El Niño impacts on human-modified tropical forests: Consequences for dung beetle diversity and associated ecological processes

Our knowledge of how tropical forest biodiversity and functioning respond to anthropogenic and climate-associated stressors is limited. Research exploring El Niño impacts are scarce or based on single post-disturbance assessments, and few studies assess forests previously affected by anthropogenic disturbance. Focusing on dung beetles and associated ecological functions, we assessed (a) the ecological effects of a strong El Niño, (b) if post-El Niño beetle responses were influenced by previous forest disturbance, and (c) how these responses compare between forests impacted only by drought and those affected by both drought and fires. We sampled 30 Amazonian forest plots distributed across a gradient of human disturbance in 2010, 2016, and 2017—approximately 5 years before, and 3–6 and 15–18 months after the 2015–16 El Niño. We found 14,451 beetles from 98 species and quantified the beetle-mediated dispersal of >8,600 seed mimics and the removal of c. 30 kg of dung. All dung beetle responses (species richness, abundance, biomass, compositional similarity to pre-El Niño condition, and rates of dung removal and seed dispersal) declined after the 2015–16 El Niño, but the greatest immediate losses (i.e., in 2016) were observed within fire-affected forests. Previous forest disturbance also influenced post-El Niño dung beetle species richness, abundance, and species composition. We demonstrate that dung beetles and their ecological functions are negatively affected by climate-associated disturbances in human-modified Amazonian forests and suggest that the interaction between local anthropogenic and climate-related stressors merits further investigation.     

Experimental evidence for the effects of dung beetle functional group richness and composition on ecosystem function in a tropical forest

Much of the literature on the relationship between species richness or functional group richness and measures of ecosystem function focuses on a restricted set of ecosystem function measures and taxonomic groups. Few such studies have been carried out under realistic levels of diversity in the field, particularly in high diversity ecosystems such as tropical forests. We used exclusion experiments to study the effects of dung beetle functional group richness and composition on two interlinked and functionally important ecological processes, dung removal and secondary seed dispersal, in evergreen tropical forest in Sabah, Malaysian Borneo. Overall, both dung and seed removal increased with dung beetle functional group richness. However, levels of ecosystem functioning were idiosyncratic depending on the identity of the functional groups present, indicating an important role for functional group composition. There was no evidence for interference or competition among functional groups. We found strong evidence for overyielding and transgressive overyielding, suggesting complementarity or facilitation among functional groups. Not all mixtures showed transgressive overyielding, so that complementarity was restricted to particular functional group combinations. Beetles in a single functional group (large nocturnal tunnellers) had a disproportionate influence on measures of ecosystem function: in their absence dung removal is reduced by approximately 75%. However, a full complement of functional groups is required to maximize ecosystem functioning. This study highlights the importance of both functional group identity and species composition in determining the ecosystem consequences of extinctions or altered patterns in the relative abundance of species.     

Dung Beetle Community and Functions along a Habitat-Disturbance Gradient in the Amazon: A Rapid Assessment of Ecological Functions Associated to Biodiversity

Although there is increasing interest in the effects of habitat disturbance on community attributes and the potential consequences for ecosystem functioning, objective approaches linking biodiversity loss to functional loss are uncommon. The objectives of this study were to implement simultaneous assessment of community attributes (richness, abundance and biomass, each calculated for total-beetle assemblages as well as small- and large-beetle assemblages) and three ecological functions of dung beetles (dung removal, soil perturbation and secondary seed dispersal), to compare the effects of habitat disturbance on both sets of response variables, and their relations. We studied dung beetle community attributes and functions in five land-use systems representing a disturbance gradient in the Brazilian Amazon: primary forest, secondary forest, agroforestry, agriculture and pasture. All response variables were affected negatively by the intensification of habitat disturbance regimes, but community attributes and ecological functions did not follow the same pattern of decline. A hierarchical partitioning analysis showed that, although all community attributes had a significant effect on the three ecological functions (except the abundance of small beetles on all three ecological functions and the biomass of small beetles on secondary dispersal of large seed mimics), species richness and abundance of large beetles were the community attributes with the highest explanatory value. Our results show the importance of measuring ecological function empirically instead of deducing it from community metrics.     

