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Identifying habitat modification by Chinese pangolin in subtropical forests of southern China

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Abstract

The excavation of Chinese pangolin ( Manis pentadactyla ) is expected to alter habitat heterogeneity and thus affect the functioning and structure of forest ecosystems. In this study, the bioturbation of Chinese pangolin on forest soils in three regions (Heping, Tianjingshan, and Wuqinzhang) across Guangdong province was quantified. Overall, a mean of 2.66 m ³ ·ha ⁻¹ and 83.1 m ² ·ha ⁻¹ of burrows and bare mounds, respectively, was excavated by Chinese pangolin; the disturbed soils had significantly lower water content and P, C, available N concentrations, but higher bulk density, pH, and microbial abundance than those undisturbed soils. The unevenness of habitat heterogeneity improvement was mainly ascribed to the stronger soil disturbance caused in resting burrows by pangolins. Patterns of altering habitat heterogeneity were site‐specific, with high‐intensity soil disturbance occurring most in shrubs, meadows, steep habitats at high elevations, and mountain tops in Heping, while in broad‐leaved, coniferous and mixed coniferous and broad‐leaved forests away from human settlements in Tianjingshan and upper mountains at high elevations far away from roads and human settlements in Wuqinzhang. Road networks are the main interference for the burrow distribution in Heping and Wuqinzhang and should be programmed.

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... The Chinese pangolin (Manis pentadactyla Linnaeus, 1758) is one of the four extant pangolin species in Asia, distributed across South Asia, Southeast Asia, and East Asia, including Hainan Island and Taiwan (Corlett, 2007;Wu et al., 2020). As an ecological engineer, the Chinese pangolin significantly influences habitat heterogeneity, impacting the structure and function of forest ecosystems (Sun et al., 2024a). Over recent decades, its population has declined drastically due to poaching for traditional medicines, dietary supplements, leather products, and decorative items (Challender et al., 2014;Shirley et al., 2023). ...
... As obligate myrmecophagous mammals, Chinese pangolins almost exclusively prey on ants and termites, often excavating burrows to access soil-inhabiting prey (Sun et al., 2020a;Chao et al., 2020). Consequently, pangolin burrows are typically classified as either resting (or resident) or feeding burrows (Sun et al., 2021b(Sun et al., , 2024aSabin et al., 2024). However, pangolins may also excavate burrows for multiple reasons. ...
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The Chinese pangolin Manis pentadactyla, a burrowing mammal, relies on burrows for foraging, shelter, and rearing its young. Pangolin burrows are typically classified as either resting (or resident) or feeding burrows. Resting burrows, in particular, are crucial for the pangolin’s survival, offering shelter, protection, and stability for the pangolin, especially during colder months and breeding seasons. Consequently, understanding the characteristics and usage patterns of resting burrows is critical for effective in-situ and ex-situ conservation efforts. This study assessed the density of resting burrows using sign surveys along line transects in a fragmented agricultural landscape. Additionally, radio telemetry was employed to specifically locate and characterize the resting burrows of seven Chinese pangolins (1 male and 6 females), as well as to analyze their usage patterns. The results revealed a burrow density of 110.8 per ha, with no significant variation across different environments. However, Chinese pangolins preferred resting burrows in areas with minimal human disturbance. Over tracking periods ranging from 52 to 327 days, female pangolins used between 12 and 32 resting burrows, while a single male pangolin used 48 resting burrows. Pangolins occupied individual burrows for up to 10 days during colder months and breeding seasons. Notably, resting burrows were frequently shared among individuals. Activity patterns showed that pangolins typically exited their burrows between 6:00 PM and 12:00 AM and entered their burrows between 10:00 PM and 4:00 AM. To conclude the findings:1) burrow count, while useful for identifying pangolin presence, is not a suitable quantitative indicator of population density. 2) Most resting burrows had compacted soil at the entrance, which could be mistaken for old or inactive burrows in other studies. 3) Resting burrows function as permanent structures that are reused across seasons instead of seasonal adaptations. 4) Preserving low-disturbance habitats will be critical for ensuring sufficient resting burrow sites, thereby supporting the long-term viability of pangolin populations. These results provide valuable insights into the usage patterns of resting burrows, thus enriching the understanding of pangolin behavior, ecological requirements and conservation strategies for this endangered species.
