OPEN ACCESS Case Report Novel Problem Solving, …The family Canidae includes different species of...

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Case Report

Novel Problem Solving, Exploration, and Socialization with Regards to Sensory Behaviors in the Family Canidae

Abigail Reid* Pawling High School, Pawling, NY, United States

*Corresponding Author: Abigail Reid, Pawling High School, Pawling, NY, United States.

Received: April 01, 2019; Published: April 30, 2019

Abstract

Keywords: Canidae; Wolves; Foxes; Exploration; Novel Problem Solving; Sensory Behaviors; Human Socialization; Puzzle Box

Canids, or animals in the Canidae family, use sensory behaviors to interact and communicate with others, gaining information about their environment. In captivity, canids are housed in several types of facilities, including zoos and conservation centers, and are exposed to various degrees of human interaction, which impact their learned behaviors. Wolves, specifically, are known to en-gage in cooperative actions, such as organized hunts and cooperative breeding, which require the use of their wolf-wolf cooperation skills. The socialization of canids other than wolves in conjunction with a problem-solving task has not been widely studied. This original research determined the relationship between sensory-behavior problem solving abilities and human socialization. The study assessed sensory abilities by presenting novel puzzle box apparatuses (one baited and one empty) to the canine subjects, while recording the exhibited exploratory behaviors. Additionally, this study examined the benefits of puzzle boxes as cognitive and sen-sory enrichment by analyzing corticosterone measures from fecal collections. The results suggest there is a correlation between an animal’s likeliness to approach a novel problem and their level of socialization, as well a difference between genre. The applications of this research extend from conservation, to captive management, to new insights on differences in canid behavior.

Citation: Abigail Reid. “Novel Problem Solving, Exploration, and Socialization with Regards to Sensory Behaviors in the Family Canidae”. EC Veterinary Science 4.3 (2019): 187-201.

Background

The family Canidae includes different species of wild and domestic dogs, wolves, jackals, dingoes and foxes. Wolves, specifically, are highly social animals that live in packs. To effectively cooperate with other pack members, they have various means of communication such as body language, scent markings, and vocalizations [1-5]. Wolves are also known to engage in cooperative actions as they are a cooperatively breeding social animal, meaning that other members in the pack help to raise the pups even if they aren’t their own [1-5]. Another example of their highly complex cooperative behaviors is that they utilize a variety of strategies during organized hunts and even territory defense [1-5].

Gray wolves or timber wolves (Canis lupus) are the most common wolf species. Mexican gray wolves (Canis lupus baileyi) are one of the subspecies that are endangered and being reintroduced into the wild. Red wolves (Canis rufus), also critically endangered, are native to North America’s east coast. Due to increased urbanization and overhunting, red wolves are now centralized in North Carolina, and are at risk of extinction [6-9]. Although they do live in packs, they are much smaller than other gray wolves, usually consisting of the mating pair and two to three other individuals. Being slimmer and smaller than the typical gray wolves, they mainly hunt smaller rodents and birds. Maned wolves (Chrysocyon brachyurus) are in a different genus than gray and red wolves. Native to South America, these animals are soli-

Introduction

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tary by nature, only pairing up for mating season [10,11]. They are also omnivores, eating a combination of fruits, vegetables, and small animals for protein. Between these taxonomic classifications, differences in their social pack interactions exist. For example, the gray wolf is a species which generally has a larger number of pack members, totaling around six to eight individuals, while the red wolf tends to live in groups of two to four wolves. Meanwhile, maned wolves exhibit more solitary social behaviors [10,11]. Foxes, genus Vulpes, are another member of the canine family. They are much smaller than wolves as they have shorter legs and bodies [12]. They are also mainly solitary, sometimes joining in groups of two or three. Foxes communicate through yelping, barking, and growls, contrasting with howling associated with wolves, genus Canis. Red foxes (Vulpes Vulpes) are native to the Northern Hemisphere, while arctic foxes (Vulpes lagopus) are mainly found in the Arctic. However, due to recent climate change and anthropological activity, their populations have expanded into more urban areas [13-15]. Both foxes and wolves are crepuscular and mainly carnivorous, yet foxes tend to hunt smaller prey, such as rats and birds, occasionally consuming primary producers [12-17].

Sensory systems allow animals to gather information about the external world. Animals use olfaction, the sense of smell, to commu-nicate with others and gain information about their environment. Canines rely mostly on their nose and sense of smell to function in the world around them [18]. Wolves and foxes use this sense to identify each other, mark their territories and to locate prey [18-24]. They rely heavily on a highly developed olfactory system to detect environmental information relevant for survival [18]. Additionally, these animals exhibit observable behaviors such as pawing and biting/pulling to explore their environment [17].

Problem solving and learning are two crucial skills for survival in an ever-changing environment. Social learning is especially impor-tant for these cooperative animals. It occurs when an organism’s learning is influenced by observation of or interactions with another animal [25,26].

Significance

Innovation is an essential concept with learning where animals solve a novel problem or find new solutions to existing problems [27,28]. It allows animals to exploit novel resources or to use current resources more efficiently. Thus, innovation improves the ability of animals to survive in complex or changing environments and to explore and create new niches [27].

