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SECTION 1 a Description of the Taxonomy, Biology, Ecology and Life Cycle of the Parasitoid Microhymenopteran Wasp, Tachardiaephagus Somervillei

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SECTION 1 a Description of the Taxonomy, Biology, Ecology and Life Cycle of the Parasitoid Microhymenopteran Wasp, Tachardiaephagus Somervillei SECTION 1 A description of the taxonomy, biology, ecology and life cycle of the parasitoid microhymenopteran wasp, Tachardiaephagus somervillei. A. Taxonomy Scientific name Tachardiaephagus somervillei (Mahdihassan, 1923) Common name None Relationships: Tachardiaephagus somervillei (Mahdihassan) is an encyrtid parasitoid: Class: Insecta Order: Hymenoptera Suborder: Apocrita Superfamily: Chalcidoidea Family: Encyrtidae Subfamily: Encyrtinae The Encyrtidae is one of the most important families of parasitic wasps (parasitoids) for the biological control of harmful insects, including a variety of scale insects infesting woody plants (Noyes and Hayat 1994, Noyes 2012). The Encyrtidae currently comprises 460 genera and 3735 species in two subfamilies. The subfamily Encyrtinae includes 353 genera and 2920 species, while the Tetracneminae includes 107 genera and 815 species. Approximately half of all encyrtid species are associated with scale insects (Hemiptera: Coccoidea)(Noyes 2012). Encyrtids are generally endoparasitoids meaning that the parasitoid egg is laid directly inside the host’s body where the hatching larva completes development while feeding on host tissues, ultimately killing the host. Encyrtids mostly parasitize immature life stages (or, rarely, adults), but some species in one genus (Microterys) are egg predators (Noyes 2012). B. Description Tachardiaephagus somervillei was first described by Mahdihassan in 1923 as Lissencyrtus somervilli but Ferrière (1928) transferred L. somervilli Mahdihassan to Tachardiaephagus. He later downgraded this taxon from specific to subspecific rank as T. tachardiae somervilli (Ferrière 1935), based on the presence of intermediate forms and the lack of defining characters. This subspecies was accepted by Mahdihassan (1957). Varshney (1976) emended spelling of the subspecific rank to somervillei. In a recent reconsideration of the taxon, Hayat et al. (2010) elevated T. tachardiae var. somervillei to T. somervillei as a valid species. A detailed, formal taxonomic description of Tachardiaephagus somervillei does not exist. Although the specific name was erected by Mahdihassan (1923) he provided no accompanying morphological 1 description. In lieu of such a specific description, we first present a genus-level description based on the type species T. tachardiae (from Prinsloo 1977), and then note how T. somervillei differs from T. tachardiae. Tachardiaephagus Ashmead Tachardiaephagus Ashmead, 1904: 303; Ferriere, 1928: 171; 1935: 396. Type-species Encyrtus tachardiae Howard, 1896. Lissencyrtus Cameron, 1913: 99; Ferrière, 1928: 171. Type-species Lissencyrtus troupi Cameron, 1913. The genus Tachardiaephagus was described from the type species, T. tachardiae (Howard) from South and Southeast Asia. For synonymy of the genus and species, descriptions and figures, consult Ferrière (1928, 1935), Prinsloo (1977), and Noyes (2012). The genus comprises seven species; four described from sub-Saharan Africa (T. absonus, T. gracilus, T. similis, and T. communis - Prinsloo 1977), and three from South and Southeast Asia (T. tachardiae, T. somervillei, and T. sarawakensis - Ferrière 1928, 1935, Hayat et al. 2010). All species are primary parasitoids known only from kerriid hosts (Kerria - the lac insect of commerce, Paratachardina, and Tachardina species) (Prinsloo 1983). FEMALE. Primary parasitoids of Kerriidae. Medium-sized encyrtids, approximately 1.5-2.0 mm in length. Colour: head and body some shade of brown to black, the head and thoracic dorsum with or without faint to moderate metallic reflections; forewing hyaline or at most very faintly infuscated partly; hind wing hyaline. Head, in dorsal view (occiput perpendicular) with posterior margin moderately concave, the anterior margin convex, deeply notched by upper scrobal confluence (except in T. absonus); frontovertex united acutely or abruptly with occiput; frontovertex moderately broad, more or less than one-third head width at median ocellus; ocelli in an acute to obtuse-angled triangle; head, in frontal view, with toruli placed well above mouth margin, their upper limits approximately level with lower eye margins; scrobes strongly sulcate (except in T. absonus), long, their lateral margins sharply angled, confluent dorsally, impressed on face as an inverted 'V'. Antenna eleven-segmented; scape subcylindrical or at most moderately expanded ventrally, not less than three times as long as its greatest width; pedicel longer than basal funicle segment; funicle six-segmented, with all segments longer than wide or at most with distal one or two segments wider than long; club