A theoretical basis for the study of predatory syrphid fly ecology

Predacious syrphid fly species, also known as flower flies or hover flies, are cosmopolitan diptera that play two important ecological roles: predator and pollinator. In decades past, syrphid flies were studied by agricultural researchers due to their larvae’s ability to function as a biological control agent. In recent years, the global decline in both honey bees and various important wild bee species has led ecological researchers to investigate the role of syrphid fly pollination in both natural systems and agriculture. While these two roles have often been considered separately, they are rarely considered together in single studies. Syrphid fly population fluctuations in natural and agricultural systems are understudied, prompting calls for further study into the fundamental drivers of population dynamics of syrphid communities. In order to develop a deeper understanding of the fundamental dynamics of syrphid ecology, the present study offers a community model where both syrphid predation and pollination are incorporated into a single dynamic model. Using populations of predacious syrphid flies, herbivorous insects, and a shared resource flowering plant, the model is used to investigate community dynamics and persistence across different levels of plant reproductive dependence on syrphid pollination. Results indicate distinct levels of community viability across different pollination relationships as well as a tendency toward chaotic dynamics inherent to the trophic interactions of the community.     

Implications of Three Biofuel Crops for Beneficial Arthropods in Agricultural Landscapes

Production of biofuel feedstocks in agricultural landscapes will result in land use changes that may have major implications for arthropod-mediated ecosystem services such as pollination and pest suppression. By comparing the abundance and diversity of insect pollinators and generalist natural enemies in three model biofuel crops: corn, switchgrass, and mixed prairie, we tested the hypothesis that biofuel crops comprised of more diverse plant communities would support increased levels of beneficial insects. These three biofuel crops contained a diverse bee community comprised of 75 species. Overall, bees were three to four times more abundant in switchgrass and prairie than in corn, with members of the sweat bee (Halictidae) and small carpenter bee (Ceratina spp.) groups the most abundant. Switchgrass and prairie had significantly greater bee species richness than corn during the July sampling period. The natural enemy community at these sites was dominated by lady beetles (Coccinellidae), long-legged flies (Dolichopodidae), and hover flies (Syrphidae) which varied in their response to crop type. Coccinellids were generally most abundant in prairie and switchgrass, with the exception of the pollen feeding Coleomegilla maculata that was most abundant in corn. Several rare or declining coccinellid species were detected in prairie and switchgrass sites. Dolichopodidae were more abundant in prairie and switchgrass while Syrphidae showed no significant response to crop type. Our results indicate that beneficial insects generally responded positively to the increased vegetational diversity of prairie and switchgrass sites; however, when managed as a dedicated biofuel crop, plant and arthropod diversity in switchgrass may decrease. Our findings support the hypothesis that vegetationally diverse biofuel crops support higher abundance and diversity of beneficial insects. Future policy regarding the production of biofuel feedstocks should consider the ecosystem services that different biofuel crops may support in agricultural landscapes.     

Opiine parasitoids (Hymenoptera: Braconidae) and biological control of fruit flies (Diptera: Tephritidae) in Australia: Past,present and future

Opiine braconids are parasitoids of the immature stages of frugivorous tephritids. The female wasp lays her eggs into the eggs or larvae of the fruit fly host, where the immature wasp develops before emerging as a next-generation adult from the now dead host pupal case. In support of a new generation of Australian fruit fly parasitoid research, this paper comprehensively reviews what is known about the Australian fruit fly infesting opiines. Based on the most recent taxonomic revision 11 fruit fly infesting opiine species are documented to occur in Australia, but we consider as doubtful the record for Diachasmimorpha longicaudata and consider the record for Fopius illusorius to be tentative without further collections. We identify that the systematics and taxonomy of the Australian native fruit fly infesting opiines are in urgent need of further work. The history of fruit fly biological control in Australia is comprehensively reviewed, including the export of native Australian opiines for fruit fly control elsewhere in the world. Australia was actively involved in three major classical biological control programmes against fruit flies from the turn of the 1900s until the mid-1960s. Despite the introduction of 11 opiine species, plus numerous other natural enemies, only Fopius arisanus established in eastern Australia, where in South-east Queensland it can now cause between 30 and 40% mean parasitism. In addition to the exotic F. arisanus, the native species Diachasmimorpha kraussii and Diachasmimorpha tryoni also cause fruit fly parasitism in agriculturally important crops: both species have also been liberated widely outside of Australia for fruit fly control. Other Australian opiines have not been reared from flies infesting commercial crops and appear biologically restricted to the fruits and environs of Australian east-coast rainforests. The biology literature for D. tryoni and D. kraussii is comprehensively reviewed, while for F. arisanus, already reviewed elsewhere, key literature only is covered. Forward looking, we consider the potential for inoculative or inundative releases of opiines in areas where they do not currently occur to be good, while conservation biological control may help to increase the impacts of parasitoids in areas where they are already established.     

