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.     

Effects of Season and Vegetation Type on Community Organization of Dung Beetles in a Tropical Dry Forest1

Dung beetles are important components of most terrestrial ecosystems. In tropical rain forests, dung beetle communities can be very rich in number of species and individuals, and they are known to be useful bioindicators of habitat disturbance. In contrast, very little is known about the organization of dung beetle communities in tropical dry forests. The aim of this study was to describe in detail the dung beetle community of a Mexican tropical dry forest and to assess the relative importance of rainfall seasonality and forest structure in affecting the temporal and spatial dynamics of this community. Dung beetles were captured with pitfall traps at the beginning of the rainy season, the middle of the rainy season, and the middle of the dry season, in two distinct forest types: deciduous forest (DF) and semideciduous forest (SDF) at the Estación de Biología Chamela. Both rainfall seasonality and forest structure affected the community organization of dung beetles. During both rainy periods, 14 species were captured, but only three during the dry season. Dung beetles captured during the dry season were only found in the SDF. When comparing the beginning and the middle of the rainy season, differences in abundance and guild structure were also observed between both periods and between forest types, but these differences were much less pronounced.     

Effects of dung and seed size on secondary dispersal,seed predation,and seedling establishment of rain forest trees

Seeds dispersed by tropical, arboreal mammals are usually deposited singly and without dung or in clumps of fecal material. After dispersal through defecation by mammals, most seeds are secondarily dispersed by dung beetles or consumed by rodents. These post-dispersal, plant-animal interactions are likely to interact themselves, as seeds buried by dung beetles are less likely to be found by rodents than unburied seeds. In a series of three experiments with seeds of 15 species in central Amazonia (Brazil), we determined (1) how presence and amount of dung associated with seeds influences long-term seed fate and seedling establishment, (2) how deeply dung beetles bury seeds and how burial depth affects seedling establishment, and (3) how seed size affects the interaction between seeds, dung beetles, and rodents. Our overall goal was to understand how post-dispersal plant-animal interactions determine the link between primary seed dispersal and seedling establishment. On average, 43% of seeds surrounded by dung were buried by dung beetles, compared to 0% of seeds not surrounded by dung (n=2,156). Seeds in dung, however, tended to be more prone than bare seeds to predation by rodents. Of seeds in dung, probability of burial was negatively related to seed size and positively related to amount of dung. Burial of seeds decreased the probability of seed predation by rodents three-fold, and increased the probability of seedling establishment two-fold. Mean burial depth was 4 cm (0.5–20 cm) and was not related to seed size, contrary to previous studies. Probability of seedling establishment was negatively correlated with burial depth and not related to seed size at 5 or 10 cm depths. These results illustrate a complex web of interactions among dung beetles, rodents, and dispersed seeds. These interactions affect the probability of seedling establishment and are themselves strongly tied to how seeds are deposited by primary dispersers. More generally, our results emphasize the importance of looking beyond a single type of plant-animal interaction (e.g., seed dispersal or seed predation) to incorporate potential effects of interacting interactions.     

Effect of forest fragmentation on dung beetle communities and functional consequences for plant regeneration

I compared dung beetle communities and assessed some of their functional effects (dung removal, seed burial, seedling establishment) in continuous forest with those in 1-ha and 10-ha forest fragments in Central Amazonia. I followed the fate of seeds until seedling establishment for three native tree species, using clean seeds and seeds surrounded by dung. The 1-ha fragments had half the number of dung beetle species captured in continuous forest and in 10-ha fragments. The continuous forest sites and the 1-ha fragments had similar number of individuals, but in the 10-ha fragments dung beetles were twice as abundant. Mean beetle size increased with increasing forest area. Dung removal and seed burial rates were higher in continuous forest than in forest fragments. Seed predation rates were higher in the forest fragments. In all sites, the proportion of seedlings established from seeds surrounded by dung vs clean seeds was the same, and it was the same in continuous forest vs fragments. When comparing seeds that remained on the forest floor with seeds buried by dung beetles, a higher percentage of seedlings established from the latter. Conservation programs that aim to maintain the regeneration ability of forest fragments must incorporate all the important components involved in seedling establishment; in Central Amazonia these include dung beetles as secondary dispersers. It is important that studies start measuring directly not only the first-order effects of forest fragmentation on species, but also the higher-order functional effects.     