Linking functional group richness and ecosystem functions of dung beetles: an experimental quantification

Dung beetles form an insect group that fulfils important functions in terrestrial ecosystems throughout the world. These include nutrient cycling through dung removal, soil bioturbation, plant growth, secondary seed dispersal and parasite control. We conducted field experiments at two sites in the northern hemisphere temperate region in which dung removal and secondary seed dispersal were assessed. Dung beetles were classified in three functional groups, depending on their size and dung manipulation method: dwellers, large and small tunnelers. Other soil inhabiting fauna were included as a fourth functional group. Dung removal and seed dispersal by each individual functional group and combinations thereof were estimated in exclusion experiments using different dung types. Dwellers were the most diverse and abundant group, but tunnelers were dominant in terms of biomass. All dung beetle functional groups had a clear preference for fresh dung. The ecosystem services in dung removal and secondary seed dispersal provided by dung beetles were significant and differed between functional groups. Although in absolute numbers more dwellers were found, large tunnelers were disproportionally important for dung burial and seed removal. In the absence of dung beetles, other soil inhabiting fauna, such as earthworms, partly took over the dung decomposing role of dung beetles while most dung was processed when all native functional groups were present. Our results, therefore, emphasize the need to conserve functionally complete dung ecosystems to maintain full ecosystem functioning.     

Post‐Dispersal Seed Removal and Germination Selected Tree Species Dispersed by Cebus capucinus on Barro Colorado Island,Panama1

Dispersal quality, an important component of seed disperser effectiveness, may strongly affect the rate of plant recruitment. Here we evaluated the quality of Cebus monkey dispersal by comparing the secondary removal fate and germination of fresh and Cebus‐ingested seeds of nine tree species on Barro Colorado Island, Panama. Overall, rates of secondary seed removal by vertebrates were low, with most Cebus defecations remaining undisturbed for extended periods on the forest floor. Only four of 30 feces were completely buried by dung beetles, and we found significantly higher vertebrate removal of defecated seeds than control seeds for only one species, Cordia bicolor. Seed germination varied greatly between plant taxa. Seeds of 3 out of 9 species showed significantly higher percent germination after monkey gut passage than control fresh seeds. Germination times tended to be shorter for defecated than for control seeds but were significantly different only for one of nine species, Cecropia insignis. Low rates of seed removal from Cebus feces, coupled with high germination probabilities, suggest high dispersal effectiveness for Cebus and contrasts strongly with patterns of post‐dispersal seed fate recorded for other primate species.     

Linking Biodiversity,the Environment and Ecosystem Functioning: Ecological Functions of Dung Beetles Along a Tropical Elevational Gradient

Biodiversity loss and anthropogenic environmental changes are known to impact ecosystem functions and services. However, there are still some uncertainties such as confounding environmental factors other than community attributes that affect ecosystem functioning. Our goal was to understand what factors influence the performance of Scarabaeinae dung beetle functions, testing the hypothesis that both community attributes and environmental variables influence the performance. Toward this aim, we collected dung beetles along an elevational gradient (800–1400 m a.s.l.) in the Espinhaço mountain range (Brazil) and quantified dung beetle functions, that is, dung removal, soil excavation and secondary seed dispersal. We recorded data on environmental factors related to climate, soil and vegetation and evaluated their effects on dung beetle functions. Dung beetle ecological functions declined with elevation and the decrease was more pronounced than richness, indicating that there are other factors involved in functions performance besides diversity of beetles. Indeed, we found that the ecological functions measured were dependent on both dung beetle community attributes and environmental factors. Climate, soil and vegetation influenced dung beetle function performance as much as richness, abundance and body size. Dung beetle functional diversity did not explain any of the functions measured. Our study demonstrates that ecological functions are directly influenced by both community attributes and environmental variables and confirms the link between biodiversity, environment and ecosystem functioning.     

