42 Plant Protection Quarterly Vol.22(2) 2007 to green to yellow, which is indicative of the Old World Taxa (Dehgan and Webster 1979). Species of Jatropha that have natural- Review ized in Australia include physic nut (J. curcas L.), peregrina (J. integerrima Jacq.), coral plant (J. multifi da L.), Buddha belly (J. podagrica Hook.), and leatherstem (J. dioica Sesse.) (HERBRECS 1998). In Australia, there appears to be several The biology of Australian weeds biotypes with morphological, phenologi- cal, and physiological differences (Pitt and 47. Jatropha gossypiifolia L. Miller 1991, Bebawi and Campbell 2004). Currently three biotypes are recognized F.F. BebawiA,B, J.S. VitelliA,B, S.D. CampbellA,B, W.D. VoglerA,B, C.J. LockettA,B, in Queensland (Bebawi and Campbell B.S. GraceC,B, B. LukitschC,B and T.A. HeardD,B 2004). These include Queensland bronze A Tropical Weeds Research Centre, Department of Natural Resources and leaf and Queensland green leaf, which oc- Water, Charters Towers, Queensland 4820, Australia. cur from Rockhampton north to Cairns and Queensland purple leaf, which oc- B CRC for Australian Weed Management. C curs from Cairns north to Cape York. Two Department of Natural Resources, Environment and the Arts, Darwin, distinct biotypes (Katherine green and Northern Territory 0831, Australia. Darwin purple) are also reported in the D CSIRO Entomology Long Pocket Labs, Indooroopilly, Queensland 4068, Northern Territory (Pitt and Miller 1991, Australia. Bebawi and Campbell 2004). Biotypes ex- isting in Western Australia have not yet been investigated. Detailed taxonomic, genetic and eco- Name purga del fraile, medicinier noir, medic- logical studies are required to verify and Botanical name cinier rouge (Caribbean, Puerto Rico) (Lio- establish differences among the Australian The genus name Jatropha combines the gier 1990), lapalapa pupa, Accra fence tree biotypes. Such studies would also help de- Greek iatros, meaning physician, with tro- (Africa) (Irvine 1961), Pinon Negro (Peru) termine whether there is only one variety pheia, mother’s milk, hinting at the me- (Pinedo et al. 1997), Purgue de fraile, tua present or if some of the noted biotypes dicinal properties of the plant (Parsons tua, piñon colarado, pinon Negro, piñon could in fact be different varieties. Any dif- and Cuthbertson 2001). The species name rojo, quelite del fraile, frailecillo (Latin ferences found may be important to weed ‘gossypiifolia’ is a combination of the Latin America), and pinhão roxo (Brazil). control, particularly with regard to the se- gossypium, meaning cotton, and folium, lection of biological control agents. suggesting that the leaves appear similar Taxonomy and related species in Different varieties of bellyache bush to those of the cotton plant (Parsons and Australia exist outside of Australia (Backer and Ba- Cuthbertson 2001). Synonyms of Jatropha The genus Jatropha belongs to the tribe khuizen van der Brink 1963, Dehgan 1982, gossypiifolia are: Adenoropium gossypiifolium Jatrophieae of Crotonoideae in the fam- Sreenivasa Rao and Raju 1994). For exam- (L.) (Pohl 1827), A. elegans (Pohl 1827), J. ily Euphorbiaceae and the genus contains ple, J. gossypiifolia L. var. elegans was listed elegans (Klotzsch 1853), and J. staphysagri- approximately 186 species (Govaerts et al. in the fl ora of Java (Backer and Bakhuizen folia (Miller 1754). 2000). Dehgan and Webster (1979) divided van der Brink (1963) and three varieties, the genus into two subgenera (Curcas and J. gossypiifolia var. elegans, J. gossypiifolia Common names Jatropha) with 10 sections and 10 subsec- var. gossypiifolia and J. gossypiifolia var. In Australia and America, Jatropha gossypi- tions. They postulated that physic nut (J. staphysagrifolia (Mill.) Müll. were identi- ifolia L. is most commonly known as belly- curcas L.) is the most primitive form of the fi ed in the United States (Dehgan 1982). ache bush (Everist 1974, Dehgan 1982, Par- genus and that J. gossypiifolia evolved from Sreenivasa Rao and Raju (1994) reported sons and Cuthbertson 2001). Other names the physic nut. The facultative annual J. gossypiifolia L. (var. gossypiifolia and var. used in countries where it is found in- growth habit (apical dominance absent) elegans (Pohl) Müll.Arg.) from India. clude: cotton-leaf physic nut, castor bean of bellyache bush is considered a more (Queensland) (Kleinschmidt and Johnson phylogenetically advanced growth habit Description 1987), red physic nut (Burkill 1994), cotton- than the arborescent growth habit (apical Bellyache bush is an erect, woody, de- leaf jatropha, purging nut, Spanish physic- dominance present) of physic nut (Dehgan ciduous, tropical or sub-tropical perennial nut tree, wild physic nut, American purg- and Webster 1979). A taxonomic character- shrub (Figure 1). It is diploid and the ba- ing nut, wild cassava (BoDD 2004), damar istic of the genus Jatropha is the occurrence sic chromosome number is 11 (2n = 22) merah; jarak kosta merah, red physic nut, of either latex-cells or latex vessels (Rao (Nanda 1962, Datta 1967, de Padua et al. jarak kling, jarak ulung, kaleke bacu (Indo- and Malaviya 1964). 1999). Plants commonly grow between 2 to nesia) (BoDD 2004), jarak beremah, jarak Although most Jatropha species are na- 3 m high (Parsons and Cuthbertson 2001), hitam, jarak merah (Malaysia) (de Padua tive to the New World (the Americas), no but can reach up to 4 m in height under et al. 1999), lansi-lansinaan, tagumbau- complete revision of the Old World (Eu- favourable conditions (Bebawi and Camp- a-nalabaga, tuba-tuba (Philippines) (de rope, Asia, and Africa) Jatropha exists (Hel- bell 2002b, Vitelli and Madigan 2004). Padua et al. 1999), nhao luat (Laos) (Padua ler 1996). Hemming and Radcliffe-Smith Bellyache bush exhibits impressive fo- et al. 1999), sabuu daeng, sabu lueat, salot (1987) revised 25 Somalian species, all of liar, fl oral and stem diversity. The stems daeng (Thailand) (de Padua et al. 1999), the subgenus Jatropha, and placed them in are thick, rather soft, coarsely hairy, 1–2 m d[aaf]u kai ti[as) (Vietnam) (de Padua et six sections and fi ve subsections. The base long and exude a watery sap when dam- al. 1999), red fi g-nut fl ower, sibidigua, Tua- colour of the petals of Australian biotypes aged (Parsons and Cuthbertson 2001). The Tua, Badigaba, Baigaba, Benderi, Lanka- is predominantly red, which is indica- number of stems per plant can range from jada, red varendra, (India) (Parrotta 2001), tive of their American origin as opposed one to two or more (Csurhes 1999, Parsons Plant Protection Quarterly Vol.22(2) 2007 43 and Cuthbertson 2001). Stems are green relatively large anthers in the upper tier are endospermic, contain oil, and ovoid when young, turning bright crimson red at and fi ve relatively small in the lower. The in shape (Figure 4) (Singh 1970, de Padua fl owering in some biotypes but invariably anthers are dorsifi xed. Pollen grains are et al. 1999), they exhibit considerable vari- turn ‘grey’ with age across all biotypes. spherical, bright yellow with a sticky, oily ation (Liogier 1990, Howard 1989, Burkill Leaves of bellyache bush are arranged coating (Reddi and Reddi 1983). 1994, Parrotta 2001, Bebawi and Campbell alternately along the stem (Parsons and Female flowers are quite similar in 2002a). Cuthbertson 2001). Leaves may be bright shape to male fl owers but they are larger, When dissected, seeds have a thin, green, dark green, bright or dark bronze, with a diameter of up to 9 mm. Sepals as outer whitish-fleshy envelope (exote- bright red or bright purplish-red depend- well as petals are larger than those on the gmen) that surrounds a thin and shell-like ing on biotype and leaf maturity (Pitt and male. Styles are 3–4, slender dilated into a testa (Figure 5). A longitudinal section of Miller 1991, Burkill 1994, Parsons and capitate, bifi d stigma. The stigmas are pale mature seeds show an exotegmen con- Cuthbertson 2001, Bebawi et al. 2005b). green and very sticky (Reddi and Reddi nected to the caruncle at the micropylar Leaf petioles are 2–7 cm long and the leaf 1983). region (proximal end) of the seed by dense blades are palmately 3–5 lobed, generally There are fi ve united sepals, which are 45–90 × 50–130 mm in size, and more or ovate to lanceolate, acuminate with stalked less elliptic (Figure 2) (Wheeler 1992). The capitate glands on the margins, distinctly lamina is glabrous and the leaf margin is imbricate and 4 mm long. They are green