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Oriental Insects, Vol. 41: 141–167, 2007.

BIOSYSTEMATICS AND BIOGEOGRAPHY OF

ORIENTAL CHALCIDOIDEA (HYMENOPTERA)

ASSOCIATED WITH PLANT GALLS

T. C. NARENDRAN, S. SANTHOSH & K. SUDHEER Systematic Entomology Laboratory, Department of Zoology,

University of Calicut, Kerala - 673 635, India Email: [email protected]; [email protected];

[email protected]

ABSTRACT. Chalcidoid wasps have exquisite life histories and diverse host relationships. They are believed to have originated in the upper Jurassic period. Among the chalcidoids it is not always easy to separate the obligatory gall inhabitants and those which have a discrete association with the galls. The fol-lowing three categories of chalcidoids are treated in the present article (i) gall inducers (ii) gall inquilines and (iii) parasitoids attacking the gall inducers and gall inquilines. Gall-inducing chalcidoids mostly belong to Agaonidae, Eulophidae, Eurytomidae, Pteromalidae, Tanaostigmatidae, and Torymidae. The families of Chalcidoidea associated with plant galls in widely varying degrees of dependence are Aphelinidae, Encyr-tidae, Trichogrammatidae, Eupelmidae, Mymaridae, Ormyridae, and Chalcididae. Several hypotheses ex-plain the phylogeny of Chalcidoidea. Morphological features tend to support the monophyletic origin of Chalcidoidea. However, no hypothesis is acceptable totally to a chalcidologist, since a comprehensive knowledge on the distribution of the character states throughout the superfamily is lacking. Gall induction arose more then once in the six families referred above. The biogeography of gall-associated Chalcidoidea is related to the distribution of their respective host plants. The Oriental chalcidoids are not isolated from the nearby Asiatic and Middle-eastern countries. Even the Palearctic and Ethiopian elements are quite fre-quently met with in the Oriental region.

Key words: Biosystematics, Biogeography, Chalcidoidea, Gall inducers, Gall inquilines, Gall associ-ates, Oriental region.

Introduction

Diverse groups of insects comprising an estimated 13,000 species induce plant galls (Shorthouse & Rohfritsch, 1992; Williams, 1994). Galls, the structures composed of plant tissue within which the insect feeds, represent distinct microhabitats that support special-ist inhabitants. Galls range in complexity from relatively open pits or folds to structures that completely enclose the inhabitants. The enclosed galls range from simple structures such as those induced by fig wasps (Agaonidae) to complex structures induced by gall wasps (Cynipidae) including those bearing extrafloral nectaries and newly developed hairs, spines, and resin canals. The ease with which galls can be collected, occupants ob-served and counted and their interactions inferred has made galls an important study sys-tem in diverse manners to understand co-evolution and population dynamics (Stone & Schönrogge, 2003).

Studies on the biosystematics and bioecology of gall-inducing (and gall associated) arthropods have long been progressing in different parts of the world. Unlike the Oriental region, their study has attracted greater attention in the Australian and Holarctic regions with the recent updates on their systematics, biogeography, and ecology. Contributions of Meyer and Maresquelle (1983), Meyer (1987), Shorthouse and Rohfritsch (1992) are monumental works on gall induction and gall morphogenesis, whereas the recently pub-

142 Oriental Insects Vol. 41

lished volume by Raman et al. (2005) relates to the biology, ecology, and evolution of gall-inducing insects. Although the classic works of Mani (1964; 1973; 2000) and Anan-thakrishnan (1984) deal largely on a global context, with the biology of gall insects, some information on the Oriental species of gall-associated Chalcidoidea can be obtained from these publications. Besides these and a few isolated papers, not much research has been carried out gall inducing and gall associated species of Chalcidoidea of the Oriental Re-gion. This has left a wide gap in our knowledge on chalcidoid-induced galls and gall-inducing chalcidoids from a region known for its extensive biological diversity.

Chalcidoid wasps (Hymenoptera: Chalcidoidea) or the chalcids are a fascinating group of insects, which show exquisite life histories and diverse types of host relation-ships. Chalcidoids represent about 22,000 species and more than 2000 genera in 19 fami-lies. In the Oriental region approximately 4200 species and 664 genera are so far known (Noyes, 2003). The Oriental region encompasses tropical Asia and associated islands, and includes an enormous diversity of gall-associated chalcidoids. Chalcidoids are micro-scopic hymenopterans, which are considered difficult for taxonomic studies. Their intri-cate host relationships and biologies include a major part associated with plant galls. Un-fortunately little is known on the biology and biogeography of Oriental chalcidoids, when compared with those from other faunal regions. However, their systematics is better known than the biology and biogeography. The work of Bouček (1988) dealing with 14 families of Chalcidoidea with keys, illustrations, and descriptions and the electronic cata-logue by Noyes (2003) have laid the foundation for reliable studies on the biosystematics of chalcidoids. Among the chalcidoids, it is difficult to separate the obligatory gall in-habitants and those which have a discrete association with galls. In the present article, we have treated chalcidoids associated with plant galls (gall-associated species) in the fol-lowing three categories: (i) gall inducers, (ii) inquilines associated with plant galls, and (iii) parasitoids attacking either gall inducers or inquilines. Categories of ‘gall inducers’ and ‘parasitoids of gall-inhabiting insects’ have been discussed in greater detail than the category of ‘inquilines associated with plant galls’, because of lack of published informa-tion on the inquilinous species in the Orient. Wherever the precise nature of the associa-tion between the chalcidoid and the gall is not explicit, we have referred to their occur-rence in galls based on available literature. The purpose of this article is to review briefly all available recent (as well as a few important old ones) works on the biosystematics and biogeography of chalcidoids associated with plant galls from the Oriental Region. [Be-cause of the difficulty in dealing with the known hundreds of chalcidoid taxa associated with plant galls in this brief review, we have enumerated them in Table II.]

Gall-inducing chalcidoids fall under six families: Agaonidae, Eulophidae, Eurytomi-dae, Pteromalidae, Tanaostigmatidae, and Torymidae (La Salle, 2005). The families of Chalcidoidea associated with plant galls in widely varying degrees of dependence are Aphelinidae, Encyrtidae, Trichogrammatidae, Eupelmidae, Mymaridae, Ormyridae, and Chalcididae (Narendran, 1984). Generally it is difficult to distinguish the gall inducers from the inquilines and parasitoids associated with a gall, which necessitates the verifica-tion of the mode of nutrition of the inhabitants within galls.

Each of the 13 families of chalcidoids associated either directly or indirectly with plant galls is treated as follows: (1) broad systematics of the family; (2) nature of associa-tion of chalcidoids with plant galls (gall inducers or gall inquilines or parasitoids of gall inhabitants). Besides these, other relevant aspects such as pest status and biogeography

2007 Narendran et al.: Chalcidoidea associated with plant galls 143

have also been discussed in the subsequent sections. Nonetheless, wherever the available information is extremely limited, all available information has been summarized.

Phylogeny and evolutionary trends

Several hypotheses explain the phylogeny of Chalcidoidea and some of the notewor-thy ones are those proposed by Gordh (1975), Rasnitsyn (1980), Ramirez (1991), Ron-quist (1994), Gibson (1986; 1990; 1999), and Gibson et al. (1999). Before Rasnitsyn pub-lished his hypothesis in 1988, Chalcidoidea was considered closely related to Cynipoidea. Rasnitsyn hypothesized that Chalcidoidea, Platygastroidea, Serphitidae and the ancient Jurassic family Jurapriidae form a monophyletic group. Recently Gibson (1999) proposed that Platygastroidea form a monophyletic group with ‘Pelecinidae+Proctotrupidae +Vanhorniidae’ and that of ‘Chalcidoidea+Mymarommatoidea’ is likely a relatively a basal clade in Apocrita with uncertain sister group relationship.

