Feeding Behaviours in Wild Wolves

Feeding Behaviours in Wild Wolves

by

Gemma Pinkard

Declaration

This dissertation is a product of my own work and is not the work of any collaboration.
I agree that this dissertation may be available for reference and photocopying at the discretion of the University and give permission for Hartpury Library Services to hold and make available an electronic copy.

Name: GEMMA PINKARD
Date: 20th March 2024
E-mail address: gemma.pinkard@hartpury.ac.uk
Faculty/ School: HARTPURY UNIVERSITY
Student name: GEMMA PINKARD
Award: BSc (Hons) CANINE TRAINING AND PERFORMANCE
Module: ANIMAL AND AGRICULTURE DISSERTATION
Project/dissertation title: FEEDING BEHAVIOURS IN WILD WOLVES (CANIS LUPUS)

Acknowledgements

Firstly, I would like to thank my dissertation supervisor, Clare Thomas-Pino for all her guidance, knowledge and support throughout my dissertation. I would also like to say a huge thank you to Tony and the wolves at wolf watch UK. They welcomed me to the sanctuary and put everything possible in place to make this study happen. Thank you to Carol Stephens for all the study skills help I received along the way. I want to say a huge thank you to my mum Jenny Pinkard for keeping me on track with continued support.

A special thank you goes out to my dogs Loki, Blade and Ghost for being my emotional support at university.

ABSTRACT

Wolves (Canis lupus) have been studied extensively for many years, however, there has been limited data collection around feeding ecology. Secretive feeding, shyness, and living in extensive landscapes, have restricted access in observing this species in proximity, and ethically has been restricted due to intrusiveness of researchers. Continuing research produces a greater understanding of wolf behaviour around feeding, and how this affects social ecology. Here we assess how wolves change actions under the influence of external and internal factors or characteristics when feeding, and whether sex is a significant influencer.

A sample of two grey wolves were used as study subjects. The wolves were siblings and included one female and one male, that live as wild in a sanctuary. The researcher was positioned in an observational hide, with the wolves unaware of the presence of the observer. Dead kills were left, and quantitative data was collected by observing each wolf feeding behaviour, and documenting on an ethogram compiled by published literature. Video cameras, a camera, and iPhone were setup, to objectively record each session, and footage was viewed to identify any other types of behaviour not recordable on the ethogram. The results demonstrate the dominance of the female wolf and the high levels of aggression that individual displays when approaching and consuming prey, compared with a male showing submission in all behavioural categories of the ethogram. The study did not reveal any findings of social pack dynamics, and the male only showed live prey hunting behaviours. Distinct feeding behaviours were identified, and the importance of understanding male and female feeding ecology, and the roles they play.

CHAPTER ONE – INTRODUCTION

1.0 Introduction

1.1 History

Wild wolves (Canis lupus) are carnivores that feed primarily on meat (Vonholdt et al. 2020). Extensive studies however have failed to clearly evaluate species feeding ecology. Dietary data is limited to published literature that is often based on one common bioregional location (Newsome et al., 2016). Published literature however supports predictions of ecological significance (Mech, 2012) and prevents miscalculations (Lindenmayer et al., 2010). Newsome et al. (2016) concluded grey wolves diet varies on location, however, findings from southern Europe indicate grey wolves switch diets from domestic species to those available (Packard, 2018).

1.2 Ecology of wolf pack dynamics

Interbreeding of Scandinavian wolves is common due to small numbers existing globally. Smeds and Ellegren (2022) however limited the study to 73 wolves over 30 years. A larger pack number would have given a preponderant data pool to a field lacking in ecological research. An ethical consideration is small sample sizes suffer repeated intrusiveness by humans over a long period of time. Habituation to humans changes natural behaviours and can cause threatening displays of behaviour in feeding to conditioned ones (Løe and Röskaft, 2004). The reliability of the research therefore is questionable. Hand fed pups are often tame, however wolves indirectly fed by humans are wild (Fritts et al., 2003). Fear of humans is a major obstacle in studies involving data collection (Mech and Janssens, 2021). Long range scopes are used, and wolves are viewed from miles away. Incorrect classifications of specific behaviours related to a specific wolf therefore are often collated, making the results unreliable. Hierarchy is established among siblings when parents leave. In a study of 19 artic wolves in Moravia, Czech-Republic, Cafazzo et al. (2016) found no substantial differences between sex. Female-male pairs were less submissive, aggressive or dominant compared with female-female, and male-male pairings. Wolves were however observed in a captive environment. Captive environments restrict inherent behaviours being performed, along with hunting behaviours of live prey (Clubb and Mason (2003); Mason, (2010).

Northern grey wolves demonstrate complex social intraspecific behaviours within packs (MacDonald et al., 2004), however each wolf makes individual decisions and weighs up the costs/benefits of fighting. Males are more aggressive than females and repel rival opponents, and females chase away pack mates (Cassidy et al., 2017). Aggression however is due to K locus genes and inherent traits (Smith, Stahler and MacNulty, 2020). Newer research needs exploring in this area. Wolves are social where pack-mates are generally related, and hunt together (Mech and Boitani, 2003). High-ranking wolves can become subordinates to their pack mates and behave more defensively and shy (Packard, 2003). However, play can lower the occurrence of aggression in established hierarchies. Dominant relationships develop in mixed aged packs, but not puppy packs (Cordoni and Palagi, 2016). There is no comprehensive contrast in play types, and future studies could help researchers appreciate the roles of play. Social play is displayed more frequently than solitary play but Cordoni (2009) used captive wolves as study subjects.

Larger body size restricts locomotor performance and restricts foraging and predator trade-offs in male wild wolves. Nevertheless, males have better handling ability of large prey. Males therefore are better attackers, selectors and killers of prey (MacNulty et al., 2009). Smaller female wolves are faster than male counterparts and demonstrate quicker bursts of acceleration and better manoeuvrability (Biewener 2003; Dial, Green and Irschick, 2008). Nilsson and Bronmark (2000) and Herrel and O’Reilly (2006) concluded however males have increased gape capacity and bite strength for successfully taking down larger prey.

1.3 Hunting

Wolves can be extremely secretive when consuming food. Observational research is difficult to conduct in the wild (Ruth et al., 2019), cumulating in a lack of research in this area. Common hunting behaviours of live prey include ambushing, relay chasing and lie in wait (Peterson and Ciucci, 2003; Mech, 2007). Relay running is a cooperative behaviour involving the continuous chase and directing of prey, and members play different roles (Peterson and Ciucci, 2003). Observational research however can be subjective and contain personal bias. Feeding consists of hunting, searching and attacking prey. Grey wolves recognise the location of prey and encircle them. The hunt is controlled by the alpha, and the beta and delta wolves come together for the attack (Mirjalili, Mirjalili and Lewis, 2014). The Grey wolf optimizer is validated by current research (Storn and Price, 1997; Askarzadeh and Rezazadeh, 2012; Gandomi and Alavi, 2012). Wolves approach prey and maintain a safe distance until contact. Once in proximity to the prey, wolves move away from each other. When the hunt is concluded wolves lay down or maintain stationary positions (Muro et al., 2011). Cordoni (2009) found wolves substantially decrease playful activity in high conflict contests, such as feeding. The study however was conducted in a captive environment and not the wild. Therefore, dead prey provided does not replicate live prey behaviours observed in the wild.

Wolves consume carrion, rich in nutrient-detritus in the form of dead animal tissue (Smith, Stahler and MacNulty, 2020). Primary food selection in wild wolves is roe deer, with less kills of smaller prey (Jędrzejewski et al., 2012). However, this is based on geographical area and prey availability. Studies on large predator-prey relations are often restricted due to the availability of the predator’s main prey (Sand, et al., 2016).
Wolves eat a diversified prey species based on prey abundance in territories.
Functional response is the relationship between kill rate and prey abundance (Smith, Stahler and MacNulty, 2020). Prey abundance on the landscape is the most important factor impacting the figure of prey killed per predator (Holling, 1959). Scandinavian wolves are flexible in changing species selection as a pack or an individual (Sand et al., 2016). Sanctuary wolves have fewer captive restrictions with home ranges in a quieter setting, and feed on live prey as consumed in a wild environment, that includes, foxes (Vulpes), magpies (Pica) and ravens (Corvus) (Wilmers et al. 2003).

