Giant otter: Difference between revisions
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==Why Did YOU DELETE MY ARTIClE== |
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{{Taxobox |
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| name = Giant Otter<ref name=msw3>{{MSW3 Wozencraft | pages = 605}}</ref> |
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| status = EN |
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| trend = unknown |
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| status_system = iucn3.1 |
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| status_ref = <ref name=iucn>{{IUCN2006|assessors=Groenendijk ''et al''|year=2004|id=18711|title=Pteronura brasiliensis|downloaded=11 May 2006}} Database entry includes justification for why this species is endangered</ref> |
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| image = giantotter.jpg |
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| image_caption = | image_width = 250px |
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| regnum = [[Animal]]ia |
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| phylum = [[Chordata]] |
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| classis = [[Mammal]]ia |
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| ordo = [[Carnivora]] |
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| familia = [[Mustelidae]] |
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| subfamilia = [[Lutrinae]] |
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| genus = '''''Pteronura''''' |
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| species = '''''P. brasiliensis''''' |
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| binomial = ''Pteronura brasiliensis'' |
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| binomial_authority = ([[Johann Friedrich Gmelin|Gmelin]], 1788) |
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| range_map = Leefgebied reuzenotter.jpg}} |
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The '''Giant Otter''' (''Pteronura brasiliensis'') is a mammalian [[carnivore]] native to [[South America]]. It is the longest member of the [[Mustelidae]], or weasel family, a globally successful group of predators. Unusually for a mustelid, the Giant Otter is a social species, with family groups typically supporting three to eight members. The groups are centred around a dominant breeding pair and are extremely cohesive and cooperative. Although generally peaceful, the species is territorial and aggression has been observed between groups. The Giant Otter is [[Diurnality|diurnal]], being exclusively active during daylight hours. It is the noisiest otter species and distinct vocalizations have been documented that indicate alarm, aggressiveness, and reassurance. |
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YOU HAVE RUINED ME YOU DICKSLAPS!!!!! |
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The Giant Otter ranges across north-central South America. Its distribution has been greatly reduced and is now discontinuous. The species was listed as [[endangered]] in 1999 and population estimates are typically below 5,000 in the wild. The [[Guiana]]s are the last real stronghold for the species. Decades of [[poaching]] for its velvety pelt, peaking in the 1950s and 1960s, decimated population numbers. Habitat degradation and loss is the greatest current threat. The Giant Otter is also rare in captivity: as of 2003, only 60 animals were held.<ref name=Cali>{{cite journal |last=Londono |first=G. Corredor |coauthors=Munoz, N. Tigreros |year=2006 |title=Reproduction, behaviour and biology of the Giant river |
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otter (''Pteronura brasiliensis'') at Cali Zoo |journal=International Zoo Yearbook |volume=40 |pages=360–371 |url= |doi=10.1111/j.1748-1090.2006.00360.x }}</ref> |
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The Giant Otter shows a variety of adaptations suitable to an amphibious lifestyle, including exceptionally dense fur, a wing-like tail, and webbed feet. The species prefers freshwater rivers and streams, which are usually seasonally flooded, and may also take to freshwater lakes and springs. It constructs extensive campsites close to feeding areas, clearing large amounts of vegetation. The Giant Otter largely subsists on a [[piscivore|diet of fish]], particularly [[characin]]s and [[catfish]], and may also take [[crab]]s. It has no serious natural predators, although it must compete with other species, including the [[Neotropical Otter]] and [[caiman]] species, for food resources. |
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==Naming== |
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The Giant Otter has a handful of other names in English. '''River Wolf''' ({{lang-es|Lobo del Río}}) and '''Water Dog''' (Spanish: ''Perro del Agua'') are used occasionally. The last of these may have been more common in the reports of explorers in the nineteenth and early twentieth centuries.<ref> See e.g., Duplaix, pg. 547.</ref> All three names are in use in Spanish and [[Portuguese language|Portuguese]], with a number of regional variations. "Giant Otter" translates as ''Nutria Gigante'' and ''Lontra Gigante'' in Spanish and Portuguese, respectively; a fourth name, ''Ariraí'' or ''Ariranha'' is also in use in South America.<ref name=FAO/> Among the [[Achuar]] people, they are known as ''Wankanim'',<ref name=Descola/> and among the [[Sanumá]] as ''Hadami''.<ref>{{cite book |title=Sanuma Memories: Yanomami Ethnography in Times of Crisis |last=Ramos |first=Alcida Rita |year=1995 |publisher=University of Wisconsin Press|isbn=0299146545 |pages=219 }}</ref><ref>{{cite journal |year=1981–1982 |title= |journal=Antropológica |volume=55–58 |pages=107 | publisher =Sociedad de Ciencias Naturales La Salle (Fundación La Salle de Ciencias Naturales)}}</ref> The genus name, '''''Pteronura''''', is derived from the [[Ancient Greek]] words ''pteron''/πτερον 'feather' or 'wing' and ''ura''/ουρά 'tail',<ref name="Liddell 1980">{{cite book | author = [[Henry George Liddell|Liddell, Henry George]] and [[Robert Scott (philologist)|Robert Scott]] | year = 1980 | title = [[A Greek-English Lexicon]] (Abridged Edition) | publisher = [[Oxford University Press]] | location = United Kingdom | id = ISBN 0-19-910207-4}}</ref> a reference to its distinctive wing-like tail.<ref name=Koepfli/> |
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==Taxonomy and evolution== |
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[[Image:Pteronura brasiliensis MG 9021.jpg|thumb|Giant Otter from the [[Museu Paraense Emílio Goeldi]] research institute]] |
