Temporal range: Miocene–Recent
Beaked whales are the members of the family Ziphiidae, which consists of 22 species. These toothed whales are notable for their elongated beaks. Among air-breathing animals, beaked whales are some of the most extreme divers: Cuvier's beaked whales regularly dive for an hour at a depth over 1,000 m (3,300 ft), and the longest and deepest foraging dive recorded is 137.5 minutes at 2,992 m (9,816 ft). This is the greatest dive-depth known for a mammal.
Beaked whales are one of the least known groups of mammals because of their deep-sea habitat, mysterious habits, and apparent low abundance. Only three to four of the 22 species are reasonably well-known. Baird's and Cuvier's beaked whales were subject to commercial exploitation, off the coast of Japan, while the northern bottlenose whale was extensively hunted in the northern part of the North Atlantic late in the 19th and early 20th centuries.
Beaked whales are moderate in size, ranging from 4 to 13 metres (13 to 43 ft) and weighing from 1 to 15 tonnes (0.98 to 14.76 long tons; 1.1 to 16.5 short tons). Their key distinguishing feature is the presence of a 'beak', somewhat similar to many dolphins. Other distinctive features include a pair of converging grooves under the throat, and the absence of a notch in the tail fluke. Although Shepherd's beaked whale is an exception, most species have only one or two pairs of teeth, and even these do not erupt in females. Beaked whale species are often sexually dimorphic – one or the other sex is significantly larger. The adult males often possess a dramatically bulging forehead. However, aside from dentition and size, very few morphological differences exist between male and female beaked whales.
Individual species are very difficult to identify in the wild, since body form varies little from one species to another. The observer must rely on often subtle differences in size, color, forehead shape, and beak length.
Beaked whales are unique among toothed whales in that most species only have one pair of teeth. The teeth are tusk-like, but are only visible in males, which are presumed to use these teeth in combat for females for reproductive rights. In females, the teeth do not develop and remain hidden in the gum tissues.
In December 2008, researchers from the Marine Mammal Institute at Oregon State University completed a DNA tree of 13 of 15 known species of Mesoplodon beaked whales (excluding spade-toothed whale which is only known from a skeletal and a few stranded specimens). Among the results of this study was the conclusion that the male's teeth are actually a secondary sexual characteristic, similar to the antlers of male deer. Each species' teeth have a characteristically unique shape. In some cases, these teeth even hinder feeding; in the strap-toothed whale, for example, the teeth curve over the upper jaw, effectively limiting the gape to a few centimeters. Females are presumed to select mates based on the shape of the teeth, because the different species are otherwise quite similar in appearance.
The social structure is not well-known, but sightings of single males accompanying multiple females suggest a polygynous mating system – which would considerably limit the number of males a female individual can choose.
Beaked whales comprise at least 22 species of small whales in the family Ziphiidae, which is one of the least-known families of large mammals; several species have been described only in the last two decades. Six genera have been identified.
The beaked whales are the second-largest family of cetaceans (after the dolphins) They were one of the first groups to diverge from the ancestral lineage. The earliest known beaked whale fossils date to the Miocene, about 15 million years ago.
- Order Cetacea
- Suborder Odontoceti: Toothed whales
- Family Ziphiidae
- Incertae sedis
- Basal forms
- Subfamily Berardiinae
- Subfamily Hyperoodontinae
- Genus †Africanacetus
- Genus Hyperoodon, bottlenose whales
- Genus Ihlengesi
- Genus Indopacetus
- I. pacificus, Longman's beaked whale
- Genus Mesoplodon, mesoplodont whales
- M. bidens, Sowerby's beaked whale
- M. bowdoini, Andrews' beaked whale
- M. carlhubbsi, Hubbs' beaked whale
- M. densirostris, Blainville's beaked whale
- M. europaeus, Gervais' beaked whale
- M. ginkgodens, ginkgo-toothed beaked whale
- M. grayi, Gray's beaked whale
- M. hectori, Hector's beaked whale
- M. layardii, strap-toothed whale
- M. mirus, True's beaked whale,
- M. peruvianus, pygmy beaked whale
- M. perrini, Perrin's beaked whale
- M. stejnegeri, Stejneger's beaked whale
- M. traversii, spade-toothed whale
- M. hotaula, Deraniyagala's beaked whale
- Subfamily Ziphiinae
- Family Ziphiidae
As many as 26 genera antedate humans.[page needed] Some included ancestors of giant beaked whales (Berardius), such as Microberardius and Cuvier's beaked whale (Ziphius) had many relatives, such as Caviziphius, Archaeoziphius, and Izikoziphius. They were probably preyed upon by predatory whales and sharks, including Carcharocles megalodon.