Pack foraging is common in wolves that hunt large ungulates. Defending territory is essential, and the risk of food loss to scavengers is great (Mech, 1999; Fuller, Mech and Cochrane, 2003; Vucetich, Peterson and Waite, 2004). Intraspecific competition affects pack dynamics in wolf populations. Smaller packs benefit from more territorial availability (Hochard, 2014). Intraspecific clashes of aggression can result in fatalities (Mech and Boitani, 2003). Nevertheless, non-reproductive wolves are not fixed to a den and travel throughout territories together (Cassidy et al., 2015). Globally the wolf home range size relates unfavourably to prey biomass, and pack density. Pack size, scope, and human density mirror larger home ranges (Ballard, Whitman and Gardner, 1987; Wydeven, Schultz and Thiel, 1995; Okarma et al. 1998; Fuller, Mech and Cochrane, 2003; Nowak, et al., 2007; Rich et al., 2012). Results however from these studies have been inconsistent due to factors influencing social and ecological circumstances (Mattisson et al., 2013; Rich et al., 2012) prey species choice (Fuller, Mech and Cochrane, 2003), and landscape characteristics equate with hunting success (Kauffman et al., 2007; Rich et al., 2012). Each play an equal role in balancing this effect. Inconsistencies between studies furthermore demonstrate home ranges are convoluted and influenced by social and ecological factors. Detailed observations of free-ranging wolves provide a more robust insight into territoriality, territory patrol, and neighbour competitiveness (Mech and Boitani, 2003). However, the study only used a Global Positioning System (GPS) on collared wolves and was not correlated across a wider wild wolf population.

1.4 Behaviour

Hunt success is based on intra-pack familiarity, cooperation, communication and protecting group resources (MacNulty et al., 2012). However, Cassidy and McIntyre (2016) concluded inter-pack relations had a higher risk of aggressive encounters, that are most frequent in the winter months leading to breeding season. Female oestrogen and male testosterone rise to a peak in February (Asa, 1997). Grey males are more aggressive compared with females, but pack size, age and being backed by pack mates greatly influences this finding (Smith, Stahler and MacNulty 2020). Cortisol levels in greys are higher than black coated wolves (Cassidy et al., 2017). However, newer research concludes black coats are highly aggressive, and aggression is directly attributable to species-specific genes. Ethically the findings of the study by Smith, Stahler and MacNulty (2020) were based on observations with Cassidy (2017) darting wolves to take saliva swabs. The question remains, how can high cortisol levels be reliably used when wolves are darted. Cassidy and McIntyre, (2016) conducted an observed study over 5,300 days into aggression in wolves. However, only six aggressive related encounters were recorded and discussed, concluding aggression between intra-pack families is low. Escalation to an attack is contact manifested as biting (Cassidy and McIntyre, 2016). However, defining behaviour through observations during data collection can be impeded by multiple aggressive encounters occurring at once (Cordoni and Palagi, 2016). Low ranking wolves submit to dominant counterparts by lowering their posture, licking the others’ mouth, wagging the tail and lowering ears and head (Baan, 2014). Behaviours were categorised by skilled animal behavioural identification specialists, and video recorded for objectivity.

Offensive behavioural patterns of play can be defined as play ambush, play bite, play jump, play paw and play run (Cordoni, 2009). The highest rank wolf commands play demonstrating competitiveness in the species (Cordoni, 2009; Smuts, 2014). Chasing is part of play and is observed when carcass feeding (McIntyre, 2019). During the absence of parents, siblings fight for alpha and beta status (Packard, Mech and Boitani, 2003). Through play, adult wolves obtain information of both the physical and cognitive skills of others for future challenges of successive position (Cordoni, 2009; Mech and Boitani, 2003). Female wolves have established themselves as alpha status in observed packs in Yellowstone (McIntyre, 2022). Many pack decisions are decided by the Alpha female (McIntyre, 2019), who dominate other females and have killed pups to retain resources for their own litters. Alpha females will fight males interspecifically and independently (McIntyre, 2022). However, other females have used cooperation over aggression in pack leadership. Alpha females start the chase and take down live prey (McIntyre, 2019). Younger wolves display kleptoparasitism and stash food away from the kill site, before returning to the carcass to feed. Subordinates challenge for food access and social relationships affect peaceful feeding and tolerance (Range and Virányi, 2015).

Tail position is the most reliable signal of social status. The baring of teeth however is not mirrored with dominant relationships (Fatjó et al. 2007). Dale et al. (2017) used animal carcasses to assess social relationships in wolves and concluded scent marking is a non-aggressive behaviour. Defecation and urination are scent marks left for other wolves to recognise conspicuous areas, and mark territories. Females urinate and males directly urinate over the top (Smith, Stahler and MacNulty, 2020).

1.5 Ecosystem effects and species interactions

Wolves will locate dens optimally near food supplies, and Alaskan wolves select sites that clear of snow expeditiously (Joly, Sorum, and Cameron, 2018). Seasonal variation can substantially affect kill rate due to limited prey abundance specifically in winter (Sand et al., 2008; Knopff et al., 2010; Metz et al., 2012). Nevertheless, Sand et al. (2016) found snow depth in winter or other predator related factors did not affect the prey kill rate in wolves. Wolves prefer to eat in the nocturnal hours, to avoid humans and increase hunting success in the twilight hours (Eggerman et al., 2009). The study by Sand et al. (2016) did not carry out observations of kill rate during the night, concluding the data of their study is not a true representation of consumption.

1.6 Rationale

The current study is unique. There appears to be much unpublished data on wolf feeding environmental science. Therefore, further research is essential to support studies that can facilitate wolf-feeding investigations (Baumann et al., 2021). The environment in which the wolves were observed in this study are unlike anywhere else globally. Furthermore, the wolves are not viewed daily unlike in zoos that restrict natural behaviours. The wolves only see the same person each day for feeding. Therefore, the data collection of the current study is distinctive, and could contribute to a significant scientific breakthrough in supporting current research, and providing new data for analysis that future studies can be built upon.

1.7 Research aims and objectives.

The overall aim was to gather scientific data through observational research on wolf feeding behaviours between male and female wolves, recorded around pre-feeding, feeding, and after feeding has finished. A further aim was to determine common behavioural trends, other behaviours, stereotypies, or those that are less frequent over three sessions.

To address the overall research, aim the objectives were:

  • to conduct three observational visits at a wolf sanctuary in England.
  • to capture data of two sibling wild wolves by completing ethograms and videoing the pair of wolves during feeding times of kill provided.
  • to complete behavioural analysis to identify behaviour types and trends.

CHAPTER TWO – METHODS

2.0 Methodology

Many researchers have utilised observations to measure behaviour in wolves (Fatjó et al. 2007; Cubaynes, Macnulty, Stahler 2014). Currently however study on species feeding ecology in wild wolves is extremely limited (Newsome et al., 2016). Many studies have focused on wolves in captive environments (Piffare et al., 2012; Irene et al., 2019) or have conducted studies viewing wolves from great distances questioning the validity of results. These differences employ the need for different methods of observation for assessing feeding behaviours in wild wolves.

2.1 Study Design

Between November and December 2023 three observational sessions took place at a sanctuary in Shropshire, lasting 60 to 90 minuets. The enclosure was environmentally wild, with rock faces, densely populated pine trees, wild grasslands, and a lake, that replicate wild wolf territories (Figure 1, Figure 2, Figure 3). Prior to commencing the study, the researcher conducted a site visit in November 2023. The advantage of a site visit prior to the study starting enabled the observer to understand the physical environment, talk with professionals onsite and to foresee any potential limitations or mistakes that may impact with the data collection. The site visit identified a den close to the feeding location. Therefore, the likelihood was greater of the wolves coming out to feed during the observations (Joly, Sorum and Cameron, 2018).
Nonetheless, no pilot study was conducted because of limited access to the research site. An advantage of a pilot study would have highlighted the need for a longitudinal study, in particular the analysis of feeding behaviours were restricted when wolves took the food into cover.

