Overview

Brief Summary

Biology

The maned wolf hunts primarily at night, and during dusk and dawn hours, while the days are often spent resting, often in areas of thick bush cover (5). The diet consists of a wide variety of fruits and small mammals, such as armadillos and rabbits, but also includes occasional pampas deer (Ozotoceros bezoarticus), birds, reptiles, insects, fish and arthropods (1). The maned wolf's main source of food is the tomato-like lobeira fruit, which grows throughout its range and is thought to provide medicinal aid against the giant kidney worm, Dioctophyme renate (1) (5). Scavenging on road-kill also occurs and free-ranging chickens are frequently stolen from farms (8). Unlike other wolves that live in cooperative breeding packs, the maned wolf is primarily solitary (10). Although the basic social unit is the male-female mated pair, which share a home range typically between 25 to 50 square kilometres (11), these individuals remain fairly independent of one another and only closely associate during the breeding season from April to June (5) (6) (8). The female gives birth to a litter of one to five pups each year (average of three) between June and September (6) (8). Originally, it was believed that the female alone cared for the young, suckling them for up to 15 weeks (3). However, in captivity males have been observed grooming and defending pups, as well as feeding them by regurgitation. Pups reach sexual maturity and disperse from their natal home range at around one year old, but do not usually reproduce until the second year (8). Captive individuals have lived up to 16 years (8).
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Description

Standing at almost a metre tall, the maned wolf is the largest Canid in South America and the only member of its genus, Chrysocyon (5) (6). With a golden-red coat, long pointed muzzle and large erect ears (7), it is similar in appearance to the red fox (Vulpes vulpes) (3). However, its extremely long, thin legs make the maned wolf immediately recognisable and, with its fox-like attributes, have earned it the epithet 'a fox on stilts' (2). This distinctive feature is thought to be an adaptation to help the animal see above the tall grass of its habitat (5). The common name, 'maned wolf', is derived from the characteristic mane-like strip of black fur running from the back of the head to the shoulders (8), which stands erect when danger is sensed (7). The muzzle and lower legs are black, while the throat, inside of the ears and tip of the tail are white (7) (8).
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Distribution

Range Description

The Maned Wolf inhabits lowland grasslands and scrublands of central South America, south of the mouth of the Parnaiba River in northeastern Brazil, throughout eastern Paraguay east of the Rio Paraguay, extending into north/northeast Rio Grande do Sul State in Brazil, south to Santa Fe and Entre Rios provinces in Argentina and west to the Pampas del Heath in Peru (Queirolo et al. 2011, Williams et al. 2012).

In Brazil, the range is limited by the Amazon forest in the north/northwest and by the arid Caatinga in the northeast (Queirolo et al. 2011). In the last five years, the Maned Wolf has been recorded in areas where no records (> 50 years) previously existed, specifically in areas where habitat degradation and conversion are occurring. In the Amazonian arc of deforestation, south of Amazonas and south/east of Para states, the species has been increasingly registered in areas that were converted from forests into pastures and agricultural fields (R. Paula pers. comm. 2015). The same is true, but to a greater degree, in southeastern Brazil where the Atlantic Forest biome has been deforested and the Cerrado has been degraded (Queirolo et al. 2011). Although these records might suggest an expansion of the species distribution range, records are scattered and populations may not be stable. In Argentina, the species is present in the northern and central region of the country (Queirolo et al. 2011) but a range contraction has occurred in the south with most recent records in the provinces of Chaco, eastern Santiago del Estero, northeastern Crdoba, northern Santa Fe, northern Entre Ros, Corrientes, and southern Misiones (Queirolo et al. 2011). The range across eastern Paraguay is considered widespread, but discontinuous, with no large concentration in any single area (Cartes et al. 2014). The Maned Wolf is distributed across the savannas of Bolivia; however, all detailed information for this country is limited to a single park in the department of Santa Cruz (Emmons 2014). The species historically ranged into northern Uruguay, but to date there have been only sporadic and unconfirmed occurrences in the north-east (Queirolo et al. 2011) since the last confirmed record in 1990 (Mones and Olazarri 1990).

Two recent evaluations of Maned Wolf distribution have provided an update on the full extent of the range (Quierolo et al. 2011), including range shifts within Brazil (Paula et al. 2013). These studies recorded that the Maned Wolf distribution covers nearly five million km2 with ~72% of the range located in Brazil, which corresponds to an increase in the total area reported 30 years ago (3,414,169km2) by Dietz (1984). This increase in the Maned Wolf distribution is likely due to both genuine change (the species expansion into new regions) as well as non-genuine change (mostly new or additional records from the field).

More recently, to seek greater precision concerning the distribution range of the species in Brazil, potential distribution and probability of presence were modelled with Maxent, a software using maximum entropy algorithms (R. Paula and E. Ferraz pers. comm. 2015). All Brazilian biomes, with the exception of the Amazon and Caatinga, neither of which are suitable for the Maned Wolf, were modelled using >1,000 presence points collected in the last 10 years combined with several variables (including topographic, bioclimatic, geophysical and anthropogenic) at a resolution of onekm2. The results indicate that the potential distribution area for Maned Wolves in Brazil is 2,815,061km2. The majority (70%) of this area represents areas with <25% probability of presence and those areas are considered unsuitable for the Maned Wolf occupancy. In addition, the highly suitable area (i.e., >75% probability of presence) accounts for only 0.4% of the total of the species distribution area within Brazil. These declines in both total area and overlap with lower probabilities of presence reflect an increase in deforestation rates in the Cerrado over the last 20 years. While the species has been documented in other biomes, the model suggests a retraction of their distribution within the Cerrado due to the ongoing conversion of intact habitat to areas with agriculture and pastures; therefore, while there is evidence that Maned Wolves can persist in mosaics of natural and disturbed areas, extensive monocultures and exotic pasturelands have been indicated as unsuitable areas.
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The maned wolf is distributed from the mouth of the Parnaiba River in northeastern Brazil west to the Pampas del Heath in Peru and South through the Chaco of Paraguay to Rio Grande do Sul, Brazil. Its former range included parts of Uruguay and Argentina.

