Deer management has become an integral part of ecosystem recovery efforts across the globe. Within Scotland, annual deer culls have been implemented to control deer browsing, with the carcasses most often removed from the landscape. Given that animal bodies concentrate large quantities of nutrients, this practice may deplete ecosystems of vital nutrients.
We quantified the nitrogen, phosphorous and calcium losses from the removal of culled deer carcasses using nationwide statutory cull reports for four deer species in Scotland between 2010 and 2021. We estimate that annual losses from carcass removal over this period averaged 195,652 kg of nitrogen, 152,834 kg of phosphorus and 251,188 kg of calcium across Scotland. While both red and roe deer were culled at a much higher rate than the other two species red deer culls accounted for approximately 70% of the nutrients lost. Further, while large quantities of all three nutrients were removed from the landscape, calcium losses were particularly high.
We then calculated nutrient losses within the three land classifications used in statutory cull reporting—agricultural areas, open range and woodlands—across Scotland's Deer Count Areas. Using data from the literature, we considered these losses in the context of other major environmental inputs and outputs within each land classification. Our results demonstrate that while open range lost more nutrients compared to the other two land classifications, culling resulted in high rates of phosphorus and calcium loss throughout all land classifications when compared to other environmental inputs.
Practical implication. Our findings suggest that current practices of carcass removal are gradually stripping nutrients from the Scottish landscape, potentially undermining habitat recovery goals. While this study offers a preliminary, coarse scale summary of the issue, the way forward requires further study of local effects from carcass removal on nutrient pools and balancing deer management with habitat function through interwoven deer and nutrient management strategies.

1 INTRODUCTION

Managing deer populations has long been a goal of wildlife ecology and conservation biology (Côté et al., 2004; Leopold, 1933). While deer are valued as keystone species in some contexts (Judziewicz & Koch, 1993) and provide essential ecosystem contributions (Bump, Webster, et al., 2009; Ferraro et al., 2023; Vellend et al., 2004), high densities of deer can have outsized impacts on natural and semi-natural systems, leading to conflicts with human land-use objectives (Putman et al., 2011; Putman & Moore, 1998). Due to the human-driven extirpation of large predators and increases in agriculture worldwide, populations of wild deer have increased in numbers and range over the last century (Côté et al., 2004). These anthropogenic changes to ecosystems have facilitated deer populations to be considered overabudant in many areas (Côté et al., 2004). Importantly, overabundance is a value judgement (Côté et al., 2004): though within ecology, animals are often considered overabundant when they ‘(a) threaten human life or livelihood, (b) are too numerous for their “own good,” (c) depress the densities of economically or aesthetically important species or (d) cause ecosystem dysfunction’ (Caughley, 1981).

In Britain, deer range expansion brings benefits and costs (Ward, 2005). In particular, the selective browsing and grazing of wild deer is of growing concern across European forestry systems, where economic losses are common due to decreased timber quality, reduced tree growth, and increased tree mortality (Gill, 1992a, 1992b; Welch et al., 1991). Additionally, there are growing concerns over damage to natural and protected habitats caused by high-pressure deer browsing (Putman & Moore, 1998; Reimoser & Putman, 2011), which can lead to woodland degradation or even loss of unfenced woodlands (Gill & Morgan, 2010; Reimoser & Putman, 2011). To combat these ecosystem impacts, deer populations are commonly managed through culling (Putman & Moore, 1998). In Scotland, deer populations have been actively managed since 1959 (The Deer Working Group, 2019). Currently, all four wild deer species present in Scotland are considered overabundant (MacMillan & Leitch, 2008), with total deer populations ranging from 593,000 to 777,000 (Pepper et al., 2019; The Deer Working Group, 2019) and densities ranging from 4 to 64 km² (Forestry and Land Scotland, 2021). The most recent population estimates indicate there are approximately 360,000–400,000 red deer (Cervus elaphus), 200,000–350,000 roe deer (Capreolus capreolus), 25,000 sika deer (Cervus nippon), and 8000 fallow deer (Dama dama) (The Deer Working Group, 2019). Between 100,000 and 180,000 deer have been culled annually over the last 20 years (The Deer Working Group, 2019), representing roughly 20% of the population each year. However, there has been mounting pressure to further reduce deer numbers across Scotland, which has led to recent changes in legislation that have removed the close season for all species of male deer in Scotland (Deer (Close Seasons) Amendment Order, 2023).