Recently, a large fossil ziphiid sample was discovered off the South African coast, confirming the extant ziphiid diversity might just be a remnant of a higher past diversity. After studying numerous fossil skulls off the shore of South Africa, researchers discovered the absence of functional maxillary teeth in all South African fossil ziphiids, which is evidence that suction feeding had already developed in several beaked whale lineages during the Miocene. Researchers also found fossil ziphiids with robust skulls, signaling that tusks were used for male-male interactions (speculated with extant beaked whales).
Beaked whales are deep divers with extreme dive profiles. They regularly dive deeper than 500 m (1,600 ft) to echolocate for food, and these deep dives are often followed by multiple shallower dives less than 500 m. This pattern is not always followed, however. Animals have been observed spending more than an hour at or near the surface breathing. Beaked whales are often seen surfacing synchronously, but asynchronous surfacing has also been observed. In March 2014, a study by Cascadia Research revealed that beaked whales were recorded to dive at least 2992 m in depth and lasting as long as 137.5 minutes, both mammalian records (Cuvier's).
Deep-diving mammals face a number of challenges related to extended breath-holding and hydrostatic pressure. Cetaceans and pinnipeds that prolong apnea must optimize the size and use of their oxygen stores, and they must deal with the accumulation of lactic acid due to anaerobic metabolism. Beaked whales have several anatomical adaptations to deep diving: large spleens, livers, and body shape. Most cetaceans have small spleens. However, beaked whales have much larger spleens than delphinids, and may have larger livers, as well. These anatomical traits, which are important for filtering blood, could be adaptations to deep diving. Another notable anatomical adaptation among beaked whales is a slight depression in the body wall that allows them to hold their pectoral flippers tightly against their bodies for increased streamlining. However, they are not invulnerable to the effects of diving so deep and so often. Cascadia Research shows that the deeper the whales dive, the less often they dive per day, cutting their efforts by at least 40%.
The challenges of deep diving are also overcome by the unique diving physiology of beaked whales. Oxygen storage during dives is mostly achieved by blood hemoglobin and muscle myoglobin. While the whale is diving, its heart rate slows and blood flow changes. This physiological dive response ensures oxygen-sensitive tissues maintain a supply of oxygen, while those tissues tolerant to hypoxia receive less blood flow. Additionally, lung collapse obviates the exchange of lung gas with blood, likely minimizing the uptake of nitrogen by tissues.
The throats of all beaked whales have a bilaterally paired set of grooves that are associated with their unique feeding mechanism, suction feeding. Instead of capturing prey with their teeth, beaked whales suck it into their oral cavity. Suction is aided by the throat grooves, which stretch and expand to accommodate food. Their tongues can move very freely. By suddenly retracting the tongue and distending the gular (throat) floor, pressure immediately drops within the mouth, sucking the prey in with the water.
Dietary information is available from stomach contents analyses of stranded beaked whales and from whaling operations. Their preferred diet is primarily deep-water squid, but also benthic and benthopelagic fish and some crustaceans, mostly taken near the sea floor. In a recent study, gouge marks in the seafloor were interpreted to be a result of feeding activities by beaked whales.
To understand the hunting and foraging behavior of beaked whales, researchers used sound and orientation recording devices (DTAGs) on two species: Cuvier's beaked whale (Ziphius cavirostris) and Blainville's beaked whale (Mesoplodon densirostris). These whales hunt by echolocation in deep water (where the majority of their prey is located) between about 200 and 1,885 m (656 and 6,184 ft) and usually catch about 30 prey per dive. Cuvier's beaked whales must forage on average at 1,070 m (3,510 ft) for 58 minutes and Blainville's beaked whales typically forage at 835 m (2,740 ft) deep for an average of 47 minutes.
