Temperature-dependent phenology of the parasitoid Fopius arisanus on the host Bactrocera dorsalis.

Fopius arisanus (Sonan), an egg-pupal parasitoid of numerous fruit fly species, was recently introduced into Africa for the control of the Oriental fruit fly, Bactrocera dorsalis (Hendel). In this study, life-table data of F. arisanus were generated under laboratory conditions at six constant temperatures (15, 20, 25, 30, 28 and 35 °C; 75% RH, L12:D12 photoperiod) and under variable conditions in a screenhouse on B. dorsalis. Several non-linear functions were fitted to model species development, mortality, longevity and oviposition using the Insect Life Cycle Modeling (ILCYM) software. The established phenology models were stochastically simulated at variable temperatures to estimate the life table parameters. Fopius arisanus completed development from egg to adult at all the temperatures tested except at 35 °C. Mean developmental time from egg to adult was inversely proportional to the temperature. The minimum temperature threshold (Tmin) from egg to adult was 8.15 °C, while the maximum temperature threshold (Tmax) was estimated at 34.2 °C. The optimal temperature for immature stages survival was predicted for 20-30 °C. The optimum fecundity estimated was 251 eggs/female at 22 °C. Following the stochastic simulations under natural conditions of the selected agro-ecological zones, it appears that the Humid Forest with Bimodal Rainfall provides a suitable thermal condition for potential population growth of F. arisanus. The present study shows the importance of temperature on the demographic parameters of F. arisanus. Implications of present findings on the biological control of B. dorsalis under climate change scenarios is discussed.

Local adaptation in thermal tolerance for a tropical butterfly across ecotone and rainforest habitats.

Thermal adaptation to habitat variability can determine species vulnerability to environmental change. For example, physiological tolerance to naturally low thermal variation in tropical forests species may alter their vulnerability to climate change impacts, compared with open habitat species. However, the extent to which habitat-specific differences in tolerance derive from within-generation versus across-generation ecological or evolutionary processes are not well characterized. Here we studied thermal tolerance limits of a Central African butterfly (Bicyclus dorothea) across two habitats in Cameroon: a thermally stable tropical forest and the more variable ecotone between rainforest and savanna. Second generation individuals originating from the ecotone, reared under conditions common to both populations, exhibited higher upper thermal limits (CTmax) than individuals originating from forest (∼3°C greater). Lower thermal limits (CTmin) were also slightly lower for the ecotone populations (∼1°C). Our results are suggestive of local adaptation driving habitat-specific differences in thermal tolerance (especially CTmax) that hold across generations. Such habitat-specific thermal limits may be widespread for tropical ectotherms and could affect species vulnerability to environmental change. However, microclimate and within-generation developmental processes (e.g. plasticity) will mediate these differences, and determining the fitness consequences of thermal variation for ecotone and rainforest species will require continued study of both within-generation and across-generation eco-evolutionary processes. This article has an associated First Person interview with the first author of the paper.

Status of the regulatory environment for utilization of insects as food and feed in Sub-Saharan Africa-a review.

A conducive regulatory environment is crucial for ensuring the safety and effective promotion of insects for direct and indirect human consumption. In this review, national and regional policies, regulations, and relevant publications in sub-Saharan Africa (SSA) were examined for their take on the use of insects as food and feed. Majority of the SSA countries (91.7%) lacked food safety policies, and of the four countries (8.3%) that had, only one considered the 'risk-based approach' for assessing food safety. Two policies, one in Malawi and the other in Tanzania respectively, recognized insects' use. The lack of regulatory frameworks in most SSA countries is partly attributed to inadequate scientific data regarding insects' biological, chemical and physical safety. This potentially exposes consumers to health hazards and limits income from insect and insect-based food and feed operations. However, some information and/or data to inform policy, and in a few cases to develop standards, has been generated by several research and development projects in the region. The need for supportive regulations toward the use of insects has been recognized and is being acted upon in a number of SSA countries. For effective promotion of insects as food and feed in SSA, countries need to generate risk assessment data as recommended by Codex Alimentarius and develop and implement relevant standards.

