Effects of carrion decomposition on litter arthropod assemblages

1. Many organisms contribute to the decomposition of carrion. For arthropods, many studies focus on the necrophagous community, those directly consuming carrion.
2. Necrophagous arthropods use carrion as a shelter or food source. Therefore, carrion generally increases the abundance and biodiversity of necrophagous species. However, it is unclear if carrion has similar effects on detrital communities.
3. This study examines changes in community structure and composition of necrophilous and detrital communities over the course of decomposition.
4. Five pig head carrion were placed at least 7 m apart under cages in temperate mixed forest. Leaf litter was sampled 0 m, 1.5 m, and 3 m from each carrion weekly during summer, and monthly during autumn, until the first frost. Arthropods were extracted from leaf litter by using Berlese funnels.
5. At the carrion site, necrophagous insects increased in abundance, species richness, and Shannon diversity during decomposition, and all decreased after dry decay.
6. Detritus arthropods displayed a sharp increase in abundance during advanced decay, decreasing during dry decay, and a general increase over time for species richness and diversity.
7. In conclusion, carrion can influence the surrounding, non-necrophagous arthropod community, highlighting the need to investigate carrion effects beyond typical necrophagous species to have a more holistic understanding of carrion ecology.

     

Delaying insect access alters community composition on small carrion: a quantitative approach

We present a study of interactions in the highly competitive insect communities inhabiting the carrion of small mammals. Via manipulation in a fully quantitative design, we delayed community development by excluding insect colonization in mouse and rat carcasses for 3 days, to study the role of early competitively dominant colonizers [burying beetles (Nicrophorus spp.; Coleoptera: Silphidae) and blowfly larvae (Diptera: Calliphoridae)] in the course of heterotrophic succession on small cadavers. Earlier studies demonstrated that in the case of large mammalian carrion, exclusion of insects’ access to the carcass in the early stages of decomposition altered the successional trajectory and species assemblages. However, the effect of such manipulation in easy monopolizable small vertebrate carrion remained unknown. Our results demonstrate that delaying insect access to carrion significantly lowered blowfly larvae abundances, while it simultaneously had no effect on colonization and carrion burial by burying beetles. Higher abundances of blowfly larvae seem to deter necrophagous beetles, whereas they are not harmful to the larvae of flesh flies, at least in larger rat carcasses. Predatory beetle species preferred the lower abundances of blowfly larvae, presumably due to better accessibility of their prey. Our results therefore suggest that in the course of the entire season, larvae of blowflies are the dominant competitors in small carcasses, and significantly affect the assembly of other insect groups, whereas burying beetles may exhibit a more temporal pattern of dominance.     

Traits reveal ecological strategies driving carrion insect community assembly

1. The succession of carrion-associated (necrophilous) insects on decomposing carrion is well documented. To exploit the changing nutritious and dynamic resources available throughout the carrion decomposition process, different species colonise and consume carrion in a predictable temporal sequence. The traits of these necrophilous insects should reflect their ecological strategies. Morphological traits of these insects, such as body size and wing size, however, have not previously been examined during active and advanced decomposition. 2. We used fourth-corner multivariate generalised linear models to identify insect community morphological trait patterns and to quantify their change through time on decomposing rabbit carcasses in grassland and woodland environments. 3. We found that larger-bodied species of flies and carrion-specialist beetles were associated with the early stages of decomposition. The morphological traits of ants, in contrast, showed no changes at carcasses through time and instead showed body size differences between grassland and woodland environments. 4. Our findings indicate that specialist flies and beetles that arrive early in the decomposition process possess traits that enable rapid discovery of carrion at a large scale. Generalist beetles and ants do not share this same trait and are instead adapted to locate and consume a wider variety of resources in their preferred habitat type at their local scale. 5. Our results provide insights into the morphological adaptations linked to the ecological strategies of distinct components of carrion insect communities. Further, our results offer insights into the community assembly dynamics that structure the communities of necrophilous insect species.     

