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Article

SARS-CoV-2 Lockdown and Ungulate Raids on Golf Courses

by
Jesús Duarte
1,*,†,
Javier Romero
2,†,
Diego Rodríguez
3 and
Miguel Ángel Farfán
2
1
Ofitecma Marbella S.L., Av. Ramón y Cajal 17, 29601 Marbella, Málaga, Spain
2
Department of Animal Biology, Faculty of Sciences, Universidad de Málaga, 29071 Málaga, Málaga, Spain
3
Rewilding Spain, 19431 Corduente, Guadalajara, Spain
*
Author to whom correspondence should be addressed.
†
These authors contributed equally to this work.
Urban Sci. 2026, 10(5), 245; https://doi.org/10.3390/urbansci10050245
Submission received: 18 February 2026 / Revised: 29 March 2026 / Accepted: 1 April 2026 / Published: 1 May 2026
(This article belongs to the Special Issue Biodiversity in Urban Landscapes)
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Abstract

The presence of wild species in urban areas is becoming increasingly common. In southern Spain, species such as wild boar cause significant damage and problems in human-dominated environments, such as peri-urban areas, sports facilities, and urban parks. Here, we used camera trapping to monitor the entry rates of ungulates (wild boar and red deer) into three golf courses located in urban areas in southern Spain. The courses are surrounded by hunting estates and other non-urban areas where species are controlled through lethal methods. Wild boars are controlled year-round, and red deer are controlled during specific hunting periods. We tested for differences during periods of normal human activity and periods of the COVID lockdown using generalised linear mixed models. We controlled ungulate raids for 2639 trapping nights, obtaining 1093 wild boar and 225 red deer independent events. During the COVID lockdown, wild boar raids on golf courses decreased significantly. Meanwhile, equivalent deer raids increased significantly during the hunting period. The results indicate that certain urban areas where control by firearms is not permitted—such as golf courses—can function as safe zones for wild species. This reserve effect is related to the structure of the urban habitat and the resources it offers in terms of security and food.

