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SPECIES PROFILE South American Coati Nasua nasua Picture 1: the South American Coati http://upload.wikimedia.org/wikipedia/commons/5/5e/Coati_roux_Amiens_4.jpg July 2014 Author: Mary Charrett BSc Last Updated: September 2014 1. SUMMARY The South American coati (Nasua nasua) is a member of the raccoon family Procyonidae ranging from southern Colombia and Venezuela to northern Argentina and Uruguay. Including Brazil, Bolivia, Ecuador, French Guiana, Guyana, Paraguay, Peru and Surinam (IUCN 2008, Gompper & Decker 1998). It has been released into Mallorca, Robinson Crusoe Island and Cambria UK. The impact on Robinson Crusoe Island has been notably on native seabird colonies where the coati has decimated numbers, due to their being no predators or limiting factors for coatis on the island. In Australia there are no coatis in the wild. There is little likelihood that this diurnal species could establish in Tasmania, the climatch modelling shows that the cold is a limiting factor. If the species did establish in Tasmania there would be concern for native bird and small mammal species. The coati is classified as least concern with the International Union for Conservation of Nature (IUCN) Red List but with a decreasing population. Uruguay has listed N. nasua solitaria, a subspecies of coati, under CITES (Convention on International Trade in Exotic Species) Appendix III. 2. NAME AND TAXONONY 1.1 Scientific Classification Kingdom: Animalia Phylum: Chordata Class: Mammalia Order: Carnivora Family: Procyonidae Genus: Nasua Species: N. nasua 1.2 Common Names Coati, Common Coati, Brown Nosed Coati, Southern Ring-tailed Coati, Ring-tailed Coati, Coatimundi, Archuni (Spanish), Tejón (Bolivia, Colombia, Ecuador), Coati rojo (Guatemala), Cusumbo, Guache (Colombian), Guati, Quatimunde (Brazil), Coachi (Guyana), Kwaskwasi (Surinam), Archuni Sehuaro (Peru) and Neusbeer (Wesapiens 2012). pg. 2 1.3 Sub-species There are 12 geographically defined subspecies of Nasua nasua (Marriner 2014). Nasua nasua nasua - North Brazil and French Guiana Nasua nasua aricana - Paraguayan Nasua nasua boliviensis - Bolivia Nasua nasua candace - Colombia Nasua nasua cinerascens – Chaco of Argentina, Bolivia, Brazil and Paraguay Nasua nasua dorsalis – Amazonian of Brazil, Ecuador, Peru and Venezuela Nasua nasua manium - Ecuador Nasua nasua montana – Mountains of Brazil and Peru Nasua nasua quichua – Azuay of Ecuador Nasua nasua solitaria – (Coatimundi) South Brazil, Argentina and Uruguay Nasua nasua spadicea – Uruguay, Brazil and Paraguay Nasua nasua vittata – Guyana, French Guiana, Surinam and Venezuela 1.4 Taxonomy The Southern American Coati belongs to the Family Procyonidae and the genus Nasua. Nasua nasua has the longest snout and claws of the Procyonidae family which includes; Bassariscus astutus (ringtail) Bassariscus sumichrasti (cacomistle) Nasuella olivacea (mountain coati) Potos flavus (kinkajou) Procyon cancrivorus (crab-eating raccoon) Procyon lotor (Guadeloupe raccoon) Procyon pygmaeus (Cozumel raccoon) Nasua narica (white nosed coati) Procyonidae is a New World family with members from southern Canada to Northern Argentina. They are typically small to medium sized, all have tails and plantigrade locomotion. They have 5 digits on hind feet and forefeet and ears are always erect. They are usually grey/brown in coloration some species have markings on their faces and rings around their tails (Braddy 2003). There are two species of the genus Nasua, Nasua nasua (South American Coati) and N. narica (White nosed Coati) including N. nelsoni. It was thought that N. nelsoni was a different species to N. nasua and N. narica mainly due to it being located solely on Cozumel Island, however the findings of Decker (1991) shows that N. nelsoni is common to N. narica. Nasua narica inhabits much of Central America and some southern North American states, there is little to no crossover of geographical boundaries. pg. 3 3. DESCRIPTION The South American coati is an arboreal and terrestrial animal which lives in large matriarchial groups. They have a long slender body of between 41 – 70 cms and a long ringed tail of 32-70 