Phlebotomine fauna (Diptera,Psychodidae) in Rio Preto State Park,Southern Espinhaço Range,Minas Gerais,Brazil

A study of the phlebotomine sand fly fauna was carried out in the Rio Preto State Park (Minas Gerais, Brazil) aiming to associate the presence of vector species with the risk of leishmaniasis transmission in that region. Sand flies were captured using HP light traps for four months from January to October 2013. The traps were exposed continuously for 40 h each month, in nine fixed sites. A total of 3129 sand flies were captured, belonging to 19 species. Lutzomyia evandroi (Costa Lima & Antunes 1936) was the predominant species (76.48%). The distribution of the phlebotomine sand flies and climatic factors (temperature, humidity, and rainfall) was evaluated. The summer was the season with higher occurrence of insects captured (82.4%). The presence of cutaneous and visceral vector species in park areas requires attention and regular monitoring of sand fly fauna in this conservation area to ensure the well-being of visitors and the regional ecotourism.     

On the complementarity of DNA barcoding and morphology to distinguish benign endemic insects from possible pests: the case of Dirioxa pornia and the tribe Acanthonevrini (Diptera: Tephritidae: Phytalmiinae) in Australia

Fruit flies are considered economically important insects due to some species being agricultural pests. However, morphological identification of fruit fly adults and larvae can be difficult requiring a high level of taxonomic expertise, with misidentifications causing problematic false-positive/negative results. While destructive molecular techniques can assist with the identification process, these often cannot be applied where it is mandatory to retain a voucher reference specimen. In this work, we non-destructively (and partial-destructively) processed larvae and adults mostly belonging to the species Dirioxa pornia (Walker, 1849), of the poorly studied nonpest fruit fly tribe Acanthonevrini (Tephritidae) from Australia, to enable molecular identifications whilst retaining morphological vouchers. By retaining the morphological features of specimens, we confirmed useful characters for genus/species-level identification, contributing to improved accuracy for future diagnostics using both molecular and morphological approaches. We provide DNA barcode information for three species of Acanthonevrini known from Australia, which prior to our study was only available for a single species, D. pornia. Our specimen examinations provide new distribution records for three nonpest species: Acanthonevroides variegatus Permkam and Hancock, 1995 in South Australia, Acanthonevroides basalis (Walker, 1853) and D. pornia in Victoria, Australia; as well as new host plant records for D. pornia, from kangaroo apple, apricot and loquat.     

Contribution of small insects to pollination of common buckwheat, a distylous crop

Crop pollination by animals is an essential ecosystem service. Among animal-pollinated crops, distylous plants strongly depend on animal pollination. In distylous pollination systems, pollinator species are usually limited, although flowers of some distylous plants are visited by diverse animals. We studied the pollination biology of common buckwheat ( Fagopyrum esculentum ), a distylous crop mainly pollinated by honeybees and visited by many insect species, to evaluate the effects of non-honeybee species on pollination services. We focused on insects smaller than honeybees to determine their contribution to pollination. We applied pollination treatments with bags of coarse mesh to exclude flower visits by honeybees and larger insects and compared the seed set of bagged plants with that of untreated plants for pin and thrum flower morphs. We found a great reduction of seed set only in bagged pin flowers. We also confirmed that small insects, including ants, bees, wasps and flies, carried pin-morph pollen. These small insects transfer pollen from the short anthers of pin flowers to the short styles of thrum flowers, leading to sufficient seed set in thrum flowers. Consequently, small, non-honeybee insects have the potential to maintain at least half of the yield of this honeybee-dependent distylous crop.     