1. Introduction

Human disturbance represents a threat to biodiversity [1,2]. Among the main threats to species survival are the loss and degradation of ecosystems [3]. However, a less well-studied aspect is the impact of human presence and outdoor recreational activities, which can induce wildlife changes at the individual level, affecting space use, resource selection, reproductive success, and behaviour [4,5,6,7]. These changes also include population-level changes such as the increasingly frequent use of urban spaces.
The infectious outbreak caused by the SARS-CoV-2 coronavirus in late 2019 (COVID-19) led to an unprecedented global confinement in 2020 aimed at reducing disease transmission, sharply restricting human mobility [8]. This period—termed the “anthropause”—offered a unique opportunity to investigate the effects of human presence and activity on wildlife [9,10,11], especially in urban environments where confinement resulted in reduced crowding, pollution, and noise levels in cities. Importantly, lockdown measures represented a rapid and transitory perturbation, allowing the evaluation of short-term behavioural responses to abrupt changes in human activity. These conditions provided a valuable framework for studying behavioural plasticity over a short temporal scale, particularly in synanthropic and highly adaptable species. The lockdown also allowed wildlife to move more freely, facilitating the observation of uncommon species in urban areas [12,13,14] and enhancing species detectability [15,16,17].
Among highly adaptable urban mammals, the wild boar (Sus scrofa) provides an ideal model in which to study behavioural responses to short-term reductions in human presence and hunting pressure. The wild boar is one of the most widely distributed mammalian species, naturally occurring across Eurasia and North Africa and introduced to North and South America and Australia [18,19]. An opportunistic and a generalist species, the boar has a highly heterogeneous diet and the ability to exploit a wide variety of habitats, from forests to agricultural landscapes [20,21,22]. In recent decades, their populations have expanded both within their natural range and in areas of introduction. This expansion process is generally attributed to a combination of factors, including insufficient hunting pressure that would control population growth on a global scale, a high reproductive capacity, large-scale favourable changes in habitat, and the absence of natural predators in many parts of its distribution range [23,24].
A remarkable feature of this expansion is the increasing presence of wild boars in urban and peri-urban areas worldwide [25,26,27]. This has generated multiple risks, including attacks on humans and pets, damage to green areas and fencing, traffic accidents, and disease transmission [24,28,29,30,31]. Their urban presence is specifically facilitated by dietary flexibility and easy access to anthropogenic food sources [22,32], the absence of natural predators, and the lack of hunting pressure; the latter has a significant influence on wild boar behaviour [33,34]. In urban areas, hunting with firearms is legally prohibited for safety reasons, whereas in peri-urban and rural areas, the wild boar is actively hunted. In response to growing populations and the associated risks, some regions—such as Andalusia in southern Spain—have adopted exceptional measures, temporarily allowing year-round hunting to control populations.
Despite the high levels of disturbance and stress associated with urban environments, cities provide wild boars with diverse habitats, offering refuge and relatively constant access to food and water [32,35,36]. Individuals capable of adjusting their behaviour to human-related stress and risk are more likely to survive and reproduce. Anthropogenic disturbances have been shown to shift activity patterns toward increased nocturnality and alter habitat use, with a preference for wooded areas during the day and herbaceous green spaces for nocturnal foraging [37]. In Mediterranean urban environments, wild boars preferentially select fragmented green areas with food availability, such as gardens, parks, and other managed green spaces [38].
The presence of red deer (Cervus elaphus) in urban environments is less well-documented. Deer are common urban exploiters in the American continent [39] and are also appearing in European urban and suburban areas [40]. In certain areas of Andalusia, animals that have escaped from hunting estates or emerged from expanding populations are also beginning to use peri-urban environments and urbanisations that integrate natural settings [41]. The factors that favour ungulates successfully settling in and using urban environments seem to apply equally to deer, including certain sports infrastructures, such as golf courses [42].
In Andalusia, golf courses represent one of the most common types of urban green spaces. The region hosts most of the golf courses in Spain, with the Costa del Sol alone accounting for 50% of these facilities [43]. The courses are located in residential and peri-urban areas, often accessible to ungulates via nearby natural vegetation and corridors such as rivers and streams [26,38]. These urban green areas meet favourable habitat conditions for these species and provide food and water throughout the year due to the presence of green, fresh grass and numerous permanent water points. The abundance of moist substrate and organic matter promotes the availability of highly valued resources for some species [44]. Patches of natural vegetation within golf courses also provide food but primarily function as daytime refuge areas when players and staff are present. At night, human activity ceases, and golf courses become quiet areas where wild boars can forage without disturbance. Finally, hunting pressure is non-existent, as hunting activities are not permitted on golf courses due to their legal classification as urban areas [45].
Urban green areas in general and golf courses in particular may exert a “reserve effect” for species that are actively hunted in adjacent peri-urban areas; this phenomenon has been described in previous studies [34,46,47,48]. Importantly, golf courses provide a well-defined and consistent habitat that allows for indirect assessment of this reserve effect, enabling comparisons of ungulate use before, during, and after periods of restricted human mobility.
The simultaneous interruption of hunting pressure and human presence and its effects on the use of urban refuges by ungulates has not yet been empirically evaluated. For this reason, we aimed to assess whether the confinement resulting from the SARS-CoV-2 outbreak affected golf course use by wild boars and red deer. We hypothesise that hunting activity ceased in peri-urban areas during confinement, temporarily removing the “reserve effect” of golf courses, resulting in ungulates reducing their use of these habitats.