While deer management in Scotland is primarily aimed at mitigating the ecological impacts of browsing, deer management practices are at risk of overlooking another key ecological process, carcass deposition. Large mammals concentrate essential nutrients within their bodies. Under natural conditions, when an animal dies, these zoogeochemical inputs (Schmitz et al., 2018) are recycled back into the ecosystem through ecological food webs (Wilmers et al., 2003) or returned to the environment through decomposition in large quantities that often exceed natural concentrations (Danell et al., 2002; Subalusky et al., 2017). Such depositions can facilitate increased plant biodiversity (Bump, Webster, et al., 2009), biogeochemical hotspots (Bump, Webster, et al., 2009; Melis et al., 2007) and landscape-level heterogeneity (Bump, Peterson, & Vucetich, 2009; Ferraro et al., 2022). However, in Scotland, culled deer carcasses are typically removed from the landscape, presumably to enter the human food chain as venison; however, there are no reliable national statistics on the actual proportion of culled deer processed for commercial meat sales (The Deer Working Group, 2019).

Deer, in particular, concentrate large quantities of nitrogen, phosphorus and calcium in their bodies and antlers; nitrogen is found primarily in crude protein within the body (McCullough & Ullrey, 1983, p. 19; Robbins et al., 1974), while phosphorus and calcium are vital structural components of bone, muscle, and antlers (McCullough & Ullrey, 1983; Moen & Pastor, 1998). In other systems, the systematic removal of large mammal carcasses can deplete these crucial nutrients (Abraham et al., 2021; Flueck, 2009; Innes, 1983; Trepel et al., 2024), which can have consequences for wildlife and ecological systems alike (Abraham et al., 2021). As such, the continual suppression of nutrient input via the removal of deer carcasses across Scotland could undermine the long-term success of the reforestation and ecosystem restoration initiatives that deer management is implemented to safeguard.

Of the four deer species in Scotland, roe deer and red deer are native, while sika deer and fallow deer are non-native introductions (The Deer Working Group, 2019). However, cull efforts in Scotland generally target deer populations based on their overall abundances and impacts, rather than discriminating between native and non-native species. Therefore, from an ecological perspective, the removal of any deer carcass from the landscape can potentially lead to a loss of nutrient inputs, regardless of the species origin.

While many northern systems tend to be nitrogen-limited (Du et al., 2020), Scottish ecosystems present mixed results. Plant growth within Scotland's regenerative woodland, for example, is limited by either nitrogen or phosphorus, depending on the presence of browsing (Carline et al., 2005). Calluna–Cladonia heathlands, on the other hand, are characterized by nitrogen-limitation, which is necessary for these systems to persist (Britton & Fisher, 2006). Throughout the country, soils are calcium-poor, and while calcium is not a limiting element to growth directly, calcium loss can impact the soil pH (Ogg & Dow, 1928). Additionally, phosphorus and calcium have relatively localized cycling systems that may be especially vulnerable to nutrient depletion. Phosphorus is virtually immobile, resulting in nearly closed local nutrient cycles (Smil, 2000). Calcium availability is governed primarily by atmospheric deposition, mineral weathering, and leaching losses (McLoughlin & Wimmer, 1999); however, increasing evidence shows that anthropogenic activity, such as acid rain (Likens et al., 1996) and timber harvesting (Reynolds & Stevens, 1998) can deplete calcium from ecosystems. Given that many commercial forestry systems rely on strict control of deer in Scotland, deer carcass removal could represent a substantially unrecognized and unaccounted-for loss of key nutrients, heavily undermining the nutrient sustainability of Scottish forestry.

Using publicly available data, we quantify the loss of nitrogen, phosphorus and calcium from deer culls throughout Scotland. Our analysis spans different spatial scales, offering nationwide estimates as well as specific assessments for localized deer count areas (DCAs). Additionally, we examine the nutrient loss in distinct habitats—woodland, open range and agriculture—following the land classifications from the statutory deer cull reports. Ultimately, we compare the loss of nutrients from carcass removal in these habitats to other inputs and outputs, such as atmospheric deposition and timber harvesting. This study represents a first step in determining whether the current carcass removal practices deplete key nutrients and raise concerns that such practices could hinder ecosystem recovery. We recognize that deer have many other positive and negative impacts on nutrient cycles and ecosystems (Bump, Webster, et al., 2009; Ferraro et al., 2023; Putman & Moore, 1998; Reimoser & Putman, 2011; Vellend et al., 2004); however, we take a value-neutral approach on the presence of deer, which allows us to assess the role of carcasses without advocating for a specific management policy within Scotland. While we do not make a value judgement on culling, our results will help guide evidence-based recommendations for integrated deer and nutrient management to better protect Scotland's plans for large-scale ecosystem recovery and sustainable wildlife management.