Temperature-dependent development, survival and reproduction of Apanteles hemara (Nixon) (Hymenoptera: Braconidae) on Spoladea recurvalis (F.) (Lepidoptera: Crambidae).

The temperature-dependent development of Apanteles hemara (Nixon), a larval endoparasitoid of the devastating amaranth pest Spoladea recurvalis (F.) was studied in the laboratory at six constant temperatures (10, 15, 20, 25, 30 and 35 °C), a photoperiod of 12L:12D and a relative humidity of 60-70%. Developmental time decreased significantly with increasing temperature within the range of 15-30 °C. The parasitoid's pupal mortality, successful parasitism rate, adult emergence rate and longevity, sex ratio and fecundity were affected by temperature. The population of A. hemara failed to develop at 10 and 35 °C. The development threshold (Tmin) and the thermal constant (K) were calculated by the linear model while the lethal temperature (Tmax) was determined by the Lactin-1 model. The estimated values of Tmin, Tmax and K by the two models were 10.3 °C, 35.0 °C and 185.18 DD respectively for the total immature development. The estimated value of the optimum temperature using the Taylor model was 30.8 °C. This is the first study to report on the effect of temperature on the developmental parameters of A. hemara giving an insight into its biology. The implications of these findings for the use of A. hemara in biological control are discussed.

Fruit Preference, Parasitism, and Offspring Fitness of Fopius arisanus (Hymenoptera: Braconidae) Exposed to Bactrocera dorsalis' (Diptera: Tephritidae) Infested Fruit Species.

Fopius arisanus (Sonan) (Hymenoptera: Braconidae) is a koinobiont solitary parasitoid of various fruit flies, particularly those in the genus Bactrocera. Researchers introduced F. arisanus into Africa for the biological control of Bactrocera dorsalis (Hendel) (Diptera: Tephritidae), a pest of a wide range of fruit trees and vegetables. However, the suitability of host fruit species as egg-laying substrates for parasitoid development remains poorly investigated in tropical Africa. The present study examines the preference and performance of F. arisanus on B. dorsalis reared on eleven fruit species through laboratory choice-test trials. We assessed the oviposition activity, parasitism rate, developmental time, and offspring fitness of F. arisanus on nine cultivated and two wild host fruits species. Oviposition attempts were higher on Psidium guajava (L.) (Myrtales: Myrtaceae) and Mangifera indica (L.) (Sapindales: Anacardiaceae) than on the other host fruits tested. The wasp parasitized host eggs in P. guajava in no-choice experiments. Psidium guajava, Irvingia wombulu (Vermoesen) (Malpighiales: Irvingiaceae), and Irvingia gabonensis (Aubry_Lecomte) Baill (Malpighiales: Irvingiaceae) were suitable for parasitism in choice tests. Of all host fruits tested, the body and hind tibia lengths of both parasitoid sexes emerging from M. indica were longer than on the others. The female ovipositor was long on Annona squamosa (L.) (Magnoliales: annonaceae) and short on Eribotrya japonica ([Thunb.] Lindl.; Rosales: Rosaceae). We obtained the longest preimaginal developmental time for both sexes on E. japonica and the shortest for females and males on Carica papaya (L.) (Brassicales: Caricaceae). These results demonstrate the ability of some tested fruit species to serve for the permanent establishment of F. arisanus in the field.

Effects of Host Age and Density on the Performance of Apanteles hemara (Hymenoptera: Braconidae), a Larval Endoparasitoid of Spoladea recurvalis (Lepidoptera: Crambidae).