... The soil heaps around these burrows were smaller, consisting of fine and loose soil resulting from superficial digging activity, whereas the resting burrows featured more circular entrances and larger soil heaps that contained a mix of fine and coarse particles, reflecting the varied soil layers excavated during construction. The soil around resting burrows also appeared more compact due to repeated use and the need for structural stability (Sun et al. 2025). The opening diameter of each burrow was measured using a measuring tape, whereas the GPS location of the burrows was recorded using a handheld Garmin GPS Map 60 CSx. ...
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The information for assessing Red List of China’s Vertebrates was mainly collected from specimens, literature and experts. Specimen data were collected from museums located at the Chinese Academy of Sciences and other universities as well as local natural history museums. Based on the specimen data collected, we analyzed the data using GIS to gain information about the extent of occurrence and area of occupancy of each vertebrate species evaluated, which provided fundamental data for the assessment. In addition to specimen data, literature review provided the main information for this assessment, in reference to new species, species distribution, ecology, conservation, and resource utilization. Furthermore, expert consultation was another important assessment method. Experts are familiar with latest status of the species they study. By referring to literature and field experience, the experts provided sufficient and updated information of relevant species, including the taxonomic definition, population size and trends, habitat conditions, threats, utilization, and conservation information. The deadline for information collection was March 31, 2015. The mammalian inventory system in the assessment followed the system of China’s Mammal Diversity (Jiang et al, 2015), including 673 species of mammals, among which 150 endemic species were assessed. The inventory systems of birds, reptiles and amphilbians followed that in Zheng (2011), Cai et al (2015) and Fei et al (2012), respectively. The assessment of the Red List of China’s Vertebrates included the following steps: information collection, database establishment, formation of an advisory committee, core experts group, and working group, and establishment of a pool of corresponding experts. Members of the advisory committee consisted of Academician Yiyu Chen (from the Endangered Species Scientific Commission of the People’s Republic of China), Academician Yaping Zhang (from the Chinese Academy of Sciences), Academician Jianming Jin (from the Ministry of Environmental Protection), Academician Jianzhang Ma (from Northeast Forestry University) and Academician Guangmei Zheng (from Beijing Normal University). The advisory commission oversaw assessment progress. Five working groups, one core expert group and corresponding expert teams of mammals, birds, reptiles, amphibians and continental fishes were formed. The core expert group assisted each working group in determining the evaluation method, criteria, and verified species data. The working groups were responsible for implementing the assessment plan as specified, including data collection and organization, initial assessment of red listed species, contacting experts by correspondence, and compiling evaluation results. The core experts discussed and reviewed the species’ living status. The working groups gathered advisory experts across the nation and formed an advisory expert pool. The advisory experts participated in the Red List evaluation by correspondence or at review meetings. After the review, we finished assessing each species using unified standards, including scientific name, Chinese name, family name, Chinese family name, status assessed, and IUCN criteria. On May 6, 2015, the Red List of China’s Vertebrates passed an evaluation jointly held by the Ministry of Environmental Protection of the People’s Republic of China and the Chinese Academy of Sciences. The Red List was officially released in the form of the 32nd Announcement of the Ministry of Environmental Protection of the People’s Republic of China and the Chinese Academy of Sciences on May 23, 2015. Detailed information and analyses of the results of the Red List can be found in Jiang ZG et al (2016), Zhang et al (2016), Cai et al (2016), Jiang JP et al (2016) and Cao et al (2016). This paper contains the Red List of China’s Vertebrates including that of mammals, birds, reptiles, amphibians and continental fishes. The references cited in this data paper can be found at the website www.biodiversity-science.net/ fileup/PDF/2016076-1.pdf.
