Goetheana shakespearei Girault, 1920

Entedoninae, Eulophidae, Chalcidoidea, Hymenoptera

Figure 1	Figure 2	Figure 3	Figure 4
Figure 5	Figure 6	Figure 7	Figure 8
Figure 9	Figure 10

Figures

Fig. 1: Mesosoma and metasoma, dorsal
Fig. 2: Antenna of female
Fig. 3: Funicle segments and clava of antenna
Fig. 4: Head dorsal with vertexal suture and frontal grooves
Fig. 5: Mesosoma dorsal with mesoscutum and scutellum
Fig. 6: Fore wing
Fig. 7: Metasoma
Fig. 8: Petiole
Fig. 9: Antenna of male
Fig. 10: Funicle segments and basal segment of antennal clava of male

Goetheana shakespearei is a larval endoparasitoid of various thrips species belonging to the family Thripidae, preferably on species belonging to the subfamily Panchaetothripinae. Female: length 0.5-0.6 mm. Head and mesosoma dark brown; antennae, legs and gaster uniformly pale yellowish, gaster sometimes with the last 2 or 3 abdominal sclerites dorsally and laterally shaded or with dark spot on each side (Fig. 1 and 7). Head with a complete and straight occipital suture across vertex just behind posterior ocelli (Fig. 4). Antenna with scape slender and about 4.5 to 5.0 x as long as wide; antennal flagellum with 2 funicle segments; F1 much smaller than F2, the latter superficially appears to be a part of clava; F1 without a sensillum, F2 with 1 long sensillum, both with many setae; 3-segmented clava about 2.5 x as long as wide, first claval segment wider than long, second a little longer than wide, third much longer than wide, usually with an apical spicula (Fig. 2 and 3). Mesosoma about as long as gaster (Fig. 1), almost smooth; midlobe of mesoscutum without setae; anterior margin of scutellum straight (Fig. 5). Fore wing very narrow, recurved along posterior margin, nearly parallel-sided below marginal vein, 5.5 to 6.5 x as long as wide; longest marginal setae 1.9 to 2.2 x as long as maximal width of fore wing; disc hyaline, with numerous setae commencing beyond base of marginal vein and scattered in 2 or 3 broken rows toward the apex of the wing (Fig. 6). Hind wing slender,18 to 20 x as long as wide; longest marginal setae 5 to 6 x as long as wing's maximal width. Petiole much wider than long (Fig. 8). Ovipositor very short, occupying about 2/5 length of gaster, not exerted.
Male similar to female, but scape of male antenna extremely swollen, 1.0 to 1.3 x as long as wide (Fig. 9 and 10).

The Genus Goetheana Girault, 1920
This genus comprises 4 described species. All known species of Goetheana are solitary, internal parasitoids of the larval stages of various Thripidae (Terebrantia), and are known from the Oriental, Australian, Ethiopian, and Neotropical regions. All species of the genus having the following features: Body size very small (less than 0.8 mm); head with a complete and straight occipital suture across vertex just behind posterior ocelli; mandible reduced and without teeth; antennal flagellum in both sexes with 2 funicle segments (F1 much smaller than F2, the latter superficially appears to be a part of clava), clava 3-segmented, usually with an apical spicula; scape of male antenna notably swollen in the majority of described species; notauli very narrow but distinct in slide-mounted specimens; midlobe of mesoscutum without setae; fore wing recurved along posterior margin, nearly parallel-sided below marginal vein, length of marginal setae exceeds width of fore wing; petiole much wider than long (Schauff 1991; Triapitsyn 2005). Gahan (1927) gave good descriptions and illustrations of two species, Goetheana incerta and Goetheana shakespearei, and Triapitsyn (2005) gave a world taxonomic revision of Goetheana and three other related entedonine genera of thrips parasitoids. Furthermore Loomans & van Lenteren (1995) provided a overview of the described thrips parasitoids and their importance for biological control of thrips pests.