Regarding conservation, red wolves and Mexican gray wolves are two of the most endangered canine species in North America [6-8,14]. In zoos, enrichment for captive animals is meant to provide stimulation and to evoke natural behaviors and enhance their well-being [29,30]. For animal husbandry, in terms of care for these captive animals, enrichment can range from habitat, sensory stimulation or food, to social and behavioral conditioning.

Currently, non-invasive measurements of captive animal hormone levels have become more popular in use [31-33]. Enrichment stud-ies used cortisol levels and behavioral observations to analyze the benefits of various enrichment types [29,30]. Several studies with facilities housing captive animals conducted case studies that examined how the effects of different types of enrichment varied between species [10,11,29,30]. The researchers measured cortisol levels through blood, urine, and feces of the animals to determine the physi-ological response of the enrichment on the animal and whether it was beneficial or not. Within several case studies of maned wolf enrich-ment [10,11], some results suggested that the maned wolves produced higher fecal cortisol levels when involved in enrichment activities in comparison to the absence of enrichment [11].

Previous studies used differently engineered apparatuses to examine the problem-solving abilities of hyenas, birds, meerkats, and oth-er mammalian carnivores [25-28,34-37]. The animals used in the studies were similar to wolves in that they were social animals and lived in communities engaging in social breeding. Recently, many researchers have conducted comparative studies to determine the differences in dog and wolf behavior for numerous abilities and cooperation tasks [5,6,19,21,22,38-45]. One specific study conducted by Polgar., et al. [22] developed the Natural Detection Task in order to quantify olfactory abilities in canines. They found that although scent dogs initially were more successful, wolves improved their success in the second test, while dogs did not. An earlier study by Udell [44] concluded that wolves spend significantly more time exploring a solvable problem-solving task and demonstrated a relationship between exploratory behaviors and completion of the task when compared to dogs. However, little research has evaluated the relationship between wolves and foxes, and exploratory behaviors associated with successful problem-solving tasks. Additionally, limited studies have aimed to examine the behaviors related to novel enrichment tasks and their correlation to human socialization experiences.


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Conservation research explored novelty behaviors and their effect on reintroduction success in different carnivore species [9,16,46]. These studies presented different endangered carnivore species with novel objects into their enclosure to explore what types of behav-iors were exhibited and to determine the “shy” personalities from the “bold” [9,16,46]. Then, when released, the researchers tracked the animals to see how well the different personalities fared in the wild. As human interaction in the wild increases, it is essential to analyze novel problem-solving success to ensure species success [47-49].

Hypothesis and Purpose

Thus, there has been minimal research on innovative problem-solving abilities in captive members of the family Canidae, regarding their exposure to human socialization, and no study has examined the combined olfactory and sensory behaviors involved with novel problem solving. This research aimed to investigate how canine problem-solving abilities are influenced by their observable sensory be-haviors and to analyze how exploratory characteristics influence success. The hypotheses were that the human socialized canids would perform better on the task, while the different genera of canids would exhibit various exploratory behaviors.

Methodology

Subjects and test sites

The subjects consisted of animals within the family Canidae as this study used wolves (n = 23) and foxes (n = 11) of different species. These animals were from seven different facilities, including conservation centers and zoos. The subjects were further categorized based on their human socialized and taxonomic groupings, specifically genus. “Socialized” animals were those who were raised by humans from birth and/or experience human interaction routinely. Also, each subject had a unique living enclosure and social situation, which evoked many variables. However, this study attempted to gather as large of a sample size as possible in order to analyze how other captive ani-mals performed the task. Appendix A includes a table of the subjects’ species, test sites, socialization, and other notable characteristics.

Apparatus

The puzzle box apparatus was made from PVC pipes and PVC boards for the experiment. This material was used so that it could be washed thoroughly between each trial and to maintain sturdiness throughout the duration of the experiment. The boxes were 7” x 11½” x 5 ½”, while the pipes used were 2” and 1½”. Other materials included were sturdy rope, steel cable, and eye hooks. The rope aided in the completion of the puzzle, while the eye hooks and cables anchored the puzzle boxes. The eye hooks and cables prevented the canines from picking the apparatus up and dragging it back to their den as well as keeping it inside the camera frame. Two GoPro Hero 3+s were used to document the trials so that human presence was not a major factor affecting the animals’ behaviors.

Figure 1: These visuals demonstrate how the apparatus worked: (A) closed puzzle box, (B) opened puzzle box, and (C) completely solved puzzle box.


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The mechanism of the apparatus consisted of two tasks. Firstly, the outer box, where the lid must be pulled off with force. The rope thus aided to get the box open. Once the top was off, the pipes on the inside could be accessed. The tubes fit inside one another, and 4” x 1½” openings were cut into each one. To obtain the treat, the subject had to rotate the pipes so that the openings lined up. Two weights, of about 25g each, were placed on the outer pipe so that it fell closed and stayed down until the animal turned it manually.

The box also had eight dissipation holes, which allowed the smell of the meat to escape the box, motivating the animals. There were four holes on the top piece: two on the outer tube and two on the inner tube. As a reward, pieces of venison were placed in the inner pipe. The gray and red wolves had about 250g of venison, while the maned wolves were only permitted 100g. For the arctic foxes, dead mice were used.

