Insect biodiversity meets ecosystem function: differential effects of habitat and insects on carrion decomposition

1. Ecological processes are maintained in different environments by different species performing similar functional roles. Yet, little is known about the role of the environment in shaping insect biodiversity associated with a process that is ephemeral and patchy. 2. In this study, the mass loss of carrion in response to contrasting habitat types (grassland or tree) was quantified experimentally, as well as the presence, diversity and composition of insect assemblages. Differences in insect assemblages between these two habitats were also examined. 3. It was found that the presence of insects led to a doubling in mass loss, but that grassland or tree habitat type had no effect on this process. By contrast, habitat type had a significant effect on the composition of generalist ant and beetle assemblages, but not on specialist fly assemblages. Given the colonisation of insects, carrion mass loss was negatively associated with increasing evenness of fly assemblages and increasing ant abundance. Variation in fly assemblage composition was also found to correlate with variation in carrion mass loss. 4. This study highlights the major role of habitat type in shaping the composition of generalist insects at carrion, but the minor role in affecting specialist and highly vagile insects. This complements the authors' findings that insect colonisation of carrion was critical to accelerated mass loss, and that fly assemblages were responsible for variation in this process, regardless of habitat. The present study sheds new light on the contribution of insect biodiversity to decomposition in variable environments, with consequences for carrion food webs and nutrient cycling.     

Ecological features and distribution of the mosquito <Emphasis Type="Italic">Aedes communis</Emphasis> (De Geer, 1776) in the Northwestern European Russia

Distribution and biotopic and seasonal associations of the mosquito Aedes communis in Northwestern European Russia is discussed.     

Decomposition rate of carrion is dependent on composition not abundance of the assemblages of insect scavengers

Environmental factors and biodiversity affect ecosystem processes. As environmental change modifies also biodiversity it is unclear whether direct effects of environmental factors on ecosystem processes are more important than indirect effects mediated by changes in biodiversity. High-quality resources like carrion occur as heterogeneous pulses of energy and nutrients. Consequently, the distribution of scavenging insects is related to resource availability. Therefore, carrion decomposition represents a suitable process from which to unravel direct effects of environmental change from indirect biodiversity-related effects on ecosystem processes. During three field seasons in 2010 we exposed traps baited with small-mammal carrion at 21 sites along a temperature gradient to explore the insect carrion fauna and decomposition rate in the Bohemian Forest, Germany. The abundance component of beetle and fly assemblages decreased with decreasing temperature. Independently, the composition component of both taxa changed with temperature and season. The change in the composition component of beetles depicted a loss of larger species at higher temperatures. Decomposition rate did not change directly along the temperature gradient but was directly influenced by season. The composition component of beetles, and to a small extent of flies, but not their abundance component, directly affected carrion decomposition. Consequently, lower decomposition rates at lower temperatures can be explained by the absence of larger beetle species. Thus, we predict that future environmental change will modify carrion fauna composition and thereby indirectly decomposition rate. Moreover, reorganizations of the insect carrion composition will directly translate into modified decomposition rates, with potential consequences for nutrient availability and carbon storage.     

Gray wolves as climate change buffers in Yellowstone

Understanding the mechanisms by which climate and predation patterns by top predators co-vary to affect community structure accrues added importance as humans exert growing influence over both climate and regional predator assemblages. In Yellowstone National Park, winter conditions and reintroduced gray wolves (Canis lupus) together determine the availability of winter carrion on which numerous scavenger species depend for survival and reproduction. As climate changes in Yellowstone, therefore, scavenger species may experience a dramatic reshuffling of food resources. As such, we analyzed 55 y of weather data from Yellowstone in order to determine trends in winter conditions. We found that winters are getting shorter, as measured by the number of days with snow on the ground, due to decreased snowfall and increased number of days with temperatures above freezing. To investigate synergistic effects of human and climatic alterations of species interactions, we used an empirically derived model to show that in the absence of wolves, early snow thaw leads to a substantial reduction in late-winter carrion, causing potential food bottlenecks for scavengers. In addition, by narrowing the window of time over which carrion is available and thereby creating a resource pulse, climate change likely favors scavengers that can quickly track food sources over great distances. Wolves, however, largely mitigate late-winter reduction in carrion due to earlier snow thaws. By buffering the effects of climate change on carrion availability, wolves allow scavengers to adapt to a changing environment over a longer time scale more commensurate with natural processes. This study illustrates the importance of restoring and maintaining intact food chains in the face of large-scale environmental perturbations such as climate change.     

