The Captive Energy Budget of Lycaon pictus (The African Painted Wolf)

The Captive Energy Budget Lycaon pictus (The African Painted Wolf) 

Adrian Raygoza (Senior undergraduate BioS major)

University of Illinois at Chicago

1200 W. Harrison St.

Chicago, IL 6060

Introduction

The African Wild Dogs, Lycaon pictus, are very unique species of wild canids that are native to the African continent. They are one of the largest canids and most successful canid hunters in the world (Tighe 2013). This species was very especially important to examine due to the lack of current studies regarding the social behavior in the lives of captive African Wild Dogs; including studies regarding possible reintroductions to the wild. There currently exists a plethora of vast studies of other large canid species, such as grey wolves. However, long-term studies of behavior, especially regarding captivity, are still considerably lacking for the Lycaon pictus (Tighe 2013).

Background:

The African Wild Dogs are known by several different common names such as the African Painted Dog and the Painted Wolf (Mulheisen et al. 2014). The latter common name is favorably the most accurate when describing this species. The reason for this is because biologists are currently making a push to disassociate this unique animal from being commonly mistaken as a feral dog, Canis familiarus (Tighe 2013). Though this canid shares a similarity, such as a pack living and social hierarchies, to several other species of canids, it is truly in its own classification, both figuratively and literally. Lycaon pictus is the only member of the Lycaon genus (Tighe 2013; Woodroffe et al. 2004). They differ significantly from common dogs and other canids. The largest difference is that this species cannot be domesticated and cannot reproduce with other members of the canid families (Woodroffe et al. 2004). For several years in the early 1900s, Lycaon pictus was classified as a species of a hyena (Woodroffe et al. 2004). This was mainly due to the similarities in color and head shape (Woodroffe et al. 2004). However, hyenas are much larger than typical Lycaon pictus (Woodroffe et al. 2004). It was not until that the distinction was made that these canids were made placed into a separate classification altogether (Tighe 2013; Woodroffe et al. 2004). The average size for this species is 39-80lbs and about 29-43 inches in length (Mulheisen et al. 2014).

In their native range, which has significantly and vastly been reduced in the last several decades, they face large pressures that other large canids typically do not face (Tighe 2013; Woodroffe et al. 2004). There are many large predators that immensely roam what encompasses the Lycaon pictus (Tighe 2013; Woodroffe et al. 2004; Rasmussen & MacDonald 2002). In comparison, grey wolves (Canis lupus), who range in North America, come into contact with other predators such as bears and occasional mountain lions, however, they generally do not directly compete against those species and have different niches (Creel and Creel 2002). Lycaon pictus does directly compete with larger predators in their natural range, including several of the large cats that reside in Africa. The largest direct competitors and predators of the Lycaon pictus are lions (Panthera leo) and hyenas (Hyaena family) (Tighe 2013).

Not only do these species present an issue of the interspecific competition to food, but they also present intraguild predation to the Lycaon pictus in the same ranges that they come in contact with (Creel and Creel 2002). It is well documented that Lycaon pictus have greater success in ranges that do not overlap with these large predators (Tighe 2013; Woodroffe et al. 2004; Rasmussen & MacDonald 2002). Lions cause a high mortality rate for the Lycaon pictus (Tighe 2013; Woodroffe et al. 2004; Rasmussen & MacDonald 2002). These predators are known to attack the Lycaon pictus, eliminating them as competition, and sometimes predating on them (Tighe 2013; Woodroffe et al. 2004; Rasmussen & MacDonald 2002). Hyenas also predate on Lycaon pictus, however, they are more notorious for kleptoparasitism to the Lycaon pictus (Tighe 2013; Woodroffe et al. 2004). Hyenas will often try to overrun the Lycaon pictus and steal their kills (Tighe 2013; Woodroffe et al. 2004). The pressures that these species have placed on the Lycaon pictus have led to the adaptation of their unique social behaviors and interactions (Woodroffe et al. 2004).