Figure 1: Wild terrain inside the sanctuary (Author’s own, 2023).
Figure 2: Densely populated pine forest (Author’s own, 2023).
Figure 3: Forest opening where the wolves first present for feeding (Author’s own).

A signed learning agreement (See appendix A), Project proposal form (See appendix B) were completed at the start of the study. Prior to commencing the study, ethical clearance was sought from Hartpury Ethics committee (ETHICS2023-88-LR) (See appendix C) and was granted. A site permission form (See appendix D), a risk assessment (See appendix E) and Hazard risk assessment completed prior to starting the study (See appendix F).

Observational field studies are a key method to collect information gathered by experimental methods to characterise ecological legitimacy (Komulainen, 2022). The present study uses an ethnographic approach including an ethogram (See appendix G, H and I) to measure the behavioural responses of wolves when feeding. The ethogram was completed manually at the time the behaviour was observed. The ethogram was selected for quantitative reliability and validity and was based on the one adopted by The International Wolf Sanctuary. Quantitative variables are known as continuous and quantitative research gathers data analysed through numbers (Hawkins, 2019, p.255). Ethograms have previously been chosen to give a detailed understanding of classifying large carnivore behaviours (Goodmann et al., 2002; Frézard and Pape, 2003; Mech and Boitani, 2003; Pifarré et al., 2012). In observational studies there is potential bias from observers. To rule out this possibility, all sessions were recorded using a Sony camera and Apple Iphone. The wolves were observed from a close vantage point (See figure 4). The design of the lookout is rustic and blends in with the environment. The lookout is elevated, and positioning stops the observer being scented or seen. Natural behaviours can then be displayed around the kill. A major advantage was the observer being able to clearly identify the behaviour being displayed, and by what wolf. Previous research has relied on scopes viewing wolves from great distances, often resulting in incorrect identification of individual behaviours, or being unable to record multiple behaviours from multiple wolves occurring at once (Janssessens and Mech, 2021).

Figure 4: The lookout point (Author’s own, 2023).

On arrival at the sanctuary the time to bring the kill to the feeding site was agreed upon by the observer and the sanctuary owner. To rule out the possibility of influence of behaviour, the same daily feeding site was used. The location is the brown feeding spot (figure 5).

Figure 5: The feeding site next to the large brown area of dead plant matter (Author’s own).

The observer entered the hide and set up the cameras The wolves came into the open when the food was brought to the site. Data was collected when the wolves started to interact in anyway with the feeding process. It was not possible to investigate the significant relationships of feeding behaviours in sessions one and two extensively due to the time the wolves spent feeding in cover. A causal factor could be the weather conditions of strong wind and heavy rain. Wolves save energy by not hunting in wind and rain. Further data collection is required to determine exactly how the weather affects feeding behaviour. The observations were not finished until the wolves had consumed all the food and had gone back into cover.

2.2 Sampling

The sample size was a wild pack of two nine-year-old sibling Norwegian Grey Wolves. Rickon, a male, and Sansa a female. A small sample size is justifiable for furthering essential research into feeding ecology in wild wolves.

Wolf one: Identifiable as smaller than brother Rickon, Sansa has black along the spine, that clusters in a square mid back, and on the tip of the tail (Figure 6).

Norwegian Grey Wolf - Gemma Pinkard
Figure 6: Sansa (Author’s own, 2023).

Wolf two: As a male Rickon is structurally bigger than Sansa. Rickon has a darker pigment on the face, but whiter block colour on the torso (Figure 6).

Norwegian Grey Wolf
Figure 7: Rickon (Photograph WolfWatch UK, n.d.)

2.3 Data analysis

The raw data from the three ethograms (See appendix G, H and I) was collected and input into an Excel spreadsheet (See appendix J). Pie charts were generated for each of the sessions to illustrate the individual behaviours of each wolf exhibited (Field, 2017). In addition, a bar chart was created to visually represent the frequency of individual behaviours (Hawkins, 2019). Subsequently, thematic analysis was conducted to identify key trends and patterns, which emerged from the raw data and descriptive statistics (Braun and Clarke, 2006). The camera footage captured was viewed and analysed in its entirety. The analysis entailed looking for any other types of specific behaviour not documented on the ethograms. This was justified to generate further added evidence of behaviour not documented in the ethograms.

A positivist approach was adopted which is appropriate for quantitative research methods (Cottrell, 2014). A positivist research philosophy takes the view only factual knowledge through observation, including measurement, is reliable. Positivism addressed the research aims in this study (Clark et al., 2021). Data collected is a natural study as data was collected in the field. Scale (counts and measures) allows the difference between values to be measured. The scale method is a strongest level of measurement compared with nominal (categories) measurement (Hawkins, 2019)

2.4 Ethics

No data was collected before ethical clearance was granted by the Hartpury Ethics Committee. For ethical reasons the decision was made that the wolves were only exposed to one experimental condition. Moreover, with the researcher observing from the hide, there was no direct contact with the wolves, and the researcher could not be seen. Therefore, the research was not intrusive and in no way could cause any physical or psychological harm (Cottrell, 2014). Anonymity and confidentiality were maintained (Whisker, 2019). The data was stored securely on the Hartpury One Drive, and password protected, in accordance with the General Data Protection Regulations (GDPR), and the Data Protection Act 2018 (GOV.UK). Researcher bias was avoided as far as possible through the quantitative ethogram data collection method supported by the objective video evidence.

CHAPTER THREE – RESULTS

3.0 Results

3.1 Ethogram results

The data from the three ethograms (See appendix G, H, and I) were analysed and identified nineteen key behaviours, and ‘other’ behaviours around feeding in wolves.

Table 1: showing the summary statistics for frequency and the mean of feeding behaviour, recorded over the three sessions.

BehaviourSession 1Session 2Session 3Mean (Average)
Relay chasing5133
Circling prey3132
Hunting live prey0183
Stalking/Hurding2122
Tall position changes4174
Aggression443815
Escape1142
Indirect approach2142
Centuary/territory checking5313
Follow3153
Vocalisations0100
Body posture changes2194
Urination/defication1052
Changes in play0000
Approach0183
Creep/crawl/crouch0052
Submissive behaviour3174
Critical reaction0031
Leave0101
Other behaviours0183
Figure 8: Graph displaying the frequency of observed behaviours of both wolves during feeding. What stands out in the table is the high rate of aggression in session 3.
Figure 9: Pie chart displaying the percentages of behaviour frequency in session 1.
Figure 10: Pie chart displaying the percentage of behaviour frequency in session 2.
Figure 11: Pie chart displaying the percentage of behaviour frequency in session 3. A striking result in this figure is the outlier of aggression.
Figure 12: Aggressive behavior displayed in each of the three sessions

Table 2: The breakdown of results obtained of pack hierarchy and feeding behaviour in sessions 1-3. Analysis of the data was compared between male and female sex.

BehaviourFemaleMaleFemaleMaleFemaleMale
Session number112233
Searching for live preyNoNoNoYesNoYes
Ambushing live preyNoNoNoNoNoYes
Relay chasing live preyNoNoNoNoNoYes
Circling Live PreyNoNoNoNoNoYes
Working as an equal packNoNoNoNoNoNo
Maintaining distance from preyNoYesYesYesNoNo
Circling when prey brought to siteNoYesYesYesYesYes
Approaching food as an equal packNoNoNoNoNoNo
Confident in approaching food from humanNoNoYesYesNoNo
Eating firstYesNoYesYesYesNo
Consumed the most foodYesNoYesNoYesNo
Aggressive around feedingYesNoYesNoYesNo
Eating in coverYesYesYesYesNoNo
Eating in the openNoNoNoNoYesYes
Decrease in playYesYesYesYesYesYes
Urinate during feedingYesYesYesYesYesYes
Defecate during feedingYesYesYesYesYesYes
Leader in the followYesNoYesNoYesNo
Leader in relay chasingYesNoYesNoYesNo
Dominant wolfYesNoYesNoYesNo
Submissive wolfNoYesNoYesNoYes
Other behaviourNoNoNoNoYesNo

Notes were made on the ethogram (See appendix G, H and I) for behaviours not listed, these are recorded as ‘other’ behaviours. Observed behaviours classed as ‘other’ included scent marking and returning to a den close to the feeding site in session 3. Both behaviours were performed by the female wolf. Session 3 also observed the female stashing food and aggressively defending it.