Area effect of distribution of Silphids (Coleoptera,Silphidae) on Korean islands

We investigated the spatial patterns in the distribution of carrion beetles on 26 islands to test the area and distance effects on species diversity. The relationships between species richness and assemblage and island area, distance from the mainland, and shortest distance from the largest island within 2?km were analyzed using linear regression, Canonical Correspondence Analysis (CCA) and Multi-Response Permutation Procedures (MRPP). We hypothesized that both area and distance from the mainland played a significant role in species richness, while geographic location in the sea influenced aggregation of carrion beetles on islands. Area was the only factor influencing species richness of carrion beetles on islands. Moreover, distance from the mainland and close islands did not significantly influence to carrion beetles. In addition, geographic location in the same Sea did not influence beetle assemblages. Taken together, the results of this study indicated that carrion beetles on Korean islands were largely affected by island size and that dispersal was stochastic.     

Periphytic algal assemblages along environmental gradients in the rivers of the Lake Ladoga basin,Northwestern Russia: implication for the water quality assessment

The purpose of this study was to examine the role of geological, hydrological, and anthropogenic factors in structuring periphytic algal assemblages in the rivers of the Lake Ladoga basin (Northwestern Russia). Canonical correspondence analysis (CCA) conducted on spring dataset showed that water temperature, color, and river morphology were primary factors shaping the structure of algal assemblages during the spring post-flood period. CCA on summer and autumn dataset revealed that conductivity and total phosphorus were the most important variables during the base flow period. Cluster analysis carried out on algal assemblages separated all rivers into two main groups that corresponded to the two geomorphological regions of the Lake Ladoga basin: the northern and the southern sub-basins. The separation of the northern and the southern groups were best explained by the pattern of conductivity reflecting geological and land use differences in the watersheds. The values of the Specific Pollution sensitivity Index (IPS) and the Biological Diatom Index (IBD) were higher, indicating better water quality, for rivers of the northern sub-basin than for rivers of the southern sub-basin. Mean values of the IPS and IBD for rivers of the northern and the southern sub-basins varied within the boundaries of a good water quality.     

Carrion mimicry in a South African orchid: flowers attract a narrow subset of the fly assemblage on animal carcasses

BACKGROUND AND AIMS: Although pollination of plants that attract flies by resembling their carrion brood and food sites has been reported in several angiosperm families, there has been very little work done on the level of specificity in carrion mimicry systems and the importance of plant cues in mediating such specialization. Specificity may be expected, as carrion-frequenting flies often exploit different niches, which has been interpreted as avoidance of interspecific competition. Interactions between the orchid Satyrium pumilum and a local assemblage of carrion flies were investigated, and the functional significance of floral traits, especially scent, tested. Pollination success and the incidence of pollinator-mediated self-pollination were measured and these were compared with values for orchids with sexual- and food-deceptive pollination systems. METHODS AND KEY RESULTS: Observations of insect visitation to animal carcasses and to flowers showed that the local assemblage of carrion flies was dominated by blow flies (Calliphoridae), house flies (Muscidae) and flesh flies (Sarcophagidae), but flowers of the orchid were pollinated exclusively by flesh flies, with a strong bias towards females that sometimes deposited live larvae on flowers. A trend towards similar partitioning of fly taxa was found in an experiment that tested the effect of large versus small carrion quantities on fly attraction. GC-MS analysis showed that floral scent is dominated by oligosulfides, 2-heptanone, p-cresol and indole, compounds that also dominate carrion scent. Flesh flies did not distinguish between floral and carrion scent in a choice experiment using olfactory cues only, which also showed that scent alone is responsible for fly attraction. Pollination success was relatively high (31·5 % of flowers), but tracking of stained pollinia also revealed that a relatively high percentage (46 %) of pollen deposited on stigmas originates from the same plant. CONCLUSIONS: Satyrium pumilum selectively attracts flesh flies, probably because its relatively weak scent resembles that of the small carrion on which these flies predominate. In this way, the plants exploit a specific subset of the insect assemblage associated with carrion. Pollination rates and levels of self-pollination were high compared with those in other deceptive orchids and it is therefore unlikely that this mimicry system evolved to promote outcrossing.     