Evolutionary trends among gall-associated species are not well known because of the lack of knowledge in their biology and their relationships. Gall-inducing capability among insects has evolved repeatedly in different insect orders (Shorthouse & Rohfritsch, 1992; Williams, 1994; Raman et al., 2005) resulting in species-rich lineages with fossil records extending back at least 300 mya (Crespi et al., 1998; Machado et al., 2001). Ap-parently phytophagy originated independently among different groups and at different times during the phylogenetic history. Gall-inducing capability arose many times in Chal-cidoidea, at least at 15 different times, seemingly more frequently than in other main groups (La Salle, 2005). All phytophagous taxa in Chalcidoidea have parasitoid progeni-tors. While it is probable that most gall inducers came from progenitors that are parasi-toids of gall inducers, the remaining gall inducers of Chalcidoidea probably arose from seed or ovule feeders and from stem borers (La Salle, 2005).

The origin of Chalcidoidea may be in the upper Jurassic (Roskam, 1992). Plant feed-ing chalcidoids belong to the seed- or ovule-feeding and gall-inducing guilds. Phyto-phagy in Chalcidoidea occurs mainly in the primitive families, Torymidae, Eurytomidae, and Agaonidae (Roskam, 1992). In the evolution of phytophagy in Chalcidoidea, the parasitoid species of Torymidae and Eurytomidae could have first attacked immature host larvae and laid eggs on them (Askew, 1975). Unlike other parasitoids, probably because they were unable to delay their larval development, to compensate insufficient food source, they commenced feeding on plant tissues including gall tissue. Since the fig tree is entirely dependent on fig wasps for pollination, it is inferred from the distribution data that the Agaonidae which belongs to the oldest known gall insects might have originated in Cretaceous period (Roskam, 1992). Gall-inducing Perilampidae and Eulophidae appear to be ancient forms, based on their tropical distribution and old age of the host plants (Ni-kolskaya, 1956).

Gall-inducing capability arose more than once in Agaonidae, Eulophidae, Eurytomi-dae, Pteromalidae, Tanaostigmatidae and Torymidae (Kjellberg et al., 2005). In Agaoni-dae (sensu Bouček, 1988), gall induction in the fig receptacles are mostly found in taxa belonging to Agaoninae. However the phylogeny of the family Agaonidae is not yet final-ized. Based on molecular evidences, Rasplus et al. (1998) have questioned the classifica-tion of Bouček (1988) and Gibson (1993). Rasplus et al. (1998) reverted Agaoninae to Agaonidae and reassigned other subfamilies (except Sycophaginae and Epichrysomalli-nae which are kept unplaced) to Pteromalidae.


In Eulophidae, Tetrastichinae includes most of the gall-inducing species. Tetrastichi-nae probably originated along with Entedoninae and Euderinae from some ancestor near the genus–group Aulogymnus Forster (Eulophinae) (Graham, 1987). The studies of Bug-bee (1936) provided the rationale for polarizing characters in Eurytomidae, but did not distinguish groups on explicit synapomorphies (Gibson 1999). Eurytomidae includes sev-eral gall inducers. So far no character, which is apomorphic, has been found in the sub-families of Eurytomidae to support monophyly of these subfamilies. Pteromalidae, which consists of several gall inducers, gall associates and other life styles, is the most difficult family to define either cladistically or phenetically (Boucek & Heydon 1997). The sub-family Ormocerinae, which also includes a majority of gall-inducing taxa, may be a mo-nophyletic group. Tanaostigmatidae is poorly represented in the Orient when compared with the Neotropical region. This family is considered to be a sister group of Encyrtidae (LaSalle 1987, Gibson 1999). Although Torymidae is well represented in Oriental region, gall inducers are hardly known. Both subfamilies of Torymidae are believed to be mono-phyletic (Grissell, 1995). Ormyridae is regarded as monophyletic and it is possible that Ormyridae renders Pteromalidae paraphyletic (Gibson, 1999). While many workers sug-gested Aphelinidae is paraphyletic with respect to either Signiphoridae or Trichogram-matidae, monophyly of the family is also suggested by some (Gibson, 1986, Gibson et al., 1999). While Encyrtidae is believed to be monophyletic, Mymaridae is probably polyphyletic. The three segmented tarsi lend support to consider Trichogrammatidae mo-nophyletic. Gibson (1989) proposed that each of three subfamilies of Eupelmidae is mo-nophyletic. While Gibson (1999) suggested that Chalcididae could be paraphyletic with respect to Leucospidae, Wijesekara (1997) has postulated four apomorphic states to sup-port monophyly of Chalcididae.

Biogeography

The Oriental region encompasses tropical Asia and closely associated continental is-lands including Sri Lanka, Sumatra, Java, Borneo, and Taiwan (and the Philippines – ac-cording to some) (Darlington, 1957). The boundaries of the region are poorly defined, as there are complex transitions of the Oriental fauna in several directions. Towards the east, in the mountains of Burma (Myanmar) and south-west China and along the coast of China a broad complex transition zone exists, wherein boundaries are hard to recognize. Mori (1936) marked the border line at the Nan Shan (Nan Ling) mountains not far from the Tropic of Cancer. The Chinese provinces coming under the Oriental region are Fujian, Guangdong, Yunnan, Hainan, Guangxi, Sichuan (except eastern parts), Zenjiang (South), and Jiangxi (South). The main cause of differentiation of the Oriental fauna from the Palearctic is evidently the zonal climatic difference, but still the mountain ranges and dry country form a barrier to the north and to the west, respectively. Largely, south Asia is a challenging region due to its antiquity, unique plate tectonic and paleoclimatic history, location at the confluence of biogeographical realms, and astounding range of physiog-nomic and habitat diversity (Mani, 1974). These factors have contributed to the diversifi-cation of biota in the region (Mani, 1974; Fernando, 1984; Morley, 2000). Changes in the extents of habitats, in space and time, have also resulted in striking disjunctions in ranges of almost all taxonomic groups (Randhawa, 1945; Hora, 1949; Dilger, 1952; Fernando, 1984; Mani, 1974; Swan, 1993). Figure 1 shows the key physiognomic and ecological features that have been considered in previous biogeographical analyses of terrestrial wet forest biota. The South-Asian region, according to Biswas & Pawar (2006) contains a complex assortment of barriers and dispersal routes varying in their nature as well as spa-


tial scale to which multiple co-distributed taxonomic groups are unlikely to show a con-gruent response. Specificity of the gall associated Chalcidoidea on their associate plants points that distribution of specific plant species could be a matter of influence for their unique dispersal.

Fig. 1. Digital elevation model of South Asia highlighting features that are important in the previous biogeographical literature on terrestrial wet forest biota. (Dotted lines rep-resent potential aquatic barriers, dash-dotted lines terrestrial ones (dry zones), and bidi-rectional arrows potentially contiguous dispersal routes.) (Courtesy of Biswas & Pawar, 2006).


Eulophidae

Eulophidae is one of the large families of chalcidoids. It currently includes 297 gen-era and 4472 species (Noyes, 2003) arranged in five subfamilies. Among these, only two tribes, Ophelimini and Tetrastichinae include gall inducers. No Ophelmini with either di-rect or indirect association with plant galls are known in the Oriental region. Ophelimini occurs only in Australia and includes Ophelimus and Australsecodes (La Salle, 2005). Ophelimus is probably the largest genus among gall-inducing eulophids and O. maskelli (Ashmead) has recently invaded the Mediterranean region and is now known from Israel, France and Italy where it induces small galls on the midribs of species of Eucalyptus (Protasov et al., 2006).