denticulate with venations ending in stipi- in the green biotype or with margins tate glandular hairs. Leaf venation is white tinged with bronze in the bronze biotype in the green biotypes and shades of red and with purple in the purple biotype. in the other biotypes (F.F. Bebawi unpub- The fi ve petals are reddish in Queensland lished observations). biotypes and Darwin biotype but dark Infl orescences are glandular and hairy purple in Katherine biotype, obovate and (de Padua et al. 1999). Flowers are pedi- 3–5 mm long. Sepals touch at the base to cellate, terminal and occur in corymbose form a short green tube and have stipitate cymes. Flower bracts are linear-lanceo- marginal glands (F.F. Bebawi unpublished late with glandular margins (Dehgan and 2006). Webster 1979). Flowers are radially sym- The fruit of bellyache bush has three metrical and loaded with nectaries. Flow- lobes (locules) (Figure 3). It is an explo- ers of the Queensland biotypes are small, sively dehiscent capsule with a single seed generally dark red or maroon with bright per locule and axile placentation. It is glo- yellow centres. Those of the Northern Ter- bose, pedicellate, generally bright green Figure 2. Digitately lobed leaves of ritory are dark purple with dull yellow and woody at maturity, turning pale green bellyache bush. centres in the Katherine biotype or light or tan when ripe (Berg 1975, Dehgan and red with bright yellow centres in the Dar- Webster 1979, Burkill 1994, Parrotta 2001). win biotype (F.F. Bebawi unpublished ob- Most fruits bear three seeds. servations). Bellyache bush seeds possess ant-at- Male fl owers are cup-shaped with a tractant substances, are carunculate, soft, diameter of 6–9 mm. There are eight sta- slippery and glossy (Bebawi and Camp- mens all fused into a tubelike structure, bell 2002a). The caruncle is a pale, spongy connate at the base, the upper portion free outgrowth rich in lipids, proteins, and with two tiers of anthers of unequal length carbohydrates (Bebawi and Campbell (Reddi and Reddi 1983). There are three 2004). While seeds have a dry testa and

Figure 3. Infructescence of bellyache bush showing trilocular capsules. Graduated scale is in millimetres.

(a)

caruncle (b)

Figure 4. Bellyache bush seeds: (a) ant-discarded without caruncle and Figure 1. Adult bellyache bush shrub (Queensland bronze leaf biotype) (b) intact with caruncle. Graduated showing sympodial growth habit. scale is in millimetres. 44 Plant Protection Quarterly Vol.22(2) 2007 parenchymatous tissue (Figure 6). When with two papery cotyledons in the centre History the seed ripens the surrounding exote- (Figure 6). Seed germination is epigeal, i.e. Though native to the drier islands of the gmen fuses to the seed tegmen. The ‘exo- the cotyledons appear above the ground Caribbean and the Venezuelan coastline tegmen’ is fully transparent, shiny, and surface. (Hickman 1974, Heard et al. 2002), the cur- slippery when mature, and securely fused Latex cells occur in the stem, leaf, peti- rent geographic range of bellyache bush to the testa (Figure 6). The testa tegmen is ole, fl ower parts, fruit, and in the seed coat (Figure 7) is large because of dispersal below the ‘exotegmen’. When the testa is (Rao and Malaviya 1964). Latex from the by humans for medicinal and ornamen- removed, a white mass called the kernel is shoot apex is nearly white. Dehgan and tal purposes (Perry 1980, Dehgan 1982, left. The kernel is enclosed in a very thin Webster (1979) also recognized four extra- Burkill 1994). membrane (endotegmen). The kernel con- fl oral gland systems in the lamina, petiole, Bellyache bush was introduced to north- sists predominantly of a thick, soft, and stipules and bracts in the genus Jatropha. ern Australia in the latter part of the 1800s oily endosperm (Singh 1970, Bebawi and (Everist 1974, Pitt and Miller 1991, Parsons Campbell 2004) that surrounds an embryo and Cuthbertson 2001). It was introduced to the Northern Territory in 1888 with other Jatropha species, probably for me- dicinal and ornamental purposes (Anon. Multifi d caruncle Exotegmen Exotegmen 1888, Pitt and Miller 1991, Pitt 1997) and was listed as a naturalized component of Endosperm Queensland’s fl ora in 1912 (Bailey 1912). The fi rst recorded specimen from Queens- Cotyledons land was collected from Townsville in 1913 (HERBRECS 2004). Embryo It has been suggested that bellyache Testa bush infestations in Queensland originat- Testa ed from the Charters Towers area, prob- ably from garden specimens planted prior Caruncle to 1900 (P. Jeffrey personal communication Figure 5. Immature bellyache bush 1998, cited in Csurhes 1999). At that time, seed showing crustaceous seed testa reticulated water was unavailable in the overlaid with a hard gelatinous Figure 6. Median-longitudinal town and local residents tended to plant tissue of exotegmen joined to a section of a mature bellyache bush very hardy plants such as bellyache bush. multifi d caruncle at the proximal seed showing regional distribution By the early 1920s bellyache bush was grown extensively in Queensland as an end of the seed. Graduated scale is of internal and external seed ornamental shrub. in millimetres. structural components.

Figure 7. World distribution map of bellyache bush indicated by (c) across all continents. Plant Protection Quarterly Vol.22(2) 2007 45 Distribution Australia The current distribution of bellyache bush in Australia is limited to the northern half of the continent (Figure 8) (Ashley 1995). In Northern Queensland, it is present in riparian and sub-riparian habitats of the Burdekin, Walsh, Palmer, Flinders, and Gregory Rivers and the headwaters of Lake Eyre Basin. Small scattered infesta- tions occur in central Queensland, partic- ularly in the Fitzroy catchment (Csurhes 1999, Barron 2004). In the Northern Territory, bellyache bush persists in several coastal areas, but is most prevalent further inland (Csurhes 1999). Among the worst infestations in the Northern Territory are those on the Daly River catchment at Willeroo Station and Scotts Creek Station, the Gulf of Carpen- taria region and the Barkly Tablelands. Other infestations occur in the McArthur River, Roper River, and Victoria River catchments (Miller and Pitt 1990). Figure 8. Current distribution of bellyache bush in Australia. In Western Australia, widespread, un- controlled populations occur in the east Kimberley, with larger infestations in the Lake Argyle catchment and near Halls Habitat dry conditions, bellyache bush is capable Creek (Parsons and Cuthbertson 2001, Bellyache bush is an opportunistic coloniz- of spreading. For example, on a property Anon 2005). Small controlled infestations er of disturbed habitats and is frequently located in the Ravenswood area of north- occur in the west Kimberly region at the found in areas where the natural vegeta- ern Queensland, a bellyache bush infesta- De Grey River, Broome, and Fitzroy River tion has been over-grazed or removed by tion increased in size by 76% over three and on the Drysdale River in the north human activity or fl oodwaters (Csurhes years, despite the area receiving only half Kimberly (King and Wirf 2005). 1999). In Java (Backer and Bakhuizen van its average annual rainfall (Vogler and The potential range of bellyache bush den Brink 1963) and Andhra Pradesh in Keir 2005). includes the entire tropical savannas India (Rao and Raju 1994), it has become In Australia, bellyache bush grows (Thorp and Lynch 2000). very common along roadsides, railway equally well in saline and non-saline soils tracks and eroded places. Similarly, Csur- but seems to prefer sandy loams (Csurhes Outside Australia hes (1999) remarked that bellyache bush 1999). It has been found growing down to Bellyache bush is cultivated widely in has become abundant in northern Aus- the high tide mark near Darwin. In Puerto tropical countries throughout the world tralia along roadsides, around abandoned Rico, it is more common in soils with high (Figure 7) (Burkill 1994, de Padua et al. homesteads and near old mine sites, sug- base saturation, such as dry areas, sites 1999). In North America it is found mainly gesting that colonization in such areas is near the ocean, and soils derived from in Florida and in Mexico (Francis 2005). probably an indicator of disturbance. limestone (Liogier 1990). In South America it occurs predominantly Bellyache bush is particularly well Bellyache bush is very sensitive to frost, in Bolivia, Venezuela, Ecuador, Peru, and