Land use influence on chacma baboon (Papio ursinus) diet in South Africa using stable isotopes

Land use influence on chacma baboon (Papio ursinus) diet in South Africa using stable isotopes | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Land use influence on chacma baboon (Papio ursinus) diet in South Africa using stable isotopes Elizabeth Kennedy Overton, Alice Bernard, Pierre-Cyril Renaud, and 5 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-3427689/v1 This work is licensed under a CC BY 4.0 License Status: Posted Version 1 posted You are reading this latest preprint version Abstract Anthropization processes affect wildlife feeding behaviours due to changes in resource availability related to land use and land cover change. To better understand the ecological responses of wildlife towards anthropogenic change, it is essential to evaluate whether human land use, characterized by high human-modified food availability, has an impact on wild animal feeding ecology. The chacma baboon ( Papio ursinus ) is interesting to study potential diet changes as it is largely present along a gradient of anthropized areas in Southern Africa. In this study, fecal samples from chacma baboon troops were collected in different land use habitats (peri-urban, agricultural and natural forest habitat) in the Garden Route, South Africa, and their isotopic ratios of carbon (δ 13 C) and nitrogen (δ 15 N) measured. Results showed significant differences between δ 15 N ratios according to land use, indicating significant higher protein intake in areas with human influence in comparison to natural forest habitats. Furthermore, the large majority of the collected samples were contained within the bracket that reflect the C 3 ecosystem of the Garden Route region, with the exception of some samples showing higher δ 13 C ratios associated with the consumption of anthropogenic foods (containing sugar, corn and wheat). The potential protein increase, as well as sources of C 4 plants present in the diets in anthropized areas suggests a visible dietary shift for this species between natural and transformed landscapes. In the future, it will be essential to determine whether and how the consumption of human-modified food could affect the health and associated fitness of chacma baboons. Carbon Nitrogen Anthropization Habitat modification Food availability Garden Route Figures Figure 1 Figure 2 Figure 3 Figure 4 Introduction The transformation of natural landscapes by human activities (Martínez-Fernández et al. 2015 ) has modified over half of the Earth’s land surface (Foley et al. 2005 ; Hooke et al. 2012). Land use and land cover change (LULCC), including extensive clearing, the creation of pastures, croplands and urban settlements (Fahrig 2003 ; McKinney 2002 ) are the main drivers impacting ecosystems and causing drastic global and regional biodiversity collapse (Ellis et al. 2021 ; Foley et al. 2005 ; Sun et al. 2021 ; Turner et al. 2007 ; Zhang et al. 2007 ). For certain species, these transformed landscapes can create new niches and provide wild animals with other resources (Dale and Polasky, 2007 ; Eriksson 2013 ; Fleming and Bateman 2018 ). In anthropized landscapes, the possible access to cultivated zones, gardens, kitchens and garbage has provided new opportunities for wildlife (Hulme-Beaman et al. 2016 ; Oro et al. 2013 ). Certain species have adapted or acclimated to these transformed areas, exploiting anthropogenic resources (Hulme-Beaman et al. 2016 ; Seoraj-Pillai and Pillay 2016 ; Webber 2017 ). For instance, wild boar ( Sus scrofa ) and vervet monkeys ( Cercopithecus aethiops ) are known to exploit croplands (Cancelliere et al. 2018 ; Lee and Lee 2019 ), carnivores such as San Joaquin kit foxes ( Vulpes macrotis mutica ) or coyotes ( Canis latrans ) exploit urban resources (e.g. garbage or compost; Newsome et al. 2010 ; Sugden et al. 2021 ) and commensal behaviour, notably as household and agricultural pests, has been observed in black rats ( Rattus rattus ) across different geographical areas on several occasions (Aplin et al. 2011 ). These changes cause direct modifications in wildlife diet, from the consumption of purely natural resources to the integration of more human-originated food (Murray et al. 2015 ). This can be a benefit to species, providing year-round food provisioning, reducing food and water deprivation during droughts or winters, or generally contributing to better individual condition. For example, a population of urban kit foxes presented higher cholesterol levels, a good indicator of energy intake (Harder and Kirk-patrick 1994), as well as reduced nutritional deprivation than rural ones in California (Cypher and Frost 1999 ). The integration of human resources into the diet of storks during periods of constrained resource availability has also been shown to lead to higher reproduction rates (Evans and Gawlik 2020 ). As another example, crop raiding behaviour in male elephants resulted in larger body size in adulthood (Chiyo et al. 2011 ), an important trait for this species during mating (Chelliah and Sukumar 2013). Despite certain benefits however, the consumption of human food has also been shown to have a negative impact on wildlife health. For instance, urban coyotes generally showed poorer average body condition as well as higher parasitic infection rates than rural coyotes in a study by Sugden et al. ( 2020 ). Reduced reproductive rates of birds (e.g. blue tits) are also associated with urban areas compared to populations in rural zones, linked to a lack of suitable diets for nestlings (Pollock et al. 2017 ; Seress et al. 2020 ). The impacts of human resources on non-human primates appear to a lesser extent in the literature, even though primates often consume human resources (Strum 2010 ). It has been shown that wild primates obtain higher levels of essential nutrients relative to body weight than humans do through their diets (Milton 1999 ), but also that varying benefits can follow their consumption of anthropic food resources depending on the type of LULCC (Maibeche et al. 2015 ; Marty et al. 2020 ). An interesting primate species to better understand diet change in the presence of human resources is the chacma baboon ( Papio ursinus ), due to its adaptive capacity (Fischer et al. 2019 ). Baboons are opportunistic, omnivorous feeders that will eat anything from fruits, grasses, leaves or roots to invertebrates and even occasionally animal matter usually resulting from opportunistic hunting (Allan et al. 2022 ; Hoffman and O’Riain 2011 ; Schreier et al. 2019 ) and are highly adaptable to a wide range of habitats and environments (Johnson et al. 2015 ). They can be found across deserts, savannahs, forests and on the outskirts or within urban zones (Hoffman and O’Riain 2011 , 2012 ; Johnson et al. 2015 ; Mormile and Hill 2017 ). With the encroachment of urban areas on natural habitats, encounters between humans and baboons have become prevalent in some areas, such as in the Western Cape of South Africa (Bracken et al. 2022a , b , 2023 ; Chowdhury et al. 2020 ; Lee and Priston 2005 ). In the Western Cape, as well as across South Africa, chacma baboons frequently raid croplands as well as people’s gardens, bins or houses (Hoffman and O’Riain 2011 ; Strum 2010 ; Walton et al. 2021), consuming resources of human origin. When the availability of rubbish is reduced, this affects the baboon’s raiding tendencies: Mazué et al. ( 2023 ) showed that when the rubbish resources were removed on regular foraging sites in a peri-urban area on the George Campus of the Nelson Mandela University (NMU), chacma baboons spent more time foraging for natural resources and less time in these more urban areas. The easy access to fruits, vegetables, meats, but also processed foods, refined sugars, added fats and oils may result in significant variations in their nutrient consumption and have unknown effects on their metabolisms. In this context, the objective of this present study was to evaluate whether human land use, characterized by high human modified food availability, has an impact on chacma baboon diet. Several methods exist to study diets in ecology, including direct observation or fecal analysis. These techniques yield accurate results but they can vary according to what is consumed and the digestibility of the latter (Matthews et al. 2020 ): some resources will appear in higher proportions in fecal samples than others depending on how easy they are to digest. Direct observation also often requires long periods of time (Jordan 2005 ), whereas non-invasive methods based on fecal samples, such as DNA metabarcoding and/or isotopic analyses, can counter this problem (Ando et al. 2020 ; Crowley et al. 2016 ; De Barba et al. 2014 ; O’Brien 2015 ). In the case of metabarcoding however, sample analysis can be costly and a broad list of plant/animal DNA sequences that could be consumed in the region would be necessary (Taberlet et al. 2018 ). Analysing isotopic ratios is therefore an interesting alternative, both less expensive and less time-consuming than other options, which explains why it is a recurrent method used in feeding ecology studies (Boecklen et al. 2011 ; Martínez del Rio et al. 2009 ; Taki et al. 2017 ). Common isotopes used for diet studies include nitrogen ( 15 N/ 14 N) and carbon ( 13 C/ 12 C). Nitrogen isotopes are a potential biomarker of protein source because tissue nitrogen derives almost entirely from dietary protein (for a review of isotopes as biomarkers see O’Brien 2015 ). Due to a stepwise increase with the trophic chain, variations of the δ 15 N isotopic ratios observed within feces may reflect consumption of dietary items with higher protein content within the different LULCC based on what resources the baboons have access to. The carbon in plants derives from atmospheric CO 2 which is fixed during the process of photosynthesis. Differences in stable carbon isotope ratios therefore reflect the type of plants consumed based on their photosynthetic pathways (C 3 , C 4 or CAM) due to differential fractionation of carbon isotopes during photosynthesis that leads to distinct δ 13 C values (Ehleringer and Cerling 2002 ; Smith and Epsten 1971 ). Examining carbon and nitrogen ratios may therefore be a suitable method of assessing the potential contribution of human-derived foods to animal diet, comparing protein intake and the type of plants consumed between natural areas and those with anthropic resources (Penick et al. 2015 ). To address the objective of this study, we obtained stable isotope ratios from fecal samples of chacma baboons, which would provide quantitative data on the types of plants and animal protein they consume. Baboons may be considered as opportunistic feeders (Allan et al. 2022 ; Schreier et al. 2019 ) and therefore the potential access to high protein sources such as meat and legumes in more anthropized areas (with crops, garbage or kitchens) may result in higher proportions of protein in their diet. We could expect that if the anthropized landscapes provide higher protein contents than the natural resources consumed by chacma baboons, the stable nitrogen isotope values would tend to be higher in these transformed landscapes. More positive δ 13 C values for samples from anthropized areas may be associated with a higher consumption of C 4 plants that pervade human-derived foods (e.g. maize, sugarcane) as well as lipids when these types of resources are included in chacma baboon diets (Post et al. 2007 ). We could then expect to observe more positive δ 13 C ratios from samples collected in more urbanized environments compared to natural landscapes. To evaluate whether transformed landscapes have an influence on chacma baboon diet, we collected fecal samples from chacma baboons along a gradient of land use in the Garden Route, South Africa, and studied stable isotopic ratio data, incorporating the surrounding landscape characteristics. Material and methods Study area The study was conducted in the Western Cape, South Africa, across the Garden Route Biosphere Reserve (listed in the UNESCO’s World Network of Biosphere Reserves; Pool-Stanvliet and Coetzer 2020 ) between George and Knysna, representing an area of 2000km² (Fig. 1a). The reserve is known for its unique and endemic biodiversity, such as afrotemperate forests, fynbos, as well as wetlands, mountains and coastal ecosystems. However, human presence has largely transformed the landscape in many places (Socio-Economic Profile: Garden Route District Municipality 2021), providing agricultural, forestry and touristic activities as well as the development of many urban settlements (e.g. Mossel Bay, George, Knysna). To protect the area, the Garden Route biosphere reserve, with the Garden Route National Park (GRNP) as a core area was created, preserving natural areas where biodiversity can prosper. Due to the complex landscape configuration that includes agriculture, logging, urban areas as well as natural vegetation, this IUCN type II protected area is very patchy and fragmented. Certain species, such as chacma baboons, can be found across all these different ecosystems, populating protected areas, rural zones as well as highly urbanized regions (Mormile and Hill 2017 ; Slater et al. 2018 ). In the study area, three main contrasted locations were selected, a peri-urban, an agricultural and a protected area, hereafter referred to as Campus, Hoekwil and Knysna throughout this study (represented in red, purple and blue respectively in Fig. 1a). These three sampling areas represent different land use types. The first location (Campus) is located on the Nelson Mandela University Campus near the city of George, where there are many people, buildings (e.g. accommodation and cafeterias) and lawns, and is surrounded by natural forest and fynbos. The second location (Hoekwil) was situated in the agricultural area bordering the Garden Route National Park to the North of the Town named Hoekwil. This area mainly consisted of livestock pastures and croplands, with occasional buildings (accommodation and infrastructure). In the region, summer crops can include lucerne, maize and vegetables (CapeFarmMapper v 2.7). The third location (Knysna) was situated within the afrotemperate forest of the Knysna Section of the Garden Route National Park (GRNP). Two subsites were established for each of these three main locations as replicates to account for troop preferences and intra-troop variation: Main and North for Campus (respectively light and dark red; Fig. 1b), Oakhurst and Woodville for Hoekwil (light and dark purple; Fig. 1b), Goudveld and Diepwalle sections of the GRNP for Knysna (light and dark blue; Fig. 1b). The two replicates for Campus represent two distinct troops, whereas the replicates for Hoekwil and Knysna were made up of one or more distinct troops each, and each replicate had varying proportions of land cover including within each of the three sites. Chacma baboon fecal sampling Sampling took place between the 14th of February and the 1st of April 2022. fecal samples from baboons were collected during regular visits to each of the three locations over this period. Each location was visited two to three times a week. Sample collection was opportunistic, resulting in different sample sizes for both temporal and spatial replicates. At the Campus, samples were collected on foot by retracing where the Main and North troops had been sighted by students or staff. At Hoekwil, footpaths and fences where the baboons were often sighted were scoured on foot. In the Knysna forest, road transects were followed by car in search of samples. Each sample was collected using gloves and inserted into plastic tubes or bags. At each sampling point the collection date, latitude, longitude, freshness of feces, location and subsite names were noted (SI 1). Samples were kept in a cooler box during the fieldwork each day before being frozen and stored at -20°C at NMU REHABS Laboratory until further analysis. Data collection and analyses were authorized by permits from SANParks (BERN-A/2020-008) and Cape Nature (CN44-87-16198), and the Section 20 of the Animal Diseases Act, 1984 (Act No 35 of 1984). Landscape characteristics surrounding each collected fecal sample The study area (latitude: 630900–725500, longitude: 6214900–6263600) was represented using QGIS (3.18), under a WGS 84 – UTM 34S projection. It was characterized according to several landscape metrics: land cover, slope, elevation and length/number of roads. In order to characterize the landscape metrics in the area used by the baboons for feeding, we recorded them within a circle of 4km² defined around each fecal sample position, based on the highest daily distance covered by chacma baboons in the literature in South Africa (Hoffman and O’Riain 2011 ; Slater et al. 2018 ) and the observations of the baboon troops on NMU Campus in George by Maud Czerwinski et al. (unpubl. Manuscript). The land cover raster layer was obtained from the South African National Land Cover Datasets (2018) of the South African Forestry, Fisheries and the Environment government website (DEFE, 2021) and clipped to the study area. The 73 classes from this layer were grouped into 10 (Fig. 1a) for simplification according to main types ( i.e . natural forest, plantations, shrubland, grassland, waterbodies, wetland, barren land, cultivated land, built up and mines). The elevation raster at 30 meters was downloaded from SRTM products (Farr and Kobrick 2000 , Shuttle Radar Topography Mission) and slopes were calculated within QGIS. The roads vector layer covering South Africa was obtained from the website of the Princeton University Library (International Steering Committee for Global Mapping, map issued in 2016). The percentage of land cover, as well as mean slope, elevation, road length and number of road segments was calculated within the buffer of each sample using QGIS (3.18). Finally, a principal component analysis (PCA) was performed considering these variables after removing those with a correlation higher than 0.8, to visualize the land use characteristics surrounding each sample. Stable isotope analysis for chacma baboon diet characterization After being placed in a fridge for 24 hours to defrost, up to 10g of each sample used for the isotopic analyses was inserted into 20mL tubes and dried for 48 hours in an oven at 60°C at the NMU REHABS laboratory (following steps used by Caut et al. 2008 ; Codron et al. 2007 ; De Carvalho et al. 2019 ). These samples were then transported to the University of Pretoria where they were ground using a mortar and pestle and passed through a ½ mm sieve. The homogenized fecal samples were weighed out in aliquots of 0.9 mg and placed in tin capsules, which were pre-cleaned in toluene to avoid any potential contamination. The isotopic ratios for carbon (C) and nitrogen (N) of these samples were analysed at the University of Pretoria, South Africa. Samples were combusted at 1,020°C using an elemental analyser (Flash EA 1112 Series) coupled to a Delta V Plus stable light isotope ratio mass spectrometer via a ConFlo IV system (all equipment supplied by Thermo Fisher, Bremen, Germany), housed at the University of Pretoria Stable Isotope Laboratory, Mammal Research Institute. Several samples were run in duplicate (every 13th sample) to verify the accuracy of the homogenization of the samples. Two laboratory running standards Merck Gel (δ 13 C = -20.26‰, δ 15 N=7.89‰, C%=41.28, N%=15.29) and DL-Valine (δ 13 C = -10.57‰, δ 15 N=-6.15‰, C%=55.50, N%=11.86) and a blank sample were run after every 11 unknown samples. The carbon and nitrogen ratios for the laboratory running (Merck and DL-Valine) standards were calibrated using the following PRIMARY standards: IAEA- CH-3 (Cellulose), IAEA-CH-6 (Sucrose), IAEA-CH-7 (Polyethylene foil), IAEA N-1 and IAEA N-2 (Ammonium sulfate), IAEA NO-3 (Potassium nitrate). All results were referenced to Vienna Pee-Dee Belemnite for carbon isotope values, and to air for nitrogen isotope values. Results are expressed in delta notation using a per mil scale using the standard equation (Coplen 2011, Rapid Comm, in Mass Spectrometry): δX(‰)=[(R sample -R standard )/R standard -1] where X = 15 N or 13 C and R represents 15 N/ 14 N or 13 C/ 12 C respectively. Statistical analysis of diet variations Statistical analyses were conducted using R (R Core Team 2020, version 3.6.3.) in RStudio (version 1.3.1093) to analyse and compare the isotopic patterns (δ 15 N and δ 13 C) of chacma baboon diets in relation to land use over the three locations (Campus, Hoekwil, Knysna). Firstly, the data for each stable isotope ratio were summarized as mean and standard deviation values according to location and subsite. After carrying out Shapiro-Wilk and Levene tests, non-parametric Kruskal-Wallis tests were used to determine the significance of the location variable on the isotopic ratios of δ 13 C and δ 15 N followed by Dunn pairwise comparison tests to establish the significance of the differences between locations characterized by different landscapes and between the subsites of each location. A biplot was also created to visualize any variation in diet according to land use by comparing the δ 13 C and δ 15 N ratios by site. Finally, the differences between chacma baboon populations were tested across the gradient of anthropization created using the resulting axes of the PCA, using a linear model (LM) for the nitrogen and distribution plots for the carbon ratios. To establish if grouping the subsites within each location had an impact on the results, an inter-troop analysis was carried out focusing on the subsites of the Campus (Main and North). The comparison between these