A key component in the operation of a biosonar system is the radiation of sound energy from the sound producing head structures of toothed whales and microbats. The current view involves a fixed transmission aperture by which the beam width can only change via changes in the frequency of radiated clicks. To test that for a porpoise, echolocation clicks were recorded with high angular resolution using a 16 hydrophone array. The beam is narrower than previously reported (DI = 24 dB) and slightly dorso-ventrally compressed (horizontal -3 dB beam width: 13°, vertical -3 dB beam width: 11°). The narrow beam indicates that all smaller toothed whales investigated so far have surprisingly similar beam widths across taxa and habitats. Obtaining high directionality may thus be at least in part an evolutionary factor that led to high centroid frequencies in a group of smaller toothed whales emitting narrow band high frequency clicks. Despite the production of stereotyped narrow band high frequency clicks, changes in the directionality by a few degrees were observed, showing that porpoises can obtain changes in sound radiation.

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a b s t r a c tManaging animal units is essential in biological conservation and requires spatial and temporal identification of such units. Since even neighbouring populations often have different conservation status and face different levels of anthropogenic pressure, detailed knowledge of population structure, seasonal range and overlap with animals from neighbouring populations is required to manage each unit separately. Previous studies on genetic structure and morphologic separation suggests three distinct populations of harbour porpoises with limited geographic overlap in the North Sea (NS), the Belt Sea (BS) and the Baltic Proper (BP) region. In this study, we aim to identify a management unit for the BS population of harbour porpoises. We use Argos satellite data and genetics from biopsies of tagged harbour porpoises as well as acoustic data from 40 passive acoustic data loggers to determine management areas with the least overlap between populations and thus the least error when abundance and population status is estimated. Discriminant analysis of the satellite tracking data from the BS and NS populations showed that the best fit of the management unit border during the summer months was an east-west line from Denmark to Sweden at latitude 56.95°N. For the border between BS and BP, satellite tracking data indicate a sharp decline in population density at 13.5°E, with 90% of the locations being west of this line. This was supported by the acoustic data with the average daily detection rate being 27.5 times higher west of 13.5°E as compared to east of 13.5°E. By using this novel multidisciplinary approach, we defined a management unit for the BS harbour porpoise population. We recommend that these boundaries are used for future monitoring efforts of this population under the EU directives. The boundaries may also be used for conservation efforts during the summer months, while seasonal movements of harbour porpoises should be considered during winter.

Toothed whales use sonar to detect, locate, and track prey. They adjust emitted sound intensity, auditory sensitivity and click rate to target range, and terminate prey pursuits with high-repetition-rate, low-intensity buzzes. However, their narrow acoustic field of view (FOV) is considered stable throughout target approach, which could facilitate prey escape at close-range. Here, we show that, like some bats, harbour porpoises can broaden their biosonar beam during the terminal phase of attack but, unlike bats, maintain the ability to change beamwidth within this phase. Based on video, MRI, and acoustic-tag recordings, we propose this flexibility is modulated by the melon and implemented to accommodate dynamic spatial relationships with prey and acoustic complexity of surroundings. Despite independent evolution and different means of sound generation and transmission, whales and bats adaptively change their FOV, suggesting that beamwidth flexibility has been an important driver in the evolution of echolocation for prey tracking.DOI: http://dx.doi.org/10.7554/eLife.05651.001

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 estimate. 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 patterns 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. Consequently, 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.