Garden Scraps: Agonistic Interactions between Hedgehogs and Sympatric Mammals in Urban Gardens

Citation: Scott, D.M.; Fowler, R.; Sanglas, A.; Tolhurst, B.A. Garden Scraps: Agonistic Interactions between Hedgehogs and Sympatric Mammals in Urban Gardens. Animals 2023,13, 590. https://doi.org/ 10.3390/ani13040590 Academic Editors: Anne Berger, Nigel Reeve and Sophie Lund Rasmussen Received: 21 December 2022 Revised: 4 February 2023 Accepted: 6 February 2023 Published: 8 February 2023 Copyright: © 2023 by the authors. Licensee MDPI, Basel, Switzerland. This article is an open access article distributed under the terms and conditions of the Creative Commons Attribution (CC BY) license (https:// creativecommons.org/licenses/by/ 4.0/). animals Article Garden Scraps: Agonistic Interactions between Hedgehogs and Sympatric Mammals in Urban Gardens Dawn Millicent Scott 1,*, Robert Fowler 2, Ariadna Sanglas 3and Bryony Anne Tolhurst 4 1School of Animal Rural and Environment Sciences, Brackenhurst Campus, Nottingham Trent University, Southwell NG25 0QF, UK 2School of Life Sciences, John Maynard Smith Building, University of Sussex, Brighton BN1 9QG, UK 3Department of Conservation Biology, Estación Biológica Doñana, CSIC, Américo Vespucio, 26, 41092 Sevilla, Spain 4School of Applied Sciences, University of Brighton, Brighton BN2 4GJ, UK *Correspondence: [email protected] Simple Summary: Hedgehogs are one of several mammals that occur in urban areas in the United Kingdom and are fed by people. Food provided by people may help wild animals but may also attract animals together that could compete, injure, or predate each other. To understand the impact of food on urban animals we need to investigate how they interact when food is available. In this study, we assessed the type of interaction between hedgehogs, foxes, badgers, and cats using videos submitted by the public. We analyzed interactions between pairs of species to determine interaction type, hierarchical relationships, and the effect of food. We found that agonistic interactions (aggression and/or submission between animals) were more common than neutral interactions, and that between-species interactions showed greater ‘agonism’ than those within the same species. Of interactions within a species, those between hedgehogs were the most agonistic (54.9%) and between badgers the least (6.7%). The species interacting affected the level of agonism, with cats and foxes showing the highest level when together (76.7%). Badgers also outcompeted cats where there were contests over food, but cats were equally as successful as foxes, which were more successful than hedgehogs. However, hedgehogs dominated access to food over cats. We discuss the need to understand interactions between urban animals and the effects of providing food, to inform practice and ensure any potential risks are minimized. Abstract: Hedgehogs occur within an urban mammal guild in the United Kingdom. This guild commonly utilizes anthropogenic food provision, which is potentially beneficial to wild animal populations, but may also bring competitors and predators into proximity, raising the question of how these species interact in urban gardens. In this study, we determined interactions between hedgehogs, foxes, badgers, and domestic cats using videos submitted via citizen science. We analyzed interactions within and between species to determine interaction type, hierarchical relationships, and effect of supplementary food presence/amount. We found that overall agonistic interactions between individuals occurred more frequently (55.4%) than neutral interactions (44.6%) and that interspecific interactions showed greater agonism (55.4%) than intraspecific ones (36%). Within intraspecific interactions, those between hedgehogs were the most agonistic (54.9%) and between badgers the least (6.7%). Species composition of the interaction affected agonism, with interactions between cats and foxes showing the highest level (76.7%). In terms of overall “wins”, where access to garden resources was gained, badgers dominated cats, which were dominant or equal to foxes, which dominated hedgehogs. However, hedgehogs exhibited a greater overall proportion of wins (39.3%) relative to cats. Our findings are important in the context of the documented impact of patchy resources on urban wildlife behavior, and we show that provision of anthropogenic food can potentially result in unintended consequences. We recommend actions to reduce proximity of guild competitors in space and time to limit negative effects. Animals 2023,13, 590. https://doi.org/10.3390/ani13040590 https://www.mdpi.com/journal/animals Animals 2023,13, 590 2 of 14 Keywords: hedgehog; Erinaceus europaeus; red fox; Vulpes vulpes; Eurasian badger; Meles meles; supplementary feeding; urban mammals; domestic cat; Felis catus; citizen science 1. Introduction Increasing urbanization, where habitats have been highly modified for intense human use and residence, typically has a negative effect on biodiversity [ 1 ], yet a few species appear to benefit, as evidenced by their urban colonization, and high relative densities (e.g., [ 2 ]). These species are termed “synurbic” [ 3 ]. Although urban development can present challenges for wildlife, it can also offer benefits. For example, thermal climatic conditions are more stable in urban environments, and ground and air temperatures can be higher than in surrounding rural areas [ 4 ], while towns and cities provide numerous sheltering opportunities, and abundant supplementary food from anthropogenic sources [5,6]. Among the many reasons to colonize urban landscapes, food availability is perhaps one of the most important [7]. The intentional provision of supplementary food to wild animals by urban residents, either as surplus household food or commercially purchased for this purpose, has increased in the UK [ 8 ], where an estimated 87% of urban dwellers have access to a back or front garden/yard of their residence [ 9 ]. Supplementary feeding can benefit wild populations by providing food sources when natural food sources are low. However, it can also create clustered, abundant, spatio-temporally predictable food resources in urban areas [ 10 ], leading to behavioral modifications of wildlife, such as changes in foraging and spatial behavior [ 6 , 11 ], diet [ 12 ], and social/territorial configuration [ 11 , 13 ] with corresponding changes in density. In the United Kingdom (UK), synurbic mammal species include the Western European hedgehog (Erinaceus europaeus) (hereafter “hedgehog”), the red fox (Vulpes vulpes) (hereafter “fox”), and the Eurasian badger (Meles meles) (hereafter “badger”). These three species form a guild within which they compete for similar food resources; however, badgers and foxes can prey on hedgehogs, i.e., intraguild predation (IGP) also occurs [ 14 , 15 ]. The presence of both badgers and foxes negatively predicts UK hedgehog presence [ 16 , 17 ], but it is unclear whether this is due to predation, competition, or a combination of both. Modeling studies suggest that intraguild predation dynamics may additionally be confounded by the presence of supplementary food [18]. Higher densities relative toruralcounterparts, have beendocumented in hedgehogs [5,16], foxes [ 6 ], and to some extent badgers [ 19 ]. Factors affecting hedgehogs in the UK are of particular interest due to recent widespread population declines [ 20 – 23 ]. Within population studies, sub/urban habitats are considered potential refuges for hedgehogs [ 16 , 24 , 25 ], typically providing a range of ecologically favorable attributes, including high food availability and low overall populations of badgers, although high badger sett (burrow network) densities have