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Home , Adenostemma, Aster amellus, Bartlettina, Cecidochares connexa, Chromolaena, Chrysanthemum indicum

Appendix 3-14

Summary of host specificity testing completed for potential biocontrol agent connexa

Eight different studies have been published detailing the results of host specificity testing of Cecidochares connexa, a biocontrol agent for Chromolaena odorata. In all, 136 taxa in 30 families have been tested, including representatives of 13 tribes of . In all cases, researchers found that C. connexa is highly specific to C. odorata. No taxa outside the tribe of Asteraceae were found to host C. connexa. Within the Eupatorieae tribe, C. connexa activity was observed on just four taxa, but could only form a sustaining population on C. odorata. In one study, oviposition was observed on Ageratum conyzoides and Austroeupatorium inulaefolium, but no larvae or gall development occurred. Cecidochares connexa was able to complete its life cycle on Praxelis clematidea, but P. clematidea could not support it for more than one generation. All three taxa are considered to be unsuitable and unsustainable hosts.

In the Hawaiian flora, there are roughly 130 taxa (depending on the latest taxonomic revisions) in 13 genera in the Asteraceae family. Hawaiian Asteraceae are diverse and include many rare and endangered . Fortunately, only one taxon sites within the Eupatorieae tribe, the indigenous Adenostemma viscosum. This is noteworthy, as all previous host specificity trials show that C. connexa can only complete its life cycle on taxa within the Eupatorieae. Thus, there is no strong biological reason to test all Hawaiian Asteraceae taxa. Adenostemma viscosum has already been tested in one host-specificity trial, and was deemed an unsuitable host.

The following table summarizes the taxa tested in all eight studies. Taxa are grouped by family. Within the Asteraceae family, subfamilies are noted by roman numeral and the suffix -oideae, and tribes are noted with numbers and the suffix -eae, following Mabberley, 2017; other taxa list only family and tribe per wikipedia. Species or genera present in Hawaii, both native and alien, are indicated with an asterisk, and were determined after review of the Smithsonian Flora of Hawaii website, online Bishop Museum herbarium records and personal communication with herbarium staff, and internet searches of local plant retailers.