Potentially all the species of the subfamily Tetrastichinae associated with plant galls are either gall inducers or inquilines or parasitoids of other insects in the gall. Gall-inducing species of Tetrastichinae belong to Oncastichus, Quadrastichoidella, Quadras-

tichus, Epichrysocharis, Ceratoneura, Exurus, Paragaleopsomyia, Trichaporus, Goethella, Aprostocetus and Tetrastichus. Among these, Oncastichus, Trichaporus and Exurus are not known from Oriental region. Although the remainder is known to occur in the Oriental region, the role of the species of these genera, which induces galls in the Ori-ent is not conclusively known. However, among the gall-inducing tetrastichines the fol-lowing species are worthy of mention. Ceratoneura indi Girault is phytophagous and also induces galls on flowers and fruits of sweet pepper (Capsicum sp., Solanaceae) and brin-jal (egg plant, Solanum melongena Linn., Solanaceae) in India (Narendran & Krishnaku-mar, 1995). Infested flowers and fruits abscise prematurely from plant. This is the only species of Ceratoneura known from Oriental region. All the other species of Ceratoneura known from other zoogeographical regions are probably parasitoids of gall midges (Dip-tera: Cecidomyiidae) (La Salle, 2005). Another gall inducer of Tetrastichinae found in Oriental region is Quadrastichus erythrinae Kim which induces galls on the leaves, stems, petioles, and young shoots of Erythrina variegata Linn, E. fusca Lam and E. in-

dica Linn. (Fabaceae). This species was described from Singapore, Mauritius and Réun-ion (Kim et al., 2004), which is presently known to have spread into several countries in the eastern Palearctic and Pacific Islands (Yang et al., 2004; Heu et al., 2005). Our recent survey in the last three years in the southern Western Ghats revealed that this species has reached an invasive pest status in the region and probably in other parts of the peninsular India (Faizal et al., 2006).An invasive species Leptocybe invasa Fisher & La Salle (Mendel et al., 2004) which induces galls on the young foliage of Eucalyptus (Myrtaceae) in Australia, Middle East, and Africa has reached India (in Karnataka state; pers. obs.; in Tamilnadu and Pondicherry states; Jacob, 2007).

Several species of Eulophidae are known from the plant galls in the Oriental region but, as pointed out earlier, the nature of their association in the galls is not clear. The fol-lowing are a few examples of these gall-associated eulophids. An undescribed species of Aprostocetus and another undescribed species of Quadrastichus have been found to emerge from unidentified leaf galls of Mangifera indica Linn. (Anacardiaceae) (Naren-dran, Santhosh & Sudheer, unpublished observations). In the galls on the leaves of Ter-

minalia cuneata Roth. (= Terminalia arjuna Linn.) (Combretaceae) induced by Trioza

fletcheri minor (Crawford) (Hemiptera: Psylloidea: Triozidae), Tamarixia sheebae Nar-endran was found to emerge, particularly during rainy seasons in southern Malabar re-gion, India (Narendran et al., 2005). It is highly likely that this eulophid could be a para-site of T. f. minor. An undescribed species from the ‘leaf’ galls of Calycopterus floribunda


Lamk. (Combretaceae) (probably induced by Austrothrips cochinchinensis Karny (Thy-sanoptera: Phlaeothripidae) from the southern Western Ghats were obtained by us. Chry-

socharis echinata (Mani) has been reared from echinate galls (probably induced by an unknown Chalcidoidea) of Ficus benghalensis Linn. (Moraceae) (Mani, 1989; Narendran & Sudheer, 2005). Chrysocharis penthea (Walker) and Chrysocharis oscinidis Ashmead are associates of cynipid gall inducers (Shorthouse & Brooks 1998; Burks 1979) in the Nearctic are also known from Oriental region (Xu et al., 1999) on Liriomyza sativae Blanchard (Agromyzidae). Closterocerus pulcherrimus (Kerrich) was recorded in the galls of Procontarinia matteiana Kieffer & Cecconi (Cecidomyiidae) galls on leaves of mango in India and Sri Lanka (Bouček, 1986), which was known from psyllid induced leaf galls in Africa (Kerrich, 1970) from the unknown leaf galls of F. benghalensis Linn. (Narendran & Sudheer, 2005). Bouček (1986) recorded C. longiscapus (Boucek) from the leaf galls of Procontarinia matteiana from southern India.

Although Euderus lividus (Ashmead) has been recorded from the Oriental region, it has been found by Grimble et al. (1971) associated with galls of Saperda inornata (Col-eoptera: Cerambycidae) on trembling aspen, Populus tremuloides in the Nearctic region (Michigan, USA). Sigmophora brevicornis Panzer usually associated with Asphondylia

sarothamni H. Loew (Diptera: Cecidomyiidae) inducing galls on broom (Sarothamnus

scoparius (Linn.) Wimmer) is a commonly occurring species in the Republic of India and Pakistan (Parnell, 1964). Sigmophora aceris Ikeda was found associated with Asphondy-

lia sp. (Cecidomyiidae) induced galls on Acer mono (Aceraceae) (Ikeda, 1999a) in China, Taiwan and Nepal. An undetermined Indian species of Pediobopsis Girault has been re-corded from the galls on Prosopis cineraria (Linn.) Druce (Mimosaceae) and Salvadora sp. (Salvadoraceae) of unknown insect origin (Bouček, 1988). Narendran & Sheela (1994b) reported Neopediobopsis Narendran from the shoot-axis galls of Prosopis sp. from New Delhi, India. Two species of Quadrastichodella (a gall-inducing genus which is represented extensively in the Australian and Nearctic regions. Q. hirsuta Ikeda, and Q. gracilis Ikeda are known from Malaysia and Thailand respectively (Ikeda, 1999b).

Agaonidae

Classification of Agaonidae followed here is that of Bouček (1988). Species of the subfamily Agaoninae are true gall-inducing insects, developing in the ovaries of ‘gall flowers’ in the fig syconia. Many of the Agaonidae act as pollinators of various species of Ficus (Moraceae), whereas others are probably parasitoids of the pollinators or gall inducers on other parts in fig syconia. Fig wasps are specialized symbiotic pollinators of Ficus with life cycles closely tied to the flowering and fruiting cycles of fig syconia (Kjellberg et al., 2005). World Agaonidae currently includes 76 genera and 757 species under 6 subfamilies (Noyes, 2003). In the Oriental region 307 species in 46 genera and in India 105 species under 33 genera occur (Noyes 2003). Narendran (1984), Boucek (1988) and Grissell (1995) have discussed various aspects of phylogenetic relationships of the agaonids.

The current distribution of fig wasps suggests a Gondwanan origin of mutualism (Corner, 1958; Murrey, 1985). During the late Cretaceous period (ca. 100 myo), Africa had already separated from South America, becoming relatively an island. The land mass composed of South America, Antarctica, Australia and India–Madagascar remained linked for most of the late-Cretaceous period, with India linked to Antarctica through the Kerguelen Plateau until at least 80 myo (Hay et al., 1999). Fig wasp mutualism was es-


tablished during this period. (Machado et al., 2001).Continental Asia would have been colonized by Ficus and fig-wasps associated with them by the subgenera of Ficus group-2 (sensu Kjellberg et al. 2005) through India and much more recently may be also through Wallacea (Kjellberg et al., 2005).