Female of Goetheana shakespearei is very similar to that of Goetheana incerta. Both may be distinguished only with difficulty: in Goetheana incerta the fore wings are shorter and smaller (5.0 to 5.5 x as long as wide) than in Goetheana shakespearei (5.5 to 6.5 x as long as wide), and the gaster is uniformly pale (in Goetheana shakespearei sometimes the last 2 or 3 abdominal sclerites dorsally and laterally shaded). But male of Goetheana shakespearei differs from Goetheana incerta in having the scape of antenna extremely swollen (1.0 to 1.3 x as long as wide). Whereas male of Goetheana incerta has a slender antennal scape (4.0 to 4.5 x as long as wide, not swollen), similar to female of Goetheana incerta.
Goetheana differs from other entedonine genera by the shape of the antennal funicle segments (in Goetheana funicle segment F1 much smaller than F2, F2 almost fused with clava; in other genera the funicle segments F1 and F2 are subequal), the shape of the fore wing (narrow and recurved below the submarginal vein in Goetheana; species of the genera Entedonomphale, Ceranisus and Thripobius with fore wings broadened beyond submarginal vein), and the length of marginal seta of fore wing (in Goetheana the longest marginal setae is much greater than width of fore wing; species of other genera with longest marginal setae at most equal to width of fore wing, but usually much less than width of fore wing). The chaetotaxy of the fore wing disc differs in the genera: in Goetheana the setae commences beyond base of marginal vein and scattered in 2 or 3 broken rows toward the apex of the wing, in Entedonomphale the fore wing is evenly setose in apical half and bare in basal half, and in Ceranisus and Thripobius the fore wing is uniformly setose beyond base of marginal vein except for a bare area at posterior margin behind base of marginal vein, which is demarcated anteriorly by a sinuate or straight line of setae. Compared to species of Ceranisus with 2-segmented and Entedonomphale with unsegmented antennal clava, species of Goetheana as well as species of Thripobius have a 3-segmented antennal clava.

Life history
G. shakespearei exhibits arrhenotoky and produces both sexes. Sex-ratio, developmental time and longevity is largely mediated by temperature. The total life cycle of this parasitoid varies from 18.6 days to 31.6 days depending on the prevailing temperature and thrips host (Loomans & van Lenteren 1995).

Major host genera/species
Goetheana shakespearei has been recorded from various Thripidae, preferably on species belonging to the subfamily Panchaetothripinae such as Caliothrips (Caliothrips insularis, Caliothrips indicus), Dinurothrips hookeri, Heliothrips haemorrhoidalis, Hercinothrips femoralis, Pseudodendrothrips mori and Selenothrips rubrocinctus and species belonging to Thripinae such as Frankliniella occidentalis, Ceratothripoides claratris and Thrips tabaci.

Biological control
Several attempts have been made to manage the red banded thrips, Selenothrips rubrocinctus and the greenhouse thrips, Heliothrips haemorrhoidalis on avocado with parasitoid specimens of G. shakespeari collected from Ghana. A lot of these biological control attempts were not successful (Loomans & van Lenteren 1995).

Additional notes
Levels of parasitism are highly variable. In Ghana 70-80% parasitism by G. shakespearei occurred on cocoa infested by Selenothrips rubrocinctus (Cotterell 1927, 1928). Parasitization levels up to 58.6% have been registered in Japan attacking Pseudodendrothrips mori on mulberry trees (Takagi 1988). In India up 92% parasitism were observed on the weed Achyranthes aspera infested by Caliothrips indicus (Mohan Daniel 1986). Natural incidence and parasitism rates of G. shakespearei depends on the thrips host species parasitized, temperature, habitat and season (Loomans et al. 1997; Loomans & van Lenteren 1995). The parastoid numbers were very low in spring and summer, and began to increase from September onwards, reaching its peak in the main dry season January to February in Ghana (Cotterell 1927), Benin and the Caribbean (Loomans & van Lenteren 1995).

Biogeography

Subtropical; apparently native to the Australasian region; likely also to the Oriental region. Australia, USA - California, Dominican Republic, Indonesia, India, Japan, Venezuela, Thailand, Trinidad and Tobago, Bahamas, Spain, Bulgaria, Africa. Benin, Ghana, Ivory Coast (Bouake), South Africa (Limpopo: Skukuza - Kruger National Park).

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