Experimental procedure

For each trial, two boxes were placed inside the enclosure. The boxes were placed about five feet away from each other, each fastened to some structure using steel cable and cable clamps through eye hooks. One box contained the reward, while the other was left empty as a control. This control condition allowed for the differentiation between the animals’ mere interest in the box and the scent of the reward. The camera was set up outside the enclosure or on an overhang fence, taking an aerial shot. This ensured that the wolves did not try to destroy/chew-up the camera.


Figure 2: This is an image of one wolf (16B) solving the puzzle box apparatus from the Wolf Conservation Center.

At two of the facilities, Fort Rickey Discovery Zoo and Wolf Conservation Center, a slightly different system was used. Since these wolves were more socialized, to avoid aggression over the possession of the boxes, one rewarded box was presented per animal. Thus, at Fort Rickey with two wolves, there were two baited boxes with one control, while at the Wolf Conservation Center there were three baited boxes with one control for the pack of three.


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The boxes were left in for 1 - 2 hrs or until animals lost interest. However, the data collected per trial was only the first 30 minutes or until the puzzle was solved and about 10 minutes after. The trials were also conducted before the animals ate so that they had the motiva-tion of hunger as well. The time between the trials differed from place to place, as there were only limited dates to conduct this experiment at certain facilities. The animals were then left alone so that human presence did not impact their behavior. Ideally, the subjects would have approached the boxes, determined which contained the reward, and then tried to solve that puzzle and obtain the treat.

For the wolves at Millbrook’s Trevor Zoo and Connecticut’s Beardsley Zoo, multiple trials ran every other day and fecal matter was collected before, during and after experimentation. These samples were then used to measure the cortisol levels, to help determine how the animals were physiologically affected by the introduction of the puzzle box.

Analysis

Behavioral

A majority of the data analysis was conducted through observations both live and on camera. The behaviors were organized into an ethogram, based on the accumulated video data. They were then divided into two main categories, active and inactive. Active behaviors were then further divided into exploratory and non-exploratory behaviors. The exploratory behaviors consisted of sniffing, pawing, bit-ing/pulling, staring, scent marking, and other actions toward the novel objects. Table 1 below shows the definitions of the categories used for coding the behaviors. The time it took for the animals to solve the box was defined as how long it took for the subject to obtain the treat and begin to consume it. Lastly, the times were divided into components of before and after based on the solved time or the halfway point, for the subjects who did not solve the puzzle.

Behavior DefinitionPawing Paws were engaged in an action to manipulate the box; a digging motionBiting Mouths/noses were engaged in an action to manipulate the box, including a pulling motion

Control Time spent purposefully at the control (empty) boxBaited Time spent purposefully at the baited box

Exploratory Actions spent purposefully investigating the boxesNon-exploratory Other normal actions spent not investigating the boxes; e.g. Pacing, playing, sitting, etc.

Staring Intentionally looking toward the boxesSniffing Action where the subject was clearly sniffing; i.e. Nose to the ground or on boxes

Scent marking Scent rubbing, urinating, defecatingInactive Laying down, sleeping, out of camera frame

Table 1: This chart defines the behaviors which were used in the ethogram and data analysis.

Physiological

To assess the physiological responses to the novel objects being presented to the animals, fecal samples were analyzed with an im-munoassay kit from Enzo Life Sciences to measure corticosterone hormone levels, or stress levels, for the three red wolves in the study. Samples were only collected for these individuals to analyze specifically the effect of the enrichment on red wolves, a critically endangered species. The fecal samples were collected before and during the experimental period, for only a select few facilities due to a time and loca-tion limitation. The line of best fit from the standard curve was used to determine the percent bound. These values and the dilution fac-tors were then incorporated in calculating the amount of corticosterone in the fecal samples using a ng (steroid) per g (feces) calculation, based on Palme., et al [31]. Appendix B contains the standard curve and line of best fit that were used.

Statistical analysis

Graphical analyses as well as Chi-Squared and Fisher’s Exact tests were conducted to analyze significance and examine relationships between genus, exploratory behaviors, and human socialization. The behavioral data analyzed was only taken from the “before” part of each initial trial, as it was investigating the various methods of approach used by the subject leading up to the time when it was solved.


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Results

Observational

Figure 3: This graph represents percent success of all the individuals tested. The data is separated into categories based on taxonomic groupings: the genus Canis, the genus Vulpes, and the overall family Canidae. The darker bars represent the human socialized animals, while

the lighter bars represent the unsocialized animals. The number of subjects for each category is placed above the bar. For each grouping, the significance value was calculated using Chi-Squared tests and Fisher’s Exact tests and placed above as well.

Figure 4: These box-and-whisker plots display the distribution of percent trial time for the exploratory behaviors comparing the socialized group and the unsocialized group. The number of subjects is placed above each plot. See Appendix C and Appendix

D for more plots broken into more categories.


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The analyses between the successes of the human socialized versus unsocialized illustrates that there was a relationship as the social-ized subjects were overall more successful. However, the Chi-Squared results, which calculated p = 0.050285, show that statistically this difference is slightly insignificant with a 95% confidence interval. Regarding genus, there was a significant relation between socialized wolves and unsocialized wolves (p = 0.0468), while there was no significant relation for the foxes between differing socialization levels (p = 0.5671). Furthermore, in all of the box and whisker plots, there was more time spent at the baited boxes than the control boxes in the first part of the trial. Fisher’s Exact tests were conducted between socialization and approach to the boxes (p = 0.0099) and between whether the puzzles were solved and the presence of scent-marking (p = 0.0057). Both of these tests demonstrated statistical significance.