Targeted detection of mammalian species using carrion fly‐derived DNA

DNA analysis from carrion flies (iDNA analysis) has recently been promoted as a powerful tool for cost‐ and time‐efficient monitoring of wildlife. While originally applied to identify any mammalian species present in an area, it should also allow for targeted detection of species and individuals. Using carrion flies captured in the Taï National Park, Côte d'Ivoire, we assessed this possibility by (i) screening carrion fly DNA extracts with nonspecific and species‐specific PCR systems, respectively, targeting mitochondrial DNA (mtDNA) fragments of any mammal or of Jentink's duiker (Cephalophus jentinki), three colobine monkeys (subfamily Colobinae) and sooty mangabey (Cercocebus atys); and (ii) genotyping carrion fly extracts containing sooty mangabey mtDNA. In comparison with the nonspecific PCR assay, the use of specific PCRs increased the frequency of detection of target species up to threefold. Detection rates partially reflected relative abundances of target species in the area. Amplification of seven microsatellite loci from carrion flies positive for sooty mangabey mtDNA yielded an average PCR success of 46%, showing that the identification of individuals is, to some extent, possible. Regression analysis of microsatellite PCR success and mtDNA concentration revealed that, among all carrion flies analysed for this study, 1% contained amounts of mammal mtDNA sufficient to attempt genotyping with potentially high success. We conclude that carrion fly‐derived DNA analysis represents a promising tool for targeted monitoring of mammals in their natural habitat.     

The nested structure of a scavenger community

Scavenging is a widespread phenomenon in vertebrate communities which has rarely been accounted for, in spite of playing an essential role in food webs by enhancing nutrient recycling and community stability. Most studies on scavenger assemblages have often presented an oversimplified view of carrion foraging. Here, we applied for the first time the concept of nestedness to the study of a species-rich scavenger community in a forest ecosystem (Białowieża Primeval Forest, Poland) following a network approach. By analysing one of the most complete datasets existing up to now in a pristine environment, we have shown that the community of facultative scavengers is not randomly assembled but highly nested. A nested pattern means that species-poor carcasses support a subset of the scavenger assemblage occurring at progressively species-rich carcasses. This result contradicts the conventional view of facultative scavenging as random and opportunistic and supports recent findings in scavenging ecology. It also suggests that factors other than competition play a major role in determining community structure. Nested patterns in scavenger communities appear to be promoted by the high diversity in carrion resources and consumers, the differential predictability of the ungulate carcass types and stressful environmental conditions.     

Are land bird assemblages functionally saturated? An empirical test in Mexico

According to the model of ideal free distribution, few or no assemblages will contain unused resources as long as species are free to colonize areas where resources are available. Consequently, because several food resources for birds are available throughout Mexico (nectar, seeds and fruits, vertebrates, carrion and different types of invertebrates), we predicted that all assemblages from Mexico contain birds from the corresponding foraging guilds. However, severe climates may prevent colonization by certain birds. Thus, a second prediction was that, if assemblages with unused resources were found, these would be from localities with climates that pose extreme stress (either extremely high temperature and low rainfall, or extremely low temperature). We tested these predictions on bird assemblages from 77 localities that represent the entire range of climatic conditions found in Mexico. Only two assemblages containing unused resources were found. These were from the localities with lowest temperature and lowest rainfall, respectively. We observed that land bird assemblages in Mexico do not contain unused resources, except when climate selectively restricts colonization by species from certain foraging guilds. Therefore, we conclude that even assemblages with almost no species in common have a common guild structure based on food resources. Null models of competition would be more realistic if they were to incorporate such patterns in their regional species pools.     

