COMMUNITYANDECOSYSTEMECOLOGY

Phytophagous Arthropods Associated with Partheniumhysterophorus (Asteraceae) in North America

A. S. McCLAY, W. A. PALMER,l F. D. BENNETf,2 ANDK. 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 forParlhenium 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'snative range in North Americawas sulVeyedfrom bases at Monterrey and Cuernavaca, Mexico,and Temple, TX. Two hundred and sixty-twophytophagous arthropod species were collectedon P. hysterophorus by various methods including hand picking,dissection, rearing, and sweep-ing; 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%), Lep-idoptera (20.2%), and Hemiptera (18.3%). Twofungal pathogens, Puccinia abrupta Dietel andHolwayvarietypartheniicola (Jackson)Parmelee and Puccinia melampodii Di'eteland Holway,were also obselVed.An index of similaritywas used to make pairwise comparisons betweenthe phytophagous arthropod communities on different plant taxa. These comparisons showedthat the fauna of P. hysterophorus is most similar to that of ragweeds (Ambrosia spp.). Sixinsect species that were shownto be stenophagouswere shipped to Australiafor further testingand possible field release. One of the fungal pathogens underwent host-range testing in theUnited Kingdom and was released in Australia.

KEY WORDS Parlhenium hysterophonJs, Asteraceae, biologicalcontrol, phytophagous, ar-thropod fauna, North America

Parthenium hysterophorus L. (Asteraceae) is a her-baceous annual weed native to the neotropics,from the southern United States to Argentina,which has become widely naturalized in othertropical and subtropical regions. It has become aparticularly serious problem in Queensland, Aus-tralia (Haseler 1976) and in India (Towers et al.1977). It also occurs in Taiwan, southern China,Vietnam, Vanuatu, New Caledonia, Kenya, Mada-gascar, Mozambique, South Africa, the Mascarenesand Seychelles, and Ethiopia. In Queensland,thousands of square kilometers of rangeland areinfested, and pure stands of P hysterophorus haveexcluded beneficial pasture species over large areas(McFadyen 1992). All parts of the plant containparthenin, a bitter, toxic sesquiterpene lactone,which makes the plant unpalatable to cattle andhas caused many cases of severe contact dermatitisin susceptible people (Towers et al. 1977). Its pro-fuse seed production enables it to spread rapidly.The plant is reported to be allelopathic (Dharma-raj et al. 1988).

lTropical Weeds Research Centre, P.O. Box 187, Charters Tow-ers, Queensland 4820, Australia.

2Department of Entomolow and Nematology, University ofFlorida, Gainesville FL 326JJ-D740.

36 Dianella Road, Port Macquarie, New South Wales 2444,Australia. .

The center of diversification of the genus Par-thenium is considered to be in northeastern Mex-ico, where at least 8 of the 16 described speciesoccur (Rollins 1950). The general climatic char-acteristics of this area are similar to those of theinfested areas of Queensland (McClay 1985).

The cost of chemical and cultural control oflarge stands of P hysterophorus on low-value pas-ture is prohibitive (McFadyen 1992). In an attemptto identify alternative control measures for this se-rious rangeland weed, the Queensland Depart-ment of Lands (QDL) started a biological controlprogram in 1975. As part of this program, scientistsof the Department of Lands and of the Common-wealth (now International) Institute of BiologicalControl (CIBC, now nBC) undertook explorationin North and South America. These staff were re-sponsible for conducting faunal studies of P hys-terophorus and selecting appropriate stenophagousspecies for host specificity and biological studies.If these tests indicated that a species was a poten-tial biological control agent, it was shipped to theSir Alan Fletcher Research Station of the Queens-land Department of Lands in Sherwood, Australia,for further testing, approval, and eventual release.

This article reports the phytophagous insectsand mites found in North America during theseinvestigations. A preliminary list of 159 phytopha-gous insects associated with the plant in Mexico

0046-225X195/0796-0809$02.00/0 © 1995 Entomological Society of America

August 1995 McCLAY ET AL.: ARTHROPODS OF P. hysterophorus 797

Fip;. 1. Searcharea for naturalenemiesof P. hysterophoros in the United Statesand Mexico(shaded).NumbersidentifYthe regionsused to indicategeographicaldistributionin Table 1.

has been published (McClay 1981) and the speciessent to Australia for further testing were discussedby McClay (1985). In addition to the surveys re-ported here, QDL staff conducted surveys ofSouth American populations of P. hysterophorus inBrazil and Argentina (McFadyen 1976; QDL, un-published data), and some collections were madein the Caribbean by F.D.B. and other CIBC staff.The biological control program for P. hysteropho-ruo\' was reviewed by McFadyen (1992).

Materials and Methods

The area covered by this survey (Fig. 1) com-prised most of Mexico and parts of the southeast-ern United States. Two preliminary surveys wereconducted over a total of 40 d in June, October,and November 1976 by F.D.B. (unpublished data,1977). During these surveys, collections weremade in the states of Veracruz, Yucatan" Campe-che, Quintana Roo, Puebla, and Nuevo Le6n,Mexico, and in the areas of Houston and CollegeStation, TX, Baton Rouge, LA, and Miami andHomestead, FL. More detailed studies were thenconducted by A.S.M. from 1978 to 1983 from abase at the Instituto Tecnol6gico y de Estudios Su-periores de Monterrey (ITESM) at Monterrey,Nuevo Le6n. During this period, extensive collec-tions were made in northeastern Mexico, particu-larly the states of Nuevo Le6n, Tamaulipas, and

eastern Coahuila. Some collections were also madealong the Gulf coastal areas of Veracruz and Ta-basco, and on the Pacific coast from Sinaloa toNayarit and from Oaxaca to Chiapas. From 1983to 1993, surveys were conducted in Texas byW.A.P.from a base at the Queensland Departmentof Lands North American Field Station (NAFS) inTemple, TX. From 1988, W.A.P and K.R.P. madeadditional surveys in southern Mexico, particularlythe state of Morelos, from a substation at Cuer-navaca, Morelos.