2. Materials and Methods

2.1. Study Area

The study was carried out simultaneously at three 18-hole golf courses located in the south of Spain, Málaga province (Figure 1). The climate in the area is temperate-subtropical Mediterranean, with average temperatures of 11 °C in January and 25 °C in July, and with an annual rainfall of less than 1000 mm [49].
The first included course is called “Los Arqueros Golf & Country Club” (hereafter, golf course A; Figure 2), located in the municipality of Benahavís (36°30′38″ N, 5°00′58″ W; Málaga, the south of Spain). The golf course is a private estate of 99.7 ha in which an urbanised area comprises residential dwellings and apartments. Course fairways and facilities, adjoining villa garden areas, and green areas composed of patches of the original forest dominate the area interspersed with the golf course. The course is surrounded by an urban matrix, except on its western boundary, which borders a forest estate.
Elevation within the property ranges from 105 m to 195 m above sea level. The vegetation is typically Mediterranean. Some forest patches are found, dominated by maritime pine (Pinus pinaster) mixed with carob (Ceratonia siliqua), wild olive (Olea europaea var. sylvestris), and cork oak (Quercus suber). However, most of the green areas inside the estate consist of Mediterranean scrubland, largely composed of rock rose (Cistus sp.), European dwarf palm (Chamaerops humilis), Labiatae, and gorse (Ulex sp.), with some heather (Erica sp.) and strawberry trees (Arbutus unedo).
The streams that typically cross the golf courses in the study area are relevant landscape features for the purposes of this study because they function as wildlife corridors. Two streams cross golf course A, the “La Leche” stream to the west and the “El Chopo” stream to the east. Both streams have a border of riparian vegetation in which willows (Salix pedicellata), oleanders (Nerium oleander), and isolated examples of gall oaks (Quercus faginea) dominate. The “La Leche” stream flows to a reservoir with the same name, 850 m from the golf course, while the “El Chopo” flows to the “Las Medranas” reservoir. The surroundings of both reservoirs are populated with pine trees, eucalyptus (Eucalyptus camaldulensis), and scrubland.
The second golf course was “Aferini Golf” (hereafter, golf course B; Figure 2), also located in the municipality of Benahavís (36°29′36″ N, 5°02′44″ W; Málaga). Course B is part of a larger complex with two other 18-hole courses, a golf academy, and a residential area called “Los Flamingos”. The course is 49.3 ha. The property is located within a developing peri-urban area. It borders urbanised areas to the northwest and south; the remaining perimeter borders an estate that was a hunting reserve until a few years ago, when it was reclassified from rural to developable land.
Elevation within the course ranges from 100 m to 156 m above sea level. The plant community surrounding the course is composed mainly of scrubland associated with peridotites (Cistus albidus, Halimium atriplicifolium, Ulex baeticus, Quercus coccifera, Chamaerops humilis, and Asparagus spp.) and micaschists (Pistacia lentiscus, Cistus monspeliensis, Cistus crispus, and Calicotome villosa). This coverage sometimes appears mixed, with isolated Q. suber and small patches of P. pinaster.
Golf course B is crossed from north to south by the “Briján” stream, which flows into the “Cancelada” reservoir, 110 m from the southern course border. The stream is associated with dense formations of tamarisk (Tamarix spp.), oleander (N. oleander), brambles (R. ulmifolius), and willows (S. pedicellata).
Two ungulate species—the wild boar (S. scrofa) and red deer (C. elaphus)—inhabit the area surrounding both golf courses. Red deer populations come from escapes from a nearby estate [42], having acclimatised to urban areas, expanded, and become abundant.
The third golf course was “Los Naranjos” Golf Club (hereafter, golf course C; Figure 2), located in the Nueva Andalucía district of Marbella (36°30′54″ N, 4°58′18″ W; Málaga, south of Spain). The course is 57.3 ha. The course is surrounded by an urban matrix and integrated urban areas.
Elevation within the course ranges from 50 m to 80 m above sea level. The course is made up of three sectors connected to each other but discontinuous due to the intersection of the “Osa Mayor” and “Sirio” streets. The course is in a residential area, surrounded by houses and dwellings with gardens. There are no patches of natural vegetation in its surroundings except for a small pine forest (Pinus halepensis) mixed with eucalyptus plantations (E. camaldulensis) and associated scrubland (Ulex spp., Cistus spp., and P. lentiscus), located 600 m to the northwest and close to the “El Ángel” reservoir.
The “Benavolá” stream crosses golf course C, connecting two reservoirs: “El Ángel” to the north and a small reservoir called “Lagomar”, 270 m south of the course border. The stream is associated with riverside oleanders (N. oleander), brambles (R. ulmifolius), ferns (Pteridium aquilinum), tamarisk (Tamarix spp.), and willows (S. pedicellata), as well as some olive trees (O. europaea var. sylvestris) mixed with carob trees (C. siliqua). There are no red deer in this area, but wild boars are abundant.
The three golf courses each have a perimeter closure consisting of metal fencing. However, in some places, the fence is neither fixed to the ground by concrete—preventing access to wild boars—nor is it high enough to prevent jumping, in the case of deer. Nevertheless, the typical access routes for ungulates are the streams that cross the courses and connect them with areas which they normally inhabit. Streams also allow ungulates a safe way to move in the urban matrix, as green corridors with the greatest vegetative coverage.