3.2 Video results

Video recorded sessions were watched in full and any ‘other’ behaviour displayed were recorded manually on the ethogram to ensure no duplication of results.

CHAPTER FOUR – DISCUSSION

4.0 Discussion

4.1 Introduction

There is global interest in feeding ecology in wild wolves. However, secretive feeding (Ruth et al., 2019) and large territories hinder the success of gathering scientific research through observational studies. This has directly resulted in a significant lack of rich and up to date data in feeding behaviours. This study aimed to provide scientifically reliable data directly related to behaviours exhibited by both male and female wolves, during the feeding process.

4.2 Pack hierarchies

The current investigation found no evidence of social pack dynamics. What is striking in table 2 is that across all 3 sessions the wolves did not work as an equal pack when food was brought to the site, or approach food as an equal pack when it was left. The discrepancy could be attributed to food being provided daily so there is little need to hunt. The study was unable to demonstrate that wolves are social and hunt together when pack mates are related in contrast to earlier findings of Mech and Boitani (2003). An interesting finding was the male wolf did not at any time challenge the female for alpha status and remained the subordinate wolf. This inconsistency maybe due to only a small sample size of two wolves being used. Packard (2003) showed that high-ranking wolves can become subordinates to their pack mates and behave in a shy manner, and that in the absence of parents, siblings fight for alpha and beta status. This differs from findings presented here. The study established the male demonstrated submissive behaviour across all three sessions to the female who remained the alpha wolf, and as siblings they did not challenge for pack status. What is interesting, is figure 9 has the highest frequency of submissive behaviours at 9% in session 1, but session one had the lowest aggression score at 11%. An alternative explanation for the result is that it was due to the wolves only being observed for a short time before they returned to cover. This study is consistent with Bann (2014) who concluded lower ranking wolves will submit to higher ranking wolves. Further research is needed to explore the mechanisms between siblings in packs.

4.3 Aggression

The results in this study found female grey wolves are more aggressive than male counterparts when consuming food. Figure 8 indicates there was a sharp increase in the number of recorded aggressive encounters in session 3. The possible explanation might be the wolves spent more time in the open so were observed for longer. However, an alternative explanation for this result is that this is a true reflection of what intraspecific aggressive behaviours around feeding occur naturally in the wild. The outcome of this result is contrary to the findings of Cassidy et al. (2017) who reported males being more aggressive than females. Another finding is the female chased away the male four times in session 3, and strikingly table 2 shows the male did not display any signs of aggression to the female across all three sessions. This intriguing result might have been attributed to the female being of alpha status in the pack. However, these results support previous observations of Cassidy et al. (2017) that females chase away pack mates, and males only fend off rival opponents, which could be a causal factor in the result. On the question of aggression this study found aggressive behaviours in all three sessions. What clearly stands out in figures 9, 10 and 11 is aggression has the highest percentage of observed behaviours. Several factors could explain this result. Firstly, aggressive encounters are highest in the winter months, when female oestrogen and male testosterone levels rise to peak in February as reported by Asa (1997) and Smith, Stahler and MacNulty (2022), who also conducted observations during the winter months. The most obvious findings to emerge from the analysis of aggressive encounters is that the lunge along with the approach were the most frequently displayed behaviours across all three sessions. This finding can be attributed to food becoming available/left at the kill site, and the consuming of food at this time. What stands out in table 2 is the female wolf eats first, consumes the most food, and was dominant in all three sessions. Table 1 clearly shows 9 incidents of body posture changes in session 3, and figure 12 shows a gradual increase of aggressive behaviours being observed across all 3 sessions. What clearly stands out in figure 12 are highly aggressive behaviours of ritualised attack, give eye and baring of teeth evident in session 3. The outcome is contrary to that of MacNulty (2012) who concluded that aggression between intra-pack families is low. No aggression was displayed when all food was consumed.

4.4 Hunting

As mentioned in the literature review common hunting behaviours of live prey include ambushing and lie in wait (Ciucci, 2003; Mech, 2007). The study further supports this idea as ambushing is seen, but in figure 2 relay chasing and searching for live prey were demonstrated. One unanticipated result was the male only displayed hunting live prey behaviours. However, the inconsistency may be due to the wolves relying on food being provided, and that the female ate most food in the sessions. The live prey species in session 3 was a magpie which supports the findings of Smith, Stahler and MacNulty (2020) and Wilmers et al. (2003), who found captive wolves feed on magpies as they would in the wild. In this study when food was brought close to the kill spot the wolves did not move away from each other. This outcome is contrary to that of Munro et al. (2011) and Cordoni (2009) who found that wolves move towards each other. The results of this study did however reflect those of Munro et al. (2011) and Cordoni (2009) observations of wolves decreasing playful activity during feeding, and maintaining stationary positions once feeding was concluded. However, these data must be interpreted with caution because food was provided by a human, and not replicable of prey consumed in the wild.

4.5 Other behaviours

When food was left, the male was observed in video analysis slinking behind the female and backing off to allow her to eat first. This supports McIntyre (2022) that females have established themselves as alphas in established packs, and pack members play different roles (Peterson and Ciucci, 2003) but is contrary to findings by Caffazzo et al. (2016) who found no substantial difference between sex.

Another important finding is changes in tail positions across all three sessions. Each ethogram (see appendix G, H, and I) significantly found that tail position 4 (tail between legs) was the only position demonstrated by the male in all feeding sessions. This finding broadly supports the work of Fatjó et al. (2007), who linked tail position changes as the most reliable signal of social status. It is possible the results in this study may underestimate the role of tail position in each session due to the partial consuming of food in cover in sessions 1 and 2, so feeding behaviour could not be fully observed.

Perhaps the most interesting finding in session 3 was the display of kleptoparasitism and stashing of food away from the kill site by the female, who aggressively defended the food from the male. This finding of stashing food by younger wolves and challenging by subordinates was reported by Range and Viranyi (2015). In contrast, however, the study did not provide evidence of male wolves stashing the food as reported by McIntyre (2019). A note of caution is due here since a sample of only two wolves was used, and the wolves were only observed consuming all food in the open in session 3. This behaviour may not have occurred in consumption of food in cover in session 1 and 2. Therefore, further investigations are needed to confirm or validate these findings.

Nothing was found in the literature review of females rubbing their necks into the ground after feeding. However, this study observed this behaviour. These findings may be taken to indicate scent marking, but it is possible that these results do not represent this behaviour. Further investigations are required to determine whether this behaviour is linked to scenting. What can clearly be seen in figure 9 and 11 is the frequency of defecation and urination in these sessions, and table 2 clearly shows both sexes performed this behaviour during feeding. However, the findings of the current study do not support previous research by McIntyre (2019) that defecation and urination are scent marks left for other wolves, and that when females urinate males will urinate directly over the top. Therefore, these findings are disappointing. To develop a full picture of urination and defecation in feeding additional studies will be needed to establish how this behaviour is linked to feeding. In summary, the results of this observational study indicate female wolves exhibit alpha status behaviours in the feeding process and show high levels of aggression to subordinate wolves when consuming food.

4.6 Study limitations

A limitation of this study is the small sample size, which means the findings cannot be generalised to a wider population (Wisker, 2019). A further limitation was not all the behaviour could be observed or filmed due to the wolves eating in cover. Observational studies can be more prone to subjectivity however, this was mitigated using video data capture and a scientifically evidenced-based ethogram.

4.7 Future research

Further research using a longitudinal observational study would establish larger data collection and allow for the development of more specific and reliable analytical method for statistical analysis. Previous research has been extremely limited due to large distances for observing wild wolves. Further investigations into sibling behaviour would evaluate the impact of relationships on behaviour. The study however is unique and allows for accurate and valid data though close and controlled observation justifying a small sample size. Researchers can use the data collected in this study for future to research into feeding ecology in wolves.
The research generated a large quantity of raw data in the form of contemporaneous notes from the researcher. The data could have facilitated further comparisons or trends in feeding behaviours of ecological significance. The data was not used due to concerns over the validity of scoring the same behaviour twice, but future studies can draw on these findings to construct new hypothesis around feeding behaviours in wolves.