Collections were made on natural stands of P.hysterophorus on roadsides, wasteland, neglectedfields, and similar sites at frequent intervalsthroughout the year. Arthropods resting on theplant were captured directly by hand or with anaspirator, or by sweeping pure stands of P hyster-ophorus with a sweep net. All parts of the plantwere dissected for endophagous species. Adults ofroot-feeding species were obtained by keepingroots in emergence boxes with glass vials fittedover the exit holes or in large glass jars coveredwith nylon gauze. Other plant parts were usuallytoo perishable to be held in the laboratory longenough for emergence of adult stages of endo-phages. Some stem and root-feeding insects werereared to the adult stage in the laboratory by feed-ing ReId-collected larvae on an artificial diet (Har-ley and Willson 1968).

798 ENVIRONMENTAL ENTOMOLOGY Vol. 24, no. 4

In addition to these collections on naturalstands, a series of plots (5 m by 5 m) were sownwith P. hysterophorus at the ITESM field station,Apodaca, Nuevo Le6n, and allowed to become in-fested with arthropods. Samples were taken fromthe plots at 2-wk intervals from 1978 to 1982 byselecting 25 plants at random and collecting all ar-thropods resting on them or feeding internally inthem.

Most specimens were initially submitted foridentification either to the United States Depart-ment of Agriculture Systematic Entomology Lab-oratory, Beltsville, MD, or to the Commonwealth(now International) Institute of Entomology, Lon-don. Specimens that could not be identified to spe-cies by these laboratories were later forwarded toother taxonomists. Many submitted specimenswere identified only to genus because of inade-quate material, unavailability of type specimens, orlack of satisfactory taxonomic revisions. These arereported here only if they represent a genus nototherwise reported or if the taxonomist explicitlystated that they are distinct from the other namedspecies of the genus reported here.

The probable level of host specificity of eachspecies was assessed using information from theliterature, specialists, and field observations. Spe-cies that appeared likely to be host specific, at leastthe subtribe level, were selected for preliminaryhost-specificity studies conducted in Mexico andTexas. Test plants were selected from the subtribeAmbrosiinae and tribe Heliantheae, to which thegenus Parthenium belongs (Stuessy 1977), with theaddition of some economic species from othertribes of the Asteraceae. Sunflower, Helianthus an-nuus L., is the most important crop species closelyrelated to Parthenium and was considered a criticaltest plant (Wapshere 1975). Experimental detailsdepended on the biology of each species tested,but many tests were conducted in multiple-choicecages as described by McClay (1987). Field obser-vations of the insects' host plant associations werealso made.

If our investigations indicated that the hostrange of an insect was within the subtribe Ambro-siinae, the insect was recommended for furtherstudy and shipped to Australia after quarantine ap-proval had been received.

In our field surveys, it was not always possibleto determine whether a captured insect was actu-ally feeding on P hysterophorus. Published faunallists for many plants frequently do not in~catewhether the species listed are known to feed onthe plant in question or were simply collected onit. We report here all the species belonging toknown phytophagous taxa that we collected on P.hysterophorus, with the exceptions of pollen ornectar gatherers, adult Lepidoptera and Diptera,species shown in laboratory tests not to feed on Physterophorus, and species whose known biolog.yclearly indicates that P. hysterophorus is an unlike-ly host. Where any life stage of an insect species

has been observed to feed on P hysterophorus, thisfact is indicated in Table 1. Records where feedingis not indicated refer only to collections of adultson the plant; this category undoubtedly includessome species which are casual visitors on P. hys-terophorus.

Specimens of many of the species identifiedwere retained by the taxonomists responsible foridentification, and were deposited mainly in theNational Museum of Natural History in Washing-ton, DC, and the Natural History Museum in Lon-don. Material of all species collected from 1978 to1983 was deposited in the collection of the IIBCCaribbean and Latin American Station, Trinidad,and at NAFS. Additional material from the years1978 to 1983 was deposited in the collection atITESM. Material collected in 1983 and afterwardwas placed in the NAFS collection, now held atthe Tropical Weeds Research Centre (TWRC),Charters Towers, Queensland. Inquiries concern-ing the location of particular material collectedduring the project may be directed to W.A.P. orA.S.M. Relevant information about each specieswas also entered into a computer database nowmaintained at TWRC (Palmer 1995); detailed da-tabase records for particular species may be ob-tained from W.A.P.

We assessed the degree of similarity betweenthe fauna of P. hysterophorus and those of 1 otherParthenium species, 6 other native North Ameri-can genera of Asteraceae, and 4 native NorthAmerican genera in other families. A pairwise in-dex of p.ercentage similarity Is was defined by

Is = 100X(llshareinpooled)

where llsharedis the number of arthropod speciesin common between the 2 plant taxa and npooledisthe total number of species in the pooled faunasof both taxa.

Results and Discussion

Phytophagous arthropods associated with P. hys-terophorus in Mexico and the United States arelisted in Table 1. Information on host specificityand pest status in this table is a synthesis of theresults of feeding tests carried out during the pro-gram, field observations, literature records, and in-formation from specialists who identified the in-sects. Species that were subjected to host rangetests during the project are indicated. The num-bered regions used to identify geographic distri-bution are shown in Fig. 1. Distribution records inTable 1 represent only data from our surveys anddo not include information from the literature.Many species may, therefore, have wider distribu-tions than indicated in the table.