cms. Adults average shoulder height is 30cms and their limbs measure 8cms long. Body mass is between 2.5kgs and 10kgs, the males are larger than the females and juveniles (Animal files 2014, Bittner et al. 2010, Gompper & Decker 1998). During the day coatis have a mean body temperature of 38.8° C and at night of 36.4°C, they have good thermoregulation which keeps them stable up to an external temperature of 35°C with a base heart rate of 70 beats per minute (Gompper & Decker 1998). Colouration of the coati varies vividly amongst the species. Their coats of thick coarse fur can be red/brown, grey/brown or yellow/brown. The upper body is usually darker, brown, grey or rust red, underpants are usually white. The non-prehensile tail is dark brown or grey ringed with yellow or white. The tail is used as an anchor when foraging and descending from trees, it is also used as a visual cue when marching with other conspecifics (Marriner 2014). The snout is dark brown or black, white spots are below, behind and above eyes, the ears have a white inner and dark outter surface, yellowish neck and black feet. Juveniles are usually lighter coloured and smaller than adults (Gompper & Decker 1998). The rostrum and rhinarium of the coati is highly specialised. The rhinarium is longer than the lower mandible, the snouts upper edge protrudes past the nostril and the is no philtrum on the upper lip. The snout is long and mobile, the protruding and pointed snout is used to get into cracks and holes to investigate and forage. Long claws, powerful limbs and a mobile snout allow the coati to break apart wood and search detritus and leaf litter for food (Braddy 2003). Their teeth are similar to that of a domestic dog, Males and females have size and shape differences in their dental structure but generally they have blade like canines, high crowned pre-molars and molars with sharp cusps, the design of their teeth is due to their omnivorous diet (Gompper & Decker 1998, Pieri et al. 2011). Coatis have well developed sight and hearing, their eyes have a reflective tapetum which allows for colour vision and they hear upto 7.4 octaves with a frequency limit of 95kHz. Compared to other carnviores this is advanced. Other adaptations they possess are acutely adapted anal glands, used for scent marking and identification, the ability to rotate their ankle bones 90° to decend trees headfirst and can grip verticle trunks with ease (Gompper & Decker 1998). Nasua nasua and N. narica are stand alone species and do not look like any other species in Australia. pg. 4 4. Conservation and Legal Status 4.1 Conservation Status South American coatis are classified as least concern by the IUCN Red List of Threatened Species, they are widespread and common in most areas, they seem to adapt well to anthropological influences. They have a decreasing population trend due to hunting and habitat loss from deforestation, road building, mining and dam building (IUCN 2008, Gompper & Decker 1998) 4.2 Legal Status The Environment Protection and Biodiversity Conservation Act 1999 (the EPBC Act) provides the legal framework to protect and manage nationally and internationally important flora, fauna, ecological communities, and regulate the import and export of wildlife. Nasua nasua is listed as a specimen ‘suitable for live import with an import permit’ issued under this Act, with the condition of import being ‘Eligible non-commercial purposes only, excluding household pets. High security facilities only’. The subspecies N. nasua solitaria is legally protected under the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES) appendix III in Uruguay (CITES 2014). 