2.2. Hunting Planning in the Study Area

Golf course A is 2.8 km away from the hunting estate MA-10837, “La Máquina”, and 2.9 km from MA-10405, “La Toma”. Golf course B is 4.1 km away from MA-10455, “El Zorzal”, 3.3 km from MA-10405, “La Toma”, and 3.8 km from MA-10771, “La Resinera”. Golf course C is 2.1 km away from MA-10778, “Sierra Real”, and 3.2 km from MA-10766, “El Meliche” (Figure 3). These hunting estates are all located north of the golf courses. A rural area of variable width exists between the courses and the hunting estates, in which damage control hunting is permitted through authorised hunting blinds. According to Andalusian hunting regulations, the use of firearms for damage control is only forbidden in urbanised areas. However, most agricultural or undeveloped forestry properties in the area request permits for hunting blinds to prevent damage from wild boars [50]. Therefore, in practice, ungulate hunting with firearms takes place along the borders of the golf courses.

2.3. Hunting Seasons in Andalucía

Red deer hunting is allowed between mid-October and early February in Andalucía [51]. However, wild boar hunting is allowed year-round due to a temporary hunting emergency declared in response to damage and health risks from wild boars and feral pigs [50]. These emergency measures for wild boar hunting have been in effect for more than 10 years and have been exacerbated by the recent risk of swine fever [52].

2.4. Sampling Period and Ungulate Monitoring

We monitored ungulate raids on the golf courses by installing 18 camera traps. Normally, the stream connects the golf course through a road underpass located at the course border. We therefore installed camera traps at these underpasses facing the closest sections of the streams, 500 m apart. Detected ungulate activities in the direction of the golf course were selected, discarding sequential events (i.e., ignoring repeated detection of animals in successive cameras along the same stretch of the stream). Only independent species detections [53] were considered, comprising pictures taken >30 min apart, thus representing independent ungulate events [54]. The rate of ungulate detections (i.e., the trapping rate) for each trap location was estimated and expressed as the number of independent events per 1000 trapping nights. Sampling included 2639 trapping nights (see Table 1).
The sampling period covered June 2020 to March 2022. In Spain, there were three main COVID lockdown periods—the so-called “anthropauses”—in which human activity was reduced to a minimum. The first (spring 2020) was not covered by our study period. However, the second (autumn 2020) and the third (winter 2020 to spring 2021) were covered. During these lockdowns, not only was human activity reduced, but hunting was also not allowed. The golf courses had no commercial activity (players), with only minimal maintenance taking place during the day. This allowed us to compare wild boar raids between these lockdowns and normal periods of activity.

2.5. Data Analysis

We tested the effects of the COVID lockdown (binomial) and seasonality on the ungulate trapping rate (the dependent variable) using generalised linear mixed models (GLMMs) [55]. For red deer, the hunting period (binomial) and the interaction between seasonality and hunting period were also included as factors; for wild boars, the hunting period comprised the entire year. Seasons were previously defined according to solstices and equinoxes of the northern hemisphere, which according to the National Geographic Institute for Spain are spring (20 March to 20 June), summer (21 June to 23 September), autumn (24 September to 20 December), and winter (21 December to 19 March). The GLMM used a normal error distribution and an identity link function. The year and golf course sampled were included as random effects in the model. We generated one model each for wild boars and red deer. The differences in trapping rate were assessed through one-way ANOVA [56]. For deer, we identified sex from photo captures whenever possible to assess whether the raids by sex and season were different, assessed by chi-squared test using the Yate’s correction for one degree of freedom [57]. Wild boar sex could not be identified. Means are given with their standard errors.