2 MATERIALS AND METHODS

2.1 Collection of cull data

We compiled national cull data for red, roe, sika and fallow deer in Scotland from 2010 to 2021 using data provided by NatureScot (The Deer Working Group, 2019). Deer culls are primarily conducted by professional stalkers and hunters hired by landowners, although recreational hunting occurs and is also represented in the deer cull statistics (The Deer Working Group, 2019). Annual cull returns are submitted by landowners to NatureScot upon request. These are initially arranged by council areas, but to provide higher spatial resolution, we requested they be categorized by the 81 DCAs, which act as spatially bound administration units to monitor and manage deer populations in Scotland. Reports were unavailable for three DCAs (East Ross, Edinburgh City and West Sutherland), so these were removed from further analysis. For each of the remaining 78 DCAs, the cull data detailed the number of males, females, and juveniles culled for each of the four deer species across three land classifications: agricultural, open range or woodland. Culls were reported over an annual period between 1 April and the following 31 March. Although the data dates back to 1996, we used annual data from 2010 to 2021 as data for culls from the ‘agriculture’ land classification is only available after 2005. To account for the underreporting of culls, we applied species-specific correction factors from the Deer Working Group (2019) to the reported cull data. For each deer species, we divided the number of reported culls in each age/sex category by the corresponding correction factor (red deer 10%, roe deer 60%, sika deer 25% and fallow deer 25%). This approach allowed us to evenly distribute the underreporting across demographic categories, enabling a more accurate representation of realized deer culls in Scotland.

2.2 Carcass mass

We conducted a literature search to determine each species' average body and antler mass, including sex and juvenile-specific averages. In ungulates, environmental factors such as population density and climatic conditions can affect adult body mass (Putman & Flueck, 2011), and the live masses of deer vary across the UK (Hewison, 1996; Putman & Flueck, 2011). Therefore, we estimated deer mass based on Scottish data where possible and used data from other regions in Britain if Scottish data were unavailable. As viscera (or gralloch) are typically left at the location where deer are shot (Putman et al., 2019), we present both gralloched mass (live mass minus the gralloch) and live mass (Table S1). Further information on how we determined each species' mass can be found in the supplement.

2.3 Nutrient budgets

To assess the nutrient loss, we calculated the estimated total mass of nitrogen, phosphorus and calcium within each population culled (Ellis-Soto et al., 2021). Given that the gralloch is nitrogen-rich, we used gralloched masses for all deer species and age/class sizes for nitrogen calculations. Live weight analyses estimate that crude protein is 21% of the mass of a white-tailed deer (McCullough & Ullrey, 1983; Robbins et al., 1974) and 22%–23% of the mass of a roe deer (Daszkiewicz et al., 2012). While nitrogen is present in several molecules in the body, including urea and creatine, estimates for the total nitrogen in deer bodies are not reported in the literature. Thus, we adopted a conservative approach and calculated total body nitrogen by estimating crude protein within each species and age/sex class (gralloched mass × 0.22) and then converting crude protein to nitrogen content (crude protein × 0.16).

Bone accounts for 89% of deer body phosphorus, while muscles and soft tissues account for 9% (McCullough & Ullrey, 1983). Modelling of red deer indicates that phosphorus in bone and soft tissue makes up 1.47% of body mass, excluding antlers (Flueck, 2009), with comparable values reported for sheep (Crisp, 1966). Consequently, we calculated the total phosphorus within each deer species and age/sex class using live mass (live mass × 0.147). Antlers also contain large reservoirs of phosphorus, constituting 9.5% of caribou antler mass (Moen & Pastor, 1998) and 13.5% of red deer antler mass (Flueck, 2009). We applied the average of these values to each deer species' antler mass (antler mass × 0.115 × number of males culled).

Like phosphorus, calcium is an essential component of bone and antlers (Moen & Pastor, 1998). Chemical analyses of white-tailed deer revealed that 99.66% of the calcium in deer bodies is stored within bone (McCullough & Ullrey, 1983). Using parameters from McCullough and Ullrey (1983), which provided the dry mass of bone, the ash mass and the calcium content found in the ash, we calculated that calcium constitutes 2.45% of deer body mass (live mass × 0.245). Calcium also constitutes a large proportion of antler mass, constituting 19% of red deer antler mass (Moen & Pastor, 1998). We applied this value to each deer species' antler mass (antler mass × 0.19 × number of males culled).