5. Life History Female South American coatis become sexually mature at 2 years, males become sexually mature at 3. The mating season is from September to February timing changes with altitude, mating times coincide with food availability, a couple of months before abundance (Favaron et al. 2014, Marriner 2014, Rodrigues da Paz et al. 2012, Russell 1981, Wesapiens 2012). Females show reproductive synchrony and they are receptive for 1-2 weeks only (Beisiegel 2001, Hirsch & Maldonado 2011), during this time they mate with multiple males, mating takes about 55 minutes on average which is a fairly long time, subsequently decreasing the opportunity for sneaky mating, there is usually one male that will accompany a group of females, otherwise males are solitary. It shows that the paternity rate for an in group male is 66-91%, however 50% of the group had extra-parented pups (Hirsch & Maldonado 2011). Gestation takes between 70-77 days. Females disband their groups during this time and nest in trees, similar to bird nesting (Gompper & Decker 1998, Olifiers et al. 2009). Females spend 5-6 weeks in the nests and have 1-7 pups (but typically 3-4) which they then bring down to join the group (Animal files 2014, Beisiegel 2001, Favaron et al. 2014, Marriner 2014, Rodrigues da Paz et al. 2012, Russell 1981, Wesapiens 2012). In captivity allonursing has been observed (Gompper & Decker 1998). pg. 5 The timing of young coming out coincides with that of the rainy season, this is due to the increased spiders and millipedes, millipedes have100 times more calcium than other arthropods and are an excellent source of calcium for mothers which are lactating (Alves-Costa et al. 2004). Neonates at 5 days old weigh 100-180 grams, at 10 days their eyes open, standing at 19 days, focusing eyes and walking well by 24 days and by 26 days they start climbing (Gompper & Decker 1998, Favaron et al. 2014, Marriner 2014). Mothers will feed until the young is about 4 months old then wean (Animal files 2014). The juvenile is classed an adult at 2 years for females and 3 years for males when they become sexually mature. In the wild coatis can live to 7 or 8 years of age, in captivity they are expected to live up to 20 years (Animal files 2014, Marriner 2014). It has been said in recent publications that the coati has no known hybridisations (Benirschke 2004). 6. Habitat Requirements and Preferences Coatis live in a wide range of habitats, Neotropical, deciduous forests, evergreen, old growth and secondary forests, they adapt well to anthropogenic influences and can become established in disturbed areas. They live in relatively stable temperature climes, minimum 3°C to maximum 29°C with an average of 18°C -20°C (Beisiegel 2001). Coatis in cold climates have to be weary about getting frostbite of the tail in temperatures below 4°C (AZA 2010). Altitude does not influence their distribution, they are found from sea level to the Andean mountains, an altitude of 2,500ft (IUCN 2008, Gompper & Decker 1998, Wesapiens 2012). Coatis use nests for sleeping, mating, giving birth and for bringing up young, they do not use tree hollows (Olifiers 2009). After mating the female coati will spend the next 5-6 weeks in the nest looking after her young, until they are old enough to re-join the group. Nests are built of twigs mainly of a diameter of 12cms, the nests are hemispherical in shape 40-50cms in diameter composed primarily of liana branches intertwined with leaves (Wesapiens 2012). The typical volume of the nest is 31.7cm³ – 47.5cm³. Nests can also be built in rock crevices and cliffs there can be multiple nests in a tree and usually above the height of 5 meters (Olifiers et al. 2009). pg. 6 7. Natural Geographic Range Figure 1: Distribution data from IUCN Red List for the South American Coati (IUCN 2008) Coatis came to South American in the late Miocene era (23 – 5 million years ago) by island hopping. Since then N. nasua has been a resident of South America, the distribution range is from Colombia and Venezuela to northern Argentina and Uruguay. Including Brazil, Bolivia, Ecuador, French Guiana, Guyana, Paraguay, Peru and Surinam (IUCN 2008, Gompper & Decker 1998). The population density of coatis in South America varies with the habitat type and resource availability. A national park in Brazil, Parque Estadual do Prosa has a density of 33.71 individuals per a km² (Costa et al. 2009). In low lying deciduous forests the population density can be as low as 6.2 individuals per km² and in tall galley forests it usually is about 13 individuals per km² (IUCN 2008). Coatis can live in numerous types of environments, they have been found in; closed in, deciduous and evergreen forests, riverine gallery