Biogeographic Regions: neotropical (Native )

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Historic Range:
Argentina, Bolivia, Brazil, Paraguay, Uruguay

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Range

The maned wolf is found in central South America, from north-eastern Brazil, south through Paraguay and west into Peru (1). It is also found in small areas of Argentina and Bolivia, and may still be present in some areas of Uruguay, despite being believed to be extinct there in the 19th century (5) (8).
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Physical Description

Morphology

Chrysocyon brachyurus is a stunning animal. The largest of all South American canids, it stands almost one meter tall at the shoulder and has a long, golden-red coat. Head and body length ranges from 1245 to 1320mm and tail length from 280 to 405mm. The long thin legs, which may serve to help the maned wolf to see above tall grass, grade from red to black at their lower portions. The anterior part of the erectile mane of long hairs is black as well. The body is narrow and the ears large and erect. The dentition of the maned wolf reflects its food habits. As this animal does not kill or eat large prey, its upper carnassials (shearing teeth) are reduced, its upper incisors weak, and its canines are long and slender.

Range mass: 20 to 23 kg.

Other Physical Features: endothermic ; bilateral symmetry

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Ecology

Habitat

Habitat and Ecology

Habitat and Ecology
In general, Maned Wolves inhabit open habitat types favouring grasslands, shrub habitats, woodland with an open canopy (cerrado), mixed forest/grassland, and wet fields (which may be seasonally flooded). Rocky fields associated with open grasslands in higher altitudes (1,000-2,000 m asl) can be an important habitat for the species in some areas (Coelho et al. 2008, R. Paula pers. obs.). In central Brazil, similar to Bestelmeyer (2000), who found Maned Wolves prefer areas with low to medium shrub density, Vynne et al. (2011) found they avoided areas where the closed canopy was 30%. Coelho et al. (2008) found the Maned Wolf strongly avoided the forested areas that border the Cerrado and Atlantic forest ecoregions. In northeastern Argentina, south/southeastern Brazil, and southeastern Paraguay, Maned Wolves have been recorded in the exotic eucalyptus, pine, soybean and sunflower plantations (Orozco et al. 2009, Soler 2009, A.J. Giordano pers. comm. 2015, R. Paula pers. obs.) and in remnants of Atlantic forest (Chiarello 2000, A.J. Giordano pers. comm. 2015). Also, the presence of Maned Wolves in populated rural and urban areas was also documented in the last ten years in Santa Fe, Corrientes, Chaco and Santiago de Estero provinces in Argentina (Pautasso et al. 2009, Orozco et al. 2013a). In addition while Maned Wolves preferred open, natural grasslands, they were as likely to use low-standing croplands within five kilometres due to the high density of rodents (Vynne et al. 2011). While Queirolo et al. (2011) proposed that Maned Wolves are expanding their former range in Brazil to include areas converted from inhospitable primary or thick secondary forest into open areas for grassland, agriculture and livestock. Muir and Emmons (2012) failed to find evidence of Maned Wolves expanding into agricultural areas in Bolivia. In Paraguay, preliminary studies with camera traps suggest that sustainable cattle-ranching was compatible with high Maned Wolf activity (A.J. Giordano pers. comm. 2015). Vynne (2010) suggests the foraging benefits of agricultural areas are negated when the crops become tall, homogenous, monoculture crops (e.g., sugarcane). Vynne et al. (2014) found that Maned Wolves may be tolerant of these habitat conversions as long as there are sufficient remnant vegetation patches for den sites, food, water and resting, including gallery forests (Dietz 1984), cerrado and marshy areas near rivers (Bestelmeyer 2000, F. Rodrigues pers. comm. 2015).

The omnivorous diet of the Maned Wolf includes a wide variety of plant material and animal matter, including fruits, arthropods and small- to medium-sized vertebrates (Dietz 1984, Carvalho and Vasconcellos 1995, Motta-Jnior et al. 1996, Azevedo and Gastal 1997, Motta-Jnior 1997, Rodrigues et al. 1998, Jcomo 1999, Santos 1999, Silveira 1999, Juarez and Marinho-Filho 2002, Amboni 2007). Although the frequency of plant and animal items found in faecal samples is approximately equal, the biomass of animal items is usually greater than that of plant items (Motta-Jnior et al. 1996, Santos 1999, Motta-Junior et al. 2014). The relative importance of specific food items may vary depending on location and season (Bueno and Motta-Jnior 2006, Amboni 2007). In areas of high food availability, the degree of overlap of home ranges increases and can reach levels of 92% between female-female and 100% for male-female (couple; Amboni 2007, Azevedo 2008, R. Paula pers. obs.).

While historically Maned Wolves were considered to be crepuscular-nocturnal in their activities, recent observations in Bolivia and Brazil reveal that their activity is actually variable or cathermeral, with activity patterns changing according to daily temperatures and the season (dry or wet; Emmons 2012). In Santa Cruz, Bolivia, Emmons (2012) found Maned Wolves tend to forage for longer distances when temperatures are cooler (between 21-28C) and show considerable decreases in activity levels when temperatures increase (>28C). In addition, while the Maned Wolf is strictly nocturnal during the dry season, it varies in its temporal activity pattern during the wet season (Emmons 2012). These patterns shift some in southeastern Brazil, an area defined by higher humidity levels, abundant permanent water sources in the form of rivers, well-defined seasons reflecting extreme temperature differences, and shifts in the seasonal food availability. While the activity pattern of Maned Wolves in Brazil is cathemeral and affected by overall temperature and seasonality, the high degree of daily variation in minimum and maximum temperature in a season means that there is additional variation in the daily pattern each season (R. Paula pers. obs.). For example, during the dry season when temperatures drop to 5C during the night, the species is more diurnal having higher activity during the day when temperatures are likely to be warmer. In contrast during the wet season activity is typically nocturnal, except after extended periods of rain when the species can be active during the middle of the day.