been documented in some sub/urban areas [ 26 ]. This raises the question of how these species interact and co-occur in towns and cities. Exploitation of supplementarily provisioned food in urban gardens has been observed in all three species, and although this often manifests in individuals feeding alone, co-occurrence and aggregation of multiple individuals and species can also result [ 8 , 27 ]. In addition to synurbic wildlife, domestic cats (Felis catus) occur at high densities in urban areas [ 28 ] and utilize anthropogenic food sources, spatially overlapping with urban wildlife [ 29 ]. Therefore, cats can also be considered part of the urban mammalian guild when investigating species interactions. Most urban mammals, including cats, forage alone, despite some living in social or family groups, except for hedgehogs, which are considered solitary [ 30 ]. Hierarchy within social groups, such as those comprised of badgers or foxes, reduces competition and the risk of aggressive interactions [ 31 ]. Evidence of aggression between conspecifics is limited in hedgehogs except for in the context of mating [ 32 ]. A garden in which supplementary food is provided could be considered a ‘high-quality’ patch containing predictable resources Animals 2023,13, 590 3 of 14 in abundance. Individuals may compete to defend, or acquire, high-quality patches and engage in intraand interspecific agonistic behavior (fighting or conflict behavior, such as threatening, aggressive, and/or submissive behaviors) to dominate access for their own benefit [ 33 ]. Each dyadic encounter (paired encounter between individuals) represents the balance of the possible fitness costs and benefits from competing with the opponent/s for access to the resource [ 34 ] and will also reflect hierarchical positioning within groups and between competing species. Currently, there is a gap in our knowledge of how these species interact when coming into proximity at focal food resources. Aggregation of multiple individuals of several species in gardens attracted by food could potentially result in higher encounter rates between competitors and predators. Increasing interference competition, aggression, stress, and predation risk may result. There have been reported incidents of urban hedgehogs admitted to rehabilitation centers with injuries from suspected encounters with urban predators, including 2.3% of admissions attributed to injuries from dogs and cats [ 35 ]. Therefore, it is essential we understand the interactions between synurbic mammals, what factors drive interactions and access to food, and how coexistence in these habitats can be supported. The aims of the study were to determine if the type of interaction between sympatric urban mammal species varied depending on which species were interacting, which of the cooccurring species was most likely to ‘win’ dyadic competitions for access to supplementary food, and finally, if presence and amount of food affected the type of interaction. We hypothesized that there would be lower levels of agonism within species, especially those with established hierarchies, and greater agonism between species. We hypothesized that the largest of the species (badger) would dominate and be the most successful at ‘winning’ access to food, and that increased food availability would increase levels of agonistic interactions between and within species. Understanding interactions between synurbic mammals and the effect of supplementary feeding on interactions can help inform best practice around food provision to prevent unintended costs to the species concerned. 2. Materials and Methods 2.1. Data Collection and Cleaning Data on urban mammal interactions were obtained from the public following a national appeal for such video footage on a UK television broadcast (British Broadcasting Commission [BBC] “Springwatch” series) in May 2017. This program is part of a longstanding seasonal series on British wildlife that has approximately 2.5 million viewers. The broadcast explained the study aims and provided details of a link to a data capture site where videos and associated metadata could be submitted. The link was also available on the associated program website and the University of Brighton website. We did not provide instructions of an experimental set-up to follow, but instead called for people who had existing footage of multiple animals within their gardens to submit their existing footage. Once on the site, the public (volunteers) could upload their videos and answer a questionnaire on the location, video content, and food provision where the video was taken. The link was available between 29 May and 15 June 2017, during which time 683 files were submitted. Video analysis at feeding sites has previously been used as a method to determine relationships and interactions between sympatric species [36]. 2.2. Data Handling All data collected were downloaded to an Excel sheet with a linking unique ID code assigned to each video. Prior to analysis, data were removed that were not in video format or did not contain a hedgehog, fox, badger, or cat. We included videos where at least two adult individuals were visible, recorded in a UK residential garden between Jan 2010 and May 2017. Urban habitat was verified by using Google Maps for the postcode data submitted with the video. Multiple files from the same address/date were assumed to be consecutive recordings, so only one representative video was used. Of the 683 files Animals 2023,13, 590 4 of 14 submitted, 586 dyads (interactions between two individuals) [ 30 ] were extracted during behavioral analysis. 2.3. Behavioral Analysis Interactions between species were assessed per dyad. When interactions involved three or more individuals, multiple dyads were derived. For each dyadic interaction, the following were recorded from the video analysis: interaction number (for videos with more than two dyadic interactions); duration of video (in seconds); visible food present (yes or no); duration of the interaction (in seconds); species; predominant behavior of animal; interaction type (neutral or agonistic) [ 36 ]; and outcome for both individuals in the dyad (“win” or “no win”) [ 31 ]. The amount of food left out (“high” or “low”, where “high” was two handfuls or more and “low” was a single handful or less) was also included from the questionnaire response. An ethogram was compiled containing detailed descriptions of six typical dyadic behaviors [ 37 ] during encounters (passive, submissive, avoidance, defensive, aggressive, attack; see Table 1). As the study focused on multiple species, some definitions included reference to a particular species. During the analysis, a predominant behavior was assigned to each member of the dyad and an interaction type then chosen to summarize the encounter. Behaviors that appeared to have no impact on either animal were deemed neutral, i.e., animals were passive towards each other, there was no defensive or aggressive behavioral change in the presence of another, and/or there was no observed agonistic behavior. Conversely, behaviors involving submission, threat, aggression, defense, or attack were classified as agonistic [ 36 ]. Dyadic outcomes for each animal were classified as either “win” (if one animal was seen eating the food or dominating the space close to the food during the video clip), “draw” if the two animals continued feeding or stayed in the garden together, or “unclear” if the video or observation ended before an outcome could be determined. A loss, draw, or unclear outcome was classified as “no win” for the individual in the dyad. 2.4. Data Analysis To test the effect of dyad composition (i.e., species), food presence, and amount on interaction type and