CHAPTER FIVE – CONCLUSION

5.0 Conclusion

The aim of this study was to explore the influence of feeding on behaviours exhibited by wild wolves. The research has identified distinct behaviour differences in both female and male wolves. One of the more significant findings clearly highlighted a female demonstrating dominant alpha behaviours especially when consuming the food, with a male beta wolf always being submissive to their counterpart. This study concluded subordinates will submit to higher ranking wolves. The results of this investigation report the wolves did not work as a pack when food was brought to the site, or on approaching the food when it was left. The wolves did not hunt together or demonstrate high levels of sociability as concluded in previous studies. However, relying on food being brought each day could be a contributory factor for the lack of sociability and the hunting of live prey. The study contributes to our understanding of the effects of prey abundance and the need for wolves to hunt. Another key finding was the inordinate nature of aggressive behaviours exhibited frequently throughout the observation periods, and the significance of these in both sex and the sibling relationship. Notwithstanding the relatively limited sample, this study offers some insight into how sex and sibling relationships affect pack hierarchy, sociability, and behaviour around feeding. This study lays the groundwork for future research that could beneficially explore the effects of sibling relationships on feeding using a bigger sample size. Prior to this study it was difficult to make predictions about how wolves behave around feeding. This is due to limitations of observing wolves at great distances over extensive landscapes. Nonetheless, this study is the first of substantial duration which examines associations between behaviour and feeding in wolves.

Author: Gemma Pinkard

EXPLORATORY STUDY OF FEEDING BEHAVIOURS IN WILD WOLVES (CANIS LUPUS). 20th March 2024

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    A Wolf in Dog’s Clothing by Mark Derr

    Emerging from the deep shade of a sandstone outcropping that shelters their flock, three skinny black-and-white dogs warily approach pieces of cantaloupe rind thrown to entice them into the open, sniff, then begin eating, their eyes fixed on the strange Anglos talking with their Navajo owner. I am amazed at how much they resemble a photograph I recently saw of the Basketmaker dog, a rare, complete mummy dating from the time of Christ that was found at White Dog Cave, not far from this hogan, in 1921 and resides at the Peabody Museum, Harvard University. Travelling through the Navajo Reservation with Hal Black, a zoologist from Brigham Young University, I will observe a dozen more of the dogs, some with buff coats and grey muzzles, but of the same physical type as the 2,000-year-old mummy, as if in this country of wind-blasted sandstone mesas there is no divide between the quick and the dead.

    Bred to no particular purpose, the Navajo dogs, that range from fifteen to sixty pounds, live with flocks of sheep and goats they protect from coyotes, other dogs, horses, mules, even strange people who come too close. Most are born among the sheep and goats they accept as members of their own pack, but others are adopted as barely weaned puppies from the ranks of feral dogs (who have severed their bond to humans and grown to fear and avoid them) living around the reservation’s garbage dumps or found on the roadside. I recognize many of them as mutts from modern breeds and dismiss them, not because they are less good as sheep guards, but because I am fascinated with the ancient ones. The latter remind me of the feists and curs of the American South, that are generally believed to descend from the dogs of Native Americans, mixed with those of seventeenth and eighteenth century colonists. It seems incredible that the type could persist for so long without change despite exposure to countless other dogs and I would like to believe that my eyes have deceived me, the way I know when my male Catahoula leopard dog sleeps on his back in a contorted pose resembling the dog from Pompeii zapped in the ash of Vesuvius that the relationship is purely visual.

    Back home in Miami Beach, I check with Stanley J. Olsen of the University of Arizona, one of the world’s foremost experts on dog palaeontology and a man given to scepticism regarding claims that certain dogs represent ancient breeds. “Oh, yes,” he says, “those little dogs on the reservation – they look just like the Basketmaker mummy.” He agrees that a comparative study would be interesting, but for now the techniques of genetic analysis are not refined enough to determine whether the sheepdogs are heir to the animals of people who lived in that land of buttes and mesas before the Navajo themselves arrived.

    Around the world, there are dogs that have apparently remained unchanged for thousands of years – bred true to type – often on islands where ancient wanderers dropped them, in jungles, parts of the Arctic or relatively remote desert environments like that of the American Southwest, where for long periods they would have come into contact with other dogs rarely, if at all, but also in regions where people have retained a strong tradition of using certain kinds of dog. Some researchers even speculate that many of these dogs are derived from an ur-dog domesticated 10,000 or more years ago from the Indian wolf and carried around the world with migrating bands of people, mixing along the way with indigenous wolves. In its effort to account for the affinities in behaviour and appearance among these unique dogs, this theory oversimplifies the process of domestication and dispersal. Foremost among them and closest to the wolf in appearance and behaviour is the dingo, that first appeared in Australia some 4,000 years ago when seafarers from Indonesia or Southeast Asia beached their dugouts to trade with the Aborigines and lost some of the dogs they carried for companionship and food. The dogs reverted to the wild and became the top carnivore, next to humans, on that island continent, joined over the centuries by other travellers who went walkabout. Although some Aboriginal tribes tamed puppies and kept them as hunting aides and camp guardians, as well as food in times of famine, the dogs bred in the wild and in general behaved so differently from those of European explorers arriving in the eighteenth century that they were called dingoes and declared a separate species.

    The New Guinea singing dog, now nearly extinct on its home island, is said to be a dingo of sorts, as are the pariahs, the ownerless dogs who live around towns and villages in Southeast Asia and even some of the Native American dogs. A number of Middle Eastern and African dogs are similar in appearance, but probably domesticated from different subspecies of wolf. The Canaan dog from Palestine was a pariah used to guard and herd sheep until the 1930s, when Rudolphina Menzel, an expert on dogs who, with her husband Rudolph, had fled Hitler’s Germany, consolidated it into a breed for use as a messenger, tracker, search-and-rescue dog and guide dog. Among the !Kung San bushmen of the Kalahari Desert, those who hunt with dogs – medium-sized buff or piebald animals – bring home 75 percent of the animal protein their band consumes. Pygmies use little hounds (refined by English and American breeders, they are called basenjis) to hunt birds and other game. On Sicily and in Portugal are graceful prick-eared hounds that appear to have changed little for several millennia.

    In fact, dogs like these are sought by collectors in increasing numbers, because they are deemed “primitive” – more quintessentially dog in their abilities and demeanour than the “refined” European and English breeds: pointers, retrievers, toys, terriers and other denizens of the show ring. Even the various curs and feists of the American South and the Arctic sled dogs are often called “primitive.” Despite all the rationalizations and examples used to support the distinction, it primarily refers to dogs that are more generalist in their talents, independent in their habits and relatively free of disabling genetic defects compared with those selectively bred for specific traits, size, colour and specialized talents like pointing. Since many are country dogs, they are deemed exotic or rare, when taken as pets. I prefer the word “basic” to “primitive”, because it bears less cultural baggage. It also recognizes that types like the Alaskan husky and curs have, over the years, received infusions of new blood without losing their distinguishing characteristics. Huskies retain their tough feet, somewhat wolfish appearance and habits as sled dogs despite the presence in their midst of individuals with lop ears and thinnish coats. Coming in a range of sizes and colours, curs are identified by their ability as herders, hunters that trail and tree, occasional pointers and guardians, as well as by their general deep-chested build.

    In Australia, dingoes are currently hybridizing freely with domestic dogs, raising concerns that they will become extinct. Hybridization occurs most frequently in areas where human predation has created a shortage of available dingo mates, meaning humans can help reverse the process by ending the senseless slaughter. To the dingo, however, hybridization has always offered life, not extinction. In the centuries before Anglo settlement, it interbred, especially along the coast, with dogs arriving, as its forebears had, with Southeast Asian and Indonesian traders. Like those early hybrids, many of the ones produced today are virtually indistinguishable from dingoes into whose society they are born. The dingo phenotype and culture prevail, leading me to conclude that the obsession with curbing interbreeding has less to do with preserving the dingo than with maintaining old notions of blood purity. Such a view is heretical in the world of wildlife protection, but the dingo is a dog that went wild because of the circumstances in which humans left it; if it changes in relationship to new human-made conditions, it is simply being a dog.

