No Indication of High Host-Plant Specificity in Afrotropical Geometrid Moths.

Specificity is one of the fundamental concepts in ecology. Host specificity of phytophagous insects has been of particular interest because of its crucial role in diversification and life-history evolution. However, the majority of tropical insects remain insufficiently explored with respect to their host-plant relations. A lack of respective data is also hindering the debate over whether higher levels of host-plant specificity prevail in tropical insects compared to temperate ones. We investigated host-plant specificity of forest geometrid moths (Lepidoptera: Geometridae) in equatorial Africa using host-plant acceptability trials with neonate larvae, with the addition of field observations. We compare our experimental data to the (well-known) host-specificity patterns of closely related temperate (hemiboreal) species. Similarly to the temperate region, there were broadly polyphagous tropical species in several clades of Geometridae utilizing hosts belonging to different plant families. Phylogenetic comparative analysis returned no significant differences in host specificity between the two regions. Our study contributes to the evidence that host-plant specificity of herbivores is not necessarily substantially higher in tropical than temperate regions.

Long-term species loss and homogenization of moth communities in Central Europe.

As global biodiversity continues to decline steeply, it is becoming increasingly important to understand diversity patterns at local and regional scales. Changes in land use and climate, nitrogen deposition and invasive species are the most important threats to global biodiversity. Because land use changes tend to benefit a few species but impede many, the expected outcome is generally decreasing population sizes, decreasing species richness at local and regional scales, and increasing similarity of species compositions across sites (biotic homogenization). Homogenization can be also driven by invasive species or effects of soil eutrophication propagating to higher trophic levels. In contrast, in the absence of increasing aridity, climate warming is predicted to generally increase abundances and species richness of poikilotherms at local and regional scales. We tested these predictions with data from one of the few existing monitoring programmes on biodiversity in the world dating to the 1960s, where the abundance of 878 species of macro-moths have been measured daily at seven sites across Hungary. Our analyses revealed a dramatic rate of regional species loss and homogenization of community compositions across sites. Species with restricted distribution range, specialized diet or dry grassland habitat were more likely than others to disappear from the community. In global context, the contrasting effects of climate change and land use changes could explain why the predicted enriching effects from climate warming are not always realized.

Determinants of parasitoid communities of willow-galling sawflies: habitat overrides physiology, host plant and space.

Studies on the determinants of plant-herbivore and herbivore-parasitoid associations provide important insights into the origin and maintenance of global and local species richness. If parasitoids are specialists on herbivore niches rather than on herbivore taxa, then alternating escape of herbivores into novel niches and delayed resource tracking by parasitoids could fuel diversification at both trophic levels. We used DNA barcoding to identify parasitoids that attack larvae of seven Pontania sawfly species that induce leaf galls on eight willow species growing in subarctic and arctic-alpine habitats in three geographic locations in northern Fennoscandia, and then applied distance- and model-based multivariate analyses and phylogenetic regression methods to evaluate the hierarchical importance of location, phylogeny and different galler niche dimensions on parasitoid host use. We found statistically significant variation in parasitoid communities across geographic locations and willow host species, but the differences were mainly quantitative due to extensive sharing of enemies among gallers within habitat types. By contrast, the divide between habitats defined two qualitatively different network compartments, because many common parasitoids exhibited strong habitat preference. Galler and parasitoid phylogenies did not explain associations, because distantly related arctic-alpine gallers were attacked by a species-poor enemy community dominated by two parasitoid species that most likely have independently tracked the gallers' evolutionary shifts into the novel habitat. Our results indicate that barcode- and phylogeny-based analyses of food webs that span forested vs. tundra or grassland environments could improve our understanding of vertical diversification effects in complex plant-herbivore-parasitoid networks.

Is climate warming more consequential towards poles? The phenology of Lepidoptera in Finland.