probability of wins, Generalized Linear Models (GLMs) were computed with binomial error structures in R Studio (RStudio, 2012) using R v3.6.1 [ 38 ] and packages MASS and lme4. Interand intraspecific dyads were modeled separately. For interaction type, both models contained dyad composition as the independent variable and interaction type (Neutral = 0, Agonistic = 1) as the dependent variable. For the intraspecific model, the highest neutral interaction dyad (badger-badger) was used as a reference in the model. For probability of wins, the independent variables comprised interaction type (Neutral = 0, Agonistic = 1), whether supplementary food was provided (No = 0, Yes = 1), and the amount of food (Low = 0, High = 1) and dyad composition (all the interspecific dyads that contain at least one of the target species for that specific model). The hedgehog-cat dyad was used as a reference for the interspecific model, as the highest neutral interaction dyad that was not deemed a potentially predatory interaction. The dependent variable in each case was win (eats the food = 1) or no win (lose, draw/shares food or is unclear = 0) for each test species. Finally, the reference level was switched for each dyad comparison (using the relevel function) so that all pairwise dyad comparisons were tested against each other. Prior to applying models, proposed explanatory variables were checked for multicollinearity using Variance Inflation Factors (VIFs). If variables had VIFs greater than 3 or correlation coefficients more than 0.6 with other variables, they were excluded from models [ 39 ]. Model residuals were assessed for normality and heteroscedasticity. Omnidirectional stepwise selection was undertaken to build the best model in each case using the step function with the command direction = both. This procedure sequentially keeps or drops variables starting with the null (intercept only) model through testing the significance of each independent variable in a linear regression model. Animals 2023,13, 590 5 of 14 Table 1. Behavior categories and types of interaction based on analysis of encounters between four species of urban mammal—Western European Hedgehog (Erinaceus europaeus), Eurasian badger (Meles meles), Red fox (Vulpes vulpes), and domestic cat (Felis catus). Symbols represent each behavior and how they relate to the two broad interaction types. Animal Behaviour Interaction Type Passive Animals 2023, 13, x 5 of 15 Table 1. Behavior categories and types of interaction based on analysis of encounters between four species of urban mammal—Western European Hedgehog (Erinaceus europaeus), Eurasian badger (Meles meles), Red fox (Vulpes vulpes), and domestic cat (Felis catus). Symbols represent each behavior and how they relate to the two broad interaction types. Animal behaviou r Interaction type Passive Animal continues feeding or remains in the area when approached by the other individual. Departs without retaliation if aggression occurs. Neutral ▬ Behaviour of both animals has no impact on the other, e.g. ignore each other. Submissive An act or posture that does not challenge the incoming animal. Body position or response indicting lower hierarchy or submission, e.g. in foxes the body and head lower. In cats crouching with ears flattened, avoiding, retreating or fleeing. Avoid ▬ Animal moves or backs away from the area or other animal b efore close proximity or physical contact. Often with body held low, casting repeated glances at the one which stays. Defensive █ A defensive posture or positioning e.g. piloerection, frowning or complete rolling in hedgehogs. In cats, hissing or piloerection. Agonistic █ ► ▬ Includes submissive, threat, attack and aggression behaviour. Could result in injury, or death, of at least one individual. Aggressive ► Vocalisation or aggressive posture. Action of initiating physical contact with another animal including lunging, b iting, scratching etc. Foxes side profile body position with arched back and head up. Attack One animals runs towards the other in an aggressive manner. Chases, lunges, bites etc. In badgers, head is lowered in a threat posture and pursues the challenger with physical contact. Cat striking with paw. 2.4. Data Analysis To test the effect of dyad composition (i.e., species), food presence, and amount on interaction type and probability of wins, Generalized Linear Models (GLMs) were computed with binomial error structures in R Studio (RStudio, 2012) using R v3.6.1 [38] and packages MASS and lme4. Interand intraspecific dyads were modeled separately. For interaction type, both models contained dyad composition as the independent variable and interaction type (Neutral = 0, Agonistic = 1) as the dependent variable. For the intraspecific model, the highest neutral interaction dyad (badger-badger) was used as a reference in the model. For probability of wins, the independent variables comprised interaction type (Neutral = 0, Agonistic = 1), whether supplementary food was provided (No = 0, Yes = 1), and the amount of food (Low = 0, High = 1) and dyad composition (all the interspecific dyads that contain at least one of the target species for that specific model). The hedgehog-cat dyad was used as a reference for the interspecific model, as the highest neutral interaction dyad that was not deemed a potentially predatory interaction. The dependent variable in each case was win (eats the food = 1) or no win (lose, draw/shares food or is unclear = 0) for each test species. Finally, the reference level was switched for each dyad comparison (using the relevel function) so that all pairwise dyad comparisons were tested against each other. Prior to applying models, proposed explanatory variables were checked for multicollinearity using Variance Inflation Factors (VIFs). If variables had VIFs greater than 3 or correlation coefficients more than 0.6 with other variables, they were excluded from models [39]. Model residuals were assessed for normality and heteroscedasticity. Omnidirectional stepwise selection was undertaken to build the best model in each case using the Animal continues feeding or remains in the area when approached by the other individual. Departs without retaliation if aggression occurs. Neutral Animals 2023, 13, x 5 of 15 Table 1. Behavior categories and types of interaction based on analysis of encounters between four species of urban mammal—Western European Hedgehog (Erinaceus europaeus), Eurasian badger (Meles meles), Red fox (Vulpes vulpes), and domestic cat (Felis catus). Symbols represent each behavior and how they relate to the two broad interaction types. Animal behaviou r Interaction type Passive Animal continues feeding or remains in the area when approached by the other individual. Departs without retaliation if aggression occurs. Neutral ▬ Behaviour of both animals has no impact on the other, e.g. ignore each other. Submissive An act or posture that does not challenge the incoming animal. Body position or response indicting lower hierarchy or submission, e.g. in foxes the body and head lower. In cats crouching with ears flattened, avoiding, retreating or fleeing. Avoid ▬ Animal moves or backs away from the area or other animal b efore close proximity or physical contact. Often with body held low, casting repeated glances at the one which stays. Defensive █ A defensive posture or positioning e.g. piloerection, frowning or complete rolling in hedgehogs. In cats, hissing or piloerection. Agonistic █ ► ▬ Includes submissive, threat, attack and aggression behaviour. Could result in injury, or death, of at least one individual. Aggressive ► Vocalisation or aggressive posture. Action of initiating physical contact with another animal including lunging, b iting, scratching etc. Foxes side profile