Understanding innate wild behaviors:

In order to combat such pressures, Lycaon pictus relies on pack effort (Tighe 2013; Woodroffe et al. 2004; Rasmussen and Macdonald 2012). As a group, they are better able to defend their kills and drive off potential kleptoparasites (Tighe 2013; Rasmussen and Macdonald 2012). They are also better able to fend off predators with larger packs in bolstered numbers (Tighe 2013; Woodroffe et al. 2004). Due to the habitat pressures, Lycaon pictus varies in their communication from other large canids (Woodroffe 2001). Their social structure also differs as well. Group living has evolved as a response to a variety of selective pressures in their native range that differ quite differently from other canids (Tighe 2013; Woodroffe et al. 2004). In the hierarchy of the Lycaon pictus, there is an alpha male and female, however, they do not assert the same type of dominance as typically known of grey wolves (Tighe 2013; Woodroffe et al. 2004). For instance, alphas are not the only breeding pair in a pack, as in grey wolves (Tighe 2013; Woodroffe et al. 2004, Woodroffe 2001). The alpha male often leads and assists the pack on hunts (Woodroffe 2001). On successful hunts, food is shared, and every individual gets usually receives a chance to feed (Rasmussen and Macdonald 2012). Whether by begging behavior or by participating in the actual kill, all individuals are afforded a chance to eat, including the young and the sick (Tighe 2013; Woodroffe et al. 2004). One behavior quite unique that has been observed in the wild is the regurgitation of food for ill members of the pack (Tighe 2013). These types of behaviors are not typical of other canids (Tighe 2013; Woodroffe 2001).

The lack of competitiveness amongst the pack individuals and the lack of persistent dominating behavior, especially when it comes to feeding, is quite unique among Lycaon pictus (Tighe 2013). In comparison to other large group hunting predators, Lycaon pictus is more successful in obtaining kills, this is due to their group cooperation and social cohesion (Tighe 2013; Woodroffe 2001). The social bonds of this species are absolutely crucial for survival and are the basis for communication and behavior (Creel and Creel 2002). Most of the communication between the species are displays within a short range of each other (Tighe 2013). This is due to the prevention of eavesdropping species, who may garner attention by intercepting signals of communication, which may cost survivorship to Lycaon pictus (Rasmussen and Macdonald 2012; Woodroffe 2001). This is very much, unlike other candids, for comparison, who use vocal communication over long distances (Woodroffe 2001). Strong odors amongst pack members are the Lycaon pictus most common form of tracking each other over long distances (Tighe 2013). Lycaon pictus practice stealth and quiet movements over long distances for their own safety (Mulheisen et al 2014; Tighe 2013). This complex phenomenon was most likely developed due to the pressures in their native range.

As in many, if not all, animal species that are held and bred in captivity, a question of how these survivorship behaviors are affected by the minimization of enclosures at zoos or other captive holdings is a fair question (Wey et al. 2008). Zoos, for instance, tend to present different stimuli and pressures on most species; this also includes the added pressures of observing humans (Wells 2005). The lack of a native range may change the behavior that has been adapted over decades or centuries. A lack of predators, a lack of hunting, a lack of large free-roaming areas, and the presence of humans in close quarters, all present a different set of pressures and stimulus that directly affect several animals held in captivity (Wells 2005). For the Lycaon pictus, who naturally have, by natural selection, adapted to several pressures of predators and competitors (Creel and Creel 2002), the question of social status and behavior comes into play. I investigated the behaviors of captive Lycaon pictus to search for answers. I hypothesized that the energy budget of the male and female Lycaon pictus would differ significantly due to restrictions of the captivity setting.

Methods

This study took place observing a pair of breeding Lycaon pictus housed at the Lincoln Park Zoo, located in Chicago IL. In order to gain a better understanding of captive behaviors, while understanding that most of Lycaon pictus studies have taken place exclusively in the wild, a formalized catalog of all observed behaviors, known as an ethogram, was constructed based on the behaviors observed in captivity at the Lincoln Park Zoo. A technique referred to as ad-lib sampling, in which all observed behaviors in a specific time-period are observed, was used to record each behavior seen from the pair of Lycaon pictus in the enclosure. All specific behaviors witnessed and interpreted during the two-hour observational span were recorded; including high energy behaviors, such as running around the enclosure, to low energy periods of rest. The continuous observations were analyzed, categorized, and constructed into the ethogram.

