Volume 26, Issue 11, Pages (June 2016)

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Volume 26, Issue 11, Pages 1441-1446 (June 2016) Ultra-High Foraging Rates of Harbor Porpoises Make Them Vulnerable to Anthropogenic Disturbance  Danuta Maria Wisniewska, Mark Johnson, Jonas Teilmann, Laia Rojano-Doñate, Jeanne Shearer, Signe Sveegaard, Lee A. Miller, Ursula Siebert, Peter Teglberg Madsen  Current Biology  Volume 26, Issue 11, Pages 1441-1446 (June 2016) DOI: 10.1016/j.cub.2016.03.069 Copyright © 2016 Elsevier Ltd Terms and Conditions

Figure 1 Buzz Rates Indicative of Prey Encounter Rates of Echolocating Harbor Porpoises (A) Example dive profile from one porpoise. Individual buzzes are marked in red. The shaded area represents twilight (gray) and night (black). (B–F) Hourly buzz counts for the five porpoises as recorded by attached tags. Numbers for the first and last incomplete hours are depicted with dashed lines. The animal’s sex, age class, standard length (SL), tagging date, and location as well as the total number of buzzes (n) and the animal’s estimated success rate (SR; mean and 95% confidence intervals) are provided in each panel. The digits in the names of the individuals indicate the year and Julian day of tag deployment. See also Supplemental Experimental Procedures, Table S1, and Movie S1. Current Biology 2016 26, 1441-1446DOI: (10.1016/j.cub.2016.03.069) Copyright © 2016 Elsevier Ltd Terms and Conditions

Figure 2 Approach and Probable Capture of a Prey by a Harbor Porpoise (A) Echogram (see also Movie S1) displaying sonar clicks and echoes recorded by a DTAG-3 tag attached to the porpoise about 5 cm behind its blowhole (i.e., about 20 cm from the tip of the animal’s rostrum). The image is a stack plot of sound envelopes synchronized to the outgoing clicks, as in an echosounder display. The y axis indicates time elapsed from emitted clicks to returning echoes, expressed as target range from the sound source below the blowhole using a sound speed of 1,500 ms−1. Clicks emitted at rates of more than 125 Hz, corresponding to inter-click intervals (ICIs) shorter than the 8-ms time window chosen here, are displayed repeatedly, making subsequent buzz clicks form a pattern akin to harmonics in the stack plot. The color scale indicates echo-to-noise ratio (ENR) on a dB scale. Amplitude variations in the prey echo track individual tail strokes of the fish when it tries to escape (see Figure 3A for details of the fish echo trace). (B) ICI color-coded for apparent output level (AOL) of echolocation clicks showing a 30-dB reduction in output energy during buzzes. (C) Norm of jerk, i.e., the vector magnitude of the rate of change of acceleration as recorded by the tag. The high magnitude peaks most likely reflect rapid movements in the gular region during generation of suction. (D) Depth (blue) and heading (green) of the tagged porpoise over the same interval. To evaluate prey capture success, we formed similar figures for a subset of buzzes for four of the tagged porpoises and presented them to four evaluators. Current Biology 2016 26, 1441-1446DOI: (10.1016/j.cub.2016.03.069) Copyright © 2016 Elsevier Ltd Terms and Conditions

Figure 3 Determination of Prey Behavior and Size (A) Expanded view of the echogram in Figure 2 showing the echo level (expressed as ENR) variation due to prey tailbeats. (B) Range (blue) and closing speed (green) to the prey extracted from the echogram using a two-state Kalman-Rauch filter to track the prey echo. Negative values of relative speed indicate when the porpoise is closing on the prey, while positive values occur when the prey and predator draw apart. (C) Received level at the tag of each prey echo tracked by the Kalman filter, expressed as root-mean-squared (RMS) ENR. The prey appears to respond to the approaching porpoise at a distance of 65 cm from the sound source (50 cm from the anterior rostrum), and oscillations in the echo level thereafter indicate tail strokes of the escaping fish. (D) Spectrogram of the echo level (interpolated to a regular time grid) showing the frequency (rate) of tail strokes. Each tail stroke requires two muscle contractions, so the 36-Hz stroke rate here implies a contraction time of 14 ms. As minimum contraction time (and therefore highest stroke rate) is a function of body length (BL) and water temperature, the maximum prey size can be deduced from the stroke rate in echograms, in this case BL < 5 cm. This is corroborated by the escape speed of the prey: assuming that the porpoise maintains its initial closing speed of 1.4 m/s throughout the chase, the prey must attain a similar speed at seconds 5.5 and 7.5 when the net speed is 0. This speed is consistent with a 5-cm fish stroking at 36 Hz with a stride of 0.8 BL. Inset in (C) shows the proportion of fish sizes targeted by the tagged porpoises as inferred from tailbeat rates in 30 randomly selected echograms per animal. Current Biology 2016 26, 1441-1446DOI: (10.1016/j.cub.2016.03.069) Copyright © 2016 Elsevier Ltd Terms and Conditions