Range and habitat
The family Ziphiidae is one of the most widespread families of cetaceans, ranging from the ice edges at both the north and south poles, to the equator in all the oceans. Specific ranges vary greatly by species, though beaked whales typically inhabit offshore waters that are at least 300 m deep.
Beaked whales are known to congregate in deep waters off the edge of continental shelves, and bottom features, such as seamounts, canyons, escarpments, and oceanic islands, including the Azores and the Canary Islands, and even off the coasts of Hawaii.
Very little is known about the life histories of beaked whales. The oldest one ever recorded was 84 years for a male Baird's beaked whale, and the oldest recorded female Baird's beaked whale is 54 years. For all other beaked whale species studied, the highest recorded age is between 27 and 39 years. Sexual maturity is reached between seven and 15 years of age in Baird's beaked whales and northern bottlenose whales. Gestation varies greatly between species, lasting 17 months for Baird's beaked whales and 12 months for the northern bottlenose whale. No data are available on their reproductive rates.
It is difficult to determine group size of beaked whales, due to their inconspicuous surfacing behavior. Groups of beaked whales, defined as all individuals found in the same location at the same time, have been reported as ranging from one to 100 individuals. Nevertheless, some populations' group size has been estimated from repeated observations. For example, northern and southern bottlenose whales (H. ampullatus and H. planifrons), Cuvier's beaked whales, and Blainville's beaked whales (Mesoplodon densirostris) have a reported maximum group size of 20 individuals, with the average ranging from 2.5 to 3.5 individuals. Berardius species and Longman's beaked whales (Indopacetus pacificus) are found in larger groups of up to 100 individuals.
Not much information is available about group composition of beaked whales. Only three species have been studied in any detail: northern bottlenose whales, Blainville's beaked whales, and Baird's beaked whales. Female northern bottlenose whales appear to form a loose network of social partners with no obvious long-term associations. In contrast to females, some male northern bottlenose whales have been repeatedly recorded together over several years, and possibly form long-term associations. Studies of Blainville's beaked whales have revealed groups usually consist of a number of females, calves, and/or juvenile animals. These whales are assumed to live in "harem-like" groups, where several females and young are accompanied by a single male. Baird's beaked whales are known to occur in multiple male groups, and in large groups consisting of adult animals of both sexes.
For many years, most beaked whale species were insulated from anthropogenic impacts because of their remote habitat. However, now several issues of concern include:
- Studies of stranded beaked whales show rising levels of toxic chemicals in their blubber.
- As a top predator, beaked whales, like raptors, are particularly vulnerable to build-up of biocontaminants. They can ingest plastic (which can be lethal).
- They more frequently become trapped in trawl nets, due to the expansion of deepwater fisheries.
- Decompression sickness
A major conservation concern for beaked whales (family Ziphiidae) is they appear to be vulnerable to modern sonar operations, which arises from recent strandings that temporally and physically coincide with naval sonar exercises. Post mortem examinations of the stranded whales in concurrence with naval exercises have reported the presence of hemorrhaging near the ears or gas and fat emboli, which could have a deleterious impact on beaked whales that is analogous to decompression sickness in humans. Gas and fat emboli have been shown to cause nervous and cardiovascular system dysfunction, respiratory distress, pain, and disorientation in both humans and animals. In the inner ear, gas embolism can cause hemorrhages, leading to disorientation or vestibular dysfunction.
Breath-holding divers, like beaked whales, can develop decompression-related problems (the "bends") when they return to the surface after deep dives. This is a possible hypothesis for the mass strandings of pelagic beaked whales associated with sonar-related activities. To illustrate, a diving beaked whale may be surfacing from a deep dive and must pass vertically through varying received sound levels. Since the whale has limited remaining oxygen supplies at the end of a long dive, it probably has limited abilities to display any normal sound avoidance behavior. Instead, the whale must continue to swim towards the surface to replenish its oxygen stores. Avoiding sonar inevitably requires a change in behavior or surfacing pattern. Therefore, sonar in close proximity to groups of beaked whales has the potential to cause hemorrhaging or to disorient the animal, eventually leading to a stranding.
