COMMUNITYAND ECOSYSTEMECOLOGY Phytophagous Arthropods Associated with Parthenium hysterophorus (Asteraceae) in North America A. S. McCLAY, W. A. PALMER,l F. D. BENNETf,2 AND K. R. PULLEN3 Alberta Environmental Centre, Bag 4000, Vegreville,AB T9C IT4 Canada Environ. Entomol. 24(4): 796-809 (1995) ABSTRACT A faunal sUlVeywas conducted to find possible biological control agents for Parlhenium hysterophorus L . (Asteraceae), a serious annual weed in many parts of the world, particularly Queensland, Australia, and India. Between 1977 and 1991, most of the plant's native range in North Americawas sulVeyedfrom bases at Monterrey and Cuernavaca, Mexico, and Temple, TX. Two hundred and sixty-twophytophagous arthropod species were collected on P . hysterophorus by various methods including hand picking,dissection, rearing, and sweeping; 144 of these species were found to feed on the plant at some stage of their life cycle. The orders represented most abundantly were Coleoptera (33.2%), Homoptera (22.9%), Lepidoptera (20.2%), and Hemiptera (18.3%). Two fungal pathogens, Puccinia abrupta Dietel and Holwayvariety partheniicola (Jackson)Parmelee and Puccinia m elam podii Di'etel and Holway, were also obselVed. An index of similarity was used to make pairwise comparisons between the phytophagous arthropod communities on different plant taxa. These comparisons showed that the fauna of P . hysterophorus is most similar to that of ragweeds (Am brosia spp.). Six insect species that were shown to be stenophagouswere shipped to Australiafor further testing and possible field release. One of the fungal pathogens underwent host-range testing in the United Kingdom and was released in Australia. KEY WORDS Parlhenium hysterophonJs, Asteraceae, biologicalcontrol, phytophagous, arthropod fauna, North America hysterophorus L. (Asteraceae) is a herbaceous annual weed native to the neotropics, from the southern United States to Argentina, which has become widely naturalized in other tropical and subtropical regions. It has become a particularly serious problem in Queensland, Australia (Haseler 1976) and in India (Towers et al. 1977). It also occurs in Taiwan, southern China, Vietnam, Vanuatu, New Caledonia, Kenya, Madagascar, Mozambique, South Africa, the Mascarenes and Seychelles, and Ethiopia. In Queensland, thousands of square kilometers of rangeland are infested, and pure stands of P hysterophorus have excluded beneficial pasture species over large areas (McFadyen 1992). All parts of the plant contain parthenin, a bitter, toxic sesquiterpene lactone, which makes the plant unpalatable to cattle and has caused many cases of severe contact dermatitis in susceptible people (Towers et al. 1977). Its profuse seed production enables it to spread rapidly. The plant is reported to be allelopathic (Dharmaraj et al. 1988). Parthenium lTropical Weeds Research Centre, P.O. Box 187, Charters Towers, Queensland 4820, Australia. 2Department of Entomolow and Nematology, University of Florida, Gainesville FL 326JJ-D740. 36 Dianella Road, Port Macquarie, New South Wales 2444, Australia. . 0046-225X195/0796-0809$02.00/0 The center of diversification of the genus Paris considered to be in northeastern Mexico, where at least 8 of the 16 described species occur (Rollins 1950). The general climatic characteristics of this area are similar to those of the infested areas of Queensland (McClay 1985). The cost of chemical and cultural control of large stands of P hysterophorus on low-value pasture is prohibitive (McFadyen 1992). In an attempt to identify alternative control measures for this serious rangeland weed, the Queensland Department of Lands (QDL) started a biological control program in 1975. As part of this program, scientists of the Department of Lands and of the Commonwealth (now International) Institute of Biological Control (CIBC, now nBC) undertook exploration in North and South America. These staff were responsible for conducting faunal studies of P hysterophorus and selecting appropriate stenophagous species for host specificity and biological studies. If these tests indicated that a species was a potential biological control agent, it was shipped to the Sir Alan Fletcher Research Station of the Queensland Department of Lands in Sherwood, Australia, for further testing, approval, and eventual release. This article reports the phytophagous insects and mites found in North America during these investigations. A preliminary list of 159 phytophagous insects associated with the plant in Mexico thenium © 1995 Entomological Society of America August 1995 M cC L A Y ET A L .: ARTHROPODS OF P. hysterophorus 797 Fip;. 1. Searcharea for natural enemies of P. hysterophoros in the United Statesand Mexico(shaded).Numbers identifYthe regionsused to indicate geographicaldistributionin Table 1. has been published (McClay 1981) and the species sent to Australia for further testing were discussed by McClay (1985). In addition to the surveys reported here, QDL staff conducted surveys of South American populations of P. hysterophorus in Brazil and Argentina (McFadyen 1976; QDL, unpublished data), and some collections were made in the Caribbean by F.D.B. and other CIBC staff. The biological control program for P. hysterophoruo\' was reviewed by McFadyen (1992). Materials and Methods The area covered by this survey (Fig. 1) comprised most of Mexico and parts of the southeastern United States. Two preliminary surveys were conducted over a total of 40 d in June, October, and November 1976 by F.D.B. (unpublished data, 1977). During these surveys, collections were made in the states of Veracruz, Yucatan" Campeche, Quintana Roo, Puebla, and Nuevo Le6n, Mexico, and in the areas of Houston and College Station, TX, Baton Rouge, LA, and Miami and