Table 1: Taxa tested as potential hosts for Cecidochares connexa. Family Taxon Host? Citation Amaranthaceae Amaranthus tricolor* No Bhumannavar et. al.; 2004; McFadyen et. al., 2003 Anacardium occidentale* No Bhumannavar et. al., 2004 Araceae Colocasia esculenta* No Esguerra, 2002 Asteraceae jamesonii* No Bhumannavar et. al., 2004; III. McFadyen et. al., 2003 1. Asteraceae Carthamus tinctorius* No Bhumannavar et. al., 2004 VII. Carduoideae Lactuca sativa* No Bhumannavar et. al., 2004 4. Cardueae Sonchus arvensis No Bhumannavar et. al., 2004 Asteraceae Gynura aurantica No McFadyen et. al., 2003 XII. 1. Senecioneae Asteraceae Calendula officinalis No Bhumannavar et. al., 2004 XII. Asteroideae 2. Calenduleae Asteraceae amellus No Bhumannavar et. al., 2004 XII. Asteroideae Aster spp.* No McFadyen et. al., 2003 4. Astereae Solidago canadensis* No Bhumannavar et. al., 2004 Asteraceae Artemisia vulgaris* No Aterrado & Bachiller, 2002 Family Taxon Host? Citation XII. Asteroideae indicum No Aterrado & Bachiller, 2002; 5. Anthemideae Bhumannavar et. al., 2004 Chrysanthemum No McFadyen et. al., 2003 morifolium Asteraceae Blumea aurita No Kernasa et. al., 2010 XII. Asteroideae Blumea balsamifera No Aterrado & Bachiller, 2002 6. Inuleae Puchea indica* No McFadyen et. al., 2003 Asteraceae Bidens pilosa* No Bhumannavar et. al., 2004; XIII. alliance Muniappan & Bamba, 2002 03. * No Bhumannavar et. al., 2004 Cosmos sulfureus* No Muniappan & Bamba, 2002 Dahlia pinnata No Bhumannavar et. al., 2004; McFadyen et. al., 2003 Asteraceae Tagetes erecta* No Bhumannavar et. al., 2004; XIII. Heliantheae alliance Kernasa et. al., 2010 05. Tageteae Asteraceae, Clibadium surinamense No McFadyen et. al., 2003 Heliantheae Guizotia abyssinica No Bhumannavar et. al., 2004 Lagasca mollis No Bhumannavar et. al., 2004 acmella No Bhumannavar et. al., 2004 Wollastonia biflora No Muniappan et. al., 2007 Asteraceae Eclipta alba (synonym E. No Bhumannavar et. al., 2004 XIII. Heliantheae alliance prostrata)* 09. Heliantheae Helianthus annuus* No Aterrado & Bachiller, 2002; Bhumannavar et. al., 2004; Kernasa et. al., 2010; McFadyen et. al., 2003; Muniappan & Bamba, 2002 Tithonia divaricata No Bhumannavar et. al., 2004 Tithonia diversifolia* No McFadyen et. al., 2003 Tridax procumbense* No Bhumannavar et. al., 2004 Xanthium strumarium* No Bhumannavar et. al., 2004 Zinnia elegans* No Bhumannavar et. al., 2004; McFadyen et. al., 2003 Asteraceae, Austroeupatorium No. Oviposition but McFadyen et. al., 2003 Eupatorieae inulaefolium no larvae or gall development. Conoclinium coelestinum No Day et. al., 2016 Mikania micrantha No Bhumannavar et. al., 2004; Day et. al., 2016 Mikania scandens No Muniappan & Bamba, 2002 Praxelis clematidea Poor host. Day et al 2016; Kernasa et al Oviposition through 2010 [No] adult emergence seen, but cannot support more than one generation. Asteraceae Adenostemma lavenia No Day et. al., 2016 XIII. Heliantheae alliance (synonym for A. 13. Eupatorieae viscosum)* Adenostemma viscosum* No Day et. al., 2016 Family Taxon Host? Citation Ageratina adenophora No Bhumannavar et. al., 2004; Day (synonym et. al., 2016; Kernasa et. al., adenophorum)* 2010 Ageratina altissima No Day et. al., 2016 Ageratina riparia* No Day et. al., 2016 Ageratum conyzoides* No. Oviposition but Bhumannavar et. al., 2004; no larvae or gall Kernasa et. al., 2010; development,only in McFadyen et. al., 2003; McFadyen study. Muniappan & Bamba, 2002 Ageratum houstonianum* No Day et. al., 2016 * No Day et. al., 2016 Chromolaena odorata* Yes Aterrado & Bachiller, 2002; Bhumannavar et. al., 2004; Day et. al., 2016; Esguerra, 2002 No Day et. al., 2016 Eupatorium cannabinum No Day et. al., 2016 Eupatorium maculatum No Day et. al., 2016 Gymnocoronis No Day et. al., 2016 spilanthoides* Liatris spicata* No Day et. al., 2016 Stevia ovata No Day et. al., 2016 Stevia rebaudiana* No Day et. al., 2016 Balsaminaceae Impatiens balsamina* No Bhumannavar et. al., 2004 Combretaceae Terminalia spp.* No Muniappan et. al., 2007 Convolvulaceae Ipomoea batatas* No Bhumannavar et. al., 2004; Esguerra, 2002 Cruciferae Brassica nigra* No Bhumannavar et. al., 2004 Brassica oleracea* No Muniappan & Bamba, 2002 Raphanus sativus* No Bhumannavar et. al., 2004 Cucurbitaceae Citrullus lanatus* No McFadyen et. al., 2003; Muniappan & Bamba, 2002 Cucumis melo* No Bhumannavar et. al., 2004; McFadyen et. al., 2003 Cucumis moschata No Bhumannavar et. al., 2004 Cucumis sativus* No Bhumannavar et. al., 2004; McFadyen et. al., 2003 Curcubita moschata No McFadyen et. al., 2003 Dioscoreaceae Dioscorea spp.