Toothed whale vocal muscles Blowhole Vertex Intermedius Right and Left Vocal Muscles Anterior externus Rostral muscle Posterior externus Anterior externus Posterior internus Anterior internus Mead (1975)

Aroyan (1990)

Harbor porpoise (Pp) Left Nasal Musculature (LNM) Fiber-type profiles Basic 100 μm Harbor porpoise (Pp) Left Nasal Musculature (LNM)

Bottlenose Dolphin (Tt) Right Nasal Musculature (RNM) Fiber-type profiles Acidic 100 μm Bottlenose Dolphin (Tt) Right Nasal Musculature (RNM)

Myosin Expression Mouse Pp Tt Pp Tt Mouse DIA LNM LNM PPS RNM DIA Fast A Fast X Fast B Slow Mouse Pp Tt Pp Tt Mouse DIA LNM LNM PPS RNM DIA

Myosin Expression Mouse Pp Tt Pp Tt Mouse DIA LNM LNM PPS RNM DIA Fast A Fast X Fast B Slow Mouse Pp Tt Pp Tt Mouse DIA LNM LNM PPS RNM DIA

Why study vocal muscles? Compare the muscles of different toothed whales Try to understand the role of these muscles in their sound production Part of a larger project Estimate the energetic costs of producing sounds

Measuring dolphin metabolic rates

Dolphin vocalizations Whistles Squawks Noren et al. (2013)

Dolphin vocalizations Whistles Longer Noren et al. (2013)

Dolphin vocalizations Whistles Longer Require a longer interval between vocalizations Noren et al. (2013)

Dolphin vocalizations Whistles Longer Require a longer interval between vocalizations Produced at a higher sound pressure level Noren et al. (2013)

 21.5 +/- 7.7% Noren et al. (2013)

 5.9 +/- 17.7% Noren et al. (2013)

Cost of vocalizations 1.2 x Resting Metabolic Rate (Noren et al., 2013) But, what about other toothed whales? Killer whales Beaked whales If morphology of vocal muscles is similar Can use Noren et al.’s (2013) study to estimate costs

Why should we care?

Increased anthropogenic noise Toothed and baleen whales will need to produce louder sounds Will cost more energy to produce Sounds are used in:

Energy budgets Important for the proper management of toothed and baleen whales Lead to better estimates of impacts of anthropogenic noise Shipping Oil exploration Navy sonar

Conclusions Toothed and baleen whales produce sound Toothed whales Both – social sounds Toothed – echolocation Toothed whales Unique structures produce sound Many muscles involved Dearolf lab – studying fiber-type profiles and myosin expression Ultimate goal - determine energetic costs of sound production Necessary to understand impacts of anthropogenic noise

Myosin Expression in Dolphin and Porpoise Muscle Acknowledgements Megan Murphy Graduated Porpoise Muscle Fiber-type profile Sarah McHugh Graduated Dolphin Muscle Fiber-type profile Elizabeth Soo Graduated Myosin Expression in Dolphin and Porpoise Muscle

References Aroyan JL. (1990) Numerical simulation of dolphin echolocation beam formation. M.S. thesis, University of California, Santa Cruz. Aroyna JL. (2002) Simplified models of the dolphin echolocation system. http://www2.cruzio.com/~jaroyan/Dolphin%20Model%204.htm. Huggenberger S, Rauschmann MA, Vogl TJ, Oelschläger. (2009) Functional morphology of the nasal complex in the harbor porpoise (Phocoena phocoena L.) Anatomical Record 292: 902-920. Kemper CM. (2002) Pygmy right whale Caperea marginata. In: Perrin WF, Würsig B, Thewissen JGM (eds.) Encyclopedia of marine mammals, 1st ed., Academic Press: San Diego, pp.1010-1012. Mead JG. (1975) Anatomy of the external nasal passages and facial complex in the Delphinidae (Mammalia: Cetacea. Smithsonian Contributions to Zoology 207: 1-35. Noren DP, Holt MM, Dunkin RC, Williams TM. (2013) The metabolic cost of communicative sound production in bottlenose dolphins (Tursiops truncatus). Journal of Experimental Biology 216: 1624- 1629. Rojas-Bracho L, Jaramillo-Legorreta A. (2002) Vaquita Phocoena sinus. In: Perrin WF, Würsig B, Thewissen JGM (eds.) Encyclopedia of marine mammals, 1st ed., Academic Press: San Diego, pp. 1277-1280. Sears R. (2002) Blue whale Balaenoptera musculus. In: Perrin WF, Würsig B, Thewissen JGM (eds.) Encyclopedia of marine mammals, 1st ed., Academic Press: San Diego, pp. 112-116. Whitehead H. (2002) Sperm whale Physeter macrocephalus. In: Perrin WF, Würsig B, Thewissen JGM (eds.) Encyclopedia of marine mammals, 1st ed., Academic Press: San Diego, pp. 1165-1172.

Beluga whale – Delphinapterus leucas Nakungayok – Naku Beluga whale – Delphinapterus leucas 14 years old In captivity for 10 years

North Pool 27 feet diam 9 feet deep South Pool 27 feet diam 9 feet deep Gate Gate Main Pool 70 feet x 40 feet 20 feet deep

Gate Gate

Aurora Kela

Gate Gate Bottlenose dolphins Tursiops truncatus

Misty Arrow Nina

Fish available per session – 1.0 kg Herring – Clupea sp. Average weight – 0.1 kg Fish available per session – 1.0 kg

Objects Five fish One fish