Many species of Sycoryctinae, Sycoecinae, Sycophaginae, and Otitesellinae are ei-ther gall inducers in the non-reproductive parts of the fig plant or parasitoids of gall-inducing species. Some species of Epichrysomallinae also induce galls on florets of figs. Out of the male flowers, female flowers with short styles, and female flowers with long styles, agaonids usually develop in female flowers with short styles, which are known as 'gall flowers'. This is because the agaonids have short ovipositor which can reach the ovi-position site only through the short styles of the female flowers (Narendran, 1984). Parthenocarpic development of the ovule [endosperm] is stimulated to produce a gall, which becomes the larval food source.

Blastophaga psenes (Linn.) (Agaoninae) induces galls in the flowers of Ficus carica

(Moraceae) of India, and Pakistan. Larvae of Philotrypesis caricae Linn. feed on the en-dosperm of Ficus carica and eventually kill the larvae of Blastophaga psenes, which uses the same food source. Walkerella kurandensis Boucek (Otitesellinae), reported from India (Kerala) and Taiwan, is an associate of Ficus microcarpa and exemplifies phytophagy within this group. Narendran (in Narendran & Sheela, 1994a) described the genus Jose-

phiella Narendran with a single species, J. malabarensis Narendran, reared from leaf galls of Ficus benghalensis in India. This wasp was the first species of Epichrysomallinae that develops within leaf galls (Beardsly & Rasplus, 2001). This species is probably a gall inducer, since Josephiella microcarpae Beardsley & Rasplus described from the Nearctic region was found inducing galls on Ficus microcarpa Linn. (Beardsley & Ras-plus, 2001). J. microcarpae was found spread to as far as Macronesian (Canarias archi-pelago) region. J. microcarpae probably originated somewhere within the native range of F. microcarpa, in south-east Asia (Beardsley & Rasplus, 2001). Abdurahiman & Jo-seph (1976) recorded Ceratosolen solmsi (Mayr) in the galled fig syconia of Ficus his-

pida Linn. from India. Godfray (1988) studied the fig wasps and recorded a species of Sycoscapter as a gall associate from Papua New Guinea. Beardsley & Rasplus (2001) postulated that this wasp is a recent introduction into Hawaii, California, Canary Islands, and from an unknown location in south-east Asia, where Ficus microcarpa is a native.

Table 1 (Courtesy: Cook & Rasplus, 2003) Subfamily Larval ecology

Epichrysomallinae induce large galls Sycophaginae gall inducers Sycoecinae gall inducers Sycoryctinae inquilines/parasitoids

(biology poorly understood) Otitesellinae gall inducers Agaonidae gall inducers

Most chalcidoids that are not associated with figs are parasitoids, but the closest rela-tives of fig pollinators are not known (Bouček, 1988; Rasplus et al., 1998). The origins of chalcidoid non-pollinating fig wasps are also unresolved and their classification has changed many times, with six subfamilies recognized currently (see Table 1; after Cook


& Rasplus, 2003). A recent molecular study supports several independent colonization of the syconium habitat by different chalcidoid lineages (Rasplus et al., 1998). These in-clude long ovipositors to penetrate the syconium from outside and male aptery to facili-tate movement within the syconium. Even the key agaonid habit of entering the syconium has arisen independently in several parasite lineages such as in Sycophaginae, Sycorict-inae, and Epichrysomallinae.

Eurytomidae

World Eurytomidae currently includes 88 genera and 1424 species placed in three subfamilies: Eurytominae, Heimbrinae, and Rileyinae. Among them 276 species in 38 genera are known from Oriental region. . From India 184 species under 26 genera are re-ported (Noyes, 2003). Eurytominae is the largest subfamily that contains species, which exhibit a range of biologies and includes all the gall-inducing species in the family. The gall-inducing habit has arisen independently in several genera (La Salle, 2005). Eury-tomids have saw-like ovipositor used to introduce eggs into the shoot axes below the api-ces of various species of Poaceae.

Eurytoma is a speciose genus with remarkably varied biology, including organisms that are parasitic, inquilinous, phytophagous, entomophagous and gall inducing. Many of the species of Eurytoma occur as parasitic in galls. E. sabiae Mani parasitizing Acroecta-

sis campanulata (Mani) (Cecidomyiidae) in India, E. dentata Mayr parasitizing As-

phondylia spp. (Cecidomyiidae) in India and the Philippines, (Parnell, 1964) and E. sia-

mensis Zerova parasitizing Ormyrus flavipes in Thailand are reported as gall associates on Litchi chinensis Sonn. Mill. (Sapindaceae) from Oriental region. Some of the species of Tetramesa Walker are stem borers, while others are gall inducers, with obviously some gradation in between these two habits (La Salle, 2005). Claridge (1961) reviewed their biology and illustrations are produced by Claridge & Dawah (1994). Close to 200 species of Tetramesa are known so far. Among them a few species are reported from the Orient, but none is reported inducing galls and nothing on their biology known.

Bruchophagus sp. is reported as probable inquilines in the galls of Psectrosema spp. (Diptera: Cecidomyiidae) on Tamarix spp. (Tamaricaceae) (Habib, 1983) from Pakistan. Austrodecatoma, related to Bruchophagus, has three species, two of them A. omninigra Girault and A. trinotata (Girault) are Australian, which induce galls in the fruits of spe-cies of Atalantia (= Eremocitrus) (Rutaceae) (La Salle, 2005) and Austrodecatoma rufi-

gastra Narendran is reported from Tamil Nadu, India. Five species of Sycophila, viz., S.

decatomoides Walker, S. dharwarensis (Joseph & Abdurahiman), S. pilosa (Joseph & Abdurahiman), S. karnatakensis (Joseph & Abdurahiman), and S. robusta (Joseph & Ab-durahiman) have been recorded from galls in the syconial receptacles of F. benghalensis

Linn. (Narendran, 1994; Narendran & Sudheer, 2005). Bruchophagus rexus Narendran, associated with leaf galls of Ficus benghalensis Linn., is probably phytophagous similar to the other species of Bruchophagus (Narendran, 1994; Narendran & Sudheer, 2005). Eurytoma rosae Nees (Zerova, 1978; Boyd, 1999) known from galls is also found re-ported from Oriental region. The African species, Syceurytoma ficus Bouček found asso-ciated with Sycophila sp. on stem galls of Ficus spp. (Boucek, 1988) is reported from In-dia by Farooqi & Subba Rao (1986) without mentioning its hosts. Sycophila quercilanae (Fitch) is found associated with cynipid galls on Quercus prinus Linn. (Fagaceae) and Peck (1963) reported this species from Oriental region. Mangoma spinidorsum Subba


Rao has been reported from Procontarinia matteianna Kieffer et Cecconi on Mangifera indica (Subba Rao, 1986b; Narendran, 1994) from India.

Pteromalidae

Pteromalidae, one of the large families of Chalcidoidea, currently includes 588 gen-era and 3506 species placed in 31 subfamilies throughout the world (Noyes, 2003). From the Oriental region, 366 species in 114 genera are known and from India 176 species in 90 genera are so far recorded (Noyes, 2003). Although many subfamilies represent evolu-tionarily meaningful groups, the monophyly of the family is still doubtful (La Salle, 2005). Most of the known gall-inducing pteromalids belong to Melanosomellini (Ormo-cerinae) with 40 genera and 176 species.

