BIOLOGICAL AND MICROBIAL CONTROL Host Specificity of Argopistes tsekooni (Coleoptera: Chrysomelidae), a Potential Biological Control Agent of Chinese Privet

YAN-ZHUO ZHANG,] JAMES L. HANULA,2 AND JIANG-EUA SUN 1,;3

J. Econ. Entoll1ol. 101(4} : 1146-1151 (2008) ABSTRACT Chinese privet, LigustrmH sinellse Lour., is a perennial semi-evergreen shrub that is it serious invasive weed in the United States. Classical biological control offers the best hope for controlling it in an economic, effective, and persistent way. Host specificity of one of the most promising biological control agents of Chinese plivet, a flea , Argopistes tsekooni Chen (Co­ leoptera: Chrysomelidae), was evaluated in China by using laboratory no-choice and choice tests on 13 species of Oleaceae and eight species in other families that have important economic v.llue. In adult no-choice survival and oviposition tests, the fed and survived for 30 d on Syringa ohlata LincH., jasminwn Hudiflom.m Lindl., and three species in the genus Ligustru.m. Females also oviposited on these species, but only larvae from eggs laid on S. oblata and Ligu.stru.m, spp. developecl successfully. In addition, the did not feed or oviposit on the species of economic importance. In choice tests, adults preferred L. siner/,se for feeding and oviposition. These results show that A. tsekooni is relatively host specific and warrants further testing as a biocontrol agent of Chinese privet in the United States.

KEY V\fORDS Argopistes tsekooni, flea beetle, weed biological control, host specificity, Ligustrum si.nense

Chinese privet, Ligustrtf.1H sinense Lour., is a perennial many nue species. For example, at least one popula­ semideciduous shrub or small tree indigenous to tion of Schweinitz's sunflower (Helianthus schweinil:­ China, Vietnam, and Laos (Wu and Raven 2003, The zii Torrey and Gray), a federally endangered endemic Nature Conservancy 2004). In its native habitat, it is plant to the piedmont of the Carolinas, has been used as an ornamental, for medicines, teas, and other pushed closer to extinction because of privet's ability uses, and it is not considered a pest (OuYang 2003, to shade out competing plant species (U.S. Fish and Ou Yang ancl Zhou 2003). Chinese privet was first Wildlife Service 2008) . L. sinense also has successfully introduced into the United States in 1852 as an orna­ invaded the limestone cedar (Jullipe,;/.tS virginianus mental shrub (Coates 196.5, Dirr 1990), but it was L.) glade-woodland complex (Quarterman 1950) of recorded as escaping from cultivation in southern the central basin of Tennessee (Morris 2001). This Louisiana by the 19:30s (Small 1933 ). During the 1950s, ecosystem is rich in plant endemism in the southeast­ 1960s, and 1970s, Chinese privet became Widespread ern United States (Estill and Cruzan 2001), but it is in natural habitats ('Wilcox and Beck 2007). Chinese globally imperiled, in part because of exotic species privet has become one of the worst invasive plants in invasions, including Chinese privet (Noss et al. 1995, the southeastern United States (Faulkner et a1. 1989, Morris et a1. 2002). L. sinense also can be directly Stone HJ97) where it is naturalized and considered a harmful to humans. Respiratory in-itation caused by severe threat to ecosystems from Texas to Florida, and floral odors is common where Chinese privet is abun­ north as far as the New England states (The Nature dant (Westbrooks and Preacher 1986). Conservancy 2004, University of Connecticut 2004). It Mowing and cutting are appropriate for small pop­ is also a pest in Australia and New Zealand (SwClrbrick ulations eh'vironmentally sensitive areas where her­ et a1. 1999), Argentina (Montaldo 199.3), and on sev­ o~' eral Pacific Islands. bicides cannot be used. Repeated mowing and cutting L. sinense is particularly damaging along sensitive controls the spread of Chinese privet, but it may not riparian areas where it forms a single-species midstory eradicate it (Tennessee Exotic Pest Plants Council that shades out native understory vegetation (Ten­ 1996). Although modern herbicides including glypho­ nessee Exotic Pest Plant Council 1996), including sate effectively kill privet (Madden and Swarbrick 1990, Batcher et al. 2000, Harrington and Miller 200.5),

I State Key Laboratory of Integrated Management of Pest environmental concerns limit the use of herbicides on & Rodents. Institute of Zoology, Chinese Academy of Science, Beijing public land or in sensitive areas. Environmental con­ 100101, P. H. China. cerns over widespread use of herbicides, combined Z USDA-Forest Service, Southern Research Station, 320 Green St.. Athens, GA ;30602-204·1. \-\lith the vast area infested in the United States, make .; Corresponding author. e-mail: [email protected]. biological control an attractive option .