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The otters form the [[Lutrinae]] subfamily within the mustelids and the Giant Otter is the [[monotypic|only member]] of the genus ''Pteronura''. Two subspecies are currently recognized by the canonical [[Mammal Species of the World]], ''P. b. brasiliensis'' and ''P. b. paraguensis''. Incorrect descriptions of the species have led to multiple synonyms (the latter subspecies is often ''P. b. paranensis'' in the literature).<ref name=msw3/> ''P. b. brasiliensis'' is distributed across the north of the Giant Otter range, including the [[Orinoco]], [[Amazon River|Amazon]], and Guianas river systems; to the south, ''P. b. paraguensis'' has been suggested in [[Paraguay]], [[Uruguay]], southern [[Brazil]], and northern [[Argentina]],<ref name=FAO/> although it may be extinct in the last three of these four. The [[World Conservation Union]] (IUCN) considers the species' presence in Argentina and Uruguay uncertain.<ref name=iucn/> In the former, investigation has shown thinly distributed population remnants.<ref>{{cite journal |last=Chehebar |first=C.|title=Searching for the Giant Otter in Northeastern Argentina |journal=IUCN Otter Specialist Group |volume=6 |issue=1 |pages=17–18 |month=February | year=1991 | url=http://www.otterspecialistgroup.org/Bulletin/Volume6/Chehebar_1991.html |accessdate= 2007-11-06}}</ref> ''P. b. paraguensis'' is supposedly smaller and more gregarious, with different [[dentition]] and skull [[morphology (biology)|morphology]]. Carter and Rosas, however, rejected the subspecific division in 1997, noting that the classification had only been validated once, in 1968, and that the ''P. b. paraguensis'' type specimen was very similar to ''P. b. brasiliensis''.<ref name=C4/> Biologist Nicole Duplaix calls the division of "doubtful value."<ref>Duplaix, pg. 511.</ref> |
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An extinct genus, ''[[Satherium]]'', is believed to be ancestral to the present species, having migrated to the New World during the [[Pliocene]] or early [[Pleistocene]].<ref name=Koepfli>{{cite journal |last= Koepfli|first=K.-P |authorlink= |coauthors= Wayne, R.K.|year= 1998|month=December |title=Phylogenetic relationships of otters (Carnivora: Mustelidae) based on mitochondrial cytochrome b sequences |journal=Journal of Zoology |volume= 246|issue=4 |pages=401–416 |url=http://www.blackwell-synergy.com/doi/abs/10.1111/j.1469-7998.1998.tb00172.x|accessdate=2007-12-29 |quote= |doi=10.1111/j.1469-7998.1998.tb00172.x |format=abstract }}</ref> The Giant Otter shares the South American continent with three of the four members of the ''[[Lontra]]'' genus of otters: the [[Neotropical River Otter]], the [[Southern River Otter]], and the [[Marine Otter]].<ref name=actionplan>{{cite journal|url=http://www.iucn.org/dbtw-wpd/html/Otter/cover.html|title=Otters: An Action Plan for their Conservation |publisher=[[International Conservation Union]]|year=1990|author=Foster-Turley, Pat; Macdonald, Sheila; Mason, Chris (eds.)|journal=IUCN/SSC Otter Specialist Group |doi=10.2305/IUCN.CH.1990.SSC-AP.3.en|pages=Sections 2 and 12 |accessdate= 2007-11-21|doi_brokendate=2008-06-23}}</ref> It seems to have evolved independently of ''Lontra'' in South America, despite the overlap. The [[Smooth-coated Otter]] (''Lutrogale perspicillata'') of Asia may be its closest extant relative: similar behaviour, vocalizations, and skull morphology have been noted.<ref name=Koepfli/> Both species also show strong [[pair bonding]] and paternal engagement in rearing cubs.<ref>Duplaix, pg. 614.</ref> |
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[[Phylogenetic]] analysis by Koepfli and Wayne in 1998 found that the Giant Otter has the highest divergence sequences within the otter subfamily, forming a distinct [[clade]] that split away 10 to 14 million years ago. They noted that the species may be the [[Basal (phylogenetics)|basal]] divergence among the otters or fall outside of them altogether, having split even before other mustelids, such as the [[Ermine]], [[Mustelinae|Polecat]], and [[Mink]].<ref name=Koepfli/> Later [[gene sequencing]] research on the mustelids, from 2005, places the divergence of the Giant Otter somewhat later, between 5 and 11 million years ago; the corresponding [[phylogenetic tree]] locates the ''Lontra'' divergence first amongst otter genera, and ''Pteronura'' second, although divergence ranges overlap.<ref>{{cite journal |last=Marmi |first=Josep |coauthors= Lopez-Giraldez, Juan F.; Domingo-Roura, Xavier |year=2004 |month= November|title=Phylogeny, evolutionary history and taxonomy of the Mustelidae based on sequences of the cytochrome b gene and a complex repetitive flanking region |journal=Zoologica Scripta |volume= 33|issue=6 |pages= 481–499|doi=10.1111/j.0300-3256.2004.00165.x}}</ref> |
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==Biology and behaviour== |
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The Giant Otter is large, gregarious, and diurnal (active through the day). Early travellers' reports describe noisy groups surrounding explorers' boats but little scientific information was available on the species until Duplaix's groundbreaking work in the late 1970s.<ref>Duplaix, pg. 497.</ref> Concern over this endangered species has since generated a corpus of research. |
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===Physical characteristics=== |
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The Giant Otter is clearly distinguished from other otters by morphological and behavioural characteristics. It has the greatest body length of any species in the mustelid family, although the [[Sea Otter]] may be heavier. Males are between 1.5 and 1.8 meters (4.9–5.9 feet) in length and females between 1.5 and 1.7 m (4.9–5.6 ft). Early reports of skins and living animals suggested exceptionally large males of up to 2.4 m (7.9 ft); intensive hunting likely reduced the occurrence of such massive specimens. Weights are between 32 and 45.3 kilograms (70–100 pounds) for males and 22 and 26 kg (48–57 lbs) for females.<ref name=C2>Carter and Rosas, pg. 2.</ref> |
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The Giant Otter has the shortest fur of all otter species; it is typically chocolate brown but may be reddish or [[fawn (color)|fawn]], and appears nearly black when wet.<ref name=C2/> The fur is extremely dense, so much so that water cannot penetrate to the skin.<ref name=Philly>{{cite web |url=http://www2.philadelphiazoo.org/zoo/Meet-Our-Animals/Mammals/Carnivores/Giant-River-Otter.htm |title=Giant Otter |accessdate=2007-11-06|work=Meet Our Animals |publisher=[[Philadelphia Zoo]]}}</ref> [[Guard hair]]s trap water and keep the inner fur dry; the guard hairs are approximately 8 millimeters (one third of an inch) in length, about twice as long as the fur of the inner coat.<ref name=Seaworld>{{cite web |url=http://www.seaworld.org/animal-info/info-books/otters/physical-characteristics.htm |title=Otters: Physical characteristics |accessdate=2007-11-06 |publisher=Anheuser-Busch Adventure Parks}}</ref> Its velvety feel makes the animal highly sought after by fur traders and has contributed to its decline.<ref name=EE>{{cite web |url=http://www.earthsendangered.com/profile.asp?ID=11&sp=311 |title=Giant Otter Facts |accessdate=2007-11-07|work=Meet Our Animals |publisher=Earth's Endangered Creatures}}</ref> Unique markings of white or cream fur colors the throat and under the chin, allowing individuals to be identified from birth.<ref name=C2/> Its well-muscled tail can account for as much as 69 centimeters (27 in) of total body length.<ref>{{cite web |url=http://www.nature.org/animals/mammals/animals/giantotter.html |title= Giant Otter (''Pteronura brasiliensis'') |accessdate=2008-01-25 |publisher=[[The Nature Conservancy]] }}</ref> |