3. Results

Golf course A was the most affected by ungulates. A total of 3288 wild boar raids were detected during the study period, of which 977 (29.7%) were considered independent events. Red deer numbers amounted to 1015 in total, of which 146 (14.4%) were independent events. At golf course B, 34 wild boar raids were detected, of which 25 (73.5%) were independent events, and 149 deer raids were observed, of which 79 (53%) were independent events. Finally, golf course C suffered 330 wild boar raids, of which 91 (27.6%) were independent events. In total, camera trapping captured 1093 independent wild boars and 225 independent red deer events (see Table 2 for more details).
Wild boar trapping rate was significantly affected by the COVID lockdowns (GLMM; F = 11.483; p = 0.001). The wild boar trapping rate was significantly lower (ANOVA; F(1451) = 29.049; p < 0.001) during lockdown (1.370 ± 0.057 captures × 1000 trapping nights) than during the normal human activity period (2.108 ± 0.094 captures × 1000 trapping nights). The seasons did not significantly affect the trapping rate (GLMM; F = 2.325; p = 0.074), although trapping rates were significantly different between seasons (ANOVA; F(3451) = 19.393; p < 0.001; see Table 3). COVID lockdowns were therefore the main factor affecting wild boar raids into golf courses, which decreased during the anthropause; there were also seasonal changes in the frequency of these raids (Figure 4), which increased during summer and autumn.
In contrast, the red deer trapping rate was not affected by the COVID lockdowns (GLMM; F = 0.358; p = 0.551), by seasonality (GLMM; F = 2.584; p = 0.056), or by the interaction between seasons and hunting period (GLMM; F = 1.118; p = 0.292). However, the hunting period did significantly affect the deer trapping rate (GLMM; F = 7.047; p = 0.009). Red deer raids on golf courses were significantly different between hunting periods (ANOVA; F(1149) = 9.958; p = 0.002), being higher during the hunting season (2.489 ± 0.128 captures × 1000 trapping nights) than during the closed period (1.848 ± 0.096 captures × 1000 trapping nights). In addition, the deer trapping rate was significantly different between seasons (ANOVA; F(3149) = 4.997; p = 0.003; see Table 3). Therefore, the main factor affecting red deer raids on golf courses was the hunting period, with deer accessing the courses more frequently during periods when they could be hunted. There were also seasonal changes in the raids (Figure 5), which decreased in summer.
Sex was determined for 173 deer (76.9% of the total deer captured). Females, usually accompanied by fawns, entered golf courses significantly more frequently than males in winter (Chi2 = 5.758; df = 1; p = 0.016) and autumn (Chi2 = 4.830; df = 1; p = 0.027). Males entered more frequently in spring (Chi2 = 27.272; df = 1; p < 0.001); there were no significant differences during summer (Chi2 = 0.493; df = 1; p = 0.482). Figure 6 presents deer raids by sex and season.