Current research reveals two species of beaked whales are most affected by sonar: Cuvier's (Z. cavirostris) and Blainville's (M. densirostris) beaked whales. These animals have been reported as stranding in correlation with military exercises in Greece, the Bahamas, Madeira, and the Canary Islands. The livers of these animals had the most damage.
Though some evidence indicates sonar-related activities can actually lead to a form of decompression sickness in beaked whales, the topic is still up for debate. A significant limiting factor in determining the likelihood of bubble formation in whale tissues and the risk of decompression sickness is the lack of information on their normal diving and surfacing patterns. More research is necessary to determine the extent of whale tissue damage caused by sonar exercises.
Four species are classified by the IUCN as "lower risk, conservation dependent": Arnoux's and Baird's beaked whales, and the northern and southern bottlenose whales. The status of the remaining species is unknown, preventing classification.
- Tyack, Peter L.; Johnson, Mark; Soto, Natacha Aguilar; Sturlese, Albert; Madsen, Peter T. (2006). "Extreme diving of beaked whales" (PDF). Journal of Experimental Biology. 209 (21): 4238–4253. doi:10.1242/jeb.02505. PMID 17050839. Retrieved January 2014. Check date values in:
- Schorr, Gregory S.; Falcone, Erin A.; Moretti, David J.; Andrews, Russel D. (2014). "First Long-Term Behavioral Records from Cuvier's Beaked Whales (Ziphius cavirostris) Reveal Record-Breaking Dives". PLOS ONE. 9: e92633. doi:10.1371/journal.pone.0092633. PMID 24670984.
- Bianucci, Giovanni; Post, Klaas; Lambert, Olivier (2008). "Beaked whale mysteries revealed by seafloor fossils trawled off South Africa". South African Journal of Science. 104 (3–4): 140–142. Retrieved January 2014. Check date values in:
- Christensen, Ivar (1984). Macdonald, David W., ed. The Encyclopedia of Mammals. New York: Facts on File. pp. 210–211. ISBN 0-87196-871-1. OCLC 10403800.
- Rommel, S. A.; Costidis, A. M.; Fernandez, A.; Jepson, P. D.; Pabst, D. A.; McLellan, W. A.; Houser, D. S.; Cranford, T. W.; van Helden, A. L.; Allen, D. M.; Barros, N. B. (2006). "Elements of beaked whale anatomy and diving physiology and some hypothetical causes of sonar-related stranding" (PDF). Journal of Cetacean Research and Management. IWC. 7 (3): 189–209. Retrieved January 2014. Check date values in:
- Litchfield, Carter; Greenberg, Anne J.; Caldwell, David K.; Caldwell, Maria C.; Sipos, J. C.; Ackman, R. G. (1975). "Comparative lipid patterns in acoustical and nonacoustical fatty tissues of dolphins, porpoises and toothed whales". Comp. Biochem. Physiol. B. 50 (4): 591–7. doi:10.1016/0305-0491(75)90095-4. OCLC 733963359. PMID 1122741.
- Dalebout, Merel L.; Steel, Debbie; Baker, C. Scott (2008). "Phylogeny of the Beaked Whale Genus Mesoplodon (Ziphiidae: Cetacea) Revealed by Nuclear Introns: Implications for the Evolution of Male Tusks" (PDF). Systematic Biology. 57 (6): 857–875. doi:10.1080/10635150802559257. PMID 19085329. Retrieved January 2014. Check date values in:
- Lambert, Olivier (2006). "Archaeoziphius microglenoideus, a new primitive beaked whale (Mammalia, Cetacea, Odontoceti) from the Middle Miocene of Belgium". Journal of Vertebrate Paleontology. 26: 182–191. doi:10.1671/0272-4634(2006)26[182:AMANPB]2.0.CO;2. Retrieved 2 August 2010.
- Dalebout, Merel L. (2014). "Resurrection of Mesoplodon hotaula Deraniyagala 1963: A new species of beaked whale in the tropical Indo-Pacific". Marine Mammal Science. 30: 1081–1108. doi:10.1111/mms.12113. Retrieved 5 February 2014.
