Exploring odontocete depredation rates in a pelagic longline fishery.

Several odontocete species depredate catch and bait from fishing gear, resulting in their bycatch and causing substantial economic costs. There are no known mitigation methods for odontocete depredation in pelagic longline fisheries that are effective, do not harm odontocetes and are commercially viable. Understanding odontocetes' depredation strategies can contribute to mitigating this human-wildlife conflict. Using observer data from the Hawaii-based tuna longline fishery, this study summarized teleost and elasmobranch species-specific mean posterior odontocete depredation rates using a simple Bayesian binomial likelihood estimator with a Bayes-Laplace prior. Depredation rates of species with sufficient sample sizes ranged from a high of 1.2% (1.1 to 1.3 95% highest posterior density interval or HDI) for shortbill spearfish to a low of 0.002% (0.001 to 0.003 95% HDI) for blue shark. Depredation of catch is a rare event in this fishery, occurring in about 6% of sets. When depredation did occur, most frequently odontocetes depredated a small proportion of the catch, however, there was large variability in depredation rates between teleost species. For example, bigeye tuna was two times more likely to be depredated than yellowfin tuna (odds ratio = 2.03, 95% CI: 1.8-2.3, P<0.0001). For sets with depredation, 10% and 2% of sets had depredation of over half of the captured bigeye tuna and combined teleosts, respectively. All elasmobranch species had relatively low depredation rates, where only 7 of almost 0.5M captured elasmobranchs were depredated. Odontocetes selectively depredate a subset of the teleost species captured within sets, possibly based on net energy value, chemical, visual, acoustic and textural characteristics and body size, but not median length, which was found to be unrelated to depredation rate (Pearson's r = 0.14, 95% CI: -0.26 to 0.50, p = 0.49). Study findings provide evidence to support the identification and innovation of effective and commercially viable methods to mitigate odontocete depredation and bycatch.

Quantifying the age structure of free-ranging delphinid populations: Testing the accuracy of Unoccupied Aerial System photogrammetry.

Understanding the population health status of long-lived and slow-reproducing species is critical for their management. However, it can take decades with traditional monitoring techniques to detect population-level changes in demographic parameters. Early detection of the effects of environmental and anthropogenic stressors on vital rates would aid in forecasting changes in population dynamics and therefore inform management efforts. Changes in vital rates strongly correlate with deviations in population growth, highlighting the need for novel approaches that can provide early warning signs of population decline (e.g., changes in age structure). We tested a novel and frequentist approach, using Unoccupied Aerial System (UAS) photogrammetry, to assess the population age structure of small delphinids. First, we measured the precision and accuracy of UAS photogrammetry in estimating total body length (TL) of trained bottlenose dolphins (Tursiops truncatus). Using a log-transformed linear model, we estimated TL using the blowhole to dorsal fin distance (BHDF) for surfacing animals. To test the performance of UAS photogrammetry to age-classify individuals, we then used length measurements from a 35-year dataset from a free-ranging bottlenose dolphin community to simulate UAS estimates of BHDF and TL. We tested five age classifiers and determined where young individuals (<10 years) were assigned when misclassified. Finally, we tested whether UAS-simulated BHDF only or the associated TL estimates provided better classifications. TL of surfacing dolphins was overestimated by 3.3% ±3.1% based on UAS-estimated BHDF. Our age classifiers performed best in predicting age-class when using broader and fewer (two and three) age-class bins with ~80% and ~72% assignment performance, respectively. Overall, 72.5%-93% of the individuals were correctly classified within 2 years of their actual age-class bin. Similar classification performances were obtained using both proxies. UAS photogrammetry is a non-invasive, inexpensive, and effective method to estimate TL and age-class of free-swimming dolphins. UAS photogrammetry can facilitate the detection of early signs of population changes, which can provide important insights for timely management decisions.

The startle reflex in echolocating odontocetes: basic physiology and practical implications.

The acoustic startle reflex is an oligo-synaptic reflex arc elicited by rapid-onset sounds. Odontocetes evolved a range of specific auditory adaptations to aquatic hearing and echolocation, e.g. the ability to downregulate their auditory sensitivity when emitting clicks. However, it remains unclear whether these adaptations also led to changes of the startle reflex. We investigated reactions to startling sounds in two bottlenose dolphins (Tursiops truncatus) and one false killer whale (Pseudorca crassidens). Animals were exposed to 50 ms, 1/3 octave band noise pulses of varying levels at frequencies of 1, 10, 25 and 32 kHz while positioned in a hoop station. Startle responses were quantified by measuring rapid muscle contractions using a three-dimensional accelerometer attached to the dolphin. Startle magnitude increased exponentially with increasing received levels. Startle thresholds were frequency dependent and ranged from 131 dB at 32 kHz to 153 dB at 1 kHz (re. 1 µPa). Startle thresholds only exceeded masked auditory AEP thresholds of the animals by 47 dB but were ∼82 dB above published behavioural audiograms for these species. We also tested the effect of stimulus rise time on startle magnitude using a broadband noise pulse. Startle responses decreased with increasing rise times from 2 to 100 ms. Models suggested that rise times of 141-220 ms were necessary to completely mitigate startle responses. Our data showed that the startle reflex is conserved in odontocetes and follows similar principles as in terrestrial mammals. These principles should be considered when assessing and mitigating the effects of anthropogenic noise on marine mammals.