Table 1 also indicates frequency of collection foreach species. Because collections were not quan-titative, the assessment of whether a given speciesis common, occasional, or rare is based on a sub-jective assessment of the collecting experience of

August 1995 MCCLAY ET AL.: ARTHROPODS OF P. hysterophorus 799

Table 1. Phytophagous arthropods associated with P. hysterophorus

Fre- Stages Feeding Feeding Host PestSpecies quenc)" feedingi' site" typed speci- Testedf Distributionf; status"6city"

AcariEriophyidae

Enophyes nr. parthenii Keifer 0 N,A F ECT ***** T

Tetmnychidae1i'tranyehlls nr. l.uleni Zacher G N,A L ECT **

OrthopteraGryllidae

Ocean thus nr. celerinictus T. walker 0 N,A L ECT

Tt'ttigoniidaeCOlwcL'}Jhalu.sfascilltuS (DeGeer) R * 8

ThysanopteraThripidat'

Clliiothrips phaseoli (Hood) G N,A L ECT * PMicrocL'}Jholothrips abdominalis (Crawford) 0 N,A F ECT **

HemipteraAlydidae

Burtinus fenwralis (Distant) R

Bt'rytidaeJalyslls reductlls Barber R 5

Cor!'idaeChariesterus lIwestus Burmeister R 5Nirooeclls cllloiger StAl R 1Pllthia picta (Drury) R 5Sagotylus conf/llens (Say) C N,A S ECT * l' 1,3,5 P

LygaeidacAcrolcucus sp. R 5Blissus IL~,copterns (Say) R 7 PNeopamem bi/obata (Say) R * 5NlJsius senecionis (Schilling) C A F ECT *** ]

N'.Isius sp. C A F ECT 1XyonysillS califomiCllS (StAl) C A F ECT 1,5

MiridaeCollana oleosa (Distant) 0 * 5Cyrtopeltis sp. 0 5HaltiClls bmctatus (Say) R * 12 P[[')(leom i'lllsta (Distant) 0 ** 5LlJg,lS [;,leolaris (Palisot de Beauvois) 0 * 5, 8 PNmrocolp.lS prob. mexicallllS Distant R 5PhlJtocoris sp. R 5Poecilocapslls nigriger (StAl) R N,A L ECT * 1Polymerus basalis (Reuter) R * 12PollJmenlS testaceipes (StAl) 0 * 5Proba distanti (Atkinson) C N,A F,L,S ECT ** l' 1,2,4,5,7Prohll sll!lei (Stll) R ** 5RCiiteroseopus omatllS (Reuter) C N,A F,L,S ECT ** T 1,2,4,8, 10Rhinacloa forticomis (Reuter) R 5 PSixeonotus sp. R 5Taedia sp. R 5TalJlorilygus pallidulus (Blanchard) C N,A F,L,S ECT * 1,5, 7, 8, 10

PentatomidaeChloroehroa ligata (Say) R * 12 PDendroeoris sp .. R 12E./.Schistus quadmtor Rolston 0 N,A L, S ECT * 1,7EuseltishlS sp. R 2, 7Momlidea notlliata (Herrich-Schaffer) R * 7Tltyanta aeeelTa McAtee R * 8 PTltyanta pemitor (F.) R * 5 P

PiesmatidaePiesma cinereum (Say) 0 P

800 ENVIHONMENTAL ENTOMOLOGY Vol. 24, no. 4

Table 1. Continued

Fre- Stages Feeding Feeding Host PestSpecies speci- Tested!" Distributio~quenc)" feedingh site" typed 6city' statmJ•

PyrrhocoridaeDysdercus mimulus Hussey R * 1 P

RhopalidaeHarrrwstes nebulOSlIS StM R ** 1,5Harrrwstes rejlexulus (Say) 0 ** 1Hannostes serratus (F.) R * 3LiorhysstlS sp. R 1

ScutelleridaeChelysomidea oariabilis (Herrich-Schaffer) R N,A L, S ECf * 1Homaemus proteus St,u 0 1,5Sphyrocons obliquus (Germar) 0 1

TingidaeCO"fthucha morrilli Osborn & Drake C N,A L,S ECT * T 1,2,5,8 PDietyla 11wnotropidia (St,u) R * 11Leptodictya tabido (Herrich-Schaffer) R * 5

HomopteraA1eyrodidae

Trialeurodes abutilonea (Haldeman) G N ·L ECf * 1 P

AphididaeAphis gossypii Glover G N,A L, S ECT * l PGeopemphigus floccOstlS (Moreira) C N,A R ECf * 1Uroleucon ambrosiae (Thomas) C N,A S,L ECf ** ••• T 1, 2, 3, 4, 7, 8, 9

CercopidaeAeneolamia sp. R 3Clastoptem xanthocephala Germar C A L ECf 8Prosapia sp. R 3

CicadellidaeAceratagallia gillettei (Osborn & Ball) C N,A L, S ECT * T 1Agallia barretti Ball R * 5Agallia quadripunetata excavata Oman C * 5Agrosoma pulchella (Guerin) R * 2,5Egidemia inf/ata Young R 3Empoasca ahmpta DeLong C N,A L, S ECT * 1 PEmpoasca or. crocostigmata Davidson

& DeLong 0 5 PEmpoasca nigra var. typhlocyboides

Gillette & Baker C N,A L, S ECT ** 1Graphocephala mjimargo (Walker) C A L ECT * 5, 7Gyponana sp. R 1,2,3Homalodisca coagulata (Say) 0 A S ECT * 1 PHomalodisca insolita (Walker) R A L, S ECT * 5 PMacrosteles sp. R 5Momoria misella (Ball) R ** 7Oncometopia elarior Walker 0 * 1,3Penestragania romlSta (Uhler) C N,A L, S ECT * T 1,2,4,5,8 PPhera lanei Young R 5Phera obtusifrons (Fowler) R 5Scaphytopius frontalis (Van Duzee) R * 8Scaphytopius jiJliginostlS (Osborn & Ball) 0 N,A L, S ECT * T 1,5 PSibooia compta (Fowler) R * 5TexananrlS ooatus (Van Duzee) 0 * 1Xerophloea viridis (F.) 0 N,A L, S ECT * T 1

