Performance of Trichogrammatoidea sp. nr lutea Girault (Hymenoptera: Trichogrammatidae) collected from Mussidia spp. in Kenya on eggs of six lepidopteran hosts

Mussidia nigrivenella Ragonot (Lepidoptera: Pyralidae), an important pest of maize ears in West Africa, has never been reported to attack crops in East and southern Africa (ESA), though it was found on various wild host plants in these regions. It was suggested that in ESA M. nigrivenella might be under natural control. In Kenya, exploration for natural enemies associated with Mussidia spp. yielded several parasitoids including a trichogrammatid egg parasitoid, Trichogrammatoidea sp. nr lutea Girault. The ability of T. sp. nr lutea to attack the eggs of several lepidopteran species found in Kenya was studied. The lepidopterans included the noctuids Busseola fusca (Fuller) and Sesamia calamistis (Hampson), the pyralids Eldana saccharina Walker, Mussidia fiorii Cecconi and de Joannis and Mussidia‘madagascariensis’, and the crambid Chilo partellus (Swinhoe). The former three species also infest cereals in West Africa. Trichogrammatoidea sp. nr lutea successfully attacked and developed in eggs of all six species indicating its potential to exploit other lepidopteran pests of maize in West Africa. Busseola fusca and S. calamistis were the most suitable hosts and had the largest number of eggs parasitized and progeny per female wasp where E. saccharina and C. partellus were the poorest hosts. The host species used to rear the parasitoid and the age of egg also significantly affected the total number of host eggs parasitized by the parasitoid. It was concluded that the ability of T. sp. nr lutea to exploit lepidopterans that are also pests of maize in West Africa may enhance biological control of M. nigrivenella and it should be considered for translocation to that area from Kenya.     

两种蚜小蜂对烟粉虱寄主的产卵选择性(英文)

双斑恩蚜小蜂和桨角蚜小蜂是华南地区烟粉虱的两种优势种寄生蜂。本文研究了两种寄生蜂对不同龄期烟粉虱寄主的产卵选择特性。结果表明:两种寄生蜂均可寄生烟粉虱的1~4龄若虫。当只有1个龄期的烟粉虱若虫存在时,双斑恩蚜小蜂更多地寄生3龄和4龄若虫,较少寄生1龄和2龄若虫;而桨角蚜小蜂则更多地寄生3龄和2龄若虫,较少寄生1龄和4龄若虫。在4个龄期若虫同时存在时,双斑恩蚜小蜂会明显增加对3龄和4龄若虫的产卵寄生,降低对1龄和2龄若虫的寄生;而桨角蚜小蜂则增加对2龄和3龄若虫的寄生,减少对1龄和4龄若虫的寄生。寄主植物的差异不影响这二种蚜小蜂对各龄期烟粉虱若虫的产卵选择倾向。结果提示,烟粉虱若虫3龄和4龄是双斑恩蚜小蜂最适宜的寄主,而桨角蚜小蜂最适宜的寄主是烟粉虱2龄和3龄若虫。     

两种蚜小蜂对烟粉虱寄主的产卵选择性（英文）

双斑恩蚜小蜂和桨角蚜小蜂是华南地区烟粉虱的两种优势种寄生蜂。本文研究了两种寄生蜂对不同龄期烟粉虱寄主的产卵选择特性。结果表明: 两种寄生蜂均可寄生烟粉虱的1～4龄若虫。当只有1个龄期的烟粉虱若虫存在时,双斑恩蚜小蜂更多地寄生3龄和4龄若虫,较少寄生1龄和2龄若虫;而桨角蚜小蜂则更多地寄生3龄和2龄若虫,较少寄生1龄和4龄若虫。在4个龄期若虫同时存在时,双斑恩蚜小蜂会明显增加对3龄和4龄若虫的产卵寄生,降低对1龄和2龄若虫的寄生;而桨角蚜小蜂则增加对2龄和3龄若虫的寄生,减少对1龄和4龄若虫的寄生。寄主植物的差异不影响这二种蚜小蜂对各龄期烟粉虱若虫的产卵选择倾向。结果提示,烟粉虱若虫3龄和4龄是双斑恩蚜小蜂最适宜的寄主,而桨角蚜小蜂最适宜的寄主是烟粉虱2龄和3龄若虫。     