Potential of endozoochorous seed dispersal by sheep in calcareous grasslands: correlations with seed traits

Questions: What is the potential of sheep to serve as seed dispersers via ingestion and defecation in calcareous grasslands? Is the presence of viable seeds from dung correlated with specific seed traits? Location: Calcareous grasslands, South Limburg, the Netherlands/Belgium. Methods: Dung samples (n=24) from sheep were collected between September 2006 and November 2007 from five sites with Mesobromion plant communities, and communities of Nardo‐Galion saxatilis. Germinability and identity of seeds in the dung samples were ascertained from germination of seedlings under glasshouse conditions. Seed traits of species with viable seeds in dung were compared with those present in the local species pool. Results: Seventy‐two plant species from 23 plant families had viable seeds in sheep dung. The plant families encountered most frequently were Gramineae and Compositae. The most abundant and frequently recorded plant species in dung samples was Urtica dioica, accounting for >80% of the total number of seeds. Mean seed density in sheep dung was 0.8 seeds g^?1 dry matter. Seeds with low seed mass and a high seed longevity index were over‐represented in dung. Viable seeds >2.5 mg were infrequent in the dung samples. Conclusions: We conclude that sheep are potentially important dispersers of plant species in Dutch calcareous grasslands. Although smaller seeds were relatively abundant in sheep dung, it cannot be excluded that this was mainly caused by differences in seed abundance.     

Ecosystem functions in natural and anthropogenic ecosystems across the East African coastal forest landscape

Habitat identity and landscape configuration significantly shape species communities and affect ecosystem functions. The conservation of natural ecosystems is of particular relevance in regions where landscapes have already been largely transformed into farmland and where habitats suffer under resource exploitation. The spillover of ecosystem functions from natural ecosystems into farmland may positively influence agricultural productivity and human livelihood quality. We measured three proxies of ecosystem functioning: Pollinator diversity (using pan traps), seed dispersal (with a seed removal experiment), and predation (using dummy caterpillars). We assessed these ecosystem functions in three forest types of the East African dry coastal forest (Brachystegia forest, Cynometra forest, and mixed forest), as well as in adjoining farmland and in plantations of exotic trees (Eucalyptus mainly). We measured ecosystem functions at 20 plots for each habitat type, and along gradients ranging from the forest into farmland. We also recorded various environmental parameters for each study plot. We did not find significant differences of ecosystem functions when combining all proxies assessed, neither among the three natural forest types, nor between natural forest and plantations. However, we found trends for single ecosystem functions. We identified highest pollinator diversity along the forest margin and in farmlands. Vegetation cover and blossom density affected the level of predation positively. Based on our findings, we suggest that flowering gardens around housings and woodlots across farmland areas support ecosystem functioning and thus improve human livelihood quality. We conclude that levels of overall ecosystem functions are affected by entire landscapes, and high landscape heterogeneity, as found in our case, might blur potential negative effects and trends arising from habitat destruction and degradation.     

Dung Beetle Communities and Seed Dispersal in Primary Forest and Disturbed Land in Amazonia1

Seeds from tropical fruiting trees ingested and defecated on the soil surface by primary dispersers (such as primates) are vulnerable to destruction from rodents, insects, and fungi. Burial by dung beedes as an incidental result of their feeding and nesting activities often provides these seeds with refugia from attack. To examine the effect of habitat disturbance on the dung beetle communities involved in this process, I surveyed dung beedes at three sites in the Amazon basin, in die states of Parí, Amazonas, and Rondónia, Brazil. Through principal component analysis on measurements of size and behavioral characters of beedes, I determined the relative quality of beetle species as seed dispersers (dispersal defined as horizontal or vertical movement of seeds) and ranked them into seed dispersal guilds. I used correspondence analysis to examine in what habitats (primary forest or varying degrees of disturbed habitat) these guilds were abundant. Most guilds decreased with increasing habitat disturbance, but one guild made up of large nocturnal burrowers (primarily Dichotomius) became more abundant with increasing disturbance (up to the level of highest disturbance surveyed), at which point all dung beede species became scarce. Clear–cuts had lower species richness, lower abundance, and lower biomass dian forested areas. These results imply that clear–cutting is detrimental to all seed dispersal dynamics in tropical rain forests, but that some levels of disturbance allow enough disperser activity to preserve this ecosystem function.     