Seed Dispersal by Monkeys and the Fate of Dispersed Seeds in a Peruvian Rain Forest1

Primary seed dispersal by two species of monkeys and the effects of rodents and dung beetles on the fate of dispersed seeds are described for a rain forest in southeastern Perú. During the six-month study period (June–November 1992) spider monkeys (Ateles paniscus) dispersed the seeds of 71 plant species, whereas howler monkeys (Alouatta seniculus) dispersed seeds of 14 species. Spider and howler monkeys also differed greatly in their ranging behavior and defecation patterns, and as a consequence, produced different seed rain patterns. Monkey defecations were visited by 27 species of dung beetles (Scarabaeidae). Dung beetles buried 41 percent of the seeds in the dung, but the number of seeds buried varied greatly, according to seed size. Removal rates of unburied seeds by rodents varied between 63–97 percent after 30 d for 8 plant species. The presence of fecal material increased the percentage of seeds removed by seed predators, but this effect became insignificant with time. Although seed predators found some seeds buried in dung balls (mimicking burial by dung beetles), depth of burial significantly affected the fate of these seeds. Less than 35 percent of Brosimum lactescens seeds buried inside dung balls at a depth of 1 cm remained undiscovered by rodents, whereas at least 75 percent of the seeds escaped rodent detection at a depth of 3 cm and 96 percent escaped at 5 cm. Both dung beetles and rodents greatly affected the fate of seeds dispersed by monkeys. It is thus important to consider postdispersal factors affecting the fate of seeds when assessing the effectiveness of frugivores as seed dispersers.     

Persistence of the effect of frugivore identity on post‐dispersal seed fate: consequences for the assessment of functional redundancy

Large frugivores play an important role as seed dispersers and their extinction may affect plant regeneration. The consequences of such extinctions depend on the likelihood of other species being functionally redundant and on how post‐dispersal events are affected. We assess the functional redundancy of two seed dispersers of the Atlantic Forest, the muriqui (Brachyteles arachnoides) and the tapir (Tapirus terrestris) through the comparison of their seed dispersal quality, taking into account post‐dispersal events. We compare tapirs and muriquis for: (1) the dung beetle community associated with their feces; (2) the seed burial probability and burial depth by dung beetles; and (3) the seed mortality due to predators or other causes according to burial depth. We determine how seed burial affects seed dispersal effectiveness (SDE) and compare the dispersal quality of four plant species dispersed by these frugivores. Muriqui feces attract 16‐fold more dung beetles per gram of fecal matter and seeds experience 10.5‐fold more burial than seeds in tapir feces. In both feces types, seed mortality due to predation decreases with burial depth but seed mortality due to other causes increases. Total seed mortality differ within plant species according to the primary disperser. Therefore, the effect of seed burial on SDE varies according to the plant species, burial depth, and primary disperser. As tapirs and muriquis differently affect the seed fate, they are not functionally redundant. Since the effect of the primary disperser persists into the post‐dispersal events, we should consider the cascading effects of these processes when assessing functional redundancy.     

Monkey and dung beetle activities influence soil seed bank structure

In Neotropical forests, dung beetles act as efficient secondary dispersers of seeds that are dispersed primarily by red howler monkeys. Here, we investigated the origins of soil seed bank variability in relation to monkey and dung beetle activity, to assess the impact of dung beetles on seed fate, and their adaptability to resource availability. This question is important to better understand the process of tree regeneration, and is especially timely in the current context of threat to primates. We characterized soil seed bank structures in sites differing in monkey frequentation, and conducted field experiments with artificial beads to monitor bead fate. We also conducted experiments on specific roller and tunneller beetle species to examine bead processing behavior and its variability among and within species. We found that seed number and diversity increased with monkey frequentation, but seed viability was optimal under moderate monkey frequentation. We showed for the first time that dung provisioning yielded higher beetle activity in sites more often visited by monkeys, which calls for further investigation to understand the mechanisms of attraction to resource and potential spatial structuration of beetle populations. Although all beetle species involved in the experiments actively excluded beads from dung reserves, selectivity was higher for small than large beetle species, and for large compared to small bead sizes. It also increased when per-capita dung resource decreased, suggesting that intraspecific competition could alter seed fate. Altogether, our results support a major role of dung beetles in soil seed bank structure and dynamics. They reveal interesting interspecific variability within the dung beetle community and a complex interplay with primary dispersal.     