adapted to the seasonally wet/dry climate which damages the apices of stems (F.F. Brazil (de Padua et al. 1999). It is also a of northern Australia (Csurhes 1999). In Bebawi personal observations). It may common plant in the Caribbean, mainly in Queensland, bellyache bush invades also suffer from waterlogging in poorly the Dominican Republic, Puerto Rico and lowland habitats such as riparian zones, drained soils (Dehgan 1982) or as a result Leeward Islands, as well as in the Hawai- ephemeral watercourses and pastures and of fl ooding. In North Queensland, fl ood- ian Islands (Csurhes 1999, de Padua et al. is also a serious invader of disturbed up- ing of the Palmer River in March and April 1999). lands around Townsville, North Queens- 2006 killed all plants that were growing in Introduced to southern Africa, the plant land. Riparian zones are most vulnerable the riverbed (F.F. Bebawi personal obser- has spread from Mozambique through to bellyache bush invasion, possibly be- vations). Zambia to the Transvaal and Natal. In cause fi re often fails to penetrate riparian West Africa, it is listed in the exotic fl ora of vegetation (Csurhes 1999). Growth and development Chad (Brundu and Camarda 2004), Cam- Most naturalized populations of belly- Morphology eroon and Ghana (Csurhes 1999). ache bush in Australia grow in areas re- Bellyache bush stem growth habit is sym- Bellyache bush is also found through- ceiving 400–1200 mm average annual podial (Figure 1) and apical dominance out the warmer parts of Asia and the Pa- rainfall, with some of the heaviest infesta- is absent because branching occurs at cifi c, particularly in Indonesia, Singapore, tions found where average annual rainfall reproductive maturity and thereafter at New Guinea and New Caledonia (Holm et is 600–1000 mm (Csurhes 1999). Overseas subsequent fl owering episodes. Follow- al. 1979, Wilson 1997, de Padua et al. 1999, in Puerto Rico, it may be found in areas ing germination, the primary stem (main IPCS INCHEM 2004). It occurs frequently receiving from 750 to about 2000 mm of axis) grows until fl owering is initiated (F.F. on plains but rarely in uplands and hilly annual precipitation (Liogier 1990). In Bebawi personal observations). areas in India and New Caledonia. wetter areas such as parts of the North- In its native environment, bellyache ern Territory, bellyache bush has shown bush seldom reaches a height of more than a 146% expansion of its area per year (Pitt 20–30 cm (Hickman 1974). In Queensland, and Miller 1991). However, even under plants are capable of growing to 4 m with 46 Plant Protection Quarterly Vol.22(2) 2007 a canopy diameter of 2 m and a basal stem Elevated CO2 had no signifi cant effect retain most of their leaves all year (Liogier diameter up to 15 cm (Bebawi and Camp- on morning xylem water potential, leaf 1990, Howard 1989, Burkill 1994, Parrotta bell 2002b, Vitelli and Madigan 2004). osmotic potential, or pressure potential 2001). Heights over 3 m have also been reported of bellyache bush grown under simulated Flowering generally occurs from June in shaded areas in the Northern Territory seasonal drought conditions (Rengifo et al. to April in riparian zones and from Sep- (Pitt and Miller 1991). The stunted form of 2002). Apparently, bellyache bush has the tember to April in sub-riparian zones of the plant in the native range has been at- capacity to respond to elevated CO2 and North Queensland (Bebawi et al. 2005b). tributed to herbivory (Heard et al. 2002). simulated drought by signifi cantly reduc- While moisture availability appears to be a In Australia, bellyache bush is capable ing the proportional thickness of its leaf key driver of fl ower production, tempera- of attaining a height of up to 2 m in a sin- photosynthetic tissue systems (Rengifo ture may infl uence timing and duration gle growing season (Dehgan and Webster et al. 2002) as well as by closing stomata (Bebawi et al. 2005b). In the Kimberley re- 1979), particularly in areas free from com- (Tezara et al. 1998). This enhances water gion of Western Australia, bellyache bush petition, such as riverbeds in Far North use effi ciency and increases the photosyn- generally fl owers and fruits from February Queensland and in the Northern Territory. thetic rate. Bellyache bush may therefore to May (Wheeler 1992). In North Queens- In a competition trial in the dry tropics of exhibit increased biomass and growth land, fruit production commences around North Queensland, bellyache bush grown rates, leading to further expansion of its September/October within riparian and on pasture-cleared plots reached average current distribution, if atmospheric CO2 sub-riparian zones and may last up to 10 heights of 64 cm and 113 cm after the fi rst increases further. months (Bebawi et al. 2005b). and second year of establishment respec- tively (F.F Bebawi unpublished results). Phenology Reproduction Similarly, in Puerto Rico bellyache bush In Australia, bellyache bush loses most of Floral biology was found to grow up to 0.5 m per year its leaves during the dry season (Figure Bellyache bush is capable of reaching re- (Liogier 1990). 9). Those that remain are generally quite productive maturity very quickly under Bellyache bush root systems are rela- small and concentrated at the apices of favourable moisture conditions. In pot tri- tively small compared with their shoot stems (Bebawi et al. 2005b). The transition als, plants fl owered 55 days after germi- systems (Pitt and Miller 1991), with stem between the two stages is fairly rapid— nation (Bebawi et al. 2005c). In the fi eld in to root ratios increasing with age. The plants are either in full leaf or almost leaf- North Queensland, the time to fi rst fl ow- stem to root dry weight biomass ratio of less (Bebawi et al. 2005b). ering averaged 74 days in cleared areas, juvenile, mature, and adult bellyache bush In North Queensland, an average of 17 294 days in rocky sites, and 454 days in plants in North Queensland averaged 5.6, and 20 leaves per stem have been recorded grazed pastures (Bebawi et al. 2005c). It 6.1 and 7.1 to 1, respectively (F.F. Bebawi during the wet season (November–April) was suggested that more plants fl owered unpublished results). on plants growing within sub-riparian in cleared areas and reached reproductive Bellyache bush has a fl eshy shallow and riparian habitats, respectively (Be- maturity earlier because of the absence root system with four short robust lateral bawi et al. 2005b). On average there is of interspecifi c competition (Bebawi et al roots and many fi ne tertiary roots (Singh one leaf per stem in the dry season (June– 2005c). Increasing bellyache bush density 1970, Liogier 1990, Howard 1989, Burkill August). However, along watercourses also caused exponential increases in time 1994, Csurhes 1999, Parrotta 2001). Fresh and dam walls bellyache bush plants may to fi rst fl owering (Bebawi et al. 2005c). At root weight of a dense infestation (20 000 plants ha-1) of juvenile (up to 20 cm height), mature (20–100 cm height) and adult (>100 25 cm height) bellyache bush plants in north Queensland averaged 0.5, 2.8 and 10.5 t Riparian ha-1 (25, 137, and 523 g plant-1), respec- Sub-riparian tively. Root moisture content of juvenile, 20 mature, and adult bellyache bush plants is relatively similar, averaging 81%, 73%, and 71%, respectively (F.F. Bebawi unpub- lished results). 15

Perennation Whilst bellyache bush is a perennial plant, there is a paucity of information on its 10 longevity. Observations of tagged plants in fi eld trials indicate that plants can live for longer than six years (F.F. Bebawi un- published 2006), with anecdotal evidence 5 suggesting greater than 20 years. Average no. of leaves per flowering branch Physiology

Bellyache bush utilizes the C3 photosyn- thetic pathway (Tezara et al. 1998, Fern- AMJ JA SOND J FM andez et al. 1999, Bebawi and Campbell

