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FINAL PROGRAMMATIC BIOLOGICAL OPINION ON U.S. NAVY ACTIVITIES IN THE HAWAII RANGE COMPLEX 2008-2013<br />

Other species experience the same phenomen<strong>on</strong> when they vocalize in the presence of high levels of background<br />

sound. Brumm (2004) studied the s<strong>on</strong>gs of territorial male nightingales (L<strong>us</strong>cinia megarhynchos) in the city of<br />

Berlin, Germany, to determine whether and to what degree background noise (from automobile traffic) produced a<br />

Lombard effect in these birds. Based <strong>on</strong> his studies, the birds increased the volume of their s<strong>on</strong>gs in resp<strong>on</strong>se to<br />

traffic noise by 14 dB (their s<strong>on</strong>gs were more than 5 times louder than birds vocalizing in quiet sites). Cynx et al.<br />

(1998) reported similar results based <strong>on</strong> their study of zebra finches (Taeniopygia guttata) exposed to white noise.<br />

Although this type of resp<strong>on</strong>se also has not been studied extensively in marine animals, Scheifele et al. (2005)<br />

reported that beluga whales in the St. Lawrence River increased the decibel levels of their vocalizati<strong>on</strong>s from 80.46-86.76<br />

dB in c<strong>on</strong>diti<strong>on</strong>s without noise to 91.74-99.10 dB when c<strong>on</strong>fr<strong>on</strong>ted with vessel noise.<br />

Holt et al. (2007) reported that endangered southern resident killer whales (Orcin<strong>us</strong> orca) in Haro Strait off the San<br />

Juan Islands in Puget Sound, Washingt<strong>on</strong>, increased the amplitude of their social calls in the face of increased<br />

sounds levels of background noise.<br />

3. Adj<strong>us</strong>t temporal structure of vocalizati<strong>on</strong>s (Box C1.2 of Figure 3). Animals resp<strong>on</strong>ding this way adj<strong>us</strong>t the<br />

temporal structure of their vocalizati<strong>on</strong>s by changing the timing of modulati<strong>on</strong>s, notes, and syllables within<br />

vocalizati<strong>on</strong>s or increasing the durati<strong>on</strong> of their calls or s<strong>on</strong>gs.<br />

Cody and Brown (1969) studied the s<strong>on</strong>gs of adult male Bewick wrens and wrentits that occupied overlapping<br />

territories and whose s<strong>on</strong>gs had similar physical characteristics (similar s<strong>on</strong>g lengths, frequency structure, and<br />

amplitude). They reported that wrentits adj<strong>us</strong>ted the timing of their s<strong>on</strong>gs so they occurred when the s<strong>on</strong>gs of the<br />

Bewick wrens subsided.<br />

Ficken et al. (1974) studied vocalizati<strong>on</strong>s of ten red-eyed vireos (Vireo olivace<strong>us</strong>) and least flycatchers (Empid<strong>on</strong>ax<br />

minim<strong>us</strong>) at Lake Itasca, Minnesota (a total of 2283 s<strong>on</strong>gs). They reported that flycatchers avoided aco<strong>us</strong>tic<br />

interference from red-eyed vireos by inserting their shorter s<strong>on</strong>gs between the l<strong>on</strong>ger s<strong>on</strong>gs of the vireos. Although<br />

there is some mutual avoidance of aco<strong>us</strong>tic interference, the flycatcher tends more str<strong>on</strong>gly to insert its short s<strong>on</strong>gs in<br />

between the l<strong>on</strong>ger s<strong>on</strong>gs of the vireo rather than vice versa. Indeed, most of the overlap occurred when the<br />

flycatcher began singing j<strong>us</strong>t after the vireo had begun, suggesting that the flycatcher had not heard the vireo begin<br />

singing.<br />

A few studies have dem<strong>on</strong>strated that marine mammals make the same kind of vocal adj<strong>us</strong>tments in the face of high<br />

levels of background noise. Rendell and Gord<strong>on</strong> (1999) reported that l<strong>on</strong>g-finned pilot whales (Globicephala melas)<br />

in the Ligurian Sea made several vocal adj<strong>us</strong>tments in call whistles when putatively exposed to active s<strong>on</strong>ar transmissi<strong>on</strong>s<br />

at frequencies of 4-5 kHz (reference and received levels were not reported).<br />

Miller et al. (2000) recorded the vocal behavior of singing humpback whales c<strong>on</strong>tinuo<strong>us</strong>ly for several hours <strong>us</strong>ing a<br />

towed, calibrated hydroph<strong>on</strong>e array. They recorded at least two s<strong>on</strong>gs in which the whales were exposed to lowfrequency<br />

active s<strong>on</strong>ar transmissi<strong>on</strong>s (42 sec<strong>on</strong>d signals at 6 minute intervals; s<strong>on</strong>ar was broadcast so that n<strong>on</strong>e of<br />

the singing whales were exposed at received levels greater than 150 dB re 1µPa). They followed sixteen singing<br />

humpback whales during 18 playbacks. In nine follows, whales sang c<strong>on</strong>tinuo<strong>us</strong>ly throughout the playback; in four<br />

follows, the whale stopped singing when he joined other whales (a normal social interacti<strong>on</strong>); and in five follows,<br />

202

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