CoccidaeCeroplastes cirripedifannis Comstock R N.A S ECT * 3 PLecanium sp. R N,A L ECT 5Parasaissetia nigra (Nietner) R N,A S ECT * 2, 3 PSaissetia miranda (Cockerell & Parrot) R N,A S ECT * 3 P

DelphacidaeStobaera concinna (St,u) C N,A L,S ECT * ••• T 1,4,5

DictyopharidaeScalops sp. C N,A L, S ECf 1

Augllst 1995 McCLAY ET AL.: ARTHROPODS OF P. hystemphorus 801

Tuble 1. Continued

Fre- Stages Feeding Feeding Host PestSpecies speci- Testedf Distribution(;quem:>", feeding" site" typed licit}" statush

Eriot'o(x'idaeAmnthocoeeus dubius (Cockerell) R N,A S ECT * 3

Flalidal'FZlltO"ucnL. sp. 0 5

Issidal'CoZpol'tem lllbavCrlosa Caldwell R 2

Margarodidaelrenja nr. littom/is Cockerell R N,A L ECT 3lcerya l'urchllsi Maskell R N,A S ECT * 3 P

Ml'mbmeidaeAcutalL. tar/area (Say) R 12Entyl/a cnnnata (Forster) C N,A S, L ECT ** 1,5,12Mien/talis sp. C 1,2,51'0/yglWtll sp. R 5Publ/lla sp. 0 1,4,5SI,L.sL.Hlus/esHnus (Say) R * 1,2,12 PStictoe17,hala sp. 0 1Umbonill prob. crass/comls Amyot& Serville 0 * 5

Vanduzcll segl1u.'fItata (Fowler) R * 1

Ortht'7jidaeOrtlwz/a pmelonga Douglas R N,A L, S ECT * 11Orthr.;/a I'SetulillSignis Morrison R * 3

PseudococcidaeFerrisia virgata (Cockerell) C N,A L, R, S, M ECT * 1,2,3,5 PParacoccus sp. R N,A S ECT 2Phenacoeeus nr. defectus Ferris G N,A S, M ECT 1Phenaeoecus gossypii Townsend& Cockerell 0 N,A S ECT * 3 P

Phenaeoceus solani Ferris 0 N,A R ECT * 1,2 PPsmdoc.llcCllSaffinls (Maskell) 0 N,A S ECT * 11 PSl'ilococclls sp. R N,A F ECT * 2,3

PlltoidaePuto barben (Cockerell) R N,A S ECT * 11 P

ColeopteraAnthribidae

Tngonorhinr •.• limbatus (Say) R 1

ApionidaeChnjSal'ion auctu", (Sharp) R 5

BntehidaeAc(mthoseelid,,.. sp. R 5

C<'rambycidaeDectes lIigril'ilus Chemsak & Linsley R 4Dect"s sayi Dillon & Dillon C L,A S, L END ** 1Dectes t.':tanr•.• LeConte 0 L,A S, L END * 4,5 PDorcasta ciril'rea (Hom) R * 1 PHippopsis lemniscata (F.) 0 L,A S, L END * 1,2,3,10 PPhaea tenrwta Bates R 5Pseudoeanidia cuemavacae Dillon 0 L S END *** T 5

ChrysomelidaeAcaZymmalJlomorum Munroe & Smith R * 3 PAgathomen~' nifi •.• Klug R 5AnopliHs sp. R L L END 2CluU'tocnel1U1homi Jacoby R 11Ghal17JUsnr. conHg"'''' Baly R A L ECT 1Ghrysodina sp. R 5C"7'idodem sp. R 5Cnjptocephall~' irrorall •.• Suffrian 0 L,A L ECT .*** T 1Cnjptocephlll, •.•prob. nigrovittall •.•Jacoby 0 L,A L ECT 1CryptocephaZus sp. 0 L,A L ECT 1

802 ENVIRONMENTAL ENTOMOLOGY Vol. 24, no. 4

Table 1. Continued

F~e- Stages Feeding Feeding Host PestSpecies speci- Testecif Distributionllquency' feedingb site" typed ficity' status"

Derocrel'is sp. R 5Diabrotica ba/teata LeConte a 1,3,5,7 PDiabrotiEa undecimpunctata Jacoby R * 11Diachlls aumtlls (F.) C L,A L ECT ** T 1,5,8Disonycha knabi Blake R 5Durallgoita chrysodinoides (Jacoby) R 1El'itrix sp. R ]

Exema prob. conspersa (Mannerheim) C L,A L ECT ** T 1,2,5,7Exema or. dispar Lacordaire a ** 5Glyptina sp. R 1G/yptosce/is nr. mexican a Jacoby R 11or. Hippllriphila sp. R 5Lobidomem sp. R 1Lexiphanes gllerini (Perbosc) C 5Lllpraea sp. R 7Microrhopa/a pulchella Baly R 3Microrhopala prob. sallei Baly R 5Nodonota nr. atm Harold R 11Omophoita aequinoctialis (L.) R * 1Ophraella communa LeSage a L,A L ECT *** T 1,2,5Ophraella prob. slobodkini Futu)'lna a L,A F ECT *** T 2Pachybrachis or. fenwmta Olivier R 11pachybrachis immacu/atus Jacoby a L,A L ECT *** T 1,4Pachybrachis nr. vestigialis Fall a 5Pachybrachis sp. a L,A L ECT *** T 1psylliodes sp. R 11Systena b/anda Melsheimer C 11 PSystena nigrop/agiata Jacoby R 5Systena nr. 1)ectom/is Clark a 1,2Systena or. semivittata Ja(.'Oby R 3Systena sp. C A L ECT 1Zygogmmma bicolorata Pallister C L,A L ECT **** T 1,2,4,5Zygogmmma ma/vae St:ll R A L ECT 5

