Ziphiidae

Superregnum: Eukaryota
Cladus: Opisthokonta
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Cladus: Deuterostomia
Phylum: Chordata
Subphylum: Vertebrata
Infraphylum: Gnathostomata
Superclassis: Tetrapoda
Classis: Mammalia
Subclassis: Theria
Infraclassis: Placentalia
Superordo: Cetartiodactyla
Ordo: Cetacea
Subordo: Odontoceti
Infraordines: Physeterida
Superfamiliae: Ziphioidea
Familia: Ziphiidae
Subfamiliae: Berardiinae - Hyperoodontinae - Ziphiinae - incertae sedis

Name

Ziphiidae Gray, 1850

References

* Ziphiidae on Mammal species of the World.
Mammal Species of the World: A Taxonomic and Geographic Reference, 2 Volume Set edited by Don E. Wilson, DeeAnn M. Reeder

Vernacular names
Internationalization
English: Beaked whale
日本語: アカボウクジラ科
Türkçe: Gagalı balinagiller
中文: 劍吻鯨科

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Beaked whales are 21 species of toothed whales, members of the family Ziphiidae that are notable for their elongated snouts. They are the only marine mammals whose evolution is believed to have been shaped by a secondary sexual characteristic (the male's teeth). Beaked whales are the world's most extreme divers. They can dive for long periods—20 to 30 minutes is common, and 85 minute dives have been recorded—and to great depths: 1,899 metres (1,038 fathoms) and possibly more.[1] To avoid getting decompression sickness—the potentially fatal build-up of nitrogen bubbles in body tissues—they must surface slowly.[2]

Beaked whales are one of the least known groups of mammals because of their deep-sea habitat, mysterious habits, and apparent low abundance. [3] Several species have yet to be formally described or named; others are known only from remains and have never been sighted alive. Only 3–4 of the 20-odd 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 in the 20th centuries.

Physical characteristics

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 15 LT; 1.1 to 17 ST). 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.[4] However, aside from dentition and size, there are very few morphological differences between male and female beaked whales. [5]

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.

Teeth

Beaked whales are unique among toothed whales in that most species only have one pair of teeth. Most species only have one pair of teeth in the lower jaw, and only males have this pair of tusk-like teeth. Males are presumed to use their teeth in combat for female reproductive rights.[6] In females, the teeth do not develop and remain hidden in the gum tissues. This characteristic helps to distinguish beaked whale species from each other.

In December 2008, researchers from the Marine Mammal Institute at Oregon State University completed a DNA tree of all 21 known species of beaked whales. 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. Females are presumed to select mates based on the shape of the teeth, because the different species are otherwise quite similar in appearance. Females may also choose mates based on the size or shape of teeth or the scars they bear.[6]

These teeth also play an important role in competing to control a harem. This is the only known instance of a secondary sexual characteristic having shaped the evolution of a marine mammal.[7]

Taxonomy

Beaked whales comprise at least twenty species of small whale 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 20 million years ago.

* ORDER CETACEA
* Suborder Odontoceti: toothed whales
o Family Ziphiidae
+ Subfamily Berardiinae
# Genus †Archaeoziphius
# Genus Berardius
* Berardius arnuxii, Arnoux's beaked whale
* Berardius bairdii, Baird's beaked whale
# Genus †Microberardius
+ Subfamily Hyperoodontinae
# Genus †Africanacetus
# Genus Hyperoodon: Bottlenose whales
* Hyperoodon ampullatus, Northern bottlenose whale
* Hyperoodon planifrons, Southern bottlenose whale
# Genus Indopacetus
* Indopacetus pacificus, Longman's beaked whale
# Genus Mesoplodon, Mesoplodont whales
* Mesoplodon bidens, Sowerby's beaked whale
* Mesoplodon bowdoini, Andrew's beaked whale
* Mesoplodon carlhubbsi, Hubbs' beaked whale
* Mesoplodon densirostris, Blainville's beaked whale
* Mesoplodon europaeus, Gervais' beaked whale
* Mesoplodon ginkgodens, Ginkgo-toothed beaked whale
* Mesoplodon grayi, Gray's beaked whale
* Mesoplodon hectori, Hector's beaked whale
* Mesoplodon layardii, Strap-toothed whale
* Mesoplodon mirus, True's beaked whale,
* Mesoplodon peruvianus, pygmy beaked whale
* Mesoplodon perrini, Perrin's beaked whale
* Mesoplodon stejnegeri, Stejneger's beaked whale
* Mesoplodon traversii, Spade toothed whale
+ Subfamily Ziphiinae
# Genus †Caviziphius
# Genus †Izikoziphius
# Genus Tasmacetus
* Tasmacetus sheperdi, Shepherd's beaked whale
# Genus Ziphius
* Ziphius cavirostris, Cuvier's beaked whale
+ Subfamily Incertae sedis
# Genus †Nenga
# Genus †Pterocetus
# Genus †Xhosacetus