The magnitude and direction of phenological shifts from climate warming could be predictably variable across the planet depending upon the nature of physiological controls on phenology, the thermal sensitivity of the developmental processes and global patterns in the climate warming. We tested this with respect to the flight phenology of adult nocturnal moths (3.33 million captures of 334 species) that were sampled at sites in southern and northern Finland during 1993-2012 (with years 2005-2012 treated as an independent model validation data set). We compared eight competing models of physiological controls on flight phenology to each species and found strong support for thermal controls of phenology in 66% of the species generations. Among species with strong thermal control of phenology in both the south and north, the average development rate was higher in northern vs. southern populations at 10 °C, but about the same at 15 and 20 °C. With a 3 °C increase in temperature (approximating A2 scenario of IPPC for 2090-2099 relative to 1980-1999) these species were predicted to advance their phenology on average by 17 (SE ± 0.3) days in the south vs. 13 (±0.4) days in the north. The higher development rates at low temperatures of poleward populations makes them less sensitive to climate warming, which opposes the tendency for stronger phenological advances in the north from greater increases in temperature.

Bottom-up impact on the cecidomyiid leaf galler and its parasitism in a tropical rainforest.

The relative importance of host-plant resources, natural enemies or their interactions in controlling the population of galling insects and their parasitism is poorly known for tropical gallers. In this study, we assessed the impacts of plant quality and density of host trees in regulating the densities of a galler species, the cecidomyiid leaf galler (Cecidomyiini sp. 1EJV) and its parasitoids and inquilines on Neoboutonia macrocalyx trees in Uganda. We manipulated the nutritional quality (or vigour) and the resource concentration with four levels each of fertilization and the group size of host tree. We then recorded the effects of these treatments on the growth rate and total leaf area of host plants, the density of gallers and their mortality by parasitoids and inquilines. Higher levels of fertilization and host density resulted in significantly higher total leaf area than did ambient nutrient levels, and lowest tree densities, respectively. Fertilization also caused significant change in the growth rate of leaf area. Both higher fertilization and host density caused higher density of gallers. Total leaf area was positively associated with galler density, but within galled replicates, the galled leaves were larger than the ungalled leaves. Although highest levels of fertilization and density of host trees caused significant change in the densities of parasitoids, the rate of parasitism did not change. However, tree-density manipulations increased the rate of inquilinism, but on a very low level. Our results demonstrate a trophic cascade in the tropical galler and its parasitoids as a response to bottom-up effects.

Three new Afrotropical species of Tersilochinae (Hymenoptera: Ichneumonidae) from the Kibale National Park, Uganda.

Seven species of Tersilochinae (Hymenoptera: Ichneumonidae) have been found from Uganda in Equatorial Africa: Allophrys budongoana Khalaim, Diaparsis interstitialis Khalaim, D. kanyawara sp. nov., D. mostovskii Khalaim, D. sinuator sp. nov., D. umbrosa sp. nov. and Tersilochus moestus Holmgren. Two species of Diaparsis, described in this paper, possess a white banded flagellum, and a key to the group of four Afrotropical species having a white-banded flagellum is provided. Tersilochus moestus Holmgren, previously known only from a single female from South Africa, is recorded from the Kibale National Park, Uganda. Antennae and ovipositor of this species, broken in the type specimen, are described for the first time. 

Herbivore community promotes trait evolution in a leaf beetle via induced plant response.

Several recent studies have emphasised that community composition alters species trait evolution. Here, we demonstrate that differences in composition of local herbivore communities lead to divergent trait evolution of the leaf beetle Plagiodera versicolora through plant-mediated indirect interactions. Our field surveys, genetic analyses and community-manipulation experiments show that herbivore community composition determines the degree of herbivore-induced regrowth of willows (Salicaceae), which in turn, promotes the divergent evolution of feeding preference in the leaf beetle from exclusive preference for new leaves to a lack of preference among leaf-age types. Regrowth intensity depends both on the differential response of willows to different herbivore species and the integration of those herbivore species in the community. Because herbivore-induced regrowth involves phenological changes in new leaf production, leaf beetle populations develop divergent feeding preferences according to local regrowth intensity. Therefore, herbivore community composition shapes the selection regime for leaf beetle evolution through trait-mediated indirect interactions.