The home range of Maned Wolves is variable, ranging from 20-115 km2 (fixed kernel, 95%). Despite considerable variation ranges average between 50-80km2(fixed kernel, 95%; Rodrigues 2002, Coelho et al. 2008, Azevedo 2008, Emmons 2012) and about 30km2in agricultural landscape of northern Argentina (L. Soler pers. comm. 2015). Food availability and habitat integrity have been linked to variation in home range size (Amboni 2007, Azevedo 2008, R. Paula pers. obs.). Sharing of home ranges has been noted in Brazil and Bolivia (Azevedo 2008, Emmons 2012). While home range overlap can be found during the breeding season, it is possible throughout the year in areas with high food availability. In these areas, the greatest sharing of home ranges is between females followed by male-female pairs with very little sharing tolerance noted among males (Azevedo 2008).

In northern Argentina the Maned Wolf shows a preference for open grassland habitats and also utilizing human disturbed areas such as cattle ranches (Soler and Carenton 2008). Direct competition with other carnivores has not been observed; however, scat analyses reveal that Maned Wolves, Crab-eating Fox (Cerdocyon thous) and Pampas Fox (Lycalopex gymnocercus) and Crab-eating Raccoon (Procyon cancrivorus) consume many of the same prey species (Soler and Carenton 2008).

Paula et al. (2008) calculated a generation time of 5.3 years in Vortex (using the exponential growth rate as another variable to account to the net reproductive rate calculations, which is the age-specific survival multiplied by fecundity over all age classes; see Lacy et al. 2015 for further details). Using the formula Sum (x lx mx)/Sum (lx mx) where lx represents the survivourship over x years and mx the maternity over x years, Paula and Desbiez (2014) calculated a generation length of 7 years. No differences on population reduction were observed when using the two different generation times.

Systems
  • Terrestrial
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Chrysocyon brachyurus is found in grassland, savanna, dry shrub forest, swampy areas, forest-edge habitat, and river areas.

Terrestrial Biomes: savanna or grassland ; scrub forest

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The maned wolf prefers open habitats in tall grasslands, low-scrub edges of forests and even swampy areas (2). In Brazil, this species is found in the cerrado, a large area of open woodland and savannah that is one of the world's most important 'hot-spots' of biodiversity (9).
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Trophic Strategy

The maned wolf is omnivorous. It eats armadillos, rabbits, rodents and other small mammals, fish, birds, bird eggs, reptiles, gastropods and other terrestrial mollusks, insects, seasonably available fruits, and other vegetation. Fruits taken include bananas, guavas, and primarily the tomato-like Solanum lycocarpum. (S. lycocarpum may provide medicinal aid against Dioctophyme renale, a worm that infects the kidneys of the maned wolf). Vegetation eaten is often in the form of roots and bulbs. Vertebrate prey do not often include large domestic stock, but an occasional newborn lamb or pig is taken by Chrysocyon. The maned wolf, much to the dislike of poultry farmers, frequently feeds upon free-ranging chickens. It stalks and pounces in a fox-like manner upon its animal prey.

Animal Foods: birds; mammals; reptiles; fish; eggs; insects; mollusks

Plant Foods: roots and tubers; fruit

Primary Diet: omnivore

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Life History and Behavior

Behavior

Perception Channels: tactile ; chemical

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Life Expectancy

Average lifespan

Status: captivity:
13.0 years.

Average lifespan

Status: captivity:
15.0 years.

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Lifespan, longevity, and ageing

Maximum longevity: 16.8 years (captivity) Observations: In captivity, these animals can live up to 16.8 years (Richard Weigl 2005).
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Reproduction

Maned wolves are monogamous, though males and females tend to live independently except during the breeding season.

Mating System: monogamous

Little is known about the reproductive patterns of wild maned wolves. Females are monoestrous. Breeding season is probably controlled by photoperiod; captives copulate between October and February in the Northern Hemisphere and between August and October in South America. The estrous lasts for a period of one to four days. Gestation in captivity is similar to that of other canids and lasts approximately 65 days. A litter usually contains one to five young. A record number of seven has been observed. Young are born weighing 340 to 430 grams and develop quickly. Their eyes and ears open by day nine, their ears stand upright and they will take regurgitated food by week four, the pelage changes from black to red by week ten, they are weaned by 15 weeks, and their bodies have the proportions of adults at one year, at which time they reach sexual maturity. Captive individuals have lived 15 years. Non-captive maned wolves give birth in natal nests hidden by thick vegetation. Wild maned wolves are rarely seen with their pups.

Range number of offspring: 1 to 5.

Average number of offspring: 2.47.

Range gestation period: 56 to 66 days.

Range weaning age: 120 to 210 days.

Key Reproductive Features: gonochoric/gonochoristic/dioecious (sexes separate); sexual

Average birth mass: 368 g.

Average number of offspring: 3.

Average age at sexual or reproductive maturity (female)

Sex: female:
730 days.

Parental Investment: altricial

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Molecular Biology and Genetics

Molecular Biology

Statistics of barcoding coverage: Chrysocyon brachyurus

Barcode of Life Data Systems (BOLDS) Stats
Public Records: 0
Specimens with Barcodes: 3
Species With Barcodes: 1
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Conservation

Conservation Status

IUCN Red List Assessment


Red List Category
NT
Near Threatened

Red List Criteria

Version
3.1

Year Assessed
2015

Assessor/s
Cunha de Paula, R. & DeMatteo, K.