Fly pollination of Linum lewish (Linaceae)

This study examines the reproductive biology of Linum lewisii Pursh. (Linaceae), a polyphilic species visited by small bees and generalist flies in montane Colorado. L. lewisii plants growing at different sites experience large temporal and spatial variations in pollinator visits. Their ability to attract both dipteran and hymenopteran pollinators allows pollination under varying conditions as pollinator pool composition changes. Although L. lewisii is self-compatible, hand-pollination studies indicate that insects are required for seed production. The relative effectiveness of fly and bee pollinators is assessed in terms of per-visit pollen deposition. Insect visitation patterns are combined with per-visit effectiveness data to evaluate the relative importance of different pollinator groups. Overall, bees tend to be more effective than flies in depositing pollen. However, in many instances flies appear to be responsible for more pollen deposition due to their higher visitation rates.     

Effects of grazing intensity on pollinator abundance and diversity,and on pollination services

1. Pollinating insects provide important ecosystem services and are influenced by the intensity of grazing. Based on the Intermediate Disturbance Hypothesis (IDH), pollinator diversity is expected to peak at intermediate grazing intensities. However, this hump‐shaped relationship is rarely found. 2. The effect of grazing intensity was tested on flower cover, on the abundance and richness of bees, hoverflies and bee flies, and on pollination services to early‐flowering bee‐pollinated Asphodelus ramosus L. For that, we used data on 11 plant–pollinator phryganic communities from Lesvos Island (Greece) widely differing in grazing intensities. 3. Flower abundance and richness showed hump‐shaped relationships with grazing intensity. Grazing affected the abundance and richness of bees and hoverflies directly and also indirectly, through changes in the flower community. Grazing influenced directly the richness but not the abundance of bee flies. Overall, pollinator abundance and richness showed hump‐shaped relationships with grazing intensity, but variations in strength (hoverfly abundance) and direction (bee community) of the effect appeared along the season. Early in the season, grazing increased bee abundance but decreased richness, resulting in increased pollen limitation in A. ramosus. 4. The effects of grazing on pollinators vary with the intensity of the disturbance, generally supporting the IDH, and the timing of land‐use activities may influence pollination services. Management strategies should include moderate grazing levels to preserve overall diversity in this area, however, the conservation of particular early bee or bee‐pollinated species may benefit from reduced grazing in early spring.     

Seasonal changes in seaweed deposition,seaweed fly abundance,and parasitism at the pupal stage along sandy beaches in central Japan

Seasonal relationships among stranded wrack quantity, seaweed fly abundances, and parasitism at the pupal stage were studied along three sandy beaches in central Japan. The seasonal occurrence patterns of puparia of seaweed flies Coelopa frigida and Fucellia spp. generally corresponded to seaweed deposition, which peaked in May–July and October–December. Parasitoids use fly puparia in these seasons. However, the occurrence of seaweed flies and their parasitoids varied among the three sandy beaches and did not correspond to the wrack amounts. These findings suggest that populations of seaweed flies and their parasitoids are seasonally, but not spatially, regulated by bottom‐up processes. The parasitoid assemblage of fly puparia was composed of two Aleochara (Coleoptera: Staphylinidae), two Trichopria (Hymenoptera: Diapriidae), and five pteromalid species (Hymenoptera), but the rate of parasitism was less than 20% and might have had little effect on fly populations.     