Homestead, FL. More detailed studies were then conducted by A.S.M. from 1978 to 1983 from a base at the Instituto Tecnol6gico y de Estudios Superiores de Monterrey (ITESM) at Monterrey, Nuevo Le6n. During this period, extensive collections were made in northeastern Mexico, particularly the states of Nuevo Le6n, Tamaulipas, and eastern Coahuila. Some collections were also made along the Gulf coastal areas of Veracruz and Tabasco, and on the Pacific coast from Sinaloa to Nayarit and from Oaxaca to Chiapas. From 1983 to 1993, surveys were conducted in Texas by W.A.P. from a base at the Queensland Department of Lands North American Field Station (NAFS) in Temple, TX. From 1988, W.A.P and K.R.P. made additional surveys in southern Mexico, particularly the state of Morelos, from a substation at Cuernavaca, Morelos. Collections were made on natural stands of P. hysterophorus on roadsides, wasteland, neglected fields, and similar sites at frequent intervals throughout the year. Arthropods resting on the plant were captured directly by hand or with an aspirator, or by sweeping pure stands of P hysterophorus with a sweep net. All parts of the plant were dissected for endophagous species. Adults of root-feeding species were obtained by keeping roots in emergence boxes with glass vials fitted over the exit holes or in large glass jars covered with nylon gauze. Other plant parts were usually too perishable to be held in the laboratory long enough for emergence of adult stages of endophages. Some stem and root-feeding insects were reared to the adult stage in the laboratory by feeding ReId-collected larvae on an artificial diet (Harley and Willson 1968). 798 E N V IR O N M E N T A L In addition to these collections on natural stands, a series of plots (5 m by 5 m) were sown with P. hysterophorus at the ITESM field station, Apodaca, Nuevo Le6n, and allowed to become infested with arthropods. Samples were taken from the plots at 2-wk intervals from 1978 to 1982 by selecting 25 plants at random and collecting all arthropods resting on them or feeding internally in them. Most specimens were initially submitted for identification either to the United States Department of Agriculture Systematic Entomology Laboratory, Beltsville, MD, or to the Commonwealth (now International) Institute of Entomology, London. Specimens that could not be identified to species by these laboratories were later forwarded to other taxonomists. Many submitted specimens were identified only to genus because of inadequate material, unavailability of type specimens, or lack of satisfactory taxonomic revisions. These are reported here only if they represent a genus not otherwise reported or if the taxonomist explicitly stated that they are distinct from the other named species of the genus reported here. The probable level of host specificity of each species was assessed using information from the literature, specialists, and field observations. Species that appeared likely to be host specific, at least the subtribe level, were selected for preliminary host-specificity studies conducted in Mexico and Texas. Test plants were selected from the subtribe Ambrosiinae and tribe Heliantheae, to which the genus Parthenium belongs (Stuessy 1977), with the addition of some economic species from other tribes of the Asteraceae. Sunflower, H elianthus annuus L., is the most important crop species closely related to Parthenium and was considered a critical test plant (Wapshere 1975). Experimental details depended on the biology of each species tested, but many tests were conducted in multiple-choice cages as described by McClay (1987). Field observations of the insects' host plant associations were also made. If our investigations indicated that the host range of an insect was within the subtribe Ambrosiinae, the insect was recommended for further study and shipped to Australia after quarantine approval had been received. In our field surveys, it was not always possible to determine whether a captured insect was actually feeding on P hysterophorus. Published faunal lists for many plants frequently do not in~cate whether the species listed are known to feed on the plant in question or were simply collected on it. We report here all the species belonging to known phytophagous taxa that we collected on P. hysterophorus, with the exceptions of pollen or nectar gatherers, adult Lepidoptera and Diptera, species shown in laboratory tests not to feed on P hysterophorus, and species whose known biolog.y clearly indicates that P. hysterophorus is an unlikely host. Where any life stage of an insect species Vol. 24, no. 4 E N T O M O LO G Y has been observed to feed on P hysterophorus, this fact is indicated in Table 1. Records where feeding is not indicated refer only to collections of adults on the plant; this category undoubtedly includes some species which are casual visitors on P. hysterophorus. Specimens of many of the species identified were retained by the taxonomists responsible for identification, and were deposited mainly in the National Museum of Natural History in Washington, DC, and the Natural History Museum in London. Material of all species collected from 1978 to 1983 was deposited in the collection of the IIBC Caribbean and Latin American Station, Trinidad, and at NAFS. Additional material from the years 1978 to 1983 was deposited in the collection at ITESM. Material collected in 1983 and afterward was placed in the NAFS collection, now held at the Tropical Weeds Research Centre (TWRC), Charters Towers, Queensland. Inquiries concerning the location of particular material collected during the project may be directed to W.A.P. or A.S.M. Relevant information about each species was also entered into a computer database now maintained at TWRC (Palmer 