0022-04!:l:3I08/1I46-1l51$04.00/0 @ 2008 Entomological Society of America August 2008 ZHANG ET AL.: HOST SPECIFICrry of A. tsekooni 1147

Surveys for potential biological control agents were TalJle 1. Plullt spedes sdef. ~leti ftH')lOslspedfidly eXpel'illumtK conducted in China during 2005 and 2006. More than wilh A. Isekuuni 100 phytophagous species were found feeding Family Species on Chinese pIivet (Zhang et a1. 2008). The most prom­ ising insect for biological control was the leaf-mining Cucurbitaceae CII(;I/mi nos 8l1t1:1)1I8 L. flea beetle Argopistes t<;e/woni Chen (Coleoptera: Eric(leeat' Rhododendron simsii Plnneh. Fabaceae Glycine max (L.) MelT. Chrysomelidae). This flea beetle caused serious dam­ Malva<:eae GO.I·SIn·liI/TII hil-WtllrJI L. age to Chinese privet when its population was high, Oleaceae Chionantl'llls mtl1s118 Lindl. & Paxt. and it was the dominant species in the insect CO~l­ FOllilll1l1sia jlwhmei Carr. munity of naturally occurring Chinese privet. Females Forsuthia oiridissi/ll(/ LincH. TaslIlinlllll lallceoiar;,ul/I Hoxb. oviposit into leaves, with only a small part visible at the ]aslIlinl/lII nlldiflonm! Lindt. surface. Newly hatched larvae begin mining between .Ta~1I1inl/lIl slllllliac (L.) AitOTl the upper and the lower surhlce of the leaves, and they Ligllstrl/m.iaponicl/TII Thunb. continue to eat the tissues as they mature. Adults feed Ligllstrltm lucie/11m Ail. Ligl/slrltTII sinen~e Lour. externally by scraping the epidermal layer and the OSTIIllnthu8 c:ooperi Hemsl. underlying cells, usually penetrating through the leaf OSTIIllllthus f/'(/gl'Ctn~ (Thunb.) Lour. and causing a small feeding hole. Holes made by adult OSlIlallthlls marginatus (Champ. ex Benth.) Hems!. beetles and numerous long mines created by hU'vae SUrillga ob/ata Li ndl. Poaccae um mays L. often result in leaf abscission (Y.-Z.Z. et aI., unpub­ Rubiaeeae Con/en';o jllsmillOid(~\' Ellis lished data). Solanaceae L!lcop(~rsicon escllientulII L. Flea beetles are a large, primarily oligophagous Theaceae Camellia ()le~rera Abel group, with several species currently beir~g use~l as biological control agents of weeds throughout the world. For example, Aphtlwna spp. were released as important agricultural or ornamental plant species biocontrol agents for leafy spurge, Euphorbia esuLa L. also were selected for testing (Table 1). (Euphorbiaceae), in North America (Lym and Nelson Test plants were obtained either from seed, cut­ 2000); Altica cardllonnn Guer. is a successful biological tings, transplants of plants growing naturally in the control agent of Cirsium arvense (L.) Scop. (Aster­ field, nursery stock, or from commercial markets. Sy­ aceae) (Wan et al. 1996); and Agm)cicles hygrophila ringa oblata Lindl. and Osnwnthus fragrans (Thunb.) Selman et Vogt was used in China for suppression of Lour. were transplanted into a nearby test field be­ Altemanthera philoxeroi.des (Mart.) Griseb., a global cause of their large size. Corn (Zea mays L.), soybean virulent weed from South AmeIica (Julien et a1.1995). [Glycine max (L.) Men. J and cotton (Gossypium hir­ At'ielwoni. is repOlted to be strictly associated with sutum. L.) were grown from seeds. Cucumber (Cu­ glossy privet, L. lucidum Ait. and Syringa oblata Lindl. cumins sativtls L.) and tomato (Lycopersicon escu len­ var. girlliclii. (Lemoine) Rehd., but more information tum L.) were obtained as potted plants from a nursery, is not available (Yu et a1. 199G). According to the as were most of the ornamental plants not within literature, the genus Argopistes is exclusively associ­ Oleaceae (Rhododendron simsii Planch., Gardenia ated with the Oleaceae (Yu et a1. 