Bird and insect pollinators differ in specialization and potential pollination services along disturbance and resource gradients

Combined studies of the communities and interaction networks of bird and insect pollinators are rare, especially along environmental gradients. Here, we determined how disturbance by fire and variation in sugar resources shape pollinator communities and interactions between plants and their pollinating insects and birds. We recorded insect and bird visits to 21 Protea species across 21 study sites and for 2 years in Fynbos ecosystems in the Western Cape, South Africa. We recorded morphological traits of all pollinator species (41 insect and nine bird species). For each site, we obtained estimates of the time since the last fire (range: 2–25 calendar years) and the Protea nectar sugar amount per hectare (range: 74–62 000 g/ha). We tested how post-fire age and sugar amount influence the total interaction frequency, species richness and functional diversity of pollinator communities, as well as pollinator specialization (the effective number of plant partners) and potential pollination services (pollination service index) of insects and birds. We found little variation in the total interaction frequency, species richness and functional diversity of insect and bird pollinator communities, but insect species richness increased with post-fire age. Pollinator specialization and potential pollination services of insects and birds varied differently along the environmental gradients. Bird pollinators visited fewer Protea species at sites with high sugar amount, while there was no such trend for insects. Potential pollination services of insect pollinators to Protea species decreased with increasing post-fire age and resource amounts, whereas potential pollination services of birds remained constant along the environmental gradients. Despite little changes in pollinator communities, our analyses reveal that insect and bird pollinators differ in their specialization on Protea species and show distinct responses to disturbance and resource gradients. Our comparative study of bird and insect pollinators demonstrates that birds may be able to provide more stable pollination services than insects.     

Differentiation of floral color and odor in two fly pollinated species ofMetrodorea (Rutaceae) from Brazil

We investigated if differences in morphological characters in two species ofMetrodorea (Rutaceae) from Brazilian semideciduous forests correspond to some pollination divergence.M. nigra andM. stipularis are sympatric species, display a similar floral morphology, are protandrous, self-incompatible, their flower periods overlap, and both are pollinated by flies.M. nigra main pollinators arePseudoptiloleps nigripoda (Muscidae) andFannia sp. (Fanniidae);M. stipularis major pollinators arePhaenicia eximia (Calliphoridae),Palpada sp. andOrnidia obesa (Syrphidae). The distinct floral odor (disagreeable inM. nigra and sweet inM. stipularis) and color (brownish violet vs. pale yellow) determine the differences on type and number of floral visitors observed. Several species from semideciduous forests initially considered to be pollinated by diverse insects, present flies as main pollinators, stressing the importance of fly pollination in such habitats.     

Fidelity of Hymenoptera and Diptera pollinators in onion (Allium cepa L.) pollination

Onion (Allium cepa L.) is protandrous in nature and requires cross‐pollination to avoid inbreeding. The pollination potential of native bees (Hymenoptera) and true flies (Diptera) was assessed in the perspective of finding the best pollinators for onion cross‐pollination and seed multiplication. The community of pollinators was composed of four bee species and twelve true fly species. Episyrphus balteatus, Eupeodes sp., Musca domestica and Eristalinus aeneus were the most abundant pollinators. The maximum pollinator activity was observed from 12 to 24 days after opening of the flowers. The pollination effectiveness of tested bees (Apis dorsata and Apis florea) was greater than true flies (E. balteatus, Eupeodes sp., M. domestica, E. aeneus and Callihoridae sp.) in terms of Spears values.     

Crane flies and microlepidoptera also function as pollinators in Epidendrum (Orchidaceae: Laeliinae): the reproductive biology of E. avicula

Crane flies and microlepidoptera have been recorded as pollinators in unrelated orchid groups, but these insects have never been recorded in Epidendroideae, the most species‐rich orchid subfamily, which includes one of the most diverse genera among Orchidaceae, Epidendrum. Based on data on phenology, floral morpho‐anatomy, pollinators, pollination mechanisms and breeding system, the reproductive biology of E. avicula was studied in south‐eastern Brazil. Epidendrum avicula possess osmophores that produce a citric fragrance at night. The flowers attract Tipulidae flies and several families of microlepidoptera that drink the nectar produced in a tube formed by the adnation of the labellum and column. As is common in Epidendrum, after removing the pollinarium, both crane flies and micro‐moths get trapped by the proboscis, which frightens the insects and inhibits any possible intent to immediately visit another flower. The behavior of the pollinators on flowers, plus the retention of the anther cap by the pollinarium, results in a reduction in the occurrence of geitonogamy. Because E. avicula is self‐incompatible, the consequence of pollinator behavior and the floral mechanisms tend to reduce the pollen loss. As far as we know, this is the first study to report the reproductive biology of a species of Epidendroideae pollinated by crane flies and microlepidoptera. Based on more recent concepts of plant–pollinator interactions, although E. avicula is pollinated by several species belonging to two distinct orders, suggesting an unspecialized pollination system is involved, nectar‐seeking microlepidoptera and Tipulidae flies can be recognized as a single functional group.     