4. Discussion

The ecological “landscape of fear” concept [58,59] proposes that animals alter their behaviour and habitat use based on perceived risk. The spatial and temporal distribution of high and low-risk areas forces species to balance their ecological requirements (foraging needs, movements, and habitat selection) with safety [60]. The concept shapes population dynamics across systems and provides a dimension for understanding how human activity can influence the creation of a landscape of fear [61].
Our results clearly indicate that hunting acts as a predatory activity, creating high-risk zones for the safety of ungulates. Conversely, urban areas, like golf courses, can function as low-risk zones where species perceive increased safety. In other words, a landscape of fear is created between areas where hunting activity is permitted and areas where it is prohibited. Wild boars accessed golf courses in the study area significantly less frequently during the COVID lockdown, while red deer used them more frequently during the hunting season. These results imply that golf courses are functioning as sanctuaries for these species. This “reserve effect” [34] is mainly explained by the lack of firearm use within these urban green spaces. Although other control methods can be used within golf courses (such as trap cages or dogs) [50], they do not appear to stop ungulate raids.
In the case of deer (which can only be hunted for a few months each year), golf courses represent lower-risk areas during hunting season. They therefore use golf courses significantly more during the hunting season (autumn and winter). The case of wild boars has an important peculiarity, in that they can be hunted year-round. This led to less frequent use of golf courses during the anthropause. In other words, the area outside golf courses ceased to be perceived as a high-risk zone for wild boars when human activity and hunting were drastically reduced.
In addition to the primary results of this study, we also show that there are raids of both ungulate species on golf courses throughout the year, more frequently in some seasons than in others. Wild boars access golf courses all year round, with a near-constant rate that increases in summer and a little less during autumn. This is logical; summer is a time of climatic stress in the Mediterranean basin, which may affect wild boars [62], as well as other species. There is less water available in the environment and, therefore, less food. At this time, golf courses provide not only refuge but also an easy larder where water, damp soil, and food may be plentiful. Autumn, especially in the early months, can function climatically like summer. Only with the arrival of the first rains and the falling of acorns (acorn-feeding season or “montanera”) does the need to search for food and water in golf courses potentially decrease [22,63], leading wild boars to use these facilities less.
Deer also access golf courses at a nearly constant rate, except during summer. Red deer are a browser species that select their habitat seasonally, seeking a balance between food and shelter [64,65]. These movements are also influenced by the reproductive season, which produces a sexual segregation in terms of nutritional requirements [65]. During certain periods, red deer select open areas and grassland [66,67]. In this context, ecotone zones, with availability of grass and woody vegetation (providing shelter and different types of food), are important for the species [65]. Golf courses may therefore become favourable habitats for deer [42].
According to our findings, the entry rate of deer into golf courses increases in winter and spring, with a marked difference between sexes. During summer, the entry rate descends, and it increases gradually during autumn. One possible hypothesis to explain the summer decline is that this is the time of greatest human activity on golf courses. Red deer have crepuscular habits that can change seasonally [68], especially in response to human activity in urban areas [69] that can force avoidance [70] and altered behaviour in urban mammals [71]. Females are more common on golf courses in winter and autumn—probably looking for easy food sources—while males are more elusive in these seasons. Females are rarely detected in spring because they are giving birth (mostly from May to June [72]. During this period, they are elusive, or their fawns are very young. Females protect their fawns; their diel activity pattern changes, prioritising safety over food [73]. Therefore, we hypothesise that they may use golf courses less frequently and look for natural areas with more vegetation cover. During summer, the critical period in the Mediterranean, both sexes access equally: males are preparing for the mating season and need to feed, while females may have fawns of several months of age, who also need food but are more capable of moving and less dependent on their mothers.

4.1. Limitations of the Study

One of the limitations of this study is the inability to provide the data on deer and wild boar hunting yields (animal catches and hunters involved) during hunting seasons in the study area. Regrettably, these data are not publicly available. In any case, reported catch figures do not usually reflect reality, as poaching is widespread and the truth about catches is rarely disclosed, so official figures are usually underestimated [74]. Another possible limitation is that there may have been some poaching during the COVID lockdown period. While this is likely, it must have been negligible, given the heavy police presence during those periods, both day and night.
Furthermore, our results associated with the cessation of hunting activity during confinement cannot be separated from the decrease in wild boar incursions, which could be due to a combination of factors such as reduced noise, human scents, or even changes in the availability of anthropogenic food resources outside the golf courses. The presence of wild boars in urban areas, even during the day, is very common in the study area. The animals are accustomed to noise, human presence, and traffic, so these factors do not seem to significantly affect their activity. Changes in food availability and anthropogenic subsidies could have had an impact (restaurant closures and less organic waste produced). However, results show that wild boars abandon safe areas (golf courses) when hunting activity ceases. Anthropogenic subsidies for these animals occur primarily in urban areas, where they scavenge for food in garbage. On golf courses organic waste production is very limited. There is only one club per golf course that offers food to the public, and it was closed during the lockdown, but existing residential properties continued to produce organic waste (i.e., subsidies availability). In any case, on golf courses, wild boars typically seek more natural food (earthworms and roots) in the fairways, which is why they cause so much damage in these facilities.