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Giant Otter muzzles are short and sloping and give the head a ball-shaped appearance.<ref name=D511>Duplaix, pg. 511.</ref> The ears are small and rounded.<ref name=Philly/> The nose (or [[rhinarium]]) is completely covered in fur, with only the two slit-like nostrils visible. The Giant Otter's highly sensitive whiskers ([[vibrissae]]) allow the animal to track changes in water pressure and currents, which aids in detecting prey.<ref name=WWF>{{cite web |url=http://www.panda.org/news_facts/education/best_place_species/current_top_10/giant_otter.cfm |title=Giant Otter |accessdate=2008-01-19 |publisher=[[World Wildlife Fund]] |work= }}</ref> The legs are short and stubby and end in large webbed feet tipped with sharp claws. Well suited for an aquatic life, it can close its ears and nose while underwater.<ref>{{cite web |url= http://www.iwokrama.org/forest/animals/giantotter.htm |title=Giant Otter, the "Water Dog" |accessdate=2007-11-07 |publisher= Iwokrama International Centre for Rainforest Conservation and Development }}</ref> |
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As of Carter and Rosas' writing, vision had not been directly studied but field observations show that the animal primarily hunts by sight; above water, it is able to recognize observers at great distances. The fact that it is exclusively active during the day further suggests its eyesight should be strong, to aid in hunting. In other otter species vision is generally [[emmetropic|normal]] or slightly [[myopic]], both on land and in water. Giant Otter hearing is acute and its sense of smell excellent.<ref name=D533>Duplaix, pg. 533.</ref><ref name=C2/> |
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===Vocalizations=== |
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The Giant Otter is an especially noisy animal, with a complex repertoire of vocalizations. All otters produce vocalizations, but by frequency and volume, the Giant Otter may be the most vocal.<ref>{{cite web|publisher=[[Seaworld]]|url=http://www.seaworld.org/Animal-info/info-books/otters/pdf/ib-otters.pdf|title=Otters: A SeaWorld Education Department Publication|year=2005|accessdate=2008-01-23|format=PDF}}</ref> Duplaix identified nine distinct sounds, with further subdivisions possible, depending on context. Quick ''HAH!'' barks or explosive snorts suggest immediate interest and possible danger. A wavering scream may be used in bluff charges against intruders, while a low growl is used for aggressive warning. Hums and coos are more reassuring within the group. Whistles may be used as advance warning of non-hostile intent between groups, although evidence is limited. Newborn cubs squeak to elicit attention, while older young whine and wail when they begin to participate in group activities.<ref>Duplaix, pp. 552–561.</ref> |
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===Social structure=== |
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The Giant Otter is a highly social animal and lives in extended family groups. Group sizes are anywhere from two to twenty members but likely average between three and eight.<ref name=FAO>{{cite web |url=http://www.fao.org/docrep/t0750e/t0750e0i.htm |title=Pteronura brasiliensis (giant otter) |accessdate=2007-11-07 |publisher=Food and agricultural organization of the United Nations |work=Carnivores }}</ref> (Larger figures may reflect two or three family groups temporarily feeding together.)<ref name=vandamme2002/> The groups are strongly cohesive: the otters sleep, play, travel, and feed together. |
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[[Image:Giant otter 3.jpg|thumb|left|Giant Otters leave a pool together at the [[Philadelphia Zoo]]. The species is extremely social, a rarity amongst mustelids, and family groups are cohesive.]] |
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Group members share roles, structured around the dominant breeding pair. The species is territorial, with groups marking their ranges with latrines, gland secretions, and vocalizations.<ref name=Ribas05>{{cite journal |last=Ribas |first=Carolina |authorlink= |coauthors=Mourão, Guilherme |year=2005 |month=January |title=Intraspecific Agonism between Giant Otter Groups |journal=IUCN Otter Specialist Group Bulletin|volume=21 |issue=2 |pages=89–93 |url=http://iucnosg.org/Bulletin/Volume21/Ribas_Mourao_2004.html |accessdate= 2007-11-07 }}</ref> At least one case of a change in alpha relationship has been reported, with a new male taking over the role; the mechanics of the transition were not determined.<ref name=Evan07/> Duplaix suggests a division between ''residents'', who are established within groups and territories, and nomadic and solitary ''transients''; the categories do not seem rigid, and both may be a normal part of the Giant Otter life cycle.<ref>Duplaix, pp. 571–572.</ref> One tentative theory for the development of sociality in mustelids is that locally abundant but unpredictably dispersed prey causes groups to form.<ref name=Johnson00/> |
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Aggression within the species ("intraspecific" conflict) has been documented. Defense against intruding animals appears to be cooperative: while adult males typically lead in aggressive encounters, cases of alpha females guarding groups has been reported.<ref name=Evan07>{{cite journal |last= Evangelista |first=Emanuela |year=2004 |month=July |title=Change Of Partners In A Giant Otter Alpha Couple |journal=IUCN Otter Specialist Group Bulletin|volume=21 |issue=1 |pages=47–51 |url=http://www.otterspecialistgroup.org/Bulletin/Volume21/Evangelista_2004.html |accessdate= 2007-11-07 }}</ref> One fight was directly observed in the Brazilian [[Pantanal]] in which three animals violently engaged a single individual near a range boundary.<ref name=Ribas05/> In another instance in Brazil, a carcass was found with clear indications of violent assault by other otters, including bites to the snout and genitals, an attack pattern similar to that exhibited by captive animals.<ref>{{cite journal |last=Rosas |first=F.C.W. |coauthors= De Mattos, G.E. |year=2003 |month= October|title=Natural Deaths Of Giant Otters (''Pteronura Brasiliensis'') In Balbina Hydroelectric Lake, Amazonas, Brazil |journal=IUCN Otter Specialist Group Bulletin |volume=20 |issue=2 |pages=62–64 |url=http://www.otterspecialistgroup.org/Bulletin/Volume20/Weber_Rosas_de_Mattos_2003.html |accessdate= 2007-11-07}}</ref> While not rare amongst large predators in general, intraspecific aggression is uncommon amongst otter species; Ribas and Mourão suggest a correlation to the animal's sociability, which is also rare amongst other otters.<ref name=Ribas05/> |