1995); detailed database records for particular species may be obtained from W.A.P. We assessed the degree of similarity between the fauna of P. hysterophorus and those of 1 other Parthenium species, 6 other native North American genera of Asteraceae, and 4 native North American genera in other families. A pairwise index of p.ercentage similarity Is was defined by Is = 100X(llshareinpooled) where llsharedis the number of arthropod species in common between the 2 plant taxa and npooledis the total number of species in the pooled faunas of both taxa. Results and Discussion Phytophagous arthropods associated with P. hysin Mexico and the United States are listed in Table 1. Information on host specificity and pest status in this table is a synthesis of the results of feeding tests carried out during the program, field observations, literature records, and information from specialists who identified the insects. Species that were subjected to host range tests during the project are indicated. The numbered regions used to identify geographic distribution are shown in Fig. 1. Distribution records in Table 1 represent only data from our surveys and do not include information from the literature. Many species may, therefore, have wider distributions than indicated in the table. Table 1 also indicates frequency of collection for each species. Because collections were not quantitative, the assessment of whether a given species is common, occasional, or rare is based on a subjective assessment of the collecting experience of terophorus August 1995 MCCLAY Table 1. Phytophagous arthropods Species 799 OF P. hysterophorus ET AL.: ARTHROPODS associated with P . hysterophorus FreStages quenc)" feedingi' Feeding site" Feeding typed Host speci6city" Testedf T Distributionf; Pest status" Acari Eriophyidae Enophyes nr. parthenii Keifer 0 N,A F ECT ***** G N,A L ECT ** L ECT Tetmnychidae 1i'tranyehlls nr. l.uleni Zacher Orthoptera Gryllidae O cean thus nr. celerinictus T. walker 0 N,A Tt'ttigoniidae C O lw cL'}Jhalu.s fascilltuS (DeGeer) R 8 * Thysanoptera Thripidat' C lliiothrips phaseoli (Hood) M icrocL'}Jholothrips abdom inalis (Crawford) G 0 N,A N,A L F ECT ECT P * ** Hemiptera Alydidae Burtinus fenw ralis (Distant) R Bt'rytidae Jalyslls reductlls Barber R 5 R R R C 5 1 5 1,3,5 Cor!'idae C hariesterus lIw estus Burmeister N irooeclls cllloiger StAl Pllthia picta (Drury) Sagotylus conf/llens (Say) N,A S ECT * l' P Lygaeidac Acrolcucus sp. Blissus IL~,copterns (Say) N eopam em bi/obata (Say) N lJsius senecionis (Schilling) N '.Isius sp. XyonysillS califom iC llS (StAl) R R R C C C A A A F F F ECT ECT ECT 5 7 5 * *** P ] 1 1,5 Miridae C ollana oleosa (Distant) C yrtopeltis sp. H altiC lls bm ctatus (Say) [[')(leom i'lllsta (Distant) LlJg,lS [;,leolaris (Palisot de Beauvois) N m rocolp.lS prob. m exicallllS Distant PhlJtocoris sp. Poecilocapslls nigriger (StAl) Polym erus basalis (Reuter) PollJm enlS testaceipes (StAl) Proba distanti (Atkinson) Prohll sll!lei (Stll) RC iiteroseopus om atllS (Reuter) Rhinacloa forticom is (Reuter) Sixeonotus sp. Taedia sp. TalJlorilygus pallidulus (Blanchard) 0 0 R 0 0 R R R R 0 C R C R R R C * * ** * N,A L ECT * * N,A F,L,S ECT * ** ** N,A F,L,S ECT ** l' T 5 5 12 5 5, 8 5 5 1 12 5 1,2,4,5,7 5 1,2,4,8, 10 5 N,A F,L,S ECT * P P P 5 5 1,5, 7, 8, 10 Pentatomidae C hloroehroa ligata (Say) D endroeoris sp .. E./.Schistus quadm tor Rolston EuseltishlS sp. M om lidea notlliata (Herrich-Schaffer) Tltyanta aeeelTa McAtee Tltyanta pem itor (F.) R R 0 R R R R * N,A L, S ECT * * * * 12 12 1,7 2, 7 7 8 5 P P P Piesmatidae Piesm a cinereum (Say) 0 P 800 ENVIHONMENTAL Vol. 24, no. 4 ENTOMOLOGY Table 1. Continued FreStages quenc)" feedingh Species Feeding site" Host speci6city' Feeding typed Tested!" Distributio~ Pest statm J• Pyrrhocoridae D ysdercus m im ulus R Hussey 1 * P Rhopalidae H arrrw stes nebulO SlIS StM H arrrw stes rejlexulus (Say) H annostes serratus (F.) LiorhysstlS sp. R 0 R R 1,5 1 3 1 ** ** * Scutelleridae C helysom idea oariabilis (Herrich-Schaffer) H om aem us proteus St,u Sphyrocons obliquus (Germar) R 0 0 N,A L, S N,A L,S ECf 1 1,5 1 * Tingidae C O "fthucha m orrilli Osborn & Drake D ietyla 11w notropidia (St,u) Leptodictya tabido (Herrich-Schaffer) C ECT * R * R * T 1,2,5,8 11 P 5 Homoptera A1eyrodidae Trialeurodes abutilonea (Haldeman) ·L ECf N,A N,A N,A L, S R S,L ECT ECf ECf A L ECf N,A L, S ECT G N G C C R C R * 1 P * l P * 1 1, 2, 3, 4, 7, 8, 9 Aphididae Aphis gossypii Glover G eopem phigus floccO stlS (Moreira) U roleucon am brosiae (Thomas) ** ••• T Cercopidae Aeneolam ia sp. C lastoptem xanthocephala Prosapia sp. Germar 3 8 3 Cicadellidae Aceratagallia gillettei (Osborn & Ball) Agallia barretti Ball Agallia quadripunetata excavata Oman Agrosom a pulchella (Guerin) Egidem ia inf/ata Young Em poasca ahm pta DeLong Em poasca or. crocostigm ata Davidson & DeLong C R C R R C * T 5 * 5 2,5 3 1 P 5 P * N,A L, S ECT 1 * * 0 Em poasca nigra var. typhlocyboides Gillette & Baker G raphocephala m jim argo (Walker) G yponana sp. H om alodisca coagulata (Say) H om alodisca insolita (Walker) M acrosteles sp. M om oria m isella (Ball) O ncom etopia elarior Walker Penestragania rom lSta (Uhler) Phera lanei Young Phera obtusifrons (Fowler) Scaphytopius frontalis (Van Duzee) Scaphytopius jiJliginostlS (Osborn & Ball) Sibooia com pta (Fowler) TexananrlS ooatus (Van Duzee) Xerophloea viridis (F.) C C N,A A L, S L ECT ECT A A S L, S ECT ECT ** * R 0 R * * R R 0 C R ** * N,A L, S ECT * T N,A L, S ECT * * T R R 0 R 0 0 * * N,A L, S ECT * S L S S ECT ECT ECT ECT * R N.A N,A N,A N,A C N,A L,S ECT * ••• C N,A L, S ECf T 1 5, 7 1,2,3 1 5 5 7 1,3 1,2,4,5,8 5 5 8 1,5 5 1 1 P P P P Coccidae C eroplastes cirripedifannis Comstock Lecanium sp. Parasaissetia nigra (Nietner) Saissetia m iranda (Cockerell & Parrot) R R R 3 5 2, 3 3 * * Delphacidae Stobaera concinna (St,u) T 1,4,5 Dictyopharidae Scalops sp. 1 P P P Augllst 1995 McCLAY ET AL.: ARTHROPODS OF P. 801 hystem phorus Tuble 1. Continued FreStages quem:>", feeding" Species Feeding site" Host specilicit}" Feeding typed Distribution(; Testedf Pest statush Eriot'o(x'idae dubius (Cockerell) Am nthocoeeus R N,A S ECT 3 * Flalidal' FZlltO "ucnL. sp. 0 5 R 2 Issidal' C oZpol'tem lllbavC rlosa Caldwell Margarodidae R R lrenja nr. littom /is Cockerell lcerya l'urchllsi Maskell N,A N,A L S ECT ECT * N,A S, L ECT ** 3 3 P Ml'mbmeidae AcutalL. tar/area (Say) Entyl/a cnnnata (Forster) M ien/talis sp. 1'0/yglW tll sp. Publ/lla sp. SI,L.sL.H lus/esH nus (Say) Stictoe17,hala sp. U m bonill prob. crass/com ls Amyot & Serville R C C R 0 R 0 * 0 R Vanduzcll segl1u.'fItata (Fowler) 12 1,5,12 1,2,5 5 1,4,5 1,2,12 1 * * 5 1 * 11 3 P Ortht'7jidae O rtlw z/a pm elonga Douglas O rthr.;/a I'SetulillSignis Morrison L, S ECT R R N,A C R G N,A N,A N,A 0 0 0 R N,A N,A N,A N,A S R S F ECT ECT ECT ECT R N,A S ECT * Pseudococcidae Ferrisia virgata (Cockerell) Paracoccus sp. Phenacoeeus nr. defectus Ferris Phenaeoecus gossypii Townsend & Cockerell Phenaeoceus solani Ferris Psm doc.llcC llS affinls (Maskell) Sl'ilococclls sp. L, R, S, M ECT ECT S S, M ECT 1,2,3,5 2 1 * * * 3 1,2 11 2,3 * 11 * * P P P P Plltoidae Puto barben (Cockerell) P Coleoptera Anthribidae Tngonorhinr •.• lim batus (Say) R 1 R 5 R 5 Apionidae C hnjSal'ion auctu", (Sharp) Bntehidae Ac(m thoseelid,,.. sp. C<'rambycidae D ectes lIigril'ilus Chemsak & Linsley D ect"s sayi Dillon & Dillon D ectes t.':tanr•.• LeConte D orcasta ciril'rea (Hom) H ippopsis lem niscata ( F . ) Phaea tenrw ta Bates Pseudoeanidia cuem avacae Dillon R C 0 R 0 R 0 L,A L,A S, L S, L END END ** * * L,A S, L END * L S END *** T 4 1 4,5 P 1 P 1,2,3,10 5 5 P Chrysomelidae AcaZymmalJlomorum Munroe & Smith Agathom en~' nifi •.• Klug AnopliH s sp. C luU 'tocnel1U 1hom i Jacoby G hal17JU snr. conH g"'''' Baly G hrysodina sp. C "7'idodem sp. C njptocephall~' irrorall •.• Suffrian C njptocephlll, •.• prob. nigrovittall •.• Jacoby C ryptocephaZus sp. R R R R R R R 0 0 0 * L L END A L ECT L,A L,A L,A L L L ECT ECT ECT .*** T 3 5 2 11 1 5 5 1 1 1 P 802 ENVIRONMENTAL Vol. 24, no. 4 ENTOMOLOGY Table 1. Continued Species D erocrel'is sp. D iabrotica ba/teata LeConte D iabrotiEa undecim punctata Jacoby D iachlls aum tlls (F.) D isonycha knabi Blake D urallgoita chrysodinoides (Jacoby) El'itrix sp. Exem a prob. conspersa (Mannerheim) Exem a or. dispar Lacordaire G lyptina sp. G /yptosce/is nr. m exican a Jacoby or. H ippllriphila sp. Lobidom em sp. Lexiphanes gllerini (Perbosc) Lllpraea sp. M icrorhopa/a pulchella Baly M icrorhopala prob. sallei Baly N odonota nr. atm Harold O m ophoita aequinoctialis ( L . ) O phraella com m una LeSage O phraella prob. slobodkini Futu)'lna Pachybrachis or. fenw m ta Olivier pachybrachis im m acu/atus Jacoby Pachybrachis nr. vestigialis Fall Pachybrachis sp. psylliodes sp. Systena b/anda Melsheimer Systena nigrop/agiata Jacoby Systena nr. 1)ectom /is Clark Systena or. sem ivittata Ja(.'Oby Systena sp. Zygogm m m a bicolorata Pallister Zygogm m m a m a/vae St:ll F~eStages quency' feedingb Feeding site" Host specificity' Feeding typed R a R C R R R C a * ** L,A L ECT L,A L ECT L,A L,A L F ECT ECT * *** *** T T L,A L ECT *** T L,A L ECT *** T T T R R R C R R R R R R a a a R C R a R C C R 5 1,3,5,7 11 1,5,8 5 1 Pest status" P ] ** ** R a a Distributionll Testecif A L,A A L L L ECT ECT ECT **** T 1,2,5,7 5 1 11 5 1 5 7 3 5 11 1 1,2,5 2 11 1,4 5 1 11 11 5 1,2 3 1 1,2,4,5 5 P Coccinellidae Ellilachna m exicana (Guerin) * R 7 P Curculionidae Baris prob. aerea Boheman Baris prob. interstitia/is Say Baris sulcipennis (Brisout) C ha/coderm us sp. C om l'sllS auricepha/us (Say) C ylindrocopturus annat!lS Champion C y/indrocopt!.rus or. m am m illatus (LeConte) C y/indrocoptunlS tetra/ob,lS Champion EpicaenlS allrifer Boheman Epicaerus prob. texanus Casey G eraellS picrm m us (Herbst) G eraeus seni/is (Boheman) Linogem eus sp. Listroderes costirostris ob/iquus Klug Lixus prob. scrobicollis Boheman M itostyl.lS SetO SllS (Sharp) O nychobaris sp. Pande/eteius sp. Rhodobaenus quinquepunctatus (Say) Rhodobaenus sanguineus (Gyllenhal) Rhodobaenus tredecim punctatus (Illiger) SciadrtlstlS propheticus Sharp Sibinia sp. Sm icronyx lutulent!.s Dietz Sm icronyx sordidus LeConte Thecestem us hirst/tus Pierce Trepobaris inonurta Champion C a L,A L,A R R END END ** ** L A L,A S L R, S END ECT END * *** T R R a a ] T R R * R a R R R R C a a L,A L,A A L S, L L END ECT * *** ** L,A L,A L,A S S S END END END ** ** *** L,A F END * ***** *** L,A R ECT L L L L S S S S END END END END T R a C R R R C R a lit •••••• R 1,2,5,6,7 1,2,6 5 1 T T 1,2,7 5 5 5 1 1,5 5 5 1 1,2,4 1,4 1 1 8 2,5 1,2 5 1,5 1,2,4,8 1 1, 8 5 P P Languriidae D asydactyZ,lS sp. Languria m ozardi Latreille Languria or. /aeta Leconte Longurites lineat!.s (Castelnau) R a R R * ** 2 1 2 1 P August 1995 M cC L A Y ET A L .: ARTHROPODS OF P. 803 hysterophorus Tobie 1. Continued FreStages quene:)'" feeding> Species Feeding site" Host specilicit}" Feeding typed Distributim,g Testedf Pest statui' Scarnblleidae Euphoria sp. R 5 Scolytidae H !lIlO thellC lIlIlSnr. beam eri Wood S END L L L L ECT ECT * * 1 2 L ECT ECT ECT ECT ECT ** 2 1 2 **** **** ] a L L L L L R C L L L L L L F,L L L F F L F,L R L 1 Lepidoptera Arctiidlle a Estigm ene (l(:rea (DnJry) H a/isidota catenulata Hubner R P Celechiidae D iehom eris nr. aenigllU ltica (Clarke) D iehom eris sim pliciella (Busck) or or. G lyphidocera sp. G 'lO rim oschaC IIU lsaphirillella Chambers lIeleystogram m a cham bersella (Murtfeldt) R a R R L L L L 1 Geometridae Anacom ptodes prob. herse (Schaus) A'lll('(lm ptodes perfectaria McDunnough Allavitrinella pam pinaria (Guenee) C M orocM am ys chloroletlcaria (Guenee) C yclopllO ra nanaria Walker EU IJithecia m isem lata (Grote) or near lIyponu:cis or. um brasaria Hubner Pleuropm cha asthenaria (Walker) Pleuropm clU l insulsaria (Guenee) Salm lodes arga Druce Synehlora frondaria Guenee TO nJos abjeG tarillS Guenee L ECT ECT ECT ECT ECT ECT ECT ECT ECT ECT ECT ECT L L END L L L L *** *** a L L END END END R G R L L a L L F L L L R L L R L L * * * * * * L L ECT ECT ECT ECT ECT ECT ECT L L ECT *** *** R C C R R R L L L L C; R L L L L L L R G R R R 5 *** * * ** * * * * T ] 1 1 1 1,8 8 5 3, 8 5 1,8 1 * **** P P P Cradllllriidlle PhyllononJcter sp. Lyonetiidae Buc,,,latrlx m nbrosiaefoliella Chambers Bucculatrix nr. criticopa Meyrick Bucm llllrix pfltthenica Bradley .