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

Oriental Insects, Vol. 41: 55–65, 2007.DIVERSITY, RICHNESS, AND PATTERNS OF RADIATIONAMONG GALL-INDUCING PSYLLIDS (HEMIPTERA:PSYLLOIDEA) IN THE ORIENT AND EASTERN PALEARCTICMAN-MIAO YANG 1 & ANANTANARAYANAN RAMAN 21 Department of Entomology, National Chung Hsing University, Taichung 40227, TaiwanE-mail: mmyang@nchu.edu.tw2 Charles Sturt University, PO Box 883, Orange, NSW 2800, AustraliaE-mail: araman@csu.edu.auABSTRACT. Psyllids, or jumping plant lice, which induce diverse and exquisite galls – from simpleleaf-margin rolls to complex two-tier galls – in the Orient and eastern Palearctic, have not been comprehensivelydocumented. Considering their richness and diversity, this article consolidates and discusses patternsof congruence and radiation among the psyllids of the two biogeographical regions.Key words: Diversity, Eastern Palearctic, Galls, Host plants, Orient, Psyllids.IntroductionPsyllids or the jumping plant lice (Sternorrhyncha) are highly host-specific, suckinginsects. Similar to their allies, the aphids and coccids, many psyllids induce galls onleaves of higher-vascular plants (Hodkinson, 1984; Burckhardt, 2005). Galls induced bypsyllids range from simple rolls (e.g., on the leaves of Strychnos nux-vomica Linn. [Loganiaceae]by Diaphorina truncata Crawford [Psyllidae: Diaphorininae], Balakrishna &Raman, 1992) to complex two-tier galls (e.g., on the foliage of Garuga pinnata Roxb.(Burseraceae) by Phacopteron lentiginosum Buckton [Phacopteronidae], Raman, 1987).Until date no gall-inducing psyllid has been reported to live on either gymnosperms orferns. Few psyllids are known to infest non-leaf organs of plants and induce telescopedstems and crinkled foliage (e.g., Euphalerus vittatus Crawford [Psyllidae: Euphalerinae]on inflorescence axes and shoot meristems of Cassia fistula Linn. [Caesalpiniaceae];Mathur 1975); whether the malformations, such as those induced by E. vittatus are to betermed galls is a matter for future debate.The highly specialized mode of feeding behavior by reaching specific points inleaves (Balakrishna & Raman, 1992; Raman, 1987; 1991; Raman et al., 1997) has contributedto the multiple origins of gall-inducing groups not only in Psylloidea (Hodkinson,1984), but also in the Hemiptera (Roskam, 1992). Gall-inducing taxa in Psylloideaoccur in all of the currently recognized families, with maximal number of species occurringwithin Triozidae, Phacopteronidae, and Calophyidae (Burckhardt, 2005). Woodyplant taxa belonging to Magnoliales, Myrtales, Rutales, Malvales, and Urticales host gallinducingspecies of Triozidae, Phacopteronidae, and Calophyidae (Burckhardt, 2005),whereas the predominantly herbaceous taxa belonging to Asteraceae and Ranunculaceaehost those of Aphalarinae and a few of Triozidae (White & Hodkinson, 1985; Burckhardt& Laurterer, 1989; Burckhardt & Mifsud, 2003). Species of Livia (Psyllidae: Liviinae)(Hodkinson, 1986) are associated with monocotyledons, particularly with species of Juncaceaeand Cyperaceae (White & Hodkinson, 1985; Hodkinson & Bird, 2000). Livia jun-

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.

