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Hymenopterous Parasitoids Associated with Diamondback Moth

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Hymenopterous Parasitoids Associated with Diamondback Moth 25 Hymenopterous Parasitoids Associated with Diamondback Moth: the Taxonomic Dilemma Mike Fitton and Annette Walker¹ Natural History Museum, London and 'International Institute of Entomology, London, SW7 5BD, England Abstract Attempts to control diamondback moth Plutella xylostella (L.) using insect parasitoids have not been entirely successful. Parasitoids which have been utilized include Diadegma species and Cotesia plutellae. A better understanding of the systematics of these Hymenoptera could lead to their more effective exploitation in biological control. Diadegma is a very large and difficult genus of lchneumonidae. There are no completely satisfactory taxonomic treatments, and from the limited work that has been done we know that some distinct biological species are almost impossible to separate using traditional, morphological characters. Nine putative species of Diadegma attack diamondback moth. So far no studies have adequately considered the taxonomic questions which are important in relation to their parasitism of this widespread pest. The microgastrine braconid Cotesia plutellae has been used with limited success in controlling diamondback moth, but recent field studies have raised suspicions that it is a complex of two or more species. We present a review of our knowledge of Diadegma and Cotesia and other microgastrines associated with diamondback moth, and attempt to outline a strategy for solving the taxonomic problems, leading to a better understanding of relationships with this host. The other parasitoids which we consider reliably recorded from diamondback moth are also noted. Introduction This paper concentrates on Diadegma and Cotesia, but briefly touches on other hymenopterous parasitoids of diamondback moth (DBM), Plutella xylostella (L.) (Lepidoptera; Yponomeutidae). In each of these three sections, the current taxonomic situation, related questions of biology, and strategies for the future are considered. Diadegma Aspects of Diadegma Taxonomy Basic morphotaxonomy. The 200 valid, described species of Diadegma, probably make up less than half the real total of this large, worldwide genus. Its taxonomy is fraught with difficulties and there are very many misidentificationsin the literature. No completely satisfactory treatments (with keys, descriptions and summaries of biologies and distributions) exist, even for limited faunas. In Europe, where over 120 species are recognized, many additional, 'cryptic' species undoubtedly await detection. For example, the morphospecies known as D. chrysostictos has been shown by Horstmann and Shaw (1984) to comprise two species which attack different hosts in different habitats, but which are both so similar and so variable as to make impossible the reliable identification of many individual specimens. 225 226 Fitton and Walker The species associated with DBM. As far as the Diadegma species associated with DBM are concerned, the situation is particularly confused because poor communication, and nomenclatural and identification problems have added to the taxonomic difficulties. For example, as recently as 1974 and 1988 new species of Diadegma parasitizing DBM were described with no reference to D. semiclausum, of which they could well turn out to be synonyms. Although there are some published data, for example, in Venkatraman (1964), no studies convincingly link distinguishing morphological characters to demonstrably separate biological species. Even in Europe, where two species, D. semiclausum and D. fenestrale, have been recognized as parasitizing DBM, many questions remain unanswered. As long ago as 1938, Hardy reported the successful production of hybrids in the laboratory and suggested that interbreeding between these species might explain the occurrence in the field in England of ‘all stages of intermediate forms. ’ The host, DBM, is cosmopolitan, its geographical origin is uncertain and it could be attacked by ‘local’ Diadegma species as well as those which have had a long association with it. In both New Zealand (Muggeridge 1930) and South Africa (Ullyett 1947) supposedly native species of Diadegma were reported as parasitizing DBM before the introduction of Diadegma species from Europe. Traditional taxonomic studies have been hampered partly because numbers of specimens in collections are low. Voucher series relating to many of the transfers, introductions and laboratory studies are inadequate or even nonexistent. Much more material is needed for proper investigations. The present confused taxonomic situation. No-one has made a special study of the Diadegma species associated with DBM. The annotated list below summarizes the present situation: it is for guidance only