2002b). C3 plants are more effi cient than 1999 2000 C4 and CAM (Crassulacean Acid Metabo- lism) plants under cool and moist condi- Months and year tions and under normal light because they require fewer enzymes and no specialized Figure 9. Monthly leaf density of bellyache bush (1999–2000) associated anatomical adaptations (Rengifo et al. with light density beneath and above (control) the canopy of bellyache 2002). bush. Vertical bars indicate the s.e. of the mean. From Bebawi et al. (2005b). Plant Protection Quarterly Vol.22(2) 2007 47 the lowest density (20 plants m-2) plants Seed production and dispersal and exotegmen of bellyache bush seeds fl owered 55 days after germination. In Seed production of bellyache bush is gen- and when fi nished discard the seed in contrast, only 6.3% of plants within the erally prolifi c, but many factors such as their middens. The middens provide an highest density (320 plants m-2) had fl ow- environmental conditions, plant biotype, improved environment (e.g. high nutrient ered 42 months after germination (Bebawi plant density and location can infl uence status and absence of fi re) for germination et al. 2005c). In a pot trial, the percentage the quantity of seeds produced. and survival of seedlings. of bellyache bush plants that fl owered de- Adult plants growing in Queensland Water, humans and other animals, par- creased exponentially with increased den- have been found to produce between 2000 ticularly the great bowerbird (Chlamydera sity (Bebawi et al. 2005c). and 12 000 seeds plant-1y-1 (Bebawi and nuchalis Jardine & Selby) are identifi ed as Once bellyache bush plants reach re- Campbell 2002a), which corresponds to potential long-distance dispersers of belly- productive maturity, they have a defi nite around 170 kg ha-1. In Western Australia ache bush seeds (APB Infonote 1994, Ash- pattern of fl owering within infl orescences. bellyache bush is reported to produce be- ley 1995, Smith 1995, F.F. Bebawi personal More female than male fl owers mature on tween 1800 and 2400 seeds plant-1y-1 (APB observations). Of these, fl oodwater is con- the fi rst day, with female fl owering show- Infonote 1994). Annual seed production of sidered most important within catchments ing a declining trend thereafter over a bellyache bush in the Indian sub-continent and humans are recognized as the main fi ve day period (Reddi and Reddi 1983). was estimated at 500 kg ha-1 (Raina and dispersers of bellyache bush at a national In contrast, the number of male fl owers Gaikwad 1987). and global scale (Csurhes 1999). gradually increases so that they dominate Dense infestations of bellyache bush by the twelfth day, after which their fre- growing in a relatively dry location in Germination quency gradually declines. In effect, the North Queensland produced four seeds Fresh bellyache bush seeds exhibit high infl orescence is protogynous (Reddi and m-2 during the 2003–2004 wet season (288 viability, but low germinability (Bebawi Reddi 1983). Dehgan and Webster (1979) mm rainfall), in comparison to 343 seeds and Campbell 2004). For example, fresh showed that there was a lag of 24–48 h in m-2 for plants growing in a wetter location intact seed collected in North Queensland anthesis of male fl owers after opening of (506 mm rainfall) (Vogler and Keir 2005). was 88% viable but only 10% of these female fl owers. Plants produce fewer seeds at high den- were readily germinable (Bebawi and The ratio of male to female fl owers is sity. Above 40 plants m-2, seed production Campbell 2004). Similarly, germination of on average 11:1 (Reddi and Reddi 1983, per plant began to decline. At very high seeds from Puerto Rico averaged just 4% Wild 2003). Dehgan (1983) found that densities (in excess of 300 plants m-2), there (Liogier 1990). Innate (primary) dormancy short days result in production of more was very little seed production (Bebawi et has been reported for other Euphorbiaceae male fl owers, while long days caused ei- al. 2005c). (Ellis et al. 1985). ther a drastic increase in or total change to A preliminary pot trial showed that Seed type, seed weight, geographical female fl owers. Consequently, the ratio of Queensland bronze biotype plants pro- location, temperature, control technique, male to female fl owers and therefore seed duced 102 seeds plant-1 month-1 compared and ants affect germination of bellyache production may be infl uenced by latitude with 62 seeds plant-1 month-1 for the Kath- bush seed (Liogier 1990, Bebawi and and season. erine green biotype (F.F Bebawi unpub- Campbell 2002a, 2002d, 2004). Seeds com- Female fl owers produce nearly 1.6 times lished results). mence germination with the start of the more nectar than male fl owers (Reddi and Seed dispersal occurs initially via de- wet season (Ashley 1995) but will germi- Reddi 1983). A positive association has hiscent capsules that are capable of cata- nate at other times of the year if environ- been detected between temperature and pulting seeds as far as 13 m (Bebawi and mental conditions are favourable. For ex- nectar concentration and an inverse one Campbell 2002a), although in dense infes- ample, in North Queensland germination between relative humidity and nectar con- tations most seeds fall close to the parent will often occur throughout the year on centration (Reddi and Reddi 1983). plant. rainfall events that exceed 25 mm (F.F Be- Bellyache bush nectar consists of glu- Once on the ground, some ants (particu- bawi unpublished results). cose, sucrose and fructose, amino acids larly native meat ants (I. spadius)) disperse Ants promote the germination of belly- and proteins (Reddi and Reddi 1983). The bellyache bush seeds (Figure 10) (Bebawi ache bush seeds (Bebawi and Campbell nectar is attractive to insects, which are and Campbell 2002a). In one study, an av- 2004). In a laboratory study 98% of ant- essential for normal seed set (Reddi and erage of 12 330 seeds were retrieved from discarded seeds were viable and read- Reddi 1983, Wild 2003). Pollination may the middens (refuse piles) of individual ily germinable (100%). Viability of intact occur through selfi ng, because the fl ow- meat ant nests over 12 months, with high- seeds that had the caruncle manually re- ers are self-compatible, or through out- est numbers recorded between February moved was on average 10% lower than crossing (Dehgan and Webster 1979). The and June (Bebawi and Campbell 2004). that of ant-discarded seeds, and only 8% of former mechanism ensures perpetuation Meat ants appear to feed on the caruncle fresh viable intact seeds were germinable of the species and the latter maintains spe- (Bebawi and Campbell 2004). Removal of cies heterozygosity (Dehgan and Webster caruncles imitates the effect of meat ants 1979). on germination but not to the same ex- Many insect species have been observed tent as in ant-discarded seeds (Bebawi and foraging on nectar of bellyache bush in Campbell 2004). Additional scarifi cation Queensland (Appendix 1). Most of the in- by ants on external seed structures may sects listed in Appendix 1 are benefi cial to also promote germination. bellyache bush because they perform vital Optimal germination temperatures for roles such as pollination, defence and dis- intact bellyache bush seeds occur between persal. For example, meat ants (Iridomyrmex 24 and 31°C. Germination generally com- spadius Schattuck), honey bees (Apis mellif- mences fi ve days after the imposition of era Lepeltier) and oleander butterfl ies (Eu- Bellyache bush seed favourable environmental conditions and ploa core-corinna Craemer) visit fl owers and reaches a maximum between days 11 and perhaps pollinate bellyache bush. Hives of 12 (F.F Bebawi unpublished results). the stingless bee Trigona carbonaria Smith Figure 10. Meat ants (Iridomyrmex Burial depth and litter cover also af- were successfully used to pollinate belly- spadius) dispersing bellyache bush fect germination and viability follow- ache bush grown in glasshouses. seed into nest entrance. ing fi re (Bebawi and Campbell 2002d). 48 Plant Protection Quarterly Vol.22(2) 2007 Germination and viability were negative- cember 1999 and crashed to 30 seedlings the Northern Territory (draft regional plan ly correlated with peak fi re temperature, m-2 one year later. In unburnt plots, seed- www.katherineweeds.nt.gov.au). which was affected by litter cover. No vi- lings peaked at 200 m-2 before crashing Lantana (Lantana camara L.) was once able seeds remained under litter cover, but to 5 m-2 during the same period (Bebawi the main shrub weed around Palu in >80% of seeds placed on bare ground or and Campbell 2002c). A fi eld trial that Central Sulawesi, but it has now been re- 2 cm below ground remained viable (Be- compared the impact of a range of four placed by Siam weed (Chromolaena odorata bawi and Campbell 2002d). Fire reduced bellyache bush seedling densities on seed- L.), which is currently being overtaken by germination and viability of seeds within ling survival showed that 12 months after bellyache bush (Partridge 2001). capsules by 31% and 35% respectively seedling emergence, seedling density was when compared with unburnt seeds (Be- reduced by 75, 89, 98 and 99% at 25, 125, Detrimental bawi and Campbell 2002d). 