Assay

The assay results were obtained using the formula from Palme., et al [31]. The graph below displays various fluctuations between the subjects; however, there is a general increase in the amount of hormone during the testing period. For the Millbrook Trevor Zoo red wolves, the stress level increases during the testing period, from 8/1/16 to 8/12/16, while it then begins to decrease afterwards. For the Connecticut Beardsley Zoo red wolf, the testing period, from 8/24/16 to 8/31/16, produced fluctuating hormone levels, but overall, there was an increase in stress levels during the experiment.

Figure 5: These box-and-whisker plots display the distribution of percent trial times for five behaviors (pawing, biting, time spent at the control, time spent at the baited box, and total exploratory behaviors) that were exhibited between the three different genera used in the

study. “Wolf” refers to Canis, “Fox” refers to Vulpes, and “Maned Wolf refers to Chrysocyon. The number of subjects is placed above each box-and-whisker plot.

Figure 6: This graph displayed the amount of stress, measured in ng corticosterone per g of feces, over time for the three red wolves used in the study. The subjects refer to 13A, 1A, and 1B respectively in order of the legend.


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Discussion

Based on the results, the significant p-value between whether the animals approached the boxes in correlation with their socialization (p = 0.0099), suggested that canids are more likely to approach novel items if they are human socialized. This trend can also be seen in figure 4, which demonstrated the humanly socialized canids as being overall more exploratory compared to the unsocialized canids. This demonstrates the significance of socialization on neophobia, or fear of new objects, of animals. The ability to approach the puzzle boxes would eventually lead to the animal’s solving the boxes. One possible reason for this overall relationship could be due to the human-socialized animals’ lowered neophobia and fear of humans from positive human experiences, like training. Some subjects, wolves mainly, that were socialized physically interacted with their keepers in playful ways. This type of human contact, while prohibited at some of the facilities, was an essential factor resulting in a lowered fear of novel objects. Secondly, the socialized animals may have been less afraid of the novel items if they had been previously exposed to similar types of enrichment. For example, some animals in the “human socialized” category were exposed to such a wide variety of enrichment, that their keepers stated it was a challenge to find new things to keep the animal’s interest. Another factor that played into their neophobia was purely subject-related. Each animal had different past experiences, influencing its current and future behaviors. For instance, 12A and 12B were afraid of the keepers due to a stressful past history. Thus, as soon as the boxes were placed, the animals were left alone except for the public viewing area. The animals were also only at that facility for five months, still adjusting, so they did not end up completing the task. However, according to their keeper, eventually, had they had been at the zoo longer, they would have actually approached the boxes. Also, since canids are pack animals, the social hierarchy within packs can create different levels of neophobia on an individual basis.

Another major aspect of the results was that human-socialized canines performed better than unsocialized canines, which was mea-sured in percent success (Figure 3). Overall, in the family Canidae, there was a slightly insignificant difference using the 95% confidence level. While this is approaching significance, statistically (as p = 0.050285), it suggests that there is some correlation between human socialization and their problem-solving abilities. The p-value right on the border could be a result of the low sample size. Thus, human interactions do affect how animals in the family Canidae may naturally approach novel items. Therefore, socialization of the animals was a factor that greatly impacted how they behaved during the trials. Human socialization on animals has various levels. At some zoos and conservation centers, animals are housed in conditions to maintain all of their natural, wild behaviors. Some public facilities attempt to keep the animals in their most wild state yet visits from the public and veterinary check-ups expose the animals to some socialization. Even further, animals at certain facilities can be trained and exposed to human interaction routinely. The last and most extreme level of human socialization would be domestication, where most “wild” behaviors have been tamed, and the animals rely heavily on human interactions daily.

Other future studies could investigate how domesticated and wild dogs would compare in similar tasks, as the level of domestication increases, in order to further examine the effects on socialization on canines. It’s possible that increased socialization of wolves may also increase their willingness to approach new items. Udell [44] had demonstrated that dogs failed to persist on problem solving tasks with regards to various human socialization levels [30]. It would be valuable to examine the fine line where socialization becomes more de-pendent on human interaction, using a range of animals from wild to completely domesticated. Furthermore, human interaction was also an influencing part of the results that varied from each facility. Most of the unsocialized subjects did best once they were left alone. The socialized animals performed the task regardless of human presence, so the keepers were present for most of the trials. Occasionally, the keepers would also encourage the animals. This may have influenced the results as the encouraged subjects may have been more moti-vated and less afraid to approach the boxes.

Differences between the genera were demonstrated in figures 3 and 5. For percent success, the significant difference regarding so-cialization for Canis (p = 0.0468) versus the insignificant difference for Vulpes (p = 0.5671) are associated with the observation that foxes were the most explorative genera regardless of socialization, which was determined in figure 5. These results have implications to the behaviors of these animals in current wild populations. With changing environments, foxes are living among humans, adapting to urban life, while several species of wolves are becoming endangered. This can be due to their generalist diets—being able to adapt to a wide variety of environmental conditions and live on a wide variety of resources, yet some of this can also be linked to exploration. This


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correlation then inquires about the distinction between beneficial exploration and negative exploration, which may provide an animal with a meal or end up killing them. Exploration and innovation is thus important to understanding conservation regarding wild popula-tions and reintroduction.