Dung beetle activity affects rain forest seed bank dynamics and seedling establishment

Dung beetles relocate vertebrate feces under the soil surface, and this behavior has many ecological consequences. In tropical forests, for example, seeds defecated by mammals that are subsequently buried by dung beetles are less likely to suffer predation. While the effects of dung beetles on the fate of defecated seeds have been relatively well studied, their effect on seeds already buried in the soil has not. To contribute to fill this gap, we designed a study with three objectives: (a) Describe the vertical re‐distribution of soil seeds that occurs due to dung beetle activity; (b) Determine if beetle activity favors establishment of seedlings from the soil seed bank; and (c) Determine if the effect of dung beetles is stronger in sites of recurrent mammal defecation. We carried out three complementary field experiments, one with artificial seeds (plastic beads) of three sizes buried at known depths, one with two species of seeds buried at those same depths, and one with the natural soil seed bank in sites of single vs. recurrent defecation. Buried beads were moved by dung beetles along the vertical axis, both upwards (9.5%) and downwards (11.5%); smaller beads were more frequently moved downwards while the contrary occurred for larger beads. Dung beetle activity caused an increase in seedling establishment, both from experimentally buried seeds and from the natural seed bank. Defecation recurrence had no effect on seedling establishment. We conclude that dung beetle activity affects seed bank dynamics with important consequences for seedling establishment in tropical forests.Abstract in Spanish is available with online material.     

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.     

Fire? They don't give a dung! The resilience of dung beetles to fire in a tropical savanna

1. Disturbance is a strong driver of community assembly and fire has long been recognised as one of the main disturbances of terrestrial ecosystems. This study tested the resilience of dung beetles to fire events in campos rupestres, which is a tropical savanna ecosystem that evolved under a frequent fire regime, by assessing the resistance and recovery of their communities. 2. Dung beetles were sampled before and after a fire event and the effect of fire on dung beetle richness, abundance, mean community biomass and composition was tested. The effects of time since last fire and fire frequency on the community were also tested. 3. No effect of fire occurrence, time since last fire and fire frequency on any community variable was found. 4. Some non‐mutually exclusive mechanisms promoting the resistance and recovery of dung beetles in campos rupestres could be acting in synergy. One potential mechanism is the mismatched seasonality between fire events and dung beetle occurrence, as fires occur during the dry season and dung beetles are present above ground during the rainy season. Furthermore, dung beetles are insects that remain buried during most of their lifetime, which could protect individuals from being burned. Another potential mechanism is the replacement of species in burned areas by the movement of individuals from unburned areas, attracted by resources and/or by metacommunity dynamics. 5. It is concluded that in this ‘fire‐dependent’ ecosystem, dung beetle communities are resilient to fire and seem not to be structured by this disturbance.     

Primary Seed Dispersal by Red Howler Monkeys and the Effect of Defecation Patterns on the Fate of Dispersed Seeds1

The effectiveness of a seed disperser depends on the quantity and quality of dispersal. The quality of dispersal depends in large part on factors that affect the post–dispersal fate of seeds, and yet this aspect of dispersal quality is rarely assessed. In the particular case of seed dispersal through endozoochory, the defecation pattern produced has the potential of affecting the fate of dispersed seeds and consequently, dispersal quality and effectiveness. In this study, I assessed the effects of dung presence and dung/seed densities on seed predation by rodents and secondary dispersal by dung beetles. In particular, I compared seed fates in clumped defecation patterns, as those produced by howler monkeys, with seed fates in scattered defecation patterns, as those produced by other frugivores. I also determined the prevalence of red howler monkeys (Alouatta seniculus) as seed dispersers at the plant community level in Central Amazonia by determining the number of species they dispersed in a 25–month period. I found that dung presence and amount affected rodent and dung beetle behavior. Seed predation rates were higher when dung was present, and when it was in higher densities. The same number of seeds was buried by dung beedes, in dumped versus scattered defecation patterns, but more seeds were buried when they were inside large dung–piles versus small piles. Seed density had no effect on rodent or dung beetle behavior. Results indicate that caution should be taken when categorizing an animal as a high or low quality seed disperser before carefully examining the factors that affect the fate of dispersed seeds. Red howler monkeys dispersed the seeds of 137 species during the study period, which is the highest yet reported number for an Alouatta species, and should thus be considered highly prevalent seed dispersers at the plant community level in Central Amazonian terra firme rain forests.     

