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1.
Vast areas of the African savanna landscapes are characterized by tree‐covered Macrotermes termite mounds embedded within a relatively open savanna matrix. In concert with termites, large herbivores are important determinants of savanna woody vegetation cover. The relative cover of woody species has considerable effects on savanna function. Despite the potentially important ecological relationships between termite mounds, woody plants, large herbivores, and birds, these associations have previously received surprisingly little attention. We experimentally studied the effects of termites and large herbivores on the avian community in Lake Mburo National Park, Uganda, where woody vegetation is essentially limited to termite mounds. Our experiment comprised of four treatments in nine replicates; unfenced termite mounds, fenced mounds (excluding large mammals), unfenced adjacent savanna, and fenced savanna. We recorded species identity, abundance, and behavior of all birds observed on these plots over a two‐month period, from late dry until wet season. Birds used termite mounds almost exclusively, with only 3.5% of observations occurring in the treeless intermound savanna matrix. Mean abundance and species richness of birds doubled on fenced (large herbivores excluded) compared to unfenced mounds. Feeding behavior increased when large mammals were excluded from mounds, both in absolute number of observed individuals, and relative to other behaviors. This study documents the fundamental positive impact of Macrotermes termites on bird abundance and diversity in an African savanna. Birds play crucial functional roles in savanna ecosystems, for example, by dispersing fruits or regulating herbivorous insect populations. Thus, the role of birds in savanna dynamics depends on the distribution and abundance of termite mounds.  相似文献   

2.
Termite mounds contribute to the spatial heterogeneity of ecological processes in many savannas, but the underlying patterns and determinants of mound distributions remain poorly understood. Using the Carnegie Airborne Observatory (CAO), we mapped the distribution of termite mounds across a rainfall gradient within a river catchment (~ 27 000 ha) of the Kruger National Park, South Africa. We assessed how different factors were associated with the distribution and height of termite mounds at three spatial scales: the entire catchment, among three broad vegetation types, and on individual hillslope crests. Abiotic factors such as the underlying geology and mean annual precipitation shaped mound densities at broad scales, while local hillslope morphology strongly influenced mound distribution at finer scales, emphasising the importance of spatial scale when assessing mound densities. Fire return period had no apparent association with mound densities or height. Mound density averaged 0.46 mounds ha?1, and exhibited a clustered pattern throughout the landscape, occurring at relatively high densities (up to 2 mounds ha?1) on crests, which are nutrient‐poor elements of the landscape. Mounds exhibited significant over‐dispersion (even spacing) at scales below 60 m so that evenly spaced aggregations of termite mounds are embedded within a landscape of varying mound densities. The tallest mounds were found in dry savanna (500 mm yr?1) and were positively correlated with mound density, suggesting that dry granitic savannas are ideal habitat for mound‐building termites. Mound activity status also varied significantly across the rainfall gradient, with a higher proportion of active (live) mounds in the drier sites. The differential spacing of mounds across landscapes provides essential nutrient hotspots in crest locations, potentially sustaining species that would otherwise not persist. The contribution to biodiversity and ecosystem functioning that mounds provide is not uniform throughout landscapes, but varies considerably with spatial scale and context.  相似文献   

3.
Knowledge of the distribution and nutrient values of key resources supporting the survival of wildlife species is integral for an effective conservation planning and management of the species. In the Miombo ecosystem of the Ugalla Game Reserve, African elephants (Loxodonta africana Blumenbach 1797), eat soil, that is geophagy, from certain termite mounds. We mapped that all the geophagic termite mounds are exclusively situated in the flood plain. To understand why soils from some termite mounds are eaten, we collected and analysed soil samples from 10 geophagic termite mounds, seven nongeophagic termite mounds and 13 samples from the surrounding flood plain. Percentage of clay content did not differ significantly among the soil samples. Soils from geophagic termite mounds were richer in mineral elements compared with other soil samples. The results demonstrate that the driver for geophagic behaviour is related to rich mineral element contents found in geophagic termite mounds made of the mineral‐enriching termites (Macrotermes). Thus, geophagic termite mounds play a role in elephant's dietary needs and possibly influence their movement patterns in Ugalla, as the elephants cannot obtain enough minerals from their feeds. Geophagic termite mounds should be protected from potential destructive land uses, such as airstrip construction.  相似文献   