Carrion fly‐derived DNA metabarcoding is an effective tool for mammal surveys: Evidence from a known tropical mammal community

Metabarcoding of vertebrate DNA derived from carrion flies has been proposed as a promising tool for biodiversity monitoring. To evaluate its efficacy, we conducted metabarcoding surveys of carrion flies on Barro Colorado Island (BCI), Panama, which has a well‐known mammal community, and compared our results against diurnal transect counts and camera trapping. We collected 1,084 flies in 29 sampling days, conducted metabarcoding with mammal‐specific (16S) and vertebrate‐specific (12S) primers, and sequenced amplicons on Illumina MiSeq. For taxonomic assignment, we compared blast with the new program protax , and we found that protax improved species identifications. We detected 20 mammal, four bird, and one lizard species from carrion fly metabarcoding, all but one of which are known from BCI. Fly metabarcoding detected more mammal species than concurrent transect counts (29 sampling days, 13 species) and concurrent camera trapping (84 sampling days, 17 species), and detected 67% of the number of mammal species documented by 8 years of transect counts and camera trapping combined, although fly metabarcoding missed several abundant species. This study demonstrates that carrion fly metabarcoding is a powerful tool for mammal biodiversity surveys and has the potential to detect a broader range of species than more commonly used methods.     

Trophic relationships and their importance for biotopic distribution of grasshoppers (Orthoptera,Acridoidea) in semi-deserts and deserts of the lower Volga river area

The trophic associations of grasshoppers in the semi-deserts and deserts of the lower Volga area are analyzed. Most of the species studied are narrow oligophages or monophages; they constitute four groups trophically associated with edificators of the commonest desert plant communities (hygro- and mesophytic grasses, xerophytic grasses, wormwoods, and chenopods). The trophic specialization is shown to contribute significantly to the formation of biotopic association and structure of grasshopper assemblages.     

A continental‐scale analysis of fish assemblage functional structure in European rivers

The theory of traits (life‐history, ecological and biological traits) states that a species’ characteristics might enable its persistence and development in given environmental conditions. If environment is the major factor controlling functional assemblage structure, species with similar attributes are expected to inhabit a similar environment. This study uses trait states in 849 European riverine fish assemblages to analyze the influence of environment, phylogeny and biogeography on the functional structure of these assemblages. European fish assemblages were highly structured and two main syndromes (a suite of coevolved traits) were observed: 1) assemblages dominated by stenothermal intolerant individuals and 2) assemblages dominated by eurythermal, eurytopic and tolerant individuals. Temperature and stream physical structure were the two main environmental factors explaining the diversity of fish assemblage functional structures, while the influence of biogeographic factors was weak, once environment was taken into account. This suggests that, whatever the regional species pool, similar assemblage functional structures will be found in similar environmental conditions. The phylogenetic relatedness between species might also explain to some extent the associations between the species traits observed among European fish assemblages.     

Community structure of carrion flies along an island gradient

species richness, abundance relations, exploitation efficiencies, and population dynamics were examined within four carrion fly communities from a small isolated island in the Baltic to the large island of Åland. A total of 23 species and 11408 Specimens were collected.
Species richness (studied by rarefaction) of potential carrion flies was lowest on the small isolated island, but the smallest number of species emerged from the second largest island. Lucilia illustris Meig, was the most numerous species at every site, comprising 46–56% of the potential breeders and 71–91% of the emerging flies. Exploitation efficiency, measured by the numbers of individuals emerging per gram of carrion, was highest on the smallest island, but faunal similarity and ordination analyses suggest that competition was more intense on the larger islands.     