2.4 Land classification

When deer culls are reported, hunters are asked to specify if the cull occurred in woodland, open range or agriculture (The Deer Working Group, 2019). Deer culling practices in Scotland are permitted equally across different land cover types, but with varying degrees of incentive and potential commercial motivations. As such, the intensity of culling efforts can vary based on the land use and management objectives. In woodland areas, culling is often aimed at preventing damage to trees and promoting successful reforestation efforts (Gill & Morgan, 2010; Reimoser & Putman, 2011). Similarly, in agricultural lands, culls may be more commercially driven, with an aim of protecting crops from deer browsing impacts (The Deer Working Group, 2019). Culling on open ground is often motivated by sporting and ecological considerations, such as maintaining a desired deer density and preventing overgrazing (Albon et al., 2017).

To estimate nutrient loss within the three land classifications (woodland, open range, agriculture) reported in statutory cull reports across Scotland, we first calculated the total area of each DCA using the raster package (Hijmans et al., 2015) in R (version 3.6.3, R Core Team, 2020). We then estimated the area of each land classification using the 2020 Scotland Habitat and Land Cover Map (Space Intelligence and NatureScot, 2021) at the national level and then within each DCA. This map categorizes land based on the European Nature Information System (EUNIS) with a resolution of 20 m × 20 m. Determining the appropriate EUNIS classifications corresponding to woodland and agriculture was straightforward, with easy analogues covering 16.8% and 22.6% of Scotland's total area, respectively (Table S2). However, determining the EUNIS classification analogue for the statutory cull report's open range classification required additional steps, as this land classification is harder to define, and we wanted to avoid simply classifying any areas not included in woodland or agriculture. We therefore used the ‘Deer Count Data’ from NatureScot (NatureScot, 2021), which reports points of deer sightings throughout the country. We extracted the EUNIS category of each reported deer sighting location, and after excluding sightings in forest or agriculture, our analysis revealed that most deer were located in the EUNIS category of ‘temperate shrub heathland’—a land classification covering 16.4% of Scotland's total area. Therefore, we used this ENUIS classification to represent the open range. All three land classifications accounted for 54.8% of total land in Scotland (Table S2).

2.5 Environmental inputs and outputs

To contextualize the loss of nutrients from deer carcasses, we gathered data from the literature detailing other significant inputs and outputs of nitrogen, phosphorus and calcium within Scotland. For nitrogen, the primary input across all ecosystems is atmospheric nitrogen deposition. Using previously published data (Tomlinson et al., 2021), we calculated atmospheric nitrogen deposition within each DCA. Phosphorus, on the other hand, is relatively conserved within ecosystems. Therefore, we compared losses from deer carcass removal to the loss of phosphorus from other resource extraction, such as wool, timber and deer carcass removal from other locations, as reported in the literature. Calcium enters ecosystems primarily through weathering and atmospheric deposition, yet large quantities are removed through timber extraction (Reynolds & Stevens, 1998). Given Scotland's calcium-poor soils, we gathered data from the literature on both inputs from weathering and atmospheric deposition, as well as estimated losses from resource extraction, including timber (Lee et al., 1999; Reynolds & Stevens, 1998).

3 RESULTS

3.1 Deer cull data

From 2010 to 2021, 2,129,744 deer were culled in Scotland after correcting for underreporting. Roe deer comprised 54% of the total cull, followed by red deer at 39%, sika deer at 5% and fallow deer at only 2%. (Table 1). The majority of culls occurred in woodland areas (66%), where roe deer dominated, accounting for 71% of the woodland cull. Open range accounted for 28% of the cull, where red deer overwhelmingly dominated the culling efforts, comprising 86% of the open range cull. Agriculture accounted for 6% of the total cull, consisting primarily of roe deer (63% of the agriculture cull) and red deer (28% of the agriculture cull) (Table 1; Figure 1).

TABLE 1. Total number of each deer species culled by land classification between 2010 and 2021.

Deer culled (2010–2021)	Agriculture	Open range	Woodland	Total
Fallow	8889	3673	24,516	37,078
Red	39,829	509,711	287,182	836,722
Roe	88,362	68,415	986,975	1,143,752
Sika	3837	8048	100,307	112,192
Total	140,917	589,847	1,398,980	2,129,744

Details are in the caption following the image — **FIGURE 1**
Open in figure viewer PowerPoint

Number of deer species culled per land classification adjusted with the species-specific correction factors reported in the Deer Working Group Report (The Deer Working Group, 2019) to account for underreporting.