250 and 500 seedlings m-2 respectively (F.F. Economic impact The economic impact Bebawi unpublished results). of bellyache bush on the livestock indus- Seed longevity Even if high mortality of seedlings oc- try has been reported in Australia (Tothill In a seed burial trial comparing germina- curs under relatively dry conditions, suf- et al. 1982), particularly in North Queens- tion and viability of intact and ant dis- fi cient recruitment for re-infestation of land, where direct losses to the pastoral persed seeds exposed to either nil (rain- treated sites and expansion of infestations industry occurred during drought due to fall excluded) or natural rainfall, no intact occurs in the absence of followup control poisoning of cattle, horses and goats (Csur- seeds exhumed after four years remained activities. Observations by the authors hes 1999). All cases of poisoning occurred viable under natural rainfall conditions, suggest that once seedlings reach about in the dry season when pasture quality whereas some ant-discarded seeds were 20 cm in size they are very hardy and will and quantity was at its lowest. Two land- still viable (3%). However, both intact generally tolerate extreme environmental holders alone have spent approximately and ant-discarded seeds exhumed after conditions. $56 240 on control of bellyache bush in the four years were 20% viable when rain Burdekin Catchment, North Queensland was excluded (F.F. Bebawi unpublished Vegetative reproduction (Csurhes 1999). A property owner within results). Bellyache bush can readily regenerate from the Burdekin Catchment spent more than In a seed bank depletion trial, seedling stem cuttings (e.g. dumped garden plant $50 000 over 10 years in an attempt to emergence was still occurring after 51 material) (Pitt and Miller 1991, Parsons prevent bellyache bush spreading along months at a rocky site away from a river, and Cuthbertson 2001) and whole plants 17 km of river frontage (Bowen Independ- whereas emergence had fi nished earlier that may be removed during control ac- ent 1996). (37 months) at a heavy clay soil site within tivities or fl ood events. The shallow root In the Northern Territory, bellyache the same period (F.F. Bebawi unpublished system of bellyache bush allows seedlings bush has spread into pastoral land, form- results). Differences in seed bank depletion and mature plants to be easily dislodged ing dense thickets with little or no grass were attributed to differences in soil mois- (Pitt and Miller 1991). underneath, and making land unsuitable ture conditions between the two sites. Several landholders have found belly- for grazing (Miller 1982). In Papua New ache bush plants reshooting several Guinea, sown pastures have become non- Seedling establishment months after they were pulled and left viable due to encroachment of several Seedling densities can be very high un- lying on the ground. Similarly, during a weeds including bellyache bush (Chand- der favourable environmental conditions. simulated slashing trial most off-cuts of hokar 1978). Averages of 247, 126, and 90 seedlings m-2 bellyache bush (particularly those cut dur- were measured within rocky, sub-riparian ing the dry season) fl owered and produced Environmental impact Bellyache bush and riparian infestation of bellyache bush, capsules with viable seed up to 12 months invasion results in a loss of biodiversity, respectively (F.F. Bebawi unpublished re- after being cut (Bebawi and Campbell wildlife habitat, changed fi re regimes, in- sults). Higher seedling densities (300–400 2002b), although a few of these had grown creased soil erosion and destabilization seedlings m-2) were recently recorded un- some new roots which were connecting of creek and river banks. Monospecifi c der dense canopies of bellyache bush (Vo- them to the ground. stands of bellyache bush inhibit establish- gler and Keir 2005). ment of native plants (Csurhes 1999). For Treatment of bellyache bush infestations Importance example, in North Queensland, there was can result in massive recruitment of seed- Bellyache bush is considered to be one of no black speargrass (Heteropogon contortus lings (Bebawi and Campbell 2002c, Vitelli the more poisonous and aggressive weeds L.) remaining under the canopy of belly- and Madigan 2002, Bebawi et al. 2004). For growing in the dry tropics of northern ache bush after two years (F.F. Bebawi un- example, bellyache bush increased in den- Australia (Csurhes 1999, Parsons and published results). sity from fi ve plants m-2 prior to aerial ap- Cuthbertson 2001). While it is in the rela- An aqueous extract of bellyache bush plication of foliar herbicides to 400 plants tively early stages of invasion, Lazarides latex is known to kill freshwater fi sh (Singh m-2 post treatment (Vitelli and Madigan and Hince (1993) indicated that bellyache and Singh 2002). The methanol and n-bu- 2002). Similarly, for every plant killed by bush has the potential to spread over tanol extracts of unripened seeds of belly- foliar spraying, slashing, stickraking and 75% of the Australian continent (includ- ache bush have also killed two species of burning as part of an integrated research ing the Northern Territory, Western Aus- freshwater snails (Lymnaea luteola L. and trial, 20, 97, 74 and 69 seedlings emerged tralia, and Queensland), where it would Indoplanorbis exustus Deshayes) overseas respectively (Bebawi et al. 2004). Treat- cause significant environmental, eco- (Amin et al. 1972, Adewunmi and Marquis ments that caused the greatest soil distur- nomic and social impacts. Bellyache bush 1980, Singh and Agarwal 1988, Sukuma- bance appeared most conducive for seed- was ranked number 21 of all widespread ran et al. 1995). ling recruitment. weeds in Australia (Thorp and Lynch High seedling mortality generally oc- 2000). It ranked number three for North Social impacts Detrimental effects on hu- curs, particularly if rainfall is limited Queensland (Bebawi et al. 2002), and four man health include seed poisoning (Kings- and/or there is strong intra- or interspe- for South Queensland (Walton and Elliot bury 1964), dermatitis (Souder 1963) and cifi c competition (Bebawi and Campbell 2003) in weed research priority exercises. sneezing (Irvine 1961). Although numer- 2002c, Vitelli and Madigan 2002). For It is also listed as a priority weed in West- ous cases of severe poisoning have been example, seedling density in burnt plots ern Australia (Pitt et al. 1990, Pitt 1999) and reported from the plant’s native range, no reached a peak of 390 seedlings m-2 in De- in the Katherine and Darwin regions of human deaths were recorded (Begg and Plant Protection Quarterly Vol.22(2) 2007 49 Gaskin 1994). However, if bellyache bush attributes. Various parts have been studied additive for plastic formulations (Ogbobe infestations become more widespread in as sources of novel pharmaceuticals, includ- and Akano 1993) and a source of insecti- northern Australia, particularly around ing potential anticancer drugs (Biehl and cides (Prasad et al. 1993, Chatterjee et al. schools, parks and other public amenity Hecker 1985, de Padua et al. 1999). Roots, 1980). In Asia, bellyache bush is used for areas, risks to human health may become stems, leaves, seeds, and fruits have been dye (Smith 1995) and for production of more serious. widely used in traditional folk medicine in biogas when used as pressed mud cake All parts of bellyache bush are con- many parts of western Africa (de Padua et (Abubacker et al. 1999). Oil extracted from sidered toxic but the seeds are especially al. 1999, IPCS INCHEM 2004). Extracts from bellyache bush seeds is also used as an so (Gardner and Bennetts 1956, Oakes the plant have been used to treat a number illuminant in Africa (Burkill 1994). It is and Butcher 1962, Kingsbury 1964, Mar- of human ailments, ranging from anaemia, possibly still sold in some nurseries in cano-Fondeur 1992, Wheeler 1992, IPCS vertigo, worms, leprosy, leukaemia, dys- Australia (Calvert 1999) and elsewhere as INCHEM 2004). Main toxins include pur- phonia, urinary complaints, ulcers, itches, an ornamental due to its colourful bronze gative oil and curcin, which is found main- conjunctivitis, dermatitis, gout, snakebite leaves (Howard 1989, Parsons and Cuth- ly in the seeds and also in the fruit