troops within site was possible due to the intimate knowledge of the troops using the area through regular monitoring and research projects. Following Shapiro-Wilk and Levene tests, a Student’s t-test was carried out on the Campus δ 15 N results and a Wilcoxon test for the δ 13 C. Results Sampling A total of 231 baboon fecal samples were collected throughout the sampling period (Fig. 1a; Table 1 ). Within the Campus, a total of 98 samples were collected, 60 for the Main and 38 for the North subsites, representing two different troops of baboons monitored on the NMU campus. At the Hoekwil site, 28 samples were found, with 14 for each subsite (Oakhurst and Woodville). In Knysna, 105 were collected in total, 67 of which for the Goudveld subsite and 38 for the Diepwalle subsite. We selected 148 samples for analysis in this study, according to their freshness and to obtain a similar number of replicates per location (Table 1 ; SI 1). A total of 60 samples were selected for both Campus and Knysna, with 30 samples per subsite. The Hoekwil samples were used in their totality due to the limitation of samples available linked to difficulties in finding samples in these areas, amounting to a total of 28 (14 per subsite). Table 1 Numbers of fecal samples collected, analyzed for land cover and analyzed for stable isotopes by location and subsite. The 146 samples chosen for the stable isotope analysis were chosen according to freshness and to obtain a representative number of replicates per location. Location Campus Hoekwil Knysna Subsite Main North Oakhurst Woodville Goudveld Diepwalle Number of collected samples 60 38 14 14 67 38 98 28 105 231 Number of samples analyzed for land cover 30 30 14 14 30 30 60 28 60 148 Number of samples analyzed for isotopes 29 29 14 14 30 30 58 28 60 146 Landscape and anthropization level characterization The PCA biplot created by combining the landscape variables used in this study revealed that the 148 samples from the three locations (Campus, Hoekwil, Knysna) were characterized by distinct land cover (Fig. 2a). Three clusters emerged within this figure, grouping together the samples from each studied location. The first axis, which explained 41.1% of the variation, opposed the flatter areas dominated by agriculture (including irrigation dams) mostly present in Hoekwil with the area to the steeper slopes dominated by natural forests and characteristic of the Knysna site. On the other hand, the second axis of this PCA, explaining 30.5% of the variation, opposed the built up location (Campus) with the more natural and less built up areas (Hoekwil and especially Knysna; Fig. 2a). A gradient, going from land rich with infrastructure (Campus) to natural environments without human settlements (Knysna) via an intermediate, agricultural area (Hoekwil), could be distinguished along the second axis (Fig. 2a). This axis was therefore defined as the gradient of anthropization of the environment for the following diet analyses of chacma baboons, with more negative values representing the more natural areas, more positive values representing more urban zones and intermediate values representing the agricultural areas. Stable isotope signatures and distributions according to the level of anthropization Out of the 148 fecal samples, 146 were analysed for δ 15 N and δ 13 C (SI 1) due to reduced quality of two samples. When combining the carbon and nitrogen ratios, the resulting biplot showed that the samples from the three locations had different nitrogen and carbon isotopic ratios (Fig. 2b). The widest range of isotopic ratios belonged to the Campus, the smallest to Knysna, with intermediate values for Hoekwil, resembling the Campus for its δ 15 N and Knysna for its δ 13 C (Fig. 2b). Low values of δ 15 N and δ 13 C tended to characterize the Knysna samples, going from − 0.02 to 3.12‰ for nitrogen and − 29.96 to -23.53‰ for carbon (Table 2 ). A wide range of δ 15 N (from 0.55 to 5.15‰) combined with low values of δ 13 C (from − 29.82 to -25.67‰) characterized those of Hoekwil (Table 2 ). A more intermediate profile characterized the Campus samples, yielding a wider range of δ 15 N (0.66 to 5.31‰) and wider ranges of δ 13 C (-29.26 to -17.92‰) than the other locations (Table 2 ). Table 2 Summary statistics (including minimum, maximum, mean and standard deviation SD) for isotopic ratios of Nitrogen and Carbon in the 146 analysed fecal samples by location and subsite. δ15N (‰) δ13C (‰) Location Min Max Mean SD Min Max Mean SD Campus 0.66 5.31 2.68 1.08 -29.26 -18.99 -26.38 1.84 Main 1.95 5.31 3.36 0.92 -29.09 -21.98 -26.43 1.42 North 0.66 3.70 1.97 0.71 -29.26 -18.99 -26.34 2.22 Hoekwil 0.55 5.15 2.55 1.21 -29.82 -25.67 -28.13 0.93 Oakhurst 2.24 5.15 3.34 0.72 -29.82 -27.54 -28.65 0.62 Woodville 0.55 4.29 1.75 1.08 -28.90 -25.67 -27.60 0.91 Knysna -0.02 3.12 1.58 0.69 -29.96 -24.24 -28.03 0.80 Goudveld 0.62 3.12 1.65 0.70 -29.96 -24.24 -27.92 0.91 Diepwalle -0.02 3.08 1.51 0.69 -29.72 -27.01 -28.14 0.67 A significant increase in the δ 15 N ratios was visible along with the level of anthropization of the environment across the land use gradient (R = 0.45, P-value = 1.4e − 08 ; Fig. 2c), created using the second axis of the PCA from Fig. 2a. The samples collected at the Campus (highest proportion of built up land cover), characterized by the highest anthropization levels, showed higher ratios of δ 15 N. The samples from Hoekwil (highest proportion of agriculture), with lower levels of built up land cover, presented intermediate δ 15 N numbers. The Knysna samples (highest proportion of forest), reflecting the lowest anthropization levels and more natural habitat, showed the lowest values for δ 15 N. The majority of the Campus samples, as well as all those from Hoekwil and Knysna, reflected an isotopic signature between about − 30‰ and − 24‰ for δ 13 C values (specific to C 3 plants), except four of the Campus samples that possessed slightly higher values (Fig. 2d). Higher values correspond to the more urban areas. The data for both δ 15 N and δ 13 C were found to be non-parametric (Shapiro-Wilk, p-value = 3.89e-4 for δ 15 N and 1.50e-12 for δ 13 C) with unequal variances between the three locations (Levene, Pr(> F) = 7.59e − 4 for δ 15 N and 1.01e − 2 for δ 13 C). Kruskal-Wallis tests indicated that at least one location was significantly different from the others for nitrogen (chi-squared = 32.97, df = 2, p-value = 6.92e − 08 ) and carbon (chi-squared = 57.92, df = 2, p-value = 2.65e − 13 ). Post-hoc Dunn tests revealed that the Knysna samples significantly differed from both the Campus (p-value = 1.04e-07) and Hoekwil (p-value = 4.77e − 04 ) samples for nitrogen (Fig. 3a), whereas for carbon ratios, the Campus samples were significantly different from both Hoekwil (p-value = 8.88e − 09 ) and Knysna (p-value = 3.28e − 11 ; Fig. 3b). According to these tests, nitrogen ratios did not significantly differ between Hoekwil and Campus samples (Dunn, p-value = 4.53e − 01 ), nor did Knysna and Hoekwil for carbon samples (Dunn, p-value = 6.77e − 01 ). Isotopic distribution variations between troops at the Campus site The resulting PCA biplot for the inter-troop analysis on the two subsites of the Campus (Main and North troops) showed little difference in distribution according to land use. The landscape surrounding the samples only differed slightly along the first axis according to the troop, which accounted for 70.9% of the variation (Fig. 4a). The Main troop samples tended towards higher levels of built up land use as well as the presence of roads, whereas the North troop spanned a wider range across the axis, going from built up to more natural landscapes (Fig. 4a). Despite these limited differences in the landscapes used by these two troops of baboons, δ 15 N nitrogen ratios of the fecal samples from these two subsites varied significantly (Student’s t-test, t = 6.50, df = 57, p-value = 1.09e-08) and according to the level of land use created using the first axis of the inter-troop PCA biplot (Fig. 4a and Fig. 4b). Higher δ 15 N values were associated with the Main troop (1.95 to 5.31‰; Table 2 ), which is represented by built up land cover and higher anthropization levels. The North troop nitrogen ratios had a wider range, spanning from ratios similar to those of the Main samples down to lower values of δ 15 N along with the anthropization level and towards areas characterized by more natural forests (0.66 to 3.70‰; Table 2 ). The carbon δ 13 C ratios of these two subsites reflected a similar distribution to those of all three locations: no significant differences were present between the two troops, whose ratios were contained between − 24‰ and − 30‰ (Wilcoxon, W = 450, p-value = 0.83; Fig. 4c; SI 1). It is noticeable that the highest δ 13 C values of two subsite samples of the Campus were specific to the North troop (Fig. 4c). Discussion In South Africa, chacma baboons are highly present in anthropized landscapes, possibly most notably in the Western Cape province. In Cape Town, as well as across the Garden Route region, chacma baboons and humans often occupy overlapping habitats, where increased conflicts have been reported associated mostly to stealing food (Chowdhury et al. 2020 ; Mormile and Hill 2017 ). Hoffman and O’Rian showed that Cape peninsula baboon troops present in human-modified habitats are associated with increased events of observed stealing from people’s houses and visiting the bins around settlements (Hoffman and O’Riain 2012 ). Mazue et al. also described that the Main troop living on the campus of Nelson Mandela University (George Campus), representing a peri-urban area, was responsible for an unexpected number of raids: 70 raids were recorded within a two-week study (unpublished data, 2016). Overall, it appears that the fragmentation and the modification of their natural habitats has pushed chacma baboons to co-exist with humans within modified landscapes, where new human-modified food resources are easily available, which could have an impact on their foraging behaviour. In the present study, the analysis of nitrogen and carbon stable isotopic ratios in chacma baboon troops inhabiting a gradient of human-modified landscapes in the Garden Route region showed significant differences in their diet in terms of protein and plant intake. A shift towards higher protein intake in anthropized areas It is well known that 15 N/ 14 N ratios in mammalian excrements are related to a combination of variables, driven by the complexity of the nitrogen cycle (Handley and Raven 1992 ). The latter leads to a variety of δ 15 N patterns in soil and plants that are consumed by animals higher up the trophic chain. When consumed, 15 N is preferably incorporated into tissues over 14 N, creating a stepwise increase with each trophic level and also increasing with protein intake (Handley and Raven 1992 ; Schoeninger and DeNiro 1984 ; Sponheimer et al. 2003 ). In the present study, the 15 N/ 14 N results showed significantly higher protein intake for chacma baboons in landscapes transformed by human activity, i.e . on the Campus with built up land cover and in Hoekwil with agricultural zones (Fig. 2c and 3a). This suggests that the chacma baboon populations known to live in close contact with human populations in the Garden Route have been shifting their diet to a richer nitrogen pattern when inhabiting more anthropized lands. This is the case whether agricultural or peri-urban, a result probably associated with the easier access to human-modified resources. Such results could be explained by the easy access to protein sources in such areas, such as garbage bins, meat detritus from the cafeteria on the Campus site or the houses and farmsteads around the Hoekwil area. Particularly on Campus, the kitchens and bins provide regular food for the baboons almost all year round, including remaining meals from students that contain large amounts of meat that would not be accessible to them in the wild. This observation was even confirmed when considering the smaller scale of the Campus site, where higher values of δ 15 N were observed to be associated with the Main troop, in comparison to the North one (Fig. 4b). This chacma baboon Main troop is followed daily by monitors and has been reported to spend considerable time on George Campus where they regularly visit buildings and bins (Mazué et al. 2023 ), which could again explain the differences observed between the Main and the North troops at the Campus location regarding nitrogen intake. This study showed that the nitrogen ratios in the fecal samples, and therefore protein intake, were significantly highest in the populations inhabiting the most anthropized areas where the most buildings, built up land cover and transformed landscape for agriculture can be observed and where a higher access to human modified food highly composed of meat is present. Some studies have also described higher protein intake by wild animals inhabiting human-modified environments, such as rats, that have been shown to consume more protein-rich resources in anthropized areas (Guiry and Buckley 2018 ). While these studies showed the same pattern and confirmed that certain wild animal populations inhabiting anthropogenic landscapes present higher protein intake in comparison to natural ones (Guiry and Buckley 2018 ), others described different patterns with either lower nitrogen intake in anthropized environments or no significant difference in relation to landscape use. Some birds and carnivorous mammal species such as foxes or coyotes have been found to possess protein-poor diets in anthropized areas in comparison to populations in rural zones (Francis et al. 2021 ; Heiss et al. 2009 ; Murray et al. 2015 ; Newsome et al. 2010 ; Scholz et al. 2020 ). Moreover, a study from Nabugabo Field Station in central Uganda showed that for vervet monkeys consuming crops, there were no resulting variations in protein, fibre or lipid composition in comparison to wild populations (Cancelliere et al. 2018 ). The same pattern has been observed for wild boars, for which similar protein levels have been described across a gradient of modified landscapes, even though the species has been proven to be present near human settlements (Cahill et al. 2012 ; Stillfried et al. 2017 ). The work from Gámez et al. (2020) also evidenced that overall, δ 15 N ratios did not differ between urban and rural or wild land predators suggesting that the protein intake was not influenced by landscape modifications. It then appears that depending on the species studied and probably the geographic location of the study, wild animals respond differently to human modified resources. In the present study, the results reveal a significant shift toward higher protein intake in chacma baboons’ diet inhabiting anthropized areas in South Africa, but this will need to be confirmed through replicates along the same landscape gradient in multiple troops. Variations in plant consumption patterns across anthropized areas It is well established that C 3 plants tend to have lower 13 C/ 12 C ratios than C 4 plants: the δ 13 C values of C 3 plants range from − 34 to -22‰ and those of C 4 plants from − 16 to -10‰ (Basu et al. 2015 ). In the Garden Route temperate region, it has been shown that most trees and shrubs follow the C 3 -photosynthetic pathway (Ehleringer and Cerling 2002 ), so we could expect chacma baboon’s diet living in natural forest to be richer in C 3 than in C 4 plants. The results obtained in this study showed that the large majority of the collected samples were contained within the bracket that reflect a C 3 ecosystem, suggesting that as expected the overall consumption of the chacma baboon troops studied was made up of C 3 plants (Fig. 2d). However, small variations have to be considered regarding the δ 13 C isotopic ratios, as some samples collected at the highly anthropized Campus location possessed a higher proportion of C 4 plants than those from Hoekwil and Knysna, suggesting more consumption of C 4 components during recent foraging by certain individuals (Fig. 2d). This could be associated with the fact that higher δ 13 C isotopic ratios can be explained by the consumption of anthropogenic foods that are known to contain large quantities of C 4 plants ( i.e . sugar, corn and wheat; Gámez et al. 2020; Nicholson and Cove 2022 ; Post et al. 2007 ). The same pattern has been observed for instance in eurasian red squirrels ( Sciurus vulgaris ) that consumed more sugar in urban zones (Wist et al. 2022 ). In the present study, the overall consumption of the chacma baboon troops studied was made up of C 3 plants with for a significantly higher δ 13 C ratios in the Campus suggesting that the troops in this area may have access to and consume a higher proportion of C 4 plants ( i.e . sugar, corn and wheat), presumably due to the access to garbage and kitchens, as described previously for nitrogen. At a smaller scale, a few samples from the most anthropized subsites (on Campus) particularly stood out, possessing higher δ 13 C values, which could be as discussed above the result of higher C 4 plant consumption by specific individuals (Fig. 4c). This raises the possible influence of other factors including social behaviour that could contribute to variations in diet in primates that can influence carbon intake. Indeed, adult males of several macaque species have been shown to consume more human resources than adult females, younger and lower ranked individuals (Maibeche et al. 2015 , Marty et al. 2020 ). Focusing management ( i.e. monitors) on certain high ranking individuals may be helpful to reduce their consumption of human resources, but can also provide lower ranking individuals with easier access to these resources (Bracken et al. 2022b ), thereby failing to reduce conflict with humans. To address such questions, future studies should integrate stable isotope analyses with complementary behavioural analyses to get a better understanding of the chacma baboons’ behavioural ecology and to establish better strategies for conflict management. Perspectives and future impacts This present study showed that land use influences chacma baboons’ diet, with a shift in anthropized environments toward a globally significant higher nitrogen and for some individuals higher carbon patterns. This quantitative analysis gives a first picture of land use impact on this species diet, however we are still far from identifying exactly what chacma baboons consume in different environments. In this context, future studies should combine stable isotope analysis with observation and/or DNA metabarcoding in order to extract the precise composition and species richness of the baboon’s diet along multiple land uses. Moreover, seasonal variations can also account for differences in carbon ratios in baboon diet across the year: Lewis et al. ( 2018 ) provided results that showed that summer and autumn δ 13 C ratios were higher than those in winter and lowest in spring. In the future, studies should therefore also focus their efforts on the analysis of the stable isotopic ratios of larger sampling size and of fecal samples collected over several seasons to avoid missing any individual or seasonal effects (Mychajliw et al. 2022 ). This study has shown that chacma baboons consume more human resourced foods in more anthropogenic landscapes, and it has previously been described that despite some benefits, the shift in trophic niche associated with human-modified resources can also be detrimental to many wild animals. Unfortunately, to date health studies tend to revolve around humans and few have focused on the extent and direct impacts that the consumption of anthropogenic food by wild animals can have on their health. Several diseases found in humans (e.g. obesity, cardio-metabolic or cardiovascular risks, cancer, type-2 diabetes) are linked to the consumption of processed foods, refined sugars, certain grains or fats (Elizabeth et al. 2020 ; Monteiro et al. 2018 ; Popkin et al. 2020 ) that, according to this study, species such as chacma baboons have increasing access to. The question is now to determine what specific human-modified resources non-human primates are consuming in urban and peri-urban areas, and if this is causing health problems (e.g. short or long term diseases; Sapolsky and Share 2004 ) as well as affecting other aspects of their lives (i.e. behavioral, social, physiological, stress; Bracken et al. 2022b ; Chowdhury et al. 2020 ; Mazué et al. 2023 ). Understanding these effects is important for future management in the growing context of human-wildlife encounters and conflict. Declarations Statements & Declarations Supplementary Information The online version contains supplementary material available at XXX Acknowledgements We are grateful to SANParks for allowing and participating in data collection throughout this study. We would especially like to thank all SANParks personnel who provided us with not only access and guidance throughout the park, but who became personally involved in the research project, particularly Nathan and Dylan. We also thank the Wildlife Ecology Lab and the students that participated in data collection and analysis, namely Eloise Krisch, Sinayo Lungile, Brittany McClear, Tessa Raw, Markus Woesner, Bianca Coulson and especially the NMU Campus Baboon Monitors, Dietre Stols, Liyabona Ntlokwana, Nusheen Brenner and Abulele Sixonxo who provided us with remarkable insight into the Campus baboon habits. Thank you to all the people who provided details on troop locations throughout the Garden Route and granted us access to their properties, including the Oakhurst Farm and the personnel of the Woodville Trading Store. Funding This work was supported by the funding of Virginie Rougeron from Centre National de la Recherche Scientifique, REHABS International Research Laboratory. Data availability and material Ethics declarations Data collection was possible according to SANParks (BERN-A/2020-008) and Cape Nature (CN44-87-16198) permits, and analysis was allowed under the Section 20 of the Animal Diseases Act, 1984 (Act No 35 of 1984). Authorship statements Conception: V.R., F.P., H.F. and C.G. Funding acquisition: V.R. Biological data acquisition and management: V.R., F.P. E.K.O. and A.B. Stable isotopic data acquisition and analysis: E.K.O. G.I.S analysis : E.K.O. and P.C.R. Interpretation of results: E.K.O, V.R., F.P., H.F. and C.G. Drafting of the manuscript: E.K.O and V.R. Reviewing and editing the manuscript: E.K.O, V.R., F.P., H.F., P.C.R., C.G., A.B. and G.H References Allan ATL, LaBarge LR, Howlett C et al (2022) Patterns of predation and meat-eating by chacma baboons in an Afromontane environment. Folia Primatol 1–24. https://doi.org/10.1163/14219980-bja10004 Ando H, Mukai H, Komura T et al (2020) Methodological trends and perspectives of animal dietary studies by noninvasive fecal DNA metabarcoding. Environ DNA 2:391–406. https://doi.org/10.1002/edn3.117 Aplin KP, Suzuki H, Chinen AA et al (2011) Multiple Geographic Origins of Commensalism and Complex Dispersal History of Black Rats. PLoS ONE 6:e26357. https://doi.org/10.1371/journal.pone.0026357 Basu S, Agrawal S, Sanyal P et al (2015) Carbon isotopic ratios of modern C3–C4 plants from the Gangetic Plain, India and its implications to paleovegetational reconstruction. Palaeogeogr Palaeoclimatol Palaeoecol 440:22–32. https://doi.org/10.1016/j.palaeo.2015.08.012 Bellon B, Zander S, De Vos A, Renaud P-C CoexistLand. In: Earth Engine Apps. https://zandersamuel.users.earthengine.app/view/coexistland . Accessed 20 Apr 2022 Boecklen WJ, Yarnes CT, Cook BA, James AC (2011) On the Use of Stable Isotopes in Trophic Ecology. Annu Rev Ecol Evol Syst 42:411–440. https://doi.org/10.1146/annurev-ecolsys-102209-144726 Bracken AM, Christensen C, O’Riain MJ et al (2022a) Flexible group cohesion and coordination, but robust leader–follower roles, in a wild social primate using urban space. Proc R Soc B 289:20212141. https://doi.org/10.1098/rspb.2021.2141 Bracken AM, Christensen C, O’Riain MJ et al (2023) Postpartum cessation of urban space use by a female baboon living at the edge of the City of Cape Town. Ecol Evol 13:e9963. https://doi.org/10.1002/ece3.9963 Bracken AM, Christensen C, O’Riain MJ et al (2022b) Socioecology Explains Individual Variation in Urban Space Use in Response to Management in Cape Chacma Baboons (Papio ursinus). Int J Primatol 43:1159–1176. https://doi.org/10.1007/s10764-021-00247-x Burbank J, Drake DAR, Power M (2022) Use of stable isotopes for assessing urbanization impacts on freshwater fishes. Front Environ Sci 10:963693. https://doi.org/10.3389/fenvs.2022.963693 Cahill S, Llimona F et al (2012) Consorci del Parc Natural de la Serra de Collserola, Barcelona, Spain, Characteristics of wild boar (Sus scrofa) habituation to urban areas in the Collserola Natural Park (Barcelona) and comparison with other locations. Anim Biodiv Conserv 35:221–233. https://doi.org/10.32800/abc.2012.35.0221 Cancelliere EC, Chapman CA, Twinomugisha D, Rothman JM (2018) The nutritional value of feeding on crops: Diets of vervet monkeys in a humanized landscape. Afr J Ecol 56:160–167. https://doi.org/10.1111/aje.12496 Caut S, Angulo E, Courchamp F (2008) Discrimination factors (∆ 15 N and ∆ 13 C) in an omnivorous consumer: effect of diet isotopic ratio. Funct Ecol 22:255–263. https://doi.org/10.1111/j.1365-2435.2007.01360.x Chiyo PI, Lee PC, Moss CJ et al (2011) No risk, no gain: effects of crop raiding and genetic diversity on body size in male elephants. Behav Ecol 22:552–558. https://doi.org/10.1093/beheco/arr016 Chowdhury S, Brown J, Swedell L (2020) Anthropogenic effects on the physiology and behaviour of chacma baboons in the Cape Peninsula of South Africa. Conserv Physiol 8:coaa066. https://doi.org/10.1093/conphys/coaa066 Codron D, Codron J, Lee-Thorp JA et al (2007) Stable isotope characterization of mammalian predator–prey relationships in a South African savanna. Eur J Wildl Res 53:161–170. https://doi.org/10.1007/s10344-006-0075-x Codron D, Lee-Thorp JA, Sponheimer M et al (2006) Inter- and intrahabitat dietary variability of chacma baboons (Papio ursinus) in South African savannas based on fecal δ13C, δ15N, and %N. Am J Phys Anthropol 129:204–214. https://doi.org/10.1002/ajpa.20253 Crowley BE, Reitsema LJ, Oelze VM, Sponheimer M (2016) Advances in primate stable isotope ecology-Achievements and future prospects: Advances in Primate Isotope Ecology. Am J Primatol 78:995–1003. https://doi.org/10.1002/ajp.22510 Cypher BL, Frost N (1999) Condition of San Joaquin Kit Foxes in Urban and Exurban Habitats. J Wildl Manag 63:930. https://doi.org/10.2307/3802807 Dale VH, Polasky S (2007) Measures of the effects of agricultural practices on ecosystem services. Ecol Econ 64:286–296. https://doi.org/10.1016/j.ecolecon.2007.05.009 De Barba M, Miquel C, Boyer F et al (2014) DNA metabarcoding multiplexing and validation of data accuracy for diet assessment: application to omnivorous diet. Mol Ecol Resour 14:306–323. https://doi.org/10.1111/1755-0998.12188 De Carvalho DR, de Castro DMP, Callisto M et al (2019) Stable isotopes and stomach content analyses indicate omnivorous habits and opportunistic feeding behavior of an invasive fish. Aquat Ecol 53:365–381. https://doi.org/10.1007/s10452-019-09695-3 Department of Forestry, Fisheries and the Environment (DFFE) (2021) SA National Land-Cover Datasets. https://egis.environment.gov.za/sa_national_land_cover_datasets . Accessed 15 Mar 2022 Ehleringer JR, Cerling TE (2002) C3 and C4 Photosynthesis. The Earth system: biological and ecological dimensions of global environmental change, 2:186–190 Elizabeth L, Machado P, Zinöcker M et al (2020) Ultra-Processed Foods and Health Outcomes: A Narrative Review. Nutrients 12:1955. https://doi.org/10.3390/nu12071955 Ellis EC, Gauthier N, Klein Goldewijk K et al (2021) People have shaped most of terrestrial nature for at least 12,000 years. Proc Natl Acad Sci USA 118:e2023483118. https://doi.org/10.1073/pnas.2023483118 Elmqvist T, Zipperer WC, Güneralp B (2016) Urbanization, habitat loss, biodiversity decline: solution pathways to break the cycle. In, Seta K, Solecki WD, Griffith CA (eds.) Routledge Handbook of Urbanization and Global Environmental Change. Routledge, London and New York. Chapter 10: 139–151 Eriksson O (2013) Species pools in cultural landscapes - niche construction, ecological opportunity and niche shifts. Ecography 36:403–413. https://doi.org/10.1111/j.1600-0587.2012.07913.x Evans BA, Gawlik DE (2020) Urban food subsidies reduce natural food limitations and reproductive costs for a wetland bird. Sci Rep 10:14021. https://doi.org/10.1038/s41598-020-70934-x Fahrig L (2003) Effects of Habitat Fragmentation on Biodiversity. Annu Rev Ecol Evol Syst 34:487–515. https://doi.org/10.1146/annurev.ecolsys.34.011802.132419 Farr TG, Kobrick M (2000) Shuttle Radar Topography Mission Fiolet T, Srour B, Sellem L et al (2018) Consumption of ultra-processed foods and cancer risk: results from NutriNet-Santé prospective cohort. https://doi.org/10.1136/bmj.k322 . BMJ k322 Fischer J, Higham JP, Alberts SC et al (2019) Insights into the evolution of social systems and species from baboon studies. eLife 8:e50989. https://doi.org/10.7554/eLife.50989 Fleming PA, Bateman PW (2018) Novel predation opportunities in anthropogenic landscapes. Anim Behav 138:145–155. https://doi.org/10.1016/j.anbehav.2018.02.011 Foley JA, DeFries R, Asner GP et al (2005) Global Consequences of Land Use. Science 309:570–574. https://doi.org/10.1126/science.1111772 Francis RJ, Kingsford RT, Murray-Hudson M, Brandis KJ (2021) Urban waste no replacement for natural foods—Marabou storks in Botswana. J Urban Ecol 7:juab003. https://doi.org/10.1093/jue/juab003 Gámez S, Potts A, Mills KL et al (2022) Downtown diet: a global meta-analysis of increased urbanization on the diets of vertebrate predators. Proc R Soc B 289:20212487. https://doi.org/10.1098/rspb.2021.2487 Guiry E, Buckley M (2018) Urban rats have less variable, higher protein diets. Proc R Soc B 285:20181441. https://doi.org/10.1098/rspb.2018.1441 Handley LL, Raven JA (1992) The use of natural abundance of nitrogen isotopes in plant physiology and ecology. Plant Cell Environ 15:965–985 Heiss RS, Clark AB, McGowan KJ (2009) Growth and nutritional state of American Crow nestlings vary between urban and rural habitats. Ecol Appl 19:829–839. https://doi.org/10.1890/08-0140.1 Hoffman TS, O’Riain MJ (2012) Landscape requirements of a primate population in a human-dominated environment. Front Zool 9:1. https://doi.org/10.1186/1742-9994-9-1 Hoffman TS, O’Riain MJ (2011) The Spatial Ecology of Chacma Baboons (Papio ursinus) in a Human-modified Environment. Int J Primatol 32:308–328. https://doi.org/10.1007/s10764-010-9467-6 Hooke RLeB, Martín-Duque JF (2012) Land transformation by humans: A review. GSAT 12:4–10. https://doi.org/10.1130/GSAT151A.1 Hulme-Beaman A, Dobney K, Cucchi T, Searle JB (2016) An Ecological and Evolutionary Framework for Commensalism in Anthropogenic Environments. Trends in Ecology and Evolution 31:633–645. https://doi.org/10.1016/j.tree.2016.05.001 Johnson C, Piel AK, Forman D et al (2015) The ecological determinants of baboon troop movements at local and continental scales. Mov Ecol 3:14. https://doi.org/10.1186/s40462-015-0040-y Johnson CA, Raubenheimer D, Rothman JM et al (2013) 30 Days in the Life: Daily Nutrient Balancing in a Wild Chacma Baboon. PLoS ONE 8:e70383. https://doi.org/10.1371/journal.pone.0070383 Jordan MJR (2005) Dietary analysis for mammals and birds: a review of field techniques and animal-management applications. Int Zoo Yearbook 39:108–116. https://doi.org/10.1111/j.1748-1090.2005.tb00010.x Lee JH, Duster M, Roberts T, Devinsky O (2022) United States Dietary Trends Since 1800: Lack of Association Between Saturated Fatty Acid Consumption and Non-communicable Diseases. Front Nutr 8:748847. https://doi.org/10.3389/fnut.2021.748847 Lee S-M, Lee E-J (2019) Diet of the wild boar ( Sus scrofa ): implications for management in forest-agricultural and urban environments in South Korea. PeerJ 7:e7835. https://doi.org/10.7717/peerj.7835 Lee P, Priston N (2005) Human attitudes to primates: Perceptions of pests, conflict and consequences for primate conservation. Commensalism and Conflict: The Human-Primate Interface. https://doi.org/10.1002/047001539X.ch1 Lewis MC, West AG, O’Riain MJ (2018) Isotopic assessment of marine food consumption by natural-foraging chacma baboons on the Cape Peninsula, South Africa. Am J Phys Anthropol 165:77–93. https://doi.org/10.1002/ajpa.23332 Maibeche Y, Moali A, Yahi N, Menard N (2015) Is Diet Flexibility an Adaptive Life Trait for Relictual and Peri-Urban Populations of the Endangered Primate Macaca. sylvanus? PLoS ONE 10:e0118596. https://doi.org/10.1371/journal.pone.0118596 Martínez del Rio C, Wolf N, Carleton SA, Gannes LZ (2009) Isotopic ecology ten years after a call for more laboratory experiments. Biol Rev 84:91–111. https://doi.org/10.1111/j.1469-185X.2008.00064.x Martínez-Fernández J, Ruiz-Benito P, Zavala MA (2015) Recent land cover changes in Spain across biogeographical regions and protection levels: Implications for conservation policies. Land Use Policy 44:62–75. https://doi.org/10.1016/j.landusepol.2014.11.021 Marty PR, Balasubramaniam KN, Kaburu SSK et al (2020) Individuals in urban dwelling primate species face unequal benefits associated with living in an anthropogenic environment. Primates 61:249–255. https://doi.org/10.1007/s10329-019-00775-4 Matos RA, Adams M, Sabaté J (2021) Review: The Consumption of Ultra-Processed Foods and Non-communicable Diseases in Latin America. Front Nutr 8:622714. https://doi.org/10.3389/fnut.2021.622714 Matthews JK, Ridley A, Kaplin BA, Grueter CC (2020) A comparison of fecal sampling and direct feeding observations for quantifying the diet of a frugivorous primate. Curr Zool 66:333–343. https://doi.org/10.1093/cz/zoz058 Mazué F, Guerbois C, Fritz H et al (2023) Less bins, less baboons: reducing access to anthropogenic food effectively decreases the urban foraging behavior of a troop of chacma baboons (Papio hamadryas ursinus) in a peri-urban area. Primates 64:91–103. https://doi.org/10.1007/s10329-022-01032-x McKinney ML (2002) Urbanization, Biodiversity, and Conservation. Bioscience 52:883. https://doi.org/10.1641/0006-3568(2002)052 [0883:UBAC]2.0.CO;2 Mendonça R, de Lopes D, Pimenta ACS AM, et al (2017) Ultra-Processed Food Consumption and the Incidence of Hypertension in a Mediterranean Cohort: The Seguimiento Universidad de Navarra Project. Am J Hypertens 30:358–366. https://doi.org/10.1093/ajh/hpw137 Mendonça R, de Pimenta D, Gea AM A, et al (2016) Ultraprocessed food consumption and risk of overweight and obesity: the University of Navarra Follow-Up (SUN) cohort study. Am J Clin Nutr 104:1433–1440. https://doi.org/10.3945/ajcn.116.135004 Milton K (1999) Nutritional characteristics of wild primate foods: do the diets of our closest living relatives have lessons for us? Nutrition 15:488–498. https://doi.org/10.1016/S0899-9007(99)00078-7 Monteiro CA, Cannon G, Moubarac J-C et al (2018) The UN Decade of Nutrition, the NOVA food classification and the trouble with ultra-processing. Public Health Nutr 21:5–17. https://doi.org/10.1017/S1368980017000234 Mormile JE, Hill CM (2017) Living With Urban Baboons: Exploring Attitudes and Their Implications for Local Baboon Conservation and Management in Knysna, South Africa. Hum Dimensions Wildl 22:99–109. https://doi.org/10.1080/10871209.2016.1255919 Murray M, Cembrowski A, Latham ADM et al (2015) Greater consumption of protein-poor anthropogenic food by urban relative to rural coyotes increases diet breadth and potential for human–wildlife conflict. Ecography 38:1235–1242 Mychajliw AM, Almonte JN, Martinez PA, Hadly EA (2022) Stable isotopes reveal seasonal dietary responses to agroforestry in a venomous mammal, the Hispaniolan solenodon ( Solenodon paradoxus ). Ecol Evol 12. https://doi.org/10.1002/ece3.8761 Newsome SD, Ralls K, Job CVH et al (2010) Stable isotopes evaluate exploitation of anthropogenic foods by the endangered San Joaquin kit fox (Vulpes macrotis mutica). J Mammal 91:1313–1321. https://doi.org/10.1644/09-MAMM-A-362.1 Nicholson M, Cove MV (2022) Stable isotopes point to anthropogenic subsidies in northern raccoons at the urban-wild interface. Food Webs 31:e00233. https://doi.org/10.1016/j.fooweb.2022.e00233 O’Brien DM (2015) Stable Isotope Ratios as Biomarkers of Diet for Health Research. Annu Rev Nutr 35:565–594. https://doi.org/10.1146/annurev-nutr-071714-034511 Oro D, Genovart M, Tavecchia G et al (2013) Ecological and evolutionary implications of food subsidies from humans. Ecol Lett 16:1501–1514. https://doi.org/10.1111/ele.12187 Penick CA, Savage AM, Dunn RR (2015) Stable isotopes reveal links between human food inputs and urban ant diets. Proc R Soc B 282:20142608. https://doi.org/10.1098/rspb.2014.2608 Pollock CJ, Capilla-Lasheras P, McGill RAR et al (2017) Integrated behavioural and stable isotope data reveal altered diet linked to low breeding success in urban-dwelling blue tits (Cyanistes caeruleus). Sci Rep 7:5014. https://doi.org/10.1038/s41598-017-04575-y Pool-Stanvliet R, Coetzer K (2020) The scientific value of UNESCO biosphere reserves. S Afr J Sci 116. https://doi.org/10.17159/sajs.2020/7432 Popkin BM, Corvalan C, Grummer-Strawn LM (2020) Dynamics of the double burden of malnutrition and the changing nutrition reality. The Lancet 395:65–74. https://doi.org/10.1016/S0140-6736(19)32497-3 Post DM, Layman CA, Arrington DA et al (2007) Getting to the fat of the matter: models, methods and assumptions for dealing with lipids in stable isotope analyses. Oecologia 152:179–189. https://doi.org/10.1007/s00442-006-0630-x QGIS Development Team (2020) QGIS Geographic Information System. Version 3.18 RStudio T (2020) RStudio: Integrated Development for R. https://www.R-project.org/ Sapolsky RM, Share LJ (2004) A Pacific Culture among Wild Baboons: Its Emergence and Transmission. PLoS Biol 2:e106. https://doi.org/10.1371/journal.pbio.0020106 Schoeninger MJ, DeNiro MJ (1984) Nitrogen and carbon isotopic composition of bone collagen from marine and terrestrial animals. Geochim Cosmochim Acta 48:625–639 Scholz C, Firozpoor J, Kramer-Schadt S et al (2020) Individual dietary specialization in a generalist predator: A stable isotope analysis of urban and rural red foxes. Ecol Evol 10:8855–8870. https://doi.org/10.1002/ece3.6584 Schreier AL, Schlaht RM, Swedell L (2019) Meat eating in wild hamadryas baboons: Opportunistic trade-offs between insects and vertebrates. Am J Primatol 81. https://doi.org/10.1002/ajp.23029 Seoraj-Pillai N, Pillay N (2016) A Meta-Analysis of Human–Wildlife Conflict: South African and Global Perspectives. Sustainability 9:34. https://doi.org/10.3390/su9010034 Seress G, Sándor K, Evans KL, Liker A (2020) Food availability limits avian reproduction in the city: An experimental study on great tits Parus major . J Anim Ecol 89:1570–1580. https://doi.org/10.1111/1365-2656.13211 Slater K, Barrett A, Brown LR (2018) Home range utilization by chacma baboon (Papio ursinus) troops on Suikerbosrand Nature Reserve, South Africa. PLoS ONE 13:e0194717. https://doi.org/10.1371/journal.pone.0194717 Smith BN, Epsten S (1971) Two Categories of 1CC/12C Ratios for Higher Plantsl. 47:5 Sponheimer M, Robinson T, Ayliffe L et al (2003) Nitrogen isotopes in mammalian herbivores: hair * 15 N values from a controlled-feeding study. Int J Osteoarchaeol 13:80–87 Stillfried M, Gras P, Busch M et al (2017) Wild inside: Urban wild boar select natural, not anthropogenic food resources. PLoS ONE 12:e0175127. https://doi.org/10.1371/journal.pone.0175127 Strum SC (2010) The Development of Primate Raiding: Implications for Management and Conservation. Int J Primatol 31:133–156. https://doi.org/10.1007/s10764-009-9387-5 Sugden S, Murray M, Edwards MA, St. Clair CC (2021) Inter-population differences in coyote diet and niche width along an urban–suburban–rural gradient. J Urban Ecol 7:juab034. https://doi.org/10.1093/jue/juab034 Sugden S, Sanderson D, Ford K et al (2020) An altered microbiome in urban coyotes mediates relationships between anthropogenic diet and poor health. Scientific Reports Sun Y, Liu S, Liu Y et al (2021) Effects of the interaction among climate, terrain and human activities on biodiversity on the Qinghai-Tibet Plateau. Sci Total Environ 794:148497. https://doi.org/10.1016/j.scitotenv.2021.148497 Taberlet P, Bonin A, Zinger L, Coissac E (2018) Reference databases. In: Taberlet P, Bonin A, Zinger L, Coissac E (eds) Environmental DNA: For Biodiversity Research and Monitoring. Oxford University Press, p 0 Taki H, Ikeda H, Nagamitsu T et al (2017) Stable nitrogen and carbon isotope ratios in wild native honeybees: the influence of land use and climate. Biodivers Conserv 26:3157–3166. https://doi.org/10.1007/s10531-016-1114-x Turner BL, Lambin EF, Reenberg A (2007) The emergence of land change science for global environmental change and sustainability. Proc Natl Acad Sci USA 104:20666–20671. https://doi.org/10.1073/pnas.0704119104 Webber AD (2017) Commensalism. The International Encyclopedia of Primatology. John Wiley and Sons, Ltd, pp 1–2 Wist B, Stolter C, Dausmann KH (2022) Sugar addicted in the city: impact of urbanisation on food choice and diet composition of the Eurasian red squirrel ( Sciurus vulgaris ). J Urban Ecol 8:juac012. https://doi.org/10.1093/jue/juac012 Zhang W, Ricketts TH, Kremen C et al (2007) Ecosystem services and dis-services to agriculture. Ecol Econ 64:253–260. https://doi.org/10.1016/j.ecolecon.2007.02.024 Additional Declarations No competing interests reported. Supplementary Files Supplementaryinformation.docx SupplementaryInformation1.ods Cite Share Download PDF Status: Posted Version 1 posted You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-3427689","acceptedTermsAndConditions":true,"allowDirectSubmit":true,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":239405054,"identity":"deab97d6-f563-498d-847c-ba79cde77c8d","order_by":0,"name":"Elizabeth Kennedy Overton","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA00lEQVRIiWNgGAWjYDACHhiDvQFIGFgQrcUAyDgAoiVI0SKRAGIQoYWf5/AxiY97/siZSz6/uuFHgQQDf3t3Al4tkr1taZIznhkYW87OKbvZA3SYxJmzG/BqMTjPY2zMc8AgccPtnLQbPEAtBhK5+LXYg7T8AWm5eSbt5h9itBjw9hg+ZgBpucF+7DZRtkicOZb4sOeAsbHBmRy22zIGEjwE/cLfk3zgwI8DcnIGx48/u/nmj40cf3svfi1IgMcATBKrHATYH5CiehSMglEwCkYQAACJWEdwEB29dwAAAABJRU5ErkJggg==","orcid":"","institution":"REHABS, CNRS-NMU-UCBL, Nelson Mandela University","correspondingAuthor":true,"submittingAuthor":false,"prefix":"","firstName":"Elizabeth","middleName":"Kennedy","lastName":"Overton","suffix":""},{"id":239405055,"identity":"5c853256-75e2-4de5-a716-0fc2292bc2fe","order_by":1,"name":"Alice Bernard","email":"","orcid":"","institution":"REHABS, CNRS-NMU-UCBL, Nelson Mandela University","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Alice","middleName":"","lastName":"Bernard","suffix":""},{"id":239405056,"identity":"8b196306-5a82-484a-a740-9c266fd240d6","order_by":2,"name":"Pierre-Cyril Renaud","email":"","orcid":"","institution":"University of Angers, BiodivAG-IRL REHABS","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Pierre-Cyril","middleName":"","lastName":"Renaud","suffix":""},{"id":239405058,"identity":"22eec156-665e-4acf-ae83-a2b12ac2914b","order_by":3,"name":"Grant Hall","email":"","orcid":"","institution":"UP Stable Isotope Laboratory, University of Pretoria","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Grant","middleName":"","lastName":"Hall","suffix":""},{"id":239405061,"identity":"4572d281-b938-427f-bb45-9c6fef2b67d0","order_by":4,"name":"Chloé Guerbois","email":"","orcid":"","institution":"Nelson Mandela University","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Chloé","middleName":"","lastName":"Guerbois","suffix":""},{"id":239405064,"identity":"0970ddcb-8d40-4b36-a693-67319a6af0fb","order_by":5,"name":"Hervé Fritz","email":"","orcid":"","institution":"REHABS, CNRS-NMU-UCBL, Nelson Mandela University","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Hervé","middleName":"","lastName":"Fritz","suffix":""},{"id":239405066,"identity":"dc525945-d5e6-4f95-9ff0-09f23772bff1","order_by":6,"name":"Franck Prugnolle","email":"","orcid":"","institution":"REHABS, CNRS-NMU-UCBL, Nelson Mandela University","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Franck","middleName":"","lastName":"Prugnolle","suffix":""},{"id":239405067,"identity":"5cb17b28-10b6-4c19-8e1c-2009949713ef","order_by":7,"name":"Virginie Rougeron","email":"","orcid":"","institution":"REHABS, CNRS-NMU-UCBL, Nelson Mandela University","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Virginie","middleName":"","lastName":"Rougeron","suffix":""}],"badges":[],"createdAt":"2023-10-10 13:44:34","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-3427689/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-3427689/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":44793613,"identity":"ab6c9b04-bb9d-4ab5-8a0a-341837a0eaef","added_by":"auto","created_at":"2023-10-17 15:02:55","extension":"jpeg","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":7712202,"visible":true,"origin":"","legend":"\u003cp\u003eStudy area and collected samples (Garden Route, Western Cape, South Africa). Map (a) shows landscape composition and the three study locations, along with the number of samples collected for each subsite. A zoom on each location (b) shows the distribution of where samples were collected for all six subsites (Main and North, Oakhurst and Woodville, Goudveld and Diepwalle). The red panel visualizes the most anthropized and built up area that is the Nelson Mandela University, named Campus. The purple panel represents the more intermediate anthropization level, the agricultural location called Hoekwil. The last two panels in blue represent the two subsites of the more natural environment within the Garden Route National Park, namely Knysna. Source: SANLC, 2018 - EPSG 32734.\u003c/p\u003e","description":"","filename":"Figure1.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-3427689/v1/f676d50617678a748c5fa931.jpeg"},{"id":44793612,"identity":"7ec94642-80b4-423d-b4ed-dbeedc340d8d","added_by":"auto","created_at":"2023-10-17 15:02:55","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":986692,"visible":true,"origin":"","legend":"\u003cp\u003e(a) PCA (Axes 1 and 2) land cover characteristics obtained around each of the 148 analyzed fecal samples across the three studied locations (Campus, Hoekwil and Knysna). Biplots of the isotopic ratios of the 146 analyzed fecal samples comparing (b) Carbon and Nitrogen (ellipse = 0.95), (c) nitrogen (δ15N) according to the anthropization level (i.e. the coordinates of the second axis of the corresponding PCA) and (d) the distribution of the carbon ratio (δ13C).\u003c/p\u003e","description":"","filename":"Figure2.png","url":"https://assets-eu.researchsquare.com/files/rs-3427689/v1/4c77fe40a3dd9743b7c99a27.png"},{"id":44793611,"identity":"bd3e802e-e06a-40f5-8519-45217fb59067","added_by":"auto","created_at":"2023-10-17 15:02:55","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":594383,"visible":true,"origin":"","legend":"\u003cp\u003eKruskal-Wallis tests for (a) nitrogen and (b) carbon including Dunn post-hoc tests comparing the three locations (Campus, Hoekwil and Knysna).\u003c/p\u003e","description":"","filename":"Figure3.png","url":"https://assets-eu.researchsquare.com/files/rs-3427689/v1/5fd3f212cb2873422da224cd.png"},{"id":44793610,"identity":"7d77556e-c932-4a2c-8c15-d813d9c9be16","added_by":"auto","created_at":"2023-10-17 15:02:55","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":604549,"visible":true,"origin":"","legend":"\u003cp\u003e(a) PCA (Axes 1 and 2) of land cover variables and the 60 analyzed fecal samples for the two subsites on Campus (Main and North). (b) Nitrogen isotopic ratios (δ15N) for the 58 fecal samples according to the anthropization level (i.e. the first axis of the corresponding PCA; ellipse = 0.95). (c) Distribution of the carbon isotopic ratios (δ13C) for the 58 fecal samples.\u003c/p\u003e","description":"","filename":"Figure4.png","url":"https://assets-eu.researchsquare.com/files/rs-3427689/v1/d10588db79a6b69650fb9788.png"},{"id":48358959,"identity":"0ec48085-dd3d-4d45-80ca-6baf48cefaa4","added_by":"auto","created_at":"2023-12-18 00:52:21","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":1129580,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-3427689/v1/d7c87988-6e44-468f-9127-6af01115397c.pdf"},{"id":44793614,"identity":"67b142a6-14da-4848-9f88-4a7e34debf41","added_by":"auto","created_at":"2023-10-17 15:02:56","extension":"docx","order_by":2,"title":"","display":"","copyAsset":false,"role":"supplement","size":733107,"visible":true,"origin":"","legend":"","description":"","filename":"Supplementaryinformation.docx","url":"https://assets-eu.researchsquare.com/files/rs-3427689/v1/385beaaa4bd96374c15fc508.docx"},{"id":44793615,"identity":"c3988a7f-1946-4f5a-b6dd-06e17a987f19","added_by":"auto","created_at":"2023-10-17 15:02:56","extension":"ods","order_by":9,"title":"","display":"","copyAsset":false,"role":"supplement","size":34919,"visible":true,"origin":"","legend":"","description":"","filename":"SupplementaryInformation1.ods","url":"https://assets-eu.researchsquare.com/files/rs-3427689/v1/49f53f1d3d421bae2ad791a6.ods"}],"financialInterests":"No competing interests reported.","formattedTitle":"Land use influence on chacma baboon (Papio ursinus) diet in South Africa using stable isotopes","fulltext":[{"header":"Introduction","content":"\u003cp\u003eThe transformation of natural landscapes by human activities (Mart\u0026iacute;nez-Fern\u0026aacute;ndez et al. \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2015\u003c/span\u003e) has modified over half of the Earth\u0026rsquo;s land surface (Foley et al. \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Hooke et al. 2012). Land use and land cover change (LULCC), including extensive clearing, the creation of pastures, croplands and urban settlements (Fahrig \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2003\u003c/span\u003e; McKinney \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2002\u003c/span\u003e) are the main drivers impacting ecosystems and causing drastic global and regional biodiversity collapse (Ellis et al. \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Foley et al. \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Sun et al. \u003cspan citationid=\"CR91\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Turner et al. \u003cspan citationid=\"CR94\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Zhang et al. \u003cspan citationid=\"CR97\" class=\"CitationRef\"\u003e2007\u003c/span\u003e). For certain species, these transformed landscapes can create new niches and provide wild animals with other resources (Dale and Polasky, \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Eriksson \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Fleming and Bateman \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2018\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eIn anthropized landscapes, the possible access to cultivated zones, gardens, kitchens and garbage has provided new opportunities for wildlife (Hulme-Beaman et al. \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Oro et al. \u003cspan citationid=\"CR70\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). Certain species have adapted or acclimated to these transformed areas, exploiting anthropogenic resources (Hulme-Beaman et al. \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Seoraj-Pillai and Pillay \u003cspan citationid=\"CR82\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Webber \u003cspan citationid=\"CR95\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). For instance, wild boar (\u003cem\u003eSus scrofa\u003c/em\u003e) and vervet monkeys (\u003cem\u003eCercopithecus aethiops\u003c/em\u003e) are known to exploit croplands (Cancelliere et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Lee and Lee \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e2019\u003c/span\u003e), carnivores such as San Joaquin kit foxes (\u003cem\u003eVulpes macrotis mutica\u003c/em\u003e) or coyotes (\u003cem\u003eCanis latrans\u003c/em\u003e) exploit urban resources (e.g. garbage or compost; Newsome et al. \u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Sugden et al. \u003cspan citationid=\"CR89\" class=\"CitationRef\"\u003e2021\u003c/span\u003e) and commensal behaviour, notably as household and agricultural pests, has been observed in black rats (\u003cem\u003eRattus rattus\u003c/em\u003e) across different geographical areas on several occasions (Aplin et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). These changes cause direct modifications in wildlife diet, from the consumption of purely natural resources to the integration of more human-originated food (Murray et al. \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). This can be a benefit to species, providing year-round food provisioning, reducing food and water deprivation during droughts or winters, or generally contributing to better individual condition. For example, a population of urban kit foxes presented higher cholesterol levels, a good indicator of energy intake (Harder and Kirk-patrick 1994), as well as reduced nutritional deprivation than rural ones in California (Cypher and Frost \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e1999\u003c/span\u003e). The integration of human resources into the diet of storks during periods of constrained resource availability has also been shown to lead to higher reproduction rates (Evans and Gawlik \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). As another example, crop raiding behaviour in male elephants resulted in larger body size in adulthood (Chiyo et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2011\u003c/span\u003e), an important trait for this species during mating (Chelliah and Sukumar 2013). Despite certain benefits however, the consumption of human food has also been shown to have a negative impact on wildlife health. For instance, urban coyotes generally showed poorer average body condition as well as higher parasitic infection rates than rural coyotes in a study by Sugden et al. (\u003cspan citationid=\"CR90\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Reduced reproductive rates of birds (e.g. blue tits) are also associated with urban areas compared to populations in rural zones, linked to a lack of suitable diets for nestlings (Pollock et al. \u003cspan citationid=\"CR72\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Seress et al. \u003cspan citationid=\"CR83\" class=\"CitationRef\"\u003e2020\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe impacts of human resources on non-human primates appear to a lesser extent in the literature, even though primates often consume human resources (Strum \u003cspan citationid=\"CR88\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). It has been shown that wild primates obtain higher levels of essential nutrients relative to body weight than humans do through their diets (Milton \u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e1999\u003c/span\u003e), but also that varying benefits can follow their consumption of anthropic food resources depending on the type of LULCC (Maibeche et al. \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Marty et al. \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). An interesting primate species to better understand diet change in the presence of human resources is the chacma baboon (\u003cem\u003ePapio ursinus\u003c/em\u003e), due to its adaptive capacity (Fischer et al. \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Baboons are opportunistic, omnivorous feeders that will eat anything from fruits, grasses, leaves or roots to invertebrates and even occasionally animal matter usually resulting from opportunistic hunting (Allan et al. \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Hoffman and O\u0026rsquo;Riain \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Schreier et al. \u003cspan citationid=\"CR81\" class=\"CitationRef\"\u003e2019\u003c/span\u003e) and are highly adaptable to a wide range of habitats and environments (Johnson et al. \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). They can be found across deserts, savannahs, forests and on the outskirts or within urban zones (Hoffman and O\u0026rsquo;Riain \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2011\u003c/span\u003e, \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Johnson et al. \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Mormile and Hill \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). With the encroachment of urban areas on natural habitats, encounters between humans and baboons have become prevalent in some areas, such as in the Western Cape of South Africa (Bracken et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2022a\u003c/span\u003e, \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003eb\u003c/span\u003e, \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Chowdhury et al. \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Lee and Priston \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2005\u003c/span\u003e). In the Western Cape, as well as across South Africa, chacma baboons frequently raid croplands as well as people\u0026rsquo;s gardens, bins or houses (Hoffman and O\u0026rsquo;Riain \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Strum \u003cspan citationid=\"CR88\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Walton et al. 2021), consuming resources of human origin. When the availability of rubbish is reduced, this affects the baboon\u0026rsquo;s raiding tendencies: Mazu\u0026eacute; et al. (\u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e2023\u003c/span\u003e) showed that when the rubbish resources were removed on regular foraging sites in a peri-urban area on the George Campus of the Nelson Mandela University (NMU), chacma baboons spent more time foraging for natural resources and less time in these more urban areas. The easy access to fruits, vegetables, meats, but also processed foods, refined sugars, added fats and oils may result in significant variations in their nutrient consumption and have unknown effects on their metabolisms. In this context, the objective of this present study was to evaluate whether human land use, characterized by high human modified food availability, has an impact on chacma baboon diet.\u003c/p\u003e \u003cp\u003eSeveral methods exist to study diets in ecology, including direct observation or fecal analysis. These techniques yield accurate results but they can vary according to what is consumed and the digestibility of the latter (Matthews et al. \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2020\u003c/span\u003e): some resources will appear in higher proportions in fecal samples than others depending on how easy they are to digest. Direct observation also often requires long periods of time (Jordan \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2005\u003c/span\u003e), whereas non-invasive methods based on fecal samples, such as DNA metabarcoding and/or isotopic analyses, can counter this problem (Ando et al. \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Crowley et al. \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; De Barba et al. \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; O\u0026rsquo;Brien \u003cspan citationid=\"CR69\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). In the case of metabarcoding however, sample analysis can be costly and a broad list of plant/animal DNA sequences that could be consumed in the region would be necessary (Taberlet et al. \u003cspan citationid=\"CR92\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Analysing isotopic ratios is therefore an interesting alternative, both less expensive and less time-consuming than other options, which explains why it is a recurrent method used in feeding ecology studies (Boecklen et al. \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Mart\u0026iacute;nez del Rio et al. \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Taki et al. \u003cspan citationid=\"CR93\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). Common isotopes used for diet studies include nitrogen (\u003csup\u003e15\u003c/sup\u003eN/\u003csup\u003e14\u003c/sup\u003eN) and carbon (\u003csup\u003e13\u003c/sup\u003eC/\u003csup\u003e12\u003c/sup\u003eC). Nitrogen isotopes are a potential biomarker of protein source because tissue nitrogen derives almost entirely from dietary protein (for a review of isotopes as biomarkers see O\u0026rsquo;Brien \u003cspan citationid=\"CR69\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). Due to a stepwise increase with the trophic chain, variations of the δ\u003csup\u003e15\u003c/sup\u003eN isotopic ratios observed within feces may reflect consumption of dietary items with higher protein content within the different LULCC based on what resources the baboons have access to. The carbon in plants derives from atmospheric CO\u003csub\u003e2\u003c/sub\u003e which is fixed during the process of photosynthesis. Differences in stable carbon isotope ratios therefore reflect the type of plants consumed based on their photosynthetic pathways (C\u003csub\u003e3\u003c/sub\u003e, C\u003csub\u003e4\u003c/sub\u003e or CAM) due to differential fractionation of carbon isotopes during photosynthesis that leads to distinct δ\u003csup\u003e13\u003c/sup\u003eC values (Ehleringer and Cerling \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2002\u003c/span\u003e; Smith and Epsten \u003cspan citationid=\"CR85\" class=\"CitationRef\"\u003e1971\u003c/span\u003e). Examining carbon and nitrogen ratios may therefore be a suitable method of assessing the potential contribution of human-derived foods to animal diet, comparing protein intake and the type of plants consumed between natural areas and those with anthropic resources (Penick et al. \u003cspan citationid=\"CR71\" class=\"CitationRef\"\u003e2015\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eTo address the objective of this study, we obtained stable isotope ratios from fecal samples of chacma baboons, which would provide quantitative data on the types of plants and animal protein they consume. Baboons may be considered as opportunistic feeders (Allan et al. \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Schreier et al. \u003cspan citationid=\"CR81\" class=\"CitationRef\"\u003e2019\u003c/span\u003e) and therefore the potential access to high protein sources such as meat and legumes in more anthropized areas (with crops, garbage or kitchens) may result in higher proportions of protein in their diet. We could expect that if the anthropized landscapes provide higher protein contents than the natural resources consumed by chacma baboons, the stable nitrogen isotope values would tend to be higher in these transformed landscapes. More positive δ\u003csup\u003e13\u003c/sup\u003eC values for samples from anthropized areas may be associated with a higher consumption of C\u003csub\u003e4\u003c/sub\u003e plants that pervade human-derived foods (e.g. maize, sugarcane) as well as lipids when these types of resources are included in chacma baboon diets (Post et al. \u003cspan citationid=\"CR75\" class=\"CitationRef\"\u003e2007\u003c/span\u003e). We could then expect to observe more positive δ\u003csup\u003e13\u003c/sup\u003eC ratios from samples collected in more urbanized environments compared to natural landscapes. To evaluate whether transformed landscapes have an influence on chacma baboon diet, we collected fecal samples from chacma baboons along a gradient of land use in the Garden Route, South Africa, and studied stable isotopic ratio data, incorporating the surrounding landscape characteristics.\u003c/p\u003e"},{"header":"Material and methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003eStudy area\u003c/h2\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eThe study was conducted in the Western Cape, South Africa, across the Garden Route Biosphere Reserve (listed in the UNESCO\u0026rsquo;s World Network of Biosphere Reserves; Pool-Stanvliet and Coetzer \u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e2020\u003c/span\u003e) between George and Knysna, representing an area of 2000km\u0026sup2; (Fig.\u0026nbsp;1a). The reserve is known for its unique and endemic biodiversity, such as afrotemperate forests, fynbos, as well as wetlands, mountains and coastal ecosystems. However, human presence has largely transformed the landscape in many places (Socio-Economic Profile: Garden Route District Municipality 2021), providing agricultural, forestry and touristic activities as well as the development of many urban settlements (e.g. Mossel Bay, George, Knysna). To protect the area, the Garden Route biosphere reserve, with the Garden Route National Park (GRNP) as a core area was created, preserving natural areas where biodiversity can prosper. Due to the complex landscape configuration that includes agriculture, logging, urban areas as well as natural vegetation, this IUCN type II protected area is very patchy and fragmented. Certain species, such as chacma baboons, can be found across all these different ecosystems, populating protected areas, rural zones as well as highly urbanized regions (Mormile and Hill \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Slater et al. \u003cspan citationid=\"CR84\" class=\"CitationRef\"\u003e2018\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eIn the study area, three main contrasted locations were selected, a peri-urban, an agricultural and a protected area, hereafter referred to as Campus, Hoekwil and Knysna throughout this study (represented in red, purple and blue respectively in Fig.\u0026nbsp;1a). These three sampling areas represent different land use types. The first location (Campus) is located on the Nelson Mandela University Campus near the city of George, where there are many people, buildings (e.g. accommodation and cafeterias) and lawns, and is surrounded by natural forest and fynbos. The second location (Hoekwil) was situated in the agricultural area bordering the Garden Route National Park to the North of the Town named Hoekwil. This area mainly consisted of livestock pastures and croplands, with occasional buildings (accommodation and infrastructure). In the region, summer crops can include lucerne, maize and vegetables (CapeFarmMapper v 2.7). The third location (Knysna) was situated within the afrotemperate forest of the Knysna Section of the Garden Route National Park (GRNP).\u003c/p\u003e \u003cp\u003eTwo subsites were established for each of these three main locations as replicates to account for troop preferences and intra-troop variation: Main and North for Campus (respectively light and dark red; Fig.\u0026nbsp;1b), Oakhurst and Woodville for Hoekwil (light and dark purple; Fig.\u0026nbsp;1b), Goudveld and Diepwalle sections of the GRNP for Knysna (light and dark blue; Fig.\u0026nbsp;1b). The two replicates for Campus represent two distinct troops, whereas the replicates for Hoekwil and Knysna were made up of one or more distinct troops each, and each replicate had varying proportions of land cover including within each of the three sites.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec4\" class=\"Section2\"\u003e \u003ch2\u003eChacma baboon fecal sampling\u003c/h2\u003e \u003cp\u003eSampling took place between the 14th of February and the 1st of April 2022. fecal samples from baboons were collected during regular visits to each of the three locations over this period. Each location was visited two to three times a week. Sample collection was opportunistic, resulting in different sample sizes for both temporal and spatial replicates.\u003c/p\u003e \u003cp\u003eAt the Campus, samples were collected on foot by retracing where the Main and North troops had been sighted by students or staff. At Hoekwil, footpaths and fences where the baboons were often sighted were scoured on foot. In the Knysna forest, road transects were followed by car in search of samples. Each sample was collected using gloves and inserted into plastic tubes or bags. At each sampling point the collection date, latitude, longitude, freshness of feces, location and subsite names were noted (SI 1). Samples were kept in a cooler box during the fieldwork each day before being frozen and stored at -20\u0026deg;C at NMU REHABS Laboratory until further analysis. Data collection and analyses were authorized by permits from SANParks (BERN-A/2020-008) and Cape Nature (CN44-87-16198), and the Section 20 of the Animal Diseases Act, 1984 (Act No 35 of 1984).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec5\" class=\"Section2\"\u003e \u003ch2\u003eLandscape characteristics surrounding each collected fecal sample\u003c/h2\u003e \u003cp\u003eThe study area (latitude: 630900\u0026ndash;725500, longitude: 6214900\u0026ndash;6263600) was represented using QGIS (3.18), under a WGS 84 \u0026ndash; UTM 34S projection. It was characterized according to several landscape metrics: land cover, slope, elevation and length/number of roads. In order to characterize the landscape metrics in the area used by the baboons for feeding, we recorded them within a circle of 4km\u0026sup2; defined around each fecal sample position, based on the highest daily distance covered by chacma baboons in the literature in South Africa (Hoffman and O\u0026rsquo;Riain \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Slater et al. \u003cspan citationid=\"CR84\" class=\"CitationRef\"\u003e2018\u003c/span\u003e) and the observations of the baboon troops on NMU Campus in George by Maud Czerwinski et al. (unpubl. Manuscript).\u003c/p\u003e \u003cp\u003eThe land cover raster layer was obtained from the South African National Land Cover Datasets (2018) of the South African Forestry, Fisheries and the Environment government website (DEFE, 2021) and clipped to the study area. The 73 classes from this layer were grouped into 10 (Fig.\u0026nbsp;1a) for simplification according to main types (\u003cem\u003ei.e\u003c/em\u003e. natural forest, plantations, shrubland, grassland, waterbodies, wetland, barren land, cultivated land, built up and mines). The elevation raster at 30 meters was downloaded from SRTM products (Farr and Kobrick \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2000\u003c/span\u003e, Shuttle Radar Topography Mission) and slopes were calculated within QGIS. The roads vector layer covering South Africa was obtained from the website of the Princeton University Library (International Steering Committee for Global Mapping, map issued in 2016). The percentage of land cover, as well as mean slope, elevation, road length and number of road segments was calculated within the buffer of each sample using QGIS (3.18). Finally, a principal component analysis (PCA) was performed considering these variables after removing those with a correlation higher than 0.8, to visualize the land use characteristics surrounding each sample.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec6\" class=\"Section2\"\u003e \u003ch2\u003eStable isotope analysis for chacma baboon diet characterization\u003c/h2\u003e \u003cp\u003eAfter being placed in a fridge for 24 hours to defrost, up to 10g of each sample used for the isotopic analyses was inserted into 20mL tubes and dried for 48 hours in an oven at 60\u0026deg;C at the NMU REHABS laboratory (following steps used by Caut et al. \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Codron et al. \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; De Carvalho et al. \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). These samples were then transported to the University of Pretoria where they were ground using a mortar and pestle and passed through a \u0026frac12; mm sieve. The homogenized fecal samples were weighed out in aliquots of 0.9 mg and placed in tin capsules, which were pre-cleaned in toluene to avoid any potential contamination.\u003c/p\u003e \u003cp\u003eThe isotopic ratios for carbon (C) and nitrogen (N) of these samples were analysed at the University of Pretoria, South Africa. Samples were combusted at 1,020\u0026deg;C using an elemental analyser (Flash EA 1112 Series) coupled to a Delta V Plus stable light isotope ratio mass spectrometer via a ConFlo IV system (all equipment supplied by Thermo Fisher, Bremen, Germany), housed at the University of Pretoria Stable Isotope Laboratory, Mammal Research Institute. Several samples were run in duplicate (every 13th sample) to verify the accuracy of the homogenization of the samples. Two laboratory running standards Merck Gel (δ\u003csup\u003e13\u003c/sup\u003eC = -20.26\u0026permil;, δ\u003csup\u003e15\u003c/sup\u003eN=7.89\u0026permil;, C%=41.28, N%=15.29) and DL-Valine (δ\u003csup\u003e13\u003c/sup\u003eC = -10.57\u0026permil;, δ\u003csup\u003e15\u003c/sup\u003eN=-6.15\u0026permil;, C%=55.50, N%=11.86) and a blank sample were run after every 11 unknown samples. The carbon and nitrogen ratios for the laboratory running (Merck and DL-Valine) standards were calibrated using the following PRIMARY standards: IAEA- CH-3 (Cellulose), IAEA-CH-6 (Sucrose), IAEA-CH-7 (Polyethylene foil), IAEA N-1 and IAEA N-2 (Ammonium sulfate), IAEA NO-3 (Potassium nitrate). All results were referenced to Vienna Pee-Dee Belemnite for carbon isotope values, and to air for nitrogen isotope values.\u003c/p\u003e \u003cp\u003eResults are expressed in delta notation using a per mil scale using the standard equation (Coplen 2011, Rapid Comm, in Mass Spectrometry): δX(\u0026permil;)=[(R\u003csub\u003esample\u003c/sub\u003e -R\u003csub\u003estandard\u003c/sub\u003e )/R\u003csub\u003estandard\u003c/sub\u003e -1] where X\u0026thinsp;=\u0026thinsp;\u003csup\u003e15\u003c/sup\u003eN or \u003csup\u003e13\u003c/sup\u003eC and R represents \u003csup\u003e15\u003c/sup\u003eN/\u003csup\u003e14\u003c/sup\u003eN or \u003csup\u003e13\u003c/sup\u003eC/\u003csup\u003e12\u003c/sup\u003eC respectively.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec7\" class=\"Section2\"\u003e \u003ch2\u003eStatistical analysis of diet variations\u003c/h2\u003e \u003cp\u003eStatistical analyses were conducted using R (R Core Team 2020, version 3.6.3.) in RStudio (version 1.3.1093) to analyse and compare the isotopic patterns (δ\u003csup\u003e15\u003c/sup\u003eN and δ\u003csup\u003e13\u003c/sup\u003eC) of chacma baboon diets in relation to land use over the three locations (Campus, Hoekwil, Knysna). Firstly, the data for each stable isotope ratio were summarized as mean and standard deviation values according to location and subsite. After carrying out Shapiro-Wilk and Levene tests, non-parametric Kruskal-Wallis tests were used to determine the significance of the location variable on the isotopic ratios of δ\u003csup\u003e13\u003c/sup\u003eC and δ\u003csup\u003e15\u003c/sup\u003eN followed by Dunn pairwise comparison tests to establish the significance of the differences between locations characterized by different landscapes and between the subsites of each location. A biplot was also created to visualize any variation in diet according to land use by comparing the δ\u003csup\u003e13\u003c/sup\u003eC and δ\u003csup\u003e15\u003c/sup\u003eN ratios by site. Finally, the differences between chacma baboon populations were tested across the gradient of anthropization created using the resulting axes of the PCA, using a linear model (LM) for the nitrogen and distribution plots for the carbon ratios.\u003c/p\u003e \u003cp\u003eTo establish if grouping the subsites within each location had an impact on the results, an inter-troop analysis was carried out focusing on the subsites of the Campus (Main and North). The comparison between these troops within site was possible due to the intimate knowledge of the troops using the area through regular monitoring and research projects. Following Shapiro-Wilk and Levene tests, a Student\u0026rsquo;s t-test was carried out on the Campus δ\u003csup\u003e15\u003c/sup\u003eN results and a Wilcoxon test for the δ\u003csup\u003e13\u003c/sup\u003eC.\u003c/p\u003e \u003c/div\u003e"},{"header":"Results","content":"\u003cdiv id=\"Sec9\" class=\"Section2\"\u003e \u003ch2\u003eSampling\u003c/h2\u003e \u003cp\u003eA total of 231 baboon fecal samples were collected throughout the sampling period (Fig.\u0026nbsp;1a; Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Within the Campus, a total of 98 samples were collected, 60 for the Main and 38 for the North subsites, representing two different troops of baboons monitored on the NMU campus. At the Hoekwil site, 28 samples were found, with 14 for each subsite (Oakhurst and Woodville). In Knysna, 105 were collected in total, 67 of which for the Goudveld subsite and 38 for the Diepwalle subsite. We selected 148 samples for analysis in this study, according to their freshness and to obtain a similar number of replicates per location (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e; SI 1). A total of 60 samples were selected for both Campus and Knysna, with 30 samples per subsite. The Hoekwil samples were used in their totality due to the limitation of samples available linked to difficulties in finding samples in these areas, amounting to a total of 28 (14 per subsite).\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eNumbers of fecal samples collected, analyzed for land cover and analyzed for stable isotopes by location and subsite. The 146 samples chosen for the stable isotope analysis were chosen according to freshness and to obtain a representative number of replicates per location.\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"7\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eLocation\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e \u003cp\u003eCampus\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colspan=\"2\" nameend=\"c5\" namest=\"c4\"\u003e \u003cp\u003eHoekwil\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colspan=\"2\" nameend=\"c7\" namest=\"c6\"\u003e \u003cp\u003eKnysna\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSubsite\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMain\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eNorth\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eOakhurst\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eWoodville\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eGoudveld\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e \u003cp\u003eDiepwalle\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003e\u003cb\u003eNumber of collected samples\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e60\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e38\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e14\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e14\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e67\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e38\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e \u003cp\u003e98\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c5\" namest=\"c4\"\u003e \u003cp\u003e28\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c7\" namest=\"c6\"\u003e \u003cp\u003e105\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colspan=\"6\" nameend=\"c7\" namest=\"c2\"\u003e \u003cp\u003e231\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003e\u003cb\u003eNumber of samples analyzed for land cover\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e30\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e30\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e14\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e14\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e30\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e30\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e \u003cp\u003e60\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c5\" namest=\"c4\"\u003e \u003cp\u003e28\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c7\" namest=\"c6\"\u003e \u003cp\u003e60\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colspan=\"6\" nameend=\"c7\" namest=\"c2\"\u003e \u003cp\u003e148\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003e\u003cb\u003eNumber of samples analyzed for isotopes\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e29\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e29\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e14\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e14\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e30\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e30\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e \u003cp\u003e58\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c5\" namest=\"c4\"\u003e \u003cp\u003e28\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c7\" namest=\"c6\"\u003e \u003cp\u003e60\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colspan=\"6\" nameend=\"c7\" namest=\"c2\"\u003e \u003cp\u003e146\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec10\" class=\"Section2\"\u003e \u003ch2\u003eLandscape and anthropization level characterization\u003c/h2\u003e \u003cp\u003eThe PCA biplot created by combining the landscape variables used in this study revealed that the 148 samples from the three locations (Campus, Hoekwil, Knysna) were characterized by distinct land cover (Fig.\u0026nbsp;2a). Three clusters emerged within this figure, grouping together the samples from each studied location. The first axis, which explained 41.1% of the variation, opposed the flatter areas dominated by agriculture (including irrigation dams) mostly present in Hoekwil with the area to the steeper slopes dominated by natural forests and characteristic of the Knysna site. On the other hand, the second axis of this PCA, explaining 30.5% of the variation, opposed the built up location (Campus) with the more natural and less built up areas (Hoekwil and especially Knysna; Fig.\u0026nbsp;2a). A gradient, going from land rich with infrastructure (Campus) to natural environments without human settlements (Knysna) via an intermediate, agricultural area (Hoekwil), could be distinguished along the second axis (Fig.\u0026nbsp;2a). This axis was therefore defined as the gradient of anthropization of the environment for the following diet analyses of chacma baboons, with more negative values representing the more natural areas, more positive values representing more urban zones and intermediate values representing the agricultural areas.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec11\" class=\"Section2\"\u003e \u003ch2\u003eStable isotope signatures and distributions according to the level of anthropization\u003c/h2\u003e \u003cp\u003eOut of the 148 fecal samples, 146 were analysed for δ\u003csup\u003e15\u003c/sup\u003eN and δ\u003csup\u003e13\u003c/sup\u003eC (SI 1) due to reduced quality of two samples. When combining the carbon and nitrogen ratios, the resulting biplot showed that the samples from the three locations had different nitrogen and carbon isotopic ratios (Fig.\u0026nbsp;2b). The widest range of isotopic ratios belonged to the Campus, the smallest to Knysna, with intermediate values for Hoekwil, resembling the Campus for its δ\u003csup\u003e15\u003c/sup\u003eN and Knysna for its δ\u003csup\u003e13\u003c/sup\u003eC (Fig.\u0026nbsp;2b). Low values of δ\u003csup\u003e15\u003c/sup\u003eN and δ\u003csup\u003e13\u003c/sup\u003eC tended to characterize the Knysna samples, going from \u0026minus;\u0026thinsp;0.02 to 3.12\u0026permil; for nitrogen and \u0026minus;\u0026thinsp;29.96 to -23.53\u0026permil; for carbon (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). A wide range of δ\u003csup\u003e15\u003c/sup\u003eN (from 0.55 to 5.15\u0026permil;) combined with low values of δ\u003csup\u003e13\u003c/sup\u003eC (from \u0026minus;\u0026thinsp;29.82 to -25.67\u0026permil;) characterized those of Hoekwil (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). A more intermediate profile characterized the Campus samples, yielding a wider range of δ\u003csup\u003e15\u003c/sup\u003eN (0.66 to 5.31\u0026permil;) and wider ranges of δ\u003csup\u003e13\u003c/sup\u003eC (-29.26 to -17.92\u0026permil;) than the other locations (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab2\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 2\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eSummary statistics (including minimum, maximum, mean and standard deviation SD) for isotopic ratios of Nitrogen and Carbon in the 146 analysed fecal samples by location and subsite.