4.2. Urban Development, Ungulates, and Human–Wildlife Coexistence

Ungulates have great capacity to colonise new environments [75]. In the case of the wild boar, the species is widely distributed and uses a great variety of habitats [76]. Human disturbance, hunting, or the presence of large predators have been shown to be factors that influence them to avoid certain areas [75,77,78,79] or to alter their behaviour between day and night [80]. However, it has also been shown that these animals can become habituated to regular human disturbances in areas of low human impact [81] and alter the nature of their responses (such as level of vigilance vs. flight response) according to available cover [70]. Both cover availability and low-level human presence are the features of golf courses. In addition, it has been suggested that hunting induces stronger fear responses regardless of coverage availability [82]. This highlights the value of areas where hunting is prohibited and where other resources allow these species to meet their needs.
In this context of the great ecological plasticity of these species [83], personality plays a role. Bolder individuals emerge [84,85], capable of venturing into and adapting to new environments [32,35,86]. The spread of fearless behaviour favours the colonisation of urban areas [87], and microevolution from shyness to boldness [81] may be supported by their success and improved fitness in these new environments.
However, the question remains of why urban environments favour ungulate success. Several reasons have been proposed; of these, the spatial structure of the urban environment and the resources it offers fit the findings of this case study. Ungulate species do not typically live permanently in urban environments; instead, a source–sink dynamic is established between rural and urban environments [88]. The connectivity between both environments is important. Species look for fragmented areas, connected by green corridors [38]. These green corridors offer shelter and food for travel between environmental niches. Urban environments connected to their rural surroundings by streams offer optimal entry routes. Meanwhile, fragmented areas—the breaking apart of continuous, natural habitats into smaller, isolated patches, usually driven by human activities like urban expansion [89]—offer ecotones and diversity of resources to small populations. This fits well with the structure of peri-urban and ex-urban landscapes, and with the golf courses included in this study area.
Ungulates tend to avoid areas with high dwelling density, but their presence is favoured when building density is low, and dwellings are integrated into a matrix of green areas with a diverse environment, with integration of forest and open areas [90]. This pattern again applies to peri- and ex-urban environments. In these environments, animals find both natural resources to satisfy their needs and subsidies they can take advantage of, such as garden pruning waste [91]. These anthropogenic food sources are predictable and can improve the species’ reproductive success [92]. The adjustment of litter size to food availability [93] creates a feedback loop that leads the animals to return to the area. More stable environmental conditions than in a natural environment can be added to these conditions, especially in terms of the availability of water and humidity; this can occur in the form of irrigation systems or artificial wetlands, further favouring reproductive performance [94,95]. Finally, there is the security factor, which, in the case of urban areas, is provided by the prohibition of hunting [96], creating small-scale refuges from human activity for ungulates [79]. Urban areas that meet all these criteria, such as golf courses, therefore become an optimal habitat for ungulates.
The expansion of ungulates into urban areas has secondary consequences for humans: human–wildlife conflicts arise, damage occurs, adaptive control strategies are required, and, in some cases, society’s view of these animals becomes more negative [97,98]. Different capture methods have been tested, especially for wild boars [99]; there is still a lack of information on alternatives for controlling red deer populations, and on the general effectiveness of other options such as deterrents, fencing, and habitat management. Managing these species in urban habitats requires new approaches beyond simply trapping them. Urban structure and the management of green spaces can play a significant role in this. These should therefore be priority lines of research in a world where wild species are becoming increasingly common in human-modified habitats.

Author Contributions

Conceptualization, J.D. and D.R.; methodology, J.D.; software, J.D. and J.R.; validation, M.Á.F. and J.D.; formal analysis, J.D. and J.R.; data curation, J.D. and J.R.; writing—original draft preparation, J.D. and J.R.; writing—review and editing, M.Á.F. All authors have read and agreed to the published version of the manuscript.

Funding

This research received no external funding.

Institutional Review Board Statement

Not applicable.

Informed Consent Statement

Not applicable.

Data Availability Statement

The original contributions presented in this study are included in the article. Further inquiries can be directed to the corresponding author.

Acknowledgments

We would like to thank the golf course managers and staff that helped us during the monitoring.

Conflicts of Interest

Author Jesús Duarte was employed by the company Ofitecma Marbella SL. The remaining authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.