3.2 Nutrient loss across Scotland

Each year, removing deer carcasses from Scotland resulted in a net loss of 195,652 kg N year⁻¹, 152,834 kg P year⁻¹, and 251,188 kg Ca year⁻¹. Red deer culls contributed substantially more to nutrient loss on the national level compared to any other species (Table 2). This is unsurprising given that the species constituted a large portion of the overall culling efforts; however, the effect is amplified by their large body mass (Figures 2 and 3; Table 2). Despite having the smallest body mass of the four deer species, roe deer culls were the second largest source of nutrient loss at the national level due to the considerable culling efforts of this species, highlighting that even small-bodied species can significantly impact nutrient budgets when present in high numbers (Figure 2; Table 2). When accounting for all deer culled, the largest overall nutrient loss occurs in open range, followed by woodland and agriculture (Table S3).

TABLE 2. Annual nutrient loss from deer culls by species in Scotland.

Species	Kg N	Kg P	Kg Ca	Kg N ha⁻¹	Kg P ha⁻¹	Kg Ca ha⁻¹
Fallow	3597	2188	3555	3597	2188	3555
Red	136,538	111,027	185,222	136,538	111,027	185,222
Roe	47,280	34,382	54,113	47,280	34,382	54,113
Sika	8237	5237	8298	8237	5237	8298
Total	195,652	152,834	251,188	195,652	152,834	251,188

3.3 Nutrient loss within DCAs

Total nitrogen loss from carcass removal within DCAs ranged from 28.55 to 14,348.43 kg N year⁻¹, with an average of 2508.36 kg N year⁻¹. By area, agriculture had the largest loss, ranging from 2.63 × 10⁻⁴ to 0.96 kg N ha⁻¹ year⁻¹, with an average of 0.06 kg N ha⁻¹ year⁻¹ (Figure 4; Table S4). In open range, nitrogen loss ranged between 5.19 × 10⁻⁴ and 0.40 kg N ha⁻¹ year⁻¹, with an average of 0.07 kg N ha⁻¹ year⁻¹ (Figure 4; Table S4). In woodland, nitrogen loss ranged from 3.21 × 10⁻³ to 0.32 kg N ha⁻¹ year⁻¹, with an average of 0.08 kg N ha⁻¹ year⁻¹ (Figure 4; Table S4).

While our results indicate large quantities of nitrogen are removed from the landscape via deer carcass removal, when contextualized with other inputs and outputs, deer carcasses may have little effect on the overall ecosystem nitrogen balance. The average atmospheric nitrogen deposition within each DCA ranged from 2.61 to 12.21 kg N ha⁻¹ year⁻¹ with an average of 6.56 kg N ha⁻¹ year⁻¹. Therefore, within each DCA, the loss of nitrogen from carcass removal constituted a fraction of what is input from atmospheric deposition (0.03%–1.36%). Even when considering the finer-scale land classifications within each DCA, in most cases, nitrogen losses from carcasses were not substantial compared to atmospheric deposition (min = 3.39 × 10⁻³%, mean = 1.28%, max = 18.15% of atmospheric deposition, respectively; Table S5).

Total phosphorus loss from carcass removal within DCAs ranged from 23.56 to 11478.59 kg P year⁻¹, averaging 1959.41 kg P year⁻¹. Averaged across agriculture areas within DCAs, phosphorus loss ranged from 1.92 × 10⁻⁴ to 0.67 kg P ha⁻¹ year⁻¹, averaging 0.05 kg P ha⁻¹ year⁻¹ (Figure 4; Table S4). In open range, phosphorus loss ranged between 3.48 × 10⁻⁴ and 0.35 kg P ha⁻¹ year⁻¹, averaging 0.06 kg P ha⁻¹ year⁻¹ (Figure 4; Table S4). In woodland, culling resulted in a loss of 1.84 × 10⁻³–0.24 kg P ha⁻¹ year⁻¹, averaging 0.06 kg P ha⁻¹ year⁻¹ (Figure 4; Table S4).