Evolutionary history

As many as eight genera predate humans.[3] Some included ancestors of giant beaked whales (Berardius), such as Microberardius. 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 of the South African coast, confirming that 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). [3]

Ecology

Diving

Beaked whales are deep divers with extreme dive profiles [8] They regularly dive deeper than 500 m to echolocate for food, and these deep dives are often followed by multiple shallower dives of less than 500 m.[9] Based on currently available data, beaked whales are thought to spend much of their lives below water.[10]

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.[8] 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 a beaked whale to hold its pectoral flippers tightly against its body for increased streamlining. [5]

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.[9] While the whale is diving, its heart rate slows and blood flow changes. This physiological dive response ensures that oxygen-sensitive tissues maintain a supply of oxygen, while those tissues that are 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. [5]

Feeding

The throats of all beaked whales have a bilaterally paired set of grooves that are associated with their unique feeding mechanism, suction feeding.[5] 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 tongue 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,[6] but also benthic and benthopelagic fish and some crustaceans, mostly taken near the sea floor.[10] In a recent study, gouge marks in the seafloor were interpreted to be a result of feeding activities by beaked whales.[11]

In order to understand the hunting and foraging behavior of beaked whales, researchers utilized 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 approximately 200 and 1885 m and usually catch about 30 prey per dive. Cuvier’s beaked whale must forage on average at 1070 m for 58 minutes and Blainville’s beaked whales typically forage at 835 m deep for an average of 47 minutes.[8]

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.[12] Specific ranges vary greatly by species; though beaked whales typically inhabit offshore waters that are at least 300 meters deep.

Beaked whales are known to congregate in deep waters off the edge of continental shelves, and bottom features like seamounts, canyons, escarpments, and oceanic islands including the Azores and the Canaries.[10]

Life history

Very little is known about the life histories of beaked whales. The oldest beaked whale 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 that have been 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.[13] There is currently no data available on the reproductive rates of beaked whales.

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 1 – 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 in size 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.[10]

There is not much information available about group composition of beaked whales. Only 3-4 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 that 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.[6] Baird’s beaked whales are known to occur in multi-male groups and in large groups, consisting of adult animals of both sexes.

Conservation

For many years most beaked whale species were insulated from anthropogenic impacts because of their remote habitat. However, there are now several issues of concern:

* Studies of stranded beaked whales show rising levels of toxic chemicals in their blubber.[14]
* As a top predator beaked whales are, like raptors, particularly vulnerable to build-up of biocontaminants. They frequently ingest plastic bags (which do not break down and can be lethal).[10]
* They more frequently become trapped in trawl nets, due to the expansion of deepwater fisheries.[15]
* Decompression sickness:

A major conservation concern for beaked whales (family Ziphiidae) is that they appear to be vulnerable to modern sonar operations. This concern arises out of recent strandings that temporally and physically coincide with naval sonar exercises.[16] Postmortem 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.[9] Gas and fat emboli have been shown cause nervous and cardiovascular system dysfunction, respiratory distress, pain, and disorientation in both humans and animals.[16] In the inner ear, gas embolism can cause hemorrhages, leading to disorientation or vestibular dysfunction.