The amaranth leaf-webber, Spoladea recurvalis (Fabricius; Lepidoptera: Crambidae), is a serious pest of Amaranthus sp. in Africa and Asia. Apanteles hemara (Nixon; Hymenoptera: Braconidae) is by far the most important larval endoparasitoid of the amaranth leaf-webber. We examined the effects of host density and age on the biological characteristics of A. hemara. The regression model of the number of hosts supplied to A. hemara against the number of larvae parasitized resulted in a curve corresponding to type II functional response, with a significant increase in the number of hosts parasitized up to the density of 30 hosts before being constant up to 40 hosts. In contrast, the parasitism rate decreased linearly with increasing host densities. Development time, sex ratio, and adult longevity were not significantly affected by host density. The immature parasitoid mortality was significantly higher at higher host densities. Apanteles hemara did not parasitize 7-d-old larvae and beyond, while parasitism was significantly higher among 1- to 2-d-old compared with 3- to 4-d-old larvae. Immature parasitoid mortality was 2.6 times higher in 1- to 2-d-old larvae compared with 5- to 6-d-old larvae. The developmental period of the parasitoid from egg to adult was longest among 1- to 2-d-old larvae and least among 5- to 6-d-old larvae. Nonreproductive mortality was markedly higher among 1- to 2-d-old larvae compared with the older larvae. Adult female A. hemara were significantly larger on 3- to 4-d-old larvae compared with either 1- to 2-d-old or 5- to 6-d-old larvae. We discuss the implications of our results for the interpretation of functional response in parasitoids, mass rearing, conservation, and augmentative biological control of S. recurvalis.

Postharvest processes of edible insects in Africa: A review of processing methods, and the implications for nutrition, safety and new products development.

In many African cultures, insects are part of the diet of humans and domesticated animals. Compared to conventional food and feed sources, insects have been associated with a low ecological foot print because fewer natural resources are required for their production. To this end, the Food and Agriculture Organization of the United Nations recognized the role that edible insects can play in improving global food and nutrition security; processing technologies, as well as packaging and storage techniques that improve shelf-life were identified as being crucial. However, knowledge of these aspects in light of nutritional value, safety, and functionality is fragmentary and needs to be consolidated. This review attempts to contribute to this effort by evaluating the available evidence on postharvest processes for edible insects in Africa, with the aim of identifying areas that need research impetus. It further draws attention to potential postharvest technology options for overcoming hurdles associated with utilization of insects for food and feed. A greater research thrust is needed in processing and this can build on traditional knowledge. The focus should be to establish optimal techniques that improve presentation, quality and safety of products, and open possibilities to diversify use of edible insects for other benefits.

Insects for Income Generation Through Animal Feed: Effect of Dietary Replacement of Soybean and Fish Meal With Black Soldier Fly Meal on Broiler Growth and Economic Performance.

The ever-increasing animal feed costs are driving many vulnerable communities involved in animal husbandry out of business. The high cost is mainly driven by the protein source, which represents the most expensive component in animal feed. In conventional feed, protein is obtained mainly from soybean and fish meal (SFM). The present study explored potential of partially replacing this SFM with black soldier fly prepupae meal (BSFPM) in Cobb 500 broiler chicken diets. A SFM-based diet was compared to three experimental diets formulated by partially substituting SFM with BSFPM at 13.8, 27.4, and 42.0% of the crude protein (CP) in the starter feed and 11.0, 37.2, and 55.5% of the CP in the finisher feed of diets D1, D2, and D3, respectively. Dietary effects on average daily feed intake, average daily body weight gain, feed conversion ratio, carcass characteristics, breast meat sensory attributes, and the economic implication of their use in broiler production were evaluated. Replacement of SFM with BSFPM did not affect daily feed intake, daily body weight gain, feed conversion ratio, aroma or taste of cooked breast meat. A 16.0% higher Cost Benefit Ratio and 25.0% better Return on Investment was recorded when the birds were reared on the highest concentration of black soldier fly (D3) compared to the conventional diet which was 19.0% more expensive. The implication of these findings for the promotion of insect mass production enterprises for animal feed protein, and their potential for income generation and job creation particularly in developing countries is discussed.

Moisture adsorption properties and shelf-life estimation of dried and pulverised edible house cricket Acheta domesticus (L.) and black soldier fly larvae Hermetia illucens (L.).