body position with arched back and head up. Attack One animals runs towards the other in an aggressive manner. Chases, lunges, bites etc. In badgers, head is lowered in a threat posture and pursues the challenger with physical contact. Cat striking with paw. 2.4. Data Analysis To test the effect of dyad composition (i.e., species), food presence, and amount on interaction type and probability of wins, Generalized Linear Models (GLMs) were computed with binomial error structures in R Studio (RStudio, 2012) using R v3.6.1 [38] and packages MASS and lme4. Interand intraspecific dyads were modeled separately. For interaction type, both models contained dyad composition as the independent variable and interaction type (Neutral = 0, Agonistic = 1) as the dependent variable. For the intraspecific model, the highest neutral interaction dyad (badger-badger) was used as a reference in the model. For probability of wins, the independent variables comprised interaction type (Neutral = 0, Agonistic = 1), whether supplementary food was provided (No = 0, Yes = 1), and the amount of food (Low = 0, High = 1) and dyad composition (all the interspecific dyads that contain at least one of the target species for that specific model). The hedgehog-cat dyad was used as a reference for the interspecific model, as the highest neutral interaction dyad that was not deemed a potentially predatory interaction. The dependent variable in each case was win (eats the food = 1) or no win (lose, draw/shares food or is unclear = 0) for each test species. Finally, the reference level was switched for each dyad comparison (using the relevel function) so that all pairwise dyad comparisons were tested against each other. Prior to applying models, proposed explanatory variables were checked for multicollinearity using Variance Inflation Factors (VIFs). If variables had VIFs greater than 3 or correlation coefficients more than 0.6 with other variables, they were excluded from models [39]. Model residuals were assessed for normality and heteroscedasticity. Omnidirectional stepwise selection was undertaken to build the best model in each case using the Animals 2023, 13, x 5 of 15 Table 1. Behavior categories and types of interaction based on analysis of encounters between four species of urban mammal—Western European Hedgehog (Erinaceus europaeus), Eurasian badger (Meles meles), Red fox (Vulpes vulpes), and domestic cat (Felis catus). Symbols represent each behavior and how they relate to the two broad interaction types. Animal behaviou r Interaction type Passive Animal continues feeding or remains in the area when approached by the other individual. Departs without retaliation if aggression occurs. Neutral ▬ Behaviour of both animals has no impact on the other, e.g. ignore each other. Submissive An act or posture that does not challenge the incoming animal. Body position or response indicting lower hierarchy or submission, e.g. in foxes the body and head lower. In cats crouching with ears flattened, avoiding, retreating or fleeing. Avoid ▬ Animal moves or backs away from the area or other animal b efore close proximity or physical contact. Often with body held low, casting repeated glances at the one which stays. Defensive █ A defensive posture or positioning e.g. piloerection, frowning or complete rolling in hedgehogs. In cats, hissing or piloerection. Agonistic █ ► ▬ Includes submissive, threat, attack and aggression behaviour. Could result in injury, or death, of at least one individual. Aggressive ► Vocalisation or aggressive posture. Action of initiating physical contact with another animal including lunging, b iting, scratching etc. Foxes side profile body position with arched back and head up. Attack One animals runs towards the other in an aggressive manner. Chases, lunges, bites etc. In badgers, head is lowered in a threat posture and pursues the challenger with physical contact. Cat striking with paw. 2.4. Data Analysis To test the effect of dyad composition (i.e., species), food presence, and amount on interaction type and probability of wins, Generalized Linear Models (GLMs) were computed with binomial error structures in R Studio (RStudio, 2012) using R v3.6.1 [38] and packages MASS and lme4. Interand intraspecific dyads were modeled separately. For interaction type, both models contained dyad composition as the independent variable and interaction type (Neutral = 0, Agonistic = 1) as the dependent variable. For the intraspecific model, the highest neutral interaction dyad (badger-badger) was used as a reference in the model. For probability of wins, the independent variables comprised interaction type (Neutral = 0, Agonistic = 1), whether supplementary food was provided (No = 0, Yes = 1), and the amount of food (Low = 0, High = 1) and dyad composition (all the interspecific dyads that contain at least one of the target species for that specific model). The hedgehog-cat dyad was used as a reference for the interspecific model, as the highest neutral interaction dyad that was not deemed a potentially predatory interaction. The dependent variable in each case was win (eats the food = 1) or no win (lose, draw/shares food or is unclear = 0) for each test species. Finally, the reference level was switched for each dyad comparison (using the relevel function) so that all pairwise dyad comparisons were tested against each other. Prior to applying models, proposed explanatory variables were checked for multicollinearity using Variance Inflation Factors (VIFs). If variables had VIFs greater than 3 or correlation coefficients more than 0.6 with other variables, they were excluded from models [39]. Model residuals were assessed for normality and heteroscedasticity. Omnidirectional stepwise selection was undertaken to build the best model in each case using the  Behaviour of both animals has no impact on the other, e.g. ignore each other. Submissive Animals 2023, 13, x 5 of 15 Table 1. Behavior categories and types of interaction based on analysis of encounters between four species of urban mammal—Western European Hedgehog (Erinaceus europaeus), Eurasian badger (Meles meles), Red fox (Vulpes vulpes), and domestic cat (Felis catus). Symbols represent each behavior and how they relate to the two broad interaction types. Animal behaviou r Interaction type Passive Animal continues feeding or remains in the area when approached by the other individual. Departs without retaliation if aggression occurs. Neutral ▬ Behaviour of both animals has no impact on the other, e.g. ignore each other. Submissive An act or posture that does not challenge the incoming animal. Body position or response indicting lower hierarchy or submission, e.g. in foxes the body and head lower. In cats crouching with ears flattened, avoiding, retreating or fleeing. Avoid ▬ Animal moves or backs away from the area or other animal b efore close proximity or physical contact. Often with body held low, casting repeated glances at the one which stays. Defensive █ A defensive posture or positioning e.g. piloerection, frowning or complete rolling in hedgehogs. In cats, hissing or piloerection. Agonistic █ ► ▬ Includes submissive, threat, attack and aggression behaviour. Could result in injury, or death, of at least one individual. Aggressive ► Vocalisation or aggressive posture. Action of initiating physical contact with another animal including lunging, b iting, scratching etc. Foxes side profile body position with arched back and head up. Attack One animals runs towards the other in an aggressive manner. Chases, lunges, bites etc. In badgers, head is lowered in a threat posture and pursues the challenger with physical contact. Cat striking with paw. 2.4. Data Analysis To test the effect of dyad composition (i.e., species), food presence, and amount on interaction type and probability of wins, Generalized Linear Models (GLMs) were computed with binomial error structures in R Studio (RStudio, 2012) using R v3.6.1 [38] and packages MASS and lme4. Interand intraspecific dyads were modeled separately. For interaction type, both models contained dyad composition as the independent variable and interaction type (Neutral = 0, Agonistic = 1) as the dependent variable. For the intraspecific model, the highest neutral interaction dyad (badger-badger) was used as a reference in the model. For probability of wins, the independent variables comprised interaction type (Neutral = 0, Agonistic = 1), whether supplementary food was provided (No = 0, Yes = 1), and the amount of food (Low = 0, High = 1) and dyad composition (all the interspecific dyads that contain at least one of the target species for that specific model). The hedgehog-cat dyad was used as a reference for the interspecific model, as the highest neutral interaction dyad that was not deemed a potentially predatory interaction. The dependent variable in each case was win (eats the food = 1) or no win (lose, draw/shares food or is unclear = 0) for each test species. Finally, the reference level was switched for each dyad comparison (using the relevel function) so that all pairwise dyad comparisons were tested against each other. Prior to applying models, proposed explanatory variables were checked for multicollinearity using Variance Inflation Factors (VIFs). If variables had VIFs greater than 3 or correlation coefficients more than 0.6 with other variables, they were excluded from models [39]. Model residuals were assessed for normality and heteroscedasticity. Omnidirectional stepwise selection was undertaken to build the best model in each case using the An act or posture that does not challenge the incoming animal. Body position or response indicting lower hierarchy or submission, e.g. in foxes the body and head lower. In cats crouching with ears flattened, avoiding, retreating or fleeing. Avoid  Animal moves or backs away from the area or other animal before close proximity or physical contact. Often with body held low, casting repeated glances at the one which stays. Defensive z A defensive posture or positioning e.g. piloerection, frowning or complete rolling in hedgehogs. In cats, hissing or piloerection. Agonistic z · Animals 2023, 13, x 5 of 15 Table 1. Behavior categories and types of interaction based on analysis of encounters between four species of urban mammal—Western European Hedgehog (Erinaceus europaeus), Eurasian badger (Meles meles), Red fox (Vulpes vulpes), and domestic cat (Felis catus). Symbols represent each behavior and how they relate to the two broad interaction types. Animal behaviou r Interaction type Passive Animal continues feeding or remains in the area when approached by the other individual. Departs without retaliation if aggression occurs. Neutral ▬ Behaviour of both animals has no impact on the other, e.g. ignore each other. Submissive An act or posture that does not challenge the incoming animal. Body position or response indicting lower hierarchy or submission, e.g. in foxes the body and head lower. In cats crouching with ears flattened, avoiding, retreating or fleeing. Avoid ▬ Animal moves or backs away from the area or other animal b efore close proximity or physical contact. Often with body held low, casting repeated glances at the one which stays. Defensive █ A defensive posture or positioning e.g. piloerection, frowning or complete rolling in hedgehogs. In cats, hissing or piloerection. Agonistic █ ► ▬ Includes submissive, threat, attack and aggression behaviour. Could result in injury, or death, of at least one individual. Aggressive ► Vocalisation or aggressive posture. Action of initiating physical contact with another animal including lunging, b iting, scratching etc. Foxes side profile body position with arched back and head up. Attack One animals runs towards the other in an aggressive manner. Chases, lunges, bites etc. In badgers, head is lowered in a threat posture and pursues the challenger with physical contact. Cat striking with paw. 2.4. Data Analysis To test the effect of dyad composition (i.e., species), food presence, and amount on interaction type and probability of wins, Generalized Linear Models (GLMs) were computed with binomial error structures in R Studio (RStudio, 2012) using R v3.6.1 [38] and packages MASS and lme4. Interand intraspecific dyads were modeled separately. For interaction type, both models contained dyad composition as the independent variable and interaction type (Neutral = 0, Agonistic = 1) as the dependent variable. For the intraspecific model, the highest neutral interaction dyad (badger-badger) was used as a reference in the model. For probability of wins, the independent variables comprised interaction type (Neutral = 0, Agonistic = 1), whether supplementary food was provided (No = 0, Yes = 1), and the amount of food (Low = 0, High = 1) and dyad composition (all the interspecific dyads that contain at least one of the target species for that specific model). The hedgehog-cat dyad was used as a reference for the interspecific model, as the highest neutral interaction dyad that was not deemed a potentially predatory interaction. The dependent variable in each case was win (eats the food = 1) or no win (lose, draw/shares food or is unclear = 0) for each test species. Finally, the reference level was switched for each dyad comparison (using the relevel function) so that all pairwise dyad comparisons were tested against each other. Prior to applying models, proposed explanatory variables were checked for multicollinearity using Variance Inflation Factors (VIFs). If variables had VIFs greater than 3 or correlation coefficients more than 0.6 with other variables, they were excluded from models [39]. Model residuals were assessed for normality and heteroscedasticity. Omnidirectional stepwise selection was undertaken to build the best model in each case using the Animals 2023, 13, x 5 of 15 Table 1. Behavior categories and types of interaction based on analysis of encounters between four species of urban mammal—Western European Hedgehog (Erinaceus europaeus), Eurasian badger (Meles meles), Red fox (Vulpes vulpes), and domestic cat (Felis catus). Symbols represent each behavior and how they relate to the two broad interaction types. Animal behaviou r Interaction type Passive Animal continues feeding or remains in the area when approached by the other individual. Departs without retaliation if aggression occurs. Neutral ▬ Behaviour of both animals has no impact on the other, e.g. ignore each other. Submissive An act or posture that does not challenge the incoming animal. Body position or response indicting lower hierarchy or submission, e.g. in foxes the body and head lower. In cats crouching with ears flattened, avoiding, retreating or fleeing. Avoid ▬ Animal moves or backs away from the area or other animal b efore close proximity or physical contact. Often with body held low, casting repeated glances at the one which stays. Defensive █ A defensive posture or positioning e.g. piloerection, frowning or complete rolling in hedgehogs. In cats, hissing or piloerection. Agonistic █ ► ▬ Includes submissive, threat, attack and aggression behaviour. Could result in injury, or death, of at least one individual. Aggressive ► Vocalisation or aggressive posture. Action of initiating physical contact with another animal including lunging, b iting, scratching etc. Foxes side profile body position with arched back and head up. Attack One animals runs towards the other in an aggressive manner. Chases, lunges, bites etc. In badgers, head is lowered