Contrasting diversity dynamics of phoretic mites and beetles associated with vertebrate carrion

Carrion is an ephemeral and nutrient-rich resource that attracts a diverse array of arthropods as it decomposes. Carrion-associated mites often disperse between animal carcasses using phoresy, the transport of one species by another. Yet few studies have contrasted the dynamics of mite assemblages with other insect taxa present at carrion. We examined and compared the changes in abundance, species richness and composition of mite and beetle assemblages sampled at kangaroo carcasses in a grassy eucalypt woodland at four different times over a 6-month period. We found that the majority of mites were phoretic, with the mesostigmatid genera Uroseius (Uropodidae), Macrocheles (Macrochelidae) and Parasitus (Parasitidae) the most abundant taxa (excluding astigmatid mites). Abundance and richness patterns of mites and beetles were very different, with mites reaching peak abundance and richness at weeks 6 and 12, and beetles at weeks 1 and 6. Both mites and beetles showed clear successional patterns via changes in species presence and relative abundance. Our study shows that mesostigmatid mite assemblages have a delay in peak abundance and richness relative to beetle assemblages. This suggests that differences in dispersal and reproductive traits of arthropods may contribute to the contrasting diversity dynamics of carrion arthropod communities, and further highlights the role of carrion as a driver of diversity and heterogeneity in ecosystems.     

Dung beetle assemblages in forests and pastures of El Salvador: a functional comparison

This study examines the functional attributes of Neotropical dung beetles, and, based on the analysis, evaluates the role of dung beetles in pastures and tropical dry forest in El Salvador. Dung beetle diversity was lower in pastures than in forests. However, the total biomass of beetles at similarly-sized dung baits was frequently higher in pastures. Diversity loss followed structured patterns: (1) carrion and fruit feeding beetles were well represented (species number) in the forests but were largely absent from pastures; (2) large ball rollers and small fast tunnellers were present in forests but were absent from pastures; (3) large fast tunnellers and small slow tunnellers were poorly represented in pastures compared to forests but the few species that survived in pastures attained extremely high populations; and (4) each functional group in the species-poor pasture assemblages was divided almost equally into day and night active species. Substrate and habitat generalists that were present in both the forests and pastures were attracted to fruit and carrion in forested habitat but not in open pastures. In open habitats, flies and other insects may be the principal decomposers of decaying fruit and carrion.     

The structure of carrion fly communities: the size and the type of carrion

Species composition and abundance relations of flies attracted to and bred from different sizes (20-100 g) and types of carrion (small mammal, cow liver, bird, and fish) were studied experimentally in natural breeding conditions in South Finland. Large pieces of carrion did not attract significantly more individuals or species than did small ones, but there was a positive correlation between size of carrion and numbers of flies bred, apparently because of competition. Altogether 10 species emerged from 50 cultures, and on average 2.78 ± 1.22 (SD) species emerged from one culture. The number of co-occurrences among the emerging flies indicated random colonization. Variation in species composition between like pieces of carrion was not less than variation between carcasses of different size or type; hence, size and type of carrion are not important determinants of community structure within the size and type ranges used.
Lucilia illustris (Meig.) was the predominant species, comprising 50% of flies trapped and 85% of flies bred. The difference indicates changing species composition in the community. Two sets of cultures were left open for egg-laying for 2 and 7 d. Competition was less Intensive in the former cultures, which produced 4.0 individuals per gram of carrion, contrasting with only 1.4 individuals per gram in the latter cultures. Competition delayed development in all species, but least in L. illustris , which suggests that fast developing species are at a competitive advantage. In agreement with this, abundance was positively correlated with the rate of development.     