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<!--http://iucnosg.org/Bulletin/Volume18/Groenendijk_et_al_2001.html-->A capacity for aggressive behaviour should not be overstated with the Giant Otter. Researchers emphasize that even between groups, conflict avoidance is generally adopted.<ref>Duplaix, pg. 563.</ref><ref name = C15/> Within groups, the animals are extremely peaceful and cooperative. Group hierarchies are not rigid and the animals easily share roles.<ref>{{cite journal |last=Schenck |first=C. |coauthors=Staib, E. |year=1992 |month=April |title=Giant Otters In Peru |journal=IUCN Otter Specialist Group |volume=7 |pages= 24–26|url=http://www.otterspecialistgroup.org/Bulletin/Volume7/Schenk_Staib_1992.html |accessdate= 2008-01-22}}</ref> |
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===Reproduction and life cycle=== |
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[[Image:Giant otter 2.JPG|thumb|right|Captive Giant Otters have provided much of the research on the species' reproduction and life cycle.]] |
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Details of Giant Otter reproduction and life cycle are scarce, and captive animals have provided much of the information. Females appear to give birth year round, although in the wild births may peak during the dry season. The [[estrous]] cycle is 21 days, with females receptive to sexual advances between 3 and 10 days.<ref name=C18>Carter and Rosas, pg. 18.</ref> Study of captive specimens has found that only males initiate copulation.<ref name=Cali/> At [[Tierpark Hagenbeck]] in Germany, long-term pair bonding and individualized mate selection were seen, with copulation most frequently taking place in water.<ref name=Germany/> Females have a gestation period of 65 to 70 days, giving birth to one to five pups, with an average of two.<ref name=Germany>{{cite journal |last=Hagenbeck |first=Carl |coauthors=Wunnemann, Claus |year=1992 |title=Breeding the Giant otter (''Pteronura brasiliensis'') at Carl Hagenbecks Tierpark |journal=International Zoo Yearbook |volume=32 |pages=240–245 }}</ref><ref name=C18/> Research over five years on a breeding pair at the [[Cali Zoo]] in Colombia found that the average interval between litters was six to seven months, but as short as 77 days when the previous litter did not survive.<ref name=Cali/> Other sources have found greater intervals, with as long as 21 to 33 months suggested for the wild.<ref name=C18/> |
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Mothers give birth to furred and blind cubs in an underground den near to the river shore and fishing sites.<ref>Duplaix, pg. 567.</ref> Males actively participate in rearing cubs and family cohesion is strong;<ref>Duplaix, pg. 576</ref> older, juvenile siblings also participate in rearing, although in the weeks immediately after birth they may temporarily leave the group.<ref name=C18/> Cubs open their eyes in their fourth week, begin walking in their fifth, and are able to swim confidently between 12 and 14 weeks.<ref name=Cali/> They are weaned by nine months and begin hunting successfully soon after.<ref name=C18/> The animal reaches sexual maturity at about two years of age and both male and female cubs leave the group permanently after two to three years.<ref name=Germany/><ref name=C18/> They then search for new territory to begin a family of their own.<ref name=Dortmund/> |
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The Giant Otter is very sensitive to human activity when rearing its young. No institution, for example, has successfully raised Giant Otter cubs unless parents were provided sufficient privacy measures; the stress caused by human visual and acoustic interference can lead to neglect, abuse and [[Infanticide (zoology)|infanticide]], as well as decreased [[lactation]]. In the wild, it has been suggested, although not systematically confirmed, that tourists cause similar stresses: disrupted lactation and denning, reduced hunting, and habitat abandonment are all risks.<ref name=Dortmund/> This sensitivity is matched by a strong protectiveness towards the young. All group members may aggressively charge intruders, including boats with humans in them.<ref>Duplaix, pp. 564–565, 570.</ref> |
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The longest documented Giant Otter lifespan in the wild is eight years. In captivity this may increase to 17, with an unconfirmed record of 19. The animal is susceptible to a variety of diseases, including the [[canine parvovirus]]. Parasites, such as the larvae of flies and a variety of intestinal worms, also afflict the Giant Otter.<ref name=Dortmund>{{cite book |title= International Giant Otter Studbook Husbandry and Management Information and Guidelines |last=Sykes-Gatz |first=Sheila |year=2005 |publisher=Zoologischer Garten Dortmund |location=Germany |edition=Second |pages=13 }}</ref> Other causes of death include accidents, [[gastroenteritis]], infanticide, and [[epileptic]] seizures.<ref name=C18/> |
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===Hunting and diet=== |
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[[Image:Riesenotter.jpg|right|thumb|A captive Giant Otter, feeding. This predator grasps prey in its forepaws and begins eating immediately, at the head.]] |