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Both land use intensification and abandonment within grasslands lead to a homogenisation of vegetation structure. Therefore, specially structured microsites such as vegetation gaps with bare ground play an important role for species conservation within grasslands. Vegetation gaps are crucial for the establishment of low-competitive plant species and offer special microclimatic conditions essential for the development of the immature stages of many invertebrate species. The influence of small-scale soil disturbance in the form of mounds created by ecosystem engineers such as ants or moles on biodiversity is therefore of special scientific concern. The effects of mound-building species on plant species diversity have been extensively studied. However, knowledge on the significance of these species for the conservation of other animals is rare. In this study we analyse the importance of mounds created by the European mole (Talpa europaea) as an oviposition habitat for the small copper (Lycaena phlaeas) within Central European mesotrophic grasslands. Our study showed that host plants occurring at molehills were preferred for oviposition. Oviposition sites were characterised by an open vegetation structure with a high proportion of bare ground (with a mean coverage of about 50 %), a low cover of herbs and low-growing vegetation (mean height: 4.5 cm). Our study clearly illustrates the importance of small-scale soil disturbance for immature stages of L. phlaeas and the conservation of this species within mesotrophic grasslands. Mound-building ecosystem engineers, such as T. europaea, act as important substitutes for missing dynamics within mesotrophic grasslands by diversifying vegetation structure and creating small patches of bare soil.
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Mammals that forage for food by biopedturbation can alter the biotic and abiotic characteristics of their habitat, influencing ecosystem structure and function. Bandicoots, bilbies, bettongs and potoroos are the primary digging marsupials in Australia, although most of these species have declined throughout their range. This study used a snapshot approach to estimate the soil turnover capacity of the southern brown bandicoot (Isoodon obesulus, Shaw 1797), a persisting digging Australian marsupial, at Yalgorup National Park, Western Australia. The number of southern brown bandicoots was estimated using mark–recapture techniques. To provide an index of digging activity per animal, we quantified the number of new foraging pits and bandicoot nose pokes across 18 plots within the same area. The amount of soil displaced and physical structure of foraging pits were examined from moulds of 47 fresh foraging pits. We estimated that an individual southern brown bandicoot could create ~45 foraging pits per day, displacing ~10.74 kg of soil, which extrapolates to ~3.9 tonnes of soil each year. The digging activities of the southern brown bandicoots are likely to be a critical component of soil ecosystem processes.
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We censused and measured armadillo burrows in ten 10 m x 40 m plots in each of four habitat types at a study site in northern Florida and one in the Atlantic coastal rainforest of Brazil. The nine-banded armadillo (Dasypus novemcinctus) was the only species of armadillo found in Florida, but several additional species were present in Brazil. Burrows were more numerous but smaller in Brazil than in the U. S., probably due to the inclusion of burrows dug by the smaller congener D. septemcinctus. In Brazil, burrows were larger and more numerous in swamp and forest habitats than in grassland or disturbed areas, suggesting that D. novemcinctus is found primarily in forests and swamps while D. septemcinctus is located in the other areas. This was supported by data from sightings of live animals. In Florida, burrows were more numerous in hardwood hammocks than in wetlands, fields or upland pine areas, but burrow dimensions did not vary across habitat types. In Florida, armadillos were seen more frequently than expected in hammocks and wetlands and less frequently than expected in fields and upland pine areas. There were also age (juvenile versus adult), sex, and yearly differences in habitat use in Florida. Biomass, abundance, and species diversity of terrestrial invertebrates did not vary significantly between habitat types in Florida, suggesting that habitat associations of armadillos were not influenced by prey availability.