Reviewer/s
Hoffmann, M. & Sillero-Zubiri, C.

Contributor/s
Giordano, A., Orozco, M., Soler, L., Gonzalez Ciccia, P., Thresher, S., Rodden, M., Rodrigues, F. & Bestelmeyer, S.V.

Justification
The current population of Maned Wolves is estimated at approximately 17,000 mature individuals ( 2 years of age), with the majority of the population (>90%) in Brazil. In the last decade or so, the species main habitats in Brazil have been subject to intense deforestation. A population viability model for Brazil generated using real and predicted deforestation rates over 15 years (three generations) resulted in an estimated reduction of ~20% in the metapopulation (based on current estimates of habitat loss ranging from 1.0 to 1.5% per year). If deforestation reaches the maximum estimated rate of loss (2% per year), then simulation suggests a national population reduction of 56% at the end of 100 years (Paula et al. 2008, Paula and Desbiez 2014). In addition to the estimated population reduction from deforestation, the species is also subject to other threats, including road kills, direct persecution by humans, and disease due to contact with domestic animals (Paula and Desbiez 2014). In other range countries (Argentina, Paraguay and Bolivia), the species status is even more precarious with small isolated populations and declining numbers due to the low quality of habitat and hunting. The Maned Wolf does not meet the thresholds for listing under criteria B, C or D, but taking into account the combined impacts of habitat loss, persecution and disease, it approximates the thresholds for listing under A3, and is therefore assessed as Near Threatened. The species situation is considered to be of major concern due to the various threats acting throughout its entire range and should be regularly re-evaluated.

History
  • Near Threatened (NT)
  • 2004
    Near Threatened (NT)
  • 1996
    Lower Risk/near threatened (LR/nt)
  • 1994
    Vulnerable (V)
  • 1990
    Vulnerable (V)
  • 1988
    Vulnerable (V)
  • 1986
    Vulnerable (V)
  • 1982
    Vulnerable (V)
  • 1965
    Status inadequately known-survey required or data sought
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Chrysocyon brachyurus is listed as CITES Appendix II, U.S. ESA-Endangered, and IUCN-Vulnerable. Habitat destruction (including the annual burning of its grasslands), persecution by angry poultry farmers, hunting for sport, and live capture are factors that threaten the maned wolf. This animal disapeared from Uruguay in the 19th Century. Its former range also included parts of Argentina south of the La Plata River.

CITES: appendix ii

IUCN Red List of Threatened Species: near threatened

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Current Listing Status Summary

Status: Endangered
Date Listed: 12/02/1970
Lead Region: Foreign (Region 10) 
Where Listed: Entire


Population detail:

Population location: Entire
Listing status: E

For most current information and documents related to the conservation status and management of Chrysocyon brachyurus , see its USFWS Species Profile

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Status

Classified as Near Threatened (NT) on the IUCN Red List (1), and listed on Appendix II of CITES (4).
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Population

Population
A Population and Habitat Viability Assessment (PHVA) workshop held in 2005 estimated the total population (newborn pups, juveniles and adults) of Maned Wolves at ~23,600 animals, including 21,746 in Brazil, 830 in Paraguay, and 660 in Argentina (Paula et al. 2008). If only mature individuals ( 2 years of age) are considered, the population size is estimated at 17,000 animals, including 15,849 in Brazil, 613 in Paraguay, and 487 in Argentina (Paula et al. 2008). Numbers in Bolivia are unlikely to exceed 1,000 animals. Numbers in Pampas del Heath, Peru are less than 40 individuals, declining, and likely geographically isolated from the adjacent Bolivian population (Williams et al. 2012). The population estimates from the PHVA workshop were generated using Vortex simulation computer models and so do require some caution (Paula et al. 2008). Even though the species has been supposedly expanding into new areas in Brazil, the species overall range has contracted in the Brazilian Cerrado (its typical habitat) and in other countries. In addition, over the last 10 years some of the areas previously defined as the most suitable in Brazil have suffered from habitat degradation, suggesting that these prior published estimates may now be considered high.

To account for the effects of habitat loss, one of the main threats to Maned Wolf, on persistence, simulations of the effects of habitat loss using detailed data on deforestation rates in Brazil were performed (and measured as permanent reduction in K or carrying capacity). This was modelled as a linear decrease (1.5% / annum) in carrying capacity over one generation or five years, which effectively reduced the maximum size a population could attain. The simulation model showed that small populations were affected more than large populations, with populations <25 individuals showing negative growth rates due to their vulnerability to stochastic processes. Populations with 25 individuals declined with a high probability of extinction within 100 years; however, populations of 50-100 individuals persisted, albeit with low levels of genetic diversity. Therefore, several hundred wolves may be needed to maintain a long-term, viable population depending upon the severity of threats, management goals, and acceptable levels of risk for wildlife managers.

In the last 10 years, the main habitats of Maned Wolf in Brazil have been subject to intense deforestation. For the current assessment, a population viability model was generated using recent observed deforestation rates, ranging from 1.0 to 1.5% per year over 15 years (three generations). These results suggested a 20% reduction in the metapopulation based on these rates of loss. If deforestation rates reach a loss of 2% per year, the simulation suggests a 56% reduction in the Brazilian population in 100 years (Paula et al. 2008, Paula and Desbiez 2014). The areas of supposed distribution expansion were not accounted for in this simulation since they represent edge areas for the distribution range and because from what is known presently these areas do not seem to hold stable and resident populations of the species.