On the population dynamics of the Australian bushfly

Each spring, the Australian bushfly (Musca vetustissima Walker) migrates from its over-wintering areas located in central Australia and repopulates the south-eastern parts of the continent. In this paper, a simple size-structured model, in the form of a reaction–advection equation with delay, is presented for the evolution of the population density of the bushfly. The model takes into account the important role of cattle dung in its life cycle and includes the effects of wind-borne migration. Bifurcation analyses of the model equations in the absence of wind advection reveals the environmental conditions under which the fly population will die out, reach a stable equilibrium or explode. Numerical simulations which include advection reveal the importance of wind-borne migration in building up the population of flies in south-eastern Australia during the summer months.     

中国农业昆虫生态调节服务价值估算

昆虫是生物多样性最丰富的物种类群,其在维持生态系统功能,维系并保持着自然界的生态平衡,满足人类需求中的具有重要作用。讨论了昆虫生态服务价值的定量估算方法,基于2007年统计数据,计算了我国农牧业生产中昆虫传粉功能、天敌昆虫控害功能和分解作用的服务价值。结果表明,昆虫在我国农业生产中传粉服务价值为6790.30×108元,占其当年农作物生产总经济价值的54.05%;天敌昆虫的控害服务价值为2621.00×108元,占其当年重要作物总经济价值的9.09%;分解昆虫(甲虫)对牧场牛羊排泄物的分解作用的价值远超过90.84×108元。由此估计出了我国2007年农业昆虫生态调节服务总价值超过9502.14×108元,相当于当年国内生产总值GDP的3.7%。显示昆虫所产生的生态调节服务价值,与我国森林或草地生态系统的直接和间接服务价值处于同一数量级,同样具有巨大的经济价值。为保护与利用昆虫生物多样性,发挥其生态服务功能奠定了基础。     

Genetic differentiation associated with host plants and geography among six widespread species of South American Blepharoneura fruit flies (Tephritidae)

Tropical herbivorous insects are astonishingly diverse, and many are highly host‐specific. Much evidence suggests that herbivorous insect diversity is a function of host plant diversity; yet, the diversity of some lineages exceeds the diversity of plants. Although most species of herbivorous fruit flies in the Neotropical genus Blepharoneura are strongly host‐specific (they deposit their eggs in a single host plant species and flower sex), some species are collected from multiple hosts or flowers and these may represent examples of lineages that are diversifying via changes in host use. Here, we investigate patterns of diversification within six geographically widespread Blepharoneura species that have been collected and reared from at least two host plant species or host plant parts. We use microsatellites to (1) test for evidence of local genetic differentiation associated with different sympatric hosts (different plant species or flower sexes) and (2) examine geographic patterns of genetic differentiation across multiple South American collection sites. In four of the six fly species, we find evidence of local genetic differences between flies collected from different hosts. All six species show evidence of geographic structure, with consistent differences between flies collected in the Guiana Shield and flies collected in Amazonia. Continent‐wide analyses reveal – in all but one instance – that genetically differentiated flies collected in sympatry from different host species or different sex flowers are not one another's closest relatives, indicating that genetic differences often arise in allopatry before, or at least coincident with, the evolution of novel host use.     