*.** T 1 1 1,4 Noctuidae C ondiea sutor (Guenee) II elieoverpa zea (Boddie) Lophoceram ica pyrrha Druce O gdoconta cinereola (Gul'nee) PscudopltlSia illc/udens (Walker Spodoptera latifascia (Walker) Spragu";a guttata Grote a 8 1 5 1,8 1,8 2 1 P 1,2,4 P P P Nymphalidae C M osy"c lacinia Geyer C T Pterophoridlle O idaem atophonls fum ivcntris (Zeller) O idaem atopllO nlS inquinatus (Zeller) O idaem atopllO nlS stram illC U S (WaIsingham) 5 R L a L L ECT R L S E N D 1,2 1 Pyralidae Achyra ranta/is (Guenee) C onchylodes ovu/alis (Guenee) R L L a L L ECT ECT * ** R L L ECT ** R L L ECT C L R END R R L R M END END 1,8 1 Hiodinidae C alephelis /avenJa Godman & Salvin or nr. Sllturniidae H ylesia sp. 2 Scsiidae C am lellta ithacae (Beutenmtiller) *** 1, 2 Tineidac Acrolophus sp. Tinea sp. L 2 7 P 804 ENVIRONMENTAL Vol. 24, no. 4 ENTOMOLOGY Table 1. Continued Species Feeding site" FreStages quen<-1' feeding/' Feeding typed Hos~ specIficity' Teste-" Distribution'l (]I Pest statu1' Tortricidae Aethesoides distigm atana Walsingham or near Am oroia em igratella Busck Am orbia or. rectangularis Meyrick Argyrotaenia m ontezum ae (Walsingham) C arolella or. buscana Comstock C ochylis subolivacea Walsingham Epiblem n strenuana (Walker) Platynota labiosana Zeller or near Platynota nigroceroina Walsingham Platynota rostTana (Walker) Platynota stultana Walsingham Sulei11U ls kinnerana Heinrich R 0 R 0 0 R C C L L L L L L L L R C L L L 0 L 0 R L L L R, S M S L L L L,F S END ECT ECT ECT END END END ECT ECT ECT ECT END * 2,5 5 2 2,3,5 1,4 7 1, 2, 3, 4, 5, 8, 10 1,2,7 8 8 * 1 *** 5 * 1 1 1 1,2,4,5 * * **** * T P P P Diptem Agromyzidae Agrom yza sp. Arnourom yza m aculosa (Malloch) Liriom yza sp. M elanagrom yza sp. C L L L L L, M L S END END END END R C L L F S END END .**** C R L L F S END END ** R 0 0 L P Cecidomyiidae C ontarinia sp. Lasioptera sp. Tephritidae Euarestoides acutangulus Thomson EutTeta augusta Banks ** C, common; 0, occasional; R, rare; G, found in greenhouse but not in field. N, nymph; L, larva; A, adult. o L, leaves; R, roots; S, stems; F, /lowers/seeds; M, apical/axillary meristems. d END, endophagous; EGr, ectophagous. e*****, genus Parthenjum; ****, subtribe Ambrosiinae; ***, tribe Heliantheae; **, family Asteraceae; *, other families. fT, species subjected to laboratory host-range tests. g I, NE Mexico (Nuevo Le6n, Tamaulipas); 2, Mexico Gulf Coast (Veracruz, Tabasco); 3, Mexico Yucatan (Campcche, Yucatan, Quintana Roo); 4, N Central Mexi<-'O(Chihuahua, Coahuila, Durango, Zacatecas, San LUISPotosI); 5, S Central Mexico (Aguascalientcs, Guanajuato, Queretaro, Hidalgo, Mexico, Puebla, Morelos, Distrito Federal, Michoac:l.n (inland), Jalisco (inland»; 6, N Pacific Mexic.'O (Sonora, Sinaloa, Nayarit, Baja California Norte, Baja California Sur); 7, S Pacific Mexico (Chiapas, Guerrero, Oaxaca, Colima, Michoac:in (coastal), Jalisco (coastal); 8, Texa~; 9, Louisiana; 10, Florida; 11, Mexico, unspecified; 12, United States, unspecified. h P, known economic pest species. a b the authors. Species listed as common can be easily found at a high proportion of collection sites within their geographic range, often in large numbers, and appear to be regularly associated with P. hysterophorus. Species listed as occasional require greater searching effort or a knowledge of particular sites at which they occur; they were found only at a limited number of sites or occasions, but still show a consistent pattern of association with the plant. Species listed as rare were collected only on one or a few occasions, usually in small numbers or as single individuals, and may not be consistently associated with the plant. We collected 262 phytophagous arthropod species, representing 108 genera and 58 families in 8 orders. The orders represented were Acari (2 species, 0.&%of species), Orthoptera (2, 0.8%), Thysanoptera (2, 0.8%), Hemiptera (48, 18.3%), Homoptera (60, 22.9%), Coleoptera (87,33.2%), Lepidoptera (53, 20.2%), and Diptera (8, 3.1%). The percentage representation of the main insect orders in the fauna of P. hysterophorus is broadly similar to those reported for other herbaceous composites in their native ranges, such as Am brosia psilostachya DC (Goeden and Ricker 1976c), C irsium califom icum Gray (Goeden and Ricker 1986), and for the woody perennial composite Baccharis halim ifolia L. (Palmer 1987), although the percentage of Diptera is lower than reported for these other composites. Fifty-three species were known agricultural pests. Of the species collected, 144 were confirmed to feed on P. hysterophorus at some stage of their life cycle. Because little has been published on the insect fauna of P. hysterophorus, many of these are new host records. In a comparable series of faunal surveys on ragweeds (Am brosia spp.), none of 9 ragweed species had an arthropod fauna of more than 130 species (Goeden and Ricker 1974a, b; 1975; 1976 a-c). Of the 144 species known to feed on P. hysterophorus, 29 (20.1%) were stenophagous as defined