The pteromalids associated with galls develop either as inquilines, or as parasitoids feeding externally on larvae, pupae or even adults of the gall inducer (Askew, 1961b). Austrosystasinae and Chromeurytominae are associated with galls on Eleocarpus sp. (Elaeocarpaceae), Eucalyptus (Myrtaceae), and Acacia (Mimosaceae) in Australia (Bouček, 1988). Trichilogaster Mayr induces hard woody galls on acacia in India and Australia. One species of Trichilogaster induces peculiarly large, spherical gall that con-sists of numerous larval chambers within a flattened cup-like depression (Mani, 1964). Mesopolobus includes a few species, which are known to induce galls in the Holarctic region, but the few species of Mesopolobus known from Oriental region are not known to be associated with plant galls. Although Notanisus versicolor Walker is known to have been reared from galls of at least two species of Tetramesa Walker, no gall-host record is available for this species from Oriental region (Bouček, 1988). Gastrancistrus cherryi Bouček is parasitic on Procontarinia matteiana that induce galls on leaves of Mangifera indica (Bouček, 1986). Pteromalus sequester Walker has been recorded from India with-out host record (Farooqi & Subba Rao, 1986b) but it is likely that this species may be associated with plant galls, since the same species is known to be associated with galls on Calicotome spinosa (Linn.) (Leguminosae) in England (Graham, 1969). The species Pachyneuron aphidis (Bouché) is known to be associated with galls (Gibson, 2001) and this species is reported from Oriental region also. Propicroscytus indicus Subba Rao is a species found in galls on Cynodon dactylon (Linn.) (Poaceae), Eleusine indica (Linn.) (Poaceae) (Bouček, 1988) and some grasses (Subba Rao, 1981). Systasis dalbergiae Mani was found parasitizing Contarinia dalbergiae Mani (Cecidomyiidae) in the galls on Dal-

bergia sissoo Roxb. (Dalbergieae) from India (Mani, 1942; Farooqi & Subba Rao, 1986b).

Tanaostigmatidae

Tanaostigmatidae currently includes nine genera and 92 species in the world and in the Oriental region five species in three genera are known (Noyes, 2003). The family is not divided into subfamilies. Tanaostigmatids are predominantly phytophagous in habit, usually either gall-inducers or inquilines in galls of other insects (La Salle, 2005). Pro-

tanaostigma derricola Ferriere has been reported from galls on Derris elliptica (Wallich) (Fabaceae) (Subba Rao, 1986a) in Indonesia. Another species Protanaostigma milletiae Ferriere is also reported from galls of the fruits of Milletia sericea Weight (Fabaceae) in Indonesia. In the catalogue of Chalcidoidea of India and adjacent countries (Subba Rao & Hayat, 1986) the host of the species Tanaostigmodes cajaninae La Salle has been re-ferred as ‘galls’ of pigeon pea Cajanus cajan (Linn.) (Fabaceae). However, La Salle (2005) states that strict seed infestation without gall induction has been reported from In-


dia, where T. cajaninae infests the seeds of pigeon pea as well as several other legumes (Lateef et al., 1985). Preferred host plants of tanaostigmatids are woody shrubs and trees of Fabaceae although other host plants are known (La Salle, 2005). In the Oriental region only members of Fabaceae are recorded as the host plants of tanaostigmatids. Chou & Huang (1994) and Yang et al. (2004) recorded T. lini and T. puerariae; the former without a host record from Taiwan and the latter as a gall-inducing species on Pueraria lobata (Fabaceae) in China.

Torymidae

This family contains 986 species placed in 68 genera in the world, and 149 species in 24 genera are reported from Oriental region (Noyes, 2003). Grissell (1995) discussed in detail the subfamily systematics of torymids and accordingly, the family includes Megas-tigminae and Toryminae.

Most of the known species of Toryminae are parasitoids of gall inducers. A few spe-cies of Torymus are known as gall inducers, which are not known to occur in the Oriental region. Megastigminae includes mainly either parasitoids, often on gall inducers, or phy-tophagous seed infesters and a few species may be inquilines, consuming the gall tissues and thus competing with the gall inducer (La Salle, 2005). In Toryminae, no reliable re-cords on gall-inducing megastigmines from the Oriental region are available. Man-

gostigmus amraeus (Kurian) and one other undescribed species of the same genus are parasitic on the cecidomyiid, Procontarinia echinogalliperda (Mani) and P. matteianna on Mangifera indica in India (Bouček, 1986; Narendran, 1994).

Ormyridae

The family currently includes three genera and 125 species in the world of which 45 species (in 3 genera) are from Oriental region. Most of the ormyrids are inhabitants of plant galls. A few (e.g. Ormyrus gopii [Narendran, 1999]) probably develop in stems of grasses or other plants. Narendran (1984) included ormyrids as a subfamily of Torymi-dae, while dealing with gall-associated chalcids. Later in the revision on Indo–Australian ormyrids, he treated them as a family (Ormyridae) as most recent authors do. Most spe-cies of Ormyridae are inhabitants of plant galls. Some may be partially phytophagous while others may be either entomophagous or phytophagous (Narendran, 1999). The ma-jority of Indo-Australian species are associated with fig-syconial galls induced by differ-ent Agaonidae. Twelve species of Ormyrus are known as associates in galls throughout the world, of which two of them reported from Oriental region. Zerova (1995) reported O. flavipes Bouček associated with Eurytoma siamensis on galls of Litchi chinensis Sonn. Mill. (Sapindaceae) in Thailand. The same species was reported by Bouček et al., (1981) from the syconial receptacles of Ficus thonningii Bl. O. sheelae Narendran and O.

zamoorini Narendran are associated with the receptacles of F. benghalensis Linn. (Nar-endran, 1999; Narendran & Sudheer, 2005).

Aphelinidae

Aphelinidae currently includes 33 genera and 1168 species placed in 7 subfamilies in the world. From Oriental region 416 species are known of which 210 species under 21 genera are recorded from India (Noyes, 2003). The only one gall-associated species from Oriental region is Aphelinus megadontus Hayat on Epipemphigus sp. (Hemiptera: Pem-phigidae) reported from Sikkim (Hayat 1998).


Encyrtidae

Encyrtidae currently includes 460 genera and 3735 species placed in two subfamilies in the world of which 813 species in 146 genera are from Oriental region and 520 species under 135 genera are from India (Noyes, 2003). Although 17 taxa are known to associate with plant galls , the exact nature of their association is not known. The only known spe-cies from Oriental region is Aloencyrtus indicus Singh & Prasad from the galls on Cas-

tanopsis hystrix A.D.C. (Fagaceae) (Singh & Prasad, 1996).

Eupelmidae

Eupelmidae includes 45 genera and 907 species placed in three subfamilies in the world, 131 species in 15 genera from Oriental region and 71 species under 13 genera from India (Noyes, 2003). Eupelmus Dalman, Brasema Cameron and Anastatus Motschulsky are the three genera known from galls, the first being the predominating gall associate. Eupelmus longicorpus Girault and E. urozonus Dalman are the two Eupelmidae species recorded from galls from Oriental region (Noyes, 2003). The Australian species, E. longicorpus, which is an associate in cecidomyiid galls on Andropogon sp. (Poaceae) (Bouček, 1988) is also known from India. Askew (1961a) reported E. urozonus as a para-site in cynipid oak galls. This species is present in India and other regions of Asia. Ac-cording to Bouček (1988) species of Neanastatus Girault are parasites of cecidomyiids especially of those species connected with grasses and galls on herbaceous plants. At least two species including Neanastatus cinctiventris Girault are found in India and south-east Asia and south China, which attack the Asian rice gall midge, Oresolia oryzae.

Trichogrammatidae

Trichogrammatidae currently includes 83 genera and 839 species in the world of which two species have so far been reported as gall associates. There are 326 species in 39 genera present in Oriental region of which 176 species in 90 genera are from India (Noyes, 2003). They are Eteroligosita tamaricis Viggiani and Thoreauia compressiven-

tris Girault, the former from Israel, and the latter from Australia. So far, no records exist pertaining to Oriental region.