The developed ethogram, with a defined catalog of behaviors for comparison, was incorporated to perform quantified behavioral observations known as focal-animal sampling. The purpose of this protocol was to obtain observational data to construct a time budget for both individual sexes. This was managed by dividing the total number of individually listed behavioral observations, listed on the ethogram, by the total observation time spent on the behavior. Animal focal-sampling was completed for a grand total of two hours of observations. The hours were broken down into 15-minute increments of direct behavioral observation. A timer was used to measure one minute, on the minute, for each individual 15-minute increment. Using the ethogram, an inventory was used to mark the observed behavior that the observed Lycaon pictus was performing during the observed minute. One total hour was spent over two days observing the male and one hour total was spent observing the female for a total of two hours of observations for the pair.

Using the data from the focal animal sampling, the time budget was constructed for the breeding Lycaon pictus pair at the Lincoln Park Zoo. The time budget was calculated by taking the number of marked behaviors during the observation period and dividing them by the total minutes of observation. Because observations took place on members of different sexes, two separate time budgets were created; one for each sex. In creating the individual time budgets, any behavior listed in the ethogram that was not observed during the focal sampling observations was removed from the time budget. A category for “other” behavior was added to the ethogram and was included in the final time budget list. This was done in case behaviors that were not originally observed during ad lib sampling were observed later during the focal sampling process.

Data

Table 1 Ethogram of Lycaon pictus

Table 2 Time budget for male Lycaon pictus

Figure 1 Distribution of Male Time Budget

Table 3 Time Budget for female Lycaon pictus

Figure 2 Distribution of female time budget

Interpretation/conclusions

The breeding pair of Lycaon pictus, at the Lincoln Park Zoo, displayed a variety of observed behaviors. Though there were some slight differences between the individual time budgets of the male and female Lycaon pictus, overall, the pair displayed similar behaviors for comparable amounts of time. The data, based on the separately generated time budgets for each sex, displayed three specific behaviors that dominated both time budgets. The dominating behaviors were referred to as the “main three” behaviors from the observational pair. These behaviors collectively exceeded 50% of the total for each time budget.

Respectively, time spent observing was the most common behavior for both sexes, at 27% for the male and 26% for the female. Followed by maintaining a neutral position, at 20% for the male and 23% for the female. The third-largest behavior observed was locomotion, at 20% for the male and 20% for the female. The total of these three specific behaviors nearly encompasses ? of the time budget (slightly above 70% for the total time) for both sexes that were observed. The following behaviors seemed to differ slightly between the pair. The male's fourth most observed behavior was play at 12%, followed by resting together at 8%. Pawing at the door, other, and object involvement all tied at 3%. Lastly, eating/drinking was observed 1% of the time. For the female, the fourth largest behavior observed and counted was resting together at 13% and play followed at 7%. Pawing the door was observed at 5%. Object involvement was observed at 3%. Lastly, eating/drinking was observed at 2%. Unfortunately, not all categories that were defined in the ethogram were observed during the focal sampling observations. Due to the lack of observance for certain behaviors during the focal sampling observations, those categories scored 0% and were, thus, removed from the time budgets.

The male was generally more physically active, displaying a larger set of active behaviors, than the female during the focal-sampling observations. The male displayed 10 out of the total 12 categorized behaviors of the ethogram. One of these behaviors were not seen by the female. This behavior was classified as “other” because it was not originally observed in the ad-lib sampling. This behavior, by the male, was the act of digging its nose into the ground. It is unknown if this behavior was a type of marking of territory or simply scratching its nose. However, because it was observed more than once during the focal sampling but not during the ab-lib sampling, it was categorized as “other”.

The male’s locomotion trend also differed from the locomotion trend observed by the female. The male had a tendency to circle the enclosure during most of the locomotion observed. The male also seemed to prompt the female into activity by being the first to get up from a neutral position and begin any locomotion. Oftentimes, the male was observed nudging the female, in which the female was non-receptive some of the time and remained in a neutral position thereafter. During the times that the female did interact with the male, it was during play-like behavior between the pair. Though both the male and female displayed some object involvement, they varied in the selection for objects of choice. The male dragged pieces of logs that were around in the enclosure. The female’s object involvement was spent dragging bedding materials, most likely hay, to the area that she was resting at. The male was observed stalking, a category that was not seen by the female. The stimulus that led the male to assume a stalking position was out of view from observation in the enclosure. The male was observed pawing the enclosure door several times, as was the female.