Maned Wolves are found at low densities throughout the range. Even in Brazilian protected areas, population densities range from 0.01 to 0.05 animals/km2 (Silveira 1999, Rodrigues 2002). The highest density reported is 0.08 individuals/km2in Serra da Canastra, Minas Gerais, Brazil (Paula et al. 2013). In Argentina, information about wild populations is scarce and the northern population is thought to be declining (Soler and Carenton 2008). Despite the lack of information on population densities across it range, high food availability may potentially support higher population densities (Amboni 2007, Azevedo 2008).

Population Trend
Unknown
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Threats

Major Threats
The four main threats affecting Maned Wolf populations throughout their distribution range are: habitat loss and alteration, especially due to conversion to crop fields and pasturelands; human persecution due to livestock losses and cultural beliefs; increasing vehicular traffic in highways resulting in road kills; and pathogens contracted from domestic animals due to increased contact in disturbed environments (Paula et al. 2008, 2013, Emmons 2012, Orozco et al. 2013a).

The intensive conversion of native habitat for agriculture results in loss of optimal habitat and creation of areas often subject to desertification. The fragmentation of highly suitable habitat causes isolation of subpopulations and often results in the remaining intact landscape become intersected by high speed roads. Many Maned Wolves are killed on roadways throughout its range. In central Brazil, road kills are a threat of major concern potentially leading to local extinction of small, isolated populations as estimates range from four to 10 individuals killed/year in some regions (Paula and Gambarini 2013, Rodrigues et al. 2014). Similar impact of vehicular collisions was detected in marginal populations from Argentina where 21 Maned Wolves were reported dead along National Route 34, concentrated between 2000 and 2005 (Orozco et al. 2013a). The same negative effect is potentially true in the humid Chaco of Paraguay where increased traffic on the Trans-Chaco highway leads to numerous road kills each year (A.J. Giordano pers. comm. 2015). Extreme climatic conditions such as uncontrolled fires and droughts are likely to exacerbate the scarcity of living resources and consequently the loss on roadways, as Maned Wolves are forced to travel farther for food and water (Pautasso et al. 2009, Orozco et al. 2013a); the latter is especially important during periods when females are lactating (Emmons 2014).

Direct interactions with humans also pose a threat to the species. Conflicts with people, road mortality, the potential for infectious diseases spread by domestic dogs and direct persecution resulting from widely held superstitions and beliefs, are the primary threats facing Maned Wolves (Songsasen and Rodden 2010). In some areas of Argentina, Maned Wolves are persecuted with hunting, trapping, and shooting due to a mix of long-standing cultural beliefs, general ignorance of the species, and trophy hunting (Orozco et al. 2009, Orozco et al. 2013a, Soler 2014). In southeastern Santiago del Estero (Argentina), 30 Maned Wolves were reported sold to game ranches in a 10-year period (Orozco et al. 2013a). In Brazil and Paraguay, this targeted persecution of Maned Wolves extends beyond the traditional and natural medicine needs to include the low public tolerance of poultry depredation, for which Maned Wolves are constantly blamed (Paula et al. 2013, Cartes et al. 2014, A.J. Giordano pers. comm. 2015). In some areas this hunting pressure is so high it is having a significant negative effect on local Maned Wolf numbers (Paula et al. 2013). While illegal trade is not common, Orozco et al. (2009) found it was widespread in Argentina, and even included the sale of young animals or pups to private zoos and collections. Of the Maned Wolves kept in captivity in Argentina during the last five years 68% were wild born and came from/were victims of illegal trade, road collisions or were orphaned after their mother was killed (P. Gonzalez Ciccia pers. comm. 2015).

Domestic dogs pose a serious threat to Maned Wolves on several levels, as often times domestic dogs accompany humans into the field for recreation and hunting. In addition, both Maned Wolves and dogs overlap in their use of habitat that has been converted to agriculture and pasture. Dogs are known to pursue Maned Wolves, which often results in the killing of the animal (Soler et al. 2005, Pautasso et al. 2009, Orozco et al. 2013a, Soler 2014, A.J. Giordano pers. comm. 2015). In a study carried out in Argentina, 26% of sightings by local villagers involved close contact between Maned Wolves and domestic dogs (Orozco et al. 2013a). Dogs are also a source of infectious diseases and parasites. However the epidemiological effects remain unclear for Maned Wolf conservation. Rural domestic dog populations in Argentina had very high seroprevalence for Neospora caninum, canine coronavirus, canine adenovirus,Dirofilaraimmitis, canine distemper virus, Toxoplasma gondii, and L. interrogans spp. In the same area, all of these agents (except N. caninum, Toxoplasma gondii and CCV) were serologically positive in local Maned Wolves (Orozco et al. 2013a). In a Bolivian population, Maned Wolves were serologically positive for two domestic dog pathogens (morbillivirus and parvovirus) that can cause pup mortality and high levels of potentially fatal (Dirofilara immitis) and debilitating (Dioctophyme renale) parasites were detected (Deem and Emmons 2005, Bronson et al. 2008, Deem et al. 2012, Emmons 2014). In Brazil, Maned Wolves were serologically positive for several dog-related infectious diseases including parvovirus, canine distemper virus, coronavirus, and leishmaniosis (Paula et al. 2014). These data from Bolivia, Brazil and Argentina suggest that other wild populations are at risk in areas where domestic dogs are present (May Jr and Felippe 2014). The frequent use of dogs for herding and hunting, along with their frequent foraging in natural habitats, could favour direct or indirect transmission of multiple pathogens (Orozco et al. 2013a).
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The most significant threat to the survival of remaining maned wolf populations is habitat loss (8). The conversion of land to agriculture has drastically reduced the available habitat for the maned wolf, with the cerrado of Brazil being reduced to about 20 percent of its original extent (8). In addition maned wolves are often killed on highways, frequently on those which border protected areas. Indeed, road kills are responsible for the death of approximately half the annual production of pups in some reserves (8). Domestic dogs also pose a threat by transferring diseases, competing for food, and even killing the maned wolf (1). Some local people attribute mystical qualities to several parts of the wolf's anatomy (eyes, skin, tail) and still hunt this threatened species in order to use these parts as 'talisman' or for medicinal remedies (6). Occasionally, this wolf is hunted for sport (5), and, due to the wolf's threat to domestic poultry, farmers also hunt it as a pest (6). As its habitat is encroached upon by ever-expanding farms, the wolf is forced into increased proximity with people, exacerbating the already-existing conflict (6).
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Management