58 Oriental Insects Vol. 41Table 1. Gall-inducing Psyllid families and their hosts in Japan 1Psyllid familyPlant group 2 Calophyidae Psyllidae TriozidaeTotalMagnoliids - 1 3 4Monocots - 1 - 1Eudicots - 4 2 6Rosids - - - -Eurosid I - 7 12 19Eurosid II 1 - - 1Euasterids II - - - -Euasterids II - - 3 3Asterids - - 7 7Unknown - 1 3 4Total 1 14 30 451 Adapted from Yukawa & Masuda, 1996.2 Phylogenetic categories of plant families are in accordance with Angiosperm Phylogeny Group (2003).Richness and diversity of gall-inducing psyllidsin the Indian subcontinent and TaiwanTaiwanNearly two decades ago, 19 gall-inducing psyllid species were recorded by Yang(1984). In this record, Triozidae included 17 species placed in Cecidotrioza, Eustenopsylla,Homotrioza, Neophacopteraon, Parastenopsylla, Pseudotrioza, and Trioza,Phacopteronidae included one species of Pseudophacopteron, and Psyllidae included onespecies of Psylla (Yang, 1984). These species have been reported to be inducing galls onleaves (e.g., pit galls, leaf folds, leaf rolls, marginal folds, spherical protruding galls) ofspecies of Daphniphyllaceae, Burseraceae, Theaceae, Lauraceae, Symplocaceae, Sapindaceae,Ericaceae, Euphorbiaceae, Pittosporaceae, Flacourtiaceae, Elaeocarpaceae, andMyrtaceae. A recent survey (Yang et al., 2006) has revealed a more extensive range ofplant families that host gall-inducing psyllids. In addition to those reported by Yang(1984), the Yang et al. (2006) report refers to the following plant families that host gallinducingpsyllids: Lardizabalacaeae, Sabiaceae, Proteaceae, Clusiaceae, Pittosporaceae,Fabaceae, Fagaceae, Moraceae, Ulmaceae, Meliaceae, Actinidiaceae, Ebenaceae, Sapotaceae,Styracaceae, Acathaceae, Aquifoliaceae, Araliaceae, Pittosporaceae, Asteraceae, andCaprifoliaceae, with 75% of the recorded galls induced on leaves, and the remainder oneither shoot axes (17.5 %) or flowers and fruits (7.5%). The Yang et al. (2006) report,however, clarifies that Lauraceae (e.g., Machilus, Cinnamomum) and Moraceae (e.g., Ficus)hosted the maximum number of gall-inducing psyllids. Notwithstanding the fact thatYang et al. (2006) have not listed the psyllid taxa, a comparison of Yang (1984) and Yanget al. (2006) reports reveals the following (Table 2):• a majority of gall-inducing psyllids occur on the relatively primitive tree speciesbelonging to Lauraceae (Magnoliids: Laurales) and Myrtaceae (Rosids: Myrtales)

2007 Yang & Raman: Gall-inducing psyllids (Hemiptera: Psylloidea) 59• recently recorded psyllids and their galls occur on trees of relatively advancedplant families belonging to Eurosids I and Euroisids II• psyllids and their galls have also been recorded on advanced plant families thatinclude predominantly herbaceous species belonging to Asterids, Euasterids I, andEuasterids II• psyllid species that are capable of inducing galls on shoot axes (stems and buds)also occur.Table 2. Gall-inducing Psyllid families and their hosts in Taiwan 1Plant group 2 Host species Gall types 3Magnoliids 25 30Monocots - -Eudicots 7 7Rosids 10 13Eurosid I 24 24Eurosid II 2 2Euasterids I 1 1Euasterids II 8 8Asterids 12 13Total 89 981 Adapted from Yang et al. (2006).2 Phylogenetic categories of plant families are in accordance with Angiosperm Phylogeny Group (2003).3 Shape of galls on each plant species and plant part has been considered as one gall type. As psyllids arehost specific, each gall type is likely to represent a specific gall-inducing species.IndiaA majority of the psyllids that induce galls (e.g., pit galls, leaf folds, leaf rolls, marginalfolds, round-protruding galls) belong to Psyllidae (e.g., Diaphorina [Diaphorininae],Livia [Liviinae], Paurocephala [Paurocephalinae]), Calophyidae (e.g., Apsylla, Calophya),Phacopterinidae (Phacopteron), and Triozidae (e.g., Cecidotrioza, Ceropsylla,Cryptotrioza, Egeirotrioza, Neotrioza, Pauropsylla, Pseudotrioza, Trioza). Nearly all theknown species of the Oriental Triozidae are gall-inducing species (e.g., Mathur, 