from the Pribilof Island flexion associated with swimming. After the epoxy had population in the Bering Sea, Alaska. In addition to the set, females were released and returned to their off- wealth of background data on this population of fur spring. Radio checks were then conducted twice daily seals, their foraging ecology is of particular interest for in order to determine whether females had departed or 2 reasons. First, foraging trips of lactating females in returned from foraging trips at sea. Each female was this species are consistently the longest in duration of subsequently recaptured after a single foraging trip, all the otariids (although single trips may be longer in and all instruments were removed. The mass of each of Arctocephalus spp. during times of depleted prey; the 4 seals was 37.5, 40.5, 47.5, and 38.0 kg at the time Gentry 1998). Females of the Pribilof population pre- of deployment, and (in the same order) 38.0, 43.5, dictably alternate 7 to 12 d foraging trips at sea with 50.5 kg at the time of recapture 11 to 12 d later (we 2 to 3 d nursing bouts on land, and while away may were not able to get a return weight on the 4th seal). travel distances in excess of 400 km (Gentry & Kooy- Each BP (Greeneridge Sciences) is fully programma- man 1986, Loughlin et al. 1987, Gentry 1998, Goebel ble and consists of a hydrophone potted in epoxy resin 2002, Robson et al. 2004). As a result, the cost of swim- that records directly to a 288 MB flash memory chip. ming is clearly important in this species. Second, the BPs (19.3 cm long × 3.2 cm diameter; 230 g) sampled at Pribilof Island population is the largest colony of north- 16 bit acoustic resolution with a maximum sampling ern fur seals in the world. Although this population has frequency of 20 kHz. Sampling frequency, which been considered stable for the past 2 decades, it now affects recording time and bandwidth, was set in 2 appears to be declining at an accelerating rate (York et units to 1560 Hz and 2 at 2048 Hz. A sampling Insley et al.: Fur seal activity and stroke rate 149 frequency of 1560 Hz results in a Nyquist of 780 Hz 3m s–1 to evaluate whether a location represented an and, after the anti-aliasing filter, an effective fre- implausible movement, or alternatively failed our filter quency bandwidth of 8 to 578 Hz, providing roughly due to an inaccurate adjacent location.
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