Effects of species evenness and dominant species identity on multiple ecosystem functions in model grassland communities

Ecosystems provide multiple services upon which humans depend. Understanding the drivers of the ecosystem functions that support these services is therefore important. Much research has investigated how species richness influences functioning, but we lack knowledge of how other community attributes affect ecosystem functioning. Species evenness, species spatial arrangement, and the identity of dominant species are three attributes that could affect ecosystem functioning, by altering the relative abundance of functional traits and the probability of synergistic species interactions such as facilitation and complementary resource use. We tested the effect of these three community attributes and their interactions on ecosystem functions over a growing season, using model grassland communities consisting of three plant species from three functional groups: a grass (Anthoxanthum odoratum), a forb (Plantago lanceolata), and a N-fixing forb (Lotus corniculatus). We measured multiple ecosystem functions that support ecosystem services, including ecosystem gas exchange, water retention, C and N loss in leachates, and plant biomass production. Species evenness and dominant species identity strongly influenced the ecosystem functions measured, but spatial arrangement had few effects. By the end of the growing season, evenness consistently enhanced ecosystem functioning and this effect occurred regardless of dominant species identity. The identity of the dominant species under which the highest level of functioning was attained varied across the growing season. Spatial arrangement had the weakest effect on functioning, but interacted with dominant species identity to affect some functions. Our results highlight the importance of understanding the role of multiple community attributes in driving ecosystem functioning.     

Tamarins and Dung Beetles: An Efficient Diplochorous Dispersal System in the Peruvian Amazonia

Dung beetles fulfill several key functions in ecosystems but their role as secondary seed dispersers is probably one of the most complex ones. Various factors, such as seed characteristics, dispersal pattern induced by the primary disperser, season, and habitat, can affect the seed–beetle interaction. Particularly little is known about the fate of seeds primarily dispersed in small feces. The aim of this study was to investigate the effects of these factors on the dung beetle community (species composition, number and size of individuals) and its consequences on burial occurrence and depth of seeds primarily dispersed by two tamarin species. We captured dung beetles in a Peruvian rain forest with 299 dung‐baited pitfall traps to characterize the dung beetle community. Seed burial occurrence and depth were assessed by marking in situ 551 dispersed seeds in feces placed in cages. Among these seeds, 22.5 percent were buried by dung beetles after 2 d. We observed a significant effect of the amount of dung, season, time of deposition, and habitat on the number of individuals and species of dung beetles, as well as on seed burial occurrence and depth, while the tamarin species significantly influenced only the number and the size of dung beetles. This seed dispersal loop is particularly important for forest regeneration: small to large seeds dispersed by tamarins in secondary forest can be buried by dung beetles. These seeds can thus benefit from a better protection against predation and a more suitable microenvironment for germination, potentially enhancing seedling recruitment.     

Functional diversity in fruit‐frugivore interactions: a field experiment with Mediterranean mammals

Using field seed sowings, we assessed how four mammal species (Meles meles, Vulpes vulpes, Sus scrofa, and Oryctolagus cuniculus) influenced seed germination in three fleshy‐fruited Mediterranean shrubs (Corema album, Pyrus bourgaeana, and Rubus ulmifolius). We predicted that gut passage and removal away from mother plants would enhance the quantity, speed, and asynchrony of seed germination. Results showed that percent germination was altered by gut passage, but that the magnitude and even the direction of such effects varied according to plant and disperser species. Likewise, dispersal away from mother plants affected the percentage and germination speed in some species but not others. Gut passage increased asynchrony of germination in Rubus and Pyrus, and removal from the mother plant increased asynchrony in Rubus, which likely enhances plant fitness in unpredictable environments. Gut passage generally had a stronger effect on germination than removal away from mother plants, but for some species both factors were similarly influential. Therefore, the combined effects of both seed dispersal services varied individually among fruit and frugivore species, leading to unusually high functional diversity in this seed dispersal mutualism.     