Potential of yellow sticky traps for lady beetle survey in cotton

A 2-yr study was conducted to investigate the potential of using yellow sticky traps to survey lady beetles in cotton and to quantify seasonal activity patterns. The performance of sticky traps was compared with that of a 2-cycle vacuum sampler. The most common lady beetle species captured by sticky traps and vacuum sampler in cotton were Hippodamia convergens Guerin-Meneville and Scymnus loewii Mulsant. Sticky traps captured significantly more of both species of lady beetles, had greater capture efficiency, and more effectively detected lady beetles compared with the vacuum sampler. These data indicate that the sticky trap can be a valuable tool in monitoring lady beetle populations in cotton. In the second part of this study, a year-round survey of lady beetle populations in the periphery of a cotton farm using sticky traps showed that lady beetles remained active throughout the year in the Texas Rolling Plains, but the activity was influenced by winter severity. Over a 2-yr period, H. convergens, S. loewii, Coccinella septempunctata (L.), and Olla v-nigrum (Mulsant) comprised 89.6, 8.2, 1.9, and 0.3% of the specimens, respectively. Sticky trap captures were affected by year, trap height, and cropping season. Traps placed at 0.75 m above ground captured significantly more (80%) lady beetles than traps placed at 1.50 m (20%) above ground; traps at 0.75 m above ground also detected the rarer species while the traps at 1.50 m above ground detected only the abundant species. Trap captures were higher during the noncotton season (November to April) compared with the cotton season (May to October). A significant positive correlation between cotton aphid abundance during the growing season and H. convergens abundance during the following noncotton season was also detected, indicating a significant movement of H. convergens from cotton to the periphery of the farm to seek refuge after cotton termination.     

A comparison of dung beetle (Coleoptera: Scarabaeidae) attraction to native and exotic mammal dung

Although the preference of dung beetles (Coleoptera: Scarabaeidae) for specific types and conditions of dung has been given substantial attention, little has been done to investigate the potential effects of exotic mammal introduction for game farms or rewilding projects. We used pitfall traps baited with various native and exotic herbivore, carnivore, and omnivore dung to evaluate dung beetle preference in the Great Plains of North America. Additionally, we analyzed of the nutrient quality of each dung type. In total, 9,089 dung beetles from 15 species were captured in 2 yr of sampling. We found significant differences (P < 0.05) in mean dung beetle capture among omnivore, herbivore, and carnivore dung, as well as differences in individual species preference for dung type. Omnivore dung was the most attractive with chimpanzee and human dung having the highest mean capture (291.1 ± 27.6 and 287.5 ± 28.5 respectively). Carrion also was highly attractive with a mean of 231.9 ± 20.6 beetles per trap (N = 8). Our results suggest definitive local preference of carrion in Phanaeus vindex Macleay and Onthophagus hecate (Panzer), while the congener, O. pennsylvanicus (Harold), was rarely captured in carrion and highly preferred omnivore dung. Preference for a specific bait type does not appear to be correlated with dung quality, mammalian diet, or origin of mammal. Results suggest niche segregation by dung type among dung beetle species.     