4.
This review is an analysis of the dimensions of termite infestation in African savannas. The aim of this work is to draw the attention of ecologists, conservationists, policy makers and farmers to the current and future threats of subterranean termites to the functioning and sustainability of such ecosystem habitats. This study analyzes and describes termite problem (questionable changes in density and assemblage structure) in selected African savannah ecosystems, synthesizes information on the effects of various human induced habitat disturbance regimes on termites’ assemblage structure, predators, nests and feed resources to generate hypotheses relating termite infestation with anthropogenic activities; it describes and critiques existing termite management practices. The review is suggestive that the infestation and resultant undesirable effects of subterranean termites in African savannahs are largely a consequence of the inappropriate savannah management practices (overgrazing, indiscriminate tree cutting and overhunting) undertaken by humans in pursuit of various livelihood options. Based on the evidence presented herein, we hypothesized that (1) human induced habitat disturbance in savanna ecosystems alters the feeding group composition of termites’ assemblages, favoring grass harvesters and polyphagus termite feeders that forage on more abundant food items, paying little attention to rarer food items and (2) habitat disturbance through activities like heavy grazing and overhunting results in decline in the populations of both macro and microscopic termite predators, which eventually enhances the proliferation of termite populations, escalates the density of termite nests particularly epigeal mounds and intensifies consumption of herbaceous savannah vegetation. The review calls for dedicated efforts to develop ecological thresholds of savannah biotic and abiotic ecosystem components in which human induced disturbance regimes trigger the destructive behavior of termites. This would provide information that will act as a precautionary savannah habitat monitoring and decision support tool to prevent future infestation of savannah habitats with termites. Also, the review shows that majority of the termite control practices are ineffective, ecologically unsustainable and above all, do not address the root cause of termite infestation and thus merely provide temporary relief to the problem. As such, termite control methods that attempt to enhance proliferation of termite predators need to be studied, developed and emphasized. This review reveals that human induced habitat disturbance depletes termites’ predator populations, leading to proliferation of termite populations particularly grass harvesters that intensify their consumption on grass biomass and eventually contribute to denudation of herbaceous vegetation cover in savannah ecosystems.  相似文献   

5.
In many ant–plant mutualisms, ants establish colonies in hollow thorns, leaf pouches, or other specialized structures on their host plants, which they then defend from herbivores. Resource heterogeneity could affect the maintenance of these mutualisms if it leads to one or both partners altering their investment in the interaction. Such a phenomenon may be especially pertinent to the Acacia–ant mutualism found in East African savannas, where termite mounds have a profound effect on the spatial structuring of resources used by both plants and ants. Here, we examined whether the proximity to termite mounds of Acacia drepanolobium trees is associated with variation in the behavior of one of their ant associates, Crematogaster nigriceps. We found that ant colonies near termite mounds had decreased aggressive responses to simulated herbivory as well as increased off‐tree movement. We hypothesize that these changes are the result of resident ant colonies near termite mounds shifting investment from defense of their host plant to foraging for nearby resources.  相似文献   

6.
Fungus‐growing termites of the subfamily Macrotermitinae together with their highly specialized fungal symbionts (Termitomyces) are primary decomposers of dead plant matter in many African savanna ecosystems. The termites provide crucial ecosystem services also by modifying soil properties, translocating nutrients, and as important drivers of plant succession. Despite their obvious ecological importance, many basic features in the biology of fungus‐growing termites and especially their fungal symbionts remain poorly known, and no studies have so far focused on possible habitat‐level differences in symbiont diversity across heterogeneous landscapes. We studied the species identities of Macrotermes termites and their Termitomyces symbionts by excavating 143 termite mounds at eight study sites in the semiarid Tsavo Ecosystem of southern Kenya. Reference specimens were identified by sequencing the COI region from termites and the ITS region from symbiotic fungi. The results demonstrate that the regional Macrotermes community in Tsavo includes two sympatric species (M. subhyalinus and M. michaelseni) which cultivate and largely share three species of Termitomyces symbionts. A single species of fungus is always found in each termite mound, but even closely adjacent colonies of the same termite species often house evolutionarily divergent fungi. The species identities of both partners vary markedly between sites, suggesting hitherto unknown differences in their ecological requirements. It is apparent that both habitat heterogeneity and disturbance history can influence the regional distribution patterns of both partners in symbiosis.  相似文献   