Decomposition and dipteran succession in pig carrion in central Argentina: ecological aspects and their importance in forensic science

Data on the insect species associated with corpse decomposition are particularly important for estimation of the post-mortem interval (PMI) in forensic science because the PMI is based on the lifecycle and behaviour of necrophagous insects, among other measures. To determine the dipteran succession on pig carrion, four experiments, one in each season, were carried out during 2004 in a rural area of Córdoba, central Argentina. Two pigs ( Sus scrofa L.) were used in each of the four experiments. At each time-point one pig was placed in the shade and the other under direct sunlight. Insects were collected daily during the first 4 weeks and thereafter every 2 or 3 days. Five stages of decomposition were observed and a total of 24 710 adult specimens were collected, belonging to the following eight families of Diptera: Calliphoridae; Muscidae; Sarcophagidae; Phoridae; Piophilidae; Fanniidae; Sphaeroceridae, and Anthomyiidae. All Calliphoridae collected in this study were considered to be of potential forensic importance because of their necrophagous behaviour and because their immature stages use carrion as a food source. Other species, such as Musca domestica L. and Ophyra aenescens (Wiedemann), were also considered to represent potential forensic indicators.     

Chemical mimicry of insect oviposition sites: a global analysis of convergence in angiosperms

Floral mimicry of decaying plant or animal material has evolved in many plant lineages and exploits, for the purpose of pollination, insects seeking oviposition sites. Existing studies suggest that volatile signals play a particularly important role in these mimicry systems. Here, we present the first large‐scale phylogenetically informed study of patterns of evolution in the volatile emissions of plants that mimic insect oviposition sites. Multivariate analyses showed strong convergent evolution, represented by distinct clusters in chemical phenotype space of plants that mimic animal carrion, decaying plant material, herbivore dung and omnivore/carnivore faeces respectively. These plants deploy universal infochemicals that serve as indicators for the main nutrients utilised by saprophagous, coprophagous and necrophagous insects. The emission of oligosulphide‐dominated volatile blends very similar to those emitted by carrion has evolved independently in at least five plant families (Annonaceae, Apocynaceae, Araceae, Orchidaceae and Rafflesiaceae) and characterises plants associated mainly with pollination by necrophagous flies and beetles.     

Matrix models for quantifying competitive intransitivity from species abundance data

In a network of competing species, a competitive intransitivity occurs when the ranking of competitive abilities does not follow a linear hierarchy (A > B > C but C > A). A variety of mathematical models suggests that intransitive networks can prevent or slow down competitive exclusion and maintain biodiversity by enhancing species coexistence. However, it has been difficult to assess empirically the relative importance of intransitive competition because a large number of pairwise species competition experiments are needed to construct a competition matrix that is used to parameterize existing models. Here we introduce a statistical framework for evaluating the contribution of intransitivity to community structure using species abundance matrices that are commonly generated from replicated sampling of species assemblages. We provide metrics and analytical methods for using abundance matrices to estimate species competition and patch transition matrices by using reverse‐engineering and a colonization–competition model. These matrices provide complementary metrics to estimate the degree of intransitivity in the competition network of the sampled communities. Benchmark tests reveal that the proposed methods could successfully detect intransitive competition networks, even in the absence of direct measures of pairwise competitive strength. To illustrate the approach, we analyzed patterns of abundance and biomass of five species of necrophagous Diptera and eight species of their hymenopteran parasitoids that co‐occur in beech forests in Germany. We found evidence for a strong competitive hierarchy within communities of flies and parasitoids. However, for parasitoids, there was a tendency towards increasing intransitivity in higher weight classes, which represented larger resource patches. These tests provide novel methods for empirically estimating the degree of intransitivity in competitive networks from observational datasets. They can be applied to experimental measures of pairwise species interactions, as well as to spatio‐temporal samples of assemblages in homogenous environments or environmental gradients.     

Oviposition preferences and antennal size in carrion flies

Carrion-feeding flies use odours emanating from the decomposing corpse as cues for oviposition and are described as generalists because the larvae feed on the corpses of diverse species. Whereas several features of the corpse may influence the oviposition choices of these flies, it is not known whether there is a preference for a particular species of corpse. We provided carrion flies with ovipositional (and feeding) choices in a field experiment, in which various odour sources were presented simultaneously. We found novel evidence of broadly consistent choices of carrion by flies from four families. Traps baited with decaying fish flesh captured the greatest number of individuals, whereas traps baited with decaying pig liver typically attracted the least. We also asked whether individuals captured in the various baits vary in antennal size, perhaps reflecting different capacities for odour detection. There was a trend for individuals of Lucilia sericata Meigen (Diptera: Calliphoridae) and the platystomatid collected from the traps baited with pig liver to have significantly larger antennae, whereas individuals of Muscina stabulans (Fallen) (Diptera: Muscidae) captured in traps baited with marine fish flesh had relatively longer antennae for their body size. Our data reveal a more nuanced pattern of oviposition behaviour in these generalist carrion flies, which may reflect differences in their preference of carrion with different nutrients, and in their capacity to detect particular odours.     