While our results indicated that, on average, the loss of phosphorus within DCAs from deer culls was smaller than that from other resource exports found in the literature, DCAs that experienced relatively high deer culls had phosphorous losses of similar magnitude to those other resource exports. For example, in a Swiss national park, migratory red deer export 0.32 kg P ha⁻¹ year⁻¹ out of their summer range via over-winter mortalities in an adjacent winter ranging area (Flueck, 2009). Similarly, the export of sheep carcasses and wool in Scotland removed 0.23 kg P ha⁻¹ year⁻¹ from local pastures, requiring 16 kg P ha⁻¹ year⁻¹ of fertilization to maintain productivity (Haygarth et al., 1998). Importantly, both of these studies consider much smaller areas where animal use is concentrated. In commercial woodlands, timber harvests remove 0.08–1.75 kg P ha⁻¹ year⁻¹, and harvesters then add 15–30 kg P ha⁻¹ year⁻¹ of fertilizer to compensate for the loss (Flueck et al., 2011). Given these examples, if we were to consider a conservative threshold of 0.1 kg P ha⁻¹ year⁻¹ as the point at which phosphorous fertilizer might be required to offset phosphorus loss, our analysis suggests that woodlands in 13% of DCAs, open range in 19% of DCAs, and agriculture in 7% of DCAs experience losses from cull carcass removal that would require phosphorous fertilization. Of note, the majority of DCAs that experienced greater phosphorus losses are in the north and northwest regions of Scotland, but the phosphorus loss in East Loch Ericht also exceeded the fertilization threshold in all three of the land classifications.

Total calcium loss from carcass removal within DCAs ranged from 39.30 to 19,003.28 kg Ca year⁻¹, averaging 3220.36 kg Ca year⁻¹. Across agriculture areas within DCAs, calcium loss ranged from 3.01 × 10⁻⁴ to 1.11 kg Ca ha⁻¹ year⁻¹, with an average of 0.08 kg Ca ha⁻¹ year⁻¹ (Figure 4; Table S4). In open range, culls resulted in a calcium loss between 5.84 × 10⁻⁴ and 0.59 kg Ca ha⁻¹ year⁻¹, with an average of 0.09 kg Ca ha⁻¹ year⁻¹ (Figure 4; Table S4). In woodland, calcium loss ranged from 3.07 × 10⁻³ to 0.40 kg Ca ha⁻¹ year⁻¹, averaging 0.10 kg Ca ha⁻¹ year⁻¹ (Figure 4; Table S4).

Weathering in Scottish uplands accumulates approximately 0.2 kg Ca ha⁻¹ year⁻¹ (Reynolds & Stevens, 1998), while atmospheric deposition in most of Scotland is less than 0.80 kg Ca ha⁻¹ year⁻¹ (Lee et al., 1999). However, the resolution of atmospheric deposition is not resolved below this threshold, and therefore, this value serves as a maximum estimate. While the average rates of calcium loss from each land classification within the DCAs are relatively low (0.08–0.10 kg Ca ha⁻¹ year⁻¹), many DCAs experienced calcium losses on the same order of magnitude as weathering and atmospheric inputs. Specifically, open range in 38% of DCAs, agriculture in 14% of DCAs, and woodland in 38% of DCAs experienced losses greater than 0.1 kg Ca ha⁻¹ year⁻¹. Furthermore, previous modelling work indicates that sheep carcass and wool removal resulted in a loss of only 0.019 kg Ca ha⁻¹ year⁻¹ (Crisp, 1966). Therefore, in most cases, deer carcass removal strips the land of more calcium than sheep farming. West Loch Shiel, East Loch Ericht, Scarba and Rum exhibited exceptionally high rates of calcium loss in agricultural areas, with 0.81, 1.11, 0.42, and 0.52 kg Ca ha⁻¹ year⁻¹, respectively.

Notably, several DCAs experienced substantial nutrient loss across land classifications: East Loch Ericht and Rum lost over 0.10 kg N, Ca and P ha⁻¹ year⁻¹ in both woodlands and agriculture. The DCAs with the highest nutrient loss were again mainly in the northwest and west coast regions of Scotland, areas characterized by less calcium-rich soils (Paterson et al., 2011).

4 DISCUSSION

Our results reveal that removing deer carcasses represents an overlooked anthropogenic impact on biogeochemical cycles (Abraham et al., 2023), actively stripping vital nutrients from landscapes across Scotland. Notably, the landscapes within Scotland are already considered nutrient-poor (Dobbie et al., 2011). To combat this, much of Scotland receives fertilizer treatments. Forests receive phosphorus and, to a lesser extent, nitrogen fertilization (Payne et al., 2018; Shah & Nisbet, 2019), while agricultural areas regularly receive nitrogen and phosphorus fertilization (Domburg et al., 2000). Additionally, lime application is employed nationwide as a strategy to address soil acidity, although this practice is experiencing a declining trend (Holland et al., 2018). Our results suggest that the removal of deer carcasses exacerbates nutrient limitations and may require fertilization to release this limitation.