7.
Abstract 1. Termites (Isoptera) in tropical savannas are known as ecosystem engineers, affecting the spatial and temporal distribution of water, carbon, cations, and nutrients through their mound structures. Their mounds, however, also offer habitation to diverse taxa and feeding guilds of other invertebrates; a keystone role that has not been properly quantified. 2. The aim of this study was to explore the ecosystem role of termitaria in determining invertebrate diversity and their potential trophic interactions. We used stable isotopes to distinguish termite‐feeding invertebrates from invertebrates merely living in termite mounds under field conditions. 3. The results suggest that inquiline spiders (Arachnida) do not feed on termites directly, but on other invertebrates within the termitaria that are termitophagous, elevating the spiders three trophic levels higher than the termites. 4. This study is the first to demonstrate food web interactions among inquiline invertebrates with a stable isotope approach. It provides evidence that termites play a keystone role in the system by providing habitat for various, trophically interacting invertebrates. These results illustrate a rather unexplored ecosystem property of savanna termites.  相似文献   

8.
A key aspect of savannah vegetation heterogeneity is mosaics formed by two functional grassland types, bunch grasslands, and grazing lawns. We investigated the role of termites, important ecosystem engineers, in creating high-nutrient patches in the form of grazing lawns. Some of the ways termites can contribute to grazing lawn development is through erosion of soil from aboveground mounds to the surrounding soil surface. This may alter the nutrient status of the surrounding soils. We hypothesize that the importance of this erosion varies with termite genera, depending on feeding strategy and mound type. To test this, we simulated erosion by applying mound soil from three termite genera (Macrotermes, Odontotermes, and Trinervitermes) in both a field experiment and a greenhouse experiment. In the greenhouse experiment, we found soils with the highest macro nutrient levels (formed by Trinervitermes) promoted the quality and biomass of both a lawn (Digitaria longiflora) and a bunch (Sporobolus pyramidalis) grass species. In the field we found that soils with the highest micro nutrient levels (formed by Macrotermes) showed the largest increase in cover of grazing lawn species. By linking the different nutrient availability of the mounds to the development of different grassland states, we conclude that the presence of termite mounds influences grassland mosaics, but that the type of mound plays a crucial role in determining the nature of the effects.  相似文献   

9.
Though insectivory by large-bodied gorillas may be unexpected, researchers have reported it in all populations of gorillas studied to date. Our study of 2 well monitored groups of western gorillas (Gorilla gorilla gorilla) at Bai Hokou in Dzanga-Ndoki National Park, Central African Republic provides information on frequency and variability of termite consumption (the most commonly eaten insect) as well as some of the first direct observations of the behavior. Pooled data from both groups indicate termite feeding on 34% and 83% of days, through fecal analysis and feeding trails, respectively. Direct observations revealed that termite feeding occurred on 91% of the days for 1 group, in which the silverback fed on termites during 13% of all feeding scans, making termites the most commonly observed food item. The group that had a higher density of termite mounds in its home range consumed termites more frequently than the other group did. A higher proportion of fecal samples from the silverbacks contained termite remains than the ones from adult females and juveniles. Termite consumption was lower during the dry season, but it does not correlate with rainfall, measures of fruit availability, or fruit consumption. Displacements at termite mounds occurred more than expected, indicating that they are a patchy, sought-after food resource. Gorillas did not use tools to extract termites, but they used 2 different techniques to remove them from the cells. Though culture or social traditions may cause the variation in termite consumption across sites, further investigation of termite availability and consumption is necessary to rule out ecological and methodological explanations for observed variations.  相似文献   

