Behavioral and physiologic responses to environmental enrichment in the maned wolf (Chrysocyon brachyurus)

The ex situ population of maned wolves is not self‐sustaining due to poor reproduction, caused primarily by parental incompetence. Studies have shown that environmental enrichment can promote natural parental behaviors in zoo animals. The objective of this study was to determine the effects of environmental enrichment on behavioral and physiological responses of maned wolves. During an 8‐week experimental period, daily behavior observations and fecal sample collection were conducted on four adult wolves (2.2) individually housed in environments without enrichment. After 2 weeks, the wolves were chronologically provided with 2‐week intervals of hiding dead mice around the exhibit, no enrichment, and introduction of boomer balls. Responses of the wolves to enrichment were assessed based on activity levels and exploratory rates, as well as the level of corticoid metabolites in fecal samples collected daily throughout the study period. Providing wolves with environmental enrichment significantly increased exploratory behaviors (P<0.05), especially when mice were hidden in the enclosure. Fecal corticoid concentrations were increased during periods of enrichment in males (P<0.05), but not in females. Overall, there were no correlations between behavioral responses to enrichment and fecal corticoid levels. Behavioral results suggest that environmental enrichment elicits positive effects on the behavior of captive maned wolves. There is evidence suggesting that providing animals with ability to forage for food is a more effective enrichment strategy than introducing objects. There is need for a longer term study to determine the impact of environmental enrichment in this species. Zoo Biol 26:331–343, 2007. © 2007 Wiley‐Liss, Inc.     

Diet of the maned wolf, Chrysocyon brachyurus, in central Brazil

Analysis of 105 scats of the maned wolf ( Chrysocyon brachyurus ) collected in central Brazil yielded 304 occurrences of food items of which fruits of Solanum lycocarpum , rodents, and birds accounted for 61.5%. By analysis of minimum number of individual animals, rodents and birds were 75% of a total of 156 prey. On the other hand, fruits of S. lycocarpum and armadillos ( Dasypus spp.) were the bulk of the total estimated biomass consumption, yielding 63.7% in a total of 73.5 kg. Rodents were mostly captured during the dry season, while miscellaneous fruits were consumed mostly in the wet season. The consumption of S. lycocarpum fruits and armadillos was aseasonal. Small rodents were taken in about the ranks of abundance in the study area, but S. lycocarpum fruits were actively searched by maned wolves, for its occurrence is limited to secondary savanna. Prey and fruits typical of savanna ('cerrado') and grassland ('campo') were the bulk of species, occurrences, prey numbers, and biomass consumed. These findings reinforce the importance of conservation of cerrado and campo in central Brazil. Despite maned wolf being a vulnerable species, its feeding habits are opportunistic, and the main reasons for its decline are likely to be habitat destruction and human disturbance.     

Crystal structure of hemoglobin from the maned wolf (Chrysocyon brachyurus) using synchrotron radiation

Crystal structure of hemoglobin isolated from the Brazilian maned wolf (Chrysocyon brachyurus) was determined using standard molecular replacement technique and refined using maximum-likelihood and simulated annealing protocols to 1.87A resolution. Structural and functional comparisons between hemoglobins from the Chrysocyon brachyurus and Homo sapiens are discussed, in order to provide further insights in the comparative biochemistry of vertebrate hemoglobins.     

Feed intake and digestion in the maned wolf (Chrysocyon brachyurus): Consequences for dietary management