The female, by contrast, was less active than the male. Only displaying the 8/12 categories based on the ethogram. After the “main three” behaviors, the largest observed category was resting together, in which both the male and female were resting in very close proximity together. Though this behavior was classified as together, it took place more during the focal sampling of the female than the male, for which it was not counted as often for the male. Though the female was observed playing with the male, it was mainly due to prompts from the male, in which she joined the play. The only other locomotion observed, that was not prompted by playing with the male, was locomotion to add bedding material to her resting area or to paw at the door of the enclosure, and briefly drinking from the water source. The female did not circle the enclosure during locomotion, as the male did. Both members of the pair were observed drinking from the water source inside the enclosure. Though there was no actual feeding observed, the female was observed chewing on something inside the enclosure at her resting area.

Neither of the pair was observed Lycaon pictus to be out-of-view during observations of the enclosure during the focal sampling portion of the study. Though the male displayed behavior that was designated “other”, the female did not display any behavior that would be classified as “other”. One category that was not seen by either sex during the focal sampling was any type of submissive behavior. Both members of the pair seemed to be more active during the warmer weather of the ad-lib observations to create the ethogram, rather than during the cooler weather during the focal sampling. It is not known, however, if the warmer temperature played any role for certain. It was also not known if the female was in gestation and whether or not that played a role in the observations.

In comparison to much of the literature that exists on the Lycaon pictus behavior and sociality, many of the studies have focused on Lycaon pictus that live in their native range and habitats. One researcher, Emily Tighe (2013), did some research on the captive Lycaon pictus residing in a zoo. In her observations, she found that many of the behaviors observed in captivity resembled the same behaviors observed in the wild (Tighe 2013). There were several differences separating her study and my observations. Ms. Tighe (2013) was fortunate to have access to an entire pack that was living in captivity and was granted access to non-public areas of viewings within the enclosures. Simultaneously, she also conducted live observations of the pack in the wild for her study; for comparison (Tighe 2013). Lastly, her observational zoo pack was all one sex, female (Tighe 2013).

In contrast to the findings of Ms. Tighe (2013), it is difficult for me to conclude that the behaviors that I have observed in captivity match or mirror the many of the same characteristics of the wild. As greatly discussed in the introduction section, several of the pressures that surround the Lycaon pictus in the wild also have driven the adaption of many of the behaviors that are normally seen by the species (Creel and Creel 2002). I do not completely contest the findings of Ms. Tighe (2013). Some of the behaviors I was able to observe, such as playing and resting together, reflect the deep sociality that is observed in the wild. For instance, laying in close proximity is a behavior of safety for the Lycaon pictus, who are normally in areas surrounded by predators and competitors (Creel and Creel 2002). Taking long moments to observe and survey their surroundings is also safety, and likely a precautionary, behavior as well (Creel and Creel 2002; Woodroffe 2001). Playing is a large form of the sociality of Lycaon pictus (Creel and Creel 2002; Tighe 2013; Woodroffe 2001).

In comparison to the literature reviews that were researched for this project, several behaviors seemed to be reduced in comparison to the wild Lycaon pictus and those observed at Lincoln Park Zoo. I believe that there are two major factors for this. One of these factors is the lack of hunting for these captive animals. Though no specific feeding time was observed for the breeding pair during my observations, it still cannot be compared to actual hunting that these animals are recognized for. Known for outrunning and tiring out their prey, Lycaon pictus are elusive hunters (Creel and Creel 2002; Tighe 2013; Woodroffe 2001).

Pack hunting requires strategic communication and group effort and also presents opportunities for observed sociality among the individuals that, unfortunately, cannot be seen in captivity. I believe that hunting not only presents opportunities for sociality but would also greatly change the energy budget of the Lycaon pictus, as the act of hunting is metabolically expensive for any predator. Specifically, for Lycaon pictus, much of their natural behavior revolves around roaming large distances for the purposes of hunting, and also evading other predators (Creel and Creel 2002; Tighe 2013; Woodroffe 2001), which contradict the large percentage of neutral and resting behaviors that were observed in my focal sampling. Though submission is not as typical of Lycaon pictus as it is for other candid species (Creel and Creel 2002; Tighe 2013; Woodroffe 2001), these opportunities of gathering to eat a capture would present more opportunities for submissive/dominance behaviors, that were not observed in my focal sampling.