1996). jasminoi.des Ellisas, and Cam.ellia oIeifem Abel). Jas­ The purpose of the study is to determine the host minum sambllc (L.) Aiton (Oleaceae) also were pur­ specificity ofA. t~'elwoni through choice and no-choice chased as potted plants. Several other plants (Fon­ feeding and oviposition trials. Establishing host spec­ tallesia fortunei CmT., Osmllnthus margi-natus (Champ. ificity is an important step in evaluating potential bi­ ex Benth.) Hemsl.; Osmanthus cooperi Hemsl.; For­ ological control agents. sUthill viridissima Lindl.; Chionanthtls retustls LineH. & Paxt.; .IasrniHu'l1l rltul(fionl'lJ'/. Lind!.; .fasmiHwn lanceo­ lariuIT! Roxb.; Ligl.lstrumsinense Lour.; Japanese privet, LigustrumjapOI;icum Thunb.; and Ugustrwn LucidwH Materials and Methods Ait.), were rooted from cuttings in sand under an Test Plants and Insects. The insects were tested on automatic misting machine in a greenhouse. Once species closely related to Chinese plivet selected ac­ successfully rooted they were transplanted to plastic cording to the now generally accepted centrifugal pots (20 cm in diameter, 15 cm in height). All plants phylogenetic method proposed by H,uris and Zw()lfer were held under natural day lengths and temperatures (1968) and 'Wapshere (1974) for host specificity test­ in an outdoor site with a shade cloth. Plants were ing of biological control agents of invasive plants inigated as needed but no pesticides or fertilizers worldwide (DeLoach et 011. 2003). According to the wel~e used to avoid any effect they might have on A literature on Oleaceae, consensus trees from sep,u-ate tselwoni biology or behavior. and combined molecular analyses were congruent and Experiments were conducted in the greenhouse of agreed well with nonmolecular data (Wallander and the Forestry Institute of Huangshan city. The green­ Albert 2000), suggesting that the plant species se­ house was maintained at 25-30°C and 40 - 60% RH, lected for host specificity testing through classical with a photoperiod of 16:8 (L:D) h. Newly emerged, were appropriate. Thirteen representative unfed adults were used to test A tselwoni survival on species from seven genera of Oleaceae were selected. different plant species. Adults were obtained by dig­ Besides the closely related species to L sinense, eight ging pupae or newly emergecl adults hom the soil 1148 JOURNAL OF ECON01... UC ENTOMOLOGY Vol. 101 , no. 4 under Chinese privet shrubs or trees in a natural area. Zero data and data produced by beetle testing or Pupae or adults were placed individually into clem' exploratory feedings, which were near zero, were ex­ beakers (12 cm in diameter, 15 cm in height) con­ cluded (SPSS Inc. 2001). In choice tests, ANOVA and taining moist fine sand and covered with a screen until independent-samples t-test were used to analyze the the n{ajOlity of A. tselwoni, had reached maturity. amount of foliage consumption and number of eggs Adults were sexed by assessing the shape of the last deposited after excluding zero data (SPSS Inc. 2001). abdominal segment (S. Y. Wang, personal communi­ cation). Adult No-Choice Survival Test. Two mating pairs of Result<; adults were selected randomly and placed in 10- by 20-cm polyester organza sleeve bags, which were then Adult No-Choice Survival and Oviposition Tests. In securely tied to branches or petioles oneaf clusters of adult no-choice tests, survival of A. tsekooni differed test plants. Two or three sleeve bags were used per significantly among host plants (F = 185.082; elf = 21, plant. The experiment was replicated 10 times for each IH8; P < 0.0001) . Some adults survived 30 d on S. plant species. Bags with adults were moved to new oblata, .J. Ht.l(l-ijforll1n, and alI three Ligustrum spp., branches on potted plants every 2-:3 d as needed for whereas no adult survived that long on the other 30 d, and the area of foliage consumed was measured species tested (Table 2). Percentage of survival on L. for each branch. Foliage ~onsumption was quantified japonicllm, L. [ueir/urn, and S, oblata did not differ by placing transparent graph paper with a I-mm2 grid significantly from the native host plant L. sinense, but over each leaf to measure the surface area damaged. survival on .J. nudifiorum was significantly lower (F = Adult survival was recorded after 30 d. 20.640; df = 4, 45; P < 0.0001). Adult No-Choice Oviposition Test. No-choice ovi­ Foliage consumption by adults also differed signif­ position tests were conducted separately from the icantly