* No Muniappan et. al., 2007 Euphorbiaceae Hevea brasiliensis* No Bhumannavar et. al., 2004; McFadyen et. al., 2003 Jatropha curcas* No Kernasa et. al., 2010 Manihot esculenta* No Bhumannavar et. al., 2004; Esguerra, 2002; Kernasa et. al., 2010; McFadyen et. al., 2003 Phyllanthus spp.* No Esguerra, 2002 Ricinus communis* No Bhumannavar et. al., 2004; McFadyen et. al., 2003 Fabaceae Phaseolus spp.* No Muniappan & Bamba, 2002 Pterocarpus indicus* No Aterrado & Bachiller, 2002 Fabaceae, Caesalpinia pulcherrima* No Bhumannavar et. al., 2004; Caesalpinaceae McFadyen et. al., 2003 Sesbania grandiflora* No McFadyen et. al., 2003 Fabaceae, Pisum sativum* No Bhumannavar et. al., 2004 Family Taxon Host? Citation Leguminosae Vigna radiata* No Kernasa et. al., 2010 Fabaceae, Albizia falcataria* No McFadyen et. al., 2003 Mimosaceae Albizia lebbeck* No Bhumannavar et. al., 2004 Calliandra No Bhumannavar et. al., 2004; haematocephala* McFadyen et. al., 2003 Leucaena glauca* No McFadyen et. al., 2003 Leucaena leucocephala* No Aterrado & Bachiller, 2002; Bhumannavar et. al., 2004 Mimosa spp.* No Esguerra, 2002 Fabaceae, Arachis hypogea* No Bhumannavar et. al., 2004; Papilionaceae McFadyen et. al., 2003 Crotolaria juncea* No Bhumannavar et. al., 2004; McFadyen et. al., 2003 Desmodium No McFadyen et. al., 2003 heterocarpon* Dolichos lablab No Bhumannavar et. al., 2004; (synonym Lablab McFadyen et. al., 2003 purpureus)* Flemingia strobilifera* No McFadyen et. al., 2003 Gliricidia sepium* No Bhumannavar et. al., 2004; McFadyen et. al., 2003 Glycine max* No Bhumannavar et. al., 2004; Kernasa et. al., 2010; McFadyen et. al., 2003 Pachyrhizus erosus* No McFadyen et. al., 2003 Psophocarpus No McFadyen et. al., 2003 tetragonolbus* Vigna unguiculata* No Bhumannavar et. al., 2004; McFadyen et. al., 2003 Labiatae Mentha arvensis* No Bhumannavar et. al., 2004 Lamiaceae Coleus blumei (synonym No Esguerra, 2002 Plectranthus scutellarioides)* Lauraceae Cinnamomum zeylanicum No Bhumannavar et. al., 2004 (synonym C. verum)* Liliaceae Allium sativum* No Bhumannavar et. al., 2004; McFadyen et. al., 2003 Malvaceae Abelmoschus esculentus* No Muniappan & Bamba, 2002 Gossypium hirsutum* No Bhumannavar et. al., 2004 Gossypium obtusifolium No McFadyen et. al., 2003 Hibiscus rosasinensis* No Bhumannavar et. al., 2004; McFadyen et. al., 2003 Meliaceae Sweitenia macrophylla* No Aterrado & Bachiller, 2002 Moraceae Morus alba* No Bhumannavar et. al., 2004 Myrtaceae Eugenia aquea (synonym No McFadyen et. al., 2003 Syzygium aqueum)* Eugenia caryophyllus No McFadyen et. al., 2003 (synonym Syzygium aromaticum)* Eugenia jambolana No Bhumannavar et. al., 2004 (synonym Syzigium cumini)* Family Taxon Host? Citation Psidium guajava* No Bhumannavar et. al., 2004; McFadyen et. al., 2003 Oleaceae Jasminum sambac* No Bhumannavar et. al., 2004 Piperaceae Piper methysticum* No Muniappan et. al., 2007 Piper nigrum* No Bhumannavar et. al., 2004 Poaceae Oryza sativa* No Bhumannavar et. al., 2004; Kernasa et. al., 2010; McFadyen et. al., 2003 Saccharum officinarum* No Kernasa et. al., 2010 Sorghum vulgare No Kernasa et. al., 2010 Zea mays* No Bhumannavar et. al., 2004; Kernasa et. al., 2010; McFadyen et. al., 2003; Muniappan & Bamba, 2002 Puniccaceae Punica granatum* No Bhumannavar et. al., 2004 Rubiaceae Coffea arabica* No Bhumannavar et. al., 2004; Kernasa et. al., 2010; McFadyen et. al., 2003 Rutaceae Citrus aurantifolia* No Muniappan & Bamba, 2002; Kernasa et. al., 2010 Citrus nobilis No McFadyen et. al., 2003 Citrus reticulata* No Bhumannavar et. al., 2004; Kernasa et. al., 2010 Murraya koenigii* No Bhumannavar et. al., 2004 Sapotaceae Achras sapota* No Bhumannavar et. al., 2004 Solanaceae Capsicum annuum* No Bhumannavar et. al., 2004; Kernasa et. al., 2010; McFadyen et. al., 2003; Muniappan & Bamba, 2002 Capsicum frutescens* No Kernasa et. al., 2010 Lycopersicon No Bhumannavar et. al., 2004; esculentum* Kernasa et. al., 2010; McFadyen et. al., 2003 Nicotiana tabacum* No Bhumannavar et. al., 2004; McFadyen et. al., 2003 Physalis spp.* No Esguerra, 2002 Solanum melongena* No Bhumannavar et. al., 2004; McFadyen et. al., 2003 Solanum tuberosum* No Bhumannavar et. al., 2004; McFadyen et. al., 2003 Sterculiaceae Camellia sinensis* No Bhumannavar et. al., 2004 Theobroma cacao* No Bhumannavar et. al., 2004; McFadyen et. al., 2003 Umbelliferae Coriandrum sativum* No Bhumannavar et. al., 2004 Verbenaceae Lantana camara* No Bhumannavar et. al., 2004; McFadyen et. al., 2003 Tectona grandis* No Bhumannavar et. al., 2004 Vitex negundo No Aterrado & Bachiller, 2002

