diversity, richness, and patterns of radiation among gall-inducing ...

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56 Oriental Insects Vol. 41corum Latr., induces organoid galls on the shoots of Juncus articulatus Linn. (Juncaceae)(Schmidt, 1966; Schmidt & Meyer, 1966) and incidence of this species exemplifies thewidth of host range of gall-inducing psyllids, more especially because extremely fewmonocotyledons host gall-inducing insects in general (Raman et al., 2005).Information on global species of gall-inducing psyllids is limited, when comparedwith the information available on gall-inducing Cecidomyiidae (Diptera) and Cynipidae(Hymenoptera). In this article, we have attempted to consolidate information on the diversityand richness of gall-inducing psyllids and their galls in the Orient and easternPalearctic, with specific examples from India and Taiwan, exploring some of the obviouspatterns of relationships among them.Diversity in gall-inducing psyllids in Eastern Palearctic and Oriental regionsInsects capable of inducing galls appear particularly rich in south-east Asia (Doctorsvan Leeuwen-Reijnvaan & Docters van Leeuwen, 1926), south Asia (Mani, 1973; 2000),and the Taiwanese bioregion (Yang & Tung, 1998). However, none of these cataloguesfocuses specifically on gall-inducing psyllids, except that of Yang et al. (2006), which hasassessed the diversity of gall-inducing psyllids and their galls in Taiwan. Information ongall-inducing psyllids remains either buried in the catalogues of gall-inducing insects referredearlier or scattered in papers dealing primarily with psyllid taxonomy and/or briefbiological details (e.g., Kandasamy & Krishnan, 1981; Kandasamy & Sharma, 1983;Kandasamy & Thenmozhi, 1985 a, b; Kandasamy, 1986; Raman & Kandasamy, 1978;Thenmozhi & Kandasamy, 1992, 1993). In an essentially taxonomic treatment of thepsyllids of the Indian subcontinent, Mathur (1975) has provided brief biological notes onsome of the gall-inducing psyllids. A comprehensive list of the psyllids of the Orientalzoogeographical region, with veiled references to the gall-inducing species, is availablein Hodkinson (1986).Taxonomic treatments of the psyllids from various provinces of China (Yang & Li,1982, 1983, Li, 1991, 1992a, b, c; 1993; Li & Huang, 1995; Wang & Wang, 2000; Li,2004) refer to gall-inducing species in the genera Cecidopsylla, Celtisaspis,Cornegenapsylla, Heterotrioza, Pauropsylla, Phacopteron, Pseudophacopteron, Rhinopsylla,Trioza, and Triozidus (Calophyidae, Psyllidae, Phacopteronidae, and Trozidae) inducinggalls on Schima argentea Pritz. (Theaceae), Celtis sinensis Pers. (Ulmaceae), Dimocarpuslongana Lour. (Sapindaceae), Chenopodium album Linn., (Chenopodiaceae),Ficus racemosa Linn. (Moraceae), Garuga sp. (Burseraceae), Alstonia scholaris (Linn.)R. Br. (Apocynaceae), Machilus vericolora S. Lee (Lauraceae), Cinnamomum austrosinesseH. T. Chang (Lauraceae), Acanthopanax brachypus Miq. (Araliaceae). Amongthese, species of Celtisaspis, reported only from the ‘China– Korea–Japan’ bioregion appearstriking because they share a disjunctive distribution with its relatives Tetragonocephalaand Pachypsylla, which are Nearctic genera (Crawford, 1914; Tuthill, 1943; Miyatake,1968, 1980; Kwon, 1983; Yang, 1995).Species of Celtisaspis, Tetragonocephala, and Pachypsylla form a monophyletictribe Pachypsyllini and feed mainly on the leaves of Celtis (Urticales: Ulmaceae) (White& Hodkinson, 1985; Yang, 1995). Species of Pachypsylla induce galls of diverse morphologieson leaves, buds, and twigs of several species of Celtis and are common inNorth America (Yang & Mitter, 1994). P. cohabitans Yang & Riemann was recentlyfound to behave as an inquiline, which on losing its ability in inducing galls, lives insidegalls induced by its sibling pachypsyllid species (Yang et al., 2001). Tetragonocephala, a
2007 Yang & Raman: Gall-inducing psyllids (Hemiptera: Psylloidea) 57monotypic genus that forms lerps, is less common and confined only to the south ofUnited States and north of Mexico. Lerp-forming species occur also in the genusEuphalerus (Hollis & Martin, 1997) but are particularly diverse among the AustralianSpondyliaspidinae (Psyllidae) (White, 1971; Morgan, 1984; Carver et al., 1991; Hollis,2004). Most species of Celtisaspis not only induce conspicuous horn galls, but also formlerps at the same time (e.g., C. quizhouana Yang & Li, C. zhejiangana Yang & Li, C.liaoningana Yang & Li); they feed on the underside of host leaves, inducing pit galls onthe upper sides of leaves and cover their bodies with lerp along the undersides of leaves(Yang & Li, 1982). C. sinica Yang & Li and C. beijingana Yang & Li have also beenknown, which either induce pit galls or modest deformations on leaves besides forminglerps. Two species of Pachypsylla (= Celtisaspis) from Japan have been reported onCeltis sinensis var. japonica (Planch.) Nakai and C. bungeana var. jessoensis (Koidz.)(Miyatake, 1968, 1980; Takagi & Miyatake, 1993). The Japanese Pachypsylla have beentreated as Celtisaspis by Hodkinson (1986): C. japonica Miyatake is bivoltine, whereasC. usubai Miyatake is univoltine. C. japonica induces both dome-shaped lerps and horngalls only in their first generations, while the second generation offspring only formlerps. C. usubai can not only form lerps, but also induce pit galls. No inquilines or anyother kind of casual visitor species in the galls of Palearctic psyllids are known so far. Inthe monograph on Korean psyllids, describing 81 psyllid species of 18 genera, Kwon(1983) has briefly referred to Pachypsylla japonica Miyatake (=Celtisaspis japonica Miyatake)on Celtis sinensis var. japonica, but no mention of gall-inducing habit of either P.japonica or any other species has been provided.Further to Celtisaspis, close to 45 species of gall-inducing psyllids placed in 19 generaand three families have been recorded from Japan (Yukawa & Masuda, 1996): Anomoneura,Aphalara, Celtisaspis, Craspedolypta, Euphalerus, Togepsylla, Livia, Psylla,Syntomoza (Psyllidae), Metapsylla (Calophyidae), Epitrioza, Leptynoptera, Pauropsylla,Petalolyma, Stenopsylla, Trichochermes, and Trioza (Triozidae) (Yukawa & Masuda,1996). Plants belonging to 22 families host gall-inducing psyllids: Aquifoliaceae, Araliaceae,Chenopodiaceae, Ebenaceae, Elaeagnaceae, Euphorbiaceae, Fagaceae, Flacourtiaceae,Clusiaceae (= Guttiferae), Juncaceae, Lardizabalaceae, Lauraceae, Leguminosae,Meliaceae, Moraceae, Pittosoraceae, Polygonaceae, Rhamnaceae, Sapotaceae, Styracaceae,Symplocaceae, and Ulmaceae. A majority of host plants of Japanese gall-inducingpsyllids (19 out of 45) fall within Eurosids I, followed by Asterids and Eudicots (Table1). Two-thirds of Japanese gall-inducing psyllids belong to Triozidae and the seven Asteridsand three Euasterids-II host plants are associated with triozids. Only Metapsylla, thesingular calophyid of Japan is associated with a plant genus belonging to Eurosids-II.
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