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Digging mammals are often considered ecosystem engineers, as they affect important properties of soils and in turn nutrient exchange, vegetation dynamics and habitat quality. Returning such species, and their functions, to areas from where they have been extirpated could help restore degraded landscapes and is increasingly being trialled as a conservation tool. Studies examining the effects of digging mammals have largely been from arid and semi‐arid environments, with little known about their impacts and importance in mesic systems. To address this knowledge gap, we investigated the ecological role of a recently introduced population of eastern barred bandicoots (Perameles gunnii) on Churchill Island, Victoria, south‐eastern Australia, from which all digging mammals have been lost. We quantified the annual rate of soil turnover by estimating the number of foraging pits bandicoots created in 100‐m2 plots over a 24‐h period. Foraging pit counts could not be completed in each season, and the overall turnover estimate assumes that autumn/winter months represent turnover rates for the entire year; however, this is likely to fluctuate between seasons. Ten fresh and ten old pits were compared to paired undug control sites to quantify the effect soil disturbance had on soil hydrophobicity, moisture content and soil strength. Plots contained between zero and 64 new foraging pits each day. We estimated that an individual eastern barred bandicoot digs ~487 (95% CI = 416–526) small foraging pits per night, displacing ~13.15 kg (95% CI = 11.2–14.2 kg) of soil, equating to ~400 kg (95% CI = 341–431 kg) of soil in a winter month. Foraging pits were associated with decreased soil compaction and increased soil moisture along the foraging pit profile. Eastern barred bandicoots likely play an important role in ecosystems through their effects on soil, which adds to an increasing body of knowledge suggesting restoration of ecosystems, via the return of ecosystem engineers and their functions, holds much promise for conserving biodiversity and ecological function.
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Ecosystem engineers have been widely studied for terrestrial systems, but global trends in research encompassing the range of taxa and functions have not previously been synthesised. We reviewed contemporary understanding of engineer fauna in terrestrial habitats and assessed the methods used to document patterns and processes, asking: (a) which species act as ecosystem engineers and with whom do they interact? (b) What are the impacts of ecosystem engineers in terrestrial habitats and how are they distributed? (c) What are the primary methods used to examine engineer effects and how have these developed over time? We considered the strengths, weaknesses and gaps in knowledge related to each of these questions and suggested a conceptual framework to delineate “significant impacts” of engineering interactions for all terrestrial animals. We collected peer‐reviewed publications examining ecosystem engineer impacts and created a database of engineer species to assess experimental approaches and any additional covariates that influenced the magnitude of engineer impacts. One hundred and twenty‐two species from 28 orders were identified as ecosystem engineers, performing five ecological functions. Burrowing mammals were the most researched group (27%). Half of all studies occurred in dry/arid habitats. Mensurative studies comparing sites with and without engineers (80%) were more common than manipulative studies (20%). These provided a broad framework for predicting engineer impacts upon abundance and species diversity. However, the roles of confounding factors, processes driving these patterns and the consequences of experimentally adjusting variables, such as engineer density, have been neglected. True spatial and temporal replication has also been limited, particularly for emerging studies of engineer reintroductions. Climate change and habitat modification will challenge the roles that engineers play in regulating ecosystems, and these will become important avenues for future research. We recommend future studies include simulation of engineer effects and experimental manipulation of engineer densities to determine the potential for ecological cascades through trophic and engineering pathways due to functional decline. We also recommend improving knowledge of long‐term engineering effects and replication of engineer reintroductions across landscapes to better understand how large‐scale ecological gradients alter the magnitude of engineering impacts.