and and venereal diseases (Irvine 1961, Kup- bertson 2001). sap (Chopra and Badhwar 1940, Simonsen chan et al. 1976, Morton 1981, Liogier 1990, In drier regions of West Africa, belly- 1945, Gardner and Bennetts 1956, Morton Das and Das 1994, Horsten et al. 1996, de ache bush is used as a hedge around vil- 1981, Joubert et al. 1984, Marcano-Fondeur Padua et al. 1999). lages to protect them against bush fi res (Ir- 1992, Burkill 1994, Parsons and Cuthbert- In Peru the leaves and latex are used vine1961, Ogbobe and Akano 1993). Some son 2001, IPCS INCHEM 2004). Curcin is to treat abscesses, tonsillitis, asthma, diar- West Africans also believe that bellyache similar to ricin, the toxic protein of castor rhoea, toothache, fever, gingivitis, fungal bush has magical powers that protect oil plant (Ricinus communis L.). Bellyache skin infections, infl ammations, burns and against snakes, lightning, and violence bush seeds have been used criminally in cough (Pinedo et al. 1997). (Burkill 1994). America (Irvine 1961). In certain African countries, people are Clinical symptoms of bellyache bush accustomed to chewing seeds of bellyache Legislation poisoning are largely associated with gas- bush when in need of a laxative (IPCS Bellyache bush is a Class 2 pest plant tar- tro-intestinal irritation (IPCS INCHEM INCHEM 2004). The seeds are oily, purga- geted for control under the Queensland 2004, Biehl and Hecker 1986, Adolf et al. tive and emetic (Irvine 1961). Tea made Land Protection (Pest and Stock Route Man- 1984, Burkhill 1994, Parrota 2001). In hu- from bark is used in Nigeria to cure in- agement) Act 2002 (Queensland Govern- mans there is acute abdominal pain and testinal worms (Irvine 1961). The leaves ment 2003). This imposes a legal respon- a burning sensation in the throat about are boiled up and used as a bath for fever sibility for control on all landowners for half an hour after ingesting seeds, fol- and the leaves are used as a purgative in land under their management and it is lowed by nausea, vomiting, and diarrhoea Jamaica (Irvine 1961, de Padua et al. 1999). an offence to keep or sell bellyache bush (Kingsbury 1964, Watt and Breyer-Brand- Roots of bellyache bush have been used without a permit. In the Northern Terri- wijk 1962, Burkhill 1994, Parrota 2001). The for treatment of leprosy (Das and Das tory, bellyache bush is a declared weed un- vomitus and faeces may contain blood. 1994, Baxter 2000). der the Weeds Management Act 2001 and is In severe intoxications, dehydration and Plant parts used for healthcare in In- categorized as Class B (growth and spread haemorrhagic gastroenteritis can occur, as dia include the young stem, root, bark to be controlled) and Class C (not to be well as central nervous system and cardio- and latex (Das and Das 1994). These parts introduced into the Northern Territory) vascular depression and collapse. Children are used either alone or with other com- (Parsons and Cuthbertson 2001). In West- are more susceptible (IPCS INCHEM 2004, ponents for the treatment of abdominal ern Australia, bellyache bush is a declared Kingsbury 1964, Watt and Breyer-Brandw- discomfort, bone fracture, toothache, con- plant under the Agriculture and Related ijk 1962), particularly as the fruit and seeds junctivitis, open wounds, diarrhoea, dys- Resources Protection Act 1976 as a P1 (in- of bellyache bush are attractive to children. entery, haemorrhoids, intra-uterine death, troduction into and movement within the Three seeds can kill a child (Gardner and muscular pain, rheumatism, tongue sores area is prohibited) and P3 (the numbers or Bennetts 1956, Oakes and Butcher 1962, and infections around fi ngernails and toe- distribution or both should be reduced) Watt and Breyer-Brandwijk 1962, Kings- nails (Banerji et al. 1993, de Padua et al. for the municipal districts of Derby-West bury 1964, Begg and Gaskin 1994). Belly- 1999). Crude hot water extract of bellyache Kimberly and Broome (Parsons and Cuth- ache bush sap or latex can also cause acute bush exhibited antimalarial properties. It bertson 2001). dermatitis on contact (Souder 1963). was capable of 100% inhibition of the ma- For Aboriginal people of Australia, laria agent Plasmodium falciparum (Gbeas- Weed management bellyache bush can reduce the availability sor et al. 1989). Individual bellyache bush plants are easy of ‘bush tucker’ and other resources by Extracts of bellyache bush have a rep- to kill with conventional control tech- displacing native plants and animals (King utation as a cancer cure (Biswanth and niques such as herbicides, machinery and Wirf 2005). Bellyache bush can also Ratna 1995, Biswanth et al. 1996, Morton and fi re (Bebawi and Campbell 2002b,c,d, disrupt spiritual and physical connections 1981, Morton 1982, Taylor et al. 1983). For Bebawi et al. 2004). However, large-scale to the country, for example by restrict- example, on the island of Aruba, people seedling recruitment generally occurs fol- ing access to sacred sites (Gardner 2005, believe that a decoction of the stems from lowing the initiation of control activities. King and Wirf 2005). Impenetrable infes- bellyache bush cures throat cancer (Mor- This highlights the need for a long term tations of bellyache bush restrict hunting, ton 1982). Derivatives of the diterpene management strategy to treat seedlings camping and bushwalking activities and jatrophone were also isolated from roots that may emerge while ever a viable seed the general physical movement of people of bellyache bush and shown to have anti- bank remains. A key priority should be (Gardner 2005, King and Wirf 2005). Belly- tumour properties in vitro (Taylor et al. the establishment of a healthy pasture that ache bush is unpleasant to touch because 1983). will compete with bellyache bush seed- it is sticky and leaves permanent reddish- lings and reduce the opportunities for es- brown stains on garments. Other benefi ts Other potential benefi ts of tablishment (Ashley 1995, Csurhes 1999). bellyache bush include its use as a source Benefi cial of oil for energy (Forni-Martins and Cruz Herbicides Medicinal The major benefi ts of belly- 1985, Burkill 1994), a source of plant food Herbicides can cause high mortal- ache bush are associated with its medicinal for human and animal consumption, an ity of bellyache bush when applied using 50 Plant Protection Quarterly Vol.22(2) 2007 hand-help equipment to treat scattered poorly, killing only 63% (Vitelli and Ma- disposal of the cut plant, as cut sections to medium infestations (Chadhokar 1978, digan 2002). Effi cacy dropped from 98 can produce seed. Pitt and Miller 1991, Vitelli et al. 1988). to 42% when bellyache bush growing in The use of residual herbicides to reduce Plants within dense bellyache bush infes- timbered country was aerially treated us- seedling recruitment has been tested. In tations have also been successfully treated ing triclopyr plus picloram (Vitelli and the Northern Territory, high mortality of by aerial application of herbicides (Vitelli Madigan 2002). In trials undertaken by mature plants and control of seedlings for and Madigan 2002). However, irrespective the Northern Territory government, three up to two years was achieved with hex- of the application technique, considerable rates of metsulfuron (50, 75 and 100 g ha-1) azinone and tebuthiuron (Table 1) (Pitt recruitment often occurs subsequently and three rates of glyphosate mixed with and Miller 1991). In a North Queensland (Vitelli and Madigan 2002). simazine (1.5 + 2.0, 1.5 + 4.0 and 3.0 + 4.0 trial, metsulfuron applied between 36 and In Papua New Guinea, foliar applica- kg ha-1) were aerially applied with carrier 1162 g ha-1 and tebuthiuron (500, 1000 and tion of 2,4-D or 2,4,5-T (5–10 g L-1) in water spray volumes of 60 L ha-1 to dense stands 1500 g ha-1) were applied to pots contain- was not effective against bellyache bush of mature bellyache bush. Twelve months ing 50 bellyache bush seeds. One hundred (Chadhokar 1978). Field studies undertak- after application, there were no visible ef- and eighty days post application, seedling en in North Queensland showed that the fects from herbicide application (Pitt and mortality plus seed mortality was record- addition of a 0.2% v/v wetting agent (poly Miller 1991). ed at greater than 80% for each treatment, dimethyl siloxane) increased the activity Cut stump and basal bark techniques compared to control pots with 35% mor- of 2,4-D acid (10 g L-1), 2,4-D amine (5 g can produce