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Beaked whales

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Madsen P.T.1*, Aguilar de Soto N.2, Tyack P.L.3, and Johnson M.3 9

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1) Zoophysiology, Dept. of Bioscience, Aarhus University, Denmark 12

2) BIOECOMAC, La Laguna University, Tenerife, Spain 13

3) SMRU, University of St. Andrews, Scotland 14

* [email protected] 15

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2 What is a beaked whale?

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The beaked whales (Ziphiidae) belong to a little known cetacean family of more than 21 species that 25

range in size from 3m and a few hundred kg to more than 10 meters and 8,000 kg. Even though beaked 26

whales all belong to the suborder toothed whales, most species, ironically, have few if any erupted teeth, 27

with these serving as tusks in male-male interactions rather than for foraging. Despite few geographical 28

boundaries in their deep ocean habitat, superficially similar, but genetically distinct, species of beaked 29

whales share what appear to be similar foraging niches. Beaked whales routinely dive deeper than 1km 30

for an hour or more, and surface for such a short time that they are very difficult to sight. This cryptic 31

lifestyle has, until recently, left us with very little information about some of the world’s biggest 32

predators beyond what can be gathered from stranded specimens. This second largest family of toothed 33

whales is so little known that two new species, Perrin's beaked whale and the pygmy beaked whale, 34

have been identified in the last 25 years, and a few more beaked whale species may be awaiting 35

discovery. It is thought-provoking that there are likely elephant-sized, mammalian predators still 36

roaming the world’s oceans that science has not yet even named. However, after many years of being a 37

largely overlooked zoological oddity, beaked whales have recently received substantial public attention 38

after a series of mass-strandings caused by mid frequency naval sonars. 39

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Why do beaked whales dive so deep? 41

The need to understand why beaked whales may respond so strongly to navy sonar has prompted the 42

development of techniques to study these elusive predators of the high seas. Deployments of electronic 43

tags have revealed the underwater behavior of bottlenose, Blainville’s and Cuvier’s beaked whales, 44

showing that these species routinely dive to more mesopelagic depths where they hunt for small, deep-45

water squid and fish. These food resources are found so deep that beaked whales often dive to more 46

than 1000m depths for around an hour (figure 1) routinely exceeding their aerobic dive limit. They 47

therefore return to the surface with a substantial oxygen debt that is paid off in a prolonged surface 48

time that includes a series of shallow non-foraging dives (figure 1). Recently, a tagged Cuvier’s beaked 49

whale was recorded diving more than 3000 meters during a 2 hour long dive that is by far the deepest 50

dive recorded for any air-breathing endotherm. How a mammal can hold its breath for so long and 51

survive a hydrostatic pressure of >300 kg/cm2 is still very much an unsolved mystery. But a 52

consequence of this deep water foraging is that beaked whales spend <20 % of their time foraging, 53

which may explain why they are often found around islands and in upwelling areas that provide stable 54

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3 How do they find and catch food in the deep sea?

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The task of locating, approaching and catching small, agile prey in the cold, dark abyss may seem almost 57

impossible to us humans. Beaked whales and other toothed whales have solved that problem by 58

emitting ultrasonic clicks and listening for returning echoes to hunt by echolocation. When the first 59

sound recording tags were deployed on Blainville's beaked whales, we were astounded to find that not 60

only could the tags record the emitted echolocation clicks, but also the echoes returning from prey, 61

allowing us a unique opportunity to tap into the sensory stream of a predator hunting in the wild. 62

Because of this, beaked whales are now among the best-studied animals that use echolocation to hunt, 63

and they have become a very unlikely model for how toothed whales in general operate their sonars in 64

the wild. Tagged whales emit some 3500 echolocation clicks to detect and approach about 25 prey per 65

dive. When whales catch their prey, they accelerate the click rate to a “buzz,” as is the case for 66

echolocating bats, revealing a remarkable functional convergence of biosonars in air and water. Analysis 67

of echo data from two species of beaked whales show that they ensonify many more organisms than 68

they try to capture, indicating careful prey selection via echo information to optimize energy returns 69

during intense bouts of foraging with durations that are limited by breath-hold dives. 70