in the no-choice test (F = 66.948; df = 21, IH8; survival test to eliminate the influence of starvation on P < 0.0001) . A. tsekooni fed normally on ]. nudijlorum" adult fecundity. Mature adults, Le., adults at least 5 d S. oblate, and Ligusf-rum, spp., whereac; it did not feed old and feeding normally on Chinese privet, were on other plants or only fed a small amount to test host collected from a natural m'ea and put in a gauze cage suitability. The amount of foliage consumed differed filled with fresh branches of Chinese privet for 48 h to significantly (F = 889.110; df = 4, 45; P < 0.0001) ensure they had fed enough for oviposition. Otherwise among plants fed upon normally. The ranking of leaf the methods were the same as the adult no-choice consumption by A. tsekooni on the vmious plants survival test and the flea beetles were allowed to tested was L. sinense > L. japonicum > L, lucidurfl > oviposit for 30 d. Eggs deposited in each plant leaf S. oblata > J nud~fiorum (Table 2). were counted using a dissecting microscope and ob­ Oviposition also differed significantly in the no­ served to determine whether eggs hatched success­ choice test (F = 204.7; df = 21, 198; P < 0.0001) . A. fully. tsekooni oviposited only on plants within the genus Adult Choice Feeding and Oviposition Tests. Plant Ligtlstrum and on S, ohlata and]. nudi;!lo'l'wn. Among \vithin the Oleaceae family \-vere used for adult choice these plant species, females deposited significantly tests. Fresh branches were inserted through holes in more eggs on L. sinense and L. lucidum than any others the rubber lid of water filled, 10-ml medical glass vials. (F = 184.716; df = 4,45; P < 0.0001), including L. These were then placed in 800-ml beakers containing japonicum. Significantly fewer eggs were laid on S. moist cotton or vermiculite and covered with a fine­ oblata compared with the Ugustrwn spp. Females mesh cloth lid. Each beaker received one L. sinense deposited an average of 0.4 2: 0.69 eggs per female on plant branch plus one or two other test plant branches. .J. nudijlortnn, but larval development was unsuccess­ Two mating pairs of A. i'iekooni adults were added to ful. In compmison, all eggs laid on Ligustrum spp. and each beaker and allowed to feed and oviposit for 1 wk. S, ohlata hatched ancllarval development was possible, Each test was replicated 10 times. Foliage consump­ at least to the point where the larvae left the mines and tion and oviposition were qualified as mentioned began looking for pupation sites. above. Adult Choice Feeding and Oviposition Tests. In Data Analysis. Data from adult no-choice and mul­ choice tests, adult A. i'iekooni consumed significantly tiple choice feeding and oviposition experiments do more L. sinense than any alternative host except for L. not generally confo~'m to the assumptions of paramet­ ja:poni.cum (Table :3). In oviposition tests where L. ric tests; thus, we used the nonparametric K-S method sinense was compm'ed with other Ligust'rum spp. the of SPSS (SPSS Inc. 2001). Survival, total amount of beetle demonstrated no preference between Ligtls­ foliage consumed in 30 d, and number of eggs depos­ trum spp. Females preferred plants within the genus ited by each female in adult no-choice tests was an­ Liguslrum for oviposition and deposited no eggs on alyzed llsing rank cases and analysis of variance oti1er test plant species (Table :3; Josepb et aI. , 1994) . (ANOVA) (SPSS Inc. 2001). Combined, these proce­ dun~s produced a test that approximated the nonpara­ metric Kruskal-Wallis test (Colpetzer et al. 2004). Discussion ANOVA and the least significant difference (LSD) multiple comparison test was used for mean sepm'a­ Demonstrating host specificity is critical before the tion in no-choice feeding and oviposition expeliments. introduction of any proposed biological control agents August 2008 ZHANG ET AL.: HOST SPECIFICfrY of A. tsekooni 1149

Tahle 2. MCllu ± SD P{~I·'. ~elltuge of sllI'vivl\l, alllOllll1 of foliage eOrJsulllcd, lIud J1\unh{~1' of t~ggs lui, I hy A. /st'lwolli on vl\rious plulIl ISpl'cies ill atiult lIu-ehukc !HII'vivulaml ovipusition lC8l!S (II = 10)