good kills of bellyache bush. tality (J.S. Vitelli unpublished results). L-1), and 2,4-D ethyl ester (5 g L-1), result- In Papua New Guinea, a complete kill was In Queensland, permits are required ing in mortality rates of 90, 100 and 90%, obtained when 2,4,5-T (5–10 g L-1) in diesel under the Vegetation Management Act respectively (Vitelli et al. 1988). In the same was applied to stumps cut at ground level 1999 when controlling weeds if off-target trial, amitrole (4 g L-1), 2,4-DP (24 g L-1), (Chadhokar 1978). Similarly, cut stump damage may result in the death of na- 2,4-D/picloram (4.8/1.2 g L-1), fl uroxypyr applications of picloram/2,4,5-T (2.5/10 g tive plants, particularly in riparian areas (4 g L-1), glyphosate (2.8 g L-1), metsulfuron L-1) in water and basal bark applications of (www.legislation.qld.gov.au/LEGISLTN/ (0.12 g L-1) and triclopyr/picloram (4.5/1.5 picloram/2,4,5-T (2/8 g L-1) in diesel killed CURRENT/V/VegetManA99.pdf). The g L-1) killed 50, 73, 97, 100, 100, 100 and 100% of plants (Pitt and Miller 1991). Cut use of gel-applied herbicides such as piclo- 100% of the treated plants, respectively stump trials in North Queensland also ram potassium salt placed onto freshly cut (Vitelli et al. 1988). Similar results occurred found 2,4-D (1.5, 3 and 30 g L-1 water), 2,4- stumps of scattered bellyache bush plants for metsulfuron (98% mortality) in trials in D/picloram (0.5/2, 1/4, and 2/8 g L-1 die- may present a safe option when using the Northern Territory, but fl uroxypyr (0.3, sel), fl uroxypyr (3, 6 and 12 g L-1 diesel), herbicides in riparian zones (M. Ferguson 0.6 and 0.9 kg ha-1) and glyphosate (0.45, triclopyr (4.8, 9.6 and 19.2 g L-1 diesel) and unpublished results). Only two chemicals 0.9 and 1.35 kg ha-1) controlled less than neat diesel killed 90 to 100% of the treated (metsulfuron and fl uroxypyr) are current- 10% of the treated plants (Pitt and Miller bellyache bush plants (Vitelli et al. 1988). ly registered for the control of bellyache 1991). The addition of wetting agents will To be effective the cut stump method re- bush in Australia (Table 2). signifi cantly increase the effectiveness of quires both treatment of the stump and herbicides when applied as foliar sprays (Csurhes 1999, Pitt and Miller 1991). In North Queensland, the aerial appli- cation of herbicides was tested as a po- Table 1. Suppression of mature and seedling bellyache bush compared to tential method for treating large areas of untreated areas 24 months after the application of residual herbicides on a dense infestations of bellyache bush grow- property in the Northern Territory (adapted from Pitt and Miller 1991). ing in non-timbered areas (Vitelli and Ma- Herbicide Rate Plant suppression (%) digan 2002). Using a carrier spray volume -1 (kg a.i. ha ) Seedlings Mature (>50 cm) of 200 litres per hectare, fi ve foliar herbi- cides (triclopyr/picloram at 450/150 g Hexazinone 5.0 78 69 ha-1, glyphosate at 2.16 kg ha-1, fl uroxypyr 10 93 91 at 400 g ha-1, metsulfuron at 72 g ha-1, and 15 94 93 metsulfuron plus glyphosate at 72 + 867 Tebuthiuron 1.0 58 52 g ha-1) killed between 92 to 100% of belly- 1.5 93 89 ache bush plants (Vitelli and Madigan 2002). 2,4-D ester at 4 kg ha-1 performed 2.0 96 94

Table 2. Herbicides registered for control of bellyache bush in Australia (adapted from Infopest Nov 2005, Biosecurity, Department of Primary Industries and Fisheries, Brisbane). Situation Active ingredient Application rate States registered in AustraliaA Agricultural land – non-crop fl uroxypyr as mhe (200 g L-1) 0.5 L 100 L-1 NSW, Q, WA Forests – timber Production fl uroxypyr as mhe (200 g L-1) 0.5 L 100 L-1 NSW, Q, WA Land – commercial/industrial fl uroxypyr as mhe (200 g L-1) 0.5 L 100 L-1 NSW, Q, WA Land – commercial/industrial metsulfuron-methyl (600 g kg-1) 10 g 100 L-1+ penetrant NSW, NT, Q, SA, T, V, WA Pastures fl uroxypyr as mhe (200 g L-1) 0.5 L 100 L-1 NSW, Q, WA Pastures metsulfuron-methyl (600 g kg-1) 10 g 100 L-1+ penetrant Q, WA Pastures – native metsulfuron-methyl (600 g kg-1) 10 g 100 L-1+ penetrant NSW, NT, Q, SA, T, V, WA Rights of way fl uroxypyr as mhe (200 g L-1) 0.5 L 100 L-1 NSW, Q, WA Rights of way metsulfuron-methyl (600 g kg-1) 10 g 100 L-1+ penetrant NSW, NT, Q, SA, T, V, WA A States where registered: NSW = New South Wales, NT = Northern Territory, Q = Queensland, SA = South Australia, T = Tasmania, V = Victoria and WA = Western Australia. Plant Protection Quarterly Vol.22(2) 2007 51 Fire often put the removed material in stacks recruitment is reduced and plants grow Burning is an effective control technique and burn it, so as to kill any reproductive more slowly (F.F Bebawi unpublished re- against bellyache bush where there is material and reduce the risk of plants re- sults). As for most tropical weeds, there suffi cient fuel to carry a fi re (Bebawi and attaching to the soil and growing (S.D. are very few data on how bellyache bush Campbell 2002a,b,c). In the Charters Tow- Campbell unpublished observations). can be managed using grazing or pasture ers region, 76% mortality occurred follow- Cutting bellyache bush close to ground management. ing burning of a riparian infestation in level can also be effective (Bebawi and September 1999. A second burn a year lat- Campbell 2002b). No plants cut off at Natural enemies er (October 2000) increased the mortality ground level regrew, whereas those cut Jatropha spp. generally have few phy- rate to 92% (Bebawi and Campbell 2002c), above ground level reacted differently tophagous insects or pathogens in their with temperatures of up to 640°C record- depending on the season. The majority of native range (Dehgan 1982). In India, ed at ground level (Bebawi and Campbell plants cut at 10, 20, and 40 cm height in the bellyache bush was reported completely 2002c). Exposure of bellyache bush to such dry season regrew, whereas only those cut free of any visible fungal or insect damage high temperatures caused them to ooze at 20 and 40 cm in the wet season regrew. (Raina and Gaikwad 1987). However, in caramelized latex and blister profusely In general, plants were more susceptible Australia, the leaf-mining moth, Epicephala due to the high sugar concentration of the when cut in the wet season (Bebawi and sp. (Wilson 1997) and the castor oil looper, latex (Bebawi and Campbell 2002b). De- Campbell 2002b). Achaea janata L. cause minor defoliation of spite the average fuel load being relatively The use of tractor-mounted slashers bellyache bush. The tenebrionids beetles, high, there was signifi cant variation across has provided similar results in areas of Lyphia australis Blackburn and Platycotylus the site and mortality tended to vary ac- suitable terrain. In one fi eld trial 100% of nitidulus Macleay, and the nitidulid bee- cordingly. Nevertheless, juvenile plants plants were killed, irrespective of whether tles, Carpophilus marginellus Motschulsky were more susceptible to fi re than mature they were juvenile, mature or adult plants and C. obsoletus Erichson, have also been plants, with old plants being the most tol- (Bebawi et al. 2004). There are, however, observed attacking the stems in the North- erant (Bebawi and Campbell 2002c). instances in other areas where not all ern Territory (F.F Bebawi unpublished re- A large portion of the seed bank is able to plants have been killed after being cut off sults). survive fi re, resulting in large scale recruit- at ground level, with re-shooting some- A soil borne fungus Scytalidium dimid- ment (Bebawi and Campbell 2002c). For times occurring (Chadhokar 1978, Pitt and iatum (Penz.) Sutton & Dyko has been ob- example, 540 seedlings m-2 emerged from Miller 1991). served to cause canker and wilt of belly- burnt plots compared with 190 seedlings The use of heavy machinery such as ache bush stems in North Queensland. m-2 in unburnt plots in a North Queens- bulldozers can be effective for directly kill- S. dimidiatum (formerly Torula dimidiata) land trial. Seedling density in burnt plots ing plants, as well as for cleaning up in- is the synanamorph of Nattrassia mangif- averaged 37 seedlings m-2 after two years, fested areas so that other forms of control erae (Syd. & P.Syd.) Sutton & Dyko (syn. compared with four seedlings m-2 in the can be implemented. In one trial, a stick Hendersonula toruloidea). It is a very com- unburnt controls (Bebawi and Campbell rake on the front of a bulldozer caused mon soil-borne fungus associated with a 2002c). Fire could therefore