10.
The building and foraging activities of termites are known to modify soil characteristics such as the heterogeneity. In tropical savannas the impact of the activity of soil-feeding termites (Cubitermes niokoloensis) has been shown to affect the properties of the soil at the aggregate level by creating new soil microenvironments (aggregate size fractions) [13]. These changes were investigated in greater depth by looking at the microbial density (AODC) and the genetic structure (automated rRNA intergenic spacer analysis: ARISA) of the communities in the different aggregate size fractions (i.e., coarse sand, fine sand, coarse silt, fine silt, and dispersible clays) separated from compartments (internal and external wall) of three Cubitermes niokoloensis mounds. The bacterial density of the mounds was significantly higher (1.5 to 3 times) than that of the surrounding soil. Within the aggregate size fractions, the termite building activity resulted in a significant increase in bacterial density within the coarser fractions (>20 m). Multivariate analysis of the ARISA profiles revealed that the bacterial genetic structures of unfractionated soil and soil aggregate size fractions of the three mounds was noticeably different from the savanna soil used as a reference. Moreover, the microbial community associated with the different microenvironments in the three termite mounds revealed three distinct clusters formed by the aggregate size fractions of each mound. Except for the 2–20 m fraction, these results suggest that the mound microbial genetic structure is more dependent upon microbial pool affiliation (the termite mound) than on the soil location (aggregate size fraction). The causes of the specificity of the microbial community structure of termite mound aggregate size fractions are discussed.This revised version was published online in November 2004 with corrections to Volume 48.  相似文献   

11.
Decomposition is a vital ecosystem process, increasingly modified by human activity. Theoretical frameworks and empirical studies that aim to understand the interplay between human land‐use, macro‐fauna and decomposition processes have primarily focused on leaf and wood litter. For a whole‐plant understanding of how land‐use and macro‐fauna influence decomposition, investigating root litter is required. Using litterbags, we quantified rates of root decomposition across contrasting tropical savanna land‐uses, namely wildlife and fire‐dominated protected areas and livestock pastureland without fire. By scanning litterbags for termite intrusion, we differentiated termite and microbial driven decomposition. Root litter was buried underneath different tree canopies (leguminous and non‐leguminous trees) and outside canopies to account for savanna landscape effects. Additionally, we established a termite cafeteria‐style experiment and common garden to explore termite selectivity of root litter and root trait relationships, respectively. After one year, we found no significant differences in root litter mass loss between wildlife dominated areas and pastureland. Instead, we found consistent species differences in root litter mass loss across land‐uses and additive and non‐additive effects of termites on root decomposition across plant species. Termite selectivity for root litter species occurred for both root and leaf litter buried near termite mounds, but was not explained by root traits measured in the common garden. Termite foraging was greater under leguminous tree canopies than other canopies; however, this did not influence rates of root decomposition. Our study suggests that land‐use has a weak direct effect on belowground processes in savannas. Instead, changes in herbaceous species composition and termite foraging have stronger impacts on belowground decomposition. Moreover, termites were not generalist decomposers of root litter, but their impact varies depending on plant species identity and likely associated root traits. This root litter selectivity by termites is likely to be an important contributor to spatial heterogeneity in savanna nutrient cycling.  相似文献   

12.
A series of papers have been published which describe the influence of vegetation and soil type on the Western Australian wheatbelt termite,Drepanotermes tamminensis (Hill), and also on its litter harvesting levels and contribution to the soil nutrient budget. This paper integrates these findings by means of a computer simulation model. The model consists of three modules which respectively describe the dynamics of litter on the ground, the dynamics of litter within termite mounds and how these in turn influence nutrient loads within the habitat. The outputs of the model suggest that this litter harvesting termite plays an important role in the nutrient dynamics of the area and it provides an estimate of the unmeasured variable, litter consumed in mounds by termites, which is consistent with measurements for other termite species with similar feeding habits.  相似文献   

13.
Some chimpanzees use 2 types of tools to extract underground termites for consumption. Chimpanzees insert thin, flexible probes into tunnels or holes in termite mounds (fishing), and sometimes use stouter, rigid sticks to first puncture the holes and also possibly to fish. Many puncturing sticks have distinctive “brushed” ends. Researchers have hypothesized that chimpanzees create the brushed ends intentionally to increase their affixibility to biting termites (Sugiyama, 1985). The results of our archaeological analysis of a large collection of puncturing sticks used by Central African chimpanzees falsifies this hypothesis, and instead agrees with the recent behavioral observations of Sanz et al. (2004; cf. Bermejo and Illera, 1999) that brushing is a coincidental result of procuring sticks from vegetation sources. The results highlight the positive contribution of an archaeological approach to problems in chimpanzee material culture and emphasize to primatologists the value of curating artifacts.  相似文献   