% surviving Foliage consumed Oviposition Plant species for :30 eI (ern2 tbeetlet:30 e1) (eggs/female/30 e1) Camel/;a ole!fera Abel o o (no feeding) o Ch;onantlws relllSlls Lindl. & Paxl. o 0.092 :!: 0.077 (test ft'ecling) o CIIClIlIlillS salit)llS L. o o (no feeding) o FOlltarw.~ia liil'llITw; Carl'. o 0.005 :!: 0.007 (test feedin~) o Forsythia ,;iridissilll() Lindl. o 0.002 :!: 0.004 (test feedin~) o Gardenia jllslIIinoides Ellis o o (no feeding) o Glycine T1/m' (L.) MelT. o 0.006 :!: O.OOH (test feeding) o COSSYP;IITI/ /'irslI/1I11I L. o 0.009 ± 0.015 (test feeding) o jasTl/illullI lanceola,-jlllll Roxb. o 0.083 :!: 0.024 (test feeding) o jaslIlillllTl/ Illldiflorlll/l Lincll. ·22.50:!: 18.45b 0.320 :!: 0.267e 0.4:!: 0.699e1 jllsm;lIuTII sambllc (L.) Aiton o 0.010 :!: 0.0]:3 (test feeding) o Ligustl'llln jll1JOllicll1l1 Thllnb. 80.00 :!: 15.81a 6.205 :!: 0.339b 24.20 :!: 4.2:nb Ligustnnn luddlnn Ait. 87.50:!: 13.17a 4.711 :!: 0.702e 28.:30 :!: :3.498a Ligustnn/l sillenst: Lour. 85.00 :!: 17.48a 9.863 :!: OA09a 30.20 :!: 2.044a Lycopersicon eSCl/lenlLlII1 L. o o (no feeding) o OSrlllllltllllS cooped Hems!. o o (no feeding) o OS/lumtlllls fmgraTis (Th Ilnb.) Lour. o 0.027 :!: 0.029 (test feeding) o OST/llIIltitUS I/lllrgirwtus (Champ. ex Benth.) Hernsl. o o (no feeding) o Rhododendmn silnsii Planch. o o (no feeding) o Syl'illgll oblata Lindl. 75.00 :!: 26.:3.5a 0.818 :!: 0.189d 17.9 :!: 2.07!:Jc Zea !lllIys L. o o (no feeding) o

" Survival and feeding tests were conducted with two newly emerged male and two newly emerged female adults enclosed in sleeve bags with a plant branch. l\'leans within eoillmns sharing the same letters are not significantly different (l' < 0.05; rank cases and ANOYA, LSD; SPSS Inc. 2001).

regardless of their control potential (Schroeder 1983). Possible harm to economic and other nontarget plants must be carefully assessed and only insects proven to Tahlc 3. I\1CIIII ± SD lent' consullIpLion (til(l number of eggs be host specific are acceptable for release (Balciunas laid 011 variolls plullt specieli ill adult A. ' ..HlkoOllj choice lests (/I = 10) et al. 1994). Larvae of A. tsekooni are leafminers that cannot Consumption move to other host plants; thus, it is important to 2 Plant species area (1010 /2 Egg no. determine the host range and oviposition behavior of pairs adults/wk) the adults, which are also herbivorous. Our tests COTl­ Test I firmed that A. tselwoni, has a narrow diet and host range UgIIstrlllll sinellse LOllI'. 176.4 :!: 44.72a 18.5 :!: 5.66 restricted to the genus Ligustrwn and S. oblata, and jaslllillulII lI11dijlorllm Lind!. 1.6 :!: 2.12b o Forsythia l)i.1iciissimll Lind!. 0.1 :!: 0.:32b o possibly other Syringa species. Of those plants tested Test 2 the genus Syringa is phylogenetically closest to Ligu­ Ligmt1'll1n silleme Lour. 134.4:3 :!: 74.82a 22.0:!: \.'3.30a stru.m (Wallander and AlbeIt 2000). The preference of Ligllstnn/l }apon;cllIIl Thunb. fiG.71 :!: 20.29a 16.0:!: 11.40a Osnullltlrllii fragrans (Thllnb.) o o A t.sekvoni supports this and suggests that more distant Lour. genera are unlikely to be acceptable as hosts for this Test 3 insect. A. L.<;eko()ni did successfully feed and oviposit on Ligllslnllll sint:lISe Loul'. 122.22 :!: 5~) . 00a 28.0:!: 18.64a S. vblata in adult no-choice survival and oviposition Ligllstrlllll {lleidlllll Ail. 44.1 J :!: 2~).74b 20.17:!: 1O.11a FonillTlCS;a for/llnei Carr. o 0 tests, but when L. sinense was present they fed very Test 4 little and did not oviposit on S. olJlata. Lilacs (S~r"inga Ligll,\tnlln silumse Lour. 10:3.5 :c: 21.40 16.0:!: ,5.62 spp.) are widely planted non-native ornamental spe­ .Iasmilllllll Imlceolar;1I111 Hoxb. o o cies in the United States, so further testing will be .Il1STlli1ll11ll slIml}{/c (L.) Aiton o o Test 5 necessary to ensure A. tse/woni, does not cause serious UglIslnml sirwnse Lour. 134:!: 70.25 15.17:!: 3.92 damage to these plants. Like Chinese privet, Japanese OSlI!mllhllS cooperi Hemsl. o o privet and glossy privet m'e considered exotic invasive Chiollllntlws "eIIlSIIS Lindl. & o o Pax I. species in the United States (Miller 2003, Munger Test G 2003); thus, the ability of A. tsekooni to feed and ovi­ Lif.!,lIstnlm sillellSI' Lour. 144.