Transmission beam pattern and dynamics of a spinner dolphin (Stenella longirostris).

Toothed whales possess a sophisticated biosonar system by which ultrasonic clicks are projected in a highly directional transmission beam. Beam directivity is an important biosonar characteristic that reduces acoustic clutter and increases the acoustic detection range. This study measured click characteristics and the transmission beam pattern from a small odontocete, the spinner dolphin (Stenella longirostis). A formerly stranded individual was rehabilitated and trained to station underwater in front of a 16-element hydrophone array. On-axis clicks showed a mean duration of 20.1 µs, with mean peak and centroid frequencies of 58 and 64 kHz [standard deviation (s.d.) ±30 and ±12 kHz], respectively. Clicks were projected in an oval, vertically compressed beam, with mean vertical and horizontal beamwidths of 14.5° (s.d. ± 3.9) and 16.3° (s.d. ± 4.6), respectively. Directivity indices ranged from 14.9 to 27.4 dB, with a mean of 21.7 dB, although this likely represents a broader beam than what is normally produced by wild individuals. A click subset with characteristics more similar to those described for wild individuals exhibited a mean directivity index of 23.3 dB. Although one of the broadest transmission beams described for a dolphin, it is similar to other small bodied odontocetes.

Modulation rate transfer functions from four species of stranded odontocete (Stenella longirostris, Feresa attenuata, Globicephala melas, and Mesoplodon densirostris).

Odontocete marine mammals explore the environment by rapidly producing echolocation signals and receiving the corresponding echoes, which likewise return at very rapid rates. Thus, it is important that the auditory system has a high temporal resolution to effectively process and extract relevant information from click echoes. This study used auditory evoked potential methods to investigate auditory temporal resolution of individuals from four different odontocete species, including a spinner dolphin (Stenella longirostris), pygmy killer whale (Feresa attenuata), long-finned pilot whale (Globicephala melas), and Blainville's beaked whale (Mesoplodon densirostris). Each individual had previously stranded and was undergoing rehabilitation. Auditory Brainstem Responses (ABRs) were elicited via acoustic stimuli consisting of a train of broadband tone pulses presented at rates between 300 and 2000 Hz. Similar to other studied species, modulation rate transfer functions (MRTFs) of the studied individuals followed the shape of a low-pass filter, with the ability to process acoustic stimuli at presentation rates up to and exceeding 1250 Hz. Auditory integration times estimated from the bandwidths of the MRTFs ranged between 250 and 333 µs. The results support the hypothesis that high temporal resolution is conserved throughout the diverse range of odontocete species.

Four odontocete species change hearing levels when warned of impending loud sound.

Hearing sensitivity change was investigated when a warning sound preceded a loud sound in the false killer whale (Pseudorca crassidens), the bottlenose dolphin (Tursiops truncatus), the beluga whale (Delphinaperus leucas) and the harbor porpoise (Phocoena phocoena). Hearing sensitivity was measured using pip-train test stimuli and auditory evoked potential recording. When the test/warning stimuli preceded a loud sound, hearing thresholds before the loud sound increased relative to the baseline by 13 to 17 dB. Experiments with multiple frequencies of exposure and shift provided evidence of different amounts of hearing change depending on frequency, indicating that the hearing sensation level changes were not likely due to a simple stapedial reflex.

Conditioned hearing sensitivity change in the harbor porpoise (Phocoena phocoena).

Hearing sensitivity, during trials in which a warning sound preceding a loud sound, was investigated in two harbor porpoises (Phocoena phocoena). Sensitivity was measured using pip-train test stimuli and auditory evoked potential recording. When a hearing test/warning stimulus, with a frequency of either 45 or 32 kHz, preceded a loud 32 kHz tone with a sound pressure level of 152 dB re 1 µPa root mean square, lasting 2 s yielding an sound exposure level (SEL) of 155 dB re 1 µPa(2)s, pooled hearing thresholds measured just before the loud sound increased relative to baseline thresholds. During two experimental sessions the threshold increased up to 17 dB for the test frequency of 45 kHz and up to 11 dB for the test frequency of 32 kHz. An extinction test revealed very rapid threshold recovery within the first two experimental sessions. The SEL producing the hearing dampening effect was low compared to previous other odontocete hearing change efforts with each individual trial equal to 155 dB re 1 µPa(2) but the cumulative SEL for each subsession may have been as high as 168 dB re 1 µPa(2). Interpretations of conditioned hearing sensation change and possible change due to temporary threshold shifts are considered for the harbor porpoise and discussed in the light of potential mechanisms and echolocation.

Expectancy and conditioned hearing levels in the bottlenose dolphin (Tursiops truncatus).