and cloud forests, rainforest, Gran Chaco, dry scrub forest and savannah. They can utilise primary or secondary forest and they can travel between vegetation islands in areas that have been logged, they are also very comfortable living in close proximity to humans and use human garbage as a source of food (Alves-Costa & Eterovick 2007, Beisiegel 2001, IUCN 2008, Gompper & Decker 1998). Coatis are adapting to human encroachment, this being said they may become more vulnerable in the future due to deforestation, mining, hunting and road kill. pg. 7 8. Introduced Geographic Range The coati has been introduced to Robinson Crusoe Island of the Juan Fernandez Islands off Chile, Mallorca and due to release from captivity by zoos or people there has been an established population in Cambria, in the UK for the last decade (Costa et al. 2009, Edgeworth 2010, IUCN 2008, Mallorcaphotoblog 2010). Robinson Crusoe Island has had coatis for over 50 years, they have led to the decline in many endemic ground nesting bird species (Independent 2009). Sightings of wild coatis in the UK have been at Windermere and Backbarrow about 16kms apart, they estimate the introduced population is 10 adults but parks and wildlife in the area are certain they are breeding so numbers will increase progressively. Coatis can travel around 2kms a day so they can travel between islands of vegetation and to new areas quickly. Mallorca is in the Balearic Islands chain south of Spain, the Mallorca government has issued fliers alerting people to the coati and to call and report sightings of the pest. It is not known how many coatis are on Mallorca (Mallorcaphotoblog 2010). Since the family Procyonidae live only in the Americas there is little chance of hybridisation in the introduced areas. Coatis thrive in disturbed areas, they are able to adapt well to human influences and interact with humans (Alves-Costa & Eterovick 2007). 9. Potential Distribution in Tasmania CLIMATCH modelling suggests that the results for Tasmania vary from 0 (very low match) to 7 (moderately high match). The source areas as shown in Figure 2, show that the native distribution of southern America was used as well as the introduced regions of Mallorca and Robinson Crusoe Island, Cambria in the United Kingdom had no available data, the closest available station was used, this being Lockerbie which is 124kms north of Windermere. It is evident in Figure 2 that these introduced areas will play a significant role in the CLIMATCH modelling. pg. 8 Figure 2: Coati distribution in South America and Mallorca, included in the comparison was the introduced populations. (Image Source: CLIMATCH - http://data.daff.gov.au:8080/Climatch/climatch.jsp) The likelihood of the coati existing in Tasmania if it should escape is moderate, due mostly to the low temperatures experienced in winter, it has been noted that for coatis held in captivity in any area that reaches less than 4°C should provide heating, the main reason being the risk of frostbite to their tails (AZA 2010). Coatis would be able to find food in rural Tasmania, not so much fruit but invertebrates and small vertebrates would be plentiful, in urban Tasmania they would scavenge off human waste as well as invertebrates and small vertebrates. Shelter would be fairly easy to come by due to the vast areas of forest in which they could build nests and forage through. There are many branches and vines they could use to build nests, they do not use specific materials. A single female can have up to 7 offspring a year, usually between 3 and 4, and this means their ability to grow a large population over time would be significant. pg. 9 Figure 3. CLIMATCH modelling of Nasua nasua native homelands in southern America and introduced areas such as Mallorca and Robinson Crusoe Island, compared to Australian environments. Tasmania being the least hospitable area. Imaged by CLIMATCH modelling. (Image Source: CLIMATCH http://data.daff.gov.au:8080/Climatch/climatch.jsp) 10. Diet and Feeding Behaviour Coatis are diurnal omnivorous generalists, their diet is seasonal and consists of fruits, invertebrates, lizards, frogs, eggs, fish, snakes and birds. They are both