Suitability of Bemisia tabaci (Hemiptera: Aleyrodidae) instars for the parasitization by Encarsia bimaculata and Eretmocerus sp. nr. furuhashii (Hymenoptera: Aphelinidae) on glabrous and hirsute host plants

The host instar preferences of Encarsia bimaculata and Eretmocerus sp. nr. furuhashii parasitizing Bemisia tabaci and their development on four host plants, collard, eggplant, cucumber and tomato, were studied in the laboratory. Both of the parasitoids accepted all nymphal stages of B. tabaci, but E. bimaculata preferred third and fourth instars while Er. sp. nr. furuhashii preferred second and third instars under both choice and no choice conditions. Regardless of host stage parasitized, adults of parasitoids emerged only from fourth instars. When given the simultaneous choice of all instars, E. bimaculata reduced parasitization of first and second instars (3.73 and 4.76%, respectively) while increasing parasitization of third and fourth instars (5.44 and 6.93%, respectively), in contrast Er. sp. nr. furuhashii increased its parasitization of second and third instar nymphs (1.27 and 3.17%, respectively) and decreased that of first and fourth instars (7.0 and 3.06%, respectively). Host plants did not significantly influence instar preference for either parasitoid. Developmental periods of both the parasitoids from egg to adult emergence were longest when first instars were parasitized and shortest when fourth instars were selected. Parasitoid developmental time was generally shorter on glabrous plants than on hirsute plants.     

Effect of temperature and host species on parasitism,development time and sex ratio of the egg parasitoid Trichogrammatoidea lutea Girault (Hymenoptera: Trichogrammatidae)

The developmental biology of Trichogrammatoidea lutea Girault (Hymenoptera: Trichogrammatidae) was studied at six constant temperatures (18, 21, 24, 27, 30 and 35 °C) on eggs of three lepidopteran host species: Helicoverpa armigera (Hübner) (Noctuidae), Chilo partellus (Swinhoe) (Crambidae) and Cadra cautella (Walker) (Pyralidae). T. lutea did not complete development at 35 °C on any of the three host species. Parasitism levels were highest on H. armigera at 27 °C (58%), C. cautella at 27 and 30 °C (31% and 28%) and C. partellus between 24 and 30 °C (13–17%). Realized progeny of T. lutea per parasitized host egg was influenced by host size. The number of progeny of T. lutea per parasitized host egg was highest on H. armigera, followed by C. partellus and lowest on C. cautella. The sex ratio was female biased on C. partellus, female biased on C. cautella with the exception of 21 °C and close to 1:1 on H. armigera. The rate of development from egg to pupa and egg to adult was fastest on H. armigera and slowest on C. partellus. Lower thresholds for development and degree days (DD) of T. lutea from egg to adult were 12.8 °C and 105.4 DD on H. armigera, 11.3 °C and 141.6 DD on C. partellus and 12.9 °C and 118.2 DD on C. cautella, respectively. Based on these results, H. armigera is the most suitable host for mass rearing of T. lutea for biological control of Lepidoptera pests because of the relatively high parasitism levels, short development time, greater clutch size and balanced sex ratio. C. cautella may also be used although longer exposure times might be required due to lower parasitism levels.     