in a threat posture and pursues the challenger with physical contact. Cat striking with paw. 2.4. Data Analysis To test the effect of dyad composition (i.e., species), food presence, and amount on interaction type and probability of wins, Generalized Linear Models (GLMs) were computed with binomial error structures in R Studio (RStudio, 2012) using R v3.6.1 [38] and packages MASS and lme4. Interand intraspecific dyads were modeled separately. For interaction type, both models contained dyad composition as the independent variable and interaction type (Neutral = 0, Agonistic = 1) as the dependent variable. For the intraspecific model, the highest neutral interaction dyad (badger-badger) was used as a reference in the model. For probability of wins, the independent variables comprised interaction type (Neutral = 0, Agonistic = 1), whether supplementary food was provided (No = 0, Yes = 1), and the amount of food (Low = 0, High = 1) and dyad composition (all the interspecific dyads that contain at least one of the target species for that specific model). The hedgehog-cat dyad was used as a reference for the interspecific model, as the highest neutral interaction dyad that was not deemed a potentially predatory interaction. The dependent variable in each case was win (eats the food = 1) or no win (lose, draw/shares food or is unclear = 0) for each test species. Finally, the reference level was switched for each dyad comparison (using the relevel function) so that all pairwise dyad comparisons were tested against each other. Prior to applying models, proposed explanatory variables were checked for multicollinearity using Variance Inflation Factors (VIFs). If variables had VIFs greater than 3 or correlation coefficients more than 0.6 with other variables, they were excluded from models [39]. Model residuals were assessed for normality and heteroscedasticity. Omnidirectional stepwise selection was undertaken to build the best model in each case using the  Includes submissive, threat, attack and aggression behaviour. Could result in injury, or death, of at least one individual. Aggressive · Vocalisation or aggressive posture. Action of initiating physical contact with another animal including lunging, biting, scratching etc. Foxes side profile body position with arched back and head up. Attack Animals 2023, 13, x 5 of 15 Table 1. Behavior categories and types of interaction based on analysis of encounters between four species of urban mammal—Western European Hedgehog (Erinaceus europaeus), Eurasian badger (Meles meles), Red fox (Vulpes vulpes), and domestic cat (Felis catus). Symbols represent each behavior and how they relate to the two broad interaction types. Animal behaviou r Interaction type Passive Animal continues feeding or remains in the area when approached by the other individual. Departs without retaliation if aggression occurs. Neutral ▬ Behaviour of both animals has no impact on the other, e.g. ignore each other. Submissive An act or posture that does not challenge the incoming animal. Body position or response indicting lower hierarchy or submission, e.g. in foxes the body and head lower. In cats crouching with ears flattened, avoiding, retreating or fleeing. Avoid ▬ Animal moves or backs away from the area or other animal b efore close proximity or physical contact. Often with body held low, casting repeated glances at the one which stays. Defensive █ A defensive posture or positioning e.g. piloerection, frowning or complete rolling in hedgehogs. In cats, hissing or piloerection. Agonistic █ ► ▬ Includes submissive, threat, attack and aggression behaviour. Could result in injury, or death, of at least one individual. Aggressive ► Vocalisation or aggressive posture. Action of initiating physical contact with another animal including lunging, b iting, scratching etc. Foxes side profile body position with arched back and head up. Attack One animals runs towards the other in an aggressive manner. Chases, lunges, bites etc. In badgers, head is lowered in a threat posture and pursues the challenger with physical contact. Cat striking with paw. 2.4. Data Analysis To test the effect of dyad composition (i.e., species), food presence, and amount on interaction type and probability of wins, Generalized Linear Models (GLMs) were computed with binomial error structures in R Studio (RStudio, 2012) using R v3.6.1 [38] and packages MASS and lme4. Interand intraspecific dyads were modeled separately. For interaction type, both models contained dyad composition as the independent variable and interaction type (Neutral = 0, Agonistic = 1) as the dependent variable. For the intraspecific model, the highest neutral interaction dyad (badger-badger) was used as a reference in the model. For probability of wins, the independent variables comprised interaction type (Neutral = 0, Agonistic = 1), whether supplementary food was provided (No = 0, Yes = 1), and the amount of food (Low = 0, High = 1) and dyad composition (all the interspecific dyads that contain at least one of the target species for that specific model). The hedgehog-cat dyad was used as a reference for the interspecific model, as the highest neutral interaction dyad that was not deemed a potentially predatory interaction. The dependent variable in each case was win (eats the food = 1) or no win (lose, draw/shares food or is unclear = 0) for each test species. Finally, the reference level was switched for each dyad comparison (using the relevel function) so that all pairwise dyad comparisons were tested against each other. Prior to applying models, proposed explanatory variables were checked for multicollinearity using Variance Inflation Factors (VIFs). If variables had VIFs greater than 3 or correlation coefficients more than 0.6 with other variables, they were excluded from models [39]. Model residuals were assessed for normality and heteroscedasticity. Omnidirectional stepwise selection was undertaken to build the best model in each case using the One animals runs towards the other in an aggressive manner. Chases, lunges, bites etc. In badgers, head is lowered in a threat posture and pursues the challenger with physical contact. Cat striking with paw. 3. Results A total of 683 files were received from volunteers. The files collected dated from May 2010 to June 2017. Using the filtering method stated in the methods section 586 separate dyads were analyzed, representing both intraand interspecific interactions. Two datasets were created with these data; the ‘all dyads’ dataset, which included all 586 separate dyads, and the ‘outcome’ dataset, which consisted of 331 interspecific dyads with wins, draws, and unclear outcomes recorded, alongside data on food presence and amount. 