Nitrogen and phosphorus limitations are common in northern systems (Carline et al., 2005; Du et al., 2020). Our results demonstrate that the removal of carcasses strips landscapes of both elements, though the loss of nitrogen from carcass removal is a fraction of that received from atmospheric deposition. However, regardless of the nitrogen loss relative to atmospheric deposition, the removal of carcasses likely exacerbates existing nitrogen constraints, given the consistent application of nitrogen fertilizers in these regions. The effect of carcass removal on phosphorus stocks, however, is much larger, which may require fertilization to offset.

Our results indicate that out of the three nutrients investigated, deer carcass removal may have the largest consequences for calcium cycles. Calcium plays an important role in each of the land classifications. Agricultural areas for crops and pasture in Scotland have a history of liming, a practice required to combat low calcium levels and high pH (Holland et al., 2018). In open range, acidic soils with low calcium levels already limit plant communities and impede natural succession (Kleijn et al., 2008). Further depletion from carcass removal could exacerbate these effects. Woodlands are recognized as important reservoirs of calcium, but the sustainability of harvesting forest products has raised concerns about increased nutrient losses associated with harvesting (de Oliveira Garcia et al., 2018; Vos et al., 2023). Low calcium soils can hinder regeneration efforts in both commercial and native woodlands (Federer et al., 1989; Huntington et al., 2000), and removing deer carcasses can exacerbate calcium loss further. For example, whole-tree harvest from 50 years of pine growth results in a net loss of 79 kg Ca ha⁻¹ year⁻¹ (Reynolds & Stevens, 1998). Over the same period, in 11% of DCAs, culling deer would result in losses of >10 kg Ca ha⁻¹ year⁻¹ in woodlands. This is especially important when considering that calcium-poor systems could have cascading effects throughout the UK. For example, passerine birds experience altered eggshell characteristics, such as shell thickness in landscapes with a scarcity of calcium (Gosler & Wilkin, 2017), highlighting the need to consider the ecological consequences of altered nutrient cycles within systems and across trophic scales. Our findings suggest that deer carcass removal may contribute to reduced calcium availability, affecting not only plants but also key components of terrestrial food webs.

Our estimates likely underestimate actual losses. This is partly because the statutory cull data does not account for all instances of deer culling in Scotland, or for all deer deaths. For example, several thousand deer are killed annually in vehicle collisions (The Deer Working Group, 2019). Additionally, due to the available data, our approach evenly distributes nutrient losses across land classifications without considering spatial variations in deer habitat-use or culls. In reality, deer habitat-use and culls are concentrated in hotspots such as specific forest patches, farms, conservation areas, and sporting estates. Our inability to account for this heterogeneity underscores an ongoing challenge in zoogeochemistry: while animals exert highly localized effects on biogeochemical cycles, these effects are often quantified at broader landscape scales, resulting in missed insights (Ferraro et al., 2022). Consequently, the country-wide and land classification-specific estimates presented here offer an initial perspective on the potential magnitude of nutrient losses from removing culled deer carcasses across Scotland. However, they fail to capture the finer-scale dynamics within specific ecosystems and likely underestimate per-hectare losses in localized deer-inhabited areas. Fine-scale distributions of nutrient losses may be of particular concern if the management of that area is aimed at restoring native woodlands, increasing agricultural production, or maintaining biodiversity. More precise spatial data on cull locations would be necessary to comprehensively understand these localized impacts. Nonetheless, our findings accentuate the importance of considering deer culling practices within sustainable nutrient budget management. Given the potential for localized impacts, landowners could use their own site-specific cull data to develop case studies, thereby facilitating a more comprehensive understanding of the impacts on nutrient dynamics at finer spatial scales.

The use of publicly available data to provide a coarse-scale estimate of the loss of nutrients from carcass removal across Scotland comes with several other limitations. First, we were unable to assess the effects of carcass removal on ecosystems. To empirically test the effects of missing carcasses on ecosystem dynamics and conservation strategies, future studies could use carcass decomposition experiments or conduct fine-scale analyses of environmental characteristics along varying culling intensities. Additionally, while we are able to point to when losses of nutrients from carcass removal may require fertilization, we are unable to predict thresholds at which nutrient losses may cause state shifts or changes to ecosystem functions. Further work on these thresholds is needed within Scottish systems to better forecast ecosystem changes.