Captive maned wolves (Chrysocyon brachyurus) often consume diets high in prey and meat items even though they are omnivorous in the wild. These soft, high protein diets may exacerbate conditions of gingivitis and cystinuria in this species. Feed intakes were monitored in wolves provided with prey and meat-based diets (3 periods) and subsequently with extruded dog chows and small amounts of prey (2 periods). Digestibilities of each type of diet were measured with the indigestible marker chromic oxide. Dry matter intakes were similar between diets (374–584 g · 30 kg^?1 · d^?1), even though dietary protein content was reduced from 44% to 29% of dry matter. Digestibility of dry matter was unaltered by these dietary changes (77% of high prey diet vs. 73% of chow diet), but fecal consistency changed to softer stools at lower intakes of prey. Estimated metabolizable energy intakes of wolves (501–674 kJ · kg^?0.75 · d^?1) were similar to the maintenance requirement for domestic dogs (525 kJ · kg^?0.75 · d^?1), suggesting a similar energy requirement for maintenance of the wolves. The metabolizable energy derived from protein in the high prey diets was 37%, whereas that of the chow diets was only 27%. These levels exceed estimates of protein requirements for maintenance (4.3–21.8%) and growth (11.5–20.3%) in domestic dogs and may thus exceed those of the wolves. Dietary management of maned wolves should minimize excess protein intake by limiting prey consumption and acclimating animals to extruded diets for domestic dogs. Further research is required to formulate diets for long-term management of cystinuria in maned wolves. © 1994 Wiley-Liss, Inc.     

Feeding habits of the maned wolf,Chrysocyon brachyurus (Carnivora: Canidae), in southeast Brazil

The feeding habits of the maned wolf were studied in southeast Brazil to understand its response towards changes in the environment and in relation to its prey. By occurrence, miscellaneous fruits, small mammals and wolf's fruit were the most consumed items. Armadillos, small mammals and wolf's fruit (Solanum lycocarpum) provided most of the ingested biomass. While wolf's fruit and small mammals were mainly consumed in the dry season, other miscellaneous fruits were taken mostly in the wet season. There was selectivity in the predation on some small mammal species. The maned wolf's diet followed patterns similar to those found in more pristine areas.

O hábito alimentar do lobo guará foi estudado no sudeste do Brasil para descrever suas respostas as alterações do ambiente e em relação as suas presas. Por frequência de ocorrência, as frutas silvestres, os pequenos mamíferos e a fruta-do-lobo foram os itens mais consumidos. Tatus, pequenos mamíferos e a frutas-do-lobo (Solanum lycocarpum) constituíram os itens mais importantes em relação à biomassa ingerida. A fruta-do-lobo e os pequenos mamíferos foram consumidos principalmente na estação seca, enquanto os demais frutos silvestres na época das chuvas. Houve seleção de algumas espécies de pequenos mamíferos. A dieta do lobo guará apresentou padrões semelhantes aqueles encontrados em áreas mais preservadas.     

Helminth infracommunity in a maned wolf,Chrysocyon brachyurus,from the humid Chaco,Argentina

Parasitism is an important factor in conservation worldwide, especially for endangered species, as it can affect host populations by reducing growth rates, fecundity and affecting nutritional status. The maned wolf Chrysocyon brachyurus, a near-threatened species, is distributed across different habitats from the south Amazonian forest in Brazil to northern Argentina, Paraguay and eastern Bolivia. Most studies on gastrointestinal parasites in wild maned wolves are based on coprological findings, and there is little information about adult parasites and parasite species richness. Therefore, this study aims to expand the knowledge about the helminth infracommunity of a maned wolf, describe adult parasites, and explore parasitic coinfections. We performed a necropsy of an adult individual found road killed in Chaco province, Argentina. We collected adult worms from subcutaneous tissues, from the right ventricle, kidneys, and intestine. The parasites were morphologically identified as Dirofilaria immitis, Dioctophyma renale and Spirometra sp. respectively. To our knowledge, this is the first record for adults of Dirofilaria immitis in maned wolves. Filarioids were located within the heart as well as in subcutaneous tissues with mature females containing uterine microfilariae; this study also adds C. brachyurus as a new host for Spirometra sp. in Argentina. Considering that the three helminths found herein are zoonotic and that the maned wolf is an endangered canid, it is essential provide baseline information to determine the risk factors involved in the transmission and extend the studies to sympatric wild and domestic canids in the area.     