Growth and Reproductive Performance of Edible Grasshopper (Ruspolia differens) on Different Artificial Diets.

Ruspolia differens (Serville) (Orthoptera: Tettigoniidae), also known as the 'edible grasshopper', 'African edible bush-cricket', and 'nsenene', is regarded as one of the most promising edible insect species that can be used for food, particularly in Sub-Saharan Africa. However, there is insufficient information on suitable diets and their effects on survival, adult weight, fecundity, and developmental time of this species, which are preconditions for large-scale production. In this study, we experimentally evaluated the effects of 12 diets (wheat bran, rice seed head, finger millet seed head, soya bran, maize bran, fresh maize comb, millet flour, chicken feed egg booster, simsim cake, sorghum seed head, powdered groundnut, and germinated finger millet), that are known to be accepted by R. differens, on their growth and reproductive parameters. The survival rate, developmental time, and adult weight varied considerably on the various diets. The highest nymphal survival rates, shortest development times, and highest adult weights were recorded for both sexes when fed fresh maize comb and germinated finger millet diet. Lifetime fecundity of females fed on germinated finger millet also was, on average, more than twice higher compared to other diets. The present study demonstrated that relatively inexpensive and locally available germinated finger millet, fresh maize seed (at the silking stage on the comb), sorghum seedhead, and finger millet seedhead could be successfully used to rear and sustain populations of R. differens. Our findings contribute to the future design of an effective mass-rearing system for this economically important edible insect.

Environmental controls on the phenology of moths: predicting plasticity and constraint under climate change.

Ecological systems have naturally high interannual variance in phenology. Component species have presumably evolved to maintain appropriate phenologies under historical climates, but cases of inappropriate phenology can be expected with climate change. Understanding controls on phenology permits predictions of ecological responses to climate change. We studied phenological control systems in Lepidoptera by analyzing flight times recorded at a network of sites in Finland. We evaluated the strength and form of controls from temperature and photoperiod, and tested for geographic variation within species. Temperature controls on phenology were evident in 51% of 112 study species and for a third of those thermal controls appear to be modified by photoperiodic cues. For 24% of the total, photoperiod by itself emerged as the most likely control system. Species with thermal control alone should be most immediately responsive in phenology to climate warming, but variably so depending upon the minimum temperature at which appreciable development occurs and the thermal responsiveness of development rate. Photoperiodic modification of thermal controls constrains phenotypic responses in phenologies to climate change, but can evolve to permit local adaptation. Our results suggest that climate change will alter the phenological structure of the Finnish Lepidoptera community in ways that are predictable with knowledge of the proximate physiological controls. Understanding how phenological controls in Lepidoptera compare to that of their host plants and enemies could permit general inferences regarding climatic effects on mid- to high-latitude ecosystems.

Northward invasion of the parasitic deer ked ( Lipoptena cervi), is there geographical variation in pupal size and development duration?

The deer ked (Lipoptena cervi) is a common ectoparasite of cervids. During the last decades the species has rapidly invaded in northern Europe, especially in Finland, towards the north and increased its prevalence on the moose population. Consequently, during this rapid invasion the deer ked has faced more severe climatic conditions. We studied whether pupal size (measured as pupal weight) and pupal development duration of the deer ked varies along historical invasion zones and temperature zones towards north in Finland. Moreover, we explored possible size- and gender-dependent variation in pupal development duration. We divided wild-collected pupae in respect to their origin in two ways: (1) temperature zones (from south-west to colder north-east) and (2) invasion history (from early to late establishment). We reared pupae in the controlled laboratory conditions in identical temperature and light conditions. Pupal size decreased towards north and the smaller pupae developed faster. However, the results do not show differences in pupal size or developmental characteristics between the invasion zones. This supports the idea of rapid developmental plasticity of the deer ked and that not the invasion history but the current temperature regime determines the life history of the deer ked when invading towards a colder environment.