Edible insects are part of the diets of a significant proportion of rural populations in the tropics especially Africa and Asia, and their use as source of key nutrients for better nutrition is re-emerging. Indigenously, elemental methods are used to process the insects before they are consumed or sold in retail outlets. In recent years, better knowledge of processing, packaging and storage has become necessary because of commercialisation needs. A common processing approach involves drying after a brief heat-treatment step, and then milling into a powdered product which is sold to manufacturers or consumers as ingredient for processing final products. The hydration properties of dried powders of edible house cricket and black soldier fly larvae (BSFL) were studied with the aim of predicting shelf-life stability under typical packaging and storage temperatures experienced in the tropics. Moisture adsorption isotherms were determined gravimetrically at 25, 30 and 35 °C, over 0.11-0.97 water activity (aW) range, and the data fitted to various models. Sorption isotherms were of type II according to Brunauer classification indicating monolayer-multilayer sorption behaviour. Cricket powder exhibited higher hydration capacity, and aW of this product was less sensitive to temperature variation as compared to BSFL powder. In the two products, water exhibited transitions from bound- to free- state at ~5 g/100 g moisture content. Based on Heiss-Eichner model, a shelf-life of 7 months at 25 °C can be achieved if the cricket and BSFL powders are dried to ca. 5 g/100 g moisture content and packaged in 80 µm thick polyethylene films. At 35 °C the shelf-life of the cricket product is shortened three- to four-fold whereas the BSFL powder is unable to store.

Performance of Apanteles hemara (Hymenoptera: Braconidae) on two Amaranth Leaf-webbers: Spoladea recurvalis and Udea ferrugalis (Lepidoptera: Crambidae).

Amaranth is an African indigenous vegetable that is gaining popularity due to its nutritional, medicinal, and economic values. In East Africa, frequent outbreaks of Lepidopteran leaf-webbers, Spoladea recurvalis F. (Lepidoptera: Crambidae) and Udea ferrugalis Hübner (1976) are reported on this crop, causing up to 100% foliage loss. The larval endoparasitoid Apanteles hemara Nixon is also frequently found associated with these pests during the outbreaks, however, its performance on both pests has never been documented. Laboratory studies were therefore carried out to assess the acceptability and suitability of S. recurvalis and U. ferrugalis to A. hemara. Both leaf-webber species were accepted by and suitable for the parasitoid. The mean host searching time and oviposition attempts were neither affected by rearing host nor test host. The total developmental time of A. hemara ranged between 10.6 ± 0.16 and 12.8 ± 0.30 days on both hosts. The sex ratio of the parasitoid was female biased when reared on S. recurvalis but male biased on U. ferrugalis. When offered 50 larvae of leaf-webbers for 24 h, a single female A. hemara achieved parasitism rates from 42.63 ± 5.80 to 44.55 ± 5.95, while a cohort of five females resulted in parasitism rates between 87.25 ± 2.70 and 94.67 ± 1.98 %. There was no significant difference between hosts in regard to progeny fitness at each parasitoid density. The parasitoid also caused significant nonreproductive larval mortalities in the hosts. The implications of these findings for mass rearing of the parasitoid as well as for conservation and augmentative biological control of amaranth lepidopteran leaf-webbers in East Africa are discussed.

Liriomyza Leafminer (Diptera: Agromyzidae) Parasitoid Complex in Different Agroecological Zones, Seasons, and Host Plants in Kenya.

Liriomyza leafminers (Diptera: Agromyzidae) are severe pests of vegetables and ornamentals worldwide. Previous studies revealed low leafminer parasitism across different agroecological zones in Kenya. The present paper reports on the composition of leafminer parasitoids at different elevations, in different seasons, and on different host crops. Surveys were conducted monthly from January to November 2012, and nine parasitoid species were recovered. Total mean parasitism in the study sites was 31.23 ± 1.03% from a total of 20 different vegetable Liriomyza-infested crops belonging to seven families. Diglyphus isaea (Walker) (Hymenoptera: Eulophidae), Phaedrotoma scabriventris, a newly released parasitoid, and Opius dissitus Muesebeck (Hymenoptera: Braconidae) were the most abundant at all elevations, accounting for 67.3, 18.6, and 9.2% of total parasitoids, respectively. Elevation, season, and host crop significantly affected the parasitoid species present and their abundance. Diglyphus isaea was more abundant at the high- and mid-elevations at all seasons compared with the low-elevation, whereas the lower-elevation favored higher abundance of P. scabriventris and O. dissitus during the long rainy season compared with the high- and mid-elevations at all seasons. Of all the host crops surveyed, parasitoids were more abundant on tomato, local kidney bean, snow pea and French bean than other crops. The total parasitism rate observed in this study suggests a considerable improvement in leafminer parasitism compared with previous surveys in Kenya. The implications of these findings for leafminer management in vegetable and ornamental production in Kenya are discussed.