3.1. Interactions between Sympatric Species Badger-badger dyads had the lowest level of agonistic interactions and were recorded to be neutral in 93.3% of interactions (Table 2). In comparison, fox-fox and hedgehoghedgehog dyads exhibited neutral behavior in 63.6% and 45.1% of interactions, respectively. Hedgehog-hedgehog dyads had the highest number of intraspecific agonistic interactions (54.9%). The most frequent interspecific dyad was fox-hedgehog, with 143 separate instances of this dyad recorded (Table 2). The split between agonistic and neutral interactions for fox-hedgehog dyads was relatively even at 49% and 51%, respectively. Interspecific interactions between cats and foxes had the highest level of agonistic interactions (76.7%). Badger-cat dyads were the least recorded, being observed only eight times, but also showed a high proportion of agonistic interactions (75%). In every dyad containing a cat, the proportion of agonistic interactions was greater than neutral; however, the incidence of these was relatively low compared to other dyads (Table 2). Hedgehogs displayed higher agonistic interactions with foxes than badgers, although these comprised <50% of interactions. Animals 2023,13, 590 6 of 14 Table 2. The number of events recorded from videos sent in by volunteers for each dyad of four species of urban mammal (Western European Hedgehog (Erinaceus europaeus), Eurasian badger (Meles meles) , Red fox (Vulpes vulpes), and domestic cat (Felis catus)) and the number and relative percentage of agonistic or neutral interactions. Dyad No. of Events No. of Agonistic Interactions N (%) No. of Neutral Interactions N (%) Intraspecific Dyads Hedgehog-Hedgehog 142 78 (54.9) 64 (45.1) Badger-Badger 95 7 (6.7) 88 (93.3) Fox-Fox 33 12 (36.4) 21 (63.6) Total 270 97 (36.0) 173 (64.0) Interspecific Dyads Fox-Hedgehog 143 70 (49.0) 73 (51.0) Badger-Hedgehog 16 5 (31.3) 11 (68.7) Cat-Hedgehog 28 16 (57.1) 12 (42.9) Badger-Fox 78 45 (57.7) 33 (42.3) Fox-Cat 43 33 (76.7) 10 (23.3) Badger-Cat 8 6 (75.0) 2 (25.0) Total 316 175 (55.4) 141 (44.6) 3.2. Comparisons between Species Dyads Statistical models revealed a greater level of agonistic behavior in all other intraand interspecific dyads compared to badger-only dyads (Table 3). Statistically, only cat-fox dyads had higher agonistic interactions than hedgehog-hedgehog dyads. Cat-fox and badger-fox showed significantly higher agonistic interactions than fox-fox. Both badgerhedgehog and badger-fox agonistic interactions were greater than badger-badger but less than cat-fox. Badger-fox agonism was less than cat-fox but greater than fox-fox. Table 3. Generalized Linear Models (GLM) of the test results investigating the effects of intraspecific and interspecific dyadic composition on interaction type (Neutral = 0; Agonistic = 1), using the relevel command in R to switch the levels to investigate all dyad comparisons against each other. Displayed are parameter estimates ± standard error, z-value and p-value for each pairwise comparison. ‘<’ agonistic interactions are greater than the reference dyad and ‘>’ are less than the reference dyad. Reference Dyad Explanatory Variables Parameter Estimate ±SE z-Value p-Value Badger-Badger <Cat-Hedgehog 2.819 ±0.548 5.147 <0.001 <Cat-Fox 3.725 ±0.533 6.984 <0.001 <Hedgehog-Hedgehog 2.729 ±0.427 6.386 <0.001 <Badger-Cat 3.63 ±0.906 4.007 <0.001 <Badger-Fox 2.845 ±0.455 6.249 <0.001 <Badger-Hedgehog 1.743 ±0.668 2.612 <0.01 <Fox-Fox 1.972 ±0.534 3.692 <0.001 <Fox-Hedgehog 2.490 ±0.427 5.832 <0.001 HedgehogHedgehog <Cat-Fox 0.996 ±0.398 2.5 <0.05 Fox-Fox <Cat-Fox 1.754 ±0.511 3.431 <0.001 <Badger-Fox 0.870 ±0.428 2.031 <0.05 BadgerHedgehog <Cat-Fox 1.982 ±0.649 3.054 <0.01 Fox-Hedgehog <Cat-Fox 1.236 ±0.398 3.106 <0.01 >Badger-Badger −2.490 ±0.427 −5.832 <0.001 Badger-Fox <Cat-Fox 0.884 ±0.428 2.067 <0.05 >Fox-Fox −0.870 ±0.428 −2.031 <0.05 Cat-Fox >Hedgehog-Hedgehog −0.996 ±0.398 −2.5 <0.05 >Badger-Badger −3.725 ±0.533 −6.984 <0.001 Animals 2023,13, 590 7 of 14 Table 3. Cont. Reference Dyad Explanatory Variables Parameter Estimate ±SE z-Value p-Value >Badger-Fox −0.884 ±0.428 −2.067 <0.05 >Badger-Hedgehog −1.982 ±0.649 −3.054 <0.01 >Fox-Fox −1.754 ±0.511 −3.431 <0.001 >Fox-Hedgehog −1.236 ±0.398 −3.106 <0.01 3.3. Species Dominance and Hierarchy In terms of overall percentage of wins (access to garden resources), the order of dominance was firstly badgers, then cats, and then foxes. Foxes dominated hedgehogs, but hedgehogs dominated cats; hence, the hierarchy was non-linear (Table 4; Figure 1 for diagrammatical representation). However, there were a large proportion of draws and unclears in these dyads, so it is not always apparent which species won in each combination. Badgers tended to be more successful at ‘winning’ interactions than the other species, winning 45.9%, 42.1%, and 66.6% of interactions against foxes, hedgehogs, and cats, respectively. Foxes had the largest proportion of wins of any species when the dyad was with hedgehogs (46.2%). While cats won 44.4% of their dyads with foxes, foxes won 31.1% of these dyads (Table 4). Unexpectedly, hedgehogs won 39.3% of the time in dyads with cats, compared to cats winning only 10.7% of these dyads. Table 4. The number of events recorded from videos sent in by volunteers between each dyad of four species of urban mammal (Western European Hedgehog (Erinaceus europaeus), Eurasian badger (Meles meles), Red fox (Vulpes vulpes), and domestic cat (Felis catus)), and the number and relative percentage (%) of ‘wins’ for each species within the dyad. Win Draw Unclear Dyad No. of Events Hedgehog Badger Fox Cat Fox-Hedgehog 145 14 (9.6) 67 (46.2) 41 (28.3) 23 (15.9) Badger-Hedgehog 19 2 (10.5) 8 (42.1) 7 (36.8) 2 (10.5) Hedgehog-Cat 28 11 (39.3) 3 (10.7) 9 (32.1) 5 (17.9) Badger-Fox 85 39 (45.9) 8 (9.4) 30 (35.3) 8 (9.4) Cat-Fox 45 14 (31.1) 20 (44.4) 7 (15.6) 4 (8.9) Badger-Cat 9 6 (66.6) 1 (11.1) 1 (11.1) 1 (11.1) Total 331 27 (14.1) 53 (46.9) 79 (28.7) 24 (29.3) 101 (30.5) 43 (12.9) 3.4. Impact of the Presence of Food When data were applied to models to test the effect of interspecific dyad, interaction type and presence/amount of food on probability of wins, badgers won more interactions when agonistic behavior was involved in the dyad (parameter est. ± S.E. = 1.395 ± 0.413; z = 3.375; p< 0.001). Zero-inflated datasets prevented the inclusion of food presence and amount in some models. However, foxes exhibited a greater chance of winning when the interaction was agonistic (parameter est. ± S.E. = 2.734 ± 0.397; z = 6.878; p< 0.001) and food was present (parameter est. ±S.E. = 2.143 ±0.830; z = 2.581; p< 0.01). Animals 2023,13, 590 8 of 14 Animals 2023, 13, x 8 of 15 Hedgehog-Cat 28 11 (39.3) 3 (10.7) 9 (32.1) 5 (17.9) Badger-Fox 85 39 (45.9) 8 (9.4) 30 (35.3) 8 (9.4) Cat-Fox 45 14 (31.1) 20 (44.4) 7 (15.6) 4 (8.9) Badger-Cat 9 6 (66.6) 1 (11.1) 1 (11.1) 1 (11.1) Total 331 27 (14.1) 53 (46.9) 79 (28.7) 24 (29.3) 101 (30.5) 43 (12.9) Figure 1. A diagrammatical representation of the hierarchical relationship in ‘winning’ access to food during paired dyads. The direction of the arrow shows the direction of the dominance is the species with the higher % of wins within dyads. 3.4. Impact of the Presence of Food When data were applied to models to test the effect of interspecific dyad, interaction type and presence/amount of food on probability of wins, badgers won more interactions when agonistic behavior was involved in the dyad (parameter est. ± S.E. = 1.395 ± 0.413; z = 3.375; p < 0.001). Zero-inflated datasets prevented the inclusion of food presence and amount in some models. However, foxes exhibited a greater chance of winning when the interaction was agonistic (parameter est. ± S.E. = 2.734 ± 0.397; z = 6.878; p < 0.001) and food was present (parameter est. ± S.E. = 2.143 ± 0.830; z = 2.581; p < 0.01). 4. Discussion Our study is the first to quantify behavioral interactions between hedgehogs and three other intraguild mammal species in sub/urban gardens in the UK. We have shown that food provided in gardens by human residents is utilized by a range of intraguild species with associated evidence of interactions within and between species. Based on the assessment of ‘wins’ during interactions over food access, we have derived a hierarchical relationship between a community of sympatric urban mammals. The study findings supported our hypothesis that the largest of the species (badgers) in the guild would be most successful at ‘winning’ access to food. We also found some support for our hypothesis that food availability affected interaction type, with higher levels of agonistic interactions when food was present than when it was absent, although analysis of this element was limited due to an unbalanced dataset. Spatially predictable and/or abundant food patches can create focal activity hotspots of resource exploitation, causing aggregation of sympatric species, which likely lead to higher occurrences of interactions [40]. The consequences Figure 1. A diagrammatical representation of the hierarchical relationship in ‘winning’ access to food during paired dyads. The direction of the arrow shows the direction of the dominance is the species with the higher % of wins within dyads. 