Extensive sampling reveals the phenology and habitat use of Afrotropical parasitoid wasps (Hymenoptera: Ichneumonidae: Rhyssinae).

Tropical invertebrates, such as the ichneumonids of tropical forests, are poorly known. Here, we report the first results of extensive sampling at Kibale National Park, Uganda, by providing some of the first tropical ecological data for the ichneumonid subfamily Rhyssinae. We sampled ichneumonids with 34 Malaise traps for a year in 10 sites, in habitats ranging from primary forest to farmland. We also gathered weather and vegetation data. The total sampling effort was 373 trap months and we caught 444 rhyssines in six species. We caught the most rhyssines in dry weather, and towards the end of the sampling year. The rhyssines showed a clear preference for decaying logs and for primary forest. We fitted a model which can be used to predict future catches at the site, and draw conclusions on when rhyssines emerge and on their adult lifespan. Sampling extensively gave us a wealth of ecological data on a poorly known parasitoid wasp subfamily. We recommend that future tropical sampling collect ecological data, and that existing data from previous large-scale surveys be used for ecological analyses.

Developing Mass Egg-Laying Medium for the Edible Ruspolia differens (Orthoptera: Tettigonidae).

The edible katydid, Ruspolia differens (Serville), is an economically and nutritionally important insect species that is common and widespread in Africa. Currently, there are aims to develop mass-rearing methods suitable for this species, but a critical first step would require a suitable oviposition medium to ensure economic and easy production of eggs with high hatchability. The objective of the present study was to determine the oviposition preference and egg hatchability in different oviposition media. Based on preliminary testing of a wide range of different materials, two oviposition media, namely, artificial (folded plastic cloth) and natural medium (leaf sheath of Pennisetum purpureum Schumach attached to a stem), were selected for a detailed experiment. A two-choice laboratory experiment was conducted in which R. differens females were able to select between the folded plastic cloth (one side covered with short fleece) and a leaf sheath of P. purpureum (wrapped in polyester fiber). The results show that R. differens laid almost three times more eggs on the plastic cloth (mean ± SE, 169.3 ± 18.0) than on the natural substrate, P. purpurem (69.0 ± 13.9). However, the hatchability (overall mean% ± SE, 56.5 ± 4.7) of eggs in the two media did not differ. Folded plastic cloth, therefore, provided a more effective medium for egg laying by R. differens, and is thus a promising oviposition medium for getting natural populations to start laying eggs in future mass rearing.

Host plants of the non-swarming edible bush cricket Ruspolia differens.

The edible Ruspolia differens (Orthoptera: Tettigoniidae) is a widely-consumed insect in East Africa but surprisingly little is known of its host plant use in the field. We studied host plants used by non-swarming R. differens for 15 months, in central Uganda. In particular, we assessed the use of host plant species with respect to host cover in the field and host parts used by R. differens, also recording their sex, developmental stages, and colour morph. Ruspolia differens were found on 19 grass and two sedge species and they were observed predominantly (99% of 20,915 observations) on seven grasses (namely, Panicum maximum, Brachiaria ruziziensis, Chloris gayana, Hyparrhenia rufa, Cynodon dactylon, Sporobolus pyramidalis, and Pennisetum purpureum). Ruspolia differens was most frequently observed on the most common grass of each study site but P. maximum, and S. pyramidalis were used more frequently than expected from their cover in the field. Furthermore, R. differens were observed predominantly on inflorescences (97% of feeding observations) and much less frequently on the leaves (3.0%), stems (0.1%), and inflorescence stalks (0.1%) of grasses and sedges. Host use was not independent of sex, developmental stage, or colour morph. Panicum maximum was the preferred host of the youngest nymphs of R. differens. Overall, our findings indicate that a continuous supply of diverse grass resources with inflorescences is necessary for the management and conservation of wild populations of R. differens.

Extensive sampling and thorough taxonomic assessment of Afrotropical Rhyssinae (Hymenoptera, Ichneumonidae) reveals two new species and demonstrates the limitations of previous sampling efforts.