The hearing sensitivity of a bottlenose dolphin for a warning sound, when the exact time of the arrival of a loud sound could or could not be predicted, was measured. Sensitivity was measured when the time of onset of the loud sound was randomly varied (random-variation sessions) and when the time of onset of the loud sound and the pattern of stimulus levels was constant (fixed-stimulus sessions). The loud sound was kept the same in both of the series. The mean duration and mean range of the levels of the test/warning signal were also kept equal across experimental sessions. Hearing sensitivity was measured using the auditory evoked potential method with rhythmic trains of short pips as test stimuli. With randomly varied warning sounds, thresholds before the loud sound were on average 10.6 dB higher than the baseline thresholds. With fixed warning signals, thresholds were on average 4.4 dB higher than the baseline thresholds. Considering that the loud sounds were identical, the difference between the random-variation and the fixed-stimulus sessions cannot be explained by a direct (unconditioned) influence of sound exposure. Therefore, the data provide reliable evidence for the conditioning nature of the hearing-dampening effect and also demonstrate that hearing sensitivity change also depends on when the animal can expect the loud sound to occur.

Learning and extinction of conditioned hearing sensation change in the beluga whale (Delphinapterus leucas).

Ice-dwelling beluga whales are increasingly being exposed to anthropogenic loud sounds. Beluga's hearing sensitivity measured during a warning sound just preceding a loud sound was tested using pip-train stimuli and auditory evoked potential recording. When the test/warning stimulus with a frequency of 32 or 45 kHz preceded the loud sound with a frequency of 32 kHz and a sound pressure level of 153 dB re 1 µPa, 2 s, hearing thresholds before the loud sound increased relative to the baseline. The threshold increased up to 15 dB for the test frequency of 45 kHz and up to 13 dB for the test frequency of 32 kHz. These threshold increases were observed during two sessions of 36 trials each. Extinction tests revealed no change during three experimental sessions followed by a jump-like return to baseline thresholds. The low exposure level producing the hearing-dampening effect (156 dB re 1 µPa(2)s in each trial), and the manner of extinction, may be considered as evidence that the observed hearing threshold increases were a demonstration of conditioned dampening of hearing when the whale anticipated the quick appearance of a loud sound in the same way demonstrated in the false killer whale and bottlenose dolphin.

Hearing in Whales and Dolphins: Relevance and Limitations.

Understanding the hearing of marine mammals has been a priority to quantify and mitigate the impact of anthropogenic sound on these apex predators. Yet our knowledge of cetacean hearing is still limited to a few dozen species, therefore compromising any attempt to design adaptive management strategies. The use of auditory evoked potentials allows scientists to rapidly and noninvasively obtain the hearing data of species rarely available in captivity. Unfortunately, many practical and ethical reasons still limit the availability of large whales, thus restricting the possibility to effectively ensure that anthropogenic sounds have minimum effects on these species. The example of a recent Blainville's beaked whale (Mesoplodon densirostris) audiogram collected after a stranding indicated, for instance, very specialized hearing between 40 and 50 kHz, which corresponded to the frequency-modulated upsweep signals used by this species during echolocation. The methods used during a stranding event are presented along with the major difficulties that have slowed down the scientific community in measuring the audition of large whales and the potential value in obtaining such results when successful.

Cognitive adaptation of sonar gain control in the bottlenose dolphin.

Echolocating animals adjust the transmit intensity and receive sensitivity of their sonar in order to regulate the sensation level of their echoes; this process is often termed automatic gain control. Gain control is considered not to be under the animal's cognitive control, but previous investigations studied animals ensonifying targets or hydrophone arrays at predictable distances. To test whether animals maintain gain control at a fixed level in uncertain conditions, we measured changes in signal intensity for a bottlenose dolphin (Tursiops truncatus) detecting a target at three target distances (2.5, 4 and 7 m) in two types of sessions: predictable and unpredictable. Predictable sessions presented the target at a constant distance; unpredictable sessions moved the target randomly between the three target positions. In the predictable sessions the dolphin demonstrated intensity distance compensation, increasing the emitted click intensity as the target distance increased. Additionally, as trials within sessions progressed, the animal adjusted its click intensity even from the first click in a click train, which is consistent with the animal expecting a target at a certain range. In the unpredictable sessions there was no significant difference of intensity with target distance until after the 7th click in a click train. Together, these results demonstrate that the bottlenose dolphin uses learning and expectation for sonar gain control.

Audiogram of a stranded Blainville's beaked whale (Mesoplodon densirostris) measured using auditory evoked potentials.

Quantifying and understanding the impact of anthropogenic sound on marine mammals has been the focus of many researchers both in laboratory settings as well as in the field. This study presents the audiogram of a sub-adult Blainville's beaked whale that stranded in Hawaii. The hearing measurements were conducted using the non-invasive auditory brainstem response technique. A total of 11 sinusoidally amplitude modulated tones were tested ranging from 5.6 to 160 kHz. The audiogram data indicated that the region of best hearing was found between 40 and 50 kHz with thresholds below 50 dB. This frequency range partially overlaps with the frequency modulated upsweep that Blainville's beaked whales have been reported to use during echolocation. These results match the frequency range obtained from the hearing measurements of a Gervais' beaked whale previously tested using contact acoustic stimulation and emphasize the importance of obtaining rapid hearing measurements on live stranded animals to improve the understanding of poorly known species.