Comparing scavenging in marine and terrestrial ecosystems: a case study with fish and gull carcasses in a small Mediterranean island

Carrion consumption by scavengers is a key component of both terrestrial and aquatic food webs. However, there are few direct comparisons of the structure and functioning of scavenging communities in different ecosystems. Here, we monitored the consumption of 23 fish (seabream Sparus aurata) and 34 bird (yellow-legged gull Larus michahellis) carcasses on a small Mediterranean island (Isla Grosa, southeastern Spain) and surrounding waters in summer to compare the structure of the scavenger assemblages and their carrion consumption efficiencies in terrestrial and shallow water habitats. Scavenging was highly efficient both in marine and terrestrial environments, especially in the presence of a highly abundant vertebrate scavenger species, the yellow-legged gull. The vertebrate scavenger community was richer in the marine environment, whereas the invertebrate community was richer on land. The scavenger network was usually well-structured (i.e., nested), with the exception of the community associated with fish terrestrial carcasses, which were almost monopolized by yellow-legged gulls. In contrast, gulls left conspecific carcasses untouched, thus allowing longer persistence of gull carcasses on land and their exploitation by a diverse insect community. Our study shows important differences in the scavenging process associated with environment and carcass type. Promising avenues for further eco-evolutionary and applied research arise from the comparison of scavenging processes in terrestrial and marine ecosystems, from small islands to continents.     

Carcass size shapes the structure and functioning of an African scavenging assemblage

The particle size of the food resource strongly determines the structure and dynamics of food webs. However, the ecological implications of carcass size variation for scavenging networks structure and functioning have been largely overlooked. Here we investigate differences in scavenging patterns due to carcass size in a complex vertebrate scavenger community, Hluhluwe‐iMfolozi Park, South Africa, while taking into account seasonality. We monitored the consumption of three types of experimental carcasses: ‘small’ (< 10 kg), ‘medium’ (10–100 kg) and ‘large’ (> 100 kg). We employed general lineal models to explore the influence of carcass size on 1) scavenging network structure (scavenger species richness per carcass) and 2) functioning (carcass detection time, consumption time, consumption rate and percentage of carrion consumed). We also tested whether the structure of the scavenging network of each carcass size was nested, i.e. whether the scavenging assemblage in species‐poor carcasses was a subset of the assemblage consuming species‐rich carcasses. We found strong evidence indicating that carcass size is a major factor governing the associated scavenger assemblage. Scavenger species richness per carcass and carcass consumption time and rate increased with carcass size, while carcass detection time and percentage of carrion biomass consumed were negatively related to carcass size. Strikingly, most of the carrion biomass was consumed by facultative scavengers, represented by large mammalian carnivores, rather than by obligate scavengers (i.e. vultures). Scavenging network nestedness tended to be higher at larger carcasses, and nestedness was sensitive to the removal of the most connected species in the network (spotted hyena) rather than vultures. When comparing scavenging and predation assemblages, crucial size‐dependent differences emerge. Also, we identified a traditionally ignored mechanism by which hunting large prey could be relatively less profitable for predators, namely the costs associated with competition from scavengers and decomposers.     

Chemical composition of polysaccharides of the red alga <Emphasis Type="Italic">Tichocarpus crinitus</Emphasis> (Tichocarpaseae) from different sites of Peter the Great Bay,Sea of Japan

A comparative analysis of the quantitative and qualitative composition of polysaccharides in the red seaweed Tichocarpus crinitus, collected at different sites of Peter the Great Bay in the Japanese Sea (Sivuch’ya Bay, Risovaya Bay and Sobol’ Bay) was carried out. The sites differed in their biotopic and anthropogenic characteristics. The main monosaccharides of the investigated algal polysaccharides were galactose and 3.6-anhydrogalactose, which are structural components of carrageenan The highest contents of protein and glucose were found in the KCl-soluble fraction of algae from Sobol Bay that were experiencing increased anthropogenic pressure. Fourier transform infrared spectroscopy revealed that the KCl-insoluble polysaccharide fractions of all investigated algae had an identical structure corresponding to κ/β-carrageenan, while the KCl-soluble polysaccharide fractions had differences in their structures. The content of heavy metals and micro- and macroelements in polysaccharide fractions differed among algae growing in areas with different biotopic and anthropogenic conditions.