Tropical forest invertebrates, such as the parasitoid wasp family Ichneumonidae, are poorly known. This work reports some of the first results of an extensive survey implemented in Kibale National Park, Uganda. A total of 456 individuals was caught of the subfamily Rhyssinae Morley, 1913, which in the Afrotropical region was previously known from only 30 specimens. Here, the six species found at the site are described and the Afrotropical Rhyssinae are reviewed. Two new species, Epirhyssa johanna Hopkins, sp. nov. and E. quagga sp. nov., are described and a key, diagnostic characters, and descriptions for all 13 known Afrotropical species are provided, including the first description of the male of Epirhyssa overlaeti Seyrig, 1937. Epirhyssa gavinbroadi Rousse & van Noort, 2014, syn. nov. is proposed to be a synonym of E. uelensis Benoit, 1951. Extensive sampling with Malaise traps gave an unprecedented sample size, and the method is recommended for other poorly known tropical areas.

The plant-based by-product diets for the mass-rearing of Acheta domesticus and Gryllus bimaculatus.

Edible insect rearing could provide one alternative for protein production by having a smaller environmental impact than traditional livestock farming due to insects' ability to convert organic side streams. Currently, the insect rearing industry utilizes soybeans as a major source of protein in the feeds. Protein-rich by-products of food industry could be used to replace them in insect feeds, but it is not known if they also meet the insects' nutritional requirements. Our study evaluated the growth performance of two widely used edible cricket species, Acheta domesticus and Gryllus bimaculatus (Orthoptera: Gryllidae), on 18 experimental diets. The experimental diets included commercial chicken feeds and cricket diets, where soybean was partly and completely replaced with by-products from food industry: potato protein, barley mash, barley feed, compressed leftover of turnip rape and mix of broad bean and pea on three levels of protein. We found that the high- and medium-protein turnip rape and barley mash diets produced the highest yield and an increase in all performance variables. Overall, the high- and medium-protein diets produced the highest yield, growth and fastest development. Our results showed that by-products of food industry could be utilized as a part of the cricket feeds and thus advance the goals of circular economy.

Fatty acid content and composition in edible Ruspolia differens feeding on mixtures of natural food plants.

OBJECTIVES: To develop successful mass-rearing programs of edible insects, knowledge of the feeds and their influence on nutritional content is critical. We assessed the influence of natural food plants (grass inflorescences) and their mixtures on fatty acid profiles of edible Ruspolia differens. We reared neonate nymphs to adult on six dietary treatments consisting of one, and mixtures of two, three, five, six and eight plants. RESULTS: The contents of saturated, monounsaturated and polyunsaturated fatty acids, omega-6/omega-3 ratio, and adult body weight did not differ among dietary treatments. However, the composition of fatty acids differed significantly among insects fed on six dietary treatments, but only for the rare fatty acids. Our results demonstrate that even if natural diets (grass inflorescences) do not strongly modify fatty acid contents or compositions of R. differens, when reared from neonate nymphs to adults, their n - 6/n - 3 fatty acid ratio is generally low and thus good for a healthy human diet.

Choice of voles among genotypes of birch seedlings: its relationship with seedling quality and preference of insects.

Selective feeding by herbivores on establishing seedlings has been suggested to affect genotype frequencies in several plant populations. The existence of genotypes susceptible to herbivores calls for an explanation in such populations. In the present study we assessed the choice of multiple herbivores, field voles (Microtus agrestis) and insects, among genotypes of silver birch (Betula pendula) representing variation occurring in a naturally regenerated stand. We examined how food choice of voles and insects is related to each other, competitive ability among the seedling genotypes and variation in soil fertility. We set up a field experiment and randomly assigned seedling populations, composed of mixed genotypes, to fully crossed insect exposure and fertilization treatments. After the first growing season we exposed a half of the seedling populations to vole herbivory. Voles selected clearly among the genotypes: they preferred the fastest growing seedlings as well as those with a low density of resin droplets on their stems. The preference of voles and insects among the genotypes was tightly correlated. We conclude that the effects of herbivory compensate those of intraspecific competition in this system and thus favor coexistence of genotypes differing in their susceptibility to herbivores.