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Understorey vegetation comprises the majority of species diversity and contributes greatly to ecosystem functioning in boreal forests. Although patterns of understorey abundance, species diversity and composition associated with forest stand development are well researched, mechanisms driving these patterns remain largely speculative. We sampled fire‐origin stands of varying stand ages and overstorey compositions on mesic sites of the boreal forest of Canada and used structural equation modelling (SEM) to link time since fire (stand age), light availability and heterogeneity, substrate heterogeneity and soil nitrogen to understorey vegetation cover and species diversity. The most parsimonious model for total understorey cover showed a positive direct effect of stand age ( r = .43) and an indirect effect via mean light level (0.18) and shrub cover (−0.11), with a positive total effect (0.50); the per cent broadleaf canopy had a direct negative effect (−0.22) and an indirect effect via shrub cover (−0.11). The model for total understorey species richness showed an indirect effect of stand age via mean light (0.24), light heterogeneity (0.10) and substrate heterogeneity (0.07), with a positive total effect (0.52); per cent broadleaf canopy had an indirect effect via light heterogeneity (0.09), and substrate heterogeneity (−0.10). Soil nitrogen did not significantly influence either understorey cover or species richness. The models for vascular plants followed similar trends to those for total understorey cover and species richness; however, there was an opposite indirect effect of light heterogeneity for both cover and species richness of non‐vascular plants. Shrub cover had positive direct and negative direct and indirect effects on both vascular and non‐vascular cover and species richness. Synthesis . Our findings indicate that understorey cover and species diversity are driven by time since disturbance, light availability as influenced by overstorey and shrub layers, but with important additional effects mediated by light and substrate heterogeneity. Non‐vascular understorey vegetation is more strongly determined by time since disturbance than vascular vegetation, and negatively affected by broadleaf tree abundance. The overall results highlight the importance of colonization, light availability and heterogeneity, substrate specialization and growth dynamics in determining successional patterns of boreal forest understorey vegetation.
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To understand the impacts of reforestation on woody species composition, species diversity and community structure, seven plantation forests in dry-hot valley of the Jinsha River in Southwest China were investigated, with adjacent wastelands, natural shrub grassland and a natural forest as references. Species importance value, species richness, species heterogeneity and Sorenson similarity index between plantations and the natural forest were analyzed. Results indicated that compared to wastelands and natural shrub grassland, reforestation improved species diversity and community structure, and more forest woody species found suitable habitats in plantations. Species diversity in understory of plantations and Sorenson similarity index were significantly negatively correlated with stem density in mature plantations (26-31 years old). Higher species diversity and Sorenson similarity index existed in mature sparse plantations due to lower stem density and more tree species planted initially. In contrast, reference natural forest, with species heterogeneity of 2.28 for shrub layer, showed the highest species diversity. It would take a long time for species composition and diversity to recover through reforestation in a dry-hot valley. Therefore, it was essential to protect remnant natural forests strictly and reforest with suitable management such as lower stem density and increasing genetic diversity of trees planted.
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Interactions between organisms are a major determinant of the distribution and abundance of species. Ecology textbooks (e.g., Ricklefs 1984, Krebs 1985, Begon et al. 1990) summarise these important interactions as intra- and interspecific competition for abiotic and biotic resources, predation, parasitism and mutualism. Conspicuously lacking from the list of key processes in most text books is the role that many organisms play in the creation, modification and maintenance of habitats. These activities do not involve direct trophic interactions between species, but they are nevertheless important and common. The ecological literature is rich in examples of habitat modification by organisms, some of which have been extensively studied (e.g. Thayer 1979, Naiman et al. 1988).
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Burrowing animals fundamentally influence the character of soils, ecosystem function, erosion, and soil management for food production and hazardous wastes. Perhaps the most important indicator of their potential impacts is the average void space created for burrow construction and maintenance per unit area per unit time. We integrated published estimates of burrow volume (m3) and excavation rates (m3 ha-1 year-1) of pocket gophers (Rodentia: Geomyidae) to assess their reliability in assessment of impacts and whether these variables are scale-dependent. Estimates were difficult to interpret and compare because research goals, assumptions, and methods varied considerably among studies, as did site conditions and reporting of results. Volume of burrows made by various species of pocket gophers averaged 0.68 m3, and averages differed significantly between species. Rate of excavation averaged 18 m3 ha-1 year-1 for Geomyidae, and it declined with increasing spatial extent of study area used to make the estimates. Counts of mounds produced the largest estimates of burrow volume and excavation rate, and injection of polyurethane foam into burrows produced low estimates. High variation in estimated rates of burrow excavation indicates the need for further research to characterize magnitude and types of impacts gophers have on natural and managed ecosystems.