Diet of the maned wolf (Chrysocyon brachyurus) and its role in seed dispersal on a cattle ranch in Brazil

The maned wolf Chrysocyon brachyurus is the largest canid inhabiting South America. Its geographic distribution includes the open fields of Brazil's central area, which is currently undergoing agricultural expansion. The diet of the maned wolf and its seasonal variation was determined on a dairy cattle ranch (São Luís farm, 566 ha) in the State of Minas Gerais, Brazil. From January to December faeces of the maned wolf were collected monthly ( n = 150 scats; 397 food item occurrences). Twenty-nine taxa were identified from scats, 18 of animal origin (46% or 183 occurrences) and 11 of plants (54% or 214 occurrences). The fruits of Solanum lycocarpum were the dominant food item in our study (29%). Mammals contributed 13%, arthropods 12%, birds 11% and reptiles 2% of the food items. Arthropods and fruits were prevalent in the rainy season and mammals in the dry season. As expected for a heavily farmed region, frugivory results were at the lower end of the diversity scale (9–33 species) and included four old garden species. No previous study of the diet of maned wolf has registered as many species of Solanaceae as this one. Although dietary richness was lower, the main food items (wolf fruit, armadillos, rodents, birds) were the same as study sites in 'cerrado' and upland meadows. In this region, the open habitats occupied by the maned wolf were previously covered by Atlantic forest, suggesting that landscape modification such as cattle ranching has opened new frontiers for distribution expansion of the maned wolf. The impact of loss of dietary richness and the increase in Solanaceae on the survival of the maned wolf need to be evaluated.     

Transit time and digestibility of two experimental diets in the maned wolf (Chrysocyon brachyurus) and domestic dog (Canis lupus)

Recent research involving nutritional management of clinical disease secondary to cystinuria in captive maned wolves (Chrysocyon brachyurus) in the United States resulted in the production of a commercially manufactured maintenance diet and subsequently, a further modified experimental diet. These two diets differed only in their protein sources and sodium concentrations. The purpose of this study was to further investigate these diets by determining their digestibility and transit time in maned wolves, and also to compare these results to the digestibility and transit time of the diets in domestic dogs. The experiment consisted of two diets (maned wolf commercial maintenance diet and experimental diet) and two species (maned wolf and domestic dog [beagle]) arranged as a 2 × 2 factorial design. For the transit time study, titanium dioxide (TiO₂, 5 g/kg diet) was administered as a pulse dose and total collection of feces was carried out for 50 hr. There was no effect of diet or species on initial Ti recovery time (IRT), the amount of Ti recovered at IRT, or the time to reach 50% of Ti excretion in the feces. Maned wolves had lower total percent recovery (P<0.01) of Ti than dogs and shorter time to last recovery of Ti in the feces (P<0.05), however, marker excretion in the maned wolves may not have been complete. For the digestibility study, in which chromic oxide was used as a marker, fecal samples were collected on 2 consecutive days immediately after 12 days of being fed the diets and then analyzed for dry matter, energy, protein, and minerals. Crude protein, calcium, phosphorus, and zinc apparent digestibilities were unaffected by species or diet (P>0.05). Apparent digestibility of dry matter (P<0.05) and energy (P<0.01) were higher in the experimental diet and in the dogs. Both dogs and maned wolves fed the maintenance diet exhibited a greater apparent retention of copper, iron, and magnesium (P<0.01). Dogs had a higher apparent retention of magnesium and sodium than the maned wolves. Zoo Biol 0:1–13, 2006. © 2006 Wiley‐Liss, Inc.     

Use of behavioral measures to assess reproductive status in maned wolves (Chrysocyon brachyurus)