The box and whisker plots in figure 5 demonstrated differences in exploratory approach to a novel problem between genera. From the initial trials, the wolves seemed to mainly use their mouths and noses more in attempting to solve the puzzle. However, the foxes were observed to exhibit both behaviors almost equally, while the maned wolves appeared to favor pawing. The initial task in the two-part puzzle box apparatus was expected to have the animals pull the top off using their jaws, while the rotating tubes were expected to be solved with pawing. Some animals were successful by only using powerful digging behaviors, like the subjects in the species Chrysocyon (14A), while others solved it by chance. Thus, as the puzzle boxes were meant to evoke naturalistic behaviors from the subjects, this dif-ference in exploratory behaviors could be a reflection in their hunting and scavenging behaviors [2,14,17], or even a reflection of their physiological strengths. The genus Canis has the strongest jaws between the three groups and tend to catch the largest prey in the wild because of it [2,14], therefore their innate behaviors utilizing their powerful jaws may be involved in their ability to solve novel problems. For the inner tubes, the wolves mainly used their noses to flick it open, or their teeth when they bit it at a certain angle. Interestingly, the foxes, mostly arctic, and the maned wolves primarily used pawing. Initially, the arctic foxes tried to dig in and around the boxes to open them; however, they did use their mouths for the other parts to the puzzle eventually. The maned wolves used only pawing to solve all parts of the task. The reliance on this scavenging behavior of the maned wolves and foxes may be related to their more omnivorous diet of primary producers and smaller-size prey.

Although there were several significant differences within the observational data, there were also important similarities. First, all the animals who interacted with the boxes exhibited biting and pulling or pawing and digging, which both representative of canine’s sense of touch. For instance, in almost all trials, there was some type of sniffing behaviors present, which demonstrates the olfactory behaviors in use when the animals smelled the rewards inside the apparatus. The box and whisker distributions in figure 4 also revealed how all the subjects spent more time at the baited box rather than the control. Even though the animals may not have first investigated the baited box, they all eventually realized which had a reward, and thus concentrated their exploration to that apparatus. The subjects also exhibited sensory behaviors related to olfaction in addition to touch. Mainly, this was observed in whether or not the animals scent marked dur-ing the trial. Those who scent-marked were primarily the animals who solved the boxes. A Fisher’s Exact test determined a significant relationship between solving the puzzle and being more likely to scent mark. This observation of sensory behaviors, specifically olfaction, can be tied to communication as well. More often, the scent marking was done by the more dominant animals who were living in a group social environment. There seemed to be scent-marking done during trials where aggression between group members was also evident. Therefore, the puzzle box apparatuses evoked possessive and somewhat aggressive behaviors between animals who lived together. Even though aggression was evident, there were some displays to teamwork. There were a few instances where the subjects almost helped each other solve the problem, as they each equally figured out how to solve the problem. In most of these cases however, it was followed by aggression over the reward, demonstrating more competitive behaviors than cooperative. For instance, a group of subjects failed to complete the task as their intense social pack dynamics over a changing hierarchy distracted them from the puzzle boxes. Either way, the presence of scent marking that represented communication, further demonstrated another important use of canids’ sense of smell.

The assay results represented more of a case study but has significance to captive management in red wolves and zoos, in general. First, prior observations of the three subjects were made during their control periods. Red wolf 1A seemed the most active out of the three, as she would often be running around or pacing throughout the enclosure; red wolf 13A was observed laying down most of the time, occasionally walking around the enclosure; and the red wolf 1B was seen running around with 1A, when nobody was nearby, but mostly hiding the rest of the day. Looking at the assay results in figure 6 red wolf 1B seemed to have the highest stress levels. This may be attributed to the fact that he is somewhat blind. The large spike in stress at the beginning of the graph could be due to the horse races near the zoo. With a loss of sight, the wolf’s hearing and olfaction would be heightened to the point where he would become stressed with novel items or sounds around his familiar environment. Anyway, during the testing period for each of the subjects, their stress levels increased. Although an increase in stress would not be ideal for an animals’ well-being, it shows how the animals were physiologically responding to


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a novel stimulus in their environment. Despite the fact that these animals did not solve the puzzles, they definitely were affected by their presence. In most of the trials, the subjects were actually observed to exhibit more beneficial active behaviors, such as running around and playing as opposed to laying down most of the day. In a previous study, Cummings., et al. [11], it was actually suggested that a higher stress level could actually indicate a “good” stressor when in correlation with more active behaviors, in which case the puzzle box is a beneficial type of enrichment. However, more research would have to examine how exactly stress levels are corresponded to and interpreted with exhibited behaviors regarding beneficial enrichment for the animals. Also, more questions arise about what a similar analysis would look like over an extended period of time where the subject actually solves the puzzle.