 The extent of the pressures faced by the Lycaon pictus in the wild is also continuously existent and the very foundation for the development of the unique behaviors seen in this species (Creel and Creel 2002; Tighe 2013; Woodroffe 2001). In captivity, the Lycaon pictus that I observed did not have to deal with the burden of becoming a meal to a larger predator and/or defending and losing their hunting kills from larger animals. The lack of predator/competitors extremely alters the life of captive species in comparison to living in the wild. It can be surmised that evading and/or defending their kills, or themselves, from predators/competitors also bears a great metabolic cost, as well as a probable direct fitness cost. It can be deduced that these activities greatly alter a time budget, as well, for the Lycaon pictus, specifically, as it relates a great deal to locomotion and resting percentages.

For future studies, the observations of a pack, of mixed-sex preferably, rather than a breeding pair may produce results that are of a better comparison to the behaviors and sociality seen in the wild. Unfortunately, it is very difficult to recreate the many variable factors that life in the wild presents to this species, as well a creating a large enough roaming range. It is crucial to understand the effects of captivity for Lycaon pictus, especially in the hopes to repopulate the species in the wild after captive breeding programs. I propose a study where a Lycaon pictus pack that was raised in captivity be reintroduced into the wild and observed as they adjust to life in the wild. I hypothesize that the pack will be able to successfully establish a territory and reproduce a new generation.

For this study, a group of mature Lycaon pictus should be reintroduced to a portion of their native ranges in Africa, preferably an area less dense with competitors and predators. Each member of the pack should be fitted with a GPS transmitting collar or device that they can be tracked within real-time. The pack should be observed and followed for a minimum of one production cycle of new generation pups that grow to reproduction maturity age. Because the species is very active diurnally, the observations should begin early in the morning and last through the evening. Recording equipment should be established in areas of high traffic for the pack. In order to fulfill this task, a research team should be established to follow the pack as they go about their daily routines, including hunts and predator evasion and defense. Along with recording equipment, the study will require photography equipment, tracking equipment, and a range of vehicles for the research team to use to safely follow the pack activities. Observations of their interactions with other species will be crucial for the determination of their survival in the reintroduction. Nocturnal observations should also be considered, as this is the time when natural enemies of the Lycaon pictus roam.

An ethogram of observed behaviors should be established using ad lib sampling. The timing of this sampling will take several weeks to allow the pack to establish a routine in their reintroduced habitat. 40 hours of total ad-lib sampling, in intervals of 2 hours, will take place over the first month of reintroduction. A time budget will be created through focal sampling. Each individual in the pack will be individually focused on for a total of 2 hours each. The ethogram and time budgets can, at a later time, be compared to the behaviors and sociality observations that were created by observing captive and free-roaming packs. These observations will also be used to determine the social hierarchy of the pack as it builds over time. This data will be used to compare and see if the overall behaviors that were not necessarily seen in captivity will be reestablished in the wild, or if a new/different set of behaviors arise. This study will be crucial and can serve as a foundation for all future reintroductions of the species to help repopulate the dwindling numbers of Lycaon pictus in their native range. The long-term goal of this study will become a precursor to successful reintroduction, in comparison to the success of other canid reintroductions, such as the grey wolves of Yellow Stone National Park. 

References

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Creel, S., N. M. Creel. 2002. "The African wild dog: behavior, ecology, and conservation."New Jersey: Princeton University Press. 

Mulheisen, M., C. Allen, and C. Allen. Lycaon pictus (African Wild Dog). Animal Diversity Web. University of Michigan Museum of Zoology. https://animaldiversity.org/accounts/Lycaon_pictus/

Tighe, EJ. 2013. The effects of captivity on display-based communication and social interaction in the captive African wild dog (Lycaon pictus). Zoology. The University of Canterbury. Biological Sciences. https://ir.canterbury.ac.nz/handle/10092/8676 

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