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The Giant Otter is an [[apex predator]] and its population status reflects the overall health of riverine ecosystems.<ref name=Barnett00>{{cite journal |last=Barnett |first=Adrian |coauthors=Shapley, Rebecca; Lehman, Shawn; et al. |year=2000 |month=October |title=Records of the Giant Otter, ''Pteronura brasiliensis'', from Guyana |journal=IUCN Otter Specialist Group |volume=17 |issue=2 |pages= 65–74}}</ref> It feeds mainly on fish, including [[cichlids]], [[characin]]s (such as [[piranha]]), [[catfish]], and [[perch]]. One full year study of Giant Otter scats in Amazonian [[Brazil]] found fish present in all fecal samples. Fish from the [[order (biology)|order]] [[Perciformes]], particularly cichlids, were seen in 97% of scats, and [[Characiformes]], such as characins, in 86%. Fish remains were of medium-sized species that seem to prefer relatively shallow water, to the advantage of the probably visually oriented Giant Otter. Prey species found were also sedentary, generally swimming only short distances, which may aid the Giant Otter in predation. The Giant Otter seems to be opportunistic, taking whatever species are most locally abundant.<ref>{{cite journal |last=Fernando |first=Rosas |coauthors= Zuanon, Jansen; et al. |year=1999 |month=September |title=Feeding Ecology of the Giant Otter, ''Pteronura brasiliensis'' |journal=Biotropica |volume= 31|issue=3 |pages= 502–506 |url= |doi=10.1111/j.1744-7429.1999.tb00393.x }}</ref> If fish are unavailable it will also take [[crab]]s, [[snake]]s, and even small [[caiman]]s and [[anaconda]]s.<ref name=OSF>{{cite web |url=http://www.otter.org/otterframes.html |title=Giant Otter (''Pteronura brasiliensis'') |accessdate=2007-11-21 |publisher=International Otter Survival Fund }}</ref> |
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The species can hunt singly, in pairs, and in groups, relying on its sharp eyesight to locate prey.<ref name=D544>Duplaix, pg. 544–546</ref> In some cases, supposed cooperative hunting may be incidental, a result of group members fishing individually in close proximity; truly coordinated hunting may only occur where the prey cannot be taken by a single Giant Otter, such as with anacondas and the [[Black Caiman]].<ref name=C15>Carter and Rosas, pg. 15.</ref> The Giant Otter seems to prefer prey fish that are generally immobile on river bottoms in clear water. Prey chase is rapid and tumultuous, with lunges and twists through the shallows and few missed targets. The otter can attack from both above and below, swiveling at the last instant to clamp the prey in its jaws. Giant Otters catch their own food and consume it immediately; they grasp the fish firmly between the forepaws and begin eating noisily at the head.<ref name=D544/> Carter and Rosas have found that captive adult animals consume around 10% of their body weight daily—about {{convert|3|kg|lb|0}}, in keeping with findings in the wild.<ref name=C17>Carter and Rosas, pg. 17.</ref> |
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==Ecology== |
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===Habitat=== |
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The species is amphibious, although primarily terrestrial.<ref name=WWF02>{{cite journal|title=Guianas Rapid River Bio-assessments and Giant Otter Conservation Project |publisher=[[World Wildlife Fund]]|url=http://www.giantotterresearch.com/articles/WWF_Giant_Otter_Report_PDFMini.pdf |format=PDF|first=Nicole|last=Duplaix|year=2002}}</ref> It is found in freshwater rivers and streams, which are generally seasonally flooded. Other water habitats include freshwater springs and permanent freshwater lakes.<ref name=iucn/> Four specific vegetation types were found on one important creek in Suriname: riverbank high forest; floodable mixed marsh and high swamp forest; floodable low marsh forest; and grass islands and floating meadows within open areas of the creek itself.<ref name=WWF02/> Duplaix identifies two critical factors in habitat selection: food abundance, which appears to positively correlate to shallow water, and low sloping banks with good cover and easy access to preferred water types. The Giant Otter seems to choose clear black waters with rocky or sandy bottoms over silty, saline, and white waters.<ref>Duplaix, pp.514–515</ref> |
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Areas adjacent to rivers are used for the construction dens, campsites, and latrines.<ref name=Utreras05>{{cite journal |last=Utreras |first=V. |coauthors= Suárez, E.; Zapata-Ríos, G.; et al. |year=2005 |month=July/December |title=Dry and Rainy Season Estimations of Giant Otter, Pteronura brasiliensis, Home-Range in the Yasuní National Park, Ecuador |journal=The Latin American Journal of Aquatic Mammals |volume= 4|issue=2 |pages=1–4 |url=http://www.giantotterresearch.com/articles/lajamareasdevidadenutriasgi.pdf |accessdate= 2007-11-07 |format=PDF}}</ref> Giant Otters clear significant amounts of vegetation in constructing their campsites. One report suggests maximum areas {{convert|28|m|ft|0|abbr=on}} long and {{convert|15|m|ft|0|abbr=on}} wide, well-marked by scent glands, urine, and feces to signal territory.<ref name=actionplan/> Carter and Rosas find average areas a third this size. Communal latrines are adopted adjacent to the campsites, and dens with a handful of entrances are dug, typically under root systems or fallen trees. One report found between three and eight campsites, clustered around feeding areas. In seasonally flooded areas, the Giant Otter may abandon campsites during the wet season, dispersing to flooded forests in search of prey.<ref name=C13>Carter and Rosas, pg. 13.</ref> Preferred locations may be adopted perennially, often on high ground. These can become quite extensive, including "backdoor" exits into forests and swamps, away from the water.<ref name=WWF02/> Not every site is visited or marked daily, but all are usually patrolled, often by a pair of otters in the morning.<ref>Duplaix, pg. 69.</ref> |
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Research generally takes place in the dry season and an understanding of the species' overall habitat use remains partial. Dry season range size analysis of three otter groups in Ecuador found areas between {{convert|0.45|and|2.79|km2|sqmi|lk=on}}. Habitat requirements and availability were presumed to be dramatically different in the rainy season: range sizes of 1.98 to as much as 19.55 square kilometres (0.76 to 7.55 sq mi) were estimated for the groups.<ref name=Utreras05/> Other researchers suggest approximately {{convert|7|km2|sqmi}} and note a strong inverse correlation between [[social behaviour|sociality]] and home range size; the highly social Giant Otter has smaller home range sizes than would be expected for a species of its mass.<ref name=Johnson00>{{cite journal |last=Johnson |first=Dominic D.P. |coauthors=MacDonald, David W.; Dickman, Amy J. |year=2000 |title=An analysis and review of the sociobiology of the Mustelidae|journal=Mammal review |volume=30 |issue=3&4 |pages=171–196 |format=PDF |url=http://www.princeton.edu/~dominic/2000%20-%20Johnson%20et%20al.%20-%20Mustelids%20-%20Mamm.%20Rev.pdf |accessdate= 2007-11-07 |doi=10.1046/j.1365-2907.2000.00066.x }} See figure three for home range size estimate.</ref> A population density high of {{convert|1.2|/km2|/sqmi|abbr=on}} has been reported in Suriname, with a low of {{convert|0.154|/km2|/sqmi|abbr=on}}<!-- one animal per {{convert|6.5|km2|sqmi|abbr=on}} --> found in Guyana.<ref name=FAO/> |
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===Predation and competition=== |