Mymaridae

Mymaridae currently includes 103 genera and 1424 species in the world, 167 species in 27 genera from Oriental region and 104 species under 20 genera from India (Noyes, 2003). Although members of this family are almost exclusively egg parasitoids, Huber et al. (2006) found that two species of Stethynium develop as parasitoids of the gall inducers Ophelimus maskelli on eucalyptus in Australia, and that this biology might pertain to other species in this genus. Two other species have so far been recorded as gall associ-ates in this family. Kieffer (1902) recorded Camptoptera dryophantae Kieffer from the cynipid (Dryophanta folii Linn.) galls in Europe. Narayanella pilipes (Subba Rao) re-corded on Lagerstroemia speciosa Linn. (Lythraceae) galls from India and Myanmar (Subba Rao, 1976).

Chalcididae

Chalcididae currently includes 87 genera and 1464 species placed in 5 subfamilies in the world, 393 species in 35 genera from Oriental region and 207 species under 28 genera from India (Noyes, 2003). Only three species are reported from galls. Hockeria tamaricis Bouček was found associated with the gall-inducing lepidopteran Amblypalpis olivierella


Ragonot (Gelechiidae) on Tamarix sp. (Bouček, 1982; Narendran, 1986) from Pakistan, H. gallicola parasitic in the galls of Grandipalpa robusta Janse and Acuticornis (Gelechiidae) is known from Namibia and Haltichella onatas (Walker) from galls of Xan-

thoteras politum (Cynipidae) in USA.

Conclusion

The phylogenetic relationships of the superfamily Chalcidoidea have been studied by a few workers. Morphological features so far analyzed tend to support the monophyletic origin of Chalcidoidea. Although some chalcidologists studied the phylogenetic relation-ships of different families of Chalcidoidea, no clear concept exists. This is primarily be-cause of lack of sufficient knowledge on the distribution of various character states throughout the superfamily.

In the family Agaonidae, the concepts developed by Bouček (1988) and Gibson (1993) were unacceptable to Rasplus and others (Rasplus et al., 1998); Rasplus and oth-ers advanced their concept based on molecular studies. However we believe that further studies involving clear cut analyses are necessary to polarize character states and to bring out evolutionary trends of character transformation in this family.

In the family Eulophidae, the concept of Graham (1987) that Tetrastichinae probably originated along with Entedoninae and Euderinae seems convincing on the basis of re-semblances in the morphology of head, mesosoma and male genitalia. The morphology of the family Eurytomidae is also not yet clearly established and their phylogenetic rela-tionships are still uncertain. Pteromalidae is probably the most difficult family to evaluate the phylogenetic relationships, because this family does not have either a single character or a set of characters to separate it from the other families of Chalcidoidea. The concept that Tanaostigmatidae is a sister group of Encyrtidae poses some problems with regard to the inclusion of Cynipencyrtus in the family from Encyrtidae. The prepectus of Cynipencyrtus was hypothesized to be an intermediate between Tanaostigmatidae [sensu

strictu] and Encyrtidae, although it is not clear how the polarity of transformation is pos-tulated. In the family Torymidae, the assessment of synapomorphy with regard to wing venation goes well with Megastigminae but poses some difficulties in defining it in Toryminae since the nature of unmodified stigma of forewing is clearly a plesiomorphic state found in most Chalcidoidea. However it is possible to propose that although the stigma retains a plesiomorphic state, the wing veins as a unit are derived with respect to the outgroup Pteromalidae (Grissell, 1995). The monophyly of Ormyridae based on two synapomorphies of nature of gastral tergites and apically curved spurs of metatibia are questionable since the genus Eubeckerella Narendran deviates from these synapomor-phies (Narendran, 1999).

The biogeography of Oriental Chalcidoidea cannot be unequivocally established since the boundaries of Oriental region are not precisely defined. The distribution of gall- associated Chalcidoidea is related to the distribution of their host plants. The chalcidoid fauna of Oriental region is not isolated from other regions of Asia. Even the Palearctic and Ethiopian elements at generic level frequently occur in the Orient. This points to a need for comparative studies of fauna of the bordering areas of the Orient for a thorough knowledge on the biogeography of Oriental Chalcidoidea including those associated with plant galls.


Acknowledgements

We thank Drs Eric Grissell (Systematic Entomology Laboratory, USDA, Washing-ton, DC., USA), John La Salle, (CSIRO, Canberra, Australia), Anantanarayanan Raman (NSW, Australia), and Virendra Gupta (Gainesville, Florida, USA) for reviewing this ar-ticle. We express our sincere gratitude to Dr S. Biswas and the editor of Journal of Bio-

sciences (Bangalore) for permitting us to reproduce the digital elevation model of south Asia. . We also thank the authorities of the University of Calicut for facilities.

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Table II page 1. 8 page table here (160-167)

1

Table II. List of Oriental Chalcidoidea associated with Plant galls

Name of the Species

Nature of Habit

Taxonomic position

Primary Host Plant Host / Plant associate

Distribution References

Blastophaga psenes (Linn.)

inducer Agaonidae: Agaoninae

unknown Ficus carica (Moraceae) India, Pakistan Boucek, 1988

Ceratosolen solmsi

(Mayr)

inducer

Agaonidae: Agaoninae

Apocrypta bakeri ; Philotrypesis pilosa

(associates)

Ficus hispida

India [KAR, KER, WB], Indonesia [Java, Bali},

Malaysia, SriLanka, Vietnam

Abdulrazak & Abdurahiman,

2002

Eupristina verticillata Waterston

phytophagous/ inquiline

Agaonidae : Agaoninae

unknown Ficus aurea; Ficus microcarpa;

India [KER], Indonesia [Sumatra], China [Taiwan, Hong kong], Philippines,

Malaysia [Sarawak]

Chen et al.,1999

Josephiella malabarensis

Narendran

phytophagous/ Iiducer

Agaonidae : Epichrysomallinae

unknown Ficus benghalensis

India [KER]

Narendran & Sheela,

1994 Aphelinus

megadontus Hayat

associate Aphelinidae : Aphelininae

Epipemphigus sp. (Aphididae)

Populus sp. (salicaceae) India [Sikkim] Hayat, 1998

Hockeria tamaricis Boucek

Associate (parasitoid)

Chalcididae : Haltichellinae

Amblypalpis olivierella (Gelechiidae)

Tamarix sp. (Tamaricaceae) Pakistan Boucek,1982; Narendran,1986

Aloencyrtus indicus Singh & Prasad


Encyrtidae: Encyrtinae

unknown Castanopsis hystrix (Fagaceae)

India Singh & Prasad,1996

Trechnites secundus (Girault)


Encyrtidae: Encyrtinae

Trioza fletcheri (Triozidae); Unspecified

Psyllidae

Terminalia sp. (Combretaceae)

India [Bihar], Sri Lanka Hayat, 1986

Aprostocetus coimbatorensis

Rohwer


Eulophidae: Tetrastichinae

Unspecified Cecidomyiidae;

Stenodiplosis sorghicola (Cecidomyiidae)

Sorghum sp. Sorghum bicolor (Poaceae)

India [AP,TN] Rohwer, 1921; Husain &

Khan,1986

Aprostocetus ilexi Sheng & Zhao

associate (inquiline)


unspecified Cecidomyiidae (cecidomyiid)

Ilex cornuta (Aquifoliaceae) China [Jiangxi]

Sheng & Zhao,1995

2

Aprostocetus psyllidis Khan &

Shafee



Unspecified Psyllidae Grewia asiatica (Tiliaceae)[Leaf galls]

India [ UP] Khan & Shfee, 1981

Aprostocetus sankarani Boucek



Procontarinia sp. (Cecidomyiidae);

Cynipidae

Mangifera indica (Anacardiaceae)