- Bianucci, Giovanni; Miján, Ismael; Lambert, Olivier; Post, Klaas; Mateus, Octávio (2013). "Bizarre fossil beaked whales (Odontoceti, Ziphiidae) fished from the Atlantic Ocean floor off the Iberian Peninsula" (PDF). Geodiversitas. 35 (1): 105–153. doi:10.5252/g2013n1a6. Retrieved January 2014. Check date values in:
- Zimmer, Walter M. X.; Tyack, Peter L. (2007). "Repetitive shallow dives pose decompression risk in deep-diving beaked whales". Marine Mammal Science. 23 (4): 888–925. doi:10.1111/j.1748-7692.2007.00152.x.
- MacLeod, Colin D.; D'Amico, Angela (2006). "A review of beaked whale behavior and ecology in relation to assessing and mitigating impacts of anthropogenic noise" (PDF). Journal of Cetacean Research and Management. IWC. 7 (3): 211–221. Retrieved January 2014. Check date values in:
- Auster, Peter J.; Watling, Les (2010). "Beaked whale foraging areas inferred by gouges in the seafloor". Marine Mammal Science. 26 (1): 226–233. doi:10.1111/j.1748-7692.2009.00325.x.
- MacLeod, Colin D.; Perrin, William F.; Pitman, Robert; Barlow, Jay; Ballance, Lisa; D'Amicon, Angela; Gerrodette, Tim; Joyce, Gerald; Mullin, Keith D.; Palka, Debra L.; Waring, Gordon T. (2006). "Known and inferred distributions of beaked whales" (PDF). Journal of Cetacean Research and Management. IWC. 7 (3): 271–286. Retrieved January 2014. Check date values in:
- Mead, James G. (1984). "Survey of reproductive data for the beaked whales (Ziphiidae)" (PDF). Reports of the International Whaling Commission (Special Issue 6): 91–96. Retrieved January 2014. Check date values in:
|access-date=(help) (PDF from IWC)
- Law, R. J.; Allchin, C. R.; Jones, B. R.; Jepson, P. D.; Baker, J. R.; Spurrier, C. J. H. (1997). "Metals and organochlorines in tissues of a Blainville's beaked whale (Mesoplodon densirostris) and a killer whale (Orcinus orca) stranded in the United Kingdom". Marine Pollution Bulletin. 34 (3): 208–212. doi:10.1016/S0025-326X(96)00148-8.
- Fertl, D.; Leatherwood, S. (1997). "Cetacean interactions with trawls: a preliminary review" (PDF). Journal of Northwest Atlantic Fishery. 22: 219–248. doi:10.2960/j.v22.a17. Retrieved January 2014. Check date values in:
- Fernández, A.; Edwards, J. F.; Rodríguez, F.; Espinosa de los Monteros, A.; Herráez, P.; Castro, P.; Jaber, J. R.; Martín, V.; Arbelo, M. (2005). ""Gas and Fat Embolic Syndrome" involving a mass stranding of beaked whales (family Ziphiidae) exposed to anthropogenic sonar signals" (PDF). Veterinary Pathology. 42 (4): 446–457. doi:10.1354/vp.42-4-446. PMID 16006604. Retrieved January 2014. Check date values in:
- Faerber, Meghan M.; Baird, Robin W. (2010). "Does a lack of observed beaked whale strandings in military exercise areas mean no impacts have occurred? A comparison of stranding and detection probabilities in the Canary and main Hawaiian Islands" (PDF). Marine Mammal Science. 26 (3): 602–613. doi:10.1111/j.1748-7692.2010.00370.x. Retrieved January 2014. Check date values in:
- Jepson, P. D.; Arbelo, M.; Deaville, R.; Patterson, I. A. P.; Castro, P.; Baker, J. R.; Degollada, E.; Ross, H. M.; Herráez, P.; Pocknell, A. M.; Rodríguez, F.; Howie, F. E.; Espinosa, A.; Reid, R. J.; Jaber, J. R.; Martin, V.; Cunningham, A. A.; Fernández, A. (2003). "Gas-bubble lesions in stranded cetaceans: was sonar responsible for a spate of whale deaths after an Atlantic military exercise?" (PDF). Nature. 425 (6958): 575–576. doi:10.1038/425575a. PMID 14534575. Retrieved January 2014. Check date values in:
- "Cetacean update of the 2008 IUCN Red List of Threatened Species" (PDF). IUCN. 2008. Retrieved January 2014. Check date values in: