Comparison of species diversity, richness, and abundance of dung beetles between wildlife and wildlife-livestock ecosystems of Namibia.

Comparison of species diversity, richness, and abundance of dung beetles between wildlife and wildlife-livestock ecosystems of Namibia. | 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 Comparison of species diversity, richness, and abundance of dung beetles between wildlife and wildlife-livestock ecosystems of Namibia. Mukendwa Hosticks Ndozi, Linnet Gohole This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-4487306/v1 This work is licensed under a CC BY 4.0 License Status: Published Journal Publication published 05 Mar, 2025 Read the published version in International Journal of Tropical Insect Science → Version 1 posted 5 You are reading this latest preprint version Abstract Assessing the species diversity, richness, and abundance of dung beetles in wildlife and wildlife-livestock ecosystems is crucial in understanding the effects of anthropogenic processes on the community structures of dung beetles to improve conservation strategies in Namibia. We tested the hypothesis that the species diversity, richness, and abundance of dung beetles in wildlife ecosystems will be better than in wildlife-livestock ecosystems. Sampling of dung beetles was carried out using baited pitfall traps for a period of 12 months. Linear transects of 1.1 km in length were installed with 12 pitfall traps separated by a distance of 100 m from each other. An independent samples test (P = 0.05) was used to compare the species diversity of dung beetles in two ecosystems. A total of 56,701 individuals were collected from both wildlife and wildlife-livestock ecosystems belonging to 44 species, 25 genera, and 8 tribes. The species diversity of the two ecosystems was similar (H’; t = 1.146, df = 22, P > 0.05). The wildlife ecosystem was more species-rich (n = 43) when compared to the wildlife-livestock ecosystem (n = 35). The species abundance and richness were significantly difference between the two ecosystems (p = < 0.05). A higher Shannon-Wiener Index (H’ = 2.63) was reported in wildlife ecosystems than in wildlife-livestock ecosystems. Different land-use systems have proven to have an impact on species assemblage of dung beetles. We concluded that wildlife ecosystems in Namibia can provide a rich ecological and functional dung beetle community. Dung beetle wildlife ecosystems wildlife-livestock ecosystem Species diversity Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Introduction The expansion of the human population has affected the biodiversity of insect communities in several ways including habitat destruction (Ceballos et al. 2015 ). As humans clear land for agriculture, urbanization, and infrastructure development, countless species lose their natural habitats and are forced to adapt or face extinction (Gardner et al. 2008 ; Barragán et al. 2011 ). Insect communities are also affected by the overexploitation of natural resources arising from increasing human interventions. As a result, natural environments have been transformed by anthropogenic activities leading to the reduction of native forests and woodland savannahs into small fragments of different sizes and various shapes (Braga et al. 2013 ). They are also major causes of climate change and escalate the losses of biological diversity (Quintero and Roslin 2005 ). The term disturbance has been used in ecology to explain the events that occur as a result of natural-related activities or human-induced factors that leads to the alteration of the environment, resulting in affecting the different communities of living organisms in their natural environment (Batisti et al. 2016). Dung beetles are a group of insects belonging to the order Coleoptera of the family Scarabaeidae and the subfamily Scarabaeinae with distinguished ecological importance during their utilization of mammalian dung. They have distinguished ecosystem services, which include the suppression of parasites that oviposit eggs in the dung; they are also responsible for seed dispersal defecated by frugivorous vertebrates (Andresen 2002 ; 2003 ) and recycling of dead materials (Hanski and Cambefort 1991 ; Andresen & Feer 2005 ; Nichols et al. 2008 ). Dung beetles contribute to soil improvement by cycling nutrients and also improving air aeration in the soil (Mittal 1993 ; Wilson 1998) and incorporating dead organic matter into the soil, which tends to improve soil structure and water-holding capacity (Feer and Hingrat 2005 ). Dung beetles have been studied as biological indicators in the assessment of altered ecosystems. High species richness and diversity of dung beetles in different habitats represent a good ecological biological indicator (Rocha et al. 2010 ). Dung beetles are good candidates for monitoring disturbed ecosystems, including forest and agricultural environments. Conversion and fragmentation of forests are quite obvious in some regions of the Continent, natural areas are turned into fragments by anthropogenic activities such as the conversion of forest areas into agriculture and pasture (Favero et al. 2011 ). The conversion of natural habitats into agriculture and pastureland shifts the abundance of mammalian communities and reducing the availability of dung resources, affecting compositions of dung beetle assemblages. Dung beetles primarily depend on the dung excreted by macro-vertebrates, and changes in the communities of mammals and other animals may influence the community structure of dung beetle assemblages (Hernández et al. 2003). Namibia, which is regarded as a semi-arid country in sub-Saharan Africa, located in the southern part of Africa. The country is experiencing different forms of land degradation, including deforestation, soil erosion, sand mining (Klintenberg 2007), and bush encroachment (De Klerk 2004 ). Bush encroachment is a serious type of land degradation that affects different land uses and ecosystems. Encroachment affects the carrying capacity of rangelands, reducing the production of livestock and consequently affecting the population of dung beetles. Dung beetles mainly depend on mammalian dung for oviposition and feeding. When mammals are reduced as a result of land degradation, the dung beetle community structures will also change. Conservation of dung beetles in Namibia is challenged by different anthropogenic activities, with the greatest pressure on conservation arising from urbanization, resource exploitation, or conversion to agroecosystems (Fairbanks et al. 2000 ). Hence, a reduction in the richness of mammalian species in any ecosystem also reduces the abundance and richness of dung beetles (Braga et al. 2013 ; Raine and Slade 2019 ). Namibian ecosystems have proven to be exceptional in dung beetle richness and abundance, with the highest richness collected in protected areas than on farms (Nependa et al. 2021 ), providing evidence of the negative effects of converting natural habitats into agricultural land (livestock farming) on dung beetle diversity. Little is known about the species diversity and community structures of dung beetles in Nkasa Rupara National Park and Dzoti Conservancy due to a lack of scientific research. In the wildlife-livestock ecosystems such as the Dzoti conservation area, people, wildlife and livestock coexist, converting natural habitats into agricultural land, which might shift the population of wildlife as well as change the community structure of dung beetles. Conversion of the landscape for crop production, which includes vegetation clearing and burning, tends to reduce the diversity of insects (Nichols and Gomez, 2014 ). In addition, in wildlife-livestock ecosystems, people are allowed to collect dung beetle larvae for self-consumption and sell the larvae to generate income. The collection of these larvae has not been quantified to determine the best management policy for sustainable ecosystem balance. Therefore, the uncontrolled collection of the dung beetle larvae might have an impact on the population structure of the beetle community, which might lead to the local extinction of some species. Thus, there is a need for comprehensive data recording and analysis on dung beetles to monitor the health of the ecosystem and to inform conservation policy and action. Hence, the results attained on the community structures and diversity of dung beetle in these two ecosystems will constitute a systematic understanding of the assemblages of dung beetle, including unraveling the consequences of anthropogenic activities on dung beetle structure. Therefore, the objective of this study was to determine the species diversity, richness, and abundance of dung beetles in wildlife and wildlife-livestock ecosystems of Namibia. We tested the hypothesis that the species diversity, richness, and abundance of dung beetles do not differ significantly between the wildlife and wildlife-livestock ecosystems. Our predictions are as follows: (i) since wildlife ecosystems are less disturbed, species diversity, richness, and abundance of dung beetles are expected to be higher than in wildlife-livestock ecosystems whereby human population cexist with wildlife, (ii) because of differences in environmental characteristics and habitat structure, this might influence changes in species diversity, richness and abundance of dung beetles in the two study areas. Materials and Methods Study area The study was conducted at Nkasa Rupara National Park (NP), which is a wildlife system, and Dzoti Conservancy (Fig. 1 ), which is a mixture of wildlife and livestock systems. Both study sites are situated in the Zambezi Region, Namibia. Nkasa Rupara NP is located at 18°25’1”S 23°39’3”E, within the strip of the Zambezi Region in the northern-eastern part of Namibia and is bordered by the Kwando River in the western part of the region. The park covers approximately 900 km 2 and is associated with grassland and riparian woodlands (MEFT, 2020)). Both study areas are dominated by mammals (including elephants, buffalo, antelopes, and many others), birds, insects, and fish. Rainfall begins in November and lasts until February, with an annual rainfall of 600–800 mm and temperatures ranging from 24–28°C. Dzoti Conservancy is located at 18°13’59”S 23°46’41”E, covers an area of 287 km 2 with a human population of 1,580 (Denker 2020 ). The conservancy borders Nkasa Rupara National Park and Mudumu National Park. The area is associated with floodplain areas and is channeled by the Linyanti River. The conservancy receives between 550 and 600 mm of rain per year (Denker 2020 ). Sampling design Data collection started in August 2022 and ended in July 2023. Dung beetles were sampled from two study areas; the wildlife ecosystem (Nkasa Rupara National Park) and the wildlife-livestock ecosystem (Dzoti Conservancy). A total of three sites (Fig. 1 ) were selected from each study area with the assistance of the Acting Park Warden, Assistant Rangers, Namibian Police officer, and game guards who had an extensive understanding of the main soil types, vegetation types, and the land use systems used in a conservancy. The selection of the sampling sites was based on the similarities of the habitats in the national park. In the conservancy, the selection of sampling sites was based on three aspects: 1) the most dominant site with livestock, 2) the site with human inhabitants, and 3) the site with wildlife-livestock dominancy. Sites in all the study areas had a minimum of 5 km from each other, this was done to avoid the process of pseudoreplication (Larsen and Forsyth 2005 ; Silva and Hernandez 2015b). Three transects were set at three sampling sites for a period of twelve (12) months. The linear transect of 1.1 km in length was installed with 12 pitfall traps separated at a distance of 100 m from each other. For each linear transect installed in the conservancy, 5 baited pitfall traps contained cattle dung (100 g), while the other 5 baited pitfall traps had buffalo dung, and 2 pitfall traps were used as a control where no bait was placed. In the national park, each linear transect had eight (8) baited pitfall traps with buffalo dung (100 g), while no bait was placed in four (4) of the pitfall traps these acted as the control traps. Dung in the pitfall traps were exposed for 24 hours and replaced immediately after collecting dung beetles, which was repeated until the end of the experiment. Sampling procedures Sampling of dung beetles was carried out using baited pitfall traps, which are regarded to be the most effective technique for capturing dung beetles (Lobo et al. 2001 ). The traps were designed from 1 L plastic containers (11.5 cm diameter and 11.5 cm deep) that were buried with the top edge at ground level and half-filled with lemon detergent water to capture beetles. To avoid the overflow of rainwater, all the traps were protected against rain using plastic plates (20 cm diameter) supported by wooded sticks, inserted approximately 12 cm above the trap. In the national park, traps were baited with fresh buffalo dung ball (100 g) collected from the park, while in the conservancy, each linear transect consisted of five buffalo-baited pitfall traps and five cattle-baited pitfall traps, and two pitfall traps were used as controls. The fresh cattle or buffalo dung was wrapped in polyethylene gauze (Jugovic et al. 2019 ) and attached to the opening of the trap with plain wire. All the collected dung were frozen (-20 ºC) until the day when they were used. The freezing of the dung was done to ensure that there is consistency in attractiveness. Collected dung beetles were preserved in 75% alcohol and transported to the Entomological Laboratory at the National Museum of Namibia in Windhoek. The identification of the specimen was done up to the species level. Identification of dung beetles Collected individuals were transported to the National Museum of Namibia, where sorting, identification and storage was finally done. The dung beetles collected from all the two study areas were identified up to the species level based on the keys and characters listed by (d’Orbigny 1913 : Ferreira 1978 ; Vaz-De-Mello et al. 2011 ; Davis et al. 2020 ). The identification of these dung beetles was mainly based on the morphological characteristics such as coloration, body size, surface sculpture and also comparing to the deposited voucher at the museum. Data analysis All the data prepared for diversity indices and statistical analysis were tested for normality using the normal probability plot in Statgraphics Centurion XVI version 16.1.11. The data that were not normally distributed were transformed using log 10 transformation. To determine the diversity indices, the Paleontological Statistics Software Package for Education and Data Analysis (PAST) version 4.04 was used to generate the diversity indices for both wildlife and wildlife-livestock ecosystems. Three indices were used: the Shannon-Weiner Index (H’), the Simpson’s Diversity Index (D), and the Shannon’s Evenness Index (E) were used to compare the species diversity of dung beetle assemblages in two different ecosystems. These indices were selected because they are popularly used in biological studies. The formula used to calculate the Shannon-Weiner Index was: H’ = - ⅀pi In(pi), where pi is the proportion of i th species in the community and S = Total number of species. The formula used to calculate Simpson’s Diversity Index was: Simpson Diversity Index = (1 – D); where D = Σni(ni-1) / N(N-1), n i represents the number of organisms belongs to species i and N, represent the total number of organisms. The Shannon’s evenness Index (E) was used to measure the evenness using the following formula: E = H’/ In(s); where H’ represents the Shannon-Weiner Index, In represents the natural log of the species richness, and s represents the number of dung beetles recorded in one ecosystem. The independent sample tests were used and the statistical analysis was accepted as significant with a p-value of < 0.05 in all the indices. The species diversity between the two ecosystems was compared using the Sample-size-based rarefaction and extrapolation curve (Margurran & McGill, 2011). The iNEXT (iNterpolation and EXTrapolation) Online software package (Hsieh et al. 2016 ; 2020) was used by considering only one Hill number (q = 0) representing the species diversity with the maximum reference sample size and confidence interval of 95%. The species composition of dung beetle was computed in Microsoft Excel version 2013. The total abundance and number of taxa (richness) of dung beetles from both ecosystems was extracted from PAST software. Richness (%) was calculated using the following formula: Richness (%) = number of species in an ecosystem/Total number of species from the two ecosystems × 100. The average means for all the diversity indices of the species composition were generated from Statgraphics software after running a paired samples T-test. Two-way permutation multivariate analysis of variance (PERMANOVA) was used to examine changes in the community structure of dung beetles between the two ecosystems. A Bray-Curtis similarity matrix (Group Average Link) of dung beetles was calculated from the species data matrix and was subjected to clustering analysis. BioDiversity Professional Software (version 5.0) was used to construct the Rank Abundance Curves (RAC) to assess the dung beetle abundance and species dominance of each ecosystem. Results A total of 56,701 individuals were collected from both Wildlife and Wildlife-livestock ecosystems belonging to 44 species, 25 genera and the following tribes (Table 1 ). The largest number of species (43 spp., 55.1% of the total) and the greatest abundance (46,302 individuals) were collected from the wildlife ecosystem. Fewer species (35 spp., 44.9% of the total) and least abundant (10,399 individuals) were collected from the wildlife-livestock ecosystem. The most abundant species in wildlife ecosystem was Sisyphus goryi (Harold, 1859) (n = 14,790) followed by Onthophagus quadrinotatus (d’Orbigny, 1905) (n = 5,437), Onthophagus lamelliger (Gerstaecker, 1871) (n = 4,907) and Allogymnopleurus splendidus (Bertolini, 1849) (n = 4,772). The Rank-abundance curve (Fig. 2 ) indicates that a high abundance of dung beetle individuals were collected from the wildlife ecosystem, while fewer species were collected from the wildlife-livestock ecosystem. S. goryi in the wildlife ecosystems was the most abundant species, exceeding 10,000 individuals. In the wildlife ecosystem, all the species collected did not exceed 5,000 individuals. The following dung beetle species were not collected in the wildlife-livestock ecosystem: Caccobius nigritulus , Cleptocaccobius viridicollis , Onthophagus ebenicolor , Onthophagus ebenus , Onthophagus fimetarius , Onthophagus flavolimbatus , Onthophagus verticalis , Phalops boschas and Copris bootes and one was exclusively from wildlife ecosystem: Catharsius tricornutus . The results on the functional groups (Fig. 5 ) demonstrated that the tunneler had the highest number species (n = 32, 72.7% of the total species), followed by the roller (n = 10, 22.7% of the total species) and least in the dweller (n = 2, 4.6% of the total species). The independent samples t-test indicated no significant difference between the wildlife ecosystem and wildlife-livestock ecosystem (H’; t = 1.146, df = 22, P > 0.05 for D). A significant difference in abundance (p ≤ 0.05) was observed between the number of individual species collected from the wildlife ecosystem and the wildlife-livestock ecosystem. The highest number of dung beetle species were collected from the wildlife ecosystem, these ecosystems had more species richness when compared to the wildlife-livestock ecosystem. The results further indicated that the wildlife ecosystem registered the highest Shannon-Wiener value (H’ = 2.633), this means elaborates the wildlife ecosystem had higher species diversity when compared to the wildlife-livestock ecosystem (H’ = 2.264). Similar results were also observed in the Simpson’s Diversity Index, with the wildlife ecosystem recording the highest diversity (D = 0.888), and the wildlife-livestock ecosystem recording the least diversity (0.849). The dung beetles in all the ecosystems were evenly and uniformly distributed (wildlife ecosystem, E = 0.844; wildlife-livestock ecosystem, E = 0.865). Table 1 Species composition of dung beetles collected from Wildlife Ecosystem and Wildlife-Livestock ecosystem in Namibia as from August 2022 to July 2023. Tribes Species G Study area Total (%) WE WLE incertae sedis Coprini Gymnopleurini Oniticellini Onitini Onthophagini Scarabaeini Sisyphini Chalconotus convexus (Boheman, 1857) Catharsius aegeus (Génier, 2017) Catharsius tricornutus (DeGeer, 1778) Copris amyntor (Klug, 1855) Copris bootes (Klug, 1855) Copris elphenor (Klug, 1855) Copris puncticollis (Boheman, 1857) Heliocopris japetus (Klug, 1855) Metacatharsius pseudoopacus (Ferreira, 1965) Allogymnopleurus splendidus (Bertolini, 1849 ) Gymnopleurus pumilus (Reiche, 1850) Euoniticellus intermedius (Péringuey, 1901) Latodrepanus laticollis ( Fåhraeus, 1857) Oniticellus formosus (Chevrolat, 1830) Tiniocellus spinipes (Roth, 1851) Onitis alexis (Klug, 1835) Onitis deceptor (Péringuey, 1901) Caccobius nigritulus (Klug, 1855) Cleptocaccobius viridicollis (Fåhraeus, 1857) Digitonthophagus gazella (Fabricius, 1787) Euonthophagus carbonarius (Klug, 1855) Kurtops quadraticeps (Harold, 1867c) Kurtops signatus (Fåhraeus, 1857) Milichus apicalis (Fåhraeus, 1857) Onthophagus aeruginosus (Roth, 1851) Onthophagus ebenicolor (d’Orbigny, 1902) Onthophagus ebenus (Péringuey, 1888) Onthophagus fimetarius (Roth, 1851) Onthophagus flavolimbatus (Klug, 1855) Onthophagus lamelliger (Gerstaecker, 1871) Onthophagus quadrinotatus (d’Orbigny, 1905) Onthophagus suffusus (Klug, 1855) Onthophagus venustulus (Erichson, 1843) Onthophagus verticalis (Fåhraeus, 1857) Onthophagus vinctus (Erichson, 1843) Phalops boschas (Klug, 1855) Phalops flavocinctus (Klug, 1855) Kheper lamarcki (Macleay, 1821) Kheper nigroaeneus (Boheman, 1857) Kheper prodigiosus (Erichson, 1843) Pachylomera femoralis (Kirby, 1828) Scarabaeus zambesianus (Péringuey, 1901) Neosisyphus calcaratus (Klug, 1855) Sisyphus goryi (Harold, 1859) R T T T T T T T T R R T D D T T T T T T T T T T T T T T T T T T T T T T T R R R R R R R 1,841 60 - 108 10 876 371 42 562 4,772 213 351 2,200 47 109 185 2 143 207 150 1,111 723 742 63 83 4 78 434 497 4907 5437 181 3,947 14 80 48 248 205 62 12 122 6 259 14,790 1,667 24 86 29 - 167 46 15 308 622 7 229 245 29 26 150 110 - - 637 279 25 50 41 15 - - - - 934 746 517 870 - 88 - 4 62 33 6 82 12 43 2,195 3,508 (6.2) 84 (0.15) 86 (0.15) 137 (0.24) 10 (0.018) 1,043 (1.84) 417 (0.74) 57 (0.1) 870 (1.53) 5,394 (9.51) 220 ( 0.39) 580 (1.02) 2,445 (4.31) 76 (0.13) 135 (0.24) 335 (0.59) 112 (0.2) 143 (0.25) 207 (0.37) 787 (1.39) 1,390 (2.45) 748 (1.32) 792 (1.4) 104 (0.18) 98 (0.17) 4 (0.01) 78 (0.14) 434 (0.77) 497 (0.88) 5,841 (10.3) 6,183 (10.9) 698 (1.23) 4,817 (8.5) 14 (0.02) 168 (0.3) 48 (0.09) 252 (0.44) 267 (0.47) 95 (0.17) 18 (0.03) 204 (0.36) 18 0.03) 302 (0.53) 16,985 (29.9) Abundance Richness Shannon Index (H’) Simpson Index Evenness (E) 46,302 43 2.633 0.888 0.844 10,399 35 2.264 0.849 0.865 56,701 44 - - - The Bray-Curtis analysis of the group average link (Fig. 3 ) indicates the twenty (20) most abundant species in all the study areas. The six most abundant species were S. goryi , C. convexus , O. venustulus , O. lamelliger , O. quadrinotatus , and A. splendidus as reflected on the Dendrogram (Fig. 3 ). The least abundant species include N. calcaratus , O. alexis , C. puncticollis , O. flavolimbatus , and O. fimetarius . However, three of the least abundant species represented tribe Onthophagini, while the other two least abundant species represented tribes Sisyphini and Onitini. The results on the sample-size-based rarefaction and extrapolation sampling curves (Fig. 4 ) indicated a significant different in the species richness of dung beetle between wildlife and wildlife-livestock ecosystems. The curves further showed that the wildlife ecosystem had the highest number of species, while the wildlife-livestock ecosystem had lower number of species. The two-way PERMANOVA indicated that the ecosystems and functional guilds had significant effects (P 0.05) between the predictors. The results of two-way PERMANOVA confirmed significant differences in the species composition of the two ecosystem types (PERMANOVA, F = 5.73, p = 0.003) and differences in the functional guilds of dung beetles (PERMANOVA, F = 2.24, p = 0.017), while 26.3% were explained by the interaction between the ecosystems and functional guilds. Discussion The results indicated a variation in the assemblages of dung beetle community structures between the two different ecosystems. The high abundance of dung beetle individuals captured in wildlife ecosystems might have been influenced by the absence of anthropogenic activities. Generally, human interventions in protected areas in Namibia are highly prohibited; this has positive benefits for the proper functioning of wildlife ecosystem services. Low human activities in the wildlife ecosystem directly increase the availability of different food resources for dung beetles, driven by a higher population of megaherbivores such as buffaloes and elephants that freely move in and out of the wildlife ecosystem. An increase in the number of megaherbivores is having a positive effect on the abundance of dung beetles. In addition, Nkasa Rupara National Park is one of the protected areas in Namibia with the highest population of buffaloes, these populations increases the availability of dung resources, which also increase the abundance of dung beetle species. On the other hand, the Conservancy was once part of the national park before its proclamation in 2009, therefore, the compositions of dung beetle in the two areas were once the same but due to anthropogenic interference, change in the compositions, abundance, richness, and diversity of dung beetles has occurred. Low dung abundance in wildlife-livestock ecosystems was influenced by the presence of anthropogenic activities such as the exploitation of dung beetle larvae from cattle kraal. During rainy seasons, when dung beetles are abundant, people harvest the larvae from the kraals of cattle, and the harvest is unlimited given the presence of larvae. The collected larvae are used for as food for people, and feeding of poultry, while some of the larvae are sold either fresh or dry in open markets. Marketing of dried and fresh dung beetle larvae in the Northern and North-Eastern parts of Namibia has been observed to be increasing in past years, affecting the life-cycle of the dung beetles. Such anthropogenic activities are responsible of shifting the compositions of dung beetle assemblages in wildlife-livestock ecosystems. Anthropogenic activities remain a major threat to the diversity, richness, and abundance of dung beetle communities in wildlife livestock ecosystems. Conservancies in Namibia are given rights over the utilization of wildlife resources through generating income which is equally distributed to the members, for example, trophy hunting. Trophy hunting of wildlife species shifts the structure of mammals, most of these mammals migrate to the wildlife ecosystems where there are minimum disturbances. The migration of mammalian species will have a negative effect on the community structure of dung beetles, which decreases the species richness, diversity, and abundance of dung beetles in wildlife livestock ecosystems. Changes in the species diversity and richness in wildlife livestock ecosystems might be associated with the practice of shifting cultivation. People who coexist with wildlife species in wildlife livestock ecosystems depend mainly on growing crops and rearing livestock. Crop farming involves the clearing of land through stumping, burning of stumps and fencing crop areas with wooden poles, all these activities increase the migration of wildlife species from these ecosystems, negatively affecting dung beetles. Habitat destruction in wildlife livestock ecosystems leads to heterogeneity of habitats causing a shift in mammalian assemblages affecting dung beetle communities as they depend on mammalian dung for their food and nesting. Moreover, the harvesting of Harpagophytum procumbens (devil’s claw) in wildlife-livestock ecosystems as a way of generating income in communal areas might pose a threat to the movement of wildlife species, alternatively affecting the species composition of dung beetle. To harvest devil’s claw in heavily encroached area, cutting of trees and bushes is a necessity in order to freely access the root system of this plant, this kind of activities disturbs the normal functioning of the ecosystem services. The study hypothesized that wildlife ecosystems have higher species diversity, richness, and abundance of dung beetle communities than wildlife-livestock ecosystems. Hence, a total of 43 species were collected in the wildlife ecosystem, while 35 were collected from the wildlife-livestock ecosystem. The interpolation and extrapolation curves confirmed that more than 43 species may ultimately be found in the wildlife ecosystem. Wildlife ecosystems in Namibia are areas where dung beetles are common and diverse due to the higher number of mammals that supply the dung beetles with sufficient food resources. Species such as C. convexus , A. splendidus , O. lamelliger, O. quadrinotatus, O. venustulus , and S. goryi were abundantly found in wildlife ecosystems. The high number of these species might be influenced by the frequent number of mammals that migrate from the neighborhood countries, and the abundance of buffaloes that roam freely in wildlife ecosystems might increase the number of dung beetles richness due to the abundant dung. However, species such as D. gazella , O. deceptor , and O. suffusus of the tunneler functional group were more abundant in wildlife livestock ecosystems, these species can be used as an indicator of a disturbed ecosystem. The absence of C. tricornutus species of dung beetle in the wildlife ecosystem might be the association of this dung species with the buffalo dung type. The buffalo dung type was the only bait used to capture dung beetles in the wildlife ecosystem, this can explain that C. tricornutus had no association with the buffalo dung type. In wildlife livestock ecosystems where both cattle and buffalo dung types were used to capture dung beetles, C. tricornutus had a higher number of individuals. Overall, the tunneler functional group dominated the species compositions in both ecosystems, this dominace can be induced by their tolerance and nesting behaviors of this group, tunnelers can utilize the dung materials as quickly as possible to minimize competition from other groups. Protected areas of Southern African ecosystems have been reported to have high species diversity, richness, and abundance of dung beetle communities (Davies et al. 1999 ; Koch et al. 2000 ; Tshikae et al. 2008 ; Nependa et al. 2021 ). Similar results were revealed in the current study, wildlife ecosystems conserve the composition of dung beetle community assemblages due to the absence of anthropogenic activities. The abundance of dung beetle communities is reduced as a result of human modification, heterogeneity of habitats, and habitat dispersion (da Silva et al. 2020 ). The other form of anthropogenic activies that alters the species composition dung beetle is land transformation occurring in tropical regions, which remains the major factor contributing to the replacement of natural systems with agricultural lands. Dung beetle density in pastureland may be influenced by the presence of cattle density, the frequency of use of pesticides, and the availability of dung beetle resources (Tonelli et al. 2017 ; Carvalho et al. 2020 a; Correa et al. 2021 ). Dung beetle communities have been used in assessing the impact of anthropogenic disturbance on the community compositions and structures of dung beetle (Nichols et al. 2007 ). Both ecosystems indicated a higher abundance in the functional group of tunnelers, hence, the tunneler beetles possess different tunneling behaviors such as beneath or sometimes alongside the dung pads, these behaviors allow the tunneling groups to use the same resources in various ways, whereby competition of dung is minimized (Silva et al. 2015; da Silva et al. 2020 ). Changes in species composition and functional guilds of dung beetle are associated with changes in land use systems, which are interconnected to the reduction of functional traits in exotic pasture habitats (Maciel et al. 2022). Species such as D. germinatus , T. heydeni , and D. gazella have been reported by Maciel et al. (2022) to be tolerant to ecosystem disturbance, consequently changing the functional composition between the habitats. Similarly, the high abundance of D. gazella in wildlife livestock ecosystems might be influenced by their tolerance of ecosystem disturbance. Conversion of landscapes into agricultural land is one of the major contributors to the loss of biodiversity in dung beetle communities (Numa et al. 2012 ). The species richness of dung beetles declines as the population of mammals is reduced, mammals are the primary food source of dung beetles (Nichols et al. 2009; Braga et al. 2013 ). A shift in the community structure of dung beetles will impose a serious impact on several ecological and ecosystem services that these insects provide to the environment. Habitat modification occurring in different ecosystems has been the major concern associated with the shifts in the community structure of dung beetle (Andresen 2003 ; Larsen et al. 2005 ). Scientific evidence has shown that changes in the population of mammal species will also change the community structure of dung beetles. Therefore, maintaining the populations of mammals in different ecosystems will positively improve the normal functioning of the ecosystem. Conclusion Different land-use systems particularly in the wildlife and wildlife-livestock ecosystems have proven to have an impact on species assemblage of dung beetles. The results of this study concluded that wildlife ecosystems in Namibia can provide a rich ecological and functional dung beetle community. Little is known about the community structure and diversity patterns of dung beetles in wildlife and wildlife-livestock ecosystems in Namibia due to a lack of interest and ongoing research activities. The results attained from this study will constitute the baseline understanding of dung beetle community structure and diversity in the wildlife and wildlife-livestock ecosystems in Namibia, concerning Nkasa Rupara National Park and Dzoti Conservancy. Declarations Acknowledgments We thank the University of Eldoret (UoE) for its support and training. We appreciate the National Commission of Research, Science, and Technology issuing the research permit. We thank the Ministry of Environment, Forestry, and Tourism for allowing this study to be conducted in their protected areas. We thank the National Museums of Kenya (NMK) and the Association of Kenya Entomologists (AKE) for the opportunity provided. Author contributions All the authors contributed to the preparation of the paper. Funding We thank Kreditanstalt für Wiederaufbau (KfW) and the University of Namibia (UNAM) for the funds provided during the data collection and other necessities. The Scholarship Programme is part of the Capacity Building Measures for the Department of Wildlife Management and Ecotourism / University of Namibia (Katima Mulilo Campus) financed under Component 2 of the Project BMZ 2015 67 015 (Expansion of the University of Namibia, Campus Katima Mulilo). Statements and Declarations The authors state and declare that there are no competing interests and this work has not been published elsewhere. 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Tropical savanna conversion to exotic pastures negatively affects taxonomic and functional diversity of dung beetle assemblages, but not dung removal. Journal of Insect Conservation and Diversity 588-599. https://doi.org/10.1111/icad.12656 Magurra AE, McGill BJ (2011). Biological diversity: Frontiers in measurement and assessment. Oxford UP. Oxford. Ministry of Environment, Forestry and Tourism. (2020). Revised National Strategy on Wildlife Protection and Law Enforcement (2021 – 2025). Mittal IC (1993). Natural manuring and soil conditioning by dung beetles. Tropical Ecology150-159. Nependa HUJ, Pryke JS, Roets F (2021). Replacing native mammal assemblages with livestock in African savannahs, impacts dung beetle diversity and reduces body size. Journal of Biological Conservation 109211. https://doi.org/10.1016/j.biocon.2021.109211 Nichols E, Gomez A (2014). Dung beetles and fecal helminth transmission: patterns, mechanisms and questions. Review. Journal of Parasitology 614–623. https://doi.org/10.1017/s0031182013002011 Nichols E, Larsen T, Spector S, Davis AL, Escobar F, Favila M, Vulinec K (2007). Global dung beetle response to tropical forest modification and fragmentation: A quantitative literature review and meta-analysis. Journal of Biological Conservation 1-19. https://doi.org/10.1016/j.biocon.2007.01.023 Nichols E, Spector S, Louzada JNC, Larsen TH, Amezquita S, Favila ME (2008). Ecological functions and ecosystem services provided by Scarabaeinae dung beetles. Journal of Biological Conservation 461–1474. https://doi.org/10.1016/j.biocon.2008.04.011 Numa C, Verdú JR, Rueda C, Galante E (2012). Comparing dung beetle species assemblages between protected areas and adjacent pasturelands in a Mediterranean savanna landscape. Rangel. Journal of Rangeland Ecology and Management 137–143. https://doi.org/10.2111/REM-D-10-00050.1 Quintero I, Roslin T (2005). Rapid recovery of dung beetle communities following habitat fragmentation in Central Amazonia. Journal of Ecology 3303-3311. https://doi.org/10.1890/04-1960 Raine EH, Slade EM (2019). Dung beetle-mammal associations: methods, research trends and future directions. Proc. R. Soc. B Biol. Sci. 286, 2018-2002. https://doi.org/10.1098/rspb.2018.2002 da Rocha JRM, Almeida JR, Lins GA, Durval A (2010). Insects as indicators of environmental changing and pollution: a review of appropriate species and their monitoring. Holos Environment 250-262. https://doi.org/10.14295/holos.v10i2.2996 Tonelli M, Verdú JR, Zunino ME (2017). Effects of grazing intensity and the use of veterinary medical products on dung beetle biodiversity in the sub-mountainous. Landscape of Central Italy. Peer Journal e2780. https://doi.org/10.7717/peerj.2780 Tshikae BP, Davis ALV, Scholtz CH (2008). Trophic associations of a dung beetle assemblage (Scarabaeidae:S carabaeinae) in a woodland Savanna of Botswana. Environmental Entomology 431–441. https://doi.org/10.1603/0046-225x(2008)37[431:taoadb]2.0.co;2 Vaz-De-Mello FZ, Edmonds WD, Ocampo FC, Schoolmeesters P (2011). A multilingual key to the genera and subgenera of the subfamily Scarabaeinae of the New World (Coleoptera: Scarabaeidae). Zootaxa 2854. https://doi.org/10.11646/zootaxa.2854.1.1 Cite Share Download PDF Status: Published Journal Publication published 05 Mar, 2025 Read the published version in International Journal of Tropical Insect Science → Version 1 posted Editorial decision: Major revisions 16 Jun, 2024 Reviewers agreed at journal 30 May, 2024 Reviewers invited by journal 29 May, 2024 Editor assigned by journal 28 May, 2024 First submitted to journal 27 May, 2024 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. 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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-4487306","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":308210696,"identity":"ad4be166-4de3-48a0-82d2-2249aca09cf9","order_by":0,"name":"Mukendwa Hosticks Ndozi","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA6klEQVRIie3PMQuCQBTA8TscGvsqOkmTH6TlHUIt3u5gcVMufoCGqK9gi9BmHJzLQWvQYkuttTn2LoggMGsLur/wfMj94CTEZvvBXDMAHzMJxPh2HNFFqHgSbQj9gJAHoTPzrYP4vfRc18ko8HuhvByXk2E/RdLERSsZZNoToCK2yU7gsqLic0kFzfSh/WL7iAomYsDFBVYoLpA4dPaOjI+GBIaUbKH4qpuAhySi+T4yS8LzTqK1Nwc1Yrk+hQRUyddItm//pUrra5OEgVuFEpcpX+7ktm7idvKavM/y4/PY9JvDNpvN9ifdALDZZm944L0/AAAAAElFTkSuQmCC","orcid":"https://orcid.org/0009-0006-0797-9460","institution":"University of Eldoret","correspondingAuthor":true,"prefix":"","firstName":"Mukendwa","middleName":"Hosticks","lastName":"Ndozi","suffix":""},{"id":308210697,"identity":"d624fd9c-bf9d-4a0a-a23d-7adf74f2a98e","order_by":1,"name":"Linnet Gohole","email":"","orcid":"","institution":"University of Eldoret","correspondingAuthor":false,"prefix":"","firstName":"Linnet","middleName":"","lastName":"Gohole","suffix":""}],"badges":[],"createdAt":"2024-05-28 00:45:44","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-4487306/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-4487306/v1","draftVersion":[],"editorialEvents":[{"content":"https://doi.org/10.1007/s42690-025-01453-3","type":"published","date":"2025-03-05T15:57:10+00:00"}],"editorialNote":"","failedWorkflow":false,"files":[{"id":58307462,"identity":"5aa0ad4e-9d11-49d2-890c-39427c6cc015","added_by":"auto","created_at":"2024-06-13 18:42:46","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":808227,"visible":true,"origin":"","legend":"\u003cp\u003eMap of the study areas showing the different sampling points in Nkasa Rupara National Park (Wildlife ecosystem) and Dzoti Conservancy (Wildlife-livestock ecosystem).\u003c/p\u003e","description":"","filename":"1.png","url":"https://assets-eu.researchsquare.com/files/rs-4487306/v1/56ee20ce6415688d0acc0949.png"},{"id":58308894,"identity":"0c1801ef-1303-4e4d-9a4d-7467bca01f37","added_by":"auto","created_at":"2024-06-13 18:50:46","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":14289,"visible":true,"origin":"","legend":"\u003cp\u003eRank-abundance curves for dung beetle species collected from wildlife and wildlife-livestock ecosystems.\u003c/p\u003e","description":"","filename":"2.png","url":"https://assets-eu.researchsquare.com/files/rs-4487306/v1/59a345cbf29db1a143062413.png"},{"id":58307464,"identity":"ca515cb4-4c52-46ef-916b-27f446af7b6e","added_by":"auto","created_at":"2024-06-13 18:42:46","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":34534,"visible":true,"origin":"","legend":"\u003cp\u003eBray-Curtis group average link cluster analysis for the dung beetle community composition (taxonomic similarity) for the wildlife and wildlife-livestock ecosystems in Namibia.\u003c/p\u003e","description":"","filename":"3.png","url":"https://assets-eu.researchsquare.com/files/rs-4487306/v1/a8633147becdb3439a69d4f9.png"},{"id":58307465,"identity":"c4921616-f3c4-423d-9bad-812058631616","added_by":"auto","created_at":"2024-06-13 18:42:46","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":628483,"visible":true,"origin":"","legend":"\u003cp\u003eSample-size-based rarefaction and extrapolation sampling curves on dung beetle species diversity (q = 0) in wildlife and wildlife-livestock ecosystems. The sample size is represented by the numbers in parentheses and the observed Hill numbers for each reference sample. Shaded areas represent 95% confidence intervals.\u003c/p\u003e","description":"","filename":"4.png","url":"https://assets-eu.researchsquare.com/files/rs-4487306/v1/f49eba908b48a54363f37240.png"},{"id":58307466,"identity":"69a4c2f3-35a6-4086-9617-88d2e893566e","added_by":"auto","created_at":"2024-06-13 18:42:46","extension":"png","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":15769,"visible":true,"origin":"","legend":"\u003cp\u003eTotal functional guild of the compositions of dung beetle assemblages in wildlife and wildlife-livestock ecosystems.\u003c/p\u003e","description":"","filename":"5.png","url":"https://assets-eu.researchsquare.com/files/rs-4487306/v1/db2f40d3c5ec4c2e61f97756.png"},{"id":78183824,"identity":"05436f02-6ce5-4986-a282-e52f788b274e","added_by":"auto","created_at":"2025-03-10 18:18:57","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":2163796,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-4487306/v1/e9f5dcc2-b4f3-4d73-ae8f-0f9870207c15.pdf"}],"financialInterests":"","formattedTitle":"Comparison of species diversity, richness, and abundance of dung beetles between wildlife and wildlife-livestock ecosystems of Namibia.","fulltext":[{"header":"Introduction","content":"\u003cp\u003eThe expansion of the human population has affected the biodiversity of insect communities in several ways including habitat destruction (Ceballos et al. \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). As humans clear land for agriculture, urbanization, and infrastructure development, countless species lose their natural habitats and are forced to adapt or face extinction (Gardner et al. \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Barrag\u0026aacute;n et al. \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). Insect communities are also affected by the overexploitation of natural resources arising from increasing human interventions. As a result, natural environments have been transformed by anthropogenic activities leading to the reduction of native forests and woodland savannahs into small fragments of different sizes and various shapes (Braga et al. \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). They are also major causes of climate change and escalate the losses of biological diversity (Quintero and Roslin \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2005\u003c/span\u003e). The term disturbance has been used in ecology to explain the events that occur as a result of natural-related activities or human-induced factors that leads to the alteration of the environment, resulting in affecting the different communities of living organisms in their natural environment (Batisti et al. 2016).\u003c/p\u003e \u003cp\u003eDung beetles are a group of insects belonging to the order \u003cem\u003eColeoptera\u003c/em\u003e of the family \u003cem\u003eScarabaeidae\u003c/em\u003e and the subfamily \u003cem\u003eScarabaeinae\u003c/em\u003e with distinguished ecological importance during their utilization of mammalian dung. They have distinguished ecosystem services, which include the suppression of parasites that oviposit eggs in the dung; they are also responsible for seed dispersal defecated by frugivorous vertebrates (Andresen \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2002\u003c/span\u003e; \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2003\u003c/span\u003e) and recycling of dead materials (Hanski and Cambefort \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e1991\u003c/span\u003e; Andresen \u0026amp; Feer \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Nichols et al. \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2008\u003c/span\u003e). Dung beetles contribute to soil improvement by cycling nutrients and also improving air aeration in the soil (Mittal \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e1993\u003c/span\u003e; Wilson 1998) and incorporating dead organic matter into the soil, which tends to improve soil structure and water-holding capacity (Feer and Hingrat \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2005\u003c/span\u003e). Dung beetles have been studied as biological indicators in the assessment of altered ecosystems.\u003c/p\u003e \u003cp\u003eHigh species richness and diversity of dung beetles in different habitats represent a good ecological biological indicator (Rocha et al. \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). Dung beetles are good candidates for monitoring disturbed ecosystems, including forest and agricultural environments. Conversion and fragmentation of forests are quite obvious in some regions of the Continent, natural areas are turned into fragments by anthropogenic activities such as the conversion of forest areas into agriculture and pasture (Favero et al. \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). The conversion of natural habitats into agriculture and pastureland shifts the abundance of mammalian communities and reducing the availability of dung resources, affecting compositions of dung beetle assemblages. Dung beetles primarily depend on the dung excreted by macro-vertebrates, and changes in the communities of mammals and other animals may influence the community structure of dung beetle assemblages (Hern\u0026aacute;ndez et al. 2003).\u003c/p\u003e \u003cp\u003eNamibia, which is regarded as a semi-arid country in sub-Saharan Africa, located in the southern part of Africa. The country is experiencing different forms of land degradation, including deforestation, soil erosion, sand mining (Klintenberg 2007), and bush encroachment (De Klerk \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2004\u003c/span\u003e). Bush encroachment is a serious type of land degradation that affects different land uses and ecosystems. Encroachment affects the carrying capacity of rangelands, reducing the production of livestock and consequently affecting the population of dung beetles. Dung beetles mainly depend on mammalian dung for oviposition and feeding. When mammals are reduced as a result of land degradation, the dung beetle community structures will also change. Conservation of dung beetles in Namibia is challenged by different anthropogenic activities, with the greatest pressure on conservation arising from urbanization, resource exploitation, or conversion to agroecosystems (Fairbanks et al. \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2000\u003c/span\u003e). Hence, a reduction in the richness of mammalian species in any ecosystem also reduces the abundance and richness of dung beetles (Braga et al. \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Raine and Slade \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Namibian ecosystems have proven to be exceptional in dung beetle richness and abundance, with the highest richness collected in protected areas than on farms (Nependa et al. \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2021\u003c/span\u003e), providing evidence of the negative effects of converting natural habitats into agricultural land (livestock farming) on dung beetle diversity.\u003c/p\u003e \u003cp\u003eLittle is known about the species diversity and community structures of dung beetles in Nkasa Rupara National Park and Dzoti Conservancy due to a lack of scientific research. In the wildlife-livestock ecosystems such as the Dzoti conservation area, people, wildlife and livestock coexist, converting natural habitats into agricultural land, which might shift the population of wildlife as well as change the community structure of dung beetles. Conversion of the landscape for crop production, which includes vegetation clearing and burning, tends to reduce the diversity of insects (Nichols and Gomez, \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). In addition, in wildlife-livestock ecosystems, people are allowed to collect dung beetle larvae for self-consumption and sell the larvae to generate income. The collection of these larvae has not been quantified to determine the best management policy for sustainable ecosystem balance. Therefore, the uncontrolled collection of the dung beetle larvae might have an impact on the population structure of the beetle community, which might lead to the local extinction of some species. Thus, there is a need for comprehensive data recording and analysis on dung beetles to monitor the health of the ecosystem and to inform conservation policy and action. Hence, the results attained on the community structures and diversity of dung beetle in these two ecosystems will constitute a systematic understanding of the assemblages of dung beetle, including unraveling the consequences of anthropogenic activities on dung beetle structure. Therefore, the objective of this study was to determine the species diversity, richness, and abundance of dung beetles in wildlife and wildlife-livestock ecosystems of Namibia. We tested the hypothesis that the species diversity, richness, and abundance of dung beetles do not differ significantly between the wildlife and wildlife-livestock ecosystems. Our predictions are as follows: (i) since wildlife ecosystems are less disturbed, species diversity, richness, and abundance of dung beetles are expected to be higher than in wildlife-livestock ecosystems whereby human population cexist with wildlife, (ii) because of differences in environmental characteristics and habitat structure, this might influence changes in species diversity, richness and abundance of dung beetles in the two study areas.\u003c/p\u003e"},{"header":"Materials and Methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003eStudy area\u003c/h2\u003e \u003cp\u003eThe study was conducted at Nkasa Rupara National Park (NP), which is a wildlife system, and Dzoti Conservancy (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e), which is a mixture of wildlife and livestock systems. Both study sites are situated in the Zambezi Region, Namibia. Nkasa Rupara NP is located at 18\u0026deg;25\u0026rsquo;1\u0026rdquo;S 23\u0026deg;39\u0026rsquo;3\u0026rdquo;E, within the strip of the Zambezi Region in the northern-eastern part of Namibia and is bordered by the Kwando River in the western part of the region. The park covers approximately 900 km\u003csup\u003e2\u003c/sup\u003e and is associated with grassland and riparian woodlands (MEFT, 2020)). Both study areas are dominated by mammals (including elephants, buffalo, antelopes, and many others), birds, insects, and fish. Rainfall begins in November and lasts until February, with an annual rainfall of 600\u0026ndash;800 mm and temperatures ranging from 24\u0026ndash;28\u0026deg;C. Dzoti Conservancy is located at 18\u0026deg;13\u0026rsquo;59\u0026rdquo;S 23\u0026deg;46\u0026rsquo;41\u0026rdquo;E, covers an area of 287 km\u003csup\u003e2\u003c/sup\u003e with a human population of 1,580 (Denker \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). The conservancy borders Nkasa Rupara National Park and Mudumu National Park. The area is associated with floodplain areas and is channeled by the Linyanti River. The conservancy receives between 550 and 600 mm of rain per year (Denker \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2020\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec4\" class=\"Section2\"\u003e \u003ch2\u003eSampling design\u003c/h2\u003e \u003cp\u003eData collection started in August 2022 and ended in July 2023. Dung beetles were sampled from two study areas; the wildlife ecosystem (Nkasa Rupara National Park) and the wildlife-livestock ecosystem (Dzoti Conservancy). A total of three sites (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e) were selected from each study area with the assistance of the Acting Park Warden, Assistant Rangers, Namibian Police officer, and game guards who had an extensive understanding of the main soil types, vegetation types, and the land use systems used in a conservancy. The selection of the sampling sites was based on the similarities of the habitats in the national park. In the conservancy, the selection of sampling sites was based on three aspects: 1) the most dominant site with livestock, 2) the site with human inhabitants, and 3) the site with wildlife-livestock dominancy. Sites in all the study areas had a minimum of 5 km from each other, this was done to avoid the process of pseudoreplication (Larsen and Forsyth \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Silva and Hernandez 2015b).\u003c/p\u003e \u003cp\u003eThree transects were set at three sampling sites for a period of twelve (12) months. The linear transect of 1.1 km in length was installed with 12 pitfall traps separated at a distance of 100 m from each other. For each linear transect installed in the conservancy, 5 baited pitfall traps contained cattle dung (100 g), while the other 5 baited pitfall traps had buffalo dung, and 2 pitfall traps were used as a control where no bait was placed. In the national park, each linear transect had eight (8) baited pitfall traps with buffalo dung (100 g), while no bait was placed in four (4) of the pitfall traps these acted as the control traps. Dung in the pitfall traps were exposed for 24 hours and replaced immediately after collecting dung beetles, which was repeated until the end of the experiment.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec5\" class=\"Section2\"\u003e \u003ch2\u003eSampling procedures\u003c/h2\u003e \u003cp\u003eSampling of dung beetles was carried out using baited pitfall traps, which are regarded to be the most effective technique for capturing dung beetles (Lobo et al. \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2001\u003c/span\u003e). The traps were designed from 1 L plastic containers (11.5 cm diameter and 11.5 cm deep) that were buried with the top edge at ground level and half-filled with lemon detergent water to capture beetles. To avoid the overflow of rainwater, all the traps were protected against rain using plastic plates (20 cm diameter) supported by wooded sticks, inserted approximately 12 cm above the trap. In the national park, traps were baited with fresh buffalo dung ball (100 g) collected from the park, while in the conservancy, each linear transect consisted of five buffalo-baited pitfall traps and five cattle-baited pitfall traps, and two pitfall traps were used as controls. The fresh cattle or buffalo dung was wrapped in polyethylene gauze (Jugovic et al. \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2019\u003c/span\u003e) and attached to the opening of the trap with plain wire. All the collected dung were frozen (-20 \u0026ordm;C) until the day when they were used. The freezing of the dung was done to ensure that there is consistency in attractiveness. Collected dung beetles were preserved in 75% alcohol and transported to the Entomological Laboratory at the National Museum of Namibia in Windhoek. The identification of the specimen was done up to the species level.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec6\" class=\"Section2\"\u003e \u003ch2\u003eIdentification of dung beetles\u003c/h2\u003e \u003cp\u003eCollected individuals were transported to the National Museum of Namibia, where sorting, identification and storage was finally done. The dung beetles collected from all the two study areas were identified up to the species level based on the keys and characters listed by (d\u0026rsquo;Orbigny \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e1913\u003c/span\u003e: Ferreira \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e1978\u003c/span\u003e; Vaz-De-Mello et al. \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Davis et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). The identification of these dung beetles was mainly based on the morphological characteristics such as coloration, body size, surface sculpture and also comparing to the deposited voucher at the museum.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec7\" class=\"Section2\"\u003e \u003ch2\u003eData analysis\u003c/h2\u003e \u003cp\u003eAll the data prepared for diversity indices and statistical analysis were tested for normality using the normal probability plot in Statgraphics Centurion XVI version 16.1.11. The data that were not normally distributed were transformed using log\u003csub\u003e10\u003c/sub\u003e transformation. To determine the diversity indices, the Paleontological Statistics Software Package for Education and Data Analysis (PAST) version 4.04 was used to generate the diversity indices for both wildlife and wildlife-livestock ecosystems. Three indices were used: the Shannon-Weiner Index (H\u0026rsquo;), the Simpson\u0026rsquo;s Diversity Index (D), and the Shannon\u0026rsquo;s Evenness Index (E) were used to compare the species diversity of dung beetle assemblages in two different ecosystems. These indices were selected because they are popularly used in biological studies.\u003c/p\u003e \u003cp\u003eThe formula used to calculate the Shannon-Weiner Index was:\u003c/p\u003e \u003cp\u003eH\u0026rsquo; = - ⅀pi In(pi), where pi is the proportion of i\u003csup\u003eth\u003c/sup\u003e species in the community and S\u0026thinsp;=\u0026thinsp;Total number of species.\u003c/p\u003e \u003cp\u003eThe formula used to calculate Simpson\u0026rsquo;s Diversity Index was:\u003c/p\u003e \u003cp\u003eSimpson Diversity Index = (1 \u0026ndash; D); where D\u0026thinsp;=\u0026thinsp;Σni(ni-1) / N(N-1), n\u003cem\u003ei\u003c/em\u003e represents the number of organisms belongs to species \u003cem\u003ei\u003c/em\u003e and N, represent the total number of organisms.\u003c/p\u003e \u003cp\u003eThe Shannon\u0026rsquo;s evenness Index (E) was used to measure the evenness using the following formula:\u003c/p\u003e \u003cp\u003eE\u0026thinsp;=\u0026thinsp;H\u0026rsquo;/ In(s); where H\u0026rsquo; represents the Shannon-Weiner Index, In represents the natural log of the species richness, and s represents the number of dung beetles recorded in one ecosystem.\u003c/p\u003e \u003cp\u003eThe independent sample tests were used and the statistical analysis was accepted as significant with a p-value of \u0026lt;\u0026thinsp;0.05 in all the indices.\u003c/p\u003e \u003cp\u003eThe species diversity between the two ecosystems was compared using the Sample-size-based rarefaction and extrapolation curve (Margurran \u0026amp; McGill, 2011). The iNEXT (iNterpolation and EXTrapolation) Online software package (Hsieh et al. \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; 2020) was used by considering only one Hill number (q\u0026thinsp;=\u0026thinsp;0) representing the species diversity with the maximum reference sample size and confidence interval of 95%. The species composition of dung beetle was computed in Microsoft Excel version 2013. The total abundance and number of taxa (richness) of dung beetles from both ecosystems was extracted from PAST software. Richness (%) was calculated using the following formula:\u003cdiv class=\"BlockQuote\"\u003e\u003cp\u003eRichness (%)\u0026thinsp;=\u0026thinsp;number of species in an ecosystem/Total number of species from the two ecosystems \u0026times; 100.\u003c/p\u003e\u003c/div\u003e\u003c/p\u003e \u003cp\u003eThe average means for all the diversity indices of the species composition were generated from Statgraphics software after running a paired samples T-test.\u003c/p\u003e \u003cp\u003eTwo-way permutation multivariate analysis of variance (PERMANOVA) was used to examine changes in the community structure of dung beetles between the two ecosystems. A Bray-Curtis similarity matrix (Group Average Link) of dung beetles was calculated from the species data matrix and was subjected to clustering analysis. BioDiversity Professional Software (version 5.0) was used to construct the Rank Abundance Curves (RAC) to assess the dung beetle abundance and species dominance of each ecosystem.\u003c/p\u003e \u003c/div\u003e"},{"header":"Results","content":"\u003cp\u003eA total of 56,701 individuals were collected from both Wildlife and Wildlife-livestock ecosystems belonging to 44 species, 25 genera and the following tribes (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). The largest number of species (43 spp., 55.1% of the total) and the greatest abundance (46,302 individuals) were collected from the wildlife ecosystem. Fewer species (35 spp., 44.9% of the total) and least abundant (10,399 individuals) were collected from the wildlife-livestock ecosystem. The most abundant species in wildlife ecosystem was \u003cem\u003eSisyphus goryi\u003c/em\u003e (Harold, 1859) (n\u0026thinsp;=\u0026thinsp;14,790) followed by \u003cem\u003eOnthophagus quadrinotatus\u003c/em\u003e (d\u0026rsquo;Orbigny, 1905) (n\u0026thinsp;=\u0026thinsp;5,437), \u003cem\u003eOnthophagus lamelliger\u003c/em\u003e (Gerstaecker, 1871) (n\u0026thinsp;=\u0026thinsp;4,907) and \u003cem\u003eAllogymnopleurus splendidus\u003c/em\u003e (Bertolini, 1849) (n\u0026thinsp;=\u0026thinsp;4,772). The Rank-abundance curve (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e) indicates that a high abundance of dung beetle individuals were collected from the wildlife ecosystem, while fewer species were collected from the wildlife-livestock ecosystem. S. goryi in the wildlife ecosystems was the most abundant species, exceeding 10,000 individuals. In the wildlife ecosystem, all the species collected did not exceed 5,000 individuals.\u003c/p\u003e \u003cp\u003eThe following dung beetle species were not collected in the wildlife-livestock ecosystem: \u003cem\u003eCaccobius nigritulus\u003c/em\u003e, \u003cem\u003eCleptocaccobius viridicollis\u003c/em\u003e, \u003cem\u003eOnthophagus ebenicolor\u003c/em\u003e, \u003cem\u003eOnthophagus ebenus\u003c/em\u003e, \u003cem\u003eOnthophagus fimetarius\u003c/em\u003e, \u003cem\u003eOnthophagus flavolimbatus\u003c/em\u003e, \u003cem\u003eOnthophagus verticalis\u003c/em\u003e, \u003cem\u003ePhalops boschas\u003c/em\u003e and \u003cem\u003eCopris bootes\u003c/em\u003e and one was exclusively from wildlife ecosystem: \u003cem\u003eCatharsius tricornutus\u003c/em\u003e. The results on the functional groups (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003e) demonstrated that the tunneler had the highest number species (n\u0026thinsp;=\u0026thinsp;32, 72.7% of the total species), followed by the roller (n\u0026thinsp;=\u0026thinsp;10, 22.7% of the total species) and least in the dweller (n\u0026thinsp;=\u0026thinsp;2, 4.6% of the total species). The independent samples t-test indicated no significant difference between the wildlife ecosystem and wildlife-livestock ecosystem (H\u0026rsquo;; t\u0026thinsp;=\u0026thinsp;1.146, df\u0026thinsp;=\u0026thinsp;22, P\u0026thinsp;\u0026gt;\u0026thinsp;0.05 for D). A significant difference in abundance (p\u0026thinsp;\u0026le;\u0026thinsp;0.05) was observed between the number of individual species collected from the wildlife ecosystem and the wildlife-livestock ecosystem. The highest number of dung beetle species were collected from the wildlife ecosystem, these ecosystems had more species richness when compared to the wildlife-livestock ecosystem. The results further indicated that the wildlife ecosystem registered the highest Shannon-Wiener value (H\u0026rsquo; = 2.633), this means elaborates the wildlife ecosystem had higher species diversity when compared to the wildlife-livestock ecosystem (H\u0026rsquo; = 2.264). Similar results were also observed in the Simpson\u0026rsquo;s Diversity Index, with the wildlife ecosystem recording the highest diversity (D\u0026thinsp;=\u0026thinsp;0.888), and the wildlife-livestock ecosystem recording the least diversity (0.849). The dung beetles in all the ecosystems were evenly and uniformly distributed (wildlife ecosystem, E\u0026thinsp;=\u0026thinsp;0.844; wildlife-livestock ecosystem, E\u0026thinsp;=\u0026thinsp;0.865).\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\u003eSpecies composition of dung beetles collected from Wildlife Ecosystem and Wildlife-Livestock ecosystem in Namibia as from August 2022 to July 2023.\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"6\"\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=\"char\" char=\".\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eTribes\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eSpecies\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eG\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colspan=\"2\" nameend=\"c5\" namest=\"c4\"\u003e \u003cp\u003eStudy area\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eTotal (%)\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eWE\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eWLE\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eincertae sedis\u003c/em\u003e\u003c/p\u003e \u003cp\u003eCoprini\u003c/p\u003e \u003cp\u003eGymnopleurini\u003c/p\u003e \u003cp\u003eOniticellini\u003c/p\u003e \u003cp\u003eOnitini\u003c/p\u003e \u003cp\u003eOnthophagini\u003c/p\u003e \u003cp\u003eScarabaeini\u003c/p\u003e \u003cp\u003eSisyphini\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eChalconotus convexus\u003c/em\u003e (Boheman, 1857)\u003c/p\u003e \u003cp\u003e\u003cem\u003eCatharsius aegeus\u003c/em\u003e (G\u0026eacute;nier, 2017)\u003c/p\u003e \u003cp\u003e\u003cem\u003eCatharsius tricornutus\u003c/em\u003e (DeGeer, 1778)\u003c/p\u003e \u003cp\u003e\u003cem\u003eCopris amyntor\u003c/em\u003e (Klug, 1855)\u003c/p\u003e \u003cp\u003e\u003cem\u003eCopris bootes\u003c/em\u003e (Klug, 1855)\u003c/p\u003e \u003cp\u003e\u003cem\u003eCopris elphenor\u003c/em\u003e (Klug, 1855)\u003c/p\u003e \u003cp\u003e\u003cem\u003eCopris puncticollis\u003c/em\u003e (Boheman, 1857)\u003c/p\u003e \u003cp\u003e\u003cem\u003eHeliocopris japetus\u003c/em\u003e (Klug, 1855)\u003c/p\u003e \u003cp\u003e\u003cem\u003eMetacatharsius pseudoopacus\u003c/em\u003e (Ferreira, 1965)\u003c/p\u003e \u003cp\u003e\u003cem\u003eAllogymnopleurus splendidus\u003c/em\u003e (Bertolini, 1849\u003cem\u003e)\u003c/em\u003e\u003c/p\u003e \u003cp\u003e\u003cem\u003eGymnopleurus pumilus\u003c/em\u003e (Reiche, 1850)\u003c/p\u003e \u003cp\u003e\u003cem\u003eEuoniticellus intermedius\u003c/em\u003e (P\u0026eacute;ringuey, 1901)\u003c/p\u003e \u003cp\u003e\u003cem\u003eLatodrepanus laticollis\u003c/em\u003e ( F\u0026aring;hraeus, 1857)\u003c/p\u003e \u003cp\u003e\u003cem\u003eOniticellus formosus\u003c/em\u003e (Chevrolat, 1830)\u003c/p\u003e \u003cp\u003e\u003cem\u003eTiniocellus spinipes\u003c/em\u003e (Roth, 1851)\u003c/p\u003e \u003cp\u003e\u003cem\u003eOnitis alexis\u003c/em\u003e (Klug, 1835)\u003c/p\u003e \u003cp\u003e\u003cem\u003eOnitis deceptor\u003c/em\u003e (P\u0026eacute;ringuey, 1901)\u003c/p\u003e \u003cp\u003e\u003cem\u003eCaccobius nigritulus\u003c/em\u003e (Klug, 1855)\u003c/p\u003e \u003cp\u003e\u003cem\u003eCleptocaccobius viridicollis\u003c/em\u003e (F\u0026aring;hraeus, 1857)\u003c/p\u003e \u003cp\u003e\u003cem\u003eDigitonthophagus gazella\u003c/em\u003e (Fabricius, 1787)\u003c/p\u003e \u003cp\u003e\u003cem\u003eEuonthophagus carbonarius\u003c/em\u003e (Klug, 1855)\u003c/p\u003e \u003cp\u003e\u003cem\u003eKurtops quadraticeps\u003c/em\u003e (Harold, 1867c)\u003c/p\u003e \u003cp\u003e\u003cem\u003eKurtops signatus\u003c/em\u003e (F\u0026aring;hraeus, 1857)\u003c/p\u003e \u003cp\u003e\u003cem\u003eMilichus apicalis\u003c/em\u003e (F\u0026aring;hraeus, 1857)\u003c/p\u003e \u003cp\u003e\u003cem\u003eOnthophagus aeruginosus\u003c/em\u003e (Roth, 1851)\u003c/p\u003e \u003cp\u003e\u003cem\u003eOnthophagus ebenicolor\u003c/em\u003e (d\u0026rsquo;Orbigny, 1902)\u003c/p\u003e \u003cp\u003e\u003cem\u003eOnthophagus ebenus\u003c/em\u003e (P\u0026eacute;ringuey, 1888)\u003c/p\u003e \u003cp\u003e\u003cem\u003eOnthophagus fimetarius\u003c/em\u003e (Roth, 1851)\u003c/p\u003e \u003cp\u003e\u003cem\u003eOnthophagus flavolimbatus\u003c/em\u003e (Klug, 1855)\u003c/p\u003e \u003cp\u003e\u003cem\u003eOnthophagus lamelliger\u003c/em\u003e (Gerstaecker, 1871)\u003c/p\u003e \u003cp\u003e\u003cem\u003eOnthophagus quadrinotatus\u003c/em\u003e (d\u0026rsquo;Orbigny, 1905)\u003c/p\u003e \u003cp\u003e\u003cem\u003eOnthophagus suffusus\u003c/em\u003e (Klug, 1855)\u003c/p\u003e \u003cp\u003e\u003cem\u003eOnthophagus venustulus\u003c/em\u003e (Erichson, 1843)\u003c/p\u003e \u003cp\u003e\u003cem\u003eOnthophagus verticalis\u003c/em\u003e (F\u0026aring;hraeus, 1857)\u003c/p\u003e \u003cp\u003e\u003cem\u003eOnthophagus vinctus\u003c/em\u003e (Erichson, 1843)\u003c/p\u003e \u003cp\u003e\u003cem\u003ePhalops boschas\u003c/em\u003e (Klug, 1855)\u003c/p\u003e \u003cp\u003e\u003cem\u003ePhalops flavocinctus\u003c/em\u003e (Klug, 1855)\u003c/p\u003e \u003cp\u003e\u003cem\u003eKheper lamarcki\u003c/em\u003e (Macleay, 1821)\u003c/p\u003e \u003cp\u003e\u003cem\u003eKheper nigroaeneus\u003c/em\u003e (Boheman, 1857)\u003c/p\u003e \u003cp\u003e\u003cem\u003eKheper prodigiosus\u003c/em\u003e (Erichson, 1843)\u003c/p\u003e \u003cp\u003e\u003cem\u003ePachylomera femoralis\u003c/em\u003e (Kirby, 1828)\u003c/p\u003e \u003cp\u003e\u003cem\u003eScarabaeus zambesianus\u003c/em\u003e (P\u0026eacute;ringuey, 1901)\u003c/p\u003e \u003cp\u003e\u003cem\u003eNeosisyphus calcaratus\u003c/em\u003e (Klug, 1855)\u003c/p\u003e \u003cp\u003e\u003cem\u003eSisyphus goryi\u003c/em\u003e (Harold, 1859)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eR\u003c/p\u003e \u003cp\u003eT\u003c/p\u003e \u003cp\u003eT\u003c/p\u003e \u003cp\u003eT\u003c/p\u003e \u003cp\u003eT\u003c/p\u003e \u003cp\u003eT\u003c/p\u003e \u003cp\u003eT\u003c/p\u003e \u003cp\u003eT\u003c/p\u003e \u003cp\u003eT\u003c/p\u003e \u003cp\u003eR\u003c/p\u003e \u003cp\u003eR\u003c/p\u003e \u003cp\u003eT\u003c/p\u003e \u003cp\u003eD\u003c/p\u003e \u003cp\u003eD\u003c/p\u003e \u003cp\u003eT\u003c/p\u003e \u003cp\u003eT\u003c/p\u003e \u003cp\u003eT\u003c/p\u003e \u003cp\u003eT\u003c/p\u003e \u003cp\u003eT\u003c/p\u003e \u003cp\u003eT\u003c/p\u003e \u003cp\u003eT\u003c/p\u003e \u003cp\u003eT\u003c/p\u003e \u003cp\u003eT\u003c/p\u003e \u003cp\u003eT\u003c/p\u003e \u003cp\u003eT\u003c/p\u003e \u003cp\u003eT\u003c/p\u003e \u003cp\u003eT\u003c/p\u003e \u003cp\u003eT\u003c/p\u003e \u003cp\u003eT\u003c/p\u003e \u003cp\u003eT\u003c/p\u003e \u003cp\u003eT\u003c/p\u003e \u003cp\u003eT\u003c/p\u003e \u003cp\u003eT\u003c/p\u003e \u003cp\u003eT\u003c/p\u003e \u003cp\u003eT\u003c/p\u003e \u003cp\u003eT\u003c/p\u003e \u003cp\u003eT\u003c/p\u003e \u003cp\u003eR\u003c/p\u003e \u003cp\u003eR\u003c/p\u003e \u003cp\u003eR\u003c/p\u003e \u003cp\u003eR\u003c/p\u003e \u003cp\u003eR\u003c/p\u003e \u003cp\u003eR\u003c/p\u003e \u003cp\u003eR\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e1,841\u003c/p\u003e \u003cp\u003e60\u003c/p\u003e \u003cp\u003e-\u003c/p\u003e \u003cp\u003e108\u003c/p\u003e \u003cp\u003e10\u003c/p\u003e \u003cp\u003e876\u003c/p\u003e \u003cp\u003e371\u003c/p\u003e \u003cp\u003e42\u003c/p\u003e \u003cp\u003e562\u003c/p\u003e \u003cp\u003e4,772\u003c/p\u003e \u003cp\u003e213\u003c/p\u003e \u003cp\u003e351\u003c/p\u003e \u003cp\u003e2,200\u003c/p\u003e \u003cp\u003e47\u003c/p\u003e \u003cp\u003e109\u003c/p\u003e \u003cp\u003e185\u003c/p\u003e \u003cp\u003e2\u003c/p\u003e \u003cp\u003e143\u003c/p\u003e \u003cp\u003e207\u003c/p\u003e \u003cp\u003e150\u003c/p\u003e \u003cp\u003e1,111\u003c/p\u003e \u003cp\u003e723\u003c/p\u003e \u003cp\u003e742\u003c/p\u003e \u003cp\u003e63\u003c/p\u003e \u003cp\u003e83\u003c/p\u003e \u003cp\u003e4\u003c/p\u003e \u003cp\u003e78\u003c/p\u003e \u003cp\u003e434\u003c/p\u003e \u003cp\u003e497\u003c/p\u003e \u003cp\u003e4907\u003c/p\u003e \u003cp\u003e5437\u003c/p\u003e \u003cp\u003e181\u003c/p\u003e \u003cp\u003e3,947\u003c/p\u003e \u003cp\u003e14\u003c/p\u003e \u003cp\u003e80\u003c/p\u003e \u003cp\u003e48\u003c/p\u003e \u003cp\u003e248\u003c/p\u003e \u003cp\u003e205\u003c/p\u003e \u003cp\u003e62\u003c/p\u003e \u003cp\u003e12\u003c/p\u003e \u003cp\u003e122\u003c/p\u003e \u003cp\u003e6\u003c/p\u003e \u003cp\u003e259\u003c/p\u003e \u003cp\u003e14,790\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1,667\u003c/p\u003e \u003cp\u003e24\u003c/p\u003e \u003cp\u003e86\u003c/p\u003e \u003cp\u003e29\u003c/p\u003e \u003cp\u003e-\u003c/p\u003e \u003cp\u003e167\u003c/p\u003e \u003cp\u003e46\u003c/p\u003e \u003cp\u003e15\u003c/p\u003e \u003cp\u003e308\u003c/p\u003e \u003cp\u003e622\u003c/p\u003e \u003cp\u003e7\u003c/p\u003e \u003cp\u003e229\u003c/p\u003e \u003cp\u003e245\u003c/p\u003e \u003cp\u003e29\u003c/p\u003e \u003cp\u003e26\u003c/p\u003e \u003cp\u003e150\u003c/p\u003e \u003cp\u003e110\u003c/p\u003e \u003cp\u003e-\u003c/p\u003e \u003cp\u003e-\u003c/p\u003e \u003cp\u003e637\u003c/p\u003e \u003cp\u003e279\u003c/p\u003e \u003cp\u003e25\u003c/p\u003e \u003cp\u003e50\u003c/p\u003e \u003cp\u003e41\u003c/p\u003e \u003cp\u003e15\u003c/p\u003e \u003cp\u003e-\u003c/p\u003e \u003cp\u003e-\u003c/p\u003e \u003cp\u003e-\u003c/p\u003e \u003cp\u003e-\u003c/p\u003e \u003cp\u003e934\u003c/p\u003e \u003cp\u003e746\u003c/p\u003e \u003cp\u003e517\u003c/p\u003e \u003cp\u003e870\u003c/p\u003e \u003cp\u003e-\u003c/p\u003e \u003cp\u003e88\u003c/p\u003e \u003cp\u003e-\u003c/p\u003e \u003cp\u003e4\u003c/p\u003e \u003cp\u003e62\u003c/p\u003e \u003cp\u003e33\u003c/p\u003e \u003cp\u003e6\u003c/p\u003e \u003cp\u003e82\u003c/p\u003e \u003cp\u003e12\u003c/p\u003e \u003cp\u003e43\u003c/p\u003e \u003cp\u003e2,195\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e3,508 (6.2)\u003c/p\u003e \u003cp\u003e84 (0.15)\u003c/p\u003e \u003cp\u003e86 (0.15)\u003c/p\u003e \u003cp\u003e137 (0.24)\u003c/p\u003e \u003cp\u003e10 (0.018)\u003c/p\u003e \u003cp\u003e1,043 (1.84)\u003c/p\u003e \u003cp\u003e417 (0.74)\u003c/p\u003e \u003cp\u003e57 (0.1)\u003c/p\u003e \u003cp\u003e870 (1.53)\u003c/p\u003e \u003cp\u003e5,394 (9.51)\u003c/p\u003e \u003cp\u003e220 ( 0.39)\u003c/p\u003e \u003cp\u003e580 (1.02)\u003c/p\u003e \u003cp\u003e2,445 (4.31)\u003c/p\u003e \u003cp\u003e76 (0.13)\u003c/p\u003e \u003cp\u003e135 (0.24)\u003c/p\u003e \u003cp\u003e335 (0.59)\u003c/p\u003e \u003cp\u003e112 (0.2)\u003c/p\u003e \u003cp\u003e143 (0.25)\u003c/p\u003e \u003cp\u003e207 (0.37)\u003c/p\u003e \u003cp\u003e787 (1.39)\u003c/p\u003e \u003cp\u003e1,390 (2.45)\u003c/p\u003e \u003cp\u003e748 (1.32)\u003c/p\u003e \u003cp\u003e792 (1.4)\u003c/p\u003e \u003cp\u003e104 (0.18)\u003c/p\u003e \u003cp\u003e98 (0.17)\u003c/p\u003e \u003cp\u003e4 (0.01)\u003c/p\u003e \u003cp\u003e78 (0.14)\u003c/p\u003e \u003cp\u003e434 (0.77)\u003c/p\u003e \u003cp\u003e497 (0.88)\u003c/p\u003e \u003cp\u003e5,841 (10.3)\u003c/p\u003e \u003cp\u003e6,183 (10.9)\u003c/p\u003e \u003cp\u003e698 (1.23)\u003c/p\u003e \u003cp\u003e4,817 (8.5)\u003c/p\u003e \u003cp\u003e14 (0.02)\u003c/p\u003e \u003cp\u003e168 (0.3)\u003c/p\u003e \u003cp\u003e48 (0.09)\u003c/p\u003e \u003cp\u003e252 (0.44)\u003c/p\u003e \u003cp\u003e267 (0.47)\u003c/p\u003e \u003cp\u003e95 (0.17)\u003c/p\u003e \u003cp\u003e18 (0.03)\u003c/p\u003e \u003cp\u003e204 (0.36)\u003c/p\u003e \u003cp\u003e18 0.03)\u003c/p\u003e \u003cp\u003e302 (0.53)\u003c/p\u003e \u003cp\u003e16,985 (29.9)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eAbundance\u003c/p\u003e \u003cp\u003eRichness\u003c/p\u003e \u003cp\u003eShannon Index (H\u0026rsquo;)\u003c/p\u003e \u003cp\u003eSimpson Index\u003c/p\u003e \u003cp\u003eEvenness (E)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e46,302\u003c/p\u003e \u003cp\u003e43\u003c/p\u003e \u003cp\u003e2.633\u003c/p\u003e \u003cp\u003e0.888\u003c/p\u003e \u003cp\u003e0.844\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e10,399\u003c/p\u003e \u003cp\u003e35\u003c/p\u003e \u003cp\u003e2.264\u003c/p\u003e \u003cp\u003e0.849\u003c/p\u003e \u003cp\u003e0.865\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e56,701\u003c/p\u003e \u003cp\u003e44\u003c/p\u003e \u003cp\u003e-\u003c/p\u003e \u003cp\u003e-\u003c/p\u003e \u003cp\u003e-\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\u003e \u003c/p\u003e \u003cp\u003eThe Bray-Curtis analysis of the group average link (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e) indicates the twenty (20) most abundant species in all the study areas. The six most abundant species were \u003cem\u003eS. goryi\u003c/em\u003e, \u003cem\u003eC. convexus\u003c/em\u003e, \u003cem\u003eO. venustulus\u003c/em\u003e, \u003cem\u003eO. lamelliger\u003c/em\u003e, \u003cem\u003eO. quadrinotatus\u003c/em\u003e, and \u003cem\u003eA. splendidus\u003c/em\u003e as reflected on the Dendrogram (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). The least abundant species include \u003cem\u003eN. calcaratus\u003c/em\u003e, \u003cem\u003eO. alexis\u003c/em\u003e, \u003cem\u003eC. puncticollis\u003c/em\u003e, \u003cem\u003eO. flavolimbatus\u003c/em\u003e, and \u003cem\u003eO. fimetarius\u003c/em\u003e. However, three of the least abundant species represented tribe Onthophagini, while the other two least abundant species represented tribes Sisyphini and Onitini. The results on the sample-size-based rarefaction and extrapolation sampling curves (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e) indicated a significant different in the species richness of dung beetle between wildlife and wildlife-livestock ecosystems. The curves further showed that the wildlife ecosystem had the highest number of species, while the wildlife-livestock ecosystem had lower number of species.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eThe two-way PERMANOVA indicated that the ecosystems and functional guilds had significant effects (P\u0026thinsp;\u0026lt;\u0026thinsp;0.05) on the community composition of dung beetles and there was no interaction (P\u0026thinsp;\u0026gt;\u0026thinsp;0.05) between the predictors. The results of two-way PERMANOVA confirmed significant differences in the species composition of the two ecosystem types (PERMANOVA, F\u0026thinsp;=\u0026thinsp;5.73, p\u0026thinsp;=\u0026thinsp;0.003) and differences in the functional guilds of dung beetles (PERMANOVA, F\u0026thinsp;=\u0026thinsp;2.24, p\u0026thinsp;=\u0026thinsp;0.017), while 26.3% were explained by the interaction between the ecosystems and functional guilds.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003eThe results indicated a variation in the assemblages of dung beetle community structures between the two different ecosystems. The high abundance of dung beetle individuals captured in wildlife ecosystems might have been influenced by the absence of anthropogenic activities. Generally, human interventions in protected areas in Namibia are highly prohibited; this has positive benefits for the proper functioning of wildlife ecosystem services. Low human activities in the wildlife ecosystem directly increase the availability of different food resources for dung beetles, driven by a higher population of megaherbivores such as buffaloes and elephants that freely move in and out of the wildlife ecosystem. An increase in the number of megaherbivores is having a positive effect on the abundance of dung beetles. In addition, Nkasa Rupara National Park is one of the protected areas in Namibia with the highest population of buffaloes, these populations increases the availability of dung resources, which also increase the abundance of dung beetle species. On the other hand, the Conservancy was once part of the national park before its proclamation in 2009, therefore, the compositions of dung beetle in the two areas were once the same but due to anthropogenic interference, change in the compositions, abundance, richness, and diversity of dung beetles has occurred. Low dung abundance in wildlife-livestock ecosystems was influenced by the presence of anthropogenic activities such as the exploitation of dung beetle larvae from cattle kraal. During rainy seasons, when dung beetles are abundant, people harvest the larvae from the kraals of cattle, and the harvest is unlimited given the presence of larvae. The collected larvae are used for as food for people, and feeding of poultry, while some of the larvae are sold either fresh or dry in open markets. Marketing of dried and fresh dung beetle larvae in the Northern and North-Eastern parts of Namibia has been observed to be increasing in past years, affecting the life-cycle of the dung beetles. Such anthropogenic activities are responsible of shifting the compositions of dung beetle assemblages in wildlife-livestock ecosystems.\u003c/p\u003e \u003cp\u003eAnthropogenic activities remain a major threat to the diversity, richness, and abundance of dung beetle communities in wildlife livestock ecosystems. Conservancies in Namibia are given rights over the utilization of wildlife resources through generating income which is equally distributed to the members, for example, trophy hunting. Trophy hunting of wildlife species shifts the structure of mammals, most of these mammals migrate to the wildlife ecosystems where there are minimum disturbances. The migration of mammalian species will have a negative effect on the community structure of dung beetles, which decreases the species richness, diversity, and abundance of dung beetles in wildlife livestock ecosystems. Changes in the species diversity and richness in wildlife livestock ecosystems might be associated with the practice of shifting cultivation. People who coexist with wildlife species in wildlife livestock ecosystems depend mainly on growing crops and rearing livestock. Crop farming involves the clearing of land through stumping, burning of stumps and fencing crop areas with wooden poles, all these activities increase the migration of wildlife species from these ecosystems, negatively affecting dung beetles. Habitat destruction in wildlife livestock ecosystems leads to heterogeneity of habitats causing a shift in mammalian assemblages affecting dung beetle communities as they depend on mammalian dung for their food and nesting. Moreover, the harvesting of \u003cem\u003eHarpagophytum procumbens\u003c/em\u003e (devil\u0026rsquo;s claw) in wildlife-livestock ecosystems as a way of generating income in communal areas might pose a threat to the movement of wildlife species, alternatively affecting the species composition of dung beetle. To harvest devil\u0026rsquo;s claw in heavily encroached area, cutting of trees and bushes is a necessity in order to freely access the root system of this plant, this kind of activities disturbs the normal functioning of the ecosystem services.\u003c/p\u003e \u003cp\u003eThe study hypothesized that wildlife ecosystems have higher species diversity, richness, and abundance of dung beetle communities than wildlife-livestock ecosystems. Hence, a total of 43 species were collected in the wildlife ecosystem, while 35 were collected from the wildlife-livestock ecosystem. The interpolation and extrapolation curves confirmed that more than 43 species may ultimately be found in the wildlife ecosystem. Wildlife ecosystems in Namibia are areas where dung beetles are common and diverse due to the higher number of mammals that supply the dung beetles with sufficient food resources. Species such as \u003cem\u003eC. convexus\u003c/em\u003e, \u003cem\u003eA. splendidus\u003c/em\u003e, \u003cem\u003eO. lamelliger, O. quadrinotatus, O. venustulus\u003c/em\u003e, and \u003cem\u003eS. goryi\u003c/em\u003e were abundantly found in wildlife ecosystems. The high number of these species might be influenced by the frequent number of mammals that migrate from the neighborhood countries, and the abundance of buffaloes that roam freely in wildlife ecosystems might increase the number of dung beetles richness due to the abundant dung. However, species such as \u003cem\u003eD. gazella\u003c/em\u003e, \u003cem\u003eO. deceptor\u003c/em\u003e, and \u003cem\u003eO. suffusus\u003c/em\u003e of the tunneler functional group were more abundant in wildlife livestock ecosystems, these species can be used as an indicator of a disturbed ecosystem. The absence of \u003cem\u003eC. tricornutus\u003c/em\u003e species of dung beetle in the wildlife ecosystem might be the association of this dung species with the buffalo dung type. The buffalo dung type was the only bait used to capture dung beetles in the wildlife ecosystem, this can explain that \u003cem\u003eC. tricornutus\u003c/em\u003e had no association with the buffalo dung type. In wildlife livestock ecosystems where both cattle and buffalo dung types were used to capture dung beetles, \u003cem\u003eC. tricornutus\u003c/em\u003e had a higher number of individuals. Overall, the tunneler functional group dominated the species compositions in both ecosystems, this dominace can be induced by their tolerance and nesting behaviors of this group, tunnelers can utilize the dung materials as quickly as possible to minimize competition from other groups.\u003c/p\u003e \u003cp\u003eProtected areas of Southern African ecosystems have been reported to have high species diversity, richness, and abundance of dung beetle communities (Davies et al. \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e1999\u003c/span\u003e; Koch et al. \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2000\u003c/span\u003e; Tshikae et al. \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Nependa et al. \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Similar results were revealed in the current study, wildlife ecosystems conserve the composition of dung beetle community assemblages due to the absence of anthropogenic activities. The abundance of dung beetle communities is reduced as a result of human modification, heterogeneity of habitats, and habitat dispersion (da Silva et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). The other form of anthropogenic activies that alters the species composition dung beetle is land transformation occurring in tropical regions, which remains the major factor contributing to the replacement of natural systems with agricultural lands. Dung beetle density in pastureland may be influenced by the presence of cattle density, the frequency of use of pesticides, and the availability of dung beetle resources (Tonelli et al. \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Carvalho et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2020\u003c/span\u003ea; Correa et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Dung beetle communities have been used in assessing the impact of anthropogenic disturbance on the community compositions and structures of dung beetle (Nichols et al. \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2007\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eBoth ecosystems indicated a higher abundance in the functional group of tunnelers, hence, the tunneler beetles possess different tunneling behaviors such as beneath or sometimes alongside the dung pads, these behaviors allow the tunneling groups to use the same resources in various ways, whereby competition of dung is minimized (Silva et al. 2015; da Silva et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Changes in species composition and functional guilds of dung beetle are associated with changes in land use systems, which are interconnected to the reduction of functional traits in exotic pasture habitats (Maciel et al. 2022). Species such as \u003cem\u003eD. germinatus\u003c/em\u003e, \u003cem\u003eT. heydeni\u003c/em\u003e, and \u003cem\u003eD. gazella\u003c/em\u003e have been reported by Maciel et al. (2022) to be tolerant to ecosystem disturbance, consequently changing the functional composition between the habitats. Similarly, the high abundance of \u003cem\u003eD. gazella\u003c/em\u003e in wildlife livestock ecosystems might be influenced by their tolerance of ecosystem disturbance. Conversion of landscapes into agricultural land is one of the major contributors to the loss of biodiversity in dung beetle communities (Numa et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). The species richness of dung beetles declines as the population of mammals is reduced, mammals are the primary food source of dung beetles (Nichols et al. 2009; Braga et al. \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). A shift in the community structure of dung beetles will impose a serious impact on several ecological and ecosystem services that these insects provide to the environment. Habitat modification occurring in different ecosystems has been the major concern associated with the shifts in the community structure of dung beetle (Andresen \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2003\u003c/span\u003e; Larsen et al. \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2005\u003c/span\u003e). Scientific evidence has shown that changes in the population of mammal species will also change the community structure of dung beetles. Therefore, maintaining the populations of mammals in different ecosystems will positively improve the normal functioning of the ecosystem.\u003c/p\u003e"},{"header":"Conclusion","content":"\u003cp\u003eDifferent land-use systems particularly in the wildlife and wildlife-livestock ecosystems have proven to have an impact on species assemblage of dung beetles. The results of this study concluded that wildlife ecosystems in Namibia can provide a rich ecological and functional dung beetle community. Little is known about the community structure and diversity patterns of dung beetles in wildlife and wildlife-livestock ecosystems in Namibia due to a lack of interest and ongoing research activities. The results attained from this study will constitute the baseline understanding of dung beetle community structure and diversity in the wildlife and wildlife-livestock ecosystems in Namibia, concerning Nkasa Rupara National Park and Dzoti Conservancy.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eAcknowledgments\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eWe thank the University of Eldoret (UoE) for its support and training. We appreciate the National Commission of Research, Science, and Technology issuing the research permit. We thank the Ministry of Environment, Forestry, and Tourism for allowing this study to be conducted in their protected areas. We thank the National Museums of Kenya (NMK) and the Association of Kenya Entomologists (AKE) for the opportunity provided.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAuthor contributions\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eAll the authors contributed to the preparation of the paper.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFunding\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eWe thank Kreditanstalt f\u0026uuml;r Wiederaufbau (KfW) and the University of Namibia (UNAM) for the funds provided during the data collection and other necessities. The Scholarship Programme is part of the Capacity Building Measures for the Department of Wildlife Management and Ecotourism / University of Namibia (Katima Mulilo Campus) financed under Component 2 of the Project BMZ 2015 67 015 (Expansion of the University of Namibia, Campus Katima Mulilo).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eStatements and Declarations\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe authors state and declare that there are no competing interests and this work has not been published elsewhere.\u0026nbsp;\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eAndresen E, Feer F (2005) The role of dung beetles as secondary seed dispersers and their effect on plant regeneration in tropical rainforests. 331-349. http://dx.doi.org/10.1079/9780851998060.0331\u003c/li\u003e\n\u003cli\u003eAndresen E (2002) Dung beetles in a Central Amazonian rainforest and their ecological role as secondary seed dispersers. 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Trophic associations of a dung beetle assemblage (Scarabaeidae:S carabaeinae) in a woodland Savanna of Botswana. Environmental Entomology 431\u0026ndash;441. https://doi.org/10.1603/0046-225x(2008)37[431:taoadb]2.0.co;2 \u003c/li\u003e\n\u003cli\u003eVaz-De-Mello FZ, Edmonds WD, Ocampo FC, Schoolmeesters P (2011). A multilingual key to the genera and subgenera of the subfamily Scarabaeinae of the New World (Coleoptera: Scarabaeidae). Zootaxa 2854. https://doi.org/10.11646/zootaxa.2854.1.1 \u003c/li\u003e\n\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":true,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":true,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"international-journal-of-tropical-insect-science","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"jtis","sideBox":"Learn more about [International Journal of Tropical Insect Science](http://link.springer.com/journal/42690)","snPcode":"42690","submissionUrl":"https://www.editorialmanager.com/jtis/default2.aspx","title":"International Journal of Tropical Insect Science","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"Dung beetle, wildlife ecosystems, wildlife-livestock ecosystem, Species diversity","lastPublishedDoi":"10.21203/rs.3.rs-4487306/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-4487306/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eAssessing the species diversity, richness, and abundance of dung beetles in wildlife and wildlife-livestock ecosystems is crucial in understanding the effects of anthropogenic processes on the community structures of dung beetles to improve conservation strategies in Namibia. We tested the hypothesis that the species diversity, richness, and abundance of dung beetles in wildlife ecosystems will be better than in wildlife-livestock ecosystems. Sampling of dung beetles was carried out using baited pitfall traps for a period of 12 months. Linear transects of 1.1 km in length were installed with 12 pitfall traps separated by a distance of 100 m from each other. An independent samples test (P\u0026thinsp;=\u0026thinsp;0.05) was used to compare the species diversity of dung beetles in two ecosystems. A total of 56,701 individuals were collected from both wildlife and wildlife-livestock ecosystems belonging to 44 species, 25 genera, and 8 tribes. The species diversity of the two ecosystems was similar (H\u0026rsquo;; t\u0026thinsp;=\u0026thinsp;1.146, df\u0026thinsp;=\u0026thinsp;22, P\u0026thinsp;\u0026gt;\u0026thinsp;0.05). The wildlife ecosystem was more species-rich (n\u0026thinsp;=\u0026thinsp;43) when compared to the wildlife-livestock ecosystem (n\u0026thinsp;=\u0026thinsp;35). The species abundance and richness were significantly difference between the two ecosystems (p\u0026thinsp;=\u0026thinsp;\u0026lt;\u0026thinsp;0.05). A higher Shannon-Wiener Index (H\u0026rsquo; = 2.63) was reported in wildlife ecosystems than in wildlife-livestock ecosystems. Different land-use systems have proven to have an impact on species assemblage of dung beetles. We concluded that wildlife ecosystems in Namibia can provide a rich ecological and functional dung beetle community.\u003c/p\u003e","manuscriptTitle":"Comparison of species diversity, richness, and abundance of dung beetles between wildlife and wildlife-livestock ecosystems of Namibia.","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2024-06-13 18:42:41","doi":"10.21203/rs.3.rs-4487306/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Major revisions","date":"2024-06-17T03:21:47+00:00","index":"","fulltext":""},{"type":"reviewerAgreed","content":"","date":"2024-05-30T05:23:11+00:00","index":0,"fulltext":""},{"type":"reviewersInvited","content":"","date":"2024-05-29T12:33:30+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2024-05-28T12:51:40+00:00","index":"","fulltext":""},{"type":"submitted","content":"International Journal of Tropical Insect Science","date":"2024-05-27T20:44:50+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"international-journal-of-tropical-insect-science","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"jtis","sideBox":"Learn more about [International Journal of Tropical Insect Science](http://link.springer.com/journal/42690)","snPcode":"42690","submissionUrl":"https://www.editorialmanager.com/jtis/default2.aspx","title":"International Journal of Tropical Insect Science","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"61a74375-a773-40cd-94b2-79349358b4e5","owner":[],"postedDate":"June 13th, 2024","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"published-in-journal","subjectAreas":[],"tags":[],"updatedAt":"2025-03-10T17:46:40+00:00","versionOfRecord":{"articleIdentity":"rs-4487306","link":"https://doi.org/10.1007/s42690-025-01453-3","journal":{"identity":"international-journal-of-tropical-insect-science","isVorOnly":false,"title":"International Journal of Tropical Insect Science"},"publishedOn":"2025-03-05 15:57:10","publishedOnDateReadable":"March 5th, 2025"},"versionCreatedAt":"2024-06-13 18:42:41","video":"","vorDoi":"10.1007/s42690-025-01453-3","vorDoiUrl":"https://doi.org/10.1007/s42690-025-01453-3","workflowStages":[]},"version":"v1","identity":"rs-4487306","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-4487306","identity":"rs-4487306","version":["v1"]},"buildId":"qtupq5eGEP_6zYnWcrvyt","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}