Seasonal composition and temporal succession of necrophagous and predator beetles on pig carrion in central Argentina

Insects are the most important components of the terrestrial fauna associated with carrion because they recycle organic matter back into the ecosystem. They can be classified into four ecological categories comprising: necrophages; parasites and predators of necrophagous species; omnivores, and incidentals. To determine the composition and temporal succession of necrophagous and predator beetles on pig carrion, four experiments, one in each season, were carried out during 2004 in a rural area of Cordoba, central Argentina. Two pigs (Sus scrofa L.), weighing approximately 8 kg each, were used in each of the four experiments. The animals were killed by a sharp blow to the head and immediately placed in an appropriate trap. One pig was placed in the shade and the other in direct sunlight. Beetle fauna were collected daily during the first 4 weeks and thereafter every 2 or 3 days. Five stages of decomposition were observed and a total of 1586 adults and 4309 immatures of Coleoptera belonging to the Staphylinidae, Nitidulidae, Cleridae, Dermestidae, Histeridae, Anthicidae and Trogidae families were collected during the four experiments. The necrophagous community was represented by Dermestes maculates (De Geer), nitidulid species and members of the Trox genus. Staphylinidae, Cleridae and Histeridae species were considered to be the main predators of the necrophagous species.     

A new dipteran forensic indicator in buried bodies

The first breeding records of miltogrammine fleshflies in buried vertebrate carrion are presented. First instars of Eumacronychia persolla Reinhard (Nearctic) (Diptera: Sarcophagidae) and Phylloteles pictipennis Loew (Palaearctic) (Diptera: Sarcophagidae) are able to penetrate dry, loose soil and reach deeply buried animal remains, an ability which distinguishes is unique to necrophagous Calyptratae. Their broad geographical distribution, fast location and colonization of carrion, complete development on buried food resources and easy identification make these species useful forensic indicators in buried bodies in dry habitats.     

Saproflorivory: A Diverse Insect Community in Fallen Flowers of Lecythidaceae in French Guiana

We investigated insects associated with flowerfalls from seven tree species belonging to the Brazil nut family (Lecythidaceae) in the lowland rain forest of French Guiana. Freshly fallen androecia were collected from the forest floor and incubated in rearing chambers. Over 1300 insects representing 21 families were collected from the caged flowers, including abundant Diptera and Coleoptera, five moth morphospecies and four species of lycaenid butterflies. These insects feed on androecia. We also documented the presence of predators and parasitoids, indicating an additional trophic level.     

Carrion flies of forensic interest: a study of seasonal community composition and succession in Lisbon,Portugal

Information on Diptera community, seasonality and successional patterns in every geographical region is fundamental for the use of flies as forensic indicators of time of death. In order to obtain these data from the Lisbon area (Portugal), experiments were conducted during the four seasons of the year, using piglet carcasses as animal models. Five stages were recognized during the decomposition process. The stages, besides visually defined, could be separated taking into account the occurrence and abundance of the specific groups of Diptera collected. In general, the bloated stage recorded higher abundance and species‐richness values. Seventy‐one species were identified, belonging to 39 families, in a total of 20 144 adult Diptera collected. Autumn yielded the highest values of species richness, whereas winter had the lowest. In all seasons of the year, Calliphoridae was the dominant family; Muscidae and Fanniidae were very abundant as well. Calliphora vicina Robineau‐Desvoidy, Calliphora vomitoria (Linnaeus), Chrysomya albiceps (Wiedemann), Lucilia ampullacea Villeneuve, Lucilia caesar (Linnaeus), Lucilia sericata (Meigen) (Calliphoridae), Hydrotaea ignava (Harris), Muscina prolapsa (Harris), Synthesiomyia nudiseta (van der Wulp) (Muscidae), Piophila megastigmata McAlpine, Stearibia nigriceps (Meigen) (Piophilidae) and Nemopoda nitidula (Fallén) (Sepsidae) were revealed to be very important members of the Diptera community collected. The necrophagous behaviour, demonstrated by their immatures, using carrion as a food source makes them useful forensic indicator species. Also of relevance is the presence of Chrysomya megacephala (Fabricius), S. nudiseta and P. megastigmata, foreign species established in the local carrion communities. This study also marks the first record of S. nudiseta in Portugal.     