1975;Kandasamy, 1986) (Table 3). Indotrioza hirsuta Kandasamy, a gall-inducing species onthe leaves of Vaccinium neilgherrensis (Ericaceae) from the semi-dry vegetation alongthe eastern slopes of the Eastern Ghats (The Shevroys; 1650 m asl; 11°40′N; 78°8′E) andCeropsylla indica Kandasamy, another gall-inducing species on the leaves of Terminaliaarjuna Wight & Arnold (Combretacae) from the coastal plains of peninsular India (Madrascity; 13°3″N; 80°13′E) have been added in the mid-1980s (Kandasamy, 1986). IndianTriozidae are, by and large, restricted to arborescent species confined to Lauraceae(Magnoliids: Laurales), Sabiaceae (Eudicots), Combretaceae (Rosids: Myrtales), Euphorbiaceae(Eurosids I: Malpighiales), Salicaceae (Eurosids I: Malpighiales), Moraceae (EurosidsI: Rosales), Urticaceae (Eurosids I: Rosales), Ericaceae (Asterids: Ericales), andEbenaceae (Asterids: Ericales). With reference to Ceropsylla indica Kandasamy, which

60 Oriental Insects Vol. 41induces galls on the flowers of Terminalia arjuna (Combretaceae), Kandasamy (1986)has remarked that species of Teminalia appear as the most preferred host species ofTriozidae. Mycopsylla gardenensis (Bhanotar, Ghosh, & Ghosh) (Homotomidae) is a singularnon-triozid species recorded as a gall inducer on Ficus religiosa (Moraceae) insouthern India (Kandasamy, 1986). Several species of Indian figs (Ficus spp., Moraceae)preferentially host several species of Homotomidae (e.g., Dynopsylla, Homotoma,Macrohomotoma) and Paurocephalinae (e.g., Paurocephala). Only two Triozidae (e.g.,Ceropsylla, Pauropsylla) are also associated with Ficus-species as gall inducers (Hodkinson,1986).A comparison of distribution and host relations of gall-inducing psyllid taxa betweenIndia and Taiwan offers several insights into the associations between particular psyllidgenera and specific plant families. For example, Calophyidae (e.g., Apsylla and Calophya)are associated with Sapindales (Anacardiaceae, Burseraceae, and Rutaceae) andCecidopsylla with Ericales (Theaceae) (Mathur, 1975; Yang, 1984; Burckhardt & Basset,2000). The diversification of the Indo-Australian genus Cecidopsylla demonstrates a distinctpattern: the Asian species of Cecidopsylla developed primarily on the Himalayanelement Schima wallichii (Theaceae; Asterids: Ericales) and has diversified simultaneouslywith the trans-Himalayan diversification and spread of S. wallichii (Hu, 1980),whereas the Australian species of Cecidopsylla developed on Proteaceae (Eudicots: Proteales)and Rutaceae (Eurosids II: Sapindales), because they occurred outside the geographicalrange of Schima, separated by the oceanic barrier. The Asian Cecidopsylla induceleaf-roll galls, whereas the Australian Cecidopsylla induce pit galls (Burckhardt,1991). Diversification of Dynopsylla (Homotomidae) appears strongly aligned with Ficus;D. grandis Crawford induces galls on the leaves of Ficus talbolti King (Moraceae) inIndia (Mani, 1973; Mathur 1975) and D. pinnativena (Enderlein) induces galls on theleaves of F. nervosa Heyne (Moraceae) in Taiwan (Yang, 1984). Livia khaziensis Heslop-Harrison (Liviinae) occurs on Juncus bufonius Linn. (Juncaeae) in Indian Himalaya; L.khaziensis (recorded as L. nigra Klimaszewski) occurs on J. prismatocarpus R.Br. in theeastern Palearctic (Hodkinson, 1986). Phacopteron lentiginosum Buckton (Phacopterinidae)induces complex, two-tier galls on the leaflets of Garuga pinnata (Burseraceae)(Raman, 1987) and is distributed extensively in India, irrespective of local climate vagaries;Neophacopteron euphoriae Yang (Phacopterinidae) induces simple pit galls on theleaves of Euphoria longan Lam. (Sapindaceae) in Taiwan (Yang, 1984; Yang et al.,2006). What is striking is that both Phacopterinidae share gall-inducing behavior and theyinfest species belonging to Sapindales (Eurosids II). Nonetheless, Pseudophacopteronalstonia Yang & Li induces galls on Alstonia scholaris R. Br. (Apocynaceae; Euasterids I:Gentianales) in China, Ps. canarium Yang & Li on