Experimental warming disrupts reproduction and dung burial by a ball‐rolling dung beetle

1. Insects are sensitive to climate change. Consequently, insect‐mediated ecosystem functions and services may be altered by changing climates. 2. Dung beetles provide multiple services by burying manure. Using climate‐controlled chambers, the effects of warming on dung burial and reproduction by the dung beetle Sisyphus rubrus Paschalidis, 1974 were investigated. Sisyphus rubrus break up dung by forming and rolling away balls of manure for burial and egg deposition. 3. To simulate warming in the chambers, 0, 2 or 4 °C offsets were added to field‐recorded, diurnally fluctuating temperatures. We measured dung ball production and burial, egg laying, survival and residence times of beetles. 4. Temperature did not affect the size or number of dung balls produced; however warming reduced dung ball burial by S. rubrus. Because buried balls were more likely to contain eggs, warming could reduce egg laying via a reduction in ball burial. Warming reduced the humidity inside the chambers, and a positive relationship was found between the number of dung balls produced and humidity in two temperature treatments. Temperature did not affect survival, or whether or not a beetle left a chamber. Beetles that did leave the chambers took longer to do so in the warmest treatment. 5. This study demonstrates that climate warming could reduce reproduction and dung burial by S. rubrus, and is an important first step to understanding warming effects on burial services. Future studies should assess warming effects in field situations, both on individual dung beetle species and on aggregate dung burial services.     

Dung beetles and their ecological functions in three agroforestry systems in the Lacandona rainforest of Mexico

There is growing interest in evaluating the impact that management intensity of agroecosystems has on animal communities and their ecological functions. Dung beetles are a highly used focal taxon for assessing the effects of anthropogenic disturbances and management practices on biodiversity. In the Lacandona rainforest region in southern Mexico, we quantified several metrics of the dung beetle community (number of species, number of individuals, total biomass, mean beetle size) and four of their ecological functions (dung removal, soil excavation, seed dispersal, seed exhumation) in conserved rainforest and three agroforestry systems with different management intensities: rustic cocoa, polyculture cocoa, and rubber monoculture. We also assessed the correlation between dung removal and the other functions, as well as the relationships between functions and community metrics. Land-use type affected the dung beetle communities as well as their functions, with negative effects on response variables in the most intensely managed agroecosystems (polyculture cocoa and rubber). Rustic cocoa had values similar to those of the conserved forest for all functions and community metrics, except the mean number of species per trap. Dung removal was correlated with the other ecological functions. The mean number of species per trap was significantly associated with all four functions. In our study region rustic cocoa plantations favor the maintenance of a high proportion of dung beetle species and maintain their ecological functions. Our findings corroborate that agroecosystems with less intense management may contribute to buffering the effects of landscape homogenization caused by more intensely managed agroecosystems, such as rubber plantations.     

Patch size and colonisation patterns: an experimental analysis using north temperate coprophagous dung beetles

The relationship between dung pad size and both adult colonisation and larval development was investigated in an assemblage of north temperate dung beetles ( Geotrupes. Aphodius and Sphaeridium ) using both dung pads and baited pitfall traps. Wet weight of 22-day-old natural dung pads was found to vary widely in the field (< 100 g – >1000 g). Across all sampling dates in field experiments, dung pad size had a significant influence on dung beetle biomass sampled from pads. Closer examination of experimental dung pads on the second day after deposition, when beetle biomass was at a maximum, revealed not only a general positive relationship between pad size and dung beetle biomass but, more importantly, a positive relationship between dung pad size and dung beetle density (dung beetle biomass per unit dung volume). There was a strong trend for Aphodius species richness to increase, and maintain higher values for longer periods of time, in larger pads. Although dung pad and pitfall trap samples could differ in the actual numbers of beetles captured, the relationship between different dung sizes and dung beetle biomass was similar, indicating that the phenomenon is largely related to immigration processes. Pat residence times oi A. rufipes in the laboratory were significantly positively correlated with dung pad size. In two field experiments, positive correlations were found between dung pad size and numbers of larvae in pads of different sizes and in one of these experiments, larval densities (numbers per unit dung volume) were significantly and positively correlated with dung pad size. In one experiment, Aphodius larvae in the early stages of development were found to preferentially occupy the basal area of dung pads. We discuss the implications of our findings in the context of resource utilisation by north temperate dung beetles.     