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[[Image:Piranha1.jpg|thumb|right|[[Characin]]s such as [[piranha]] species are prey for the Giant Otter, but these aggressive fish may also pose a danger. Duplaix speculates that piranhas may bite the Giant Otter.]] |
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Adult Giant Otters have no serious natural enemies, beyond human beings. "Possible and occasional" predation by the [[Jaguar]], [[Cougar]], and anacondas has been suggested by Duplaix, but based on historical reports, not direct observation.<ref>Duplaix, pp. 523, 529.</ref> Cubs are more vulnerable, and may be taken by the Black Caiman and other large predators,<ref name=Dortmund/> although adults are constantly mindful of stray young. The [[Spectacled Caiman]] is another potential competitor, but Duplaix found no conflict with the species in Suriname. When in the water, the Giant Otter faces danger from animals that are not strictly preying upon it: the [[Electric eel]] and [[stingray]]s are potentially deadly if stumbled upon, and piranha may be capable of at least taking bites out of a Giant Otter, as evidenced by scarring on individuals.<ref>Duplaix, pp. 529–530.</ref> |
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Even if without direct predation, the Giant Otter must still compete with other predators for food resources. Duplaix documented interaction with the [[Neotropical Otter]].<ref>''Lontra longicaudis.'' In Duplaix (1980) it was listed as the Guiana Otter under the older binomial ''Lutra enudris''.</ref> While the two species are [[sympatric]] (related, with overlapping ranges) during certain seasons, there appeared to be no serious conflict. The smaller Neotropical Otter is far more shy, less noisy, and less social; at about a third the weight of the Giant Otter, it is more vulnerable to predation and, hence, a lack of conspicuousness is to its advantage. The Neotropical Otter is active during twilight and darkness, reducing the likelihood of conflict with the diurnal Giant Otter.<ref>Duplaix, pp. 527–529.</ref> Its smaller prey, different denning habits, and different preferred water types also reduce interaction.<ref name=C17/> |
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Other species that prey upon similar food resources include the caimans and large fish that are themselves [[piscivore]]s. [[Gymnotid]]s, such as the Electric Eel, and the large [[Silurid]] catfish are among aquatic competitors. Two river dolphins, the [[Tucuxi]] and [[Boto]], might potentially compete with the Giant Otter but different spatial use and dietary preferences suggest minimal overlap.<ref name=C17/> |
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==Conservation status== |
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The IUCN listed the Giant Otter as "endangered" in 1999; it had been considered "[[Vulnerable species|vulnerable]]" under all previous listings from 1982 when sufficient data had first become available. It is regulated internationally under Appendix I of the [[Convention on International Trade in Endangered Species of Wild Fauna and Flora]] (CITES): all trade in specimens and parts is illegal.<ref>{{cite web |url=http://www.cites.org/eng/app/appendices.shtml |title=Appendices I, II and III |accessdate=2008-01-22 |publisher=[[Convention on International Trade in Endangered Species of Wild Fauna and Flora]]}}</ref> |
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===Threats=== |
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The animal faces a variety of critical threats. [[Poaching]] has long been a problem. Statistics show that between 1959 and 1969 Amazonian Brazil alone accounted for 1,000 to 3,000 pelts annually. The species was so thoroughly decimated that the number dropped to just 12 in 1971. The implementation of CITES in 1973 finally brought about significant hunting reductions,<ref name=FAO/> although demand did not disappear entirely: in the 1980s, pelt prices were as high as 250USD on the European market. The threat has been exacerbated by the otters' relative fearlessness and tendency to approach human beings. They are extremely easy to hunt, being active through the day and highly inquisitive.<ref name=Defenders>{{cite web|last=Ridgley|first=Heidi |url=http://www.giantotterresearch.com/articles/GO_article_Defenders_of_Wildlife_Winter_2007.pdf |format=PDF|title=The Wolf of the River |accessdate=2007-11-09 |date=Winter 2007 |publisher=Defenders of Wildlife |work=Defenders Magazine }}</ref> The animal's relatively late sexual maturity and complex social life makes hunting especially disastrous.<ref name=FAO/> |
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More recently, habitat destruction and degradation has become the principal danger and a further reduction of 50% is expected in Giant Otter numbers within the 20 years from 2004 (about the span of three generations of Giant Otters).<ref name=iucn/> Typically, loggers first move into rainforest, clearing the vegetation along riverbanks. Farmers follow, creating depleted soil and disrupted habitats. As human activity expands, Giant Otter home ranges become increasingly isolated. Sub-adults leaving in search of new territory find it impossible to set up family groups.<ref name=Joy>{{cite web |url=http://www.otterjoy.com/otterinfo/pteronura/brasiliensis/brasiliensis_threats.html |title=Threats to the Giant Otter |accessdate=2008-01-25 |publisher=Otterjoy.com |first=Lesley|last=Wright }}</ref> Specific threats from human industry include unsustainable [[mahogany]] logging in parts of the Giant Otter range,<ref name=Defenders/> and concentrations of [[mercury (element)|mercury]] in its diet of fish, a by-product of gold mining.<ref>{{cite journal |last=Fonseca |first=Fabrizio R.D. |coauthors=Malm, Olaf; Waldemarin, Helen F. |title=Mercury levels in tissues of Giant otters (''Pteronura brasiliensis'') from the Rio Negro, Pantanal, Brazil |journal=Environmental Research|volume=98 |pages=368–371 | year=2005 |url=http://www.giantotterresearch.com/articles/Articleariranhas.pdf |format=PDF |accessdate= 2007-11-09 |doi=10.1016/j.envres.2004.11.008}}</ref><ref>{{cite journal |last=Gutleb |first=A.C. |coauthors=Schenck, C; Staib, E |title=Giant otter (''Pteronura brasiliensis'') at risk? Total mercury and methylmercury levels in fish and otter scats, Peru |journal=Ambio |volume=26 |issue=8 |pages=511–514 | year=1997|month=December |url=http://md1.csa.com/partners/viewrecord.php?requester=gs&collection=ENV&recid=4314870&q=Pteronura+brasiliensis&uid=1023712&setcookie=yes |accessdate= 2007-11-09}}</ref> Water pollution from mining, fossil fuel extraction, and agriculture is a serious danger: concentrations of [[pesticide]]s and other chemicals are magnified at each step in the food chain, and can poison top predators like the Giant Otter. |