CoccinellidaeEllilachna mexicana (Guerin) R * 7 P

CurculionidaeBaris prob. aerea Boheman C L,A R END ** T 1,2,5,6,7Baris prob. interstitia/is Say a L,A R END ** 1,2,6Baris sulcipennis (Brisout) R 5Cha/codermus sp. R L S END 1Coml'sllS auricepha/us (Say) a A L ECT * ]

Cylindrocopturus annat!lS Champion a L,A R, S END *** T 1,2,7Cy/indrocopt!.rus or. mammillatus (LeConte) R 5Cy/indrocoptunlS tetra/ob,lS Champion R 5EpicaenlS allrifer Boheman R * 5 PEpicaerus prob. texanus Casey a 1GeraellS picrmmus (Herbst) R 1,5Geraeus seni/is (Boheman) R 5Linogemeus sp. R 5Listroderes costirostris ob/iquus Klug R L,A L * 1 PLixus prob. scrobicollis Boheman C L,A S, L END *** T 1,2,4Mitostyl.lS SetOSllS(Sharp) a A L ECT ** 1,4Onychobaris sp. a 1Pande/eteius sp. R 1Rhodobaenus quinquepunctatus (Say) a L,A S END ** 8Rhodobaenus sanguineus (Gyllenhal) C L,A S END ** 2,5Rhodobaenus tredecimpunctatus (Illiger) R L,A S END *** 1,2SciadrtlstlS propheticus Sharp R 5Sibinia sp. R * 1,5Smicronyx lutulent!.s Dietz C L,A F END ***** T 1,2,4,8Smicronyx sordidus LeConte R *** 1Thecestemus hirst/tus Pierce a L,A R ECT lit •••••• T 1, 8Trepobaris inonurta Champion R 5

LanguriidaeDasydactyZ,lS sp. R L S END 2Languria mozardi Latreille a L S END * 1 PLanguria or. /aeta Leconte R L S END 2Longurites lineat!.s (Castelnau) R L S END ** 1

August 1995 McCLAY ET AL.: ARTHROPODS OF P. hysterophorus 803

Tobie 1. Continued

Fre- Stages Feeding Feeding Host PestSpecies speci- Testedf Distributim,gquene:)'" feeding> site" typedlicit}" statui'

ScarnblleidaeEuphoria sp. R 5

ScolytidaeH!lIlOthellClIlIlSnr. beameri Wood R L S END 1

LepidopteraArctiidlle

Estigmene (l(:rea (DnJry) a L L ECT * 1 PHa/isidota catenulata Hubner R L L ECT * 2

CelechiidaeDiehomeris nr. aenigllUltica (Clarke) R L L ECT ** 2Diehomeris simpliciella (Busck) or or. a L L ECT 1Glyphidocera sp. R L L ECT 2G'lOrimoschaCIIUlsaphirillella Chambers R L L ECT **** ]

lIeleystogramma chambersella (Murtfeldt) a L L ECT **** 1Geometridae

Anacomptodes prob. herse (Schaus) R L L ECT 5A'lll('(lmptodes perfectaria McDunnough C L L ECT *** T ]

Allavitrinella pampinaria (Guenee) R L L ECT * 1 PCMorocMamys chloroletlcaria (Guenee) C L L ECT * 1CyclopllOra nanaria Walker C L F,L ECT ** 1EUIJithecia misemlata (Grote) or near R L L ECT * 1,8lIyponu:cis or. umbrasaria Hubner R L L ECT * 8Pleuropmcha asthenaria (Walker) R L F ECT * 5 PPleuropmclUl insulsaria (Guenee) R L F ECT * 3, 8 PSalmlodes arga Druce R L L ECT 5Synehlora frondaria Guenee C; L F,L ECT * 1,8TOnJos abjeGtarillS Guenee R L L ECT **** 1

CradllllriidllePhyllononJcter sp. R L L END

LyonetiidaeBuc,,,latrlx mnbrosiaefoliella Chambers G L L END *** 1Bucculatrix nr. criticopa Meyrick R L L END *** 1Bucmllllrix pfltthenica Bradley a L L END .*.** T 1,4

NoctuidaeCondiea sutor (Guenee) R L L ECT * 8IIelieoverpa zea (Boddie) G L F ECT * 1 PLophoceramica pyrrha Druce R L L ECT * 5Ogdoconta cinereola (Gul'nee) a L L ECT * 1,8PscudopltlSia illc/udens (Walker R L L ECT * 1,8 PSpodoptera latifascia (Walker) R L L ECT * 2 PSpragu";a guttata Grote a L L ECT 1

NymphalidaeCMosy"c lacinia Geyer C L L ECT *** T 1,2,4 P

PterophoridlleOidaematophonls fumivcntris (Zeller) R L 5OidaematopllOnlS inquinatus (Zeller) a L L ECT *** 1,2OidaematopllOnlS stramillCUS (WaIsingham) R L S END 1

PyralidaeAchyra ranta/is (Guenee) R L L ECT * 1,8 PConchylodes ovu/alis (Guenee) a L L ECT ** 1

HiodinidaeCalephelis /avenJa Godman & R L L ECT **

Salvin or nr.

SllturniidaeHylesia sp. R L L ECT 2

ScsiidaeCamlellta ithacae (Beutenmtiller) C L R END *** 1, 2

TineidacAcrolophus sp. R L R END 2Tinea sp. R L M END 7

804

Table 1. Continued

ENVIRONMENTAL ENTOMOLOGY Vol. 24, no. 4

Species Fre- Stagesquen<-1' feeding/'

Feedingsite"

Feeding Hos~ Teste-"typed specI- (]I

ficity'Distribution'l Pest

statu1'

TortricidaeAethesoides distigmatana Walsingham

or nearAmoroia emigratella BusckAmorbia or. rectangularis MeyrickArgyrotaenia montezumae (Walsingham)Carolella or. buscana ComstockCochylis subolivacea WalsinghamEpiblemn strenuana (Walker)Platynota labiosana Zeller or nearPlatynota nigroceroina WalsinghamPlatynota rostTana (Walker)Platynota stultana WalsinghamSulei11Ulskinnerana Heinrich

AgromyzidaeAgromyza sp.Arnouromyza maculosa (Malloch)Liriomyza sp.Melanagromyza sp.