Conservation Actions

Conservation Actions
Legislation
It is included on CITES Appendix II. Maned Wolves are protected by law in many parts of their range, but enforcement is frequently problematic. This species is classified as Endangered in Argentina (Ojeda et al. 2012) and declared a Provincial Natural Monument in select areas, including Santa Fe, Corrientes, and Chaco (Soler 2014). It is classified as Vulnerable in Brazil (Paula et al. 2013). Hunting is prohibited in Brazil, Paraguay, Bolivia and in Argentina. The species is included in the United States Endangered Species list.

Presence in protected areas
This species occurs in many protected areas in Argentina, Bolivia, Brazil, Paraguay and Peru

Occurrence in captivity
As of January 2012, 150 institutions held a total of 394 Maned Wolves (182 males, 211 females, one unknown) in captivity (Holland 2014).

Other
In Brazil, conservation actions to reverse population decline have focused on addressing the issues associated with the human dimension. Raising public awareness and using proactive methods to prevent conflict are among the main tools used in increasing tolerance towards the species. In addition, campaigns to vaccinate domestic dogs in areas that border or overlap with Maned Wolf populations have been ongoing for several years (Paula et al. 2014). New proposals to reduce the high number of road kills in southeast Brazil have been discussed at a political level. The 2005 Population and Habitat Viability Assessment workshop for Maned Wolves generated an Action Plan aimed at addressing the five main themes that effect the long-term conservation of the species across its distribution: 1) threats and habitat management; 2) distribution and status; 3) environmental education, social aspects, and economic alternatives; 4) ex-situ conservation; and 5) population dynamics and modelling (Paula et al. 2008). Each of these themes address gaps in knowledge for the species or specific problems associated with the species survival. Each theme was broken into multiple goals with specific actions set to address the associated problems, names of personnel responsible for seeing the action is completed, timeline for when these actions should be completed, associated costs, potential obstacles, and expected outcome defined (Paula et al. 2008). As a follow up to this workshop, Brazil and Argentina used information on recent and ongoing changes in the species status and threats to modify the international Action plan and generate specific National Action Plans that could help direct local efforts in Maned Wolf conservation. In 2014, the Brazilian government officially recognized the technical advisory group (originally formed in 2010) aimed at assisting the government on implementing the strategies developed in the National Action Plan. A Maned Wolf working group has also been implemented in continental level under the coordination of the IUCN SSC Canid Specialist Group, with an aim to ensure long-term coordination among researchers across the Maned Wolves distribution and work together to share strategies directed at reversing population decline.

In Argentina, the GAAG (Grupo Argentino Aguar Guaz or Maned Wolf Argentine Group) was founded as a national strategy to develop, manage, execute, and monitor the Action Plan for the Maned Wolfs conservation in Argentina. The group comprises 16 institutional members, which includes provincial and national government agencies, zoos, NGOs and research groups from universities and museums. Between 2002 and 2011, GAAG carried out 10 regional workshops aimed at: 1) mapping threats for Maned Wolf in natural habitats; 2) prioritizing conflicts in the wild and problems in captivity; 3) prioritizing strategies and actions for its ex situ and in situ conservation; 4) developing recommendations for conservation in the wild and management in captivity; 5) developing efficient education strategies for Maned Wolf conservation both in situ and ex situ, and 6) validating methodologies and strategies for conservation education. Since 2005, five species-directed projects have been developed and include work in five of the eight provinces in the Maned Wolfs distribution (Orozco et al. 2013b). Extensive surveys of farmers and ranchers have provided valuable information about the distribution of the species and also about people's attitudes towards Maned Wolves (Songsasen and Rodden 2010). Ongoing education programs are aimed at changing negative perceptions of this wild carnivore (Soler 2008, Orozco et al. 2013b).
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Conservation

The maned wolf occurs in a number of protected areas across its range. Although protected by law in certain countries, with hunting prohibited in Brazil, Paraguay and Bolivia, law enforcement is often problematic. At present, there are no known conservation actions specific to the maned wolf, but there are broader attempts to protect parts of its habitat and reduce the impact of animal road kills in Brazil (1). Encouragingly, observations indicate that the maned wolf is able to colonize different habitats and that the species' range has altered in configuration in recent years rather than diminished (12). This has, however, led these wolves into areas of greater proximity and conflict with humans, and education programmes have therefore been started to dissuade farmers from shooting this rare species (2). As of 2003, 146 institutions reported a total of 431 maned wolves in captivity, including 208 males and 222 females (8). However, for unknown reasons, canids breed poorly in captivity. Research has therefore been conducted into behaviour affecting hormones, nutrition and stress in captivity, as well as the use of modern reproductive technologies to aid the process (10). Future studies need to focus on population surveys throughout the species' range, as well as research into how human encroachment and habitat loss is impacting this distinctive canid (1).
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Relevance to Humans and Ecosystems

Benefits

As mentioned above, the maned wolf takes domestic poultry and the occasional lamb or newborn pig.

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The maned wolf eats crop pests such as rabbits and small rodents.

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Wikipedia

Maned wolf

The maned wolf (Chrysocyon brachyurus) is the largest canid of South America. Its markings resemble those of foxes, but it is not a fox, nor is it a wolf, as it is not closely related to other canids. It is the only species in the genus Chrysocyon (meaning "golden dog").