卷蛾分索赤眼蜂和拟澳洲赤眼蜂对小菜蛾利他素各成分的嗅觉反应

卷蛾分索赤眼蜂Trichogrammatoidea bactrae Nagaraja和拟澳洲赤眼蜂Trichogramma confusum Viggiani是小菜蛾Plutella xylostella (L.)的优势寄生蜂。本研究利用“Y”型管测定了2种赤眼蜂对小菜蛾P. xylostella利他素-卵表和腹部鳞片13种饱和烷烃的嗅觉反应。结果显示: 卷蛾分索赤眼蜂交配雌蜂进入2, 6, 10, 14-四甲基十五烷、正十五烷、正十七烷处理区的百分数分别为80.65%, 68.75%和66.67%, 表明这3种烷烃对卷蛾分索赤眼蜂交配雌蜂有显著的吸引作用, 其他10种饱和烷烃对卷蛾分索赤眼蜂雌蜂无明显作用。拟澳洲赤眼蜂进入2, 6, 10, 14-四甲基十五烷、正三十五烷和正十五烷处理区的数目分别占84.38%, 70%和62.16%, 表明其对拟澳洲赤眼蜂交配雌蜂起着显著的吸引作用, 而另外10种烷烃对拟澳洲赤眼蜂雌蜂无作用。卷蛾分索赤眼蜂和拟澳洲赤眼蜂未交配雄蜂进入13种烷烃处理区和对照区的时间均无显著差异, 表明利他素各组分对2种赤眼蜂未交配雄蜂均无吸引作用。     

Suitability of five mealybug species (Hemiptera,Pseudococcidae) as hosts for the solitary parasitoid Anagyrus sp. nr. pseudococci (Girault) (Hymenoptera: Encyrtidae)

Anagyrus sp. nr. pseudococci is an endoparasitoid which has been used as a biological control agent of mealybug pests. In this study, we compared the suitability of five mealybugs species with different phylogenetic relationships and geographical origins as hosts of this parasitoid. The selected mealybugs were: (1) a Mediterranean-native species, Planococcus ficus, sharing a long co-evolutionary history with the parasitoid; (2) three exotic species, the Afrotropical Planococcus citri, the Australasian Pseudococcus calceolariae and the Neotropical Pseudococcus viburni, with a recent history; and (3) the Neotropical Phenacoccus peruvianus, with no previous common history with the parasitoid. Host suitability was assessed based on different fitness parameters, such as body size, developmental time, emergence rate and sex ratio. The parasitoid was able to complete development in all mealybug species. Nevertheless, its emergence rate significantly varied among mealybug species, with the highest values observed in Pl. ficus and Pl. citri, intermediate values in Ps. calceolariae and the lowest ones in Ps. viburni and Ph. peruvianus. The body size of adult wasp females varied with host suitability and was positively correlated with other measures of parasitoid fitness, including the emergence rate and the sex ratio. The parasitoid developmental time differed among mealybug species but did not correlate with any other measure of fitness. A female biased sex ratio was found in the parasitoid progeny emerged from all mealybug species, except in Ps. viburni and Ph. peruvianus. There was a direct relationship between the proportion of females in the parasitoid progeny and the emergence rate.     

Host selection behaviour and specificity of the solitary parasitoid of mealybugs Anagyrus sp. nr. pseudococci (Girault) (Hymenoptera,Encyrtidae)

The host selection behaviour of Anagyrus sp. nr. pseudococci was compared in no-choice tests among five mealybug species of different geographical and phylogenetic origin, including the Mediterranean native host, Planococcus ficus and four exotic mealybug species, one of the same genus, Pl. citri, two Pseudococcus species, Ps. calceolariae and Ps. viburni and a more distant one, Phenacoccus peruvianus. All five studied mealybug species were recognised by the parasitoid as potential hosts and parasitised, but the behavioural pattern of host recognition, host handling and the level of host acceptance of Anagyrus sp. nr. pseudococci significantly varied among the five studied species, indicating a clear preference for the two Planococcus species, Pl. ficus in particular. The results suggest that A. sp. nr. pseudococci has a broader host range and a more generalist behaviour in comparison with other Anagyrus species. Practical implications of the findings are discussed.     