Pest management through tropical tree conservation

Tropical trees can provide various ecological services to adjacent agricultural environments, including maintaining and amplifying the numbers of beneficial insects. In Mexico, certain tree species harbor a diverse guild of hymenopteran parasitoids that attack pest fruit flies (Diptera: Tephritidae) and are at the same time sources of valuable hardwood timber. Indigenous trees and their associated fauna are slowly disappearing due to forest clearance and the expansion of crop monocultures. Here we explore the relationship among pest and non-pest fruit flies, their fruit-hosts and parasitoids in the context of mango orchards and surrounding patches of uncultivated vegetation and propose a novel mechanism to use these associations in favor of conservation purposes and pest management. Trees of conservation biological control interest are classified as: (1) parasitoid multiplier plants, species that serve as alternate hosts for key fruit fly pests when their commercial hosts are not available, but in which they are unusually vulnerable to parasitism; (2) parasitoid reservoir plants, native or introduced trees in whose fruits non-pest fruit flies serve as hosts to generalist parasitoids that are able to attack pest tephritids in other species of commercially grown fruit; and (3) pest-based parasitoid reservoir plants, native or introduced species that are not economically important locally, but which harbor fruit flies that would be pests in other circumstances and that serve as hosts for parasitoids of the important pests in the vicinity. Protection, multiplication and dissemination of such tree species has the potential to increase the number of naturally produced fruit fly parasitoids and could assist in the management of tephritid pests in areas where destruction of forests has impoverished the historical sources of fruit fly natural enemies. Tropical forest conservation may help resource-poor farmers reduce crop losses, increase biodiversity within fruit-growing regions and conserve native forests for both conservation purposes and commercial use of native hardwoods.     

Histories of host shifts and cospeciation among free‐living parasitoids of Rhagoletis flies

Host shifts by specialist insects can lead to reproductive isolation between insect populations that use different hosts, promoting diversification. When both a phytophagous insect and its ancestrally associated parasitoid shift to the same novel host plant, they may cospeciate. However, because adult parasitoids are free living, they can also colonize novel host insects and diversify independent of their ancestral host insect. Although shifts of parasitoids to new insect hosts have been documented in ecological time, the long‐term importance of such shifts to parasitoid diversity has not been evaluated. We used a genus of flies with a history of speciation via host shifting (Rhagoletis [Diptera: Tephritidae]) and three associated hymenopteran parasitoid genera (Diachasma, Coptera and Utetes) to examine cophylogenetic relationships between parasitoids and their host insects. We inferred phylogenies of Rhagoletis, Diachasma, Coptera and Utetes and used distance‐based cophylogenetic methods (ParaFit and PACo) to assess congruence between fly and parasitoid trees. We used an event‐based method with a free‐living parasitoid cost model to reconstruct cophylogenetic histories of each parasitoid genus and Rhagoletis. We found that the current species diversity and host–parasitoid associations between the Rhagoletis flies and parasitoids are the primary result of ancient cospeciation events. Parasitoid shifts to ancestrally unrelated hosts primarily occur near the branch tips, suggesting that host shifts contribute to recent parasitoid species diversity but that these lineages may not persist over longer time periods. Our analyses also stress the importance of biologically informed cost models when investigating the coevolutionary histories of hosts and free‐living parasitoids.     