This mammal is found in open and semi-open habitats, especially grasslands with scattered bushes and trees, in south, central-west, and southeastern Brazil (Mato Grosso, Mato Grosso do Sul, Minas Gerais, Goiás, São Paulo, Federal District, and recently, Rio Grande do Sul), Paraguay, northern Argentina, Bolivia east and north of the Andes,[4] and far southeastern Peru (Pampas del Heath only).[5] It is very rare in Uruguay, possibly being displaced completely through loss of habitat.[2] IUCN lists it as near threatened,[2] while it is considered vulnerable by the Brazilian government (IBAMA).

It is known locally as aguará guazú (meaning "large fox" in the Guarani language), or "kalak" by the Toba, lobo de crin, lobo de los esteros, or lobo colorado, and as lobo-guará in Brazil. It also is called borochi in Bolivia.

Description[edit]

Video of captive maned wolves at Ueno Zoo, in Japan
Drawing of the skull of a maned wolf

The maned wolf bears minor similarities to the red fox, although it belongs to a different genus. The average adult weighs 23 kg (51 lb) and stands 90 cm (35 in) tall at the shoulder, has a head-body length of 100 cm (39 in) with the tail adding another 45 cm (18 in).[6]

The maned wolf is the tallest of the wild canids, its long legs probably are an adaptation to the tall grasslands of its native habitat.[7] Fur of the maned wolf may be reddish brown to golden orange on the sides with long, black legs, and a distinctive black mane. The coat is marked further with a whitish tuft at the tip of the tail and a white "bib" beneath the throat. The mane is erectile, and typically, is used to enlarge the wolf's profile when threatened or when displaying aggression.

The maned wolf also is known for the distinctive odor of its territory markings, which has earned it the nickname "skunk wolf."

Habits[edit]

Hunting and territoriality[edit]

Unlike other large canids (such as the gray wolf, the African hunting dog, or the dhole) the maned wolf does not form packs.[6] It hunts alone, usually between sundown and midnight. It kills its prey by biting on the neck or back, and shaking the prey violently if necessary.[8] Monogamous pairs may defend a shared territory of approximately 30 km2 (12 sq mi), although outside of mating, the individuals may meet seldom. The territory is crisscrossed by paths that the maned wolves create as they patrol at night. Several adults may congregate in the presence of a plentiful food source, for example, a fire-cleared patch of grassland that would leave small vertebrate prey exposed while foraging.

A maned wolf and pup at White Oak Conservation

Both female and male maned wolves use their urine to communicate, e.g. to mark their hunting paths or the places where they have buried hunted prey.[8] The urine has a very distinctive odor, which some people liken to hops or cannabis. The responsible substance very likely is a pyrazine, which also occurs in both plants.[9] (At the Rotterdam Zoo, this smell once set the police on a hunt for cannabis smokers.[9][10]) The preferred habitat of the maned wolf include grasslands, scrub prairies, and forests.

Reproduction[edit]

Maned wolf pup

Their mating season ranges from November to April. Gestation lasts 60 to 65 days and a litter may have from two to six black-furred pups, each weighing approximately 450 g (16 oz). Pups are fully grown when one year old. During that first year, the pups are known to rely on their parents for food.[8]

Diet[edit]

The maned wolf is omnivorous. It specializes in small and medium-sized prey, including small mammals (typically rodents and rabbits), birds, and even fish,[11][8] but a large portion of its diet (more than 50%, according to some studies) is vegetable matter, including sugarcane, tubers, and fruit (especially the wolf apple, Solanum lycocarpum, a tomato-like fruit).[12] Traditionally, captive maned wolves were fed meat-heavy diets, but that caused them to develop bladder stones. Zoo diets for them now feature fruits and vegetables, as well as meat and dog chow.

Relations with other species[edit]

The maned wolf participates in symbiotic relationships. It contributes to the propagation and dissemination of the plants that it feeds on, through excretion. Often maned wolves defecate on the nests of leafcutter ants. The ants then use the dung to fertilize their fungus gardens, but they discard the seeds contained in the dung onto refuse piles just outside their nests. This process significantly increases the germination rate of the seeds.[13]

The maned wolf is not a common prey species for any predator, although it may be attacked or killed by feral dogs. An additional threat to the maned wolf exists from sharing territory with domestic dogs. The maned wolf is particularly susceptible to infection by the giant kidney worm, a potentially fatal parasite that also may infect domestic dogs.

Relations with humans[edit]

Generally, the maned wolf is shy and flees when alarmed, so it poses little direct threat to humans. Popularly, the maned wolf is thought to have the potential of being a chicken thief. It once was considered a similar threat to cattle and sheep, although this now is known to be false.

Historically, in a few parts of Brazil, these animals were hunted down for some body parts, notably the eyes, that were believed to be good luck charms. Since its classification as a vulnerable species by the Brazilian government, it has received greater consideration and protection.

They are threatened by habitat loss and being run over by automobiles. Feral and domestic dogs pass on diseases to them, and have been known to attack them.

The species occurs in several protected areas, including the national parks of Caraça and Emas in Brazil. The maned wolf is well represented in captivity and has been bred successfully at a number of zoos, particularly in Argentina.

Taxonomy[edit]

Although the maned wolf displays many fox-like characteristics, it is not closely related to foxes. It lacks the elliptical pupils found distinctively in foxes. The maned wolf's evolutionary relationship to the other members of the canid family makes it a unique animal.