by Goeden and Teerink (1993), feeding on species within the tribe Heliantheae; of these 6 (4.2%) were considered to be monopha- August 1995 M cC LA Y ET A L .: ARTHROPODS I!;0us,feeding only on species within the genus Parthl'nillnl. In comparison, the insect faunas of Ambrosiinae in California reviewed by Goeden and Teerink (1993) included 27.0% stenophagous and 18.2% monophagous species. The insect fauna of P. hysterophorus thus appears to be in general less host specific than for other species of North American Ambrosiinae, being particularly deficient in monophagous species. This may be related to the status of P hysterophorus as an annual species whose abundance fluctuates strongly in response to rainfall and habitat disturbance, thus providing a relatively unreliable food resource and discoura!!;inl!;the evolution of specialized phytophages (Bernays and Chapman 1994). Forty (27.8%) of the species feeding on P hystl'rophorus were endophagous, a similar value to that observed by Goeden and Teerink (1993) for Californian Ambrosiinae. Internally feeding species tended to be more host specific; 30.0% of endophagous species and only 16.3% of ectophagous species were restricted to the tribe Heliantheae. Thirteen species were considered to be restricted to the subtribe Ambrosiinae. This level of specificity was considered sufficient to quality a species as a potential biological control agent for use in Australia, because the only members of this subtribe occurring in Australia are introduced weeds in the genera Am brosia, Parthenium , Xanthium , and Iva. Six species were shipped to Australia for further testing. These were a defoliating beetle, Zygogram m a bicolorata Pallister, a stem-galling (Walker) (McClay moth, Epiblem a strenuana 1987), a leaf-mining moth, Bucculatrix parthenica Bradley (McClay et aI. 1990), a sap-sucking planthopper, Stobaera concinna (Stal) (McClay 1983), a seed-feeding weevil, Sm icronyx lutulentus Dietz, and a root-galling weevil, Thecestem us hirsutus Pierce (McClay and Anderson 1985). All except T. hirslltus were subsequently released (McFadyen 1992). Zygogram m a bicolorata also has been released and established in India (Jayanth 1987, Jayanth and Bali 1990). Epiblem a strenuana has been introduced to China for the biological control of annnal ragweed, Am brosia artem isiifolia L. (Wan and Wang 1995). Two mst fungi were also found on P hysterophorus. Puccinia abrupta Dietel and Holway var. partheniicola (Jackson) Parmelee was found on the stems and abaxial leaf surfaces, most commonly in Coahuila and other areas of Mexico at altitudes of 1.'500 m or higher. It was subsequently screened under quarantine in the United Kingdom and released in Queensland (Evans 1987, Parker 1990, McFadyen 1992, Parker et aI. 1994). Puccinia m elam podii Dietel and Holway was found on the adaxial leaf surfaces on plants in south Texas and Nuevo Leon. The 6 insect species shipped to Australia for further testing covered a range of feeding niches on P hysterophorus. Although a root-mining insect would have been a desirable addition to this list, OF P. hysterophorus 805 no suitable candidates were identified. The 2 most common root-feeding species were Baris prob. aerea Boheman, which was tested and found to be insufficiently specific, and C arm enta ithacae (Beutenmiiller). Larvae of C. ithacae were common in the roots of P hysterophorus in Mexico and were reared to the adult stage on many occasions. C arm enta ithacae is found on species of H elenium and H eliopsis (Asteraceae) in the United States (Eichlin and Duckworth 1988). However, because no fertile eggs were ever laid in the laboratory, we were unable to -conduct host-specificity tests to determine whether C. ithacae from P hysterophorus would also accept these host plants. Other species that may have a narrow host range on P hysterophorus include the leaf-rolling moth H elcystogram m a cham bersella (Murtfeldt) and the aphid U roleucon am brosiae (Thomas). The only other host records for H. cham bersdla are from Am brosia spp. (Hodges 1986). It was never obtained in sufficient numbers for host-specificity tests to be conducted. U roleucon am brosiae is recorded from many species of Asteraceae (Robinson 1985), but material collected from P hysterophorus in Mexico would breed only on P hysterophorus and P confertum Gray in cage tests, suggesting the existence of sibling species or host races. This bears out the suggestion of Robinson (1985) that the host range of U . am brosiae may be more limited than appears from literature records. Numerous species of Cicadellidae (Homoptera) and Miridae (Hemiptera) were collected. Few feeding tests were conducted witll these species, because most did not appear sufficiently damaging to be useful as biological control agents. Many of these species are therefore indicated in Table 1 as collection records only. With further tests, it is likely that some of these could be shown to feed and develop on P hysterophorus. Adults of several species of Chlamisinae and Cryptocephalinae (Coleoptera: Chrysomelidae) were frequently collected. Larvae of these subfamilies live in a case constmcted from fecal material. Some of these species, including C ryptocephalus irroratus Suffrian, Pachybrachisim m aculatus Jacoby, Exem a conspersa Mannerheim, and D iachus auratus F., could be reared from eggs to adults on living foliage of P hysterophorus, but larvae were rarely encountered in the field. Other members of these subfamilies feed as larvae primarily on dead foliage and detritus (LeSage 1985). The indices of percentage similarity between the fauna of P hysterophorus and the 11 other plant taxa included in our comparisons ranged from 1.1 to 8.3% (Table 2). Although these are not high overall levels of