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Other threats to the Giant Otter include conflict with fishermen, who often view the species as a nuisance ([[#Interactions with indigenious peoples|see below]]). [[Eco-tourism]] also presents challenges: while it raises money and awareness for the animals, by its nature it also increases human effect on the species, both through associated development and direct disturbance in the field.<ref name=Joy/> A number of restrictions on land use and human intrusion are required to properly maintain wild populations. Schenck ''et al.'', who undertook extensive fieldwork in Peru in the 1990s, suggest specific "no-go" zones where the species is most frequently observed, offset by observation towers and platforms to allow viewing. Limits on the number of tourists at any one time, fishing prohibitions, and a minimum safe distance of {{convert|50|m|ft|0|abbr=off}} are proposed to offer further protection.<ref>{{cite journal |last=Schenck |first=Christof |coauthors=Jessica, Groenendijk; et al. |year=1999 |month=April |title=Giant Otter Project In Peru: Field Trip And Activity Report, 1998 |journal=IUCN Otter Specialist Group |volume=16 |issue=1 |pages= 33–43|url=http://www.otterspecialistgroup.org/Bulletin/Volume16/Schenck_et_al_1999.html |accessdate= 2008-01-22}}</ref> |
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===Distribution and population=== |
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The Giant Otter has lost as much as 80% of its South American range.<ref name=Defenders/> While still present in a number of north-central countries, Giant Otter populations are under considerable stress. The IUCN lists [[Bolivia]], [[Brazil]], [[Colombia]], [[Ecuador]], [[French Guiana]], [[Guyana]], [[Paraguay]], [[Peru]], [[Suriname]], and [[Venezuela]] as current range countries.<ref name=iucn/> Given local extinctions, the species' range has become discontinuous.<ref name=FAO/> Total population numbers are difficult to estimate. Research in 1988 suggested 1,000 to 3,000 individuals remain, although this may have been an underestimate.<ref name=C4>Carter and Rosas, pg. 4.</ref> One conservation group suggests 2,000 to 5,000 otters.<ref name=EE/> Populations in Bolivia were once widespread but the country became a "black spot" on distribution maps after poaching between the 1940s and 1970s; a relatively healthy but still small population of 350 was estimated in the country in 2002.<ref name=vandamme2002>{{cite journal |last=van Damme |first=Paul |coauthors=Wallace, Rob; et al. |title=Distribution and Population Status of the Giant Otter Pteronura brasiliensis in Bolivia |journal=IUCN Otter Specialist Group |volume=19 |issue=2 |pages=87–96 |month=October | year=2002 |url=http://www.otterspecialistgroup.org/Bulletin/Volume19/Van_Damme_et_al_2002.html |accessdate= 2007-11-05}}</ref> The species has likely been extirpated from southern Brazil, but in the west of the country decreased hunting pressure in the critical [[Pantanal]] may have led to recolonization; a rough estimate suggests 1,000 animals in the region.<ref name=C8>Carter and Rosas, pg. 8.</ref><ref name=Defenders/> |
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[[Image:LocationSuriname.svg|left|thumb|200px|The [[Guiana]]s are the last real stronghold of the Giant Otter. Tiny [[Suriname]] retains extensive forest cover and many protected areas. It is centred above. [[Guyana]] is immediately to the left and [[French Guiana]] immediately to the right.]] |
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As of 1997, only Suriname, French Guiana, and Guyana (the Guianas) showed strong population numbers.<ref name=C4/> In the last of these, interviews and surveys in 2000 showed relatively widespread populations.<ref name=Barnett00/> Seven years earlier, the IUCN's Otter Specialist Group reported that Colombia was still a stronghold for the species.<ref name=actionplan/> Tiny Suriname, still has significant forest cover and an extensive system of protected areas, much of which protect the Giant Otter.<ref>{{cite web |url= http://www.unesco-suriname.org/natural%20heritage%20in%20suriname.htm|title=Natural Heritage in Suriname |accessdate=2008-01-21 |publisher=[[UNESCO]] |work=Suriname Natcom |accessdate=2007-01-21 }}</ref> Duplaix returned to the country in 2000 and found the Giant Otter still present on the [[Kaburi Creek]], a "jewel" of [[biodiversity]], although increased human presence and land use suggests that, sooner or later, the species may not be able to find suitable habitat for campsites.<ref name=Duplaix2>{{cite web |url=http://www.2docstock.com/Suriname/Reports/Kaburi%20Report/Contents.html |title=A Survey of Kaburi Creek, West Suriname, and its Conservation Implications |first=Nicole|last=Duplaix| coauthors=Lingaard, Marchal; ''et al.'' |accessdate=2008-01-22 |year=2001 |publisher=2DocStock Photography}}</ref> In a report for [[World Wildlife Fund]] in 2002, Duplaix was emphatic about the importance of Suriname and the other Guianas:<ref name=WWF02/> {{cquote|The three Guianas remain the last stronghold of Giant otters in South America, with pristine Giant otter habitat on some rivers and good Giant otter densities overall—still, but for how long? The survival of the Giant otter populations in the Guianas is essential to the survival of this endangered species in South America.}} |
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Other countries have taken a lead in designating protected areas in South America. In 2004, Peru created one of the largest conservation areas in the world, [[Alto Purús National Park]], with an area similar in size to Belgium. The park harbors many endangered plants and animals, including the Giant Otter, and holds the world record for mammal diversity.<ref>{{es icon}} {{cite news|url=http://news.bbc.co.uk/hi/spanish/science/newsid_4399000/4399507.stm|title=Perú creará inmensa reserva amazónica|publisher=BBC Mundo|date=2005-04-01|accessdate=2008-01-07}}</ref><ref>{{cite web|url=http://www.roundriver.org/Peru.html|title=The Alto Purús Conservation Project|publisher=Round River Conservation Studies|accessdate=2008-01-25}}</ref> Bolivia designated wetlands larger than the size of Switzerland as a freshwater protected area in 2001; these are also home to the Giant Otter.<ref>{{cite press release|url=http://www.ramsar.org/wn/w.n.bolivia_wwf3.htm|title=WWF welcomes Latin America's largest freshwater protected area|date=2001-09-18|publisher=The Ramsar Convention on Wetlands|accessdate=2008-01-27}}</ref> |
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==Interactions with indigenous peoples== |