and it is not meant to imply that there are nine separate biological species of Diadegma attacking DBM. The species are in alphabetical order and the currently accepted synonymy (indented) follows the valid species name. Diadegma fenestrale (Holmgren, 1860) gracilis (Gravenhorst, 1829) This European species probably has Cnephasia stephensiana (a tortricid which feeds on a variety of herbaceous plants) as its main host, but is able to move on to DBM in suitable situations. Available information suggests that it may not be able to sustain populations on DBM in the absence of its usual host. Lloyd (1942), for instance, found it difficult to get females to oviposit in DBM larvae. It may interbreed with D. semiclausum (Hardy 1938). Diadegma insulare (Cresson, 1865) polynesialis (Cameron, 1883) hellulae (Viereck, 1912) plutellae (Viereck, 1912) pygmaeus (Viereck, 1925) congregator (Walley, 1926) This New World species (Carlson 1979) is recorded from southern Canada south to Venezuela and west to Hawaii, and from the pyralid Hellula undalis and Plutella armoricae as well as DBM. Most of the synonymy shown above seems to be the result only of examination of dry pinned material. It needs to be critically reassessed. The wide geographical distribution and the early dates of description of some the synonymized species begs the question of the real number of species involved. Even if there is only one, potential for use in biological control may vary considerably between populations. D. plutellae has sometimes been recognized as Hymenopterous Parasitoids Taxonomy 227 a distinct species (for example, by Gupta 1987) but without any justification or characters for its separation being given. There have been few attempts to introduce D. insulare to new areas to control DBM. Diadegma leontiniae (Brèthes, 1923) This species was described from specimens reared from DBM in Argentina and has subsequently been recorded (also from reared material) from Uruguay. It may be the Diadegma species reported from DBM in Chile (Rojas, 1965). It may also be a synonym of insulare but (see discussion above) no-one seems to have considered that possibility. Diadegma rapi (Cameron, 1912) D. rapi from Australia is recognized easily because it lacks one small cross-vein in the forewing. There seems to be reproductive isolation of this species (Venkatraman 1964). There are no records of hosts other than P. xylostella, although no effort appears to have been made to find any. It has been moved within Australia to help control P. xylostella (Wilson 1960), but was outperformed by D. semiclausum. Diadegma semiclausum (Hellen, 1949) eucerophaga Horstmann, 1969 cerophaga - misidentification tibialis - misidentification This is the best-known of the Diadegma which parasitize DBM. After 1969 the name eucerophaga quickly came into general use, but in 1980 Horstmann synonymized it with semiclausum (a species described from the Azores) and unfortunately this name change has taken a long time to get into the literature relating to DBM. D. semiclausum seems originally to have had a Eurasian distribution but it has been widely introduced for biological control. However, the identity of material used for introductions requires further investigation (see below). Diadegma varuna Gupta, 1974 Described from India, compared in the original description to fenestrale, but not semiclausum. It may be a synonym of semiclausum. Diadegma xylostellae Kusigemati, 1988 Described from a single male from Nepal. Said to be very close to D. varuna. Diadegma sp. indet. A. lateralis - misidentification This is the species, supposedly native, which was found attacking DBM in New Zealand before the introduction of Diadegma species from England (Muggeridge 1930). Diadegma sp. indet. B. This is the species, supposedly native, which was found attacking DBM in South Africa before the introduction of Diadegma semiclausum from Europe (Ullyett 1947). Ullyett reported that the two species interbred. Biology of Diadegma Some aspects of the biology of Diadegma deserve attention because they are important in relation to both taxonomy and biological control. Their significance has been overlooked because of the confused taxonomic situation. 228 Fitton and Walker Host range. Data in the literature suggest that many Diadegma species have wide host ranges. For example, Hardy (1938) apparently accepted that D. fenestrale attacked 24 species of Lepidoptera and one coleopteran, describing it as ‘very polyphagous’. Although such lists do include accurate records, many of the records need verification and they are potentially extremely misleading (see, for example, Shaw 1981). Critical appraisal of what is now known of the patterns of host associations and the mechanisms involved (see, for example, Dijkerman 1990) leads us to conclude that most species of Diadegma are relatively host-specific.
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