India [UP, KAR] Boucek,1986

* Ceranisus menes

(Walker)

Associate

(parasitoid)

Eulophidae: Entedoninae

Callirhytis glandium (Cynipidae) (galls in

Palearctic)

unknown

India[TN,KAR, UP], Indonesia, Malaysia, Sri

Lanka, Philippines, Taiwan, Thailand

Boucek & Askew, 1968

Ceratoneura indi

Girault

inducer/ inquiline/ parasitoid

Eulophidae:

Tetrastichinae

Unspecified

Cecidomyiidae;

Capsicum annuum; Capsicum sp.; Solanum melongena (Solanaceae)

(Flower bud galls)

India[TN,KAR, MP,MR]; Malaysia [Sabah,

Sarawak]China [Hong Kong], Sri Lanka

Boucek,1988; Narendran &

Krishnakumar, 1995; Husain &

Khan, 1986 Chrysocharis echinata (Mani)



unknown Ficus benghalensis L. (echinate galls)

India [TN] Mani, 1989; Narendran &

Sudheer, 2005

* Chrysocharis

oscinidis Ashmead



Liriomyza sativae (Agromyzidae) in China [Reported from galls in

USA]

unknown China [Guangdong] Burks,1979; Xu et al., 1999

* Chrysocharis

pentheus (Walker)



Liriomyza sativae Blanchard (Agromyzidae)

in Oriental; Diplolepis rosaefolii (Cynipidae)

[Reported gall associate in Canada]

Rosa sp. 85 plants (Noyes,2003)

Malaysia, Taiwan, China [Guangdong,

Jiangxi]

Shorthouse & Brooks,1998

Chrysonotomyia obesula Boucek



Procontarinia sp. (Cecidomyiidae);

Cynipidae (leaf gall)


India [UP] Boucek,1986

Chrysonotomyia postmarginaloides

(Saraswat)



Procontarinia matteianna (Cecidomyiidae),

Unspecified Psyllidae

Mangifera indica (Anacardiaceae) Grewia

asiatica (Tiliaceae)

India [KAR,TN,UP] Boucek,1986

3

Closterocerus longiscapus Boucek



Procontarinia matteianna (Cecidomyiidae)


India [TN, KAR] Boucek,1986

Closterocerus pulcherrimus

(Kerrich)



Procontarinia matteianna (Cecidomyiidae);

unspecified Psyllidae

Ficus benghalensis Linn; Mangifera indica

(Anacardiaceae) (leaf galls)

India [GU, UP, KAR, KER], Sri Lanka,

Pakistan

Kerrich, 1970; Narendran &

Sudheer, 2005

* Euderus lividus

(Ashmead)


Eulophidae: Euderinae

Melanagromyza obtusa (Malloch) in India; Saperda inornata (Cerambycidae)

(galls reported in USA)

Cajanus cajan (L.) in Oriental region; Populus

tremuloides in USA

India [KER,UP,DEL, Bihar,Haryana]

Singh,1992; Sebastian,1993; Grimble et al.,

1971

Goethella asulcata Girault

# inducer/

associate

Eulophidae: Tetreastichinae

unknown Capsicum annuum Solanaceae)

India [KAR,MP,MR] Boucek,1988

Leptoybe invasa Fisher & La Salle

iducer Eulophidae: Tetreastichinae

----------- Eucalyptus spp. India [KAR] Mendel et al., 2004

Neopediobopsis setosa Narendran



unknown Prosopis sp. (Fabaceae) India [DEL] Narendran & Sheela, 1994

Pediobopsis sp. Associate (parasitoid)


unknown Prosopis cineraria (Fabaceae), Salvadora sp.

(Salvadoraceae)

India [Rajasthan] Boucek, 1988; Parihar, 1994.

Quadrastchodella gracilis Ikeda

Iiducer # Eulophidae: Tetrastichinae

unknown Ulmus davidiana Thailand Ikeda, 1999b

Quadrastichus erythrinae Kim

inducer Eulophidae: Tetrastichinae

unknown Erythrina fusca Erythrina indica

Erythrina variegata (Fabaceae)

India [KER], Singapore Kim et al., 2004

Sigmophora aceris Ikeda


Eulophidae : Tetrastichinae

Asphondylia sp. (Cecidomyiidae)

Acer mono (Aceraceae) China [Fujian, Hunan]; Taiwan; Nepal

Ikeda,1999a

Sigmophora brevicornis (Panzer)


Eulophidae : Tetrastichinae

Asphondylia sarothamni H. Loew; unspecified

Cecidomyiidae

Sarothamnus scoparius (Linn.) (Leaf galls)

India [TN]; Pakistan Parnell,1964; Husain &

Khan,1986 Aprostocetus

longicauda (Kieffer) Associate

(parasitoid) Eulophidae:

Tetrastichinae Unspecified

Cecidomyiidae Artemisia sp. (Asteraceae)

India [TN,WB] Husain & Khan, 1986

4

Tetrastichus

tanjorensis Husain & Khan



Asphondylia riveae (Cecidomyiidae)

Unknown India [TN] Husain & Khan, 1986

Anastatus leithi (Walker)


Eupelmidae: Eupelminae

Unknown (blister galls on leaves)

Duranta sp. (Verbenaceae)

India [MR], Sri Lanka

Islam & Hayat, 1986

Eupelmus argentinotatus

Girault #


Eupelmidae Procontarinia matteianna (Cecidomyidae)


India [KAR] Boucek,1986

Eupelmus carinatus Kieffer



Unspecified Cecidomyiidae

Artemisia sp. (Asteraceae) India [WB] Islam & Hayat, 1986

Eupelmus longicorpus Girault




Andropogon sp. (Poaceae)

India Boucek, 1988

Eupelmus testaceiventris (Motschulsky)



Contarinia sorghicola; Procontarinia

matteiana;;Unspecified Cecidomyiidae

Oryza sativa; Hyparrhenia hirta (Poaceae)

India, Sri Lanka Herting,1978

Eupelmus urozonus Dalman



Psectrosema spp., Asphondylia sp.

(Cecidomyiidae) in Oriental; parasitoids of

Cynipidae in Holarctic & Palaearctic regions

Tamarix spp., (Tamaricaceae) in Oriental; Quercus spp.,and Quercus cerris in Holarctic &

Palaearctic regions

India [DEL,GU,WB, TN}, China, Pakistan

Askew,1961a; Habib,1983;

Chari & Patel,1974; Islam

& Hayat, 1986

Neanastatus indicus Shafee


Eupelmidae : Neanastatinae

Unspecified Psyllids (Triozidae, Psyllidae)

Grewia asiatica (Tiliaceae) India [UP] Shafee, 1973

Neanastatus pulchricorpus

(Girault)



Unknown Unknown India [TN] Islam & Hayat, 1986

Neanastatus trochantericus

Gahan




Unknown India [TN] Islam & Hayat, 1986

5

Eurytoma dentata Mayr


Eurytomidae : Eurytominae

Asphondylia sarothamni H. Loew ; Asphondylia

sesami (Cecidomyiidae)

Sarothamnus scoparius (Linn.) Sesamum indicum, Zea mays (gingly galls)

India [TN,UP]; Philippines

Farooqi & Subba Rao,1986a;

Parnell, 1964; Baltazar,1966

Eurytoma poroensis Mukerjee


Eurytomidae: Eurytominae

unknown Oryza sativa India [WB] Mukerjee, 1981

Eurytoma rosae Nees



Diplolepis rosae (Cecidomyiidae)

Rosa sp. Malaysia; China Boyd,1999; Zerova,1978

Eurytoma sabiae Mani



Acroectasis campanulata Sabia campanulata (stem galls)