()O :!: :35.48a 14.00:!: 2.83 posit on them should be of less concern. The intro­ OSlIIlIllthllS mll1"f~;I1(/tlls o o duction of A. t~'ekooni also may provide some control (Champ. ex Benth.) Hemsl. S!lringa oblata Lindl. 2.90:!: 1.7:3b o of these other invasive exotic weeds. Recently, ecologists have expressed concerns about Means within a test group and column sharing the same leiters are the ecological risk of classical biological control (Mc­ not significantly different (1' < 0.05; ANOVA, LSD). Evoy 1996; Simberloff and Stiling 1996; Sh'ong 1997; Eadl container had two newly emerged male and female A. tsek­ (lOlli, one L sinellse plant branch, and branches of two other plant Pembelton 2000; Londa et al. 2003a,b). Some unpre­ species. dieted side effects have been documented as an un- 1150 JOUHNAL OF ECONmvtlC ENTm,,10LOGY Vol. 101 , no. 4

desired consequence of a highly limited number of tn.tlS lalipus Korotyaev (Coleoptera: Curculionidae), a weed biological control programs (Carruthers and potential biological control agent of mile-a-minute weed, D'Antonio 2005). In some eases, biological control Pof"gollum peljoliatllm L. (Polygonales: Polygonaceae). agents have potential to reduce biodiversitv of native BioI. Control :30: .511-522. species (Louda et al. 1997). HO\vever, whe;'e damage Cullen, J. M. H)90. Current problems in host-specificity to nontarget plant species has occurred, it has resulted screening, pp. 27-36. In E. S. Delfosse [ed.], Proceedings of the VII International Symposium on Biological Control from imported insects that adapted to eat physiolog­ of Weeds. G-ll March 1988. Home, Italy. CSIHO Publi- ically acceptable but less preferred and less suitable cations, Melbourne, A lIstralia. . host, in situations where the "preferred" host was not DeLoueh, C. J., P. A. Lewis, J. C. Herr, R. I. Carn! thers, J. 1.. present (Louda et al. 200:3a, Colpetzer et al. 2004), In Tl"Hcy, and J. Johnson. 200:3. Host specificity of the our tests, the physiological hosts of A. tsekooni were , Diorhabda elollgata deserlkola (Coleoptera: Li.gllstnlm spp. and S. oblata, which are non-native Chrysomelidae) from Asian, a biological control agent for species in the United States. Generally, most practi­ saltcedars (Tamarix: Tamaricaceae) in the western tioners of biological control believe the ecological host United States. BioI. Control 27: 117-147. range in the field is nan-ower than physiological host Din', M. 1990. Manual of woody landscape plants: their range (Wapshere 1989, Cullen 1990, Hm'ley and Forno identification, ornamental characteristics, culture, prop­ 1992). During observations of A. t<;ekooni 'in the field, agation, and uses, 4th ed. Stipes Publishing Co., Cham­ the flea beetles rarely fed on L. japonicml1 adjacent to paign, IL. Estill, J. C., and M. n. Cruzan. 2()01. Phytogeography of rare heavily infested and damaged L sinense (Y,-Z.Z., un­ plant species endemic to the southeastern United States. published data). Thus, A tsekooni seems to be a good Castanea 66: 3-23. candidate for biological control of L. si.llense in North Faulkner, J. L., E.E.C. Clcbsch, and W. L. Sandel'S. 198H. America and shoul~l be evaluated further. Use of prescribed burning for managing natural and his­ Our results demonstrate the narrow host range of A. toric resources in Chickamauga and Chattanooga Na­ tsekooni, but we were unable to test North An; eric an tional Military Park, USA. Environ. Manage. 13: G03-612. species; thus, further testing will be necessary. How­ Harley, K.L.S., and I. W. Forno. 19H2. Biological control of ever, this and other studies (Y.-Z.Z. et aI., unp{lblisbed weeds: a handbook for practitioners and students. In kata data) demonstrate the potential of this beetle as a Press, Melbourne, Australia. biological control agent for L. si.nense. Harrington, T., and J. H. Miller. 