    Whatever terms we use, the attempt to draw clear distinctions between basic and pedigree show dogs or even between breeds, reflects our continuing attempts to understand the animal that shares our lives more intimately than any other. Under funded and assigned low priority by palaeontologists, archaeologists and evolutionary biologists, whose efforts are directed more toward examining issues relating to humans, extinct and endangered species, and those efforts proceed in fits and starts, like a dog trying to fix on a cold trail.

    Whether read on cuneiform tablets, scrolls, bas-reliefs, paintings, books, film, or the flickering pixels of cyberspace, divined from bones or mummified flesh, deciphered from the genes, what we know remains an unstable mixture of fact and received wisdom, which is too often accepted as revealed truth. As biologists decipher the dog genome – the genetic blueprint that makes it unique – archaeologists open new sites and behaviourists deepen their knowledge of dog and wolf behaviour, the story will doubtless become, paradoxically, more clear and complex. On a practical level, I hope that this knowledge will lead to a revolution in breeding that will bring an end to the production of mutant animals fit only to serve human vanity and create animals of good health and temperament, sound minds and abundant talent. Bred to type, like the sheep guards of the Navajo, the curs and huskies, these dogs would show considerably more variability than is allowed in the narrowly prescribed physical standards of show dogs, like the Pekingese, malamute or any of the other 140 or so pure breeds recognised by the American Kennel Club.

    Enough has been learned over the past three decades to allow concerned breeders and trainers to make dramatic improvements, but more must be done. The chief drawback to that reform, one expert told me, lies in inadequate dissemination of the information at hand and continued reliance on folk wisdom that views inheritance and behaviour in overly simplistic terms. I would add to that list an unwillingness among many people involved with dogs to change their ways.

    Defining Dog
    What we know is this: The dog is a subspecies of the wolf altered over more than fifteen millennia by selective breeding. Analyses of mitochondrial DNA, which is inherited only from the mother, have shown no distinctive differences between wolf and dog or even between breeds of dog, no matter their shape and size. (DNA fingerprinting does allow scientists to identify individual dogs, but not their breed or type.) The New Guinea singing dog and dingo appear to have one or two distinctive genetic markers, perhaps due to thousands of years of island isolation, but they are not significant enough to distinguish them as separate species. Contrary to theories set forth in the past and still repeated in some quarters, no contributions were made by jackals, coyotes, foxes, otters or bears, nor were there any ur-dogs that appeared suddenly on the earth and then vanished into the bosom of domesticity, like a dog Adam and Eve. Our dog is formally Canis lupus familiaris.

    Canis means “dog” in Latin, so the dog is technically a domesticated wolf, which is a wild dog. Canis lupus is one of thirty-four living species grouped in Canidae (the dog family) of the order Carnivora, which also includes Ursidae (bears), Mustelidae (weasels), Procyonidae (raccoons), Ailuropoda (pandas), Otariidae (sea lions), Odobenidae (walruses), Phocidae (seals), Felidae (cats), Viverridae (civets) and Hyaenidae (hyenas).

    Collectively the carnivores are intelligent animals that care for their young and possess relatively large dogs for killing, carnassials – the first molar on the lower jaw and last premolar on the upper – for rending flesh and molars for crushing bones. They have four to five toes with claws that are retractable in the cats, except the cheetah, and not in the others. All lack the opposable thumb, even those with five digits. In dogs, the fifth toe of the fore and hind feet has become a dewclaw, although some breeds have no rear dewclaws while others, especially among the French sheepdogs and some yellow blackmouth curs, have two on each foot. Dogs and cats walk on their toes; bears on their heels and soles. Classification, being a less than exact science, some of these carnivores are omnivores and one, the panda, eats bamboo. Still, among this group are the top terrestrial predators, next to humans – the only natural enemy of many of them.

    Canids – members of the dog family – began to distinguish themselves from other mammalian carnivores some 50 to 60 million years ago, almost immediately upon their first appearance following extinction of the dinosaurs. These animals were miacids – ferret – to fox-size creatures with a longer body than legs, tails and those mashing and cutting teeth. Miacids gave way to larger creodants with five distinctive toes. Around 15 million years ago in the Western Hemisphere, another fox like animal, Hesperocyon, appeared, walking on its toes. From there the line passes through Leptocyon, believed to be the common ancestor of wolves and foxes. Canis lepophagus, whose remains were found in Texas and dated to the Pliocene some 5 million years ago, might be the forerunner of the wolf like canids. From their origins in what is now North America, early canids migrated to Eurasia, Africa and South America.

    By the best current estimates, 7 to 10 million years ago the dog family began to divide into the broad groupings we see today: the wolf like canids, South American canids, red foxes, and miscellaneous. The foxes, miscellaneous and South American canids have different numbers of chromosomes from the wolf like canids and do not figure in the evolution of the wolf, although the South American bushdog (Speothos venaticus), which dives under water, has been domesticated occasionally.

    The wolf like canids have seventy-eight chromosomes and could conceivably all be classed as Canis, but two are not: Lyacon pictus, the African wild dog, with four toes front and back and the highly variable markings usually associated with domestic dogs and Cuon alpinus, the dhole or red “dog”, native to Asia and India. Those grouped in Canis are the wolf (lupus); golden jackal (aureus); side-striped jackal (adustus); black-backed jackal (mesomelas); Simien jackal or Ethiopian wolf (simensis); coyote (latrans) and red wolf (rufus). The huge dire wolf (Canis dirus) rose and fell during the Pleistocene, while its cousin, the gray wolf, flourished.

    Although the wolf, coyote and golden jackal probably diverged 3 to 4 million years ago, they can mate and produce fertile offspring. Largely because of its geographic isolation in eastern and southern Africa, the African wild dog (also known as the Cape hunting dog) went its separate way about the same time. All of these canids have strong jaws and the relatively big teeth typical of carnivores, as Little Red Riding Hood discovered! Their legs are adapted for loping or trotting long distances, with the exception of the mutant domestic dog breeds and running for shorter periods with bursts of speed. As a general rule, they show a marked propensity toward pack or group behaviour. They also communicate vocally through a variety of calls, physical posturing and scent marking. Their olfactory abilities are superb, as is their hearing. They have excellent peripheral and night vision, as well as high sensitivity to light and movement. Dogs, wolves and perhaps other canids, see fairly well at a distance and discern colours, although not as acutely as humans.

    Observers have long argued that wolves and dogs possess some sort of extrasensory perception that allows them to sense the moods of humans or prey, to locate someone at a distance, to anticipate the arrival of a master, pack member or quarry, to discern when they are nearing their destination, even if riding in a closed car. Of particular fascination to a number of experts is “psi trailing,” the apparent ability of an animal to find its owners after they have left it and moved to a place it has never been before. ESP is, of course, a term human’s use for any psychic phenomenon beyond their explanation and so its use with canids is probably irrelevant. It is fairer to say that canids live in a perceptual universe far different from ours and that we are unaware of many of the olfactory and auditory signals they detect. Both dogs and wolves respond to higher frequencies than humans; wolves reportedly can hear sounds on the Alaska tundra from a distance up to ten miles.

    No one knows how many subspecies of Canis lupus have existed. Estimates range from twenty to forty. Part of the difficulty, as with defining breeds of dogs, is that wolves are highly variable in size, coloration and behaviour. In addition, heavy human predation has seriously diminished their numbers worldwide, making it difficult even to determine with accuracy what has been lost. Due primarily to heat and parasites, wolves tend to be smaller in southern than in northern latitudes, so that the little Arabian wolf and the red wolf are in the forty-five-pound range, while the Arctic grey wolf regularly exceeds one hundred pounds. The Arabian wolf seems to howl rarely and generally hunts alone or in small groups. Indeed, many of these subspecies have been studied little; more than a few cannot be examined at all, except in their remains. Thus, we will probably never know how the behaviour of specific wolves is reflected in the dogs derived from them millennia ago.