Captive breeding efforts of the threatened maned wolf (Chrysocyon brachyurus) have been plagued by a lack of reliable reproduction and a high rate of neonatal mortality. A particular problem for animal managers has been the ability to detect pregnancy in a noninvasive manner. Pseudopregnancies are common, and many staff hours are expended preparing for a birth that may not occur. The objectives of our study were to document changes in behavior during the breeding season in captive maned wolves in order to determine if behaviors other than sexual (i.e., copulation) could be used to distinguish nonbreeding from breeding pairs and, further, whether successful breeding pairs (young born) could be distinguished behaviorally from breeding pairs that did not produce young (pseudopregnant). Between 1988 and 1994, behavioral data were collected during the annual reproductive season from 52 maned wolf pairings (27 males, 26 females) housed at 17 North American institutions. Breeding animals showed significant increases in rates of affiliative behaviors (approach, friendly) and the amount of time spent in close association (social) during the estrous period, compared to pre- and post-estrous periods. In contrast, the behavior of nonbreeding pairs varied little throughout the breeding season. Discriminant function analysis (DFA) showed that several behavioral measurements during certain reproductive periods were useful in discriminating between nonbreeding and breeding animals (rates of friendly and social behavior for males, marking and social behavior for females). Unsuccessful (pseudopregnant) and successful breeding pairs could be further distinguished on the basis of agonistic and approach behavior frequencies (successful females showed higher rates of agonistic and lower rates of approach behavior during certain periods; successful males had higher rates of both behaviors). Discriminant equations for key behavioral measurements and examples of their practical application are presented. DFA provides animal managers with an effective, noninvasive technique for assessing the reproductive status of maned wolf pairs. © 1996 Wiley-Liss, Inc.     

Behavioral changes associated with introductions of male maned wolves (Chrysocyon brachyurus) to females with pups

Three male maned wolves were successfully introduced to their mates and 7‐ to 12‐week‐old pups at the Houston Zoological Gardens and Fossil Rim Wildlife Center in 1994 and 1995. The introductions took 3–7 weeks and had three stages: 1) allowing the male to see the female and pups through a chain‐link panel or a full chain‐link fence, 2) introducing the male and female without the pups present, and 3) introducing the male to the pups. For 7 days before Stage 2 and 7 days during Stage 3, the adults were observed for 150 hr. The amount of time they spent active and the amount of time they spent near the pups were recorded, as well as all behaviors they directed to the pups. After the males were introduced, the females' activity decreased, and the males' activity increased. The females spent less time near the pups after the introduction. The introduction did not affect the females' rates of affiliative or aggressive behaviors to the pups, and there was no difference between males' and females' rates of affiliative or aggressive behaviors. This introduction procedure offers an alternative to leaving the male with the pups from the time they are born. The adults' behavioral changes after the introduction show the benefits that occur when captive male maned wolves are allowed to help rear their pups. Zoo Biol 18:189–197, 1999. © 1999 Wiley‐Liss, Inc.     

Secret lives of maned wolves (Chrysocyon brachyurus Illiger 1815): as revealed by GPS tracking collars

The maned wolf Chrysocyon brachyurus is a nocturnal and shy species, which has proven difficult to study in the field; consequently, data about its behavioural biology are almost absent from the scientific literature. However, recent advances in global positioning system (GPS) technology mean that it is now possible to study the socio-spatial dynamics of this species in the field. In the present study three related maned wolves (an adult pair and their juvenile female offspring) were monitored by GPS tracking collars for a 6-month period, which included the birth of a litter of pups to the adult pair. The three GPS collars were programmed to record the position of the wearer every 2 h (simultaneously for all three individuals). Analyses of the data from the three individuals showed that the female had the largest home range, then the male and finally the juvenile. Furthermore, there was considerable overlap in home-range use by all three individuals. The home ranges of the adult pair also varied in relation to the birth of pups (decreasing dramatically and then showing a slow increase). During the night, which is the normal period of activity for this species, the three individuals avoided contact with each other. However, during the day, the adult pair usually slept together and their juvenile slept within a few hundred metres of their location. The distance travelled at night was greatest for the female, then the male and finally the juvenile. These data suggest that (1) the social bond between the male and female is strong, (2) maned wolves are highly tolerant of their juvenile offspring, even in the case of new pups being born, and (3) that maned wolves do not hunt together and, in fact, avoid each other when hunting.     