Literature referenced:

Aterrado, E. D., and N. S. J. Bachiller. 2002. Biological control of Chromolaena odorata: Preliminary studies on the use of the gallforming Cecidochares connexa in the Philippines. In C. Zachariades, R. Muniappan, & L. W. Strathie (Eds.). 2002. Proceedings of The Fifth International Workshop on Biological Control and Management of Chromolaena Odorata, Durban, South Africa, 23-25 October 2000, pp. 137-139. ARC-PPRI Pretoria.

Bhumannavar B. S., S. Ramani, S. K. Rajeshwari, and B. K. Chaubey. 2004. Host-specificity and biology of Cecidochares connexa (Macquart) (Diptera: Tethritidae) introduced into India for the biologica suppression of Chromolaena odorata (Linnaeus) King & Robinson. Journal of Biological Control, 18(2): 111-120.

Bishop Museum. 2019. Natural Sciences Taxonomic Name and Specimen Search. http://nsdb.bishopmuseum.org/. Accessed June 24, 2019.

Day, M. D., N. Riding, and K. A. D. W. Senaratne. 2016. The host specificity and climatic suitability of the gall fly Cecidochares connexa (Diptera: ), a potential biological control agent for Chromolaena odorata (Asteraceae) in Australia. Biological Science and Technology, 26:5: 691-706.

Esguerra, N. 2002. Introduction and establishment of the Tephritid gall fly Cecidochares connexa on Siam Weed, Chromolaena odorata, in the Republic of Palau. In C. Zachariades, R. Muniappan, and L.W. Strathie (Eds.), Proceedings of The Fifth International Workshop on Biological Control and Management of Chromolaena Odorata, Durban, South Africa, 23-25 October 2000, pp. 148-151. ARC-PPRI Pretoria (2002).

Kernasa, O., P. Buchatian, and W. Suasa-ard. 2013. Biological control of Chromolaena odorata in Thailand. In C. Zachariades, L.W. Strathie, M.D. Day, and R. Muniappan (Eds.). 2013. Proceedings of the Eighth International Workshop on Biological Control and Management of Chromolaena odorata and other Eupatorieae, Nairobi, Kenya, 1-2 November 2010, pp. 141-145. ARC-PPRI Pretoria.

Mabberly, D. 2017. Mabberley’s Plant-Book, A portable dictionary of , their classifications and uses, Fourth Edition. Cambridge University Press.

McFadyen R., R. Desmier de Chenon, and A. Sipayung. 2003. Biology and host specificity of the chromolaena stem gall fly, Cediochares connexa (Macquart) (Diptera: Tephritidae). Australian Journal of Entomology, 42: 294-297.

Muniappan, R., K. Englberger, and G. V. P. Reddy. 2007. Biological control of Chromolaena odorata in the American Pacific Micronesian Islands. In P. Y. Lai, GVP Reddy, and R Muniappan (Eds.), Proceedings of the Seventh International Workshop on Biological Control and Management of Chromolaena odorata and Mikania micrantha, PingTung, Taiwan, 12-15 September 2006, pp. 49-52. National Pingtung University of Science and Technology.

Muniappan, R., and J. Bamba. 2002. Host specificity testing of Cecidochares connexa, a biological control agent for Chromolaena odorata. In C. Zachariades, R. Muniappan, & L. W. Strathie (Eds.), Proceedings of the Fifth International Workshop on Biological Control and Management of Chromolaena odorata, Durban, South Africa, 23-25 October 2000, pp. 134–136. ARC-PPRI Pretoria.

Smithsonian National Museum of Natural History. 2019. Flora of the Hawaiian Islands. https://naturalhistory2.si.edu/botany/hawaiianflora/. Accessed June 24, 2019.

Wikipedia, The Free Encyclopedia. 2019. Asteraceae. https://en.wikipedia.org/wiki/Asteraceae. Accessed June 24, 2019.