How varying pest and trap densities affect Tribolium castaneum capture in pheromone traps

The red flour beetle, Tribolium castaneum (Herbst) (Coleoptera: Tenebrionidae), is an important insect pest in food processing facilities. Pheromone trapping is frequently used to monitor red flour beetle populations in structures; however, the optimal trap density and the relationship between trap captures and beetle density is not known. Two experiments were performed concurrently in environmentally controlled 30‐m² walk‐in chambers to determine the relationship between aggregation pheromone trap captures of red flour beetles and beetle and trap number. In one experiment, beetle density was kept constant at 200 individuals per chamber while trap number was varied from 1 to 8, and in the other experiment trap number remained constant at one per chamber while beetle density varied from 20 to 800 individuals. Results indicated that approximately one out of 23 red flour beetles were captured in a trap. Number of beetles captured in traps increased significantly as beetle density increased; however, the proportion of beetles captured remained consistent across beetle densities with a mean of 4.7 ± 0.6% of individuals captured. Trap captures varied significantly with trap placement within experimental chambers, indicating that subtle differences in the trapping environment can influence trap captures. Data suggested that trap densities of 0.07–0.10 m^?2 (2–3 traps per chamber) would maximize trap capture, whereas a trap density of 0.13 m^?2 (four traps per chamber) would maximize the predictive ability of a trapping equation estimating beetle density from trap captures. Results provide information needed to more thoroughly explore how environmental factors might influence red flour beetle trap capture in the absence of changes in beetle density. Further understanding of these relationships will allow for more accurate assessments of absolute beetle density from pheromone trap capture data.     

Invasion and retreat: shifting assemblages of dung beetles amidst changing agricultural landscapes in central Peru

This study examines the effects of deforestation, and the habitat value of coffee and regenerated forest for tropical dung beetles, a functionally significant insect group. Pitfall trapping was conducted at 22 sites in a montane region of central Peru during April and November/December of 2002. Sites included primary and secondary forest, shade-coffee, regenerated forest and open farms (mainly with banana, yuca, and corn). Ordination techniques indicated that beetle assemblages in forests, regenerated forest and coffee were relatively similar. However, assemblage compositions in forested areas differed even at similar altitudes under the influence of biogeographical factors, and the assemblages at disturbed sites (farms/coffee) were influenced by beetle dispersal from adjacent forests. During dry months, when beetle activity is low, communities at all habitat types tended to converge because fewer unique species were recorded in forests at that time and habitat/season generalists were dominant. Preliminary results also indicate that beetles in shady crops such as bananas responded to plant growth: as the banana canopy closed-in, producing more shade, open-habitat specialists retreated and forest/shade specialists invaded the sites. Chronosequence data at two of the sites demonstrate the rapid and dramatic changes in species richness and assemblage composition caused by deforestation. As forests become increasingly fragmented, and open farms continue to expand, dung beetles will become more restricted to the remaining fragments and reserves. In the mosaic landscape studied here, shade crops, like coffee, act as habitat and corridors for many dung beetle species. Small farm size and the consequent magnitude of edge effects, likely contributed to beetle movement between habitat types and determined the apparent generalist nature of many of the dung beetle species in this study.     

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.     

Response of a dung beetle assemblage along a reforestation gradient in Restinga forest

Scarabaeinae dung beetles are indicator insects used in the evaluation of the ecological effects caused by changes in habitat structure and ecosystem integrity resulting from environmental degradation. We compared dung beetle diversity in conserved restinga forests (coastal tropical moist broadleaf forest) and in reforested areas of various ages during the rainy and dry seasons, on the coast of Paraíba State, Brazil. A total of 3,634 individuals comprising 14 species were collected. In the reforested areas there was a gradual increase in species abundance relative to the area’s age, but in the conserved restinga the abundance of individuals was 10–20 times higher than that recorded in areas of recent reforestation. The highest species richness was found in the conserved restinga and in the oldest reforested area (16 years old) during the rainy season. During the dry season, when environmental conditions do not seem to favor adult survival, most of the species were found in the conserved restinga forest. The dung beetle community structure was related to the increases in habitat heterogeneity in the successional processes of the reforested areas. Our results suggest that reforested areas act as a source of and refuge for dung beetle species.