arboreal and terrestrial feeders even foraging under bark and in dens. They especially like epiphytic bromeliads which are a rich food source. When eating tarantulas, venomous frogs and millipedes’ coatis use intense prey manipulation, they rub the prey on the ground to either get rid of venom or irritating setae (Alves-Costa et al. 2004, Beisiegel 2001, Bittner et al. 2010, IUCN 2008, Ferreira et al. 2013, Gompper & Decker 1998). In areas of human influence the diet of the coati is altered to include anthropogenic waste, food scraps and animals attracted to food waste such as cockroaches (Ferreira et al. 2013). They have a growing role in the ecosystem with the reduction of large herbivore numbers the seed spreading of coatis is keeping flora diversity growing (Alves-Costa & Eterovick 2007). Coatis find their food primarily by olfaction and pg. 10 they can travel for about 2 kilometres each day (Di Blanco & Hirsch 2006). When feeding coatis forage individually, there is no sentry or lookout each coati keeps vigil. It is known that 2 bird species forage simultaneously with coatis, the Mantled hawk and the Plain brown woodcreeper, they would eat prey flushed out by the coati, and it is thought the hawk may act as a sentinel, spotting danger and alerting the coatis to it (Beisiegel 2007). If coatis were freed around Tasmania they would feed off fruit trees, they would also generalist feed on small invertebrates and vertebrates, finding enough food would be difficult. Because they feed mainly on fruits the rest of the tree is left intact so amenity trees and shrubs would be safe. 11. Social Behaviour and Groupings Coatis live in matriarchal family groups, there can be up to 70 individuals in a group, the group is made up of adult females and juveniles of both sexes. Sometimes there will be a single male in the group however it shows that he adds no fitness to the group and provide no parental care. The band is made up usually of family members but some unrelated members are found too, after mating has finished the adult females will go off solitarily and build nests that they will stay in for 4 months while they rear their young, during this time the juveniles will stay in a band together. Males are solitary year round and will only approach females when it is time to mate, they are received with hostility and aggression year round. Coatis are not aggressively territorial, females will have large ranges which will have several male territories within. Males can become aggressive when defending their territories (Alves-Costa & Eterovick 2007, AZA 2010, Beisiegel 2001, IUCN 2008, Gompper & Decker 1998, Hirsch & Maldonado 2011, Russell 1981). Band co-operation is extremely important due to the females needing to protect the juveniles from the physically bigger males, males will prey on juvenile coatis but not their own offspring which they are able to recognise. This is one reason why all but one male is excluded from a social group, he will have fathered most of the young (Russell 1981). pg. 11 12. Natural Predators and Disease 12.1 Natural Predators The main predators of coatis are jaguar, puma, ocelot, jaguarmundi, snakes, domestic dog, foxes, large birds of prey and humans. Adult monkeys will predate on young in the nest while the mother feeds (Alves-Costa et al. 2004, Gompper & Decker 1998, Olifiers et al. 2009, Perry & Rose 1994, Wesapiens 2012). Coatis vocalise when stressed or in danger so that will attract other predators to the area and alert other coatis to the danger. They have no defences apart from their claws and teeth, they will try to bite and scratch an attacker and chatter shrilly (Janzen 1970). There are no predators to full grown coatis in Tasmania, there are predators to juveniles, such as birds of prey, quolls and possums. 12.2 Disease Coatis have many parasites in the wild, tapeworms, kidney worms, ticks, fleas, lice, nematodes and acanthocephalan (hook worms passed on by crustaceans) (Gompper & Decker 1998, Milanelo et al. 2009, Rodrigues et al. 2006). They are known to carry other pathogens that both humans and animals can be subject to such as, mange, tuberculosis, canine distemper, feline panleukopaenia, toxoplasmosis, dermatophytosis, histoplasmosis, and sporotrichosis. Myobacterium bovis can cause tuberculosis in humans, air droplet transmission though very rare can be a public health issue (Murakami et al. 2012). 