14.
Termites through mound construction and foraging activities contribute significantly to carbon and nutrient fluxes in nutrient-poor savannas. Despite this recognition, studies on the influence of termite mounds on carbon and nitrogen dynamics in sub-tropical savannas are limited. In this regard, we examined soil nutrient concentrations, organic carbon and nitrogen mineralization in incubation experiments in mounds of Macrotermes falciger and surrounding soils of sub-tropical savanna, northeast Zimbabwe. We also addressed whether termite mounds altered the plant community and if effects were similar across functional groups i.e. grasses, forbs or woody plants. Mound soils had significantly higher silt and clay content, pH and concentrations of calcium (Ca), magnesium (Mg), potassium (K), organic carbon (C), ammonium (NH4+) and nitrate (NO3) than surrounding soils, with marginal differences in phosphorus (P) and sodium (Na) between mounds and matrix soils. Nutrient enrichment increased by a factor ranging from 1.5 for C, 4.9 for Mg up to 10.3 for Ca. Although C mineralization, nitrification and nitrification fraction were similar between mounds and matrix soils, nitrogen mineralization was elevated on mounds relative to surrounding matrix soils. As a result, termite mounds supported unique plant communities rich and abundant in woody species but less diverse in grasses and forbs than the surrounding savanna matrix in response to mound-induced shifts in soil parameters specifically increased clay content, drainage and water availability, nutrient status and base cation (mainly Ca, Mg and Na) concentration. In conclusion, by altering soil properties such as texture, moisture content and nutrient status, termite mounds can alter the structure and composition of sub-tropical savanna plant communities, and these results are consistent with findings in other savanna systems suggesting that increase in soil clay content, nutrient status and associated changes in the plant community assemblage may be a general property of mound building termites.  相似文献   

15.
Question: What are the genesis and development of thicket clumps within a savanna landscape at geomorphically different locations and what are the driving forces? Location: The Kagera Region, in the border area of Rwanda, Uganda and Tanzania. Methods: The vegetation of 32 dry evergreen thicket clumps and their surrounding savannas have been analysed at different geomorphic locations. At each vegetation plot Na+, K+, Mg2+, Ca2+, Al3+, Fe2+/3+, H+, P, C, N, bulk density and particle size were determined for each soil horizon. The impact of soil and termite mounds on thicket clump dynamics on seasonally waterlogged plains, gentle slopes and stony hillsides were assessed. Results: Thicket clumps and their surrounding savannas have a distinct structure and floristic composition. They also have distinct soil properties although parent materials are the same. On seasonally waterlogged plains, new thicket clumps can develop on Macrotermitinae mounds; on stony hillsides, Trinervitermes and Macrotermes show a uniform distribution pattern and may initiate the genesis of thicket clumps. Conclusions: Geomorphology broadly determines the significance and interactions of the main factors affecting site‐specific vegetation dynamics. On seasonally waterlogged plains, thicket clumps are restricted to termite mounds. Since intra‐species competition dictates a minimal distance between neighbouring Macrotermitinae colonies, thicket clumps do not coalescence. By contrast, on stony hillsides, the vegetation mosaic is highly dynamic and determined by the interplay of several factors. The growth of thicket clumps is mainly a function of the fire regime and the browsing intensity. At the present time, frequent cool, early dry season fires and the near absence of large browsers have favoured the advance and coalescence of thicket clumps and forest patches on stony hillsides.  相似文献   

16.
Termite mounds are a widespread feature in most African savannas. These structures exhibit high nutrient contents and often host a special vegetation composition. In this study, we analysed mound distribution patterns of a fungus-growing termite species, Macrotermes michaelseni, an important ecosystem engineer in the savannas of Namibia. Inhabited mounds taller than 0.7 m were regularly distributed. We view this pattern as a result of intraspecific competition. The heights of mounds taller than 0.7 m were correlated positively with their distance, such that mounds closer together, i.e. up to inter-mound distances of approximately 50 m, tended to be smaller than average. This indicates that intraspecific competition for foraging areas controls mound distribution pattern and colony size. Differences between mound heights increased on the spatial scale up to inter-mound distances of 80 m. We assume that the foundation of new colonies is only possible in unoccupied patches. In such patches, young colonies are able to occur close together as they have a relatively low foraging demand and therefore a low spatial demand. In contrast, their critical distance to taller colonies with higher foraging demands is rather large, which leads to the observed increasing difference of mound heights with increasing distances between them.  相似文献   