However, looking at other carcass studies, the ecosystem contributions lost due to the removal of deer carcasses can be predicted. Large mammal carcasses create vital nutrient hotspots on the landscape (Bump, Peterson, & Vucetich, 2009; Ferraro et al., 2022), which can have lasting effects on nutrient cycling (Danell et al., 2002). For example, soil calcium concentrations remained elevated near European bison carcasses for up to 7 years (Melis et al., 2007). Modelling work has demonstrated such effects scale to the landscape level, and without nutrients returning from carcasses, deer create nutritionally homogenous landscapes (Ferraro et al., 2022). Importantly for Scotland, promoting forested areas may require geochemical hotspots, as higher soil fertility can favour late-successional species over early-successional species (Marrs, 1993; Marrs & Gough, 1989). Yet the interaction between deer and wild predators plays an important role in how carcasses are distributed on the landscape, often mediating landscape heterogeneity (Bump, Peterson, & Vucetich, 2009; Johnson-Bice et al., 2023; Monk & Schmitz, 2021). Therefore, even if carcasses were purposefully left on the landscape by human hunters, human deposition of carcasses will likely not create the same levels of heterogeneity compared to wild predators, as human hunting is constrained by logistical factors such as access via roads/paths and visibility.

Carcasses have numerous other benefits for ecosystems as well, including directly increasing plant biodiversity after decomposition and increasing forage quality (Bump, Webster, et al., 2009; Danell et al., 2002). Within Scotland, deer carcasses can also serve as a source of carrion for mesopredators such as pine martens (Martes martes) (Balharry, 1993). In other systems during the breeding season for ground-nesting birds, carcass presence significantly reduces predation of clutches, and work to test this applicability for Capercaillie conservation has been promising as an employable method to reduce depredation (Bamber et al., 2023; Finne et al., 2019).

In January 2024, the Scottish Government launched the ‘Managing deer for climate and nature consultation’ to consider amendments to the Deer (Scotland) Act 1996 in light of the biodiversity and climate crises. While there is evidence that deer can have direct effects on climate mitigation strategies, such as carbon sequestration (Hirst, 2021), it is important to recognize the broader impacts that deer management strategies could have. The removal of deer carcasses and the resulting nutrient loss in ecosystems may reduce plant productivity. Such consequences could undermine ongoing ecosystem restoration, thereby compromising carbon sequestration goals. As deer management practices intensify across Scotland, it is important to fully evaluate the potential effects of wildlife management on a large scale.

5 CONCLUSIONS

Our results provide a coarse understanding of the nutrient loss due to deer culls throughout Scotland. Although it is difficult to contextualize the scale of this loss, our analysis shows that removing deer carcasses is gradually stripping vital nutrients from landscapes. Given the scale of deer culled in Scotland and current ambitions by landowners and the Government to increase deer management efforts, over time, the continued removal of deer carcasses could undermine Scotland's habitat recovery goals as these losses represent un-accounted depletions of critical nutrients required to support ecosystem function, plant growth and wildlife populations.

AUTHOR CONTRIBUTIONS

Kristy M. Ferraro and Chris Hirst conceived the ideas. Kristy M. Ferraro designed the methodology and conducted data analysis. Kristy M. Ferraro and Chris Hirst wrote the manuscript. All authors contributed critically to the drafts and gave final approval for publication.

ACKNOWLEDGEMENTS

We thank NatureScot and Space Intelligence for their help gathering the data needed for this project. We also thank L. Seivwright for her insightful conversations. This work was supported by the National Science Foundation Graduate Research Fellowship (DGE-1752134), awarded to KF. CH received grants from the Natural Environment Research Council (NE/S007407/1), the British Deer Society, and Forest Research while conducting this work. Any opinions, findings, and conclusions or recommendations expressed in this material are those of the authors(s) and do not necessarily reflect the views of the National Science Foundation.

CONFLICT OF INTEREST STATEMENT

The authors declare no conflicts of interest.

Open Research

DATA AVAILABILITY STATEMENT

Data on deer culls were retrieved from The Deer Working Group (2019), data on deer habitat use was retrieved from ‘Deer Count Data’ from NatureScot (NatureScot, 2021), data on atmospheric deposition was retrieved from Tomlinson et al. (2021) and land classification through the country was retrieved from the Scotland Habitat and Land Cover Map from NatureScot and Space Intelligence (2021). Code is available in the data repository https://github.com/kristymferraro/MissingDeerCarcassesScotland.git (Ferraro, 2024).

Supporting Information

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Citing Literature

Volume5, Issue3

July–September 2024

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Georgina Mace Prize 2024

Missing carcasses, lost nutrients: Quantifying nutrient losses from deer culling practices in Scotland

Abstract

1 INTRODUCTION