India [HP] Mani, 1969; Farooqi & Subba

Rao,1986a Eurytoma setitibia

Gahan Associate

(parasitoid) Eurytomidae: Eurytominae

Orseolia oryzae, Trioza fletcheri; unspecified

Psyllidae

Oryza sativa, Cordia myxa & Trewia nudiflora

(leaf galls)

India [TN,AP,WB, Bihar], Indonesia [Java]

Farooqi, & Subba

Rao,1986a Prodecatoma

pongamiae Mani & Kurian



unknown Pongamia glabra (flower galls)

India [KAR] Narendran, 1994; Mani, & Kurian,1953

Prodecatoma sabiae Mani



Acroectasis campanulata Sabia campanulata India [HP] Mani, 1970

Mangoma spinidorsum SubbaRao





India Subba Rao,1986b;

Narendran, 1994

Syceurytoma ficus Boucek



Sycophila sp. Ficus burkei; Ficus sycomorus (stem galls)

India Boucek, et al.. 1981

Sycophila inquilina Mohendro & Prasad

Associate (inquiline)


unknown

Castanopsis hystrix (Fagaceae)

India [Manipur] Mohendro & Prasad, 2004

Sycophila floribundae Narendran



Unspecified Psyllidae (leaf galls)

Calycopteris floribunda India [KER] Narendran, 1994

Sycophila quercilanae (Fitch)



Acraspis spp., Callirhytis spp., Neuroterus spp.

(Cynipidae)

Quercus prinus (Fagaceae) India [DEL] Peck,1963

6

Systole calycopterae Narendran



Unspecified Psyllidae (leaf galls)

Calycopteris floribunda India [KER] Narendran, 1994

Narayanella pilipes (Subba Rao)


Mymaridae unknown Lagerstroemia flosreginae (Lythraceae)

India [WB], Myanmar Subba Rao, 1976

Ormyrulus gibbus Boucek


Ormyridae Procontarinia matteianna (Cecidomyiidae)


India {AP,UP,KAR] Boucek,1986

Ormyrus calycopteridis

Narendran


Ormyridae Austrothrips cochinchinensis (Phlaeothripidae)

Calycopteris floribunda (Combretaceae)

India [KER] Narendran, 1999

* Ormyrus flavipes

Boucek


Ormyridae Eurytoma siamense Zerova; Syceurytoma ficus (associate only);

[unspecified Cynipidae & Syceurytoma ficus (In

Zimbabwe)]

Litchi chinensis (Sapindaceae)

Ficus burkei (Moraceae)

Thailand Boucek et. al., 1981

Ormyrus gopii Narendran


Ormyridae Unidentified stem galls Unidentified plant India [KER] Narendran, 1999; Xu et al,

2002 Ormyrus pomaceus

(Geoffroy) Associate

(parasitoid) Ormyridae Andricus sp., Biorhiza sp.,

Cynipidae (galls) Phyllostachys sp. (bamboo)

(Poaceae) China [Zhejiang] Narendran,

1999; Xu et al., 2002

Gastrancistrus (Mangistrus) cherryi

Boucek


Pteromalidae : Pireninae



India [KAR] Boucek,1986

Gastrancistrus muneswari Yadav


Pteromalidae : Pireninae

Oligotrophus mangiferae (Kieffer); Cecidomyiidae

Mangifera indica (Anacardiaceae) (galls)

India [Bihar] Boucek,1986; Subba Rao,1981

Notanisus versicolor Walker


Pteromalidae: Cleonyminae

Unspecified Aphididae; Tetramesa sp. ( Eurytomidae)

Gramineae (grass stem) India [Bihar, TN] Boucek,1988

Pachyneuron aphidis

(Bouché)

Associate

(parasitoid)

Pteromalidae: Pteromalinae

Aphididae (galls) in

Australia

Sonchus oleraceus; Sisymbrium officinale; Gossypium; Triticum

aestivum

India [HP,KAR,DEL,JK,Harya

na,PB,UP]; Pakistan

Gibson,2001; Sureshan, 2003

7

Propicroscytus indicus Subba Rao


Pteromalidae Unspecified Cynipidae (galls)

Cynodon dactylon India {MR,PB] Subba Rao, 1981; Farooqi &

Subba Rao,1986b

* Pteromalus

sequester Walker

Associate

(parasitoid)

Pteromalidae: Pteromalinae

Apion fuscirostre; Dinarmus italicus;

Asphondylia sarothamni; Apion cerdo;

Bruchophagus (in England)

Sarothamnus scoparius; Calicotome spinosa (in

England)

India [DEL,PB]

Graham,1969

Systasis dalbergiae Mani


Pteromalidae Contarinia dalbergiae Dalbergia sissoo India [DEL,UP] Mani,1942; Farooqi&Subba

Rao, 1986b Protanaostigma derricola Ferrière


Tanaostigmatidae Unknown Derris elliptica (Fabaceae) Indonesia [Sumatra] Ferrière, 1932; Subba Rao,

1986a Protanaostigma milletiae Ferrière


Tanaostigmatidae Unknown (fruit galls) Millettia sericea (Fabaceae) Indonesia [Java, Bali] Ferrière, 1932; Subba Rao,

1986a Tanaostigmodes cajaninae LaSalle


Tanaostigmatidae Unspecified cynipidae Cajanus cajan (Fabaceae) India [AP,Orissa,TN] Boucek, 1988; Lateef et al, 1985

Mangostigmus amraeus (Chandy

Kurian)


Torymidae: Megastigminae

Amradiplosis echinogalliperda Mani, Pachydiplosis oryzae;

Procontarinia echinogalliperda;

Procontarinia matteiana (Cecidomyiidae)


India [UP], Malaysia, Thailand, Brunei

Chandy Kurian, 1953;

Boucek,1986; Farooqi,1986

Megastigmus dorsalis (Fabricius)

phytophagous/ inquiline

Torymidae: Megastigminae

Andricus sp.,Cynips sp. (Cynipidae); Syntomaspis

sp. (Torymidae)

Quercus spp.; Quercus semecarpifolia; Quercus

suber (associate)

India [UP] Farooqi,1986; Mathur,1962

Megastigmus viggianii Narendran

& Sureshan


Torymidae : Megastigminae

unknown Calycopteris floribunda (leaf bud galls)

India [KER] Narendran & Sureshan, 1988

8

Pseudotorymus indicus (Mani)


Torymidae Unknown (flower bud galls)

Dalbergia sissoo India [UP,KER,TN] Farooqi,1986; Mani, 1942

Torymoides kiesenwetteri (Mayr)


Torymidae: Toryminae

Cynipidae (galls) Eupatorium glandulosum (galls)

India [KER], Nepal Grissell, 1995; Narendran, 1994

* Torymus

calcaratus Nees


Torymidae: Toryminae:

Ectemnius rubicola; Adleria quercustozae; Ectemnius rubicola; Pemphredon sp.;

Andricus kollari ; Stigmus solskyi; Cynips kollari (Cynipidae) in Europe

Quercus sp.; Rubus sp.

India [KER] Grissell,1995; Narendran et al.,

2005

Table II. KAR: Karnataka; KER: Kerala; WB: West Bengal; AP: Andhra Praesh; TN: Tamil Nadu; UP: Uttar Pradesh;MP: Madhya Pradesh; MR: Maharashtra; GU: Gujarat; DEL: Delhi; HP: Himachal Pradesh; JK: Jammu and Kashmir; PB: Punjab. # Need to be verified

* Reported from Oriental region but reported as gall associate some where outside

biosystematics and biogeography of · pdf fileoriental insects, vol. 41: 141–167, 2007....

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