similarity, the values appear to reflect the taxonomic relationships of the host plants, at least at the tribal level. The fauna of P hysterophorus is most closely related to those of other genera within the tribe Heliantheae, such as Ambrosia, H elianthus, and Xanthium . Characteristic stenophagous insect genera shared among these 806 ENVIRONMENTAL Table 2. Similarities between phytophagous arthropod faWlas of P . hysterophorus and other plant taxa Total phytophagous Plant taxon (no. species included) Am bm sia"-g (9) H elianth1/.J> ,h , i ( 1 0 ) Xanthium o.j (2) Baccharisk""P (ll) Lantana'" q ( 4 ) Parthenium argentatm n r, insect 8, b(l) Senna" (5) G utierrezitl (1) Parkinsonia'" (1 ) Prosopis" (5) CirsiumlO-<j (9) II Phytophagous speIndex of cies in common similarity , with P. % Is species hljsteropharos 274 224 197 577 550 194 121 149 120 41 433 117 37 32 44 38 20 11 11 10 13 4 8.28 8.24 7.49 5.53 4.91 4.59 2.96 2.75 2.69 1.91 1.07 o W.A.P., unpublished data. b A.S.M., unpublished data. C Harris and Piper (1970). d Goeden and Ricker (1975). C Goeden and Ricker (1976a). f Goeden and Ricker (1976b). g Goeden and Ricker (1976c). h Hilgendorf and Goeden (1981). i Rogers (1988). j Hilgendorf and Goeden (1982). k Boldt and Robbins (1987). I Boldt and Robbins (1994). Palmer (1987). ••Palmer and Bennett (1988). o Palmer and Pullen (1994). p Tilden (1951). q Perkins and Swezey (1924). r Aguirre-Uribe et aI. (1988). Stone and Fries 1986. t Foster et al. (1981). Wood (1992). "Ward et aI. (1977). Goeden and Ricker (1986). x Goeden and Ricker (1987a). Y Goeden and Ricker (1987b). m S II W plant taxa include Exem a, O phraella, Zygogram m a, C ylindrocopturus, Sm icronyx, C arolella, H elcystogram m a, and Epiblem a. The relatively high overlap with Am brosia occurs despite the fact that most of the faunal data on Am brosia are from southern California, well outside the range of our survey and the native range of P hysterophorus. There was a lower level of similarity between the fauna of P. hysterophorus and that of the congeneric species Parthenium argentatum Gray (guayule), with only 20 insect species in common. This may be because of the marked differences of plant architecture, secondary chemistry, and habitat between the 2 species. P argentatum is a small mbber-producing shmb of xeric habitats in northern Mexico, and rarely co-occurs with P. hysterophorus. In contrast, the genera Am brosia, Xanthium , and H elianthus include numerous herbaceous annual or perennial species occurring in mderal habitats similar to those occupied by P hysterophorus. Genera in the tribes Astereae (Baccharis and G utierrezia) and Cynareae (C irsium ) and the families Caesalpini- Vol. 24, no. 4 ENTOMOLOGY aceae (Parkinsonia, Senna) and Mimosaceae (Prosopis) had lower similarities with the fauna of P hysterophorus, reflecting their greater taxonomic distance from Parlhenium . Lantana cam ara had a surprisingly high degree of similarity (4.91%) with P hysterophorus; the reasons for this are unknown. These faunal relationships are of importance to the practical aspects of selecting weed biological control agents. The relatively high degree of similarity between the faunas of P. hysterophorus and H elianthus validated the selection of H. annuus as a critical test plant. Thirty-nine species (14.8%) of the fauna of P hysterophorus have also been recorded from H elianthus, making them ineligible for further consideration as biological control agents. This close association between the faunas of P hysterophorus and H. annuus was probably the most important factor limiting the number of biological control agents obtained from this project. The choice of Monterrey in northeastern Mexico as the main project base seems to have been fully justified. All 6 of the species sent to Australia were found in the immediate vicinity of Monterrey, whereas only S. lutulentus, E. strenuana, and S. concinna were found in proximity to the base at Temple, TX, and only E. strenuana and S. concinna were found near Cuernavaca. One indicator of potentially good biological control agents is their effect on the host plant in its native range (Wapshere 1985, Crawley 1989). Severe insect damage on P hljsterophorus was rarely observed in our surveys. Zljgogram m a bicolorata occasionally became abundant enough to cause defoliation, and on one occasion a young stand was severely stunted by heavy E. strenuana infestation. These 2 species have been the most effective biological control agents when released in India and Australia (McFadyen 1992, Jayanth and Bali 1990). During the cooler winter months, plants were often coated with dense colonies of U . am brosiae. Occasional defoliation by polyphagous Lepidoptera was seen in Texas. The absence of native or economic species of Ambrosiinae in Australia facilitated the introduction of biological control agents into Queensland against P. hysterophorus, as it was possible to release several species which were oligophagous on Parlhenium , Am brosia, or Xanthium , rather than strictly monophagous. This implies, however, that caution should be exercised in releasing any of these agents in other countries where P hysterophorus is a problem. Any such releases should be preceded by a careful review of the status of potential host plants present in the proposed area of introduction, and, if necessary, additional host specificity tests should be performed. Acknowledgments We thank Hugo Miranda Sanchez, Hugo Perales Rivera, and Manuel Valencia for technical assistance, Dieter August 1995 M cC LA Y ET A L .: ARTHROPODS Enkerlin ,md Juan D. Vega for providing facilities at ITESM, Marion Herbut for assistance with graphics, and L. Dosdall, R. Casey, R. E. McFadyen, and F. D. Panetta for their review of the manuscript. We