exacerbate the greater than 90% mortality of plants with- wide range of hosts, including crop spe- bellyache bush problem if followup con- in a dense infestation. Those that survived cies (such as mangoes in Queensland) and trol is not implemented. were generally smaller plants that escaped eucalypts, and causing stem end rot (Tom- Fire can only be used when there is a being ripped out (Bebawi et al. 2004). For ley 2003). suffi cient fuel load (Vitelli 2000). In drier mechanical control of woody weeds such Bellyache bush has been a target for areas, burns may only occur following as bellyache bush, effi cacy is generally biological control in Australia since 1996 several years of above average rainfall. highest for treatments that completely re- (Heard and Chan 2002). A total of 170 loca- There are also substantial costs in using move the whole plant or sever the root tions in nine countries (Mexico, Venezuela, fi re, particularly as stock must be excluded system underground (20–30 cm). Season Dominican Republic, Puerto Rico, Nica- before and after a burn, fi re cannot always of application, soil texture, soil moisture ragua, Netherlands Antilles, Guatemala, be used in fi re sensitive areas (Vitelli 2000) and weed density also infl uence survival Trinidad and Cuba) have been investi- and pasture may be lost. and regrowth (Vitelli 2000). gated for potential agents against Jatropha. Flame throwers have been tested for Physical disturbance that occurs when Over 1000 specimens were collected and treatment of bellyache bush where chemi- using mechanical control will generally sent for identifi cation. cal and mechanical control is inappropri- result in massive recruitment of seedlings Of the insects investigated, Pachycoris ate or ineffective (Vitelli and Madigan (Bebawi et al. 2004). This can be a means klugii Burmeister (Scutelleridae) was the 2004). Flaming for 10 seconds around the of depleting the seed bank more quickly, most damaging herbivore of plantations entire circumference of the base of indi- provided regrowth is controlled before it of J. curcas in Nicaragua, but unfortu- vidual bellyache bush plants (5 cm above reaches reproductive maturity (Campbell nately failed to develop on J. gossypiifolia. ground level) at a maximum temperature and Grice 2000). Cylindrocopturus jatrophae Fall (Curculio- of 820°C killed 92% of the treated plants at There are some legislative restrictions nidae), a stem-boring weevil, Colaspis sp. a cost of 7.5 cents per plant (Vitelli and Ma- that may infl uence whether mechanical (Chrysomelidae) and Styloleptus sp. and digan 2004). This practice would be most techniques can be used in certain situa- Parmenonta sp. (Cerambycidae) were appropriate for treating small and/or scat- tions. For example, in Queensland, it is a imported into quarantine but could not tered infestations. requirement to obtain a permit under the be reared. While Lagocheirus sp. (Ceram- Vegetation Management Act 1999 to use ma- bycidae) established in quarantine, host Mechanical control chinery in riparian areas (Vitelli 2000). testing revealed it fed on cassava and was Since bellyache bush has a shallow root rejected. Preliminary studies on the rust system it can be fairly easily removed by Pasture management fungus, Phakopsora jatrophicola Cummins hand, especially if the soil is moist (Csur- Preliminary results from a fi eld trial in- ex Cummins (Uridenales) are promising, hes 1999). A number of landholders with vestigating the impact of fi ve simulated but limited funds have prevented further small patches in the Burdekin region pre- grazing regimes on four bellyache bush host testing. fer to use this practice. By physically re- population densities suggest that belly- To date, the seed feeding jewel bug, Ago- moving the plants they are confi dent that ache bush grows best in areas void of pas- nosoma trilineatum Fabricius (Scutelleridae) plants have been killed. Landholders also ture. Where there is grass cover, seedling is the only agent approved for release in 52 Plant Protection Quarterly Vol.22(2) 2007 Australia against bellyache bush (Figure 11). The jewel bug was fi rst released in the Northern Territory in March 2003 at Wil- leroo Station and in North Queensland in June 2004 at Barkla Station (Heard 2003, Bebawi 2004). The potential impact of jewel bugs on bellyache bush has been quantifi ed in a greenhouse experiment (Bebawi et al. 2005a). Potted bellyache bush plants were exposed to 0, 6 or 24 jewel bugs per plant. Jewel bugs significantly increased the level of abortion of both immature and mature capsules, particularly at the high- er density of insects. Immature capsules were the most susceptible, averaging 80% capsule abortion, compared with only 21% for more mature capsules (Bebawi et al. 2005a). After 30 days exposure to the jewel bug, seed production was reduced by 22% and 75% respectively at low and high jew- el bug density (Bebawi et al. 2005a). Fur- thermore, nearly 60% and 83% of the seeds Figure 11. Male (left) and female (right) jewel bug (Agonosoma trilineatum) produced at low- and high jewel bug den- sity were damaged (Bebawi et al. 2005a). In mating on a bellyache bush capsule. the laboratory, feeding by adult jewel bugs completely destroyed seeds of bellyache bush (Heard and Chan 2002). Table 3. Recruitment of bellyache bush seedlings as affected by singular To date, there have been limited signs (initial treatment, 2003), fi rst follow-up (2004) and second follow-up (2005) of A. trilineatum establishment at three control techniques at Almora Cattle Station, north Queensland. of the nine release sites in Queensland. The most promising signs of establish- Singular (initial 1st followup 2nd followup ment were seen in early 2005, when Control treatment 2003) (2004) (2005) nymphs were observed near one release technique Seedling recruitment for every plant killed site and in early 2006, when both adults Foliar spray 20 6 3 and nymphs were seen at a further two Burn 69 12 9 release sites. Evidence of feeding damage on seed capsules was also noted. It is un- Slash 97 19 11 certain yet whether the insect will persist Stick-rake 74 19 11 at these sites (C.J. Lockett unpublished data).

Integrated control killed (Table 3) (F.F. Bebawi unpublished Jatropha species grown in Nigeria. Quar- No single control method provides effec- results). This means that with two follow- terly Journal of Crude Drug Research 18, tive management of bellyache bush at a ups, seedling recruitment can be reduced 141-5. reasonable cost (Vitelli 2000). Results from up to 85%, making control in subsequent Adolf, W., Opferkuch, H.J. and Hecker, a trial of integrated control methods have years much easier (F.F. Bebawi unpub- E. (1984). Irritant phorbol derivatives shown that high kill rates of bellyache lished results). from four Jatropha species. Phytochemis- bush can be obtained with a single appli- try 23, 129-32. cation of techniques such as foliar spray- Acknowledgments Amin, M.A., Daffala, A.A. and El Moneim, ing, slashing, fi re and stickraking (Bebawi We wish to thank Susan King, Laura Wirf, O.A. (1972). Preliminary report of the et al. 2004). However, followup control will and Ian Miller of the Department of Natu- molluscicidal properties of Habat El- generally be needed for several years to ral Resources, Environment and the Arts Mollok, Jatropha sp. Transactions of the control remaining plants and seedling re- (NT) for their literature contribution to Royal Society of Tropical Medicine Hygiene growth. Selective foliar spraying appears this publication. The technical assistance 66, 805-6. to be one of the most effective options for provided by the Queensland Museum Anon. (1888). List of useful plants grown followup as it allows the maintenance of in insect identifi cation is also gratefully in the Government Botanical Garden at a grass cover to compete with seedlings acknowledged. A special thankyou is ex- Port Darwin. Government Resident’s that emerge afterwards. In the integrat- tended to Tony Grice and Bob Mayer for Report on Northern Territory for the ed control trial, for every plant killed by reviewing the manuscript. Year 1887, pp. 33-40. a single foliar spray treatment, 20 plants Anon (2005). Draft ‘Weed management were recruited from the seed bank, much References plan for bellyache bush (Jatropha gossyp- less than in slashing, mechanical and fi re Abubacker, M.N., Rao, G.R. and Kumare- iifolia) in the Northern Territory (2005– treatments