CecidomyiidaeContarinia sp.Lasioptera sp.

TephritidaeEuarestoides acutangulus ThomsonEutTeta augusta Banks

R L R END 2,50 L L ECT * 5R L L ECT 20 L L ECT * 2,3,50 L R, S END 1,4R L M END 7C L S END **** T 1, 2, 3, 4, 5, 8, 10C L L ECT 1,2,70 L L ECT * 8R L L ECT * 8C L L,F ECT * 10 L S END *** 5

Diptem

R L L END 10 L L, M END * 10 L L END 1C L S END 1,2,4,5

R L F ENDC L S END .****

C L F END **R L S END **

P

PP

P

a C, common; 0, occasional; R, rare; G, found in greenhouse but not in field.b 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, Mi-choac:in (coastal), Jalisco (coastal); 8, Texa~; 9, Louisiana; 10, Florida; 11, Mexico, unspecified; 12, United States, unspecified.

h P, known economic pest species.

the authors. Species listed as common can be easilyfound at a high proportion of collection sites withintheir geographic range, often in large numbers,and appear to be regularly associated with P. hys-terophorus. Species listed as occasional requiregreater searching effort or a knowledge of partic-ular sites at which they occur; they were foundonly at a limited number of sites or occasions, butstill show a consistent pattern of association withthe plant. Species listed as rare were collected onlyon one or a few occasions, usually in small num-bers or as single individuals, and may not be con-sistently associated with the plant.

We collected 262 phytophagous arthropod spe-cies, representing 108 genera and 58 families in 8orders. The orders represented were Acari (2 spe-cies, 0.&%of species), Orthoptera (2, 0.8%), Thysa-noptera (2, 0.8%), Hemiptera (48, 18.3%), Ho-moptera (60, 22.9%), Coleoptera (87,33.2%), Lep-idoptera (53, 20.2%), and Diptera (8, 3.1%). Thepercentage representation of the main insect or-ders in the fauna of P. hysterophorus is broadly

similar to those reported for other herbaceouscomposites in their native ranges, such as Ambro-sia psilostachya DC (Goeden and Ricker 1976c),Cirsium califomicum Gray (Goeden and Ricker1986), and for the woody perennial composite Bac-charis halimifolia L. (Palmer 1987), although thepercentage of Diptera is lower than reported forthese other composites. Fifty-three species wereknown agricultural pests.

Of the species collected, 144 were confirmed tofeed on P. hysterophorus at some stage of their lifecycle. Because little has been published on the in-sect fauna of P. hysterophorus, many of these arenew host records. In a comparable series of faunalsurveys on ragweeds (Ambrosia spp.), none of 9ragweed species had an arthropod fauna of morethan 130 species (Goeden and Ricker 1974a, b;1975; 1976 a-c). Of the 144 species known to feedon P. hysterophorus, 29 (20.1%) were stenopha-gous as defined by Goeden and Teerink (1993),feeding on species within the tribe Heliantheae; ofthese 6 (4.2%) were considered to be monopha-

August 1995 McCLAY ET AL.: ARTHROPODS OF P. hysterophorus 805

I!;0us,feeding only on species within the genus Par-thl'nillnl. In comparison, the insect faunas of Am-brosiinae in California reviewed by Goeden andTeerink (1993) included 27.0% stenophagous and18.2% monophagous species. The insect fauna ofP. hysterophorus thus appears to be in general lesshost specific than for other species of North Amer-ican Ambrosiinae, being particularly deficient inmonophagous species. This may be related to thestatus of P hysterophorus as an annual specieswhose abundance fluctuates strongly in responseto rainfall and habitat disturbance, thus providinga relatively unreliable food resource and discour-a!!;inl!;the evolution of specialized phytophages(Bernays and Chapman 1994).

Forty (27.8%) of the species feeding on P hys-tl'rophorus were endophagous, a similar value tothat observed by Goeden and Teerink (1993) forCalifornian Ambrosiinae. Internally feeding spe-cies tended to be more host specific; 30.0% of en-dophagous species and only 16.3% of ectophagousspecies were restricted to the tribe Heliantheae.

Thirteen species were considered to be restrict-ed to the subtribe Ambrosiinae. This level of spec-ificitywas considered sufficient to quality a speciesas a potential biological control agent for use inAustralia, because the only members of this sub-tribe occurring in Australia are introduced weedsin the genera Ambrosia, Parthenium, Xanthium,and Iva. Six species were shipped to Australia forfurther testing. These were a defoliating beetle,Zygogramma bicolorata Pallister, a stem-gallingmoth, Epiblema strenuana (Walker) (McClay1987), a leaf-mining moth, Bucculatrix parthenicaBradley (McClay et aI. 1990), a sap-sucking plant-hopper, Stobaera concinna (Stal) (McClay 1983), aseed-feeding weevil, Smicronyx lutulentus Dietz,and a root-galling weevil, Thecestemus hirsutusPierce (McClay and Anderson 1985). All except T.hirslltus were subsequently released (McFadyen1992). Zygogramma bicolorata also has been re-leased and established in India (Jayanth 1987, Jay-anth and Bali 1990). Epiblema strenuana has beenintroduced to China for the biological control ofannnal ragweed, Ambrosia artemisiifolia L. (Wanand Wang 1995).