also thank the lJ niversidad Aut6noma del Estado de Morelos for providing facilities in Cuernavaca. We particularly thank the expert ta'(onomists who identified the arthropods, and often also offered valuable advice on their biology and habits: D. M. Anderson (USDA Systematic Entomology Laboratory, Washington DC [SEL], Coleoptera; Curculioni<lae); J. D. Bradley (International Institute of Entomology, London [IlE], Lepidoptera); R. L. Brown (Mississippi Entomological Museum, Lepidoptera; Tortricidae); J. A. Chemsak (University of California, Berkeley, Coleoptera; Cerambycidae); M. L. Cox (IlE, Coleoptera: Curculionidae, Scolytidae); D. R. Davis (National Museum of Natural History, Washington, DC [NMNH], Lepidoptera; Lyonetiidae); C. H. Dietrich (SEL, Homoptera: Auchenorrhyncha); E.A.J. DuffY (IIE, Coleoptera: Chrysomelidae); T. D. Eichlin (California Department of Food and Agriculture, Lepidoptera: Sesiidae); D. C. Ferguson (SEL, Lepidoptera; Geometridae, Pterophoridae, Pyralidae); R. H. Foote (SEL, Diptera: Tephritidae); R. Gagne (SEL, Diptera: Cecidomyiidae); M.S.K. Ghauri (IIE, Hemiptera, Homoptera); R. D. Gordon (SEL, Coleoptera; Scarabaeidae, Coccinellidae); K. M. Harris (IIE, Diptera: Agromyzidae); T. J. Henry (SEL, Hemiptera); H. A. Hespenheide (University of California, Los Angeles, C ylindrocopturus); R. W. Hodges (SEL, Lepidoptera: Gelechiidae); J. D. Holloway (IIE, Lepidoptera); F. Hovore (Placerita Canyon Nature Center, [Coleoptera: Cerambycidae); J. M. Kingsolver (SEL, Coleoptera: Bruchidae); J. P. Kramer (SEL, Homoptera; Auchcnorrhyncha); D. Macfarlane (lIE, Acari; Tetranychidae); R. B. Madge (lIE, Coleoptera; Languriidae); J. A. Marshall (lIE, Orthoptera: Gryllidae); J. H. Martin (lIE, Homoptera; Aleyrodidae, Aphididae); D. R. Miller (SEL, Homoptera; Coccoideae); D. A. Nickle (SEL, Orthoptera: Acrididae); A. Norrhom (SEL, Diptera; Tephritidae); C. W. O'Brien (Florida A&M University, Coleoptera; Curculionidae); J . M. Palmer (lIE, Thysanoptera); R. W. Poole (SEL, Lepidoptera: Noctuidae); J. Razowski (Polish Academy of Sciences, C arolella); F. H. Rindge (American Museum of Natural History, Lepidoptera; Geometridae); J. Schaffner (Texas A&M University, Hemiptera: Miridae); T. J. Spilman (SEL, Coleoptera; Cerambycidae, Elateridae); M. B. Stoetze1 (SEL, Homoptera: Aphididae); M. R. Wilson (lIE, Hemiptera: Miridae); R. E. White (SEL, Coleoptera: Chrysomelidae, Cerambycidae); the late D. R. Whitehead (SEL, Coleoptera; Curculionidae); and D. J. Williams (IlE, Homoptera: Pseudococcidae). Funding for the study was provided by the Rural Lands Protection Board, Queensland, Australia. References Cited Aguirre-Uribe, L. A., J. Corrales-Reynaga, and A. Lozoya-Saldana. 1988. Catalogo de insectos asociados aI guayule Parthenium argentatum Gray. Agraria, Universidad Aut6noma Agraria Antonio Narro, Saltillo, Mexico. 4: 60-71. Bernays, E. A. and R. F. Chapman. 1994. Host-plant selection by phytophagous insects. Chapman & Hall, New York. Boldt, P. E. and T. O. Robbins. 1987. Phytophagous and pollinating insect fauna of Baccharis neglecta OF P. hysterophorus 807 (Compositae) in Texas. Environ. Entomol. 16; 887895. 1994. Phytophagous insect faunas of Baccharis salicina, B. pteronioides, and B. bigelovii (Asteraceae) in the southwestern United States and northern Mexico. Environ. Entomol. 23; 47-57. Crawley, M. J. 1989. Insect herbivores for the biological control of weeds. Plants Today 2: 59-64. Dharmaraj, G., R. Chandra Babu, N. Natarajatnam, and S. Subramaniam. 1988. Allelopathy of certain weed species. Madras Agric. J. 75: 147-148. Eichlin, T. D. and W. D. Duckworth. 1988. Sesioidea; Sesiidae. In R. B. Dominick, D. R. Davis, T. Dominick, D. C. Ferguson, J. G. Franclemont, R. W. Hodges, E. G. Munroe, and J. A. Powell [eds.], The moths of America north of Mexico, fascicle 5.1. The Wedge Entomological Research Foundation, Washington, DC. Evans, H. C. 1987. The life cycle of Puccinia abrupta val'. partheniicola, a potential biological control agent of Parthenium hysterophorus. Trans. Br. Mycol. Soc. 88; 105--111. Foster, D. E., D. N. Ueckert, and C. J. DeLoach. 1981. Insects associated with broom snakeweed (Xanthocephalum sarothroe) and thread-leaf snakeweed (Xanthocephalum m icrocephala) in west Texas and eastern New Mexico. J. Range Manage. 34; 446454. Goeden, R. D. and D. W. Ricker. 1974a. The phytophagous insect fauna of the ragweed, Am brosia acanthicarpa, in southern California. Environ. Entomol. 5; 827-834. 1974b. The phytophagous insect fauna of the ragweed, Am brosia cham issonis, in southern California. Environ. Entomol. 5; 835-839. 1975. The phytophagous insect fauna of the ragweed, Am brosia confertiflora, in southern California. Environ. Entomol. 4; 301-306. 1976a. 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Phytophagous insects reported worldwide from the noxious weeds spiny c1otbur, Xanthium spinosum , and cocklebur, X. stn.t1Tw rium . Bull. Entomol. Soc. Am. 287: 147-152. Hodges, R. W. 1986. Gelechioidea: Gelechiidae (part). In R. B. Dominick, D. R. Davis, T. Dominick, D. C. Ferguson, J . G. Franclemont, R. W. Hodges, E. G. Munroe, and J . A. Powell [eds.], The moths of America north of Mexico, fascicle 7.1. The Wedge Entomological Research Foundation, Washington, DC. Jayanth, K. P. 1987. Introduction and establishment of Zygogram m a bicolorata on Parthenium hysterophoros at Bangalore, India. Curro Sci. (Bangalore) 56: 310-311. Jayanth, K. P. and G. Bali. 1990. Evaluation of the efficacy of Zygogram m a bicolorata in suppressing Parthenium hysterophoros at a field release site in Bangalore. National Symposium on Conservation and Management of Living Resources, 10-12 January 1990, Bangalore, India. LeSage, L. 1985. 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