    Taming Wolf
    Fossil evidence from Zhoukoudian, China, shows Homo erectus pekinensis, the elusive Peking or Beijing man, was sharing time and space, food and shelter with wolves (generally classed as Canis lupus variabilis) at least 500,000 years ago. Remains of Homo erectus and wolves have also turned up in Boxgrove in Kent, England, dated to 400,000 years ago and Lazeret in the south of France, 150,000 years ago. It is more likely that throughout the Northern Hemisphere these precursors of modern humans and wolves lived and hunted in close proximity than that these three sites represent an accidental accumulation of old bones. Beyond that, we have only questions and surmise, especially since we know less about our prehistoric forebears than we do about wolves.

    Relatively short, with slightly smaller brains, flatter skulls, more prominent brow ridges and a noticeably more protruding jaw holding larger teeth than humans, these hominids were probably semi-nomadic hunter-gatherers who colonized much of the world. They had stone tools to help them butcher their kill for cooking. The fossils found at Zhoukoudian indicate that the brains of their compatriots – or competitors – made up at least part of their diet. In the main, however, early hominids were omnivores, deriving an estimated 60 to 80 percent of their calories and protein from nuts and vegetables.

    Even estimated dates are in dispute, however, so it seems fair to say that sometime around 200,000 years ago archaic humans, Homo sapiens, emerged in Africa. They possessed significantly larger brains than Homo erectus, whom they supplanted and made superior stone weapons with which they hunted big game. Whether Neanderthals, who emerged around 100,000 years ago and vanished 70,000 years later, were a separate human species or a stocky, heavy-browed, big-brained cousin of Homo sapiens – the way the dog is a subspecies of wolf – is not yet clear, but these powerful Ice Age hunters were also found throughout Europe, Asia and Africa. Around 40,000 to 50,000 years ago, slightly different beings arose, modern humans (Homo sapiens sapiens) with brains wired for invention and the drive to remake the world. More precisely, our ancestors showed up with a more highly developed and enlarged basal neocortex (believed to be involved in ethical and social behaviour, as well as formation of personality) than their predecessors.

    As humans colonized the world, some of them became – especially in the Arctic, Patagonia, the Great Plains of North America and steppes of Asia – predominately carnivorous in response to ecological conditions. (The polar bear, which evolved as a separate species 20,000 to 40,000 years ago, shows a similar adaptation, becoming the only solely carnivorous and semi-aquatic bear.) In the main, however, they moved in small bands of approximately twenty-five men, women, and children, taking most of their calories from plants and nuts.

    From my childhood in the 1950s and 1960s, I remember portrayals of early humans as people terrified of the world and its animals, living a marginal existence on the edge of death by starvation, exposure or assault. Since then, we have come increasingly to perceive those ancient hunters and gatherers as having had a rich culture and diet. They moved through their world as easily as we navigate ours, only in that world the boundary between human camps and nature was highly porous. Hunter-gatherers viewed animals as beings with their own habits, cultures and souls, making many of them totemic figures, the way we invest in material objects like cars and houses or famous humans with special status, not to mention God in his various guises. Early humans tamed nearly every animal they came into contact with and that impulse to collect animals has remained as strong as the related impulse to hunt them; in fact, it is no mistake that some of the most ardent conservationists have been hunters, which is not the same as saying that all hunters are conservationists – far too many of them are not.

    Even with weapons, hominids and early humans were not natural hunters and so they would have scavenged carnivores’ kills and also looked to them for guidance on how to bring down their own meat. They turned not to the bear, another omnivore, nor to the cats, but to animals – wolves and African wild dogs – that, like them, hunted in packs to bring down game much larger than themselves. Humans wanted those heavy animals for the same reasons wolves did: They provided enough meat to feed the group for days.

    Wolves and humans do not talk the same language – I assume, as do many enlightened naturalists, that all animals possess language, defined here as the ability to communicate through verbal or visual signs – but they understand each other to a remarkable degree. By the look on their faces, the tilt of their ears, position of their tails and bodies, wolves convey a great deal about their mood and intent that humans can interpret. Like humans, wolves possess associative minds and wanderlust. The social structure of their packs and their habits of nurturing and educating their young parallel those of human groups.

    People adopted wolf puppies that were orphaned or that they or their children lifted from dens during explorations. Women nursed the youngest of those puppies the way they suckled their own children. Not surprisingly, some of those hand-raised wolves hung around their adoptive family, becoming companions to the children or even the young men who played with them and learned to hunt with them. The tamed wolf took to the village as its home, alerting people to danger, the way it warned its own kind if a stranger approached the den – by barking. In some regions – for this process was occurring in many parts of the world – when food got scarce or if a spirit needed to be propitiated, people sacrificed and ate the wolf; if it proved a foul-tempered ingrate, it was driven off or killed.

    The wolves who became the tamest and lingered around the camps were those that were in personality the most social and least fearful. Mating with each other and free-ranging animals living near the camps, the tamed wolves produced, over time, a population with a high overall level of sociability, a group of fellow-travelling wolves. Under no breeding pressure from humans, allowed to come and go as they wished, they retained their wolfish look and demeanour.

    Becoming Dog
    Near the end of the Palaeolithic (Old Stone) Age, our direct forebears developed better, sharper stone blades, the atlatl for throwing spears and around 18,000 to 20,000 years ago, the boomerang (subsequently isolated in Australia) and the bow and arrow. These weapons allowed hunters to kill larger animals with greater ease from a longer range. Around the same time, in many parts of the world humans took up fishing and established semi-permanent villages with populations larger than their traditional bands, constructing their homes of the materials at hand: wood, earth, stone, skins, mammoth tusks. They developed better ways to carry water, food, firewood and pelts back to camp: baskets, ceramic pots, sledges, toboggans and travois. Boats extended the distances they could travel in search of food and in trade for furs, tools or ceramics. These humans also turned the tamed wolf into a dog, the first fully domesticated animal, meaning its evolution and breeding became directed more by humans than by nature.

    The circumstances in which our forebears found themselves changed dramatically – in part because of their activities – between the last glacial advance, which peaked around 18,000 years ago and the end of the Pleistocene some 8,000 years later. At their maximum, glaciers in eastern North America extended south over what are now the Middle Atlantic states and in the west covered Alaska, western Canada, Idaho, Washington and Montana. In Europe, Scandinavia, Denmark, most of Great Britain, Poland, Germany and Russia were under ice. Glaciers embraced the Alps and Dolomites, covering what are now Switzerland and sections of Austria, France and Italy. Bordering the ice sheets were dry steppes and grasslands supporting herds of animals, including mammoths, reindeer and giant bison. Among the predators hunting them were sabre-toothed tigers, scimitar cats, dire wolves, grey wolves and humans with their wolf dogs. In some areas, the wolf dogs resembled short-faced wolves, that is, they were barely distinguishable from dogs.

    As glaciers retreated, the earth warmed and sea levels rose, reconfiguring shorelines, flooding the Bering land bridge between the Americas and Eurasia. Established ecosystems collapsed while new ones emerged. Heavy rains turned solid land to marsh, lakes dried up, steppes and grasslands turned to forests and the great inland sea of North America, with its lush marshes, became a high desert, the Great Plains. As many as forty species of mammals vanished, especially of the huge predators and prey, among them mammoths, mastodons, great-horned bison, giant rhinoceroses, giant sloths, cave bears, dire wolves, all the sabre-toothed cats and the armadillo-like glyptodonts. Others, like the horse and camel, disappeared from North America, finding refuge in Eurasia or the Southern Hemisphere.

    To a degree we cannot yet determine that paleo-hunters contributed to the extinction of some of those large animals, like the mammoths, giant bison and rhinos, with hunting techniques that included driving them off cliffs or into bogs where they were slaughtered, baying them up with wolf dogs so they could be filled with arrows. In turn, their demise hastened that of the giant predators who fed on them. But many of those animals, especially the predators, also appear to have reached an evolutionary dead end, because they were unable to adapt to a world that had turned suddenly warmer and in some cases, to the loss of their preferred food. Their populations stressed, they were pushed over the brink by human activities, but we must not overestimate the force behind the shove. Humans with bows and arrows and atlatls, no matter how skilled, cannot drive a vibrant population to extinction, as we can see by observing how little impact the Plains Indians of North America had on the bison herds during the centuries they hunted them without horses and guns – and that is just one example. Even with those weapons, the bison endured until white commercial and sport hunters slaughtered them by the thousand for their skins. (Curiously, the Plains Indians do not seem to have used dogs in hunting bison, although they kept hundreds in their villages and donned wolf pelts while stalking their prey.)