麦田昆虫群落的结构与时序动态

采用目测法和网捕法 ,通过对小麦整个生长季节内昆虫群落的系统调查 ,共查得昆虫11目 2 3科 36种 ,其中捕食性昆虫有 4目 5科 12种、寄生性昆虫有 2目 3科 4种、植食性昆虫有 5目 12科 15种、粪食性等其他类昆虫有 1目 3科 5种。在捕食性天敌昆虫中 ,瓢虫的种类和数量最多 ,其次为草蛉和食蚜蝇 ;在寄生性天敌昆虫中 ,蚜茧蜂为优势种 ,其次为寄蝇 ;在植食性昆虫中 ,麦长管蚜为绝对的优势种 ,其次为赤须盲蝽、灰飞虱和麦蓟马。天敌昆虫对害虫具有明显的跟随现象 ,不能有效地控制麦长管蚜的发生与危害。依据昆虫群落的组成与结构 ,结合小麦生育期 ,采用最优分割法将麦田昆虫群落动态划分为 5个阶段 ,并指出不同阶段昆虫群落特点及相应的害虫防治策略。     

Structure of a herbivore–parasitoid community: Are parasitoids shared by different herbivore guilds?

Plant–herbivore–parasitoid interactions are a common occurrence in terrestrial food webs. Few parasitoids are thought to be shared by host insects of different feeding guilds because different parasitism strategies are required to use hosts of different feeding types. However, this assumption has rarely been tested using data from nature. To clarify whether parasitoids are shared among host guilds, I examined the structure of parasitoid communities on herbivore guilds associated with two Rhododendron species (Ericaceae) in a temperate secondary forest in central Japan. Leaf- and flower-feeding insects were collected from Rhododendron reticulatum and Rhododendron macrosepalum shrubs and reared in the laboratory for 3 years from April 1999 to March 2002. In total, 79 species of holometabolous herbivores (Lepidoptera, Diptera, Coleoptera, and Hymenoptera) were recorded, with 62 species on R. reticulatum and 51 species on R. macrosepalum. A total of 81 parasitoid species (Hymenoptera and Diptera) was recorded from the sampled herbivores, with 48 species from those on R. reticulatum and 50 species from those on R. macrosepalum. In total, 36 herbivore species were parasitised by 1–18 parasitoid species per host species, although the number of parasitoid species was strongly affected by sample size. Parasitoids that had two or more host species frequently attacked herbivore species from different families or on different host plants, whereas they did not attack species from different herbivore guilds; no parasitoids were shared between external feeders and rollers. Therefore, my results support the hypothesis that few parasitoids are shared among herbivores of different feeding guilds.     

Aphid suppression by natural enemies in mulched cereals

Large populations of natural enemies are the basis for natural pest control. Effects of mulch on predator–prey interactions in arable fields are poorly known, despite its potential to enhance ground‐dwelling predators and thereby reduce pest infestations. We studied the densities of predators and parasitoids, and their impact on cereal aphids in the presence and absence of mulch. Released populations of the bird cherry aphid, Rhopalosiphum padi (L.) (Homoptera: Aphididae), and two naturally occurring aphid species, were monitored under experimentally reduced densities of: (i) ground‐dwelling predators, (ii) flying predators and parasitoids, and (iii) with straw mulch. The three treatments were applied in a 2 × 2 × 2 factorial design in a field of spring wheat (Triticum aestivum L.). The exclusion of ground‐dwelling predators increased aphid populations by 55% in June and 40% in July, respectively. Mulched plots had 25% lower aphid densities in June. This was presumably due to enhanced densities of spiders (Araneida) in mulched plots. The exclusion of flying predators and parasitoids led to 94% higher aphid populations in late July (109 vs. 56 individuals per 100 shoots), irrespective of mulch or ground predator manipulation. This was attributed to the larvae of gall midges Aphidoletes cf. aphidimyza (Rondani) (Diptera: Cecidomyiidae) and hoverflies (Diptera: Syrphidae). The results indicate that a scarcity of predators and a bare soil surface renders crops more susceptible to arthropod pests. Farming schemes should aim at enhancing both ground‐dwelling and flying predators for elevated levels of natural pest control.     