Differential Effects of Pesticide Applications on Liriomyza huidobrensis (Diptera: Agromyzidae) and its Parasitoids on Pea in Central Kenya.

Three Liriomyza species [Liriomyza huidobrensis (Blanchard), Liriomyza trifolii (Burgess), and Liriomyza sativae Blanchard] have been reported as the most important leafminer pests in vegetable production systems in Africa. In Kenya, farmers rely on indiscriminate synthetic insecticides use. On-farm field investigations were set up at three different locations (Sagana, Kabaru, and Naromoru) in central Kenya to determine the effect of pesticide application on the abundance of leafminers and their parasitoids under three management practices, namely: farmer practice (FP), reduced pesticide use (RP), and a control with no use of pesticides (CO). In addition, laboratory experiments were designed to test the effect of commonly used pesticides in pea production systems in central Kenya--Dimethoate, Dynamec, Thunder, Cyclone, Bestox, Folicur, Milraz, and Bulldock--on L. huidobrensis and two of its parasitoids, Diglyphus isaea Walker and Phaedrotoma scabriventris Nixon. The mean numbers of leafminer flies in control treatment were higher than in RP and FP in both first and second seasons across all sites, but RP and FP did not differ significantly. Parasitoid numbers were very low and there was no much variation between treatments at each location in both first and second seasons. No significant differences were observed between the three management practices with regards to the yield measurements. In the laboratory, the estimated LD50 values for L. huidobrensis larvae were all more than two times higher than the recommended dosages, while the LD50 of adults were below the recommended dosages. The estimated LD50 values for the parasitoids were much lower than recommended dosages for all pesticides except Thunder. This study, therefore, demonstrates that the pesticides currently used do not control the Liriomyza leafminer larvae that constitute the most destructive stage of the pest, but are rather detrimental to their parasitoids. In addition, the current low level of parasitoids recorded under field conditions even where no pesticide was used during this study, warrants consideration of classical biological control programs.

Species Composition, Distribution, and Seasonal Abundance of Liriomyza Leafminers (Diptera: Agromyzidae) Under Different Vegetable Production Systems and Agroecological Zones in Kenya.

A longitudinal study to identify the species of Liriomyza leafminer, their distribution, relative abundance, and seasonal variation, including their host range, was conducted in vegetable fields at three altitudes in Kenya from November 2011 to November 2012. Three main species were identified: Liriomyza huidobrensis (Blanchard), Liriomyza sativae Blanchard, and Liriomyza trifolii (Burgess), of which L. huidobrensis was the most abundant across all altitudes irrespective of the cropping season and accounting for over 90% of the total Liriomyza specimens collected. Liriomyza species were collected from all infested incubated leaves of 20 crops surveyed belonging to seven families: Fabaceae, Solanaceae, Cucurbitaceae, Malvaceae, Brassicaceae, Amaranthaceae, and Amaryllidaceae. However, more than 87.5% of the Liriomyza species were obtained from only four of these crops: Pisum sativum L., Phaseolus vulgaris L., Solanum lycopersicum L., and Solanum tuberosum, thereby demonstrating that Fabaceae and Solonaceae crops are the most important hosts with regard to Liriomyza species richness and relative abundance. L. huidobrensis had the widest host range (20 crops), followed by L. sativae (18 crops) and L. trifolii (12 crops). Although L. trifolii has been considered the dominant Liriomyza leafminer in Kenya, this study suggests that this may not be the case anymore, as L. huidobrensis dominates at all altitudes.