13. Threat to Human Safety The coatis can do limited damage to a person, they have fairly large canines which in the reports, they use freely to puncture skin and the occasional clawing is reported. One such attack of two children in a small Brazilian town left the children with open wounds up to 3cms long. It was thought that the coati attacked them because it felt threatened. They were treated by firefighters at the scene and taken to the Vila hospital to have the wounds flushed thoroughly and dressed, they also received a rabies prophylaxis. Attacking humans does not seem to be premeditated but a split decision act which can, from reports, sometimes cause the coati to go into a catatonic state afterwards. Females it has been noted become unpredictable when reaching sexual maturity. Multiple posts on how the jingling of keys and the sound of metal clinking can cause split second attacks (Bittner et al. 2010). pg. 12 Coatis in captivity are more likely to have tuberculosis due to the frequency of human interactions, tuberculosis is present in Tasmania. Canine distemper is also present in Tasmania, it is thought to have caused part of the decline in Tasmanian tigers (Australian Government 2007). Feline panleukopenia and toxoplasmosis are worldwide. Myobaterium bovis is found Australia wide. Histoplasmosis and sporotrichosis are found in Tasmania. Coatis being imported will be subject to quarantine checks and will not be bringing diseases into Tasmania. 14. History as a Pest The number of coatis is growing rapidly in South America due to a loss of larger predators like Jaguars. They have adapted so well to human influences that they have incorporated stealing food and rummaging through garbage as viable food sources. The coati has become a semi-common pet in America but accidental or purposeful releases of coatis have been on the rise, releasing captive coatis is thought to be behind the recent Mallorca and Cambria UK populations. Introducing a non-native species can cause all sorts of ecological changes, the intentional introduction onto Robinson Crusoe Island has resulted in the devastation of many seabird colonies (Independent 2009). 15. Potential Impact in Tasmania If coatis were to enter the ecosystem of Tasmania it would not have a natural predator to keep numbers in check and it could harm the native wildlife. That being said it would not be able to survive over winter in Tasmania, the minus temperatures are so different from its native weather. The available diet for this mainly frugivorous animal would not be sustainable in the wild. If N. nasua were to establish in Tasmania, it would be a competitor with animals that consume a similar diet and sleep in trees but not tree hollows (Olifiers 2009), mainly the brushtailed possum (Trichosurus vulpecula), to a lesser extent, ringtail possum (Pseudocheirus peregrinus), eastern quoll (Dasyurus viverrinus) and spotted tailed quoll (Dasyurus maculatus) may compete for sleeping trees. pg. 13 16. Previous Risk Assessments There is a previous risk assessment from Great Britian, the result was a low risk with moderate uncertainty. Due mostly to the temperature restrictions and complex social structure with low reproductivity. The negative aspect highlighted was the generalist omnivore diet which comes with high environmental plasticity (GBNNSS 2011). The coati is listed on the Vertebrate Pest Committee a 2/Extreme, which means it is able to be kept in zoos or endorsed collections and it has a extreme threat rating (Feral 2007). 17. Risk Management A complete Species Risk Assessment will be undertaken by the Tasmanian Department of Primary Industries, Parks, Water and Environment (DPIPWE), following the submission of this Species Profile. Utilising the Bomford (2008) Modelling System, a four-factor analysis will be applied to assess the likelihood of establishment of coatis, should import of the species be permitted and an escape or release occurs. Should the risk classification process allow the import of coatis into Tasmania, a detailed Species Management Plan will be developed to cover all aspects of transport, keeping and disposal risk management, in consideration of the associated classified level of risk for the species. 