Breath-hold divers, like beaked whales, can develop decompression-related problems (the “bends”) when they return to the surface after deep dives.[8] 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 in order to replenish its oxygen stores.[10] 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 that two species of beaked whales are most affected by sonar: Cuvier’s beaked whales (Ziphius cavirostris) and Blainville’s (Mesoplodon densirostris) beaked whales. These animals have been reported as stranding in correlation with military exercises in Greece, the Bahamas, Madeira, and the Canary Islands.[17] The livers of these animals had the most damage. [18]

Though some evidence indicates that 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 the normal diving patterns and surfacing patterns of beaked whales.[8] 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.[19]

Film

Tracking their silent voices (original title: Auf der leisen Spur der Schnabelwale) is a documentary about an Atlantic scientific expedition. A team of marine biologists tries to find beaked whales and to analyze their behavior using acoustic methods only.[20][vague]

References

1. ^ Lewis Smith (October 17, 2006). "It's official: New free-diving record is 1,899 meters (6,230 ft)". CDNN. http://www.cdnn.info/news/industry/i061017a.html.
2. ^ Does Sonar Disrupt Deep Divers? March 26, 2010. Science (5973): 1561.
3. ^ a b c Bianucci, G., K. Post, and O. Lambert. 2008. "Beaked whale mysteries revealed by seafloor fossils trawled off South Africa", South African Journal of Science 104(3-4):140-142.
4. ^ Christensen, Ivar (1984). Macdonald, D.. ed. The Encyclopedia of Mammals. New York: Facts on File. pp. 210–211. ISBN 0-87196-871-1.
5. ^ a b c d Rommel et al. 2006. Elements of beaked whale anatomy and diving physiology and some hypothetical causes of sonar-related stranding. Journal of Cetacean Resource Management 7(3): 189-209.
6. ^ a b c d Dalebout, M. L., D. Steel, and C. S. Baker. 2008. Phylogeny of the Beaked Whale Genus Mesoplodon (Ziphiidae: Cetacea) Revealed by Nuclear Introns: Implications for the Evolution of Male Tusks. Systematic Biology 57(6):857-875.
7. ^ "Whale's teeth are aid to mating". BBC. December 2008. http://news.bbc.co.uk/2/hi/science/nature/7783517.stm.
8. ^ a b c d e Tyack, P. L. et al. 2006. Extreme diving of beaked whales. The Journal of Experimental Biology 209:4238-4253.
9. ^ a b c Zimmer, W. M. X. and P. L. Tyack. 2007. Repetitive shallow dives pose decompression risk in deep-diving beaked whales. Marine Mammal Science 23(4):888-925.
10. ^ a b c d e f MacLeod, C. D. and A. D’Amico. 2006. A review of beaked whale behavior and ecology in relation to assessing and mitigating impacts of anthropogenic noise. Journal of Cetacean Restoration and Management 7(3):211-221.
11. ^ Auster, P. J. and L. Watling. 2010. Beaked whale foraging areas inferred by gouges in the seafloor. Marine Mammal Science 26(1):226-233.
12. ^ MacLeod, C. D. et al. 2006. Known and inferred distributions of beaked whales. Journal of Cetacean Restoration and Management 7(3):271-286.
13. ^ Mead, J. G. 1984. Survey of reproductive data for the beaked whales (Ziphiidae),. Rep. int. Whal. Commn (special issue), 6:91-6.
14. ^ Law, R. J., Allchin, C. R., Jones, B. R., Jepson, P. D., Baker, J. R. and 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 34:208.
15. ^ Fertl, D. and S. Leatherwood. 1997. Cetacean interactions with trawls: a preliminary review. Journal of Northwest Atlantic Fishery 22:219-248.
16. ^ a b Fernandez, A. et al. 2005. ‘‘Gas and Fat Embolic Syndrome’’ involving a mass stranding of beaked whales (family Ziphiidae) exposed to anthropogenic sonar signals. Veterinary Pathology 42:446-457.
17. ^ Faerber, M. M. and R.W. Baird. (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. Marine Mammal Science 26(3):602-613.
18. ^ Jepson, P. D. et al. 2003. Gas bubble lesions in stranded cetaceans- was sonar responsible for a spate of whale deaths afteram Atlandtic military exercise? Nature 425:575-576.
19. ^ http://cmsdata.iucn.org/downloads/cetacean_table_for_website.pdf
20. ^ Documentary, 2010, 44 Min., Germany. Production: Bayerischer Rundfunk, Germany