Genetic and environmental determinants of insect herbivore community structure in a Betula pendula population.

A number of recent studies have shown that intraspecific genetic variation of plants may have a profound effect on the herbivorous communities which depend on them. However less is known about the relative importance of intraspecific variation compared to other ecological factors, for example environmental variation or the effects of herbivore damage. We randomly selected 22 Betula pendula genotypes from a local population (< 0.9 ha), cloned them and planted cloned seedlings on two study sites separated at a regional scale (distance between sites about 30 km) to examine an insect community of 23-27 species on these genotypes. B. pendula genotypes did not differ in their species richness, but the total mean abundance and the structure of the insect herbivore community was significantly affected by the genotype, which could account for up to 27% of the total variation in community structure. B. pendula genotype accounted for two to four times more variation in the arthropod community structure than did environmental (block) variation on a local scale, while on a regional scale, genotypic and environmental (site) variation accounted for 4-14% of the arthropod community structure. The genetic effects were modified by environmental variation on both a local and regional scale over one study year, and locally, the largest part of the variation (38%) could be explained by the genotype × environment (block) interactions. Suppression of insect herbivores during one growing season led to changed arthropod community structure in the following growing season, but this effect was minimal and could explain only 4% of the total variation in insect community structure. Our results suggest that both genetic and environmental factors are important determinants of the community structure of herbivorous insects. Together these mechanisms appear to maintain the high diversity of insects in B. pendula forest ecosystems.

Rapid estimation of the oxidative activities of individual phenolics in crude plant extracts.

Previous studies of purified phenolic compounds have revealed that some phenolics, especially ellagitannins, can autoxidise under alkaline conditions, which predominate in the midgut of lepidopteran larvae. To facilitate screening for the pro-oxidant activities of all types of phenolic compounds from crude plant extracts, we developed a method that combined our recent spectrophotometric bioactivity method with an additional chromatographic step via UPLC-DAD-MS. This method allowed us to estimate the total pro-oxidant capacities of crude extracts from 12 plant species and to identify the individual phenolic compounds that were responsible for the detected activities. It was found that the pro-oxidant capacities of the plant species (i.e., the concentrations of the easily-oxidised phenolics) varied from 0 to 57 mg/g dry wt, representing from 0% to 46% of the total phenolics from different species. UPLC-DAD-MS analysis revealed that most flavonol and flavone glycosides were only slightly affected by alkaline conditions, thus indicating their low pro-oxidant activity. Interestingly, myricetin-type compounds differed from the other flavonoids, as their concentrations decreased strongly due to alkaline incubation. The same effect was detected for hydrolysable tannins and prodelphinidins, suggesting that a pyrogallol sub-structure could be a key structural component that partially explains their easy oxidation at high pH. Other types of phenolic compounds, such as hydroxycinnamic acids, were relatively active, as well. These findings demonstrate that this method displays the potential to identify most of the active and inactive pro-oxidant phenolic compounds in various plant species.

Leaf litter decomposition differs among genotypes in a local Betula pendula population.

Ecosystem processes, such as plant litter decomposition, are known to be partly genetically determined, but the magnitude of genetic variation within local populations is still poorly known. We used micropropagated field-grown saplings of 19 Betula pendula genotypes, representing genetic variation in a natural birch population, to examine (1) whether genotype can explain variation in leaf litter decomposition within a local plant population, and (2) whether genotypic variation in litter decomposition is associated with genotypic variation in other plant attributes. We found that a local B. pendula population can have substantial genotypic variation in leaf litter mass loss at the early stages of the decomposition process and that this variation can be associated with genotypic variation in herbivore resistance and leaf concentrations of soluble proteins and total nitrogen (N). Our results are among the first to show that fundamental ecosystem processes can be significantly affected by truly intraspecific genetic variation of a plant species.