Canarium album Raeusch (Burseraceae;Asterids: Ericales), Ps. floccosum (Crawford) on Aglaia roxburghiana Wight &Arnold (Meliaceae; Eurosids II: Sapindales) in Sri Lanka and the central-Pacific Islands,and Ps. tuberculatum (Crawford) on AlstoniaTable 3. Gall-inducing Triozidae in the Indian SubcontinentTrioza species Host plant Plant family ReferenceT. eugenoides Not known Ramakrishna, 1924T. hirsuta Terminalia alata Combretaceae Mani, 1948T. lobata Duabanga grandiflora Mathur, 1975T. malloticola Mallotus phillippiensis Euphorbiacceae Mani, 1973T. serrata* Sabia paniculata Sabiaceae Mathur, 1975

2007 Yang & Raman: Gall-inducing psyllids (Hemiptera: Psylloidea) 61T. pitiformis Mallotus phillippiensis Euphorbiacceae Mathur, 1975T. vitiensis Syzygium jambolana Myrtaceae Mani, 1959T. analis Not known Mathur, 1975T. beesoni* Litsea spp. Lauraceae Mathur, 1975T. bifurcate* Populus sp. Salicaceae Mathur, 1975T. brevigenae Ficus sp. Moraceae Mathur, 1975T. camphorae Cinnamomum camphora Lauraceae Mani, 1973T. fletcheri Trewia nudiflora Euphorbiaceae Mathur, 1975T. fletcheri minor Terminalia spp. Combretaceae Mathur, 1975T. fusca Syzygium cumini Myrtaceae Mathur, 1975T. gigantea Vaccinium neilgherrense Ericaceae Kandasamy, 1986T. jambolanae Syzygium cumini Myrtaceae Mani, 1973T. longiantennata* Populus euphratica Salicaceae Mathur, 1975T. obsolete Diospyros melenoxylon Ebenaceae Mathur, 1975T. spinulata Syzygium cumini Myrtaceae Mathur, 1975T. simplifica* Terminalia travancorensis Combretaceae Kandasamy, 1986T. urticae* Urtica spp. Urticaceae Mathur, 1975*Species that occur restricted to Indo-Gangetic plains and the Himalaya. Unasterisked species occurthroughout the Indian subcontinent irrespective of climatic variations.scholaris R. Br. (Apocynaceae; Euasterids I: Gentianales) in India (Mathur, 1975; Hodkinson,1983; Yang & Li, 1983). Whereas gall-inducing behavior is retained among theOriental taxa of Phacopteridae * , Ps. tuberculatum and Ps. alstonia have radiated on toplant families that are not related to Sapindales. The capability of Ps. tuberculatum andPs. alstonia to invade A. scholaris, a taxon of Apocynaceae, and induce galls on them isstriking, given that the latex of Apocynaceae includes strong compounds that are toxic tophytophagous insects (Uysal et al., 2005).ConclusionDocumented information on gall-inducing psyllids of the Orient and the easternPalearctic points them to be highly specific to their recorded host plants. However, thiscomment needs to be viewed with considerable caution, because modest oligophagyamong adult insects of central-European Triozidae and Neotropical Calophyidae isknown (Burckhardt & Basset, 2000; Burckhardt & Lauterer, 2002). Moreover, severalpredatory insects from Anthocoridae, Miridae, Syrphidae, and Chrysopidae (Hodkinson& Flint, 1971) and parasitic insects from Encyrtidae, Eulophidae, Eupelmidae, Eurytomidae,Pteromalidae, Torymidae, and Chalcodoidea (Jansen, 1957) are associated with gallinducingpsyllids and their galls in the western Palearctic and Nearctic; unfortunately, noreliable data from the Orient and eastern Palearctic are available. Triozidae appears todominate as the principal gall-inducing family in the Orient and eastern Palearctic, whichis followed by relatively fewer species of the Calophyidae and Phacoteronidae. Variationin morphologies of galls induced by the Oriental psyllids ranges from simple leaf-marginrolls induced by Diaphorininae (e.g., Diaphorina truncata on Strychnos nux-vomica) atone extreme to complex two-tier galls induced by Phacopteronidae (e.g., Phacopteronlentiginosum on Garuga pinnata) on the other. Species of Triozidae induce galls that fitbetween the two above-identified extremes by inducing spherical and protruding galls on* D. Burckhardt, while reviewing this manuscript (pers. comm.. email: 14 June 2007) commented: “This is not true forthe world fauna: nearly half of the species of Pseudophacopteron from Cameroon induce galls and the other half donot.”