Developmental time,longevity, reproductive capacity and sex ratio of the mealybug parasitoid Anagyrus sp. nr. pseudococci (Girault) (Hymenoptera: Encyrtidae)

Key biological parameters of Anagyrus sp. nr. pseudococci reared on the citrus mealybug Planococcus citri (Risso) were determined. The mean longevity of female wasps was 21.1 days, the median lethal time (LT₅₀) was 22 days, the mean number of offspring per female was 30.2 wasps and the sex ratio of the progeny was slightly male biased (1.1 males per female). Wasps fed with water showed a significantly lower longevity compared to that recorded for wasps fed with syrup for bee nutrition mixed with water. The longevity of female wasps was significantly higher compared to males when the syrup-water solution was provided as food. Longevity was significantly higher for female wasps fed with syrup and water in comparison with those exposed to host mealybugs. The mean development time of female and male wasps was 14.7±0.1 and 14.5±0.1 days, respectively.     

Response to host age of the egg parasitoid Anaphes n.sp. (Hymenoptera: Mymaridae)

The mymarid Anaphes n.sp. was observed to parasitize the eggs of the carrot weevil Listronotus oregonensis (LeConte) up to 138 h old at 23±1°C and 18L:6D. Adult emergence of the parasitoid was observed in 85%‐97.5% of the weevil eggs 42–138 h old and host embryogenesis was interrupted in nearly 100% of these eggs. In host eggs of 18 h old, parasitism was very high but not all parasitoids completed their development and their subsequent emergence was generally retarded. An.sp. could parasitize host eggs 162 h old, but such an attack rarely retarded the normal development of the host. The parasitoid was unable to complete its development in sterile carrot weevil eggs.     

Defense Response of Native and Alien Mealybugs (Hemiptera: Pseudococcidae) Against the Solitary Parasitoid Anagyrus sp. nr. pseudococci (Girault) (Hymenoptera: Encyrtidae)

The host behavioral and immune (encapsulation) defenses against the parasitoid Anagyrus sp. nr. pseudococci were compared for five mealybug species with different phylogenetic relationships and geographical origins: i) a Mediterranean native mealybug species, Planococcus ficus, with a long co-evolutionary history with the parasitoid; ii) three alien mealybugs species, Planococcus citri, Pseudococcus calceolariae and Pseudococcus viburni, with a more recent co-evolutionary history; and iii) a fourth alien mealybug species, Phenacoccus peruvianus, with no previous common history with the parasitoid. Three host defense behaviors were registered: abdominal flipping, reflex bleeding and walking away. The native host Pl. ficus and its congener Pl. citri exhibited the lowest probability of defense behavior (0.11?±?0.01 and 0.09?±?0.01 respectively), whereas the highest value was observed in P. viburni (0.31?±?0.02). Intermediate levels of defense behavior were registered for Ps. calceolariae, and Ph. peruvianus. The probability of parasitoid encapsulation was lowest and highest for two alien host species, Ph. peruvianus (0.20?±?0.07) and Ps. viburni (0.86?±?0.05), respectively. The native host Pl. ficus, its congener Pl. citri and Ps. calceolariae showed intermediate values (0.43?±?0.07, 0.52?±?0.06, and 0.45?±?0.09, respectively). The results are relevant with respect to biological control and to understand possible evolutionary processes involved in host range of A. sp. nr. pseudococci.     

Continuous rearing on Ephestia kuehniella reshaped quality of the parasitoid wasp Trichogramma brassicae (Hymenoptera: Trichogrammatidae)

Continuous mass rearing of Trichogramma brassicae (Bezdenko) at commercial mass-rearing insectaries may affect both quality and performance of natural enemies. In the present study, we studied the quality and performance of a colony of T. brassicae reared for over 45 generations (G) on Ephestia kuehniella Zeller using two-sex life table parameters and parasitism capacity. Our results revealed that although different generations showed no significant difference in terms of female longevity or total life span until G35, G5 and G10 had the highest values of fecundity, gross reproductive rate (GRR), net reproductive rate (R₀), intrinsic rate of natural increase (r), and finite rate of increase (λ). No significant difference in male adult longevity was found among different generations. The longest and shortest mean generation times (T) were found in G10 (13.65 ± 2.31 d) and G45 (13.25 ± 3.37 d), respectively. The finite rate of parasitism (ω) ranged from 0.355 ± 2.332 host/parasitoid/day in G5 to 0.242 ± 0.017 host/parasitoid/day in G45. However, ω did not show any significant difference until G20. These results indicate that T. brassicae wasps held under continuous laboratory rearing declined in quality after 20 generations, and therefore periodical rejuvenation of the colony by adding feral parasitoids is strongly recommended.     