Electrophoretic studies did not link Chrysocyon with any of the other living canids studied. One conclusion of this study is that the maned wolf is the only species among the large South American canids that survived the late Pleistocene extinction. Fossils of the maned wolf from the Holocene and the late Pleistocene have been excavated from the Brazilian Highlands.[14]

A study, published in 2003,[15] on the brain anatomy of several canids, placed the maned wolf together with the Falkland Islands wolf and with pseudo-foxes of the genus Pseudalopex. One study based on DNA evidence, published in 2009, showed that the extinct genus Dusicyon, the Falkland Islands wolf and its mainland relative, was the most closely related species to the maned wolf in historical times, and that about seven million years ago it shared a common ancestor with that genus.[16]

The maned wolf is not closely related to any other living canid. It is not a fox, wolf, coyote, dog, or jackal, but a distinct canid. Although based only on morphological similarities, it previously had been placed in Canis and Vulpes genera.[3] Its closest living relative is the bush dog (genus Speothos) and it has a more distant relationship to other South American canines (the short-eared dog, the crab-eating fox, and the 'false foxes' or Pseudalopex).[17]



Short-eared dog




Crab-eating fox







Sechuran fox



Culpeo fox





Pampas fox



South American gray fox





Darwin's fox




Hoary fox







Maned wolf[18](Fig. 10)



Bush dog



Gallery[edit]

References[edit]

  1. ^ Wozencraft, W. C. (2005). "Order Carnivora". In Wilson, D. E.; Reeder, D. M. Mammal Species of the World (3rd ed.). Johns Hopkins University Press. pp. 532–628. ISBN 978-0-8018-8221-0. OCLC 62265494. 
  2. ^ a b c Rodden, M., Rodrigues , F. & Bestelmeyer, S. (2008). Chrysocyon brachyurus. In: IUCN 2008. IUCN Red List of Threatened Species. Retrieved 22 March 2009. Database entry includes justification for why this species is near threatened.
  3. ^ a b Osgood, Wilfred H. (1919). "Names of Some South American Mammals". Journal of Mammalogy 1 (1): 35. doi:10.2307/1373718. JSTOR 1373718. 
  4. ^ Langguth, A. (1975). "Ecology and evolution in the South American canids". In M. W. Fox, ed. The wild canids: their systematics, behavioral ecology and evolution. New York: Van Nostrand Reinhold Company. pp. 192–206. ISBN 0442224303. 
  5. ^ Sillero-Zubiri, Hoffmann, & Macdonald (eds). 2004.Canids: Foxes, Wolves, Jackals and Dogs – 2004 Status Survey and Conservation Action Plan. IUCN/SSC Canid Specialist Group.
  6. ^ a b Dietz, J. M. (1984). "Ecology and social organization of the maned wolf (Chrysocyon brachyurus)". Smithsonian Contributions to Zoology 392 (392): 1–51. doi:10.5479/si.00810282.392. 
  7. ^ Dietz, James (1984). Macdonald, D., ed. The Encyclopedia of Mammals. New York: Facts on File. p. 31. ISBN 0-87196-871-1. 
  8. ^ a b c d Cristian Frers. "Un lobo de crin llamado Aguará Guazú". Retrieved 2007-04-23. 
  9. ^ a b Brian Switek (2011-03-10). "Maned Wolf Pee Demystified". Wired. Retrieved 2011-06-05. 
  10. ^ Süddeutsche Zeitung, 2006-09-02, p3
  11. ^ Juarez, Keila Macfadem; Jader Marinho-Filho (November 2002). "Diet, habitat use, and home ranges of sympatric canids in central Brazil". Journal of Mammalogy 83 (4): 925–934. doi:10.1644/1545-1542(2002)083<0925:DHUAHR>2.0.CO;2. 
  12. ^ Motta-Junior, J. C., S. A. Talamon, J. A. Lombardi, AND K. Simokomaki (1996). "Diet of maned wolf, Chrysocyon brachyurus, in central Brazil". Journal of Zoology (London) 240 (2): 277–284. doi:10.1111/j.1469-7998.1996.tb05284.x. 
  13. ^ Courtenay, O. (1994). "Conservation of the Maned Wolf: fruitful relationships in a changing environment". Canid News 2. 
  14. ^ [1]
  15. ^ Lyras, G.A., Van der Geer, A.A.E. 2003. External brain anatomy of the Canidae. Zoological Journal of the Linnean Society 138: 505–522. London. doi:10.1046/j.1096-3642.2003.00067.x
  16. ^ Scientists solve the 320-year-old mystery of how the Falklands wolf ended up on the island: It skated across a frozen sea chasing a penguin
  17. ^ Kerstin, Lindblad-Toh; Claire M Wade, Tarjei S. Mikkelsen, Elinor K. Karlsson, David B. Jaffe, Michael Kamal, Michele Clamp, Jean L. Chang, Edward J. Kulbokas III, Michael C. Zody, Evan Mauceli, Xiaohui Xie, Matthew Breen, Robert K. Wayne, Elaine A. Ostrander, Chris P. Ponting, Francis Galibert, Douglas R. Smith, Pieter J. deJong, Ewen Kirkness, Pablo Alvarez, Tara Biagi, William Brockman, Jonathan Butler, Chee-Wye Chin, April Cook, James Cuff, Mark J. Daly, David DeCaprio, Sante Gnerre, Manfred Grabherr, Manolis Kellis, Michael Kleber, Carolyne Bardeleben, Leo Goodstadt, Andreas Heger, Christophe Hitte, Lisa Kim, Klaus-Peter Koepfli, Heidi G. Parker, John P. Pollinger, Stephen M. J. Searle, Nathan B. Sutter, Rachael Thomas, Caleb Webber (2005-12-08). "Genome sequence, comparative analysis and haplotype structure of the domestic dog". Nature 438 (7069): 803–819. doi:10.1038/nature04338. PMID 16341006. 
  18. ^ Lindblad-Toh et al. (2005). "Genome sequence, comparative analysis and haplotype structure of the domestic dog". Nature 438 (7069): 803–819. doi:10.1038/nature04338. PMID 16341006. 
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