Nesting strategies and disease risk in necrophagous beetles

While the effects of carcass decomposition on microorganisms have been demonstrated in recent years, little is known of how this impacts necrophagous insects. A common assumption is that insects that exploit carcasses are exposed to a high density of potentially harmful microorganisms, but no field data have so far validated this. Necrophagous beetles such as the Scarabaeinae have complex nesting behaviors with elaborate parental care. So here, we begin to explore whether this conjunction of life history and nesting behavior represents an adaptive response to the threat posed by microbes in these environments, mainly by entomopathogens. We evaluated the density and distribution of fungi and bacteria from soil near the carcasses, and their ability to infect and kill insects that are in contact with this soil during the decomposition process. Our data showed an increase in the density and activity of opportunistic or facultative pathogens during the apex of decomposition, when there is a predominance of necrophagous insects. Meanwhile, the survivorship of bait insects decreased when in contact with soil from this period of decomposition, indicating a potential risk of infection. However, the density and activity of these microorganisms decreased with distance from the carcass, mainly with depth, which would benefit tunneller beetles in particular. We have thus provided the first field data to show that necrophagous insects are indeed exposed to high densities of potentially harmful microorganisms. Furthermore, we propose that some parental care strategies may have arisen not only as a response to competition, but also as adaptations that reduce the risks of disease. Although we have focused on carrion feeders, we suggest that the same occurs with coprophagous beetles, as both carrion and dung are nutrient‐rich resources.     

Susceptibility of house flies (Diptera: Muscidae) and five pupal parasitoids (Hymenoptera: Pteromalidae) to abamectin and seven commercial insecticides.

Assays of five commercial insecticides applied as residual sprays at label rates to plywood indicated the most toxic insecticide overall for pteromalid parasitoids of house flies, Musca domestica L., was Atroban (permethrin), followed by Ciodrin (crotoxyphos), Rabon (tetrachlorvinphos), Ectrin (fenvalerate), and Cygon (dimethoate). Insecticide-susceptible house flies were susceptible to all five insecticides (mortality, 62-100%). Flies that were recently colonized from populations on dairy farms in New York were susceptible only to Rabon. Urolepis rufipes (Ashmead) was the most susceptible parasitoid species overall to these insecticides, followed by Muscidifurax raptor Girault & Sanders, Nasonia vitripennis Walker, Pachycrepoideus vindemmiae (Rondani), and Spalangia cameroni Perkins. Compared with susceptible flies, newly colonized flies showed moderate resistance to avermectin B1a (abamectin). Abamectin was more toxic to all of the parasitoids except N. vitripennis and S. cameroni than to newly colonized house flies when exposed for 90 min to plywood boards treated with 0.001-0.1% abamectin. Space sprays with Vapona (dichlorvos) killed all of the parasitoids and susceptible flies and 64% of the newly colonized flies when insects were placed directly in the path of the spray; mortality was substantially lower among flies and parasitoids protected under 5 cm of wheat straw. Space sprays with Pyrenone (pyrethrins) killed greater than 86% of all insects exposed to the spray path except for the newly colonized flies (1% mortality); mortality of insects protected under straw was low (less than 12%) except for S. cameroni (76%). Because responses of the five parasitoids to the different insecticides varied considerably, general conclusions about parasitoid susceptibility to active ingredients, insecticide class, or method of application were not possible.     

Host ontogeny determines parasitoid use of a forest caterpillar

For most organisms, patterns of natural enemy‐mediated mortality change over the course of development. Shifts in enemy pressure are particularly relevant for organisms that exhibit exponential growth during development, such as juvenile insects that increase their mass by several orders of magnitude. As one of the dominant groups of insect herbivores in most terrestrial plant communities, larval lepidopterans (caterpillars) are host to a diverse array of parasitoids. Previous research has described how the frequency of herbivore parasitism varies among host plants or habitats, but much less is known about how parasitism pressure changes during host development. To test whether the two major parasitoid taxa, wasps and flies, differentially attack shared hosts based on host developmental stage, we simultaneously exposed early‐ and late‐instar Euclea delphinii Boisduval (Lepidoptera: Limacodidae) caterpillars to parasitism in the field. We found strong evidence that parasitoids partition hosts by size; adult female wasps preferentially parasitized small caterpillars, whereas adult female flies preferred to attack large caterpillars. Our results demonstrate that host ontogeny is a major determinant of parasitoid host selection. Documenting how shifts in enemy pressure vary with development is important to understanding both the population biology and evolutionary ecology of prey species and their enemies.