Seed dispersal and predation in the endemic Atlantic rainforest palm <Emphasis Type="Italic">Astrocaryum aculeatissimum</Emphasis> across a gradient of seed disperser abundance

Tropical forests have been subject to intense hunting of medium and large frugivores that are important in dispersing large-seeded species. It has been hypothesized that in areas with extinction or low abundance of medium and large-bodied animals the density of small rodents may increase. Therefore, this increment in the density of small rodents may compensate for the absence or low abundance of medium and large frugivores on seed removal and seed dispersal. Here, we fill up this gap in the literature by determining if seed removal, seed dispersal, and seed predation by small rodents (spiny rats, Trinomys inheringi and squirrels, Sciurus ingrami) are maintained in defaunated areas. We accessed seed removal, seed dispersal, seed predation, and seedling recruitment of an endemic Atlantic rainforest palm, Astrocaryum aculeatissimum, in a gradient of abundance of agoutis. We found that seed removal, scatter hoarding, and seed predation increase with the abundance of agoutis. In contrast, the proportion of dispersed but non-cached seeds decreased with the abundance of agoutis. We did not find any effect of the abundance of agoutis on seed dispersal distance, but we did find a positive trend on the density of seedlings. We concluded that small rodents do not compensate the low abundance of agoutis on seed removal, scatter hoarding, and seed predation of this palm tree. Moreover, areas in which agoutis are already extinct did not present any seed removal or scatter hoarding, not even by small rodents. This study emphasizes both the importance of agoutis in dispersing seeds of A. aculeatissimum and the collapse in seed dispersal of this palm in areas where agoutis are already extinct.     

Seed dispersal by Galápagos tortoises

Aim Large‐bodied vertebrates often have a dramatic role in ecosystem function through herbivory, trampling, seed dispersal and nutrient cycling. The iconic Galápagos tortoises (Chelonoidis nigra) are the largest extant terrestrial ectotherms, yet their ecology is poorly known. Large body size should confer a generalist diet, benign digestive processes and long‐distance ranging ability, rendering giant tortoises adept seed dispersers. We sought to determine the extent of seed dispersal by Galápagos tortoises and their impact on seed germination for selected species, and to assess potential impacts of tortoise dispersal on the vegetation dynamics of the Galápagos. Location Galápagos, Ecuador. Methods To determine the number of seeds dispersed we identified and counted intact seeds from 120 fresh dung piles in both agricultural and national park land. To estimate the distance over which tortoises move seeds we used estimated digesta retention times from captive tortoises as a proxy for retention times of wild tortoises and tortoise movement data obtained from GPS telemetry. We conducted germination trials for five plant species to determine whether tortoise processing influenced germination success. Results In our dung sample, we found intact seeds from > 45 plant species, of which 11 were from introduced species. Tortoises defecated, on average, 464 (SE 95) seeds and 2.8 (SE 0.2) species per dung pile. Seed numbers were dominated by introduced species, particularly in agricultural land. Tortoises frequently moved seeds over long distances; during mean digesta retention times (12 days) tortoises moved an average of 394 m (SE 34) and a maximum of 4355 m over the longest recorded retention time (28 days). We did not find evidence that tortoise ingestion or the presence of dung influenced seed germination success. Main conclusions Galápagos tortoises are prodigious seed dispersers, regularly moving large quantities of seeds over long distances. This may confer important advantages to tortoise‐dispersed species, including transport of seeds away from the parent plants into sites favourable for germination. More extensive research is needed to quantify germination success, recruitment to adulthood and demography of plants under natural conditions, with and without tortoise dispersal, to determine the seed dispersal effectiveness of Galápagos tortoises.     

Do riparian reserves support dung beetle biodiversity and ecosystem services in oil palm-dominated tropical landscapes?