62 Oriental Insects Vol. 41leaves (e.g., Pauropsylla depressa on Ficus recemosa Linn. [= F. glomerata Roxb.](Mani, 1973; Mathur, 1975), Trioza jambolanae Crawford on Syzigium cumini (Linn)Skeels (Raman, 1991), Trioza fletcheri minor Crawford on Terminalia tomentosa Wight& Arnold and T. arjuna Wight & Arnold (Raman et al., 1997). Host relations of IndianTriozidae have been experimentally tested with Trioza fletcheri minor; karyology of thepopulations of the triozids reared from mature galls of Terminalia tomentosa and T. arjunaindicated them to be of identical chromosomal configuration (Raman et al., 1997)pointing that the same triozid induces galls (of similar morphology) on two species ofTerminalia. T. f. minor has been recorded as a gall inducer not only on T. tomentosa andT. arjuna, but also on T. catappa Linn., T. paniculata Roth, and T. tomentosa x T. arjuna,which are distributed almost uniformly throughout India (Champion & Seth 1968;Mathur, 1975; Hodkinson, 1986). Galls on the flowers of T. arjuna were reported to beinduced by Trioza sp (T. f. minor[?], Sokhi & Kapil, 1984; 1985), but we currently knowthat these galls are induced by Cerospylla indica (Triozidae) (Kandasamy, 1986).A cursory evaluation of the pattern of association of gall-inducing psyllids of theOrient and eastern Palearctic with their respective host-plant taxa indicates that themaximum number of psyllids are associated with arborescent species of Eurosids (AngiospermPhylogeny Group, The, 2003), whereas relatively few are associated with theprimitive Magnoliids and relatively advanced Asterids; whether such a pattern has anevolutionary context and meaning, is an issue remaining to be explored.AcknowledgementsWe thank Daniel Burckhardt (Naturhistorisches Museum, Basel, Switzerland) forreviewing this article. We are grateful to V. V. Ramamurthy (Indian Agricultural ResearchInstitute, New Delhi, India) and B. V. David (Sun Agrochemicals, Madras, India) forsupplying a few key papers on Indian psyllids.ReferencesANGIOSPERM PHYLOGENY GROUP, THE, 2003. An update of the Angiosperm Phylogeny Groupclassification for the orders and families of flowering plants: APG II. Botanical Journal of the LinneanSociety, 141: 399–436.BALAKRISHNA, P. & RAMAN, A., 1992. Cecidogenesis of leaf galls of Strychnos nux-vomica(Loganiaceae) induced by the jumping plant louse species Diaphorina truncata (Homoptera:Psylloidea: Psyllidae). Entomologia Generalis, 17: 285–292.BURCKHARDT, D., 1991. Notes on the Indo-Australian plant-louse genus Cecidopsylla (Homoptera:Psylloidea) with descriptions of two new species. Phytophaga (Madras), 3: 73–86.BURCKHARDT, D., 2005. Biology, ecology, and evolution of gall-inducing psyllids (Hemiptera: Psylloidea).In: Raman, A., Schaefer C. W., & Withers T. M. (eds) Biology, ecology, and evolution of gallinducingarthropods. Science Publishers, New Hampshire, USA. Pp. 143–158.BURCKHARDT, D. & LAURTERER, P., 1989. Systematics and biology of the Rhinocolinae (Homoptera:Psylloidea). Journal of Natural History, 223: 643–712.BURCKHARDT, D. & LAURTERER, P., 2002. Revision of the Central European Trioza rotundata Florcomplex (Hemiptera: Psylloidea): taxonomy and bionymy. Mittelungen Schweizerischen EntomologischenGesselschaft, 75: 21–34.BURCKHARDT, D. & MIFSUD, D., 2003 Jumping plant lice of the Paurocephalinae (Insecta, Hemiptera,Psylloidea): systematics and phylogeny. Contributions to Natural History, Bern, 2: 3–34.BURCKHARDT, D. & BASSET, Y., 2000. The jumping plant-lice (Hemiptera, Psylloidea) associated withSchinus (Anacardiaceae): systematics, biogeography and host plant relationships. Journal of NaturalHistory, 34: 57-155CARVER, M., GROSS, G..F. & WOODWARD, T. E., 1991. Hemiptera. In: CSIRO. The Insects ofAustralia, Vol. 1, 2nd edition, Melbourne University Press, Melbourne, Australia.

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