What is their social structure? 71

Little is known about the social structure of beaked whales, but the smaller species seem to live in 72

groups of females and young with a single mature male. The heavily scarred older males may be seen 73

with different groups of females over time suggesting that they fight over access to the females using 74

their tusks. Females are often larger than males, which may relate to the large birth weight of beaked 75

whale calves enabling them to achieve deep diving capabilities sooner. Beaked whale groups remain 76

close together but, unlike in other toothed whales, this is not mediated by social calls at the surface, at 77

least in the smaller species, which seem to keep track of one another at depth by eavesdropping on one 78

another’s echolocation clicks. Blainville’s beaked whales also produce social sounds but these are made 79

at depths below 200 m, which together with long silent ascents from deep dives suggest a strategy of 80

acoustic crypsis perhaps to reduce the risk of predation from killer whales that seldom dive deep. 81

Species with larger body size, such as the bottlenose and Baird’s beaked whales, may have a more 82

relaxed crypsis, forming larger groups while making some sounds near the surface. 83

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4 Why do some beaked whales strand when exposed to navy sonars?

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The anti-predation strategies of beaked whales may also explain why some smaller beaked whale 88

species are prone to strand in conjunction with naval sonar exercises. Recent playback studies suggest 89

that the strandings happen as a result of strong behavioral responses that appear to be elicited by low 90

level sonar pulses at frequencies around 3-4 kHz. Playbacks of killer whale calls in a similar frequency 91

range also evoke strong flight responses, suggesting that the sonar pulses may trigger an antipredator 92

avoidance behaviour with potential physiological consequences that may include decompression 93

sickness from repeated dives to escape the perceived threat. 94

What is their conservation status? 95

Although mass-strandings of beaked whales provide a dramatic example of the vulnerability of these 96

species to human activities, sonar exposure is not the only, nor necessarily the most important, 97

conservation threat to these large predators. Beaked whales live in areas that are hard to survey and 98

their diving behavior precludes reliable visual counts, leading the IUCN to list most species as “data 99

deficient.” Currently, we know next to nothing about how beaked whale populations are affected by 100

potential environmental stressors such as toxins, shipping noise, or bycatch in fisheries, nor do we 101

know anything about their global population sizes. However, their distinctive frequency-modulated 102

clicks with potential species-specific differences now enable acoustic surveys to estimate habitat use 103

and population sizes. The echolocation clicks, vital for their foraging, may thus also provide researchers 104

with a unique window to study and protect some of the largest and most cryptic predators alive. 105

Where can I learn more? 106

Aguilar Soto N., Madsen P. T., Tyack P., Arranz P., Marrero J., Fais A., Revelli E. and Johnson M. P. (2011) , "No 107

shallow talk: Cryptic strategy in the vocal communication of Blainville’s beaked whales". Marine Mammal Science, 108

DOI: 10.1111/j.1748-7692.2011.00495.x 109

Cox, T. M., Ragen, T. J., Read, A. J., Vos, E., Baird, R. W., Balcomb, K., Barlow, J., Caldwell, J., Cranford, T., Crum, L. et 110

al. (2006). Why do beaked whales strand? Report of workshop to understand the impacts of anthropogenic 111

sound. J. Cetacean Res. Manag. 7, 177-187. 112

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Dalebout, M.L, Baker, C.S., Cockcroft ,V.G., Mead, J.G., and Yamada, T.K. 2004. A comprehensive and validated

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molecular taxonomy of beaked whales, family Ziphiidae. Journal of Heredity 96, 459-473.

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Hooker, S.K., and R.W. Baird. 1999. Deep-diving behaviour of the northern bottlenose whale, Hyperoodon

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ampullatus (Cetacea: Ziphiidae). Proceedings of the Royal Society, London B. 266: 671-676. 117

Madsen P.T., Aguilar Soto N., Arranz P. and Johnson M.(2013) "Echolocation in Blainville’s beaked whales 118

(Mesoplodon densirostris)". J. Comp. Physiol. A 199: 451-469. 119

Tyack, P.L., Johnson, M., Aguilar Soto, N., Sturlese, A., Madsen, P.T. (2006), "Extreme diving of beaked whales", 120

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Figure 1. 125

A) Dive profile of a Blainville’s beaked whale. Echolocation clicks (yellow line) are only made during the 126

deep foraging dive where red dots mark foraging buzzes. B) Male Blainville’s beaked whale with 127

barnacles on its large tusks (Photo credit ULL group). C) Breaching Cuviers beaked whale of El Hierro in 128

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