Taurine deficiency in maned wolves (Chrysocyon brachyurus) maintained on two diets manufactured for prevention of cystine urolithiasis

This study assesses the long‐term effects of an experimental diet vs. a commercially available manufactured diet, intended to reduce clinical disease related to cystinuria, on the taurine status of captive maned wolves. For 13 weeks, two pairs of maned wolves were maintained on the commercially available maintenance diet, whereas two individually housed wolves were maintained on the experimental diet. All six wolves, at the beginning and at the end of the diet trial, had severely decreased plasma concentrations of taurine (as compared to the normal canine reference range of 60–120 nmol/ml) (National Research Council [2003] National Academies Press) with average taurine concentrations of 16 nmol/ml at the beginning of the study and 3 nmol/ml at the end of the study. There was no statistically significant difference in the taurine concentrations between animals on the maintenance vs. experimental diets. Both diets were supplemented subsequently with taurine at a concentration of 0.3%. All study animals were eventually switched to the taurine‐supplemented version of the commercially manufactured maintenance diet and subsequent samplings were carried out to monitor plasma taurine concentrations. A final sampling, carried out approximately 5 months after the initiation of taurine supplementation, showed an average taurine concentration within the target canine reference range (90.25 nmol/ml). There are numerous physiologic (e.g., possible unique metabolism and requirements for taurine in this species as compared to other canids) and dietary factors (e.g., effects of the types and concentrations of fiber and protein on nutrient availability, taurine metabolism, and enterohepatic circulation of taurine‐conjugated bile salts; impaired taurine synthesis secondary to low cysteine availability) that could be potential contributors to the development of taurine deficiency in the maned wolves in this study. Taurine supplementation should be considered in maned wolves maintained on diets intended for reduction of cystinuria‐related complications. Zoo Biol 0:1–14, 2005. © 2005 Wiley‐Liss, Inc.     

First observations of nest attendance behavior by wild maned wolves,Chrysocyon brachyurus

In many species of canids the male's role in reproduction extends to providing extensive or some parental care to his offspring. Maned wolves are a monogamous canid species whose males have been observed providing parental care to their offspring in captivity, but no field observations exist. We observed a wild pair of maned wolves at their nest site in a period soon after the female had given birth for a total period of 65 days. We made five observation sessions with an average of 3–4 days each separated by approximately 2 weeks. Direct visual observations of maned wolves were made each 30 min during the hours of darkness (17:00–07:00) using night-vision binoculars and confirmed by VHF radiotelemetry. During observations we recorded the location of the male and the female in relation to the nest (i.e., in the nest, nearby or long way from the nest). The results showed that the female spent more than 60.44% of her time in or near the nest. The male spent 28.90% of his time in or near the nest. There was a positive significant correlation between the female and the male in terms of the amount of time spent in or near the nest (P<0.01). The maned wolves showed a strong temporal variation in time spent in or near the nest. In conclusion, our data show that wild male maned wolves—probably—provide parental care through provision of food to their female and presumed offspring rather than “babysitting.” Zoo Biol 28:69–74, 2009. © 2008 Wiley-Liss, Inc.     

An evaluation of manual and automated methods for detecting sounds of maned wolves (Chrysocyon brachyurus Illiger 1815)

Although bioacoustics is increasingly used to study species and environments for their monitoring and conservation, detecting calls produced by species of interest is prohibitively time consuming when done manually. Here we compared four methods for detecting and identifying roar-barks of maned wolves (Chrysocyon brachyurus) within long sound recordings: (1) a manual method, (2) an automated detector method using Raven Pro 1.4, (3) an automated detector method using XBAT and (4) a mixed method using XBAT's detector followed by manual verification. Recordings were done using a song meter installed at the Serra da Canastra National Park (Minas Gerais, Brazil). For each method we evaluated the following variables in a 24-h recording: (1) total time required analysing files, (2) number of false positives identified and (3) number of true positives identified compared to total number of target sounds. Automated methods required less time to analyse the recordings (77–93 min) when compared to manual method (189 min), but consistently presented more false positives and were less efficient in identifying true positives (manual = 91.89%, Raven = 32.43% and XBAT = 84.86%). Adding a manual verification after XBAT detection dramatically increased efficiency in identifying target sounds (XBAT+manual = 100% true positives). Manual verification of XBAT detections seems to be the best way out of the proposed methods to collect target sound data for studies where large amounts of audio data need to be analysed in a reasonable time (111 min, 58.73% of the time required to find calls manually).     