three-segmented, as long as or longer than the distal three funicle segments together, rounded apically, at most a little wider than funicle segment VI; maxillary palpi each with four segments, the labial with three; mandibles tridentate (sub-Saharan species) or with two teeth and a truncation (in T. tachardiae from South and Southeast Asia). Sculpture of head cellulate-reticulate, the cells relatively small, not raised, the frontovertex with fine setigerous punctations. Thorax moderately convex from side to side and anteroposteriorly, the mesoscutum plainly wider than long, the scutellum approximately as wide as long; mesoscutum without parapsidal sulci; mesoscutum with posterior margin overlapping mesal union of axillae partly or entirely, the latter not raised, separated medially by a narrow sulcus; sculpture of mesonotum cellulate-reticulate, the cells with very fine aciculations in some species; mesonotum moderately densely setose, the scutellum with one pair of suberect setae near apex. Legs not especially modified. Forewing 2 moderately broad, approximately 2.5 times as long as broad; venation well developed: submarginal vein slender; postmarginal vein usually longer than marginal, the former reaching to a level near apex of stigmal; wing disc evenly and rather densely setose from speculum to apex, the basal triangle never devoid of setae. Abdomen as long as or longer than thorax, the gaster usually heart-shaped, pointed or truncate at apex; cercal plates not strongly advanced towards base of gaster; gaster without paratergites; ovipositor varying in length, less than one-half to as long as gaster, as seen through the derm in cleared slide-mounted specimens; ovipositor protruding at most slightly caudally. MALE. Differing mainly from the female as follows: head and body generally black, with stronger metallic reflections (yellow variant in Oriental tachardiae). Head with frontovertex broader, approximately one-half head width at median ocellus; toruli placed higher on face, their lower margins about level with lower eye margins; ocelli in an obtuse-angled triangle. Antenna nine- segmented, the club not segmented, approximately as long as the distal two funicle segments together; flagellum with moderate to rather long, curved setae. Tachardiaephagus somervillei (Mahdihassan 1923)(Fig. 1.1, Fig. 1.3A) Lissencyrtus somervilli Mahdihassan, 1923: 71. Female. India, Karnataka. Tachardiaephagus somervilli (Mahdihassan): Ferrière, 1928: 71, taxonomy; Namkum (India) record. Tachardiaephagus tachardiae var. somervilli (Mahdihassan): Ferrière, 1935: 396, taxonomy; Malaysia record. Mahdihassan, 1957: 73-74. Tachardiaephagus tachardiae somervillei (Mahdihassan): Varshney, 1976: 61-62, emendation of specific name as it was based upon the name of Prof. Somerville. No detailed species description is available for T. somervillei. However, Hayat et al. (2010) treat T. somervillei as a valid species as it differs from T. tachardiae not only in the yellow to orange-yellow colour of the head, but also in having F6 or F5 and F6 quadrate to slightly broader than long; and in having third valvula 0.73x mid-tibial spur. (In T. tachardiae: F5 and F6 longer than broad; and third valvula subequal in length to mid-tibial spur). Ferrière (1935) indicated that he found intermediate forms in two females from West Malaysia] (Rawang and Kuala Selangor Road) in which "the head has the vertex, the temples and cheeks dark green like T. tachardiae, and the face and inner margin of eyes reddish, as in somervilli". He, therefore, downgraded the species somervillei to a variety of tachardiae. However, Hayat et al. (2010) reinstated the species-level status for T. somervillei after examining specimens from Thailand and Malaysia and failing to find any specimens intermediate in head colour to T. somervillei and T. tachardiae. C. Distribution, biology, behaviour and life cycle of the agent Tachardiaephagus somervillei (Hymenoptera: Encyrtidae) is a primary parasitoid of kerriid lac scale species, including the yellow lac scale Tachardina aurantiaca (Hemiptera: Kerriidae), the target species for biological control on the Australian external territory of Christmas Island. It is an 3 endoparasitic species, meaning that it uses an ovipositor to insert eggs within the body of its female lac hosts. These eggs hatch and the larvae slowly consume the lac insect as it develops, finally pupating and emerging as winged adults, which results in the death of the host lac insect. Unlike the vast majority of encyrtids, T. somervillei attacks the mature stages of its hosts. The genus Tachardiaephagus has been known for almost a century due to the pest status of T. tachardiae in lac scale cultures in India and parts of Southeast Asia. Lac is produced from the lac scale, Kerria lacca, and T. tachardiae and T. somervillei parasitize K. lacca, often causing economic injury (Narayanan 1962, Sharma 2006). Surprisingly, while the taxonomy
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