Agricultural expansion and intensification are major threats to global biodiversity, ecological functions, and ecosystem services. The rapid expansion of oil palm in forested tropical landscapes is of particular concern given their high biodiversity. Identifying management approaches that maintain native species and associated ecological processes within oil palm plantations is therefore a priority. Riparian reserves are strips of forest retained alongside rivers in cultivated areas, primarily for their positive hydrological impact. However, they can also support a range of forest‐dependent species or ecosystem services. We surveyed communities of dung beetles and measured dung removal activity in an oil palm‐dominated landscape in Sabah, Malaysian Borneo. The species richness, diversity, and functional group richness of dung beetles in riparian reserves were significantly higher than in oil palm, but lower than in adjacent logged forests. The community composition of the riparian reserves was more similar to logged forest than oil palm. Despite the pronounced differences in biodiversity, we did not find significant differences in dung removal rates among land uses. We also found no evidence that riparian reserves enhance dung removal rates within surrounding oil palm. These results contrast previous studies showing positive relationships between dung beetle species richness and dung removal in tropical forests. We found weak but significant positive relationships between riparian reserve width and dung beetle diversity, and between reserve vegetation complexity and dung beetle abundance, suggesting that these features may increase the conservation value of riparian reserves. Synthesis and applications: The similarity between riparian reserves and logged forest demonstrates that retaining riparian reserves increases biodiversity within oil palm landscapes. However, the lack of correlation between dung beetle community characteristics and dung removal highlights the need for further research into spatial variation in biodiversity–ecosystem function relationships and how the results of such studies are affected by methodological choices.     

The role of nocturnal omnivorous lemurs as seed dispersers in Malagasy rain forests

Fruit-eating animals play important roles as seed dispersal agents in terrestrial systems. Yet, the extent to which seed dispersal by nocturnal omnivores may facilitate germination and the recruitment of plant communities has rarely been investigated. Characterizing their roles in seed dispersal is necessary to provide a more complete picture of how seed dispersal processes affect ecosystem functioning. We investigated the roles and impacts of two species of nocturnal omnivorous lemur species, Microcebus jollyae and M. rufus, on seed dispersal in Madagascar's rain forests, through analysis of fecal samples and germination experiments. Data show that these lemur species, which are among the world's smallest primates, dispersed 22 plant species from various forest strata and that the defecated seeds germinated faster and at higher rates than control seeds for the eight plant species we tested. Even though mouse lemurs dispersed both native and non-native plant species, non-native plant species represented a relatively small proportion (17%). These results demonstrate that overlooked nocturnal omnivores can act as important seed dispersers, which may have critical implications for forest regeneration and the maintenance of plant diversity in fragmented/degraded forests. Finally, we provide critical insights into the previously unobserved behavior and diet of endangered nocturnal lemurs for their effective conservation.     

Intertwined effects of defaunation,increased tree mortality and density compensation on seed dispersal

Contemporary defaunation has profound ecological consequences ranging from local or even global co-extinctions of interacting species to the loss of ecosystem functions and services critical for humanity. Other components of global change (climate change, introduced pests, land use changes) are also harming ecosystem functioning by augmenting tree mortality worldwide. Defaunation and increased tree mortality often coincide in many human-altered ecosystems but whether they interact, leading to non-additive effects on ecosystem functioning, remains largely unknown. However, under some ecological circumstances, the decline or extirpation of one species due to defaunation can be neutralized by increases in the abundance of some functionally similar species (i.e. ‘density compensation’). We combined long-term field data with individual-based modelling to investigate the potential interactive effects of seed disperser loss, increased tree mortality and density compensation on seed dispersal in a heterogeneous landscape. Our simulation experiments showed that both stressors markedly limit not only the quantity of seed dispersal but also its quality since the impact on seed dispersal strongly varied among habitat types that differ strikingly in suitability for tree establishment. Density compensation had a marked positive effect on seed dispersal which, however, was largely limited under increased tree mortality. The combined negative effects of defaunation and increased tree mortality on seed dispersal were lower than the expected additive effect. This highlights the need to account for the joint operation of multiple stressors to accurately predict the impacts of global change on the link between biodiversity and ecosystem functioning.     