17.
Herbivores do not forage uniformly across landscapes, but select for patches of higher nutrition and lower predation risk. Macrotermes mounds contain higher concentrations of soil nutrients and support grasses of higher nutritional value than the surrounding savanna matrix, attracting mammalian grazers that preferentially forage on termite mound vegetation. However, little is known about the spatial extent of such termite influence on grazing patterns and how it might differ in time and space. We measured grazing intensity in three African savanna types differing in rainfall and foliar nutrients and predicted that the functional importance of mounds for grazing herbivores would increase as the difference in foliar nutrient levels between mound and savanna matrix grasses increases and the mounds become more attractive. We expected this to occur in nutrient‐poor areas and during the dry season when savanna matrix grass nutrient levels are lower. Tuft use and grass N and P content were measured along transects away from termite mounds, enabling calculation of the spatial extent of termite influence on mammalian grazing. Using termite mound densities estimated from airborne light detection and ranging (LiDAR), we further upscaled field‐based results to determine the percentage of the landscape influenced by termite activity. Grasses in close proximity to termite mounds were preferentially grazed at all sites and in both seasons, but the strength of mound influence varied between savanna types and seasons. In the wet season, mounds had a relatively larger effect on grazers at the landscape scale in the nutrient‐poor, wetter savanna, whereas in the dry season the pattern was reversed with more of the landscape influenced at the nutrient‐rich, driest site. Our results reveal that termite mounds enhance the value of savanna landscapes for herbivores, but that their functional importance varies across savanna types and seasons.  相似文献   

18.
In tropical ecosystems, termite mound soils constitute an important soil compartment covering around 10% of African soils. Previous studies have shown (S. Fall, S. Nazaret, J. L. Chotte, and A. Brauman, Microb. Ecol. 28:191-199, 2004) that the bacterial genetic structure of the mounds of soil-feeding termites (Cubitermes niokoloensis) is different from that of their surrounding soil. The aim of this study was to characterize the specificity of bacterial communities within mounds with respect to the digestive and soil origins of the mound. We have compared the bacterial community structures of a termite mound, termite gut sections, and surrounding soil using PCR-denaturing gradient gel electrophoresis (DGGE) analysis and cloning and sequencing of PCR-amplified 16S rRNA gene fragments. DGGE analysis revealed a drastic difference between the genetic structures of the bacterial communities of the termite gut and the mound. Analysis of 266 clones, including 54 from excised bands, revealed a high level of diversity in each biota investigated. The soil-feeding termite mound was dominated by the Actinobacteria phylum, whereas the Firmicutes and Proteobacteria phyla dominate the gut sections of termites and the surrounding soil, respectively. Phylogenetic analyses revealed a distinct clustering of Actinobacteria phylotypes between the mound and the surrounding soil. The Actinobacteria clones of the termite mound were diverse, distributed among 10 distinct families, and like those in the termite gut environment lightly dominated by the Nocardioidaceae family. Our findings confirmed that the soil-feeding termite mound (C. niokoloensis) represents a specific bacterial habitat in the tropics.  相似文献   

19.
In African savannas, termite mounds usually serve as browsing hotspots for mammals because of their soil fertility. Van der Plas et al., in this issue, describe that browsers avoid the unpalatable, evergreen tree species on mounds of Macrotermes natalensis in a mesic savanna, preferring mainly leguminous species with high leaf N and P concentrations in the matrix. This exception is probably a consequence of the fertile soils of the study area, and highlights the importance of environmental context for assessing ecological interactions.  相似文献   

20.
Abstract Two years after an intense fire burnt large parts of the Barrens in Mediterranean Western Australia, its effects on wood-eating and litter-harvesting termites were investigated. Nine vegetation types varying in height, structure and floristics, were used for paired unburnt/burnt comparisons. Wood-eating termites were significantly less abundant and diverse in burnt stands, apparently by perishing in the fire rather than through food limitation. The harvester Tumulitermes westraliensis was not significantly affected by fire. Strategies enhancing persistence despite intense fire were construction of hard, protective clay mounds (Coptotermes frenchi, Amitermes obeuntis, T. westraliensis) and flexibility to site nests in diverse microhabitats, including the mounds of other termite species (Heterotermes).  相似文献   

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