2005. Effects of application rate, timing and formulation of glyphosate and tric10pyr for control of Chinese privet (Ligustrurn sine/l.se). Weed Aclmowledgments Technol. 19: 47-54. Hanis, P., and H. Zwtilfcl'. 1968. Screening of phytopha­ vVe are grateful to Prof. Shuyong Wang for assistance with gous insects for biological control of weeds. Can. Ento­ identification of A. t,'>elwolli; Prof. Xinjian Pan (Forestry In­ mol. 100: 295-303. stitute of Huangshan city) for identification of test plants; Julien, M. H., B. Skarratt, and G. F. MaywaJd. 1995. Poten­ Prof. Jianqing Ding for the suggestion of the test design; and tial geographical distIibution of alligator weed and its Huaijun Xue for invaluable advice. We also thank Fang Fang biological control by Agasicles hygrophila. J. Aquat. Plant and Shengli Xiang for field assistance. We appreciate the Manag. 33: 55-60. advice of Shuguang Hao and Jianxtl Zhang on data analysis. Louda, S. M., A. E. Arnett, T. A. Rand, and F. L. Russell. We are especially grateful to Judith Hough-Goldstein (Uni­ 200:3a. Invasiveness of some biological control insects versity of Delaware) and two anonymous reviewers for re­ viewing the manuscript. This research is palt of an ongoing and adequacy of their ecological risk assessment and Sino-U.S. Chinese privet biological control cooperative pro­ regulation. Conserv. Bio1. 17: 73-82. gram funded by the USDA-Forest Service Southern Re­ LomIa, S. M., D. Kendall, J. Connor, and D. Sim berl 0 fl. IHH7. search Station and partially supported by the National Nat­ Ecological effects of an insect introduced for the biolog­ ural Science Foundation of China (:30.52.5009) and the ical control of weeds. Science ('Vash., D.e.) 277: J088- National Key R&D Program (200(JBAD08A17). 1090. Luuda, S. M., R. W. IJembcl'ton, M. T. Johnson, and P. A. Follett. 200:Jb. Nontarget efTects-the Achilles' heel of References Cited biological control? Retrospective analyses to reduce risk associated with biocontrol introduction. Annu. Rcv. En­ Balciunas, J. K., W. B. Damien, and F. P. Matthew. ]994. tOIllOI. 48: :36.5-396. Field and laboratory host ranges of the Australian weevil, Lym, H. G., andJ. A. Nelson. 2000. Biological control ofleafy Oxyops vitiasa (Coleoptera: Curculionidae), a potential spurge (ElIphorbia esufa) with Aphtlwna spp. along rail­ biological control agent for the paperbark tree, Melafeuca road right-of-ways. Weed Techno\. 14: 642-646. qllinqLlcnervia .. BioI. Control 4: :3.51-360. Hatchcr, M. S.,M. Tu, B. l'vlcyel's-Hicc, and J. M. Randall. Madden, J. E., and.J. T. Swm'brick. Hmo. Chemical control 2000. Element stewardship (abstr.) for Ligtlstnun spp. of Ugustrtl1n luciclwn. Plant Prot. Q. 5: 145-147. plivet. (http:// tncweed~.ucdavis.edul esadocsl documntsl McEvoy, P. n. 19H6. Host specificity in biological contro\. ligu~sp.pdf) . Bioscience 46: 401-405. Carl'Uthcrs, R. I., and C. 1\-1. D'Antonio. 2005. Science and MilleI', J. H. 2003. Nonnative invasive plants of southern decision making in biological control of weeds: benefits forests: a field guide for identjfication and control. Gen­ and risks of biological control. BioI. Control .3.5: 181-182. eral Technology Report. SRS-62. U.S. Department of Ag­ Coates, A. M. 1965. Garden shrubs and their histories. E. P. riculture. Forest Service, Southern Research Station, Dutton, New York. Asheville. NG Colpctzer, K,.l. Hough-Goldstein,J. Q. Ding, and "V. D. Fu. Montaldo, N. H. 199:3. Avian dispersal and reproductive suc­ 2004. Host specificity of the Asian weevil, Rhil}oncomi- cess of two species of Ug/lstl'llm (Oleaceae) in a sub- August 2008 ZHANG ET AL.: HOST SPEcIFICrry of A. tsekooni 1151