Seed dispersal distance,seed morphology,and recruitment in the Chilean sclerophyllous tree Quillaja saponaria: implications for passive restoration in a semiarid ecosystem

Recolonization of wind-dispersed tree species in degraded areas may decline with distance from remnant forest fragments because seed rain frequently decreases with distance from the seed source. However, regeneration of these species may be even more limited to sites close to the seed source if dispersal distance is negatively affected by seed mass, and germination probability is positively affected by seed mass. We evaluated these hypotheses in a Mediterranean-type ecosystem of central Chile, using the wind-dispersed tree species Quillaja saponaria. We assessed the seed rain curve in a degraded open area adjacent to a remnant forest fragment of this species, and related seed mass with dispersal distance from the seed source. Then, we evaluated the relationship between seed mass, germination, and seedling growth, and if seeds that fall nearer the seed source have greater germination probability. We found a decreasing seed rain with the distance from the seed source. Seed mass was not related to dispersal distance, although seeds with higher wing area dispersed further. Germination probability was significantly and positively related to the seed mass. We observed no significant relationship between distance and germination probability. We conclude that germination probability of this species does not vary along the seed rain curve, and that the recruitment density would be greater near the seed source only due to decreasing seed rain with distance. Our results suggest that this species has the potential to be passively restored in degraded areas, especially within the first 70 m from the remnant forest fragments.

     

Antelope mating strategies facilitate invasion of grasslands by a woody weed

Intra and interspecific variation in frugivore behaviour can have important consequences for seed dispersal outcomes. However, most information comes from among‐species comparisons, and within‐species variation is relatively poorly understood. We examined how large intraspecific differences in the behaviour of a native disperser, blackbuck antelope Antilope cervicapra, influence dispersal of a woody invasive, Prosopis juliflora, in a grassland ecosystem. Blackbuck disperse P. juliflora seeds through their dung. In lekking blackbuck populations, males defend clustered or dispersed mating territories. Territorial male movement is restricted, and within their territories males defecate on dung‐piles. In contrast, mixed‐sex herds range over large areas and do not create dung‐piles. We expected territorial males to shape seed dispersal patterns, and seed deposition and seedling recruitment to be spatially localized. Territorial males had a disproportionately large influence on seed dispersal. Adult males removed twice as much fruit as females, and seed arrival was disproportionately high on territories. Also, because lek‐territories are clustered, seed arrival was spatially highly concentrated. Seedling recruitment was also substantially higher on territories compared with random sites, indicating that the local concentration of seeds created by territorial males continued into high local recruitment of seedlings. Territorial male behaviour may, thus, result in a distinct spatial pattern of invasion of grasslands by the woody P. juliflora. An ex situ experiment showed no beneficial effect of dung and a negative effect of light on seed germination. We conclude that large intraspecific behavioural differences within frugivore populations can result in significant variation in their effectiveness as seed dispersers. Mating strategies in a disperser could shape seed dispersal, seedling recruitment and potentially plant distribution patterns. These mating strategies may aid in the spread of invasives, such as P. juliflora, which could, in turn, negatively influence the behaviour and ecology of native dispersers.     

Dispersión Primaria de Semillas por Primates y Dispersión Secundaria por Escarabajos Coprófagos en Tikal,Guatemala1

We linked primary dispersal by spider monkeys (Ateles geoffroyi) and howler monkeys (Alouatta pigra) to post‐dispersal seed fate by studying the effects of dung type and defecation pattern on secondary seed dispersal by dung beetles. First, we described the defecation patterns for both primate species. Howler monkeys generally defecated in groups (88% of observed defecations), with each individual producing on average 31 g of dung, resulting in a large area of the forest floor (31 m²) covered by large amounts of dung (clumped spatial pattern). Spider monkeys generally (96% of observed defecations) defecated individually, each individual producing an average of 11 g of dung, resulting in a small area of the forest floor (2 m²) covered by small amounts of dung (scattered spatial pattern). Secondly, we captured dung beetles using as bait the dung of both primate species, to detect differences in the assemblages of these secondary seed dispersers attracted to the dung of both primates. More individual dung beetles, but not more species, were attracted to howler monkey dung than to spider monkey dung. Finally, we assessed experimentally (using plastic beads as seed mimics) how dung type (Ateles vs. Alouatta) and defecation pattern (scattered vs. clumped) affect secondary seed dispersal by dung beetles. We found that post‐dispersal seed fate was affected by dung type, with more seeds being buried when present in howler monkey dung, than in spider monkey dung, but was not affected by defecation pattern. It is important to consider post‐dispersal processes, such as secondary seed dispersal by dung beetles, when comparing species of primary dispersers.