4. Discussion Our study is the first to quantify behavioral interactions between hedgehogs and three other intraguild mammal species in sub/urban gardens in the UK. We have shown that food provided in gardens by human residents is utilized by a range of intraguild species with associated evidence of interactions within and between species. Based on the assessment of ‘wins’ during interactions over food access, we have derived a hierarchical relationship between a community of sympatric urban mammals. The study findings supported our hypothesis that the largest of the species (badgers) in the guild would be most successful at ‘winning’ access to food. We also found some support for our hypothesis that food availability affected interaction type, with higher levels of agonistic interactions when food was present than when it was absent, although analysis of this element was limited due to an unbalanced dataset. Spatially predictable and/or abundant food patches can create focal activity hotspots of resource exploitation, causing aggregation of sympatric species, which likely lead to higher occurrences of interactions [ 40 ]. The consequences of interactions between garden mammals are potentially numerous. Direct interactions between competing species can be aggressive, leading to injury or death, with increased competition or competitive exclusion reducing access to resources for subordinate species or individuals, with knock-on welfare effects. Furthermore, the dynamics of space use overlap and interaction rates within wild populations and between domestic and wild species has implications for pathogen transmission, particularly in urban areas at the interface between humans and wildlife, where zoonoses can emerge [41]. 4.1. Intraspecific Interactions Our findings support the hypothesis that agonistic behavior would be higher between species compared to within species. As expected from previous studies [ 31 ], we also showed there were few agonistic interactions between badgers at feeding sites (7%). It appears that this observation is consistent between urban and rural badger populations, despite differences in urban badger group size, density, and territory size compared to rural [ 19 ]. As expected, foxes showed higher intraspecific agonism than badgers. Although foxes typically forage alone, there is a hierarchy within fox social groups [ 42 ]. Subordinate foxes have been shown to use supplementary food patches in urban gardens in a different way to dominant foxes, using fewer patches, spending less time in predictable patches, and feeding later [ 43 ]. This is explained as an evolved behavioral strategy to reduce competition/antagonism within social groups. Animals 2023,13, 590 9 of 14 Hedgehogs are considered solitary and non-territorial [ 32 ]. Field studies of behavior have shown that hedgehogs tend to avoid each other, and adults are usually only found together during courtship, or when attracted to a localized food source. Other than during courtship, when females are typically aggressive towards males, and competing male suitors may fight each other, overt aggression is rarely seen [ 44 ]. Contrastingly, we observed high levels of agonism between wild hedgehogs. This included a characteristic behavior where one hedgehog attacked another by running at it, causing the victim to roll up, after which the attacker pushed it away. Typically, the function of this behavior appeared to involve moving a competitor away from the food source, such as to the edge of the garden. In one case, an individual was pushed down a flight of concrete, and another into water. We termed this behavior ‘barge and roll’ and deemed it to be competitive. Many of these observations of agonism between hedgehogs occurred outside of the breeding period. Anecdotal evidence from rescue centers suggests aggression can occur when food is provisioned to a group; thus, the most likely explanation for agonism in our study is defense of food patches. This behavioral disparity may be a consequence of patchy distribution and abundance of urban food resources relative to rural or wilderness areas. Previous studies on red deer (Cervus elaphus) have shown increased aggression at food patches when provided with spatially and temporally predictable supplementary food during winter [ 45 ]. Access to food that will allow for an increase in body weight may be more critical for hibernating species such as hedgehogs than species that are active all year, as very low body weight could affect overwinter survival [ 46 , 47 ]. As the focus of our study was to investigate intraguild interactions of urban wildlife, we did not request videos of cat-cat interactions in this study, and thus, could not determine such interactions at garden feeding sites. 4.2. Interspecific Interactions and Hierarchy Between-species interactions were more agonistic overall than those within species, and badgers tended to be more successful at securing food than all other species. In our study, badgers were dominant over foxes where contests occurred, but if initial interactions did not escalate to aggression, each species was unaffected by the presence, proximity, or orientation of the other (consistent with [ 34 ]). Nonetheless, our study is a temporal snapshot, and previous experience may affect what was observed [ 36 ]. Interspecific interactions involving hedgehogs showed the highest levels of agonism with cats, although cats were predominantly observed to be submissive during these interactions, allowing hedgehogs access to food without contest. Hedgehogs co-occurred in urban gardens with potential predators (badgers and foxes) and approximately half of encounters between hedgehogs and foxes were agonistic. Hedgehogs have previously been reported to avoid predators [ 48 ], but in urban gardens where food is provided, they may not always do so, with the benefits of access to food provision potentially outweighing predation risk. As cats are typically fed by owners, access to additional food is less critical to their survival or fitness compared to wild counterparts, although they are highly territorial [ 49 ] and likely to defend the gardens within their territories. We observed high levels of agonism between cats and foxes, with the former appearing dominant over the latter overall. The relatively high proportion of hedgehog wins over cats was unexpected but may relate to hedgehog spines, which domestic cats are not physically or behaviorally adapted to defend themselves against, as compared to wild predators. Although competition could impact access to food, previous studies have shown that there is more food available in urban areas than some species metabolically require (e.g., foxes [ 12 , 42 , 43 ]), which suggests that if an animal unsuccessfully forages at one patch, there are likely to be multiple alternative patches where competition could be lower due to temporal activity partitioning [ 43 ]. These may be readily accessible to more mobile species, such as foxes [ 50 ], whereas hedgehog movement can be affected by barriers in urban areas [ 51 ]. This may account for the higherthan-expected agonism observed between hedgehogs, i.e., if the value of a feeding patch is perceived to be high and movement between patches is restricted, defense may be adaptive.