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Figure 1. The location of the study area in the south of Spain, Andalucía, Málaga province. Golf courses: (A) Los Arqueros; (B) Alferini; and (C) Los Naranjos.
Figure 1. The location of the study area in the south of Spain, Andalucía, Málaga province. Golf courses: (A) Los Arqueros; (B) Alferini; and (C) Los Naranjos.
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Figure 2. The three golf courses sampled for estimating ungulate raids ((A) Los Arqueros; (B) Alferini; and (C) Los Naranjos), all of them located in the south of Spain, Andalucía, Málaga province. Courses (A,C) are in the Benahavís and Marbella municipalities, respectively, and are integrated within the urban matrix. Course (B) is in the Benahavís municipality, in a peri-urban context. The northern area in course (C) is still under urban development. The red dot indicates the access point from which camera traps were installed, downstream of the streams (marked in blue).
Figure 2. The three golf courses sampled for estimating ungulate raids ((A) Los Arqueros; (B) Alferini; and (C) Los Naranjos), all of them located in the south of Spain, Andalucía, Málaga province. Courses (A,C) are in the Benahavís and Marbella municipalities, respectively, and are integrated within the urban matrix. Course (B) is in the Benahavís municipality, in a peri-urban context. The northern area in course (C) is still under urban development. The red dot indicates the access point from which camera traps were installed, downstream of the streams (marked in blue).
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Figure 3. Hunting estates (shaded in pink) to the north and west of the golf courses (green) and the developed urban matrix (purple) in the study area. The spaces between the courses, the urban matrix, and the hunting estates are rural or agricultural properties, where damage control with firearms is permitted subject to authorisation. (A) Los Arqueros; (B) Alferini; and (C) Los Naranjos.
Figure 3. Hunting estates (shaded in pink) to the north and west of the golf courses (green) and the developed urban matrix (purple) in the study area. The spaces between the courses, the urban matrix, and the hunting estates are rural or agricultural properties, where damage control with firearms is permitted subject to authorisation. (A) Los Arqueros; (B) Alferini; and (C) Los Naranjos.
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Figure 4. Wild boar trapping rate in relation to the COVID lockdowns and season in the study area. The cross represents the median.
Figure 4. Wild boar trapping rate in relation to the COVID lockdowns and season in the study area. The cross represents the median.
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Figure 5. Red deer trapping rate in relation to the COVID lockdowns, hunting period, and season in the study area. The cross represents the median.
Figure 5. Red deer trapping rate in relation to the COVID lockdowns, hunting period, and season in the study area. The cross represents the median.
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Figure 6. Red deer raid frequency on golf courses by sex and season in the study area.
Figure 6. Red deer raid frequency on golf courses by sex and season in the study area.
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Table 1. Sampling effort and periods in the study area.
Table 1. Sampling effort and periods in the study area.
Golf CourseCams InstalledTrapping NightsStartFinish
A. Los Arqueros91464June 2020March 2022
B. Alferini2442March 2021March 2022
C. Los Naranjos7733July 2020March 2022
Total182639
Table 2. Independent ungulate events during the sampling period in the study area across different periods.
Table 2. Independent ungulate events during the sampling period in the study area across different periods.
PeriodWild BoarRed Deer
Independent events (total)1093225
COVID lockdowns76741
Normal activity period326184
Hunting season---55
Non-hunting season---170
Table 3. Seasonal trapping rates (ungulate × 1000 trapping nights; mean ± SE) and effort in the study area.
Table 3. Seasonal trapping rates (ungulate × 1000 trapping nights; mean ± SE) and effort in the study area.
WinterSpringSummerAutumn
Wild boar1.456 ± 0.0821.299 ± 0.1262.279 ± 0.1501.889 ± 0.107
Red deer2.388 ± 0.2122.316 ± 0.1271.631 ± 0.1422.094 ± 0.158
Trapping nights562546738793
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MDPI and ACS Style

Duarte, J.; Romero, J.; Rodríguez, D.; Farfán, M.Á. SARS-CoV-2 Lockdown and Ungulate Raids on Golf Courses. Urban Sci. 2026, 10, 245. https://doi.org/10.3390/urbansci10050245

AMA Style

Duarte J, Romero J, Rodríguez D, Farfán MÁ. SARS-CoV-2 Lockdown and Ungulate Raids on Golf Courses. Urban Science. 2026; 10(5):245. https://doi.org/10.3390/urbansci10050245

Chicago/Turabian Style

Duarte, Jesús, Javier Romero, Diego Rodríguez, and Miguel Ángel Farfán. 2026. "SARS-CoV-2 Lockdown and Ungulate Raids on Golf Courses" Urban Science 10, no. 5: 245. https://doi.org/10.3390/urbansci10050245

APA Style

Duarte, J., Romero, J., Rodríguez, D., & Farfán, M. Á. (2026). SARS-CoV-2 Lockdown and Ungulate Raids on Golf Courses. Urban Science, 10(5), 245. https://doi.org/10.3390/urbansci10050245

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