    The animals that survived the turmoil at the end of the Pleistocene were the smaller, less specialized, more mobile ones: humans, grey wolves, lions, the smaller ungulates, downsized elephants, rhinos and horses. Their size left them better suited to the warmer, damper world emerging with the retreat of the ice.

    Disruptions caused by the changing climate and vanishing game fuelled the trend toward different settlement and dietary patterns. In some regions, groups of people realized that in the midden heaps and latrine areas of their camps, food plants they usually harvested from the wild were sprouting and flourishing. Combined with diminishing wild supplies, the bounty reinforced their inclination to return to the same campsites repeatedly to prolong their stays; humans, like other animals, being creatures of habit and territory.

    Coincident with these cultural developments, humans began deliberately breeding their wolf dogs. They culled those that were unsocial or overly timid, thereby increasing the likelihood that subsequent generations would be as easily socialized. In the process, they turned the wolf into a dog. The humans wanted a guaranteed supply of reliable animals; the wolf dogs wanted security and society.

    In many parts of Eurasia, North America and northern Africa, tamed wolves had proven themselves as hunting partners, but they became more difficult to obtain as people settled into permanent villages, were prone to moving off when they felt the call to mate and were maimed or killed in combat with large, fierce animals. At a time when hunters had to turn to other species, they needed, more than ever, to be guaranteed the assistance of an animal that excelled at scenting, tracking, and holding game or driving it into ambush. With their speed and agility, the dogs could handle anything from bears to birds, deer, elk, sheep, oxen and buffalo. They also could help guard the village against marauders.

    Because no one had many tame wolf dogs – the entire human population of the world at the time was probably around 10 million – efforts to breed them dramatically narrowed the gene pool. For reasons we do not yet understand, that constriction had the effect of releasing the phenotypic variability inherent in the wolf, creating smaller, larger, differently marked animals. Slight genetic mutations – those for lop ears or a particular coat, for example – could rapidly be fixed in a line of dogs and then passed on, allowing bands to develop distinctive animals they could easily differentiate from wolves, a necessity after domestication of sheep and goats.

    Although involving a biological process, creation of the dog was fundamentally a cultural act, like making tools, weapons and baskets. Bands in one region turned their captive wolves into dogs and then traded them, the way they bartered other goods or gave them as gifts during ceremonial exchanges. The knowledge of how to tame wolves was transmitted by people who were travelling. They also mated one of the dogs accompanying them to an animal in another village. Within a few generations, a general type of dog could have become well established and spread fairly widely.

    Dogs were valued precisely because they possessed the stellar abilities of the tame wolf, but were less inclined to go their own way. The dog was as adaptable as the wolf to different climates and it was versatile enough to fit a range of needs. In addition to hunting and serving as dinner, dogs sounded a warning when someone approached, helped keep the camp clean of garbage and their people warm. They were playmates for children, totem objects for adults, as were nearly all animals that figured prominently in people’s lives. They exhibited a talent for finding their way home no matter what the conditions, which made them in some societies valued guides for the dead to the next world and for helping people in times of need – pulling them from the water, protecting them from attack by other people or animals. Wounds they licked seemed to heal miraculously, a fact that finds expression to this day in the saying “as clean as a hound’s tooth.” They also would breed with tame wolves that were still brought into camp – a bonus. It is not surprising that people domesticated the wolf thousands of years before any other animal and that many of them, especially the hunter-gatherers, kept only dogs.

    For centuries, Americans and Europeans have underestimated the importance of the emergent dog as a food source, although that was probably one of its earliest functions. Many Native Americans ate puppies, considered the most delectable, on feast days or to honour special visitors and a number of traditionalists continue the practice. The Aztec and other people in South and Central American and the Caribbean also relied heavily on dog meat for their animal protein, frequently from animals that were castrated and fattened for the purpose. Throughout Asia and Oceania, the dog has remained a highly desirable meat, frequently the primary source of animal protein. During the 1988 Summer Olympics, the South Korean government requested butchers to move their dogs, who can sell for $200 apiece – the price of some hunting dog puppies in the United States – from display in their windows so as not to offend American and European sensibilities. On walks through New York’s Chinatown, I have seen dog carcasses hanging in the windows of butcher shops.

    Throughout Europe, prehistoric people appear to have eaten dog, although at some point their descendants stopped, taking it up again only when no other food was available. Even then, they often did so reluctantly. Travelling along the Columbia River to explore land the young United States had acquired from France as part of the Louisiana Purchase, Meriwether Lewis, William Clark and their men ate dogs provided by local Indians in the winter of 1806 to supplement their meagre rations. Clark wrote that after overcoming their cultural bias, many of them became “extremely [sic] fond of their flesh.” Lewis preferred it to venison or elk; although not personally “reconciled” to the taste, Clark admitted that he and the men were stronger and healthier for having lived on dogs than they had been for months. Other travellers filed similar reports.

    By 15,000 years ago, people around the world were raising dogs, with the centres of activity being northern Europe, including England, northern North America, especially the Arctic region, the Middle East, China, Japan and Siberia. Presently, the earliest fossil called a dog comes from Obercassel, Germany and dates to 14,000 years ago, the late Pleistocene or upper Palaeolithic. (Pleistocene refers to the geological age; Palaeolithic to the human culture.)

    Trying to piece together the puzzle of simultaneous domestication around the world, experts assigned certain broad types of dog to specific subspecies of wolf based on perceived morphological similarities and assumed areas of origin. It is a rough evolutionary tree that we hope will be refined as the tools of genetic analysis become more sophisticated.

    Canis lupus pallipes, the small Indian wolf, probably gave rise to the dingo and its kin: the Asian pariah dogs, the New Guinea singing dog and related Pacific Island dogs. It could also have contributed to a few of the Native American dogs. Despite exposure to other dogs, the pariah has bred true to its original dingo type for at least 5,000 years.

    Canis lupus arabs, the equally small and closely related Arabian or desert wolf – it and the Indian wolf are now sometimes considered the same subspecies – might have been progenitor of the sight hounds, the basenji and small-game hunters of southern Europe and a number of dogs indigenous to the Middle East, like the Canaan dog of Israel and pariahs who hang around villages as scavengers and guards. Many of these animals are similar to dingoes in size and appearance, leading some people to suggest that they might, in fact, have a common origin.

    Canis lupus chanco, the woolly Chinese wolf, is the possible source of the chow chow and assorted Asian toy breeds, as well as the mastiffs, believed to have originated in the Himalayas, whose bloodlines were ultimately joined by descendants of the European wolf. Although this association is the most speculative, Canis lupus hodophilax, the extinct little Japanese wolf, probably figured in the creation of dogs like the shikoku, kai, the shiba inu and other indigenous breeds.

    Canis lupus lupus, the European grey wolf, lies at the foundation of various herding, guard, and spitz-type dogs indigenous to Europe, as well as some of the terriers, believed to have originated in the British Isles. Along with the North American grey wolf, it is also progenitor to the Eskimo dogs and many Native American dogs, with an assist in some cases from animals crossing the Bering land bridge with migrating people.

    The one apparent exception to this rule of wolf origin, which nonetheless proves that domestication was a process occurring around the world, is the Falkland wolf (Dusicyon australis). In The Voyage of the Beagle (1839), Charles Darwin described Falkland wolves as so fearless and tame that they would invade campsites at night and steal meat from under the heads of sleeping shepherds and sealers. Taking advantage of that behaviour, the men would offer each visitor a piece of meat with one hand and knife it with the other. By the turn of the century, the little twenty to thirty-pound animals, which had fed on birds until the arrival of white men, were all dead. Within the past decade, however, Robert K. Wayne, an evolutionary biologist at the University of California at Los Angeles who has contributed greatly to understanding canid evolution, conducted mitochondrial DNA analyses of one of the few pelts still in existence and his results indicated that this extinct animal is most closely related to the coyote. Since coyotes, a North American native, could not have gotten to those remote islands by themselves, the findings lend support to a theory that the little canids were brought to the Falklands by humans some 6,000 years ago.

    © 1997 Mark Derr All rights reserved. ISBN: 0-8050-4063-3