Two mst fungi were also found on P hystero-phorus. Puccinia abrupta Dietel and Holway var.partheniicola (Jackson) Parmelee was found on thestems and abaxial leaf surfaces, most commonly inCoahuila and other areas of Mexico at altitudes of1.'500 m or higher. It was subsequently screenedunder quarantine in the United Kingdom and re-leased in Queensland (Evans 1987, Parker 1990,McFadyen 1992, Parker et aI. 1994). Puccinia me-lampodii Dietel and Holway was found on the ad-axial leaf surfaces on plants in south Texas andNuevo Leon.

The 6 insect species shipped to Australia for fur-ther testing covered a range of feeding niches onP hysterophorus. Although a root-mining insectwould have been a desirable addition to this list,

no suitable candidates were identified. The 2 mostcommon root-feeding species were Baris prob.aerea Boheman, which was tested and found to beinsufficiently specific, and Carmenta ithacae (Beu-tenmiiller). Larvae of C. ithacae were common inthe roots of P hysterophorus in Mexico and werereared to the adult stage on many occasions. Car-menta ithacae is found on species of Helenium andHeliopsis (Asteraceae) in the United States (Ei-chlin and Duckworth 1988). However, because nofertile eggs were ever laid in the laboratory, wewere unable to -conduct host-specificity tests to de-termine whether C. ithacae from P hysterophoruswould also accept these host plants.

Other species that may have a narrow host rangeon P hysterophorus include the leaf-rolling mothHelcystogramma chambersella (Murtfeldt) and theaphid Uroleucon ambrosiae (Thomas). The onlyother host records for H. chambersdla are fromAmbrosia spp. (Hodges 1986). It was never ob-tained in sufficient numbers for host-specificitytests to be conducted. Uroleucon ambrosiae is re-corded from many species of Asteraceae (Robinson1985), but material collected from P hysteropho-rus in Mexico would breed only on P hysteropho-rus and P confertum Gray in cage tests, suggestingthe existence of sibling species or host races. Thisbears out the suggestion of Robinson (1985) thatthe host range of U. ambrosiae may be more lim-ited than appears from literature records.

Numerous species of Cicadellidae (Homoptera)and Miridae (Hemiptera) were collected. Fewfeeding tests were conducted witll these species,because most did not appear sufficiently damagingto be useful as biological control agents. Many ofthese species are therefore indicated in Table 1 ascollection records only. With further tests, it is like-ly that some of these could be shown to feed anddevelop on P hysterophorus.

Adults of several species of Chlamisinae andCryptocephalinae (Coleoptera: Chrysomelidae)were frequently collected. Larvae of these subfam-ilies live in a case constmcted from fecal material.Some of these species, including Cryptocephalusirroratus Suffrian, Pachybrachisimmaculatus Ja-coby, Exema conspersa Mannerheim, and Diachusauratus F., could be reared from eggs to adults onliving foliage of P hysterophorus, but larvae wererarely encountered in the field. Other members ofthese subfamilies feed as larvae primarily on deadfoliage and detritus (LeSage 1985).

The indices of percentage similarity between thefauna of P hysterophorus and the 11 other planttaxa included in our comparisons ranged from 1.1to 8.3% (Table 2). Although these are not highoverall levels of similarity, the values appear to re-flect the taxonomic relationships of the host plants,at least at the tribal level. The fauna of P hyster-ophorus is most closely related to those of othergenera within the tribe Heliantheae, such as Am-brosia, Helianthus, and Xanthium. Characteristicstenophagous insect genera shared among these

806 ENVIRONMENTAL ENTOMOLOGY Vol. 24, no. 4

Table 2. Similarities between phytophagous arthro-pod faWlas of P. hysterophorus and other plant taxa

plant taxa include Exema, Ophraella, Zygogramma,Cylindrocopturus, Smicronyx, Carolella, Helcysto-gramma, and Epiblema. The relatively high overlapwith Ambrosia occurs despite the fact that most ofthe faunal data on Ambrosia are from southernCalifornia, well outside the range of our survey andthe native range of P hysterophorus. There was alower level of similarity between the fauna of P.hysterophorus and that of the congeneric speciesParthenium argentatum Gray (guayule), with only20 insect species in common. This may be becauseof the marked differences of plant architecture,secondary chemistry, and habitat between the 2species. P argentatum is a small mbber-producingshmb of xeric habitats in northern Mexico, andrarely co-occurs with P. hysterophorus. In contrast,the genera Ambrosia, Xanthium, and Helianthusinclude numerous herbaceous annual or perennialspecies occurring in mderal habitats similar tothose occupied by P hysterophorus. Genera in thetribes Astereae (Baccharis and Gutierrezia) andCynareae (Cirsium) and the families Caesalpini-

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).iRogers (1988).j Hilgendorf and Goeden (1982).k Boldt and Robbins (1987).I Boldt and Robbins (1994).m 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).S Stone and Fries 1986.t Foster et al. (1981).II Wood (1992)."Ward et aI. (1977).W Goeden and Ricker (1986).x Goeden and Ricker (1987a).Y Goeden and Ricker (1987b).

Plant taxon (no.species included)

Ambmsia"-g (9)Helianth1/.J>,h, i (10)Xanthiumo.j (2)Baccharisk""P(ll)Lantana'" q (4)Parthenium argentatmnr, 8, b(l)Senna" (5)Gutierrezitl (1)Parkinsonia'" II (1)Prosopis" (5)CirsiumlO-<j (9)

Totalphyto-

phagousinsect

species

274224197577550194121149120433117

Phytopha-gous spe-

cies incommonwith P.hljstero-pharos

413732443820111110134

Index ofsimilarity ,% Is

8.288.247.495.534.914.592.962.752.691.911.07

aceae (Parkinsonia, Senna) and Mimosaceae (Pro-sopis) had lower similarities with the fauna of Physterophorus, reflecting their greater taxonomicdistance from Parlhenium. Lantana camara had asurprisingly high degree of similarity (4.91%) withP hysterophorus; the reasons for this are unknown.