The small sample size limitation caused several other variables within the study, which may have influenced the results. This influence can directly be seen in the assay results as only three subjects were used in this part of the study, which was the most that could be ob-tained. In the ethogram, small sample sizes also came into play as some of the categories ended up only having a few subjects. Additionally, this limitation made statistical analyses difficult as it was hard to make conclusions representative of the whole population with a small sample. Only a select few facilities agreed to conduct the study, and each facility had its own set of rules and schedules which had to be followed. Already this caused variations in the animals’ enclosure, diet schedule, individuality, and social environment. Thus, in trying to obtain a larger sample size, there were also more variables. In general, small sample sized studies are at a disadvantage with statistical analyses because many of the different tests make assumptions about your data set, which shouldn’t be violated.

Due to variables and limitations to this novel research study, future research should try to obtain a more constant and controlled sample size. However, there is a point in research with zoo animals where there will always be some limitations and uncontrolled vari-ables. Currently, a second study is beginning to examine the results of this novel problem-solving experiment over multiple trials with the same subjects. So far, some of the observable results suggested that novelty for some animals would wear off. While one subject has shown improvement in learning, others, who were initially very interested, experienced decreasing exploratory behaviors to the point where the animals didn’t even investigate the boxes at all. Another intriguing result suggested that foxes rely more on memory than on their olfac-tion after the first trial. In the second trials, the location of the rewards was switched and the foxes seemed to investigate the box where the reward was initially, rather than going directly to the one containing the reward. Thus, extending the testing period, and conducting multiple trials should also be a focus of future research. Other studies should examine more specifically the roles of innate sensory be-haviors on problem solving and cognitive sensory enrichment, with regard to their physiological responses using more quantifiable data.

Finally, this study has major implications for conservation. Although few statistical tests could quantitatively demonstrate clear sig-nificance, the suggestions about novel problem-solving and human socialization in canids can nonetheless be applied to the current ever-changing environment. The results had demonstrated that the more socialized wolves were the ones who were more likely to approach the boxes. This means that the more “wild” wolves were more afraid of approaching something new and different. Thus, wolves in their wild populations are diminishing because they are failing to adapt to the spread of human populations. Unlike foxes, who have adapted to living among humans, wolves are much larger and have more specific diets, making transitions like these much more difficult. Thus, hu-mans must stop encroaching on endangered canine habitats to ensure their species’ survival on earth. Furthermore, the high exploratory behaviors expressed by the foxes in comparison to the wolves can provide insight into current changes in canid habitats. With tempera-tures in the arctic rising, arctic fox habitats are shrinking and moving. Red fox success surviving in the wild also had led to their habitat range increasing. As habitats begin to overlap, it presents the animals with new competition in which the findings about their exploratory behavior could assist in predicting possible outcomes to the interactions in the wild. Even though the statistical analyses are approaching significance, the differences are still present and still have a relationship to the conservation of species persistence.

Conclusion

In summary, canids, members of the family Canidae, utilize specific sensory behaviors to solve novel problems. These behaviors were found to have differed among genus in their frequency of use. However, olfactory behaviors specifically were exhibited throughout most


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of the trials (such as scent marking and sniffing), as olfaction is used in finding prey and communicating with others. Human socializa-tion of the subjects played a crucial role in how the animals approached the boxes as well. The various levels of domestication influenced the neophobia towards the novel objects and human interactions impacted innate behaviors of the canids. The described neophobia and innate exploratory behaviors of canines are also important in understanding how to preserve and maintain their species in both healthy wild and captive populations, in terms of reintroduction success and captive management.

This study was the first to investigate the differences between foxes and wolves regarding behaviors toward an original novel problem solving task, that incorporated both senses and cognition. The hypothesis was mainly supported as there was a relationship determined between socialization and success on a novel problem, and the different genera exhibited exploratory behaviors in different ways. How-ever, the study was inconclusive on specifically how olfaction would play a role in problem solving, which is where future research can examine. The findings from the results are significant as they may have applications to conservation and improve our understanding of the various differences between canid genera. Future investigations should focus on the use of a larger and more constant sample size, reducing the number of variables between subjects and allowing for thorough data analysis of consistent results, which can be generalized to population conservation.

Appendix A: This table displays all the subjects included in the study.


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Appendix B: This graph is the standard curve used in the analysis of the assay results.

Appendix C: These box and whisker plots are representing the distribution of data for the

subjects grouped into the four categories.

Appendix D: These box and whisker plots represent the separation of the subjects into

even more refined categories.


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Bibliography

1. Boitani Luigi and Paolo Ciucci. “Comparative social ecology of feral dogs and wolves”. Ethology Ecology and Evolution 7.1 (1995): ‏.49-72

2. Lopez BH. “Of wolves and men”. New York: Scribner (1978).

3. Mech L David. “Possible use of foresight, understanding, and planning by wolves hunting muskoxen”. Arctic 60.2 (2007): 145-149.‏

4. Muro Cristian., et al. “Wolf-pack (Canis lupus) hunting strategies emerge from simple rules in computational simulations”. Behav-ioural Processes 88.3 (2011): 192-197.‏

5. Range Friederike and Zsófia Virányi. “Tracking the evolutionary origins of dog-human cooperation: the “Canine Cooperation Hypoth-esis”. Frontiers in Psychology 5 (2015): 1582.