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Throughout its range, the Giant Otter interacts with [[indigenous peoples of the Americas|indigenous groups]], who often practice traditional hunting and fishing. A study of five indigenous communities in Colombia suggests that native attitudes toward the animal are a threat: the otters are often viewed as a nuisance that interferes with fishing, and are sometimes killed. Even when told of the importance of the species to ecosystems and the danger of extinction, interviewees showed little interest in continuing to coexist with the species. Schoolchildren, however, had a more positive impression of the animal.<ref name="Velasco">{{es icon}} Velasco, Diana Marcela (2005). [http://www.giantotterresearch.com/articles/Velasco_Inirida_Colombia.pdf "Estudio preliminar sobre el estado de conservación de la nutria gigante (Pteronura brasiliensis) en la zona de influencia de Inírida (Bajo río Inírida) Guainía, Colombia"] (PDF). Giant Otter Research. Retrieved on [[2008-01-27]].</ref> |
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In Suriname, the Giant Otter is not a traditional prey species for human hunters, which affords some protection.<ref name=Duplaix2/> (One researcher has suggested that the Giant Otter is hunted only in desperation due to its horrible taste.)<ref name=Joy/> The animal sometimes drowns in nets set across rivers and machete attacks by fishermen have been noted, according to Duplaix, but "tolerance is the rule" in Suriname.<ref name=D529>Duplaix, pp. 529–530</ref> One difference in behaviour was seen in the country in 2002: the normally inquisitive Giant Otters showed "active avoidance behavior with visible panic" when boats appeared. Logging, hunting, and cub seizure may have led groups to be far more wary of human activity.<ref name=WWF02/> |
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Local people sometimes take cubs for the [[exotic pet|exotic pet trade]] or as pets for themselves, but the animal rapidly grows to become unmanageable.<ref name=Joy/> Duplaix relates the story of an [[Arawak]] Indian who took two cubs from its parents. While revealing of the affection held for the animal, the seizure was a profound blow to the breeding pair, which went on to lose their territory to competitors.<ref name=D529/> Diane McTurk manages Giant Otter rehabilitation work with rescued and orphaned Giant Otter cubs at Karanambo Ranch in Guyana.<ref>{{cite web|url=http://www.nationalgeographic.co.in/watch/program_details.aspx?id_program=3257|title=Mission: Giant Otters|publisher=National Geographic Channel|accessdate=2008-01-25}}</ref> |
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The species has also appeared in the folklore of the region. It plays an important role in the mythology of the [[Achuar]] people, where Giant Otters are seen as a form of the ''Tsunki'', or water spirits: they are a sort of "water people" who feed on fish. They appear in a [[Fishing techniques|fish poisoning]] legend where they assist a man who has wasted his sexual energy, creating the [[anaconda]]s of the world from his distressed and extended genitals.<ref name="Descola">{{cite book |title=In the Society of Nature: A Native Ecology in Amazonia |last=Descola |first=Philippe |year=1994 |publisher=Cambridge University |
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Press|isbn=0521411033 |pages=280–282 }}</ref> The [[Bororo people|Bororo]] have a legend on the origin of [[tobacco smoking]]: those who used the leaf improperly by swallowing it were punished by being transformed into Giant Otters; the Bororo also associate the Giant Otter with fish and with fire.<ref>{{cite book |title=The Raw and the Cooked |last=Lévi-Strauss |first=Claude |others= trans. John Weightman and Doreen Weightman|year=1983 |publisher=University of Chicago Press|isbn=0226474879 |pages=104–108 }}</ref> A [[Ticuna language|Ticuna]] legend has it that the Giant Otter exchanged places with the Jaguar: the story says Jaguar formerly lived in the water and the Giant Otter came to the land only to eat.<ref>{{cite book |title=El ojo que cuenta: Mitos y costumbres de la Amazonía indígena ilustrados |last=Landolt |first=Gredna |year=2005 |publisher=International Work Group for Indigenous Affairs|isbn=9972264009 |pages=81 }}</ref> The indigenous [[Kichwa]] peoples from Amazonian Peru believed in a world of water where Yaku runa reigned as mother of the water and was charged with caring for fish and animals. Giant Otters served as Yaku runa's canoes.<ref>{{es icon}} Ching, César (October 2006). [http://www.premioreportagem.org.br/article.sub?docId=24668&c=Peru&cRef=Peru&year=2007&date=outubro%202006 PER-I38: El mundo del agua temido y poco conocido.] BioDiversity Reporting Award. Semanario Kanatari, Iquitos, Iquitos. Retrieved on [[2008-01-27]].</ref> |
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==Notes== |
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{{reflist|2}} |
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==References== |
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* {{cite journal |last=Carter |first=S.K. |coauthors=Rosas, F.C.W. |year=1997 |title=Biology and conservation of Giant Otter (''Pteronura brasiliensis'') |journal= Mammal review |volume=27 |issue=1 |pages=1–26 |url=http://www.giantotterresearch.com/articles/Carter_and_Rosas_1997.pdf |format=PDF |accessdate= 2007-11-06 |doi=10.1111/j.1365-2907.1997.tb00370.x}} |
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* {{cite journal |last=Duplaix |first=Nicole |year=1980 |title=Observations on the ecology and behavior of the giant river otter ''Pteronura brasiliensis'' in Suriname |journal=Revue d'Ecologie (Terre Vie) |volume=34 |pages= 495–620 |quote= }} |
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==External links== |
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{{commons|Pteronura brasiliensis}} |
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{{wikispecies|Pteronura brasiliensis}} |
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*ARKive – [http://www.arkive.org/species/GES/mammals/Pteronura_brasiliensis/ images and movies of the giant otter ''(Pteronura brasiliensis)''] |
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* {{cite journal|title=Guianas Rapid River Bio-assessments and Giant Otter Conservation Project |publisher=[[World Wildlife Fund]]|url=http://www.giantotterresearch.com/articles/WWF_Giant_Otter_Report_PDFMini.pdf |format=PDF|first=Nicole|last=Duplaix|year=2002}} ''Includes excellent photography''. |
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* {{cite web |url=http://www.2docstock.com/Suriname/Reports/Bibliography/bibliography.html |title=Giant Otter Bibliography |accessdate=2008-01-27 |publisher=2DocStock Photography|last=Duplaix|first=Nicole |coauthors=Groenendijk, Jessica; et al. }} ''A very complete bibliography on giant otters.'' |
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[[Category:Otters]] |
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[[Category:Mammals of South America]] |
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