These faunal relationships are of importance tothe practical aspects of selecting weed biologicalcontrol agents. The relatively high degree of sim-ilarity between the faunas of P. hysterophorus andHelianthus validated the selection of H. annuus asa critical test plant. Thirty-nine species (14.8%) ofthe fauna of P hysterophorus have also been re-corded from Helianthus, making them ineligiblefor further consideration as biological controlagents. This close association between the faunasof P hysterophorus and H. annuus was probablythe most important factor limiting the number ofbiological control agents obtained from this pro-ject.

The choice of Monterrey in northeastern Mexi-co as the main project base seems to have beenfully justified. All 6 of the species sent to Australiawere found in the immediate vicinity of Monterrey,whereas only S. lutulentus, E. strenuana, and S.concinna were found in proximity to the base atTemple, TX, and only E. strenuana and S. concin-na were found near Cuernavaca.

One indicator of potentially good biological con-trol agents is their effect on the host plant in itsnative range (Wapshere 1985, Crawley 1989). Se-vere insect damage on P hljsterophorus was rarelyobserved in our surveys. Zljgogramma bicolorataoccasionally became abundant enough to cause de-foliation, and on one occasion a young stand wasseverely stunted by heavy E. strenuana infestation.These 2 species have been the most effective bi-ological control agents when released in India andAustralia (McFadyen 1992, Jayanth and Bali 1990).During the cooler winter months, plants were of-ten coated with dense colonies of U. ambrosiae.Occasional defoliation by polyphagous Lepidop-tera was seen in Texas.

The absence of native or economic species ofAmbrosiinae in Australia facilitated the introduc-tion of biological control agents into Queenslandagainst P. hysterophorus, as it was possible to re-lease several species which were oligophagous onParlhenium, Ambrosia, or Xanthium, rather thanstrictly monophagous. This implies, however, thatcaution should be exercised in releasing any ofthese agents in other countries where P hystero-phorus is a problem. Any such releases should bepreceded by a careful review of the status of po-tential host plants present in the proposed area ofintroduction, and, if necessary, additional hostspecificity tests should be performed.

Acknowledgments

We thank Hugo Miranda Sanchez, Hugo Perales Ri-vera, and Manuel Valencia for technical assistance, Dieter

August 1995 McCLAY ET AL.: ARTHROPODS OF P. hysterophorus 807

Enkerlin ,md Juan D. Vega for providing facilities atITESM, Marion Herbut for assistance with graphics, andL. Dosdall, R. Casey, R. E. McFadyen, and F. D. Panettafor their review of the manuscript. We also thank thelJ niversidad Aut6noma del Estado de Morelos for pro-viding facilities in Cuernavaca. We particularly thank theexpert ta'(onomists who identified the arthropods, and of-ten also offered valuable advice on their biology and hab-its: D. M. Anderson (USDA Systematic Entomology Lab-oratory, Washington DC [SEL], Coleoptera; Curculioni-<lae); J. D. Bradley (International Institute of Entomol-ogy, London [IlE], Lepidoptera); R. L. Brown(Mississippi Entomological Museum, Lepidoptera; Tor-tricidae); J. A. Chemsak (University of California, Berke-ley, Coleoptera; Cerambycidae); M. L. Cox (IlE, Cole-optera: Curculionidae, Scolytidae); D. R. Davis (NationalMuseum of Natural History, Washington, DC [NMNH],Lepidoptera; Lyonetiidae); C. H. Dietrich (SEL, Ho-moptera: Auchenorrhyncha); E.A.J. DuffY (IIE, Coleop-tera: Chrysomelidae); T. D. Eichlin (California Depart-ment of Food and Agriculture, Lepidoptera: Sesiidae); D.C. Ferguson (SEL, Lepidoptera; Geometridae, Ptero-phoridae, Pyralidae); R. H. Foote (SEL, Diptera: Te-phritidae); R. Gagne (SEL, Diptera: Cecidomyiidae);M.S.K. Ghauri (IIE, Hemiptera, Homoptera); R. D. Gor-don (SEL, Coleoptera; Scarabaeidae, Coccinellidae); K.M. Harris (IIE, Diptera: Agromyzidae); T. J. Henry(SEL, Hemiptera); H. A. Hespenheide (University ofCalifornia, Los Angeles, Cylindrocopturus); R. W. Hodg-es (SEL, Lepidoptera: Gelechiidae); J. D. Holloway (IIE,Lepidoptera); F. Hovore (Placerita Canyon Nature Cen-ter, [Coleoptera: Cerambycidae); J. M. Kingsolver (SEL,Coleoptera: Bruchidae); J. P. Kramer (SEL, Homoptera;Auchcnorrhyncha); D. Macfarlane (lIE, Acari; Tetra-nychidae); 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, Or-thoptera: Acrididae); A. Norrhom (SEL, Diptera; Te-phritidae); C. W. O'Brien (Florida A&M University, Co-leoptera; Curculionidae); J. M. Palmer (lIE, Thysanop-tera); R. W. Poole (SEL, Lepidoptera: Noctuidae); J. Ra-zowski (Polish Academy of Sciences, Carolella); F. H.Rindge (American Museum of Natural History, Lepidop-tera; Geometridae); J. Schaffner (Texas A&M University,Hemiptera: Miridae); T. J. Spilman (SEL, Coleoptera;Cerambycidae, Elateridae); M. B. Stoetze1 (SEL, Ho-moptera: Aphididae); M. R. Wilson (lIE, Hemiptera:Miridae); R. E. White (SEL, Coleoptera: Chrysomelidae,Cerambycidae); the late D. R. Whitehead (SEL, Cole-optera; Curculionidae); and D. J. Williams (IlE, Homop-tera: Pseudococcidae). Funding for the study was provid-ed by the Rural Lands Protection Board, Queensland,Australia.

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Received for publication 17 May 1994; accepted 6March 1995.