Hedrick Philip W and RJ Fredrickson. “Captive breeding and the reintroduction of Mexican and red wolves”. Molecular Ecology 17.1‏ .6(2008): 344-350.

‏.Phillips Michael K and Warren T Parker. “Red wolf recovery: a progress report”. Conservation Biology 2.2 (1988): 139-141‏ .7

8. Sparkman Amanda M., et al. “Helper effects on pup lifetime fitness in the cooperatively breeding red wolf (Canis rufus)”. Proceedings of the Royal Society B: Biological Sciences 278.1710 (2010): 1381-1389.‏

9. Watters Jason V and Cheryl L Meehan. “Different strokes: can managing behavioral types increase post-release success?” Applied Animal Behaviour Science 102.3-4 (2007): 364-379.‏

10. Coelho Carlyle Mendes., et al. “Behavioral responses of maned wolves (Chrysocyon brachyurus, Canidae) to different categories of environmental enrichment stimuli and their implications for successful reintroduction”. Zoo Biology 31.4 (2012): 453-469.‏

11. Cummings Dawn., et al. “Behavioral and physiologic responses to environmental enrichment in the maned wolf (Chrysocyon brachy-urus)”. Zoo Biology 26.5 (2007): 331-343.‏

12. Hare Brian., et al. “Social cognitive evolution in captive foxes is a correlated by-product of experimental domestication”. Current Biol-ogy 15.3 (2005): 226-230.‏

13. Hersteinsson Pall and David W Macdonald. “Interspecific competition and the geographical distribution of red and arctic foxes Vulpes vulpes and Alopex lagopus”. Oikos 64.3 (1992): 505-515.‏

14. Sillero-Zubiri Claudio., et al. “Canids: foxes, wolves, jackals, and dogs: status survey and conservation action plan”. Gland, Switzerland: IUCN (2004).‏

15. Stickney Alice. “Seasonal patterns of prey availability and the foraging behavior of arctic foxes (Alopex lagopus) in a waterfowl nest-ing area”. Canadian Journal of Zoology 69.11 (1991): 2853-2859.‏

16. Bremner-Harrison Samantha., et al. “Behavioural trait assessment as a release criterion: boldness predicts early death in a reintro-duction programme of captive-bred swift fox (Vulpes velox)”. Animal Conservation Forum 7.3 (2004): 313-320.‏

17. Gadbois Simon., et al. “Revisiting the concept of behavior patterns in animal behavior with an example from food-caching sequences in Wolves (Canis lupus), Coyotes (Canis latrans), and Red Foxes (Vulpes vulpes)”. Behavioural Processes 110 (2015): 3-14.‏

18. Craven Brent A., et al. “The fluid dynamics of canine olfaction: unique nasal airflow patterns as an explanation of macrosmia”. Journal of The Royal Society Interface 7.47 (2009): 933-943.‏

19. Lord, Kathryn. “A comparison of the sensory development of wolves (Canis lupus lupus) and dogs (Canis lupus familiaris)”. Ethology ‏.110-120 :(2013) 119.2

20. Peters Roger and L David Mech. “Scent-marking in wolves”. Wolf and Man. Academic Press (1978): 133-147.‏


https://www.tandfonline.com/doi/abs/10.1080/08927014.1995.9522969

https://www.tandfonline.com/doi/abs/10.1080/08927014.1995.9522969

http://pubs.aina.ucalgary.ca/arctic/Arctic60-2-145.pdf

https://www.sciencedirect.com/science/article/abs/pii/S0376635711001884


https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4295532/


https://www.ncbi.nlm.nih.gov/pubmed/18173506


https://www.jstor.org/stable/2386098













http://www.carnivoreconservation.org/files/actionplans/canids.pdf

http://www.carnivoreconservation.org/files/actionplans/canids.pdf

https://www.nrcresearchpress.com/doi/10.1139/z91-402

https://www.nrcresearchpress.com/doi/10.1139/z91-402

https://www.uwosh.edu/facstaff/mcpheem/documents/personalities/Bremner-Harrison%201744.pdf

https://www.uwosh.edu/facstaff/mcpheem/documents/personalities/Bremner-Harrison%201744.pdf





https://onlinelibrary.wiley.com/doi/abs/10.1111/eth.12044

https://onlinelibrary.wiley.com/doi/abs/10.1111/eth.12044

200


21. Polgár Zita., et al. “Strategies used by pet dogs for solving olfaction-based problems at various distances”. PloS one 10.7 (2015): e0131610.‏

22. Polgár Zita., et al. “A test of canine olfactory capacity: comparing various dog breeds and wolves in a natural detection task”. PloS one 11.5 (2016): e0154087.‏

23. Rothman Russell J and L David Mech. “Scent-marking in lone wolves and newly formed pairs”. Animal Behaviour 27.3 (1979): 750-‏.760

24. Ryon Jenny., et al. “Scent rubbing in wolves (Canis lupus): the effect of novelty”. Canadian Journal of Zoology 64.3 (1986): 573-577.‏

25. Benson-Amram Sarah., et al. “Limited social learning of a novel technical problem by spotted hyenas”. Behavioural Processes 109 (2014): 111-120.