Ecosystem services provided by dung beetles in Australia

Although there are nearly 500 species of native dung beetles in Australia, most are adapted to small, hard, dry, pelletised marsupial droppings and not to dealing with the large, moist deposits of cattle. In 1788, Governor Arthur Phillip arrived at Botany Bay with five cows, two bulls, 44 sheep and seven horses: this signalled major changes in Australia. Now there are about 27 million cattle, whose annual dung production has a dry matter content of about 42 million tonnes. Until CSIRO introduced exotic dung beetles in the 1960s, the dung of these herbivores sat on the soil surface, sometimes for years, locking up organic matter, smothering pasture and polluting waterways. CSIRO introduced 53 exotic dung beetle species, of which 43 were released to the Australian mainland between 1965 and 1985. Twenty-three of these have become established, many of which have reached the natural limits of their distribution. I consider the reason for the failure of the other 30 species to establish and briefly review previous contributions to examining the role of dung beetles in delivering ecosystem services, noting that much of the published literature concerns laboratory studies. New field data are then examined on the way in which introduced species are transforming dung communities and the ecosystem services they provide. The capacity of deep-tunnelling dung beetles to transform the soil profile is examined along with their effects on pasture production and the flow of nutrients from dung on pasture. The biocontrol capacity of dung beetle activity is considered in relation to the native bush fly, Musca vetustissima, the introduced buffalo fly, Haematobia irritans exigua, and dung-borne intestinal parasites (helminths and Cryptosporidium). The rationale for introducing additional species to Australia is considered.     

Pollination of Aristolochia pallida Willd. (Aristolochiaceae) in the Mediterranean

A first study of the pollination biology of a Mediterranean Aristolochia species in its natural habitat is presented. In all, 183 flowers of Aristolochia pallida Willd. were investigated, which in total contained 73 arthropods, dominated by two groups of Diptera, black fungus gnats (Sciaridae representing 37%) and scuttle flies (Phoridae representing 19%), respectively. However, only Phoridae are regarded as potential pollinators, since pollen has been found exclusively on the body of these insects. All Phoridae belong to the genus Megaselia and are recognised as three morpho-species. The measurements of flower and insect dimensions suggest that size is an important constraint for successful pollination: (a) the insects must have a definitive size for being able to enter the flower and (b) must be able to get in touch with the pollen. Only very few insect groups found in A. pallida fulfil these size requirements. However, size alone is not a sufficient filter as too many fly species of the same size might be trapped but not function as pollinators. Instead, specific attraction is required as otherwise pollen is lost. Since all trapped Phoridae are males, a chemical attraction (pheromones) is proposed as an additional constraint. Since the flowers are protogynous, the record of Megaselia loaded with pollen found in a flower during its female stage proves that this insect must have had visited at least one different flower during its male stage before. Further on, this observation provides strong evidence that the flowers are cross-pollinated. All these factors indicate a highly specialised pollination of A. pallida by Megaselia species.     

Pollination mutualism between Alocasia macrorrhizos (Araceae) and two taxonomically undescribed Colocasiomyia species (Diptera: Drosophilidae) in Sabah, Borneo

Two taxonomically undescribed Colocasiomyia species were discovered from inflorescences of Alocasia macrorrhizos in Kota Kinabalu City, Sabah, Borneo, Malaysia. The aims of this study were to investigate the reproductive ecology of the flies and the plant, ascertain the importance of the flies as pollinators and examine the intimate association between flowering events and life history of the flies. We conducted sampling, observations and field pollination experiments. The flies were attracted by the odour of female‐phase inflorescences in the early morning on the first day of anthesis. They fed, mated and oviposited in the inflorescences for 1 day. On the second day, the flies, covered with pollen grains, left the male‐phase inflorescences for the next female‐phase inflorescences. The immature forms of both fly species hatched, developed and pupated within the infructescences without damaging the fruits, and developed adults emerged when the mature infructescences dehisced. The flowering events and fly behaviours were well synchronized. In field pollination experiments, inflorescences bagged with a fine mesh (insect exclusion) produced almost no fruits, whereas those bagged with a coarse mesh (bee exclusion) produced as many fruits as the open‐pollinated controls. These results indicate that these flies are the most efficient and specialised pollinators for their host, A. macrorrhizos. These flies, in return, depend on A. macrorrhizos for food and habitat through most of their life cycle. This study provides a deeper insight into the less recognised, highly intimate pollination mutualism between Araceae plants and Colocasiomyia flies.