\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"9\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c9\" colnum=\"9\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colspan=\"4\" nameend=\"c5\" namest=\"c2\"\u003e \u003cp\u003eδ15N (\u0026permil;)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colspan=\"4\" nameend=\"c9\" namest=\"c6\"\u003e \u003cp\u003eδ13C (\u0026permil;)\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eLocation\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMin\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eMax\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eMean\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eSD\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eMin\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e \u003cp\u003eMax\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c8\"\u003e \u003cp\u003eMean\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c9\"\u003e \u003cp\u003eSD\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eCampus\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.66\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003e5.31\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003e2.68\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1.08\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003e-29.26\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e \u003cp\u003e-18.99\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c8\"\u003e \u003cp\u003e-26.38\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c9\"\u003e \u003cp\u003e1.84\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eMain\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e1.95\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e5.31\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e3.36\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.92\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e-29.09\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e-21.98\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e-26.43\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e1.42\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eNorth\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.66\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e3.70\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1.97\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.71\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e-29.26\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e-18.99\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e-26.34\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e2.22\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eHoekwil\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003e0.55\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cb\u003e5.15\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cb\u003e2.55\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003e1.21\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e\u003cb\u003e-29.82\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cb\u003e-25.67\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e\u003cb\u003e-28.13\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e\u003cb\u003e0.93\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eOakhurst\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e2.24\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e5.15\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e3.34\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.72\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e-29.82\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e-27.54\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e-28.65\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e0.62\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eWoodville\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.55\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e4.29\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1.75\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1.08\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e-28.90\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e-25.67\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e-27.60\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e0.91\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eKnysna\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003e-0.02\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cb\u003e3.12\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cb\u003e1.58\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003e0.69\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e\u003cb\u003e-29.96\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cb\u003e-24.24\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e\u003cb\u003e-28.03\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e\u003cb\u003e0.80\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eGoudveld\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.62\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e3.12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1.65\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.70\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e-29.96\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e-24.24\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e-27.92\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e0.91\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eDiepwalle\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e-0.02\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e3.08\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1.51\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.69\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e-29.72\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e-27.01\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e-28.14\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e0.67\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003eA significant increase in the δ\u003csup\u003e15\u003c/sup\u003eN ratios was visible along with the level of anthropization of the environment across the land use gradient (R\u0026thinsp;=\u0026thinsp;0.45, P-value\u0026thinsp;=\u0026thinsp;1.4e\u003csup\u003e\u0026minus;\u0026thinsp;08\u003c/sup\u003e; Fig.\u0026nbsp;2c), created using the second axis of the PCA from Fig.\u0026nbsp;2a. The samples collected at the Campus (highest proportion of built up land cover), characterized by the highest anthropization levels, showed higher ratios of δ\u003csup\u003e15\u003c/sup\u003eN. The samples from Hoekwil (highest proportion of agriculture), with lower levels of built up land cover, presented intermediate δ\u003csup\u003e15\u003c/sup\u003eN numbers. The Knysna samples (highest proportion of forest), reflecting the lowest anthropization levels and more natural habitat, showed the lowest values for δ\u003csup\u003e15\u003c/sup\u003eN. The majority of the Campus samples, as well as all those from Hoekwil and Knysna, reflected an isotopic signature between about \u0026minus;\u0026thinsp;30\u0026permil; and \u0026minus;\u0026thinsp;24\u0026permil; for δ\u003csup\u003e13\u003c/sup\u003eC values (specific to C\u003csub\u003e3\u003c/sub\u003e plants), except four of the Campus samples that possessed slightly higher values (Fig.\u0026nbsp;2d). Higher values correspond to the more urban areas.\u003c/p\u003e \u003cp\u003eThe data for both δ\u003csup\u003e15\u003c/sup\u003eN and δ\u003csup\u003e13\u003c/sup\u003eC were found to be non-parametric (Shapiro-Wilk, p-value\u0026thinsp;=\u0026thinsp;3.89e-4 for δ\u003csup\u003e15\u003c/sup\u003eN and 1.50e-12 for δ\u003csup\u003e13\u003c/sup\u003eC) with unequal variances between the three locations (Levene, Pr(\u0026gt;\u0026thinsp;F)\u0026thinsp;=\u0026thinsp;7.59e\u003csup\u003e\u0026minus;\u0026thinsp;4\u003c/sup\u003e for δ\u003csup\u003e15\u003c/sup\u003eN and 1.01e\u003csup\u003e\u0026minus;\u0026thinsp;2\u003c/sup\u003e for δ\u003csup\u003e13\u003c/sup\u003eC). Kruskal-Wallis tests indicated that at least one location was significantly different from the others for nitrogen (chi-squared\u0026thinsp;=\u0026thinsp;32.97, df\u0026thinsp;=\u0026thinsp;2, p-value\u0026thinsp;=\u0026thinsp;6.92e\u003csup\u003e\u0026minus;\u0026thinsp;08\u003c/sup\u003e) and carbon (chi-squared\u0026thinsp;=\u0026thinsp;57.92, df\u0026thinsp;=\u0026thinsp;2, p-value\u0026thinsp;=\u0026thinsp;2.65e\u003csup\u003e\u0026minus;\u0026thinsp;13\u003c/sup\u003e). Post-hoc Dunn tests revealed that the Knysna samples significantly differed from both the Campus (p-value\u0026thinsp;=\u0026thinsp;1.04e-07) and Hoekwil (p-value\u0026thinsp;=\u0026thinsp;4.77e\u003csup\u003e\u0026minus;\u0026thinsp;04\u003c/sup\u003e) samples for nitrogen (Fig.\u0026nbsp;3a), whereas for carbon ratios, the Campus samples were significantly different from both Hoekwil (p-value\u0026thinsp;=\u0026thinsp;8.88e\u003csup\u003e\u0026minus;\u0026thinsp;09\u003c/sup\u003e) and Knysna (p-value\u0026thinsp;=\u0026thinsp;3.28e\u003csup\u003e\u0026minus;\u0026thinsp;11\u003c/sup\u003e; Fig.\u0026nbsp;3b). According to these tests, nitrogen ratios did not significantly differ between Hoekwil and Campus samples (Dunn, p-value\u0026thinsp;=\u0026thinsp;4.53e\u003csup\u003e\u0026minus;\u0026thinsp;01\u003c/sup\u003e), nor did Knysna and Hoekwil for carbon samples (Dunn, p-value\u0026thinsp;=\u0026thinsp;6.77e\u003csup\u003e\u0026minus;\u0026thinsp;01\u003c/sup\u003e).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec12\" class=\"Section2\"\u003e \u003ch2\u003eIsotopic distribution variations between troops at the Campus site\u003c/h2\u003e \u003cp\u003eThe resulting PCA biplot for the inter-troop analysis on the two subsites of the Campus (Main and North troops) showed little difference in distribution according to land use. The landscape surrounding the samples only differed slightly along the first axis according to the troop, which accounted for 70.9% of the variation (Fig.\u0026nbsp;4a). The Main troop samples tended towards higher levels of built up land use as well as the presence of roads, whereas the North troop spanned a wider range across the axis, going from built up to more natural landscapes (Fig.\u0026nbsp;4a).\u003c/p\u003e \u003cp\u003eDespite these limited differences in the landscapes used by these two troops of baboons, δ\u003csup\u003e15\u003c/sup\u003eN nitrogen ratios of the fecal samples from these two subsites varied significantly (Student\u0026rsquo;s t-test, t\u0026thinsp;=\u0026thinsp;6.50, df\u0026thinsp;=\u0026thinsp;57, p-value\u0026thinsp;=\u0026thinsp;1.09e-08) and according to the level of land use created using the first axis of the inter-troop PCA biplot (Fig.\u0026nbsp;4a and Fig.\u0026nbsp;4b). Higher δ\u003csup\u003e15\u003c/sup\u003eN values were associated with the Main troop (1.95 to 5.31\u0026permil;; Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e), which is represented by built up land cover and higher anthropization levels. The North troop nitrogen ratios had a wider range, spanning from ratios similar to those of the Main samples down to lower values of δ\u003csup\u003e15\u003c/sup\u003eN along with the anthropization level and towards areas characterized by more natural forests (0.66 to 3.70\u0026permil;; Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe carbon δ\u003csup\u003e13\u003c/sup\u003eC ratios of these two subsites reflected a similar distribution to those of all three locations: no significant differences were present between the two troops, whose ratios were contained between \u0026minus;\u0026thinsp;24\u0026permil; and \u0026minus;\u0026thinsp;30\u0026permil; (Wilcoxon, W\u0026thinsp;=\u0026thinsp;450, p-value\u0026thinsp;=\u0026thinsp;0.83; Fig.\u0026nbsp;4c; SI 1). It is noticeable that the highest δ\u003csup\u003e13\u003c/sup\u003eC values of two subsite samples of the Campus were specific to the North troop (Fig.\u0026nbsp;4c).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e"},{"header":"Discussion","content":"\u003cp\u003eIn South Africa, chacma baboons are highly present in anthropized landscapes, possibly most notably in the Western Cape province. In Cape Town, as well as across the Garden Route region, chacma baboons and humans often occupy overlapping habitats, where increased conflicts have been reported associated mostly to stealing food (Chowdhury et al. \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Mormile and Hill \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). Hoffman and O\u0026rsquo;Rian showed that Cape peninsula baboon troops present in human-modified habitats are associated with increased events of observed stealing from people\u0026rsquo;s houses and visiting the bins around settlements (Hoffman and O\u0026rsquo;Riain \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). Mazue et al. also described that the Main troop living on the campus of Nelson Mandela University (George Campus), representing a peri-urban area, was responsible for an unexpected number of raids: 70 raids were recorded within a two-week study (unpublished data, 2016). Overall, it appears that the fragmentation and the modification of their natural habitats has pushed chacma baboons to co-exist with humans within modified landscapes, where new human-modified food resources are easily available, which could have an impact on their foraging behaviour. In the present study, the analysis of nitrogen and carbon stable isotopic ratios in chacma baboon troops inhabiting a gradient of human-modified landscapes in the Garden Route region showed significant differences in their diet in terms of protein and plant intake.\u003c/p\u003e \u003cdiv id=\"Sec14\" class=\"Section2\"\u003e \u003ch2\u003eA shift towards higher protein intake in anthropized areas\u003c/h2\u003e \u003cp\u003eIt is well known that \u003csup\u003e15\u003c/sup\u003eN/\u003csup\u003e14\u003c/sup\u003eN ratios in mammalian excrements are related to a combination of variables, driven by the complexity of the nitrogen cycle (Handley and Raven \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e1992\u003c/span\u003e). The latter leads to a variety of δ\u003csup\u003e15\u003c/sup\u003eN patterns in soil and plants that are consumed by animals higher up the trophic chain. When consumed, \u003csup\u003e15\u003c/sup\u003eN is preferably incorporated into tissues over \u003csup\u003e14\u003c/sup\u003eN, creating a stepwise increase with each trophic level and also increasing with protein intake (Handley and Raven \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e1992\u003c/span\u003e; Schoeninger and DeNiro \u003cspan citationid=\"CR79\" class=\"CitationRef\"\u003e1984\u003c/span\u003e; Sponheimer et al. \u003cspan citationid=\"CR86\" class=\"CitationRef\"\u003e2003\u003c/span\u003e). In the present study, the \u003csup\u003e15\u003c/sup\u003eN/\u003csup\u003e14\u003c/sup\u003eN results showed significantly higher protein intake for chacma baboons in landscapes transformed by human activity, \u003cem\u003ei.e\u003c/em\u003e. on the Campus with built up land cover and in Hoekwil with agricultural zones (Fig.\u0026nbsp;2c and 3a). This suggests that the chacma baboon populations known to live in close contact with human populations in the Garden Route have been shifting their diet to a richer nitrogen pattern when inhabiting more anthropized lands. This is the case whether agricultural or peri-urban, a result probably associated with the easier access to human-modified resources. Such results could be explained by the easy access to protein sources in such areas, such as garbage bins, meat detritus from the cafeteria on the Campus site or the houses and farmsteads around the Hoekwil area. Particularly on Campus, the kitchens and bins provide regular food for the baboons almost all year round, including remaining meals from students that contain large amounts of meat that would not be accessible to them in the wild. This observation was even confirmed when considering the smaller scale of the Campus site, where higher values of δ\u003csup\u003e15\u003c/sup\u003eN were observed to be associated with the Main troop, in comparison to the North one (Fig.\u0026nbsp;4b). This chacma baboon Main troop is followed daily by monitors and has been reported to spend considerable time on George Campus where they regularly visit buildings and bins (Mazu\u0026eacute; et al. \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e2023\u003c/span\u003e), which could again explain the differences observed between the Main and the North troops at the Campus location regarding nitrogen intake. This study showed that the nitrogen ratios in the fecal samples, and therefore protein intake, were significantly highest in the populations inhabiting the most anthropized areas where the most buildings, built up land cover and transformed landscape for agriculture can be observed and where a higher access to human modified food highly composed of meat is present.\u003c/p\u003e \u003cp\u003eSome studies have also described higher protein intake by wild animals inhabiting human-modified environments, such as rats, that have been shown to consume more protein-rich resources in anthropized areas (Guiry and Buckley \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). While these studies showed the same pattern and confirmed that certain wild animal populations inhabiting anthropogenic landscapes present higher protein intake in comparison to natural ones (Guiry and Buckley \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2018\u003c/span\u003e), others described different patterns with either lower nitrogen intake in anthropized environments or no significant difference in relation to landscape use. Some birds and carnivorous mammal species such as foxes or coyotes have been found to possess protein-poor diets in anthropized areas in comparison to populations in rural zones (Francis et al. \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Heiss et al. \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Murray et al. \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Newsome et al. \u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Scholz et al. \u003cspan citationid=\"CR80\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Moreover, a study from Nabugabo Field Station in central Uganda showed that for vervet monkeys consuming crops, there were no resulting variations in protein, fibre or lipid composition in comparison to wild populations (Cancelliere et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). The same pattern has been observed for wild boars, for which similar protein levels have been described across a gradient of modified landscapes, even though the species has been proven to be present near human settlements (Cahill et al. \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Stillfried et al. \u003cspan citationid=\"CR87\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). The work from G\u0026aacute;mez et al. (2020) also evidenced that overall, δ\u003csup\u003e15\u003c/sup\u003eN ratios did not differ between urban and rural or wild land predators suggesting that the protein intake was not influenced by landscape modifications. It then appears that depending on the species studied and probably the geographic location of the study, wild animals respond differently to human modified resources. In the present study, the results reveal a significant shift toward higher protein intake in chacma baboons\u0026rsquo; diet inhabiting anthropized areas in South Africa, but this will need to be confirmed through replicates along the same landscape gradient in multiple troops.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec15\" class=\"Section2\"\u003e \u003ch2\u003eVariations in plant consumption patterns across anthropized areas\u003c/h2\u003e \u003cp\u003eIt is well established that C\u003csub\u003e3\u003c/sub\u003e plants tend to have lower \u003csup\u003e13\u003c/sup\u003eC/\u003csup\u003e12\u003c/sup\u003eC ratios than C\u003csub\u003e4\u003c/sub\u003e plants: the δ\u003csup\u003e13\u003c/sup\u003eC values of C\u003csub\u003e3\u003c/sub\u003e plants range from \u0026minus;\u0026thinsp;34 to -22\u0026permil; and those of C\u003csub\u003e4\u003c/sub\u003e plants from \u0026minus;\u0026thinsp;16 to -10\u0026permil; (Basu et al. \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). In the Garden Route temperate region, it has been shown that most trees and shrubs follow the C\u003csub\u003e3\u003c/sub\u003e-photosynthetic pathway (Ehleringer and Cerling \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2002\u003c/span\u003e), so we could expect chacma baboon\u0026rsquo;s diet living in natural forest to be richer in C\u003csub\u003e3\u003c/sub\u003e than in C\u003csub\u003e4\u003c/sub\u003e plants. The results obtained in this study showed that the large majority of the collected samples were contained within the bracket that reflect a C\u003csub\u003e3\u003c/sub\u003e ecosystem, suggesting that as expected the overall consumption of the chacma baboon troops studied was made up of C\u003csub\u003e3\u003c/sub\u003e plants (Fig.\u0026nbsp;2d). However, small variations have to be considered regarding the δ\u003csup\u003e13\u003c/sup\u003eC isotopic ratios, as some samples collected at the highly anthropized Campus location possessed a higher proportion of C\u003csub\u003e4\u003c/sub\u003e plants than those from Hoekwil and Knysna, suggesting more consumption of C\u003csub\u003e4\u003c/sub\u003e components during recent foraging by certain individuals (Fig.\u0026nbsp;2d). This could be associated with the fact that higher δ\u003csup\u003e13\u003c/sup\u003eC isotopic ratios can be explained by the consumption of anthropogenic foods that are known to contain large quantities of C\u003csub\u003e4\u003c/sub\u003e plants (\u003cem\u003ei.e\u003c/em\u003e. sugar, corn and wheat; G\u0026aacute;mez et al. 2020; Nicholson and Cove \u003cspan citationid=\"CR68\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Post et al. \u003cspan citationid=\"CR75\" class=\"CitationRef\"\u003e2007\u003c/span\u003e). The same pattern has been observed for instance in eurasian red squirrels (\u003cem\u003eSciurus vulgaris\u003c/em\u003e) that consumed more sugar in urban zones (Wist et al. \u003cspan citationid=\"CR96\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). In the present study, the overall consumption of the chacma baboon troops studied was made up of C\u003csub\u003e3\u003c/sub\u003e plants with for a significantly higher δ\u003csup\u003e13\u003c/sup\u003eC ratios in the Campus suggesting that the troops in this area may have access to and consume a higher proportion of C\u003csub\u003e4\u003c/sub\u003e plants (\u003cem\u003ei.e\u003c/em\u003e. sugar, corn and wheat), presumably due to the access to garbage and kitchens, as described previously for nitrogen.\u003c/p\u003e \u003cp\u003eAt a smaller scale, a few samples from the most anthropized subsites (on Campus) particularly stood out, possessing higher δ\u003csup\u003e13\u003c/sup\u003eC values, which could be as discussed above the result of higher C\u003csub\u003e4\u003c/sub\u003e plant consumption by specific individuals (Fig.\u0026nbsp;4c). This raises the possible influence of other factors including social behaviour that could contribute to variations in diet in primates that can influence carbon intake. Indeed, adult males of several macaque species have been shown to consume more human resources than adult females, younger and lower ranked individuals (Maibeche et al. \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2015\u003c/span\u003e, Marty et al. \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Focusing management (\u003cem\u003ei.e.\u003c/em\u003e monitors) on certain high ranking individuals may be helpful to reduce their consumption of human resources, but can also provide lower ranking individuals with easier access to these resources (Bracken et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2022b\u003c/span\u003e), thereby failing to reduce conflict with humans. To address such questions, future studies should integrate stable isotope analyses with complementary behavioural analyses to get a better understanding of the chacma baboons\u0026rsquo; behavioural ecology and to establish better strategies for conflict management.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec16\" class=\"Section2\"\u003e \u003ch2\u003ePerspectives and future impacts\u003c/h2\u003e \u003cp\u003eThis present study showed that land use influences chacma baboons\u0026rsquo; diet, with a shift in anthropized environments toward a globally significant higher nitrogen and for some individuals higher carbon patterns. This quantitative analysis gives a first picture of land use impact on this species diet, however we are still far from identifying exactly what chacma baboons consume in different environments. In this context, future studies should combine stable isotope analysis with observation and/or DNA metabarcoding in order to extract the precise composition and species richness of the baboon\u0026rsquo;s diet along multiple land uses.\u003c/p\u003e \u003cp\u003eMoreover, seasonal variations can also account for differences in carbon ratios in baboon diet across the year: Lewis et al. (\u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2018\u003c/span\u003e) provided results that showed that summer and autumn δ\u003csup\u003e13\u003c/sup\u003eC ratios were higher than those in winter and lowest in spring. In the future, studies should therefore also focus their efforts on the analysis of the stable isotopic ratios of larger sampling size and of fecal samples collected over several seasons to avoid missing any individual or seasonal effects (Mychajliw et al. \u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e2022\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThis study has shown that chacma baboons consume more human resourced foods in more anthropogenic landscapes, and it has previously been described that despite some benefits, the shift in trophic niche associated with human-modified resources can also be detrimental to many wild animals. Unfortunately, to date health studies tend to revolve around humans and few have focused on the extent and direct impacts that the consumption of anthropogenic food by wild animals can have on their health. Several diseases found in humans (e.g. obesity, cardio-metabolic or cardiovascular risks, cancer, type-2 diabetes) are linked to the consumption of processed foods, refined sugars, certain grains or fats (Elizabeth et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Monteiro et al. \u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Popkin et al. \u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e2020\u003c/span\u003e) that, according to this study, species such as chacma baboons have increasing access to. The question is now to determine what specific human-modified resources non-human primates are consuming in urban and peri-urban areas, and if this is causing health problems (e.g. short or long term diseases; Sapolsky and Share \u003cspan citationid=\"CR78\" class=\"CitationRef\"\u003e2004\u003c/span\u003e) as well as affecting other aspects of their lives (i.e. behavioral, social, physiological, stress; Bracken et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2022b\u003c/span\u003e; Chowdhury et al. \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Mazu\u0026eacute; et al. \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Understanding these effects is important for future management in the growing context of human-wildlife encounters and conflict.\u003c/p\u003e \u003c/div\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eStatements \u0026amp; Declarations\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eSupplementary Information\u0026nbsp;\u003c/strong\u003eThe online version contains supplementary material available at XXX\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAcknowledgements\u0026nbsp;\u003c/strong\u003eWe are grateful to SANParks for allowing and participating in data collection throughout this study. We would especially like to thank all SANParks personnel who provided us with not only access and guidance throughout the park, but who became personally involved in the research project, particularly Nathan and Dylan. We also thank the Wildlife Ecology Lab and the students that participated in data collection and analysis, namely Eloise Krisch, Sinayo Lungile, Brittany McClear, Tessa Raw, Markus Woesner, Bianca Coulson and especially the NMU Campus Baboon Monitors, Dietre Stols, Liyabona Ntlokwana, Nusheen Brenner and Abulele Sixonxo who provided us with remarkable insight into the Campus baboon habits. Thank you to all the people who provided details on troop locations throughout the Garden Route and granted us access to their properties, including the Oakhurst Farm and the personnel of the Woodville Trading Store.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFunding\u0026nbsp;\u003c/strong\u003eThis work was supported by the funding of Virginie Rougeron from Centre National de la Recherche Scientifique, REHABS International Research Laboratory.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eData availability and material\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eEthics declarations\u0026nbsp;\u003c/strong\u003eData collection was possible according to SANParks (BERN-A/2020-008) and Cape Nature (CN44-87-16198) permits, and analysis was allowed under the Section 20 of the Animal Diseases Act, 1984 (Act No 35 of 1984).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAuthorship statements\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eConception: V.R., F.P., H.F. and C.G.\u003c/p\u003e\n\u003cp\u003eFunding acquisition: V.R.\u003c/p\u003e\n\u003cp\u003eBiological data acquisition and management: V.R., F.P. E.K.O. and A.B.\u003c/p\u003e\n\u003cp\u003eStable isotopic data acquisition and analysis: E.K.O.\u003c/p\u003e\n\u003cp\u003eG.I.S analysis : E.K.O. and P.C.R.\u003c/p\u003e\n\u003cp\u003eInterpretation of results: E.K.O, V.R., F.P., H.F. and C.G.\u003c/p\u003e\n\u003cp\u003eDrafting of the manuscript: E.K.O and V.R.\u003c/p\u003e\n\u003cp\u003eReviewing and editing the manuscript: E.K.O, V.R., F.P., H.F., P.C.R., C.G., A.B. and G.H\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eAllan ATL, LaBarge LR, Howlett C et al (2022) Patterns of predation and meat-eating by chacma baboons in an Afromontane environment. Folia Primatol 1\u0026ndash;24. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1163/14219980-bja10004\u003c/span\u003e\u003cspan address=\"10.1163/14219980-bja10004\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eAndo H, Mukai H, Komura T et al (2020) Methodological trends and perspectives of animal dietary studies by noninvasive fecal DNA metabarcoding. Environ DNA 2:391\u0026ndash;406. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1002/edn3.117\u003c/span\u003e\u003cspan address=\"10.1002/edn3.117\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eAplin KP, Suzuki H, Chinen AA et al (2011) Multiple Geographic Origins of Commensalism and Complex Dispersal History of Black Rats. PLoS ONE 6:e26357. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1371/journal.pone.0026357\u003c/span\u003e\u003cspan address=\"10.1371/journal.pone.0026357\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eBasu S, Agrawal S, Sanyal P et al (2015) Carbon isotopic ratios of modern C3\u0026ndash;C4 plants from the Gangetic Plain, India and its implications to paleovegetational reconstruction. Palaeogeogr Palaeoclimatol Palaeoecol 440:22\u0026ndash;32. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.palaeo.2015.08.012\u003c/span\u003e\u003cspan address=\"10.1016/j.palaeo.2015.08.012\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eBellon B, Zander S, De Vos A, Renaud P-C CoexistLand. In: Earth Engine Apps. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://zandersamuel.users.earthengine.app/view/coexistland\u003c/span\u003e\u003cspan address=\"https://zandersamuel.users.earthengine.app/view/coexistland\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e. Accessed 20 Apr 2022\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eBoecklen WJ, Yarnes CT, Cook BA, James AC (2011) On the Use of Stable Isotopes in Trophic Ecology. Annu Rev Ecol Evol Syst 42:411\u0026ndash;440. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1146/annurev-ecolsys-102209-144726\u003c/span\u003e\u003cspan address=\"10.1146/annurev-ecolsys-102209-144726\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eBracken AM, Christensen C, O\u0026rsquo;Riain MJ et al (2022a) Flexible group cohesion and coordination, but robust leader\u0026ndash;follower roles, in a wild social primate using urban space. Proc R Soc B 289:20212141. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1098/rspb.2021.2141\u003c/span\u003e\u003cspan address=\"10.1098/rspb.2021.2141\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eBracken AM, Christensen C, O\u0026rsquo;Riain MJ et al (2023) Postpartum cessation of urban space use by a female baboon living at the edge of the City of Cape Town. Ecol Evol 13:e9963. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1002/ece3.9963\u003c/span\u003e\u003cspan address=\"10.1002/ece3.9963\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eBracken AM, Christensen C, O\u0026rsquo;Riain MJ et al (2022b) Socioecology Explains Individual Variation in Urban Space Use in Response to Management in Cape Chacma Baboons (Papio ursinus). Int J Primatol 43:1159\u0026ndash;1176. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s10764-021-00247-x\u003c/span\u003e\u003cspan address=\"10.1007/s10764-021-00247-x\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eBurbank J, Drake DAR, Power M (2022) Use of stable isotopes for assessing urbanization impacts on freshwater fishes. Front Environ Sci 10:963693. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.3389/fenvs.2022.963693\u003c/span\u003e\u003cspan address=\"10.3389/fenvs.2022.963693\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eCahill S, Llimona F et al (2012) Consorci del Parc Natural de la Serra de Collserola, Barcelona, Spain, Characteristics of wild boar (Sus scrofa) habituation to urban areas in the Collserola Natural Park (Barcelona) and comparison with other locations. Anim Biodiv Conserv 35:221\u0026ndash;233. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.32800/abc.2012.35.0221\u003c/span\u003e\u003cspan address=\"10.32800/abc.2012.35.0221\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eCancelliere EC, Chapman CA, Twinomugisha D, Rothman JM (2018) The nutritional value of feeding on crops: Diets of vervet monkeys in a humanized landscape. Afr J Ecol 56:160\u0026ndash;167. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1111/aje.12496\u003c/span\u003e\u003cspan address=\"10.1111/aje.12496\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eCaut S, Angulo E, Courchamp F (2008) Discrimination factors (∆ \u003csup\u003e15\u003c/sup\u003e N and ∆ \u003csup\u003e13\u003c/sup\u003e C) in an omnivorous consumer: effect of diet isotopic ratio. Funct Ecol 22:255\u0026ndash;263. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1111/j.1365-2435.2007.01360.x\u003c/span\u003e\u003cspan address=\"10.1111/j.1365-2435.2007.01360.x\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eChiyo PI, Lee PC, Moss CJ et al (2011) No risk, no gain: effects of crop raiding and genetic diversity on body size in male elephants. Behav Ecol 22:552\u0026ndash;558. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1093/beheco/arr016\u003c/span\u003e\u003cspan address=\"10.1093/beheco/arr016\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eChowdhury S, Brown J, Swedell L (2020) Anthropogenic effects on the physiology and behaviour of chacma baboons in the Cape Peninsula of South Africa. Conserv Physiol 8:coaa066. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1093/conphys/coaa066\u003c/span\u003e\u003cspan address=\"10.1093/conphys/coaa066\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eCodron D, Codron J, Lee-Thorp JA et al (2007) Stable isotope characterization of mammalian predator\u0026ndash;prey relationships in a South African savanna. Eur J Wildl Res 53:161\u0026ndash;170. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s10344-006-0075-x\u003c/span\u003e\u003cspan address=\"10.1007/s10344-006-0075-x\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eCodron D, Lee-Thorp JA, Sponheimer M et al (2006) Inter- and intrahabitat dietary variability of chacma baboons (Papio ursinus) in South African savannas based on fecal δ13C, δ15N, and %N. Am J Phys Anthropol 129:204\u0026ndash;214. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1002/ajpa.20253\u003c/span\u003e\u003cspan address=\"10.1002/ajpa.20253\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eCrowley BE, Reitsema LJ, Oelze VM, Sponheimer M (2016) Advances in primate stable isotope ecology-Achievements and future prospects: Advances in Primate Isotope Ecology. Am J Primatol 78:995\u0026ndash;1003. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1002/ajp.22510\u003c/span\u003e\u003cspan address=\"10.1002/ajp.22510\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eCypher BL, Frost N (1999) Condition of San Joaquin Kit Foxes in Urban and Exurban Habitats. J Wildl Manag 63:930. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.2307/3802807\u003c/span\u003e\u003cspan address=\"10.2307/3802807\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eDale VH, Polasky S (2007) Measures of the effects of agricultural practices on ecosystem services. Ecol Econ 64:286\u0026ndash;296. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.ecolecon.2007.05.009\u003c/span\u003e\u003cspan address=\"10.1016/j.ecolecon.2007.05.009\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eDe Barba M, Miquel C, Boyer F et al (2014) DNA metabarcoding multiplexing and validation of data accuracy for diet assessment: application to omnivorous diet. Mol Ecol Resour 14:306\u0026ndash;323. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1111/1755-0998.12188\u003c/span\u003e\u003cspan address=\"10.1111/1755-0998.12188\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eDe Carvalho DR, de Castro DMP, Callisto M et al (2019) Stable isotopes and stomach content analyses indicate omnivorous habits and opportunistic feeding behavior of an invasive fish. Aquat Ecol 53:365\u0026ndash;381. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s10452-019-09695-3\u003c/span\u003e\u003cspan address=\"10.1007/s10452-019-09695-3\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eDepartment of Forestry, Fisheries and the Environment (DFFE) (2021) SA National Land-Cover Datasets. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://egis.environment.gov.za/sa_national_land_cover_datasets\u003c/span\u003e\u003cspan address=\"https://egis.environment.gov.za/sa_national_land_cover_datasets\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e. Accessed 15 Mar 2022\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eEhleringer JR, Cerling TE (2002) C3 and C4 Photosynthesis. The Earth system: biological and ecological dimensions of global environmental change, 2:186\u0026ndash;190\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eElizabeth L, Machado P, Zin\u0026ouml;cker M et al (2020) Ultra-Processed Foods and Health Outcomes: A Narrative Review. Nutrients 12:1955. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.3390/nu12071955\u003c/span\u003e\u003cspan address=\"10.3390/nu12071955\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eEllis EC, Gauthier N, Klein Goldewijk K et al (2021) People have shaped most of terrestrial nature for at least 12,000 years. Proc Natl Acad Sci USA 118:e2023483118. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1073/pnas.2023483118\u003c/span\u003e\u003cspan address=\"10.1073/pnas.2023483118\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eElmqvist T, Zipperer WC, G\u0026uuml;neralp B (2016) Urbanization, habitat loss, biodiversity decline: solution pathways to break the cycle. In, Seta K, Solecki WD, Griffith CA (eds.) Routledge Handbook of Urbanization and Global Environmental Change. Routledge, London and New York. Chapter 10: 139\u0026ndash;151\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eEriksson O (2013) Species pools in cultural landscapes - niche construction, ecological opportunity and niche shifts. Ecography 36:403\u0026ndash;413. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1111/j.1600-0587.2012.07913.x\u003c/span\u003e\u003cspan address=\"10.1111/j.1600-0587.2012.07913.x\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eEvans BA, Gawlik DE (2020) Urban food subsidies reduce natural food limitations and reproductive costs for a wetland bird. Sci Rep 10:14021. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1038/s41598-020-70934-x\u003c/span\u003e\u003cspan address=\"10.1038/s41598-020-70934-x\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eFahrig L (2003) Effects of Habitat Fragmentation on Biodiversity. Annu Rev Ecol Evol Syst 34:487\u0026ndash;515. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1146/annurev.ecolsys.34.011802.132419\u003c/span\u003e\u003cspan address=\"10.1146/annurev.ecolsys.34.011802.132419\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eFarr TG, Kobrick M (2000) Shuttle Radar Topography Mission\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eFiolet T, Srour B, Sellem L et al (2018) Consumption of ultra-processed foods and cancer risk: results from NutriNet-Sant\u0026eacute; prospective cohort. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1136/bmj.k322\u003c/span\u003e\u003cspan address=\"10.1136/bmj.k322\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e. BMJ k322\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eFischer J, Higham JP, Alberts SC et al (2019) Insights into the evolution of social systems and species from baboon studies. eLife 8:e50989. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.7554/eLife.50989\u003c/span\u003e\u003cspan address=\"10.7554/eLife.50989\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eFleming PA, Bateman PW (2018) Novel predation opportunities in anthropogenic landscapes. Anim Behav 138:145\u0026ndash;155. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.anbehav.2018.02.011\u003c/span\u003e\u003cspan address=\"10.1016/j.anbehav.2018.02.011\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eFoley JA, DeFries R, Asner GP et al (2005) Global Consequences of Land Use. Science 309:570\u0026ndash;574. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1126/science.1111772\u003c/span\u003e\u003cspan address=\"10.1126/science.1111772\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eFrancis RJ, Kingsford RT, Murray-Hudson M, Brandis KJ (2021) Urban waste no replacement for natural foods\u0026mdash;Marabou storks in Botswana. J Urban Ecol 7:juab003. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1093/jue/juab003\u003c/span\u003e\u003cspan address=\"10.1093/jue/juab003\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eG\u0026aacute;mez S, Potts A, Mills KL et al (2022) Downtown diet: a global meta-analysis of increased urbanization on the diets of vertebrate predators. Proc R Soc B 289:20212487. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1098/rspb.2021.2487\u003c/span\u003e\u003cspan address=\"10.1098/rspb.2021.2487\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eGuiry E, Buckley M (2018) Urban rats have less variable, higher protein diets. Proc R Soc B 285:20181441. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1098/rspb.2018.1441\u003c/span\u003e\u003cspan address=\"10.1098/rspb.2018.1441\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eHandley LL, Raven JA (1992) The use of natural abundance of nitrogen isotopes in plant physiology and ecology. Plant Cell Environ 15:965\u0026ndash;985\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eHeiss RS, Clark AB, McGowan KJ (2009) Growth and nutritional state of American Crow nestlings vary between urban and rural habitats. Ecol Appl 19:829\u0026ndash;839. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1890/08-0140.1\u003c/span\u003e\u003cspan address=\"10.1890/08-0140.1\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eHoffman TS, O\u0026rsquo;Riain MJ (2012) Landscape requirements of a primate population in a human-dominated environment. Front Zool 9:1. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1186/1742-9994-9-1\u003c/span\u003e\u003cspan address=\"10.1186/1742-9994-9-1\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eHoffman TS, O\u0026rsquo;Riain MJ (2011) The Spatial Ecology of Chacma Baboons (Papio ursinus) in a Human-modified Environment. Int J Primatol 32:308\u0026ndash;328. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s10764-010-9467-6\u003c/span\u003e\u003cspan address=\"10.1007/s10764-010-9467-6\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eHooke RLeB, Mart\u0026iacute;n-Duque JF (2012) Land transformation by humans: A review. GSAT 12:4\u0026ndash;10. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1130/GSAT151A.1\u003c/span\u003e\u003cspan address=\"10.1130/GSAT151A.1\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eHulme-Beaman A, Dobney K, Cucchi T, Searle JB (2016) An Ecological and Evolutionary Framework for Commensalism in Anthropogenic Environments. Trends in Ecology and Evolution 31:633\u0026ndash;645. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.tree.2016.05.001\u003c/span\u003e\u003cspan address=\"10.1016/j.tree.2016.05.001\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eJohnson C, Piel AK, Forman D et al (2015) The ecological determinants of baboon troop movements at local and continental scales. Mov Ecol 3:14. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1186/s40462-015-0040-y\u003c/span\u003e\u003cspan address=\"10.1186/s40462-015-0040-y\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eJohnson CA, Raubenheimer D, Rothman JM et al (2013) 30 Days in the Life: Daily Nutrient Balancing in a Wild Chacma Baboon. PLoS ONE 8:e70383. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1371/journal.pone.0070383\u003c/span\u003e\u003cspan address=\"10.1371/journal.pone.0070383\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eJordan MJR (2005) Dietary analysis for mammals and birds: a review of field techniques and animal-management applications. Int Zoo Yearbook 39:108\u0026ndash;116. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1111/j.1748-1090.2005.tb00010.x\u003c/span\u003e\u003cspan address=\"10.1111/j.1748-1090.2005.tb00010.x\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eLee JH, Duster M, Roberts T, Devinsky O (2022) United States Dietary Trends Since 1800: Lack of Association Between Saturated Fatty Acid Consumption and Non-communicable Diseases. Front Nutr 8:748847. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.3389/fnut.2021.748847\u003c/span\u003e\u003cspan address=\"10.3389/fnut.2021.748847\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eLee S-M, Lee E-J (2019) Diet of the wild boar (\u003cem\u003eSus scrofa\u003c/em\u003e): implications for management in forest-agricultural and urban environments in South Korea. PeerJ 7:e7835. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.7717/peerj.7835\u003c/span\u003e\u003cspan address=\"10.7717/peerj.7835\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eLee P, Priston N (2005) Human attitudes to primates: Perceptions of pests, conflict and consequences for primate conservation. Commensalism and Conflict: The Human-Primate Interface. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1002/047001539X.ch1\u003c/span\u003e\u003cspan address=\"10.1002/047001539X.ch1\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eLewis MC, West AG, O\u0026rsquo;Riain MJ (2018) Isotopic assessment of marine food consumption by natural-foraging chacma baboons on the Cape Peninsula, South Africa. Am J Phys Anthropol 165:77\u0026ndash;93. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1002/ajpa.23332\u003c/span\u003e\u003cspan address=\"10.1002/ajpa.23332\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMaibeche Y, Moali A, Yahi N, Menard N (2015) Is Diet Flexibility an Adaptive Life Trait for Relictual and Peri-Urban Populations of the Endangered Primate Macaca. sylvanus? PLoS ONE 10:e0118596. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1371/journal.pone.0118596\u003c/span\u003e\u003cspan address=\"10.1371/journal.pone.0118596\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMart\u0026iacute;nez del Rio C, Wolf N, Carleton SA, Gannes LZ (2009) Isotopic ecology ten years after a call for more laboratory experiments. Biol Rev 84:91\u0026ndash;111. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1111/j.1469-185X.2008.00064.x\u003c/span\u003e\u003cspan address=\"10.1111/j.1469-185X.2008.00064.x\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMart\u0026iacute;nez-Fern\u0026aacute;ndez J, Ruiz-Benito P, Zavala MA (2015) Recent land cover changes in Spain across biogeographical regions and protection levels: Implications for conservation policies. Land Use Policy 44:62\u0026ndash;75. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.landusepol.2014.11.021\u003c/span\u003e\u003cspan address=\"10.1016/j.landusepol.2014.11.021\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMarty PR, Balasubramaniam KN, Kaburu SSK et al (2020) Individuals in urban dwelling primate species face unequal benefits associated with living in an anthropogenic environment. Primates 61:249\u0026ndash;255. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s10329-019-00775-4\u003c/span\u003e\u003cspan address=\"10.1007/s10329-019-00775-4\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMatos RA, Adams M, Sabat\u0026eacute; J (2021) Review: The Consumption of Ultra-Processed Foods and Non-communicable Diseases in Latin America. Front Nutr 8:622714. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.3389/fnut.2021.622714\u003c/span\u003e\u003cspan address=\"10.3389/fnut.2021.622714\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMatthews JK, Ridley A, Kaplin BA, Grueter CC (2020) A comparison of fecal sampling and direct feeding observations for quantifying the diet of a frugivorous primate. Curr Zool 66:333\u0026ndash;343. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1093/cz/zoz058\u003c/span\u003e\u003cspan address=\"10.1093/cz/zoz058\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMazu\u0026eacute; F, Guerbois C, Fritz H et al (2023) Less bins, less baboons: reducing access to anthropogenic food effectively decreases the urban foraging behavior of a troop of chacma baboons (Papio hamadryas ursinus) in a peri-urban area. Primates 64:91\u0026ndash;103. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s10329-022-01032-x\u003c/span\u003e\u003cspan address=\"10.1007/s10329-022-01032-x\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMcKinney ML (2002) Urbanization, Biodiversity, and Conservation. Bioscience 52:883. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1641/0006-3568(2002)052\u003c/span\u003e\u003cspan address=\"10.1641/0006-3568(2002)052\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e[0883:UBAC]2.0.CO;2\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMendon\u0026ccedil;a R, de Lopes D, Pimenta ACS AM, et al (2017) Ultra-Processed Food Consumption and the Incidence of Hypertension in a Mediterranean Cohort: The Seguimiento Universidad de Navarra Project. Am J Hypertens 30:358\u0026ndash;366. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1093/ajh/hpw137\u003c/span\u003e\u003cspan address=\"10.1093/ajh/hpw137\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMendon\u0026ccedil;a R, de Pimenta D, Gea AM A, et al (2016) Ultraprocessed food consumption and risk of overweight and obesity: the University of Navarra Follow-Up (SUN) cohort study. Am J Clin Nutr 104:1433\u0026ndash;1440. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.3945/ajcn.116.135004\u003c/span\u003e\u003cspan address=\"10.3945/ajcn.116.135004\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMilton K (1999) Nutritional characteristics of wild primate foods: do the diets of our closest living relatives have lessons for us? Nutrition 15:488\u0026ndash;498. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/S0899-9007(99)00078-7\u003c/span\u003e\u003cspan address=\"10.1016/S0899-9007(99)00078-7\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMonteiro CA, Cannon G, Moubarac J-C et al (2018) The UN Decade of Nutrition, the NOVA food classification and the trouble with ultra-processing. Public Health Nutr 21:5\u0026ndash;17. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1017/S1368980017000234\u003c/span\u003e\u003cspan address=\"10.1017/S1368980017000234\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMormile JE, Hill CM (2017) Living With Urban Baboons: Exploring Attitudes and Their Implications for Local Baboon Conservation and Management in Knysna, South Africa. Hum Dimensions Wildl 22:99\u0026ndash;109. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1080/10871209.2016.1255919\u003c/span\u003e\u003cspan address=\"10.1080/10871209.2016.1255919\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMurray M, Cembrowski A, Latham ADM et al (2015) Greater consumption of protein-poor anthropogenic food by urban relative to rural coyotes increases diet breadth and potential for human\u0026ndash;wildlife conflict. Ecography 38:1235\u0026ndash;1242\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMychajliw AM, Almonte JN, Martinez PA, Hadly EA (2022) Stable isotopes reveal seasonal dietary responses to agroforestry in a venomous mammal, the Hispaniolan solenodon (\u003cem\u003eSolenodon paradoxus\u003c/em\u003e). Ecol Evol 12. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1002/ece3.8761\u003c/span\u003e\u003cspan address=\"10.1002/ece3.8761\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eNewsome SD, Ralls K, Job CVH et al (2010) Stable isotopes evaluate exploitation of anthropogenic foods by the endangered San Joaquin kit fox (Vulpes macrotis mutica). J Mammal 91:1313\u0026ndash;1321. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1644/09-MAMM-A-362.1\u003c/span\u003e\u003cspan address=\"10.1644/09-MAMM-A-362.1\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eNicholson M, Cove MV (2022) Stable isotopes point to anthropogenic subsidies in northern raccoons at the urban-wild interface. Food Webs 31:e00233. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.fooweb.2022.e00233\u003c/span\u003e\u003cspan address=\"10.1016/j.fooweb.2022.e00233\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eO\u0026rsquo;Brien DM (2015) Stable Isotope Ratios as Biomarkers of Diet for Health Research. Annu Rev Nutr 35:565\u0026ndash;594. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1146/annurev-nutr-071714-034511\u003c/span\u003e\u003cspan address=\"10.1146/annurev-nutr-071714-034511\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eOro D, Genovart M, Tavecchia G et al (2013) Ecological and evolutionary implications of food subsidies from humans. Ecol Lett 16:1501\u0026ndash;1514. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1111/ele.12187\u003c/span\u003e\u003cspan address=\"10.1111/ele.12187\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003ePenick CA, Savage AM, Dunn RR (2015) Stable isotopes reveal links between human food inputs and urban ant diets. Proc R Soc B 282:20142608. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1098/rspb.2014.2608\u003c/span\u003e\u003cspan address=\"10.1098/rspb.2014.2608\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003ePollock CJ, Capilla-Lasheras P, McGill RAR et al (2017) Integrated behavioural and stable isotope data reveal altered diet linked to low breeding success in urban-dwelling blue tits (Cyanistes caeruleus). Sci Rep 7:5014. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1038/s41598-017-04575-y\u003c/span\u003e\u003cspan address=\"10.1038/s41598-017-04575-y\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003ePool-Stanvliet R, Coetzer K (2020) The scientific value of UNESCO biosphere reserves. S Afr J Sci 116. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.17159/sajs.2020/7432\u003c/span\u003e\u003cspan address=\"10.17159/sajs.2020/7432\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003ePopkin BM, Corvalan C, Grummer-Strawn LM (2020) Dynamics of the double burden of malnutrition and the changing nutrition reality. The Lancet 395:65\u0026ndash;74. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/S0140-6736(19)32497-3\u003c/span\u003e\u003cspan address=\"10.1016/S0140-6736(19)32497-3\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003ePost DM, Layman CA, Arrington DA et al (2007) Getting to the fat of the matter: models, methods and assumptions for dealing with lipids in stable isotope analyses. Oecologia 152:179\u0026ndash;189. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s00442-006-0630-x\u003c/span\u003e\u003cspan address=\"10.1007/s00442-006-0630-x\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eQGIS Development Team (2020) QGIS Geographic Information System. Version 3.18\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eRStudio T (2020) RStudio: Integrated Development for R. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://www.R-project.org/\u003c/span\u003e\u003cspan address=\"https://www.R-project.org/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eSapolsky RM, Share LJ (2004) A Pacific Culture among Wild Baboons: Its Emergence and Transmission. PLoS Biol 2:e106. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1371/journal.pbio.0020106\u003c/span\u003e\u003cspan address=\"10.1371/journal.pbio.0020106\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eSchoeninger MJ, DeNiro MJ (1984) Nitrogen and carbon isotopic composition of bone collagen from marine and terrestrial animals. Geochim Cosmochim Acta 48:625\u0026ndash;639\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eScholz C, Firozpoor J, Kramer-Schadt S et al (2020) Individual dietary specialization in a generalist predator: A stable isotope analysis of urban and rural red foxes. Ecol Evol 10:8855\u0026ndash;8870. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1002/ece3.6584\u003c/span\u003e\u003cspan address=\"10.1002/ece3.6584\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eSchreier AL, Schlaht RM, Swedell L (2019) Meat eating in wild hamadryas baboons: Opportunistic trade-offs between insects and vertebrates. Am J Primatol 81. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1002/ajp.23029\u003c/span\u003e\u003cspan address=\"10.1002/ajp.23029\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eSeoraj-Pillai N, Pillay N (2016) A Meta-Analysis of Human\u0026ndash;Wildlife Conflict: South African and Global Perspectives. Sustainability 9:34. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.3390/su9010034\u003c/span\u003e\u003cspan address=\"10.3390/su9010034\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eSeress G, S\u0026aacute;ndor K, Evans KL, Liker A (2020) Food availability limits avian reproduction in the city: An experimental study on great tits \u003cem\u003eParus major\u003c/em\u003e. J Anim Ecol 89:1570\u0026ndash;1580. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1111/1365-2656.13211\u003c/span\u003e\u003cspan address=\"10.1111/1365-2656.13211\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eSlater K, Barrett A, Brown LR (2018) Home range utilization by chacma baboon (Papio ursinus) troops on Suikerbosrand Nature Reserve, South Africa. PLoS ONE 13:e0194717. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1371/journal.pone.0194717\u003c/span\u003e\u003cspan address=\"10.1371/journal.pone.0194717\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eSmith BN, Epsten S (1971) Two Categories of 1CC/12C Ratios for Higher Plantsl. 47:5\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eSponheimer M, Robinson T, Ayliffe L et al (2003) Nitrogen isotopes in mammalian herbivores: hair * 15 N values from a controlled-feeding study. Int J Osteoarchaeol 13:80\u0026ndash;87\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eStillfried M, Gras P, Busch M et al (2017) Wild inside: Urban wild boar select natural, not anthropogenic food resources. PLoS ONE 12:e0175127. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1371/journal.pone.0175127\u003c/span\u003e\u003cspan address=\"10.1371/journal.pone.0175127\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eStrum SC (2010) The Development of Primate Raiding: Implications for Management and Conservation. Int J Primatol 31:133\u0026ndash;156. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s10764-009-9387-5\u003c/span\u003e\u003cspan address=\"10.1007/s10764-009-9387-5\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eSugden S, Murray M, Edwards MA, St. Clair CC (2021) Inter-population differences in coyote diet and niche width along an urban\u0026ndash;suburban\u0026ndash;rural gradient. J Urban Ecol 7:juab034. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1093/jue/juab034\u003c/span\u003e\u003cspan address=\"10.1093/jue/juab034\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eSugden S, Sanderson D, Ford K et al (2020) An altered microbiome in urban coyotes mediates relationships between anthropogenic diet and poor health. Scientific Reports\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eSun Y, Liu S, Liu Y et al (2021) Effects of the interaction among climate, terrain and human activities on biodiversity on the Qinghai-Tibet Plateau. Sci Total Environ 794:148497. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.scitotenv.2021.148497\u003c/span\u003e\u003cspan address=\"10.1016/j.scitotenv.2021.148497\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eTaberlet P, Bonin A, Zinger L, Coissac E (2018) Reference databases. In: Taberlet P, Bonin A, Zinger L, Coissac E (eds) Environmental DNA: For Biodiversity Research and Monitoring. Oxford University Press, p 0\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eTaki H, Ikeda H, Nagamitsu T et al (2017) Stable nitrogen and carbon isotope ratios in wild native honeybees: the influence of land use and climate. Biodivers Conserv 26:3157\u0026ndash;3166. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s10531-016-1114-x\u003c/span\u003e\u003cspan address=\"10.1007/s10531-016-1114-x\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eTurner BL, Lambin EF, Reenberg A (2007) The emergence of land change science for global environmental change and sustainability. Proc Natl Acad Sci USA 104:20666\u0026ndash;20671. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1073/pnas.0704119104\u003c/span\u003e\u003cspan address=\"10.1073/pnas.0704119104\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eWebber AD (2017) Commensalism. The International Encyclopedia of Primatology. John Wiley and Sons, Ltd, pp 1\u0026ndash;2\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eWist B, Stolter C, Dausmann KH (2022) Sugar addicted in the city: impact of urbanisation on food choice and diet composition of the Eurasian red squirrel (\u003cem\u003eSciurus vulgaris\u003c/em\u003e). J Urban Ecol 8:juac012. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1093/jue/juac012\u003c/span\u003e\u003cspan address=\"10.1093/jue/juac012\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eZhang W, Ricketts TH, Kremen C et al (2007) Ecosystem services and dis-services to agriculture. Ecol Econ 64:253\u0026ndash;260. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.ecolecon.2007.02.024\u003c/span\u003e\u003cspan address=\"10.1016/j.ecolecon.2007.02.024\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":true,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true},"keywords":"Carbon, Nitrogen, Anthropization, Habitat modification, Food availability, Garden Route","lastPublishedDoi":"10.21203/rs.3.rs-3427689/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-3427689/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eAnthropization processes affect wildlife feeding behaviours due to changes in resource availability related to land use and land cover change. To better understand the ecological responses of wildlife towards anthropogenic change, it is essential to evaluate whether human land use, characterized by high human-modified food availability, has an impact on wild animal feeding ecology. The chacma baboon (\u003cem\u003ePapio ursinus\u003c/em\u003e) is interesting to study potential diet changes as it is largely present along a gradient of anthropized areas in Southern Africa. In this study, fecal samples from chacma baboon troops were collected in different land use habitats (peri-urban, agricultural and natural forest habitat) in the Garden Route, South Africa, and their isotopic ratios of carbon (δ\u003csup\u003e13\u003c/sup\u003eC) and nitrogen (δ\u003csup\u003e15\u003c/sup\u003eN) measured. Results showed significant differences between δ\u003csup\u003e15\u003c/sup\u003eN ratios according to land use, indicating significant higher protein intake in areas with human influence in comparison to natural forest habitats. Furthermore, the large majority of the collected samples were contained within the bracket that reflect the C\u003csub\u003e3\u003c/sub\u003e ecosystem of the Garden Route region, with the exception of some samples showing higher δ\u003csup\u003e13\u003c/sup\u003eC ratios associated with the consumption of anthropogenic foods (containing sugar, corn and wheat). The potential protein increase, as well as sources of C\u003csub\u003e4\u003c/sub\u003e plants present in the diets in anthropized areas suggests a visible dietary shift for this species between natural and transformed landscapes. In the future, it will be essential to determine whether and how the consumption of human-modified food could affect the health and associated fitness of chacma baboons.\u003c/p\u003e","manuscriptTitle":"Land use influence on chacma baboon (Papio ursinus) diet in South Africa using stable isotopes","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2023-10-17 15:02:48","doi":"10.21203/rs.3.rs-3427689/v1","editorialEvents":[{"type":"communityComments","content":0}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"dcb1ee1c-a8de-4d76-9439-7897a2e4cf6e","owner":[],"postedDate":"October 17th, 2023","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[],"tags":[],"updatedAt":"2023-12-18T00:44:14+00:00","versionOfRecord":[],"versionCreatedAt":"2023-10-17 15:02:48","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-3427689","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-3427689","identity":"rs-3427689","version":["v1"]},"buildId":"cBFmMYwuxLRRLfASyISRj","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}