tropical forest relict in Argentina. Rev. Chil. Hist. Nat. 66: Tennessee Exotic Pest Plants Coundl. 199(i. Invasive exotic 75-8.5. pest plants of Tennessee. (http://www.Webdriver.com/ MOlTis, L. L. 2001. A comparison of growth and reproduc­ tn-eppcl exlist.htm) . tive characteristics of the invasive Ligtlsfrum sirumse and The Nature Conservancy. 2004. Element stewardship (ab­ the native Foresliera ligllstrina (Oleaceae). M.S. thesis, stract) for Ligl/stnnnspp. privet. (http://tncweeds.ucdavis. Middle Tennessee State University, Murfreesboro, TN. eelu/ esadocs/ documnts/ liglUp.pdf) . MOITis, L. L.,J. L. Walck, and S. N. Hidayati. 2002. Growth University of Connecticut. 2004. Invasive plant atlas of New and reproduction of the invasive J..AgIIslrllm sinellse and England. (http://www.lih.uconn.edu/webapps/ipane/ native Foresi.iera ligllslrirw (Oleaceae): implication for browsing.cfm?descriptionid=(3) . the invasion and persistence of a nonnative shrub. lnt. J. U.S. Fish and Wildlife Sendce. 2008. Schweinitz's sun­ Plant Sci . ]():3; 1001-1010. flower (Heliantlws schweinitzii) . (http://www.fws.gov/ Munger, T. 2003. Ligllstnnn spp. In Fire effects information nc-es/ plant/ schwsun.htllll) . system (online). U.S. Dep. Agric.-Forest Service, Rocky 'Wallander, E., and V. A. Albel·t. 2000. Phylogeny and clas­ Mountain Researeh Station, Fire Sciences. Laboratory sification of Oleaceae based on rpsl(j and trnLF sequence (producer). (http://www.fs.fed.us/database/feis/ ). data. Am. }. Bot. 87: 1827-]841. Noss, R. F., E. T. LaRoe III, and J. M. Scott. 1H95. Endan­ Wan, F. H., P. Han-is, 1.. M. Cai, and M. X. Zhang. H)96. gered ecosysterns of the United States: a preliminary Biology and suitability ofAllica carduon.tlrt (Chrysomeli­ assessment of loss and degradation. Biological report 28. dae: Coleoptera). a defoliator of Cirsiwn (lmense (L.) U.S. Department of the Interior, Washington, DC. Scop. Biocontrol Sci . Technol. 6: 59 -519. OuYang, M. A. 200:J. Studies on lignans and flavonoid gly­ Wapsherc, A. J. 1974. A strategy for evaluating the safety of cosides of Ligl.lslr1l11l sinense. Chin Tract Herb. Drugs 34: organisms for biological weed control. Ann. Appl. BioI. 77: 196-199. 201-211. OuYang, M. A., andJ. N. Zhou. 200:3. Studies on the phenol \Vapsherc, A. J. 198H. A testing sequence for reducing re­ glycoside constituents of Ugllsirum sinun<;e. Guihaia 23: jection of potential biological control agents for weeds. 276-278. Ann. Appl. BioI. 114: .515-.526. Pemberton, R. W. 2000. Predictable risk to native plants in Westbrooks, R G., and J. W. Preacher. U)86. Poisonous weed biological control. Oecologia (Ber!.) 12.5: 489 - 494. plants of eastern North America. University of South Quarterman, E. 1950. Major plant communities of Tennes­ Carolina Press, Columbia, Sc. see cedar glades. Ecology 31: 234-2.54. 'Wilcox, J., and C. W. Beck. 2007. Effects of Ligtlstmm si­ Schl'Oeder, D. 198:J. Biological control of weeds, pp. 41-78. nense Lour. (Chinese privet) on abundance and diversity III W. E. Fletcher led. J, Recent advances in weed re­ of songbirds and native plants in a southeastern nature search. Commonwealth Agricultlll'al Bureaux, Farnham preserve. Southeast. Nat. 6: .'535-.550. Royal. United Kingdom. Wu,Z.,andP.H. Raven. 2003. Flora of China. Science Press, Silllherl()fl~ D., and P. Stiling. 199(i. Risks of species intro­ Beijing, and lvlissouri Botanical Garden Press, St. Louis, duced for biological control. BioI. Conserv. 78: 18.5-192. MO. Multiple volumes. some in draft form. (http://£)ora. Small, J. K. 19:~3. Manual of the southeastern flora. The huh.harvard.edu/ china/). University of North Carolina Press, Chapel Hill, NC. YlI, P. Y., S. Y. Wang, and X. K. Yang. U)96. Economic insect SPSS Inc. 2001. SPSS] I for Windows. SPSS Inc., Chicago, IL. hUlIla of' China, vol. 54. Coleoptera: Chrysomelidae (II). Stone, S. L. 1997. Privet removed from Austin Nature Pre­ Science Press, Beijing, China. serves (Texas) . Restor. Manag. Notes. 1.5: 93. Zhang, Y. Z., J. I. Hanula, and J. H. Sun. 2008. Survey for Strong, D. R. 1997. Fear no weevil? Science (Wash., D.C.) potential insect biological control agents of Ligllslrum 277: 1058-10.59. sincmse (Scrophulariales: Oleaceae) in China. Fla. Ento­ Swarbdck,J. T., S. M. Timmins, and K. M. Bullen. 1999. The mol. (in press) . biology of Australian weeds. :1G. LigustnlTn lueidl/rn Aiton and Ugllslruill sinense Lour. Plant Prot. Q . 14: 122-130. Heceived 4 December 2007; accepted 1:3 April 2008.