Food Provisioning by Adult Maned Wolves (Chrysocyon brachyurus)

Book reviewed in this article: Bücher/Books: Tyler, M., ed. (1983): The gastric brooding frog (Der magenbrütende Frosch). Paepke, H.-J. (1983): Die Stichlinge (Sticklebacks). Die Neue Brehm-Bücherei, A. Eisenberg, J. F., and D. G. Kleiman, eds. (1983): Advances in the study of mammalian behavior (Fortschritte der Säugetierforschung). Ankel-Simons, F. (1983): A survey of living primates and their anatomy (Eine Übersicht über die heute lebenden Primaten und ihre Anatomie). Wood-Gush, D. G. M., M. Dawkins and R. Ewbank, eds. (1981): Self-awareness in domesticated animals (Zum Ich-Bewußtsein domestizierter Tiere).     

A note on the effect of the full moon on the activity of wild maned wolves, Chrysocyon brachyurus

It is well established in the scientific literature that animal prey species reduce their activity at times of high predation risk. In the case of nocturnal animals this occurs when there is a full moon; however, the response of predators to the changes in their prey behaviour is relatively unknown. Two responses are possible: (1) increase in search effort to maintain food intake; or (2) decrease in distance travelled due to either: (a) an effort to conserve energy or (b) increased kill efficiency. Using GPS tracking collars we monitored the distances travelled (which is representative of search effort) by three maned wolves during the night of the full and new moon for five lunar cycles (during the dry season). A Wilcoxon matched-paired test showed that the maned wolves travelled significantly less during the full compared to the new moon (p < 0.05). On average, during the 10 h of darkness during a full moon maned wolves travelled 1.88 km less than on a new moon. These data suggest that maned wolves respond to temporally reduced prey availability by reducing their distance travelled.     

Evaluation of homologous, heterologous, and affinity conjugates for the serodiagnosis of Toxoplasma gondii and Neospora caninum in maned wolves (Chrysocyon brachyurus)

Use of serological tests in the diagnosis of infectious diseases in wild animals has several limitations, primarily the difficulty of obtaining species-specific reagents. Wild canids, such as maned wolves (Chrysocyon brachyurus), are highly predisposed to infection by Toxoplasma gondii and, to a lesser extent, to Neospora caninum. The aim of the present study was to evaluate homologous, heterologous, and affinity conjugates in enzyme-linked immunosorbent assays (ELISAs) and indirect fluorescent antibody tests (IFATs) for detecting immunoglobulin (Ig) G antibodies against T. gondii and N. caninum in maned wolves. Serum samples were obtained from 59 captive animals in Brazil and tested by ELISA for T. gondii serology and IFAT for N. caninum serology using 3 different enzymatic and fluorescent conjugates: homologous (guinea pig anti-maned wolf IgG-peroxidase and -fluorescein isothiocyanate [FITC]), heterologous (rabbit anti-dog IgG-peroxidase and -FITC), and affinity (protein A-peroxidase and -FITC). Seropositivity to T. gondii was comparable among the homologous (69.5%), heterologous (74.6%), and affinity (71.2%) enzymatic conjugates. A significant positive correlation was found between the antibody levels determined by the 3 enzymatic conjugates. The highest mean antibody levels (ELISA index = 4.5) were observed with the protein A-peroxidase conjugate. The same seropositivity to N. caninum (8.5%) was found with the homologous and heterologous fluorescent conjugates, but protein A-FITC was not able to detect or confirm any positive samples with homologous or heterologous conjugates. Our results demonstrate that homologous, heterologous, and affinity conjugates might be used in ELISA for serological assays of T. gondii in wild canids, whereas for N. caninum infection, only the homologous or heterologous fluorescent conjugates have been shown to be useful.