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Dimensions of the burrow systems of four small mammals common to southeastern Idaho (Spermophilus townsendii, Dipodomys ordii, Microtus montanus and Peromyscus maniculatus) were determined in undisturbed soils. Spermophilus constructed two distinct burrow systems: over 120 cm deep and less than 60 cm deep. The deeper systems were significantly longer and had larger volume than the shallower burrows and the systems constructed by the other species. Burrow parameters for Dipodomys were bimodal, suggesting deep and shallow burrows, but this was not demonstrated statistically. All parameters for Dipodomys burrows were similar to shallow Spermophilus burrows. Volumes of both were significantly greater than volumes for Peromyscus and Microtus. A significant portion of the variability of all parameters for Microtus and shallow Spermophilus burrows was explained by the distribution of soil particle sizes, but equations based on these were only of limited value in predicting burrow parameters.
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Chinese Pangolin is nocturnal, shy, non-aggressive, solitary, strange and burrowing mammal which has received low scientific attention. Little information is known about its biology, distribution and status. In Nepal it is distributed in many districts and protected areas. The status of this mammal is decreasing in the country but there is no any research regarding its biology, status and distribution. Although, this mammal is protected nationally and internationally, it is facing too much problems due to habitat destruction and illegal trade. This article mainly focuses on the biology, status, distribution and conservation threats of this ecologically beneficial handsome creature. DOI: http://dx.doi.org/10.3126/init.v5i0.10267 The Initiation 2013 Vol.5; 164-170
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The zoogeomorphological impact of burrowing animals varies in time and space as a result of the particular life history traits of the organisms involved, the patchy distribution of habitat resources, and fluctuations in population size. Such ecological complexity presents a major challenge for biogeomorphologists wishing to upscale from individuals to populations. Using a unique ecological data set for Eurasian badgers (Meles meles L.) in Wytham Woods, Oxfordshire, UK, we show that direct zoogeomorphological impact (soil displacement during sett excavation) is constrained by fluctuations in overall population size. Modeled digging rates for individual badgers (0.19–4.51 m3 yr−1) varied depending on the ecological function of the sett they are associated with, and we estimate that the whole population has displaced 304–601 ± 72 m3 of soil during the construction of 64 setts. This represents an overall excavation rate of 6.7–19.4 m3 (6.0–17.5 t) yr−1 in sett areas or 1.42–4.12 g m−2 yr−1 when averaged over the whole 424 ha woodland. As well as direct soil displacement, badger digging exposes material that is initially susceptible to erosion by water relative to undisturbed and litter-covered soils. Over time, setts become stabilized, representing unique landforms that persist in the landscape for decades to centuries.
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Habitat selection by Chinese pangolin (Manis pentadactyla) in winter was studied in Dawuling Natural Reserve, Maoming City of Guangdong Province, China, from December 1999 through February 2001.The objective of this study was to determine the pangolin's habitat requirement in winter during poor environmental conditions. The dominant habitat of this Reserve was classified into 4 types, i.e. mixed coniferous and broadleaf forest (MCBF), evergreen broadleaf forest (EBF), coniferous forest (CF) and shrub forest (SF). The results showed that the ranking of vegetation types selected by pangolins in winter was MCBF > SF > EBF > CF. Pangolins preferred MCBF, and avoided CF. The environmental factors preferred by pangolins in winter were 30°-60° steep slopes, middle of slopes and bottom of slopes, sunny slopes, distance from human disturbance source exceeding 1 000 m with a minor disturbance degree, heavy (81 % - 100 %) undergrowth with good shelter conditions, moderate (760 - 1 500 m) elevation, and medium (31 % - 70 %) closure of arbor canopy. The surroundings factors avoided by pangolins were sharp slopes steeper than 60° or gentle slopes less than 30°, shady slopes, distance from human disturbance source within 1 000 m, dense (71 % - 100 %) or sparse (0 % - 30 %) closure of arbor canopy, medium or lower coverage (0 % - 50 %) of undergrowth, and the top of the slope. Pangolins preferred south-facing burrow entrances with thick cover, and avoided north-facing burrow entrances with bare or poor shelter. The thick layer of shrub and herbs growing under the tree canopy appeared to be especially important to pangolins during winter.
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