18. References Alves-Costa, CP Fonseca, GAB & Christofaro, C 2004, ‘Variation in the diet of the brown nosed coati (Nasua nasua) in southeastern Brazil’, Journal of Mammalogy, vol. 85, no. 3, pp. 478-482. Alves-Costa, CP & Eterovick, PC 2007, ‘Seed dispersal services by coatis (Nasua nasua, Procyonidae) and their redundancy with other frugivores in southeastern Brazil’, acta oecologica, vol. 32, pp. 77–92. Animal Files 2014, ‘South American Coati’, viewed 7 July 2014, <http://www.theanimalfiles.com/mammals/carnivores/coati_south_american.html> pg. 14 Association of Zoos and Aquariums (AZA) Small Carnivore Taxon Advisory Group 2010, Procyonid (Procyonidae) Care Manual. Association of Zoos and Aquariums, Silver Spring, MD. Australian Government 2007, ‘The Tasmanian Tiger’, viewed 11 August 2014, <http://australia.gov.au/about-australia/australian-story/tasmanian-tiger> Beisiegel, BM 2001, ‘Notes on the coati, Nasua nasua (Carnivora: Procyonidae) in an Atlantic forest area’, Brazillian Journal of Biology, vol. 61, no. 4. Beisiegel, BDM 2007, ‘Foraging Association between Coatis (Nasua nasua) and Birds of the Atlantic Forest, Brazil’, Biotropica, vol. 39, no. 2, pp. 283-285. Benirschke, K 2004, ‘Coatimundi’, Comparitive Placentation, viewed 12 August 2014, <http://placentation.ucsd.edu/nasua.html> Bittner, GC Hans, NR Neto, GH Morais, MO Filho, GH & Haddad, V 2010, ‘Coati (Nasua nasua) Attacks on Humans: Case Report’, Wilderness & Environmental Medicine, vol. 21, no. 4, pp. 349-352. Bomford, M 2008, ‘Risk assessment models for establishment of exotic vertebrates in Australia and New Zealand’, Invasive Animals Cooperative Research Centre, Canberra. Braddy, S 2003, ‘Nasua nasua’, Animal Diversity Web, viewed 8 July 2014, <http://animaldiversity.ummz.umich.edu/accounts/Nasua_nasua/> Bureau of Rural Sciences (BRS) 2011, ‘Climatch’, Department of Agriculture, Fisheries and Forestry, viewed 1 August 2014, <http://adl.brs.gov.au:8080/Climatch/> Chevillard-Hugot, MC Muller, EF & Kulzen, E 1980, ‘Oxygen consumption, body temperature and heart rate in the coati (Nasua nasua)’, Comparative Biochemistry and Physiology Part A: Physiology, vol. 65, no. 3, pp. 305-309. 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Durrell Wildlife Conservation Trust (DWCT) 2006, ‘Ring-tailed coati species factsheet’, viewed 8 August 2014, <http://www.durrell.org/animals/mammals/ringtailed-coati/> Edgeworth, A 2010, ‘Coati numbers on the increase in Cumbria’, The Westmorland Gazette, Westmorland UK. Favaron, PO Morini, JC Mess, AM Miglino, MA & Ambrosio, CE 2014, ‘Placentation and fetal membrane development in the South American coati, Nasua nasua (Mammalia, Carnivora, Procyonidae)’, Reproductive Biology and Endocrinology, vol. 12, no. 57. Feral 2007, ‘List of Exotic Vertebrate Animals in Australia’, Vertebrate Pests Committee, viewed 8 August, < http://www.feral.org.au/list-of-exotic-vertebrate-animals-in-australia/> Ferreira, GA Nankano-Oliveira, E Genaro, G & Lacerda-Chaves, AK 2013, ‘Diet of the coati Nasua nasua (Carnivora: Procyonidae) in an area of woodland inserted in an urban environment in Brazil’, Revista chilena de historia natural, vol. 86, no. 1. Freitas, EP Rahal, SC Teixeira, CR Teixeira, RH Mendes, GM & Gioso, MA 2008, ‘Oral cavity evaluation and dental chart registration of coati (Nasua nasua) in captivity’, Journal of Veterinary Dentistry, vol. 25, no. 2, pp. 110-117. Gompper, ME & Decker, DM 1998, ‘Nasua nasua’, Mammalian Species, no. 580, pp 1-9. Great Britain non-native species secretariat (GBNNSS) 2011, Risk assessments, Nasua nasua (Coati), viewed 7 August 2014, <http://www.nonnativespecies.org//index.cfm?sectionid=51> Hirsch, BT & Maldonado, JE 2011, ‘Familiarity breeds progeny: sociality increases reproductive success in adult male ring-tailed coatis (Nasua nasua)’, Molecular Ecology, vol. 20, no. 2, pp. 409–419. 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