Composite asymmetry as an indicator of quality in the beneficial wasp Trichogramma nr. brassicae (Hymenoptera: Trichogrammatidae)

Trichogramma are egg parasitoids that are widely cultured for the control of moth pests in commercial insectaries. Stock quality is often maintained by releasing wasps from a mass culture in an area with target host eggs and initiating fresh stocks from parasitized eggs. Here we test if variability among individuals as measured by fluctuating asymmetry can be used as an additional method of quality control in Trichogramma nr. brassicae by comparing fluctuating asymmetry levels between a random sample of wasps before release and wasps that successfully reach host eggs. Although the asymmetry of individual traits was not associated with wasp fitness, a total asymmetry measure based on combining the uncorrelated asymmetries of nine traits did predict the ability of wasps to reach the eggs.     

Interactions between the ant Tapinoma nigerrimum (Hymenoptera: Formicidae) and the main natural enemies of the vine and citrus mealybugs (Hemiptera: Pseudococcidae)

Honeydew-excreting hemipterans, such as mealybug pests, can be protected from their natural enemies by tending ants in return for honeydew, thereby compromising the aims of biological control. In this respect, antagonistic interactions between the ant Tapinoma nigerrimum, native to the Mediterranean basin, and the main natural enemies of both the vine mealybug (VMB), Planococcus ficus, and the citrus mealybug (CM), Planococcus citri, were assessed in laboratory conditions. Parasitism of vine and CMs by their respective parasitoids, Anagyrus sp. nr. pseudococci and Leptomastix dactylopii, was negatively affected by the ant T. nigerrimum. Similarly, T. nigerrimum was shown to significantly disrupt the predatory potential of ladybird larvae, Cryptolaemus montrouzieri, when foraging on host CMs. By contrast, the presence of the ant did not negatively influence the predatory activity of C. montrouzieri adults when feeding on CMs. Consequently, the encyrtid parasitoids A. sp. nr. pseudococci and L. dactylopii and the larval stage of the predator C. montrouzieri may be considered as T. nigerrimum-sensitive, whereas the adults of C. montrouzieri may be regarded as T. nigerrimum-resistant predators. Accordingly, the ant T. nigerrimum constitutes a threat to the biological control of mealybugs by either the encyrtids A. sp. nr. pseudococci and L. dactylopii or the larval stage of the ladybird C. montrouzieri. Hence, adequate control of T. nigerrimum is highly recommended before any release of these mealybugs' natural enemies.     

Selection and evaluation of RT-qPCR reference genes for expression analysis in the tiny egg parasitoid wasp,Trichogramma dendrolimi matsumura (Hymenoptera: Trichogrammatidae)

The egg parasitoid, Trichogramma spp., is an important biological control agent used against a broad range of Lepidopteran pests in agriculture and forestry. The biology of Trichogramma has been studied in details. Further studies should focus on the molecular mechanisms of Trichogramma by qualifying the expression of related genes It is critical to select appropriate reference genes for normalizing RT-qPCR results and establishing a robust method for quantifying target gene expression. This study aimed to identify and validate appropriate reference genes for use in RT-qPCR analysis of Trichogramma dendrolimi. Ten candidate housekeeping genes, namely beta-actin (ACTIN), forkhead box O (FOXO), glyceraldehyde-3-phosphate dehydrogenase (GAPDH), heat shock protein 90 (HSP90), ribosomal protein L10a (RPL10a), L18 (RPL18), L28 (RPL28), S13 (RPS13), S15 (RPS15), and superoxide dismutase (SOD), were tested for their suitability as reference genes for developmental stage (3rd, 4th, 5th, 6th, 7th, 8th, 9th, and 10th day after parasitization), tissue (head, thorax, and abdomen of adults), sex of adults (male and female), and temperature (17℃, 25℃, and 32℃). According to the GeNorm analysis, a robust analysis should involve using an appropriate combination of reference genes, namely, at least three genes for different development stages, two genes for different tissues, two genes for different sex, and two genes for different temperatures, respectively. According to the RelFinder method by the integrated results of GeNorm, NormFinder, BestKeeper, and the ΔCt method, we identified the developmental stage-specific reference genes SOD, GAPDH, and ACTIN; tissue-specific reference genes RPL18 and RPS15; sex-specific reference genes RPL18 and SOD; and temperature-specific reference genes RPL18 and RPL10a. This study provides a standardized procedure for the quantification of gene expression in T. dendrolimi and will be helpful for future biological control programs using Trichogramma wasps.     