Range Friederike and Zsófia Virányi. “Social learning from humans or conspecifics: differences and similarities between wolves and‏ .26dogs”. Frontiers in Psychology 4 (2013): 868.‏

27. Benson-Amram Sarah and Kay E Holekamp. “Innovative problem solving by wild spotted hyenas”. Proceedings of the Royal Society B: Biological Sciences 279.1744 (2012): 4087-4095.‏

28. Thornton Alex and Jamie Samson. “Innovative problem solving in wild meerkats”. Animal Behaviour 83.6 (2012): 1459-1468.‏

29. Swaisgood Ronald R and David J Shepherdson. “Scientific approaches to enrichment and stereotypies in zoo animals: what’s been done and where should we go next?” Zoo Biology 24.6 (2005): 499-518.

Wells Deborah L. “Sensory stimulation as environmental enrichment for captive animals: a review”. Applied Animal Behaviour Science‏ .30‏.1-11 :(2009) 118.1-2

31. Palme R., et al. “Steroid extraction: get the best out of faecal samples”. Wiener Tierärztliche Monatsschrift 100.9-10 (2013): 238-246.‏

32. Walker SL., et al. “Reproductive endocrine patterns in captive female and male red wolves (Canis rufus) assessed by fecal and serum hormone analysis”. Zoo Biology 21.4 (2002): 321-335.‏

33. Young KM., et al. “Noninvasive monitoring of adrenocortical activity in carnivores by fecal glucocorticoid analyses”. General and Com-parative Endocrinology 137.2 (2004): 148-165.

-Benson-Amram Sarah., et al. “Brain size predicts problem-solving ability in mammalian carnivores”. Proceedings of the National Acad‏ .34emy of Sciences 113.9 (2016): 2532-2537.‏

35. Benson-Amram S., et al. “Problem-solving in Captive and Wild Spotted Hyenas”. Psyc EXTRA Dataset.

36. Liker András and Veronika Bókony. “Larger groups are more successful in innovative problem solving in house sparrows”. Proceed-ings of the National Academy of Sciences 106.19 (2009): 7893-7898.‏

37. Morand-Ferron Julie and John L Quinn. “Larger groups of passerines are more efficient problem solvers in the wild”. Proceedings of the National Academy of Sciences 108.38 (2011): 15898-15903.‏

38. Range Friederike., et al. “Testing the myth: tolerant dogs and aggressive wolves”. Proceedings of the Royal Society B: Biological Sciences 282.1807 (2015): 20150220.

‏.MacLean Evan L., et al. “The evolution of self-control”. Proceedings of the National Academy of Sciences 111.20 (2014): E2140-E2148‏ .39

40. Marshall-Pescini Sarah., et al. “The effect of domestication on inhibitory control: wolves and dogs compared”. PloS one 10.2 (2015): e0118469.‏

41. Moretti Lucia., et al. “The influence of relationships on neophobia and exploration in wolves and dogs”. Animal Behaviour 107 (2015): ‏.159-173






https://www.sciencedirect.com/science/article/pii/0003347279900101

https://www.sciencedirect.com/science/article/pii/0003347279900101







https://www.sciencedirect.com/science/article/pii/S0003347212001339

https://onlinelibrary.wiley.com/doi/abs/10.1002/zoo.20066




https://www.wtm.at/explorer/WTM/Archiv/2013/WTM_09-10-2013_Artikel_3_Palme.pdf





https://www.pnas.org/content/113/9/2532

https://www.pnas.org/content/113/9/2532












201


Volume 4 Issue 3 May 2019©All rights reserved by Abigail Reid.

42. Range Friederike and Zsófia Virányi. “Development of gaze following abilities in wolves (Canis lupus)”. PLoS One 6.2 (2011): e16888.‏

43. Range Friederike and Zsófia Virányi. “Wolves are better imitators of conspecifics than dogs”. PLoS One 9.1 (2014): e86559.‏

44. Udell Monique AR. “When dogs look back: inhibition of independent problem-solving behaviour in domestic dogs (Canis lupus famil-iaris) compared with wolves (Canis lupus)”. Biology Letters 11.9 (2015): 20150489.

45. da Silva Vasconcellos Angélica., et al. “Training reduces stress in human-socialised wolves to the same degree as in dogs”. PloS one 11.9 (2016): e0162389.

46. Sinn David L., et al. “Boldness towards novelty and translocation success in captive-raised, orphaned Tasmanian devils”. Zoo Biology 33.1 (2014): 36-48.

47. Berger-Tal Oded., et al. “Integrating animal behavior and conservation biology: a conceptual framework”. Behavioral Ecology 22.2 (2011): 236-239.

48. Greggor AL., et al. “Comparative cognition for conservationists”. Trends in Ecology and Evolution 29.9 (2014): 489-495.

49. Roth II., et al. “Cognition-centered conservation as a means of advancing integrative animal behavior”. Current Opinion in Behavioral Sciences 6 (2015): 1-6.

50. Gácsi Márta., et al. “Explaining dog wolf differences in utilizing human pointing gestures: selection for synergistic shifts in the devel-opment of some social skills”. PLoS One 4.8 (2009): e6584.










https://academic.oup.com/beheco/article/22/2/236/208826

https://academic.oup.com/beheco/article/22/2/236/208826






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