Changes in Biological Characteristics of Trichogramma pretiosum (Hym.: Trichogrammatidae) Reared on Eggs of Anagasta kuehniella (Lep.: Pyralidae) for 23 Generations

Possible changes in biological characteristics of Trichogramma pretiosum (Riley) (Hymenoptera: Trichogrammatidae) were monitored during 23 generations reared on eggs of Anagasta kuehniella (Zeller) (Lepidoptera: Pyralidae), to determine quality control parameters in rearing procedures of this parasitoid. Parasitism rates (percent of eggs parasitized) of T. pretiosum varied from 10 to 76%, with lower values for the first four generations. From the 17th generation, the variability in the parasitism rate was lower. Viability (percent adult emergence rate) of T. pretiosum in eggs of A. kuehniella varied from 22 to 100%, again with lower values in earlier generations. Adult longevity of T. pretiosum varied from 5.3 to 15.9 days, and showed a reduced longevity with increasing number of generations. Sex ratio of the offspring of this parasitoid was not affected by the timing of parasitism on eggs of A. kuehniella.     

Morphology and taxonomy of five marine sand-dwelling Thecadinium species (Dinophyceae) from Japan, including four new species: Thecadinium arenarium sp. nov., Thecadinium ovatum sp. nov., Thecadinium striatum sp. nov. and Thecadinium yashimaense sp. nov.

Thecadinium inclinatum Balech and four new marine sand‐dwelling species of the dinoflagellate genus Thecadinium are described from the sandy beaches along the coast of Shikoku, Japan. Thecadinium inclinatum is thecate, bilaterally flattened, elliptical in shape, non‐photosynthetic, and measures 55–75 μ in length and 43–59 μ in depth. The epi‐ and hypotheca theca are semielliptical and the thecal surface is smooth with small pores. The plate formula is Po (pore plate), 3′, 7″,?c,?s, 5″′1″′.Thecadinium ovatum sp. nov. is thecate, non‐photosynthetic, bilaterally flattened and almost oval in lateral view. The cell measures 40–50 μm in length and 33–40 μm in depth. The hypotheca has two or three strong antapical spines. The plate formula is 3′, 6″,6c, 5s?, 5″′, 1″′. Thecadinium striatum sp. nov. is thecate, non‐photosynthetic, bilaterally flattened and somewhat elliptical in lateral view. The cell is 33–41 μm long and 23–30 μm deep. Several striae are present on the hypotheca. The plate formula is 3′, 6″, 6c, 5s?, 5″′, 1″″. Thecadinium yashimaense sp. nov. is bilaterally flattened, photosynthetic and elliptical in ventral view. The cell is 44–65 μm long and 23–36 μm wide. The thecal surface is smooth with small pores. he cingulum forms a steep left–handed spiral. The plate formula is Po, 3′, la, 6″, 5c, 4s, 5″′, 1″′. Thecadinium arenarium sp. nov. is somewhat wedge‐shaped in ventral view, photosynthetic with brownish chloroplasts and almost rounded in cross section. The cingulum forms a steep left‐handed spiral. The cell measures 35–41 μm in length and 25–30 μm in width. The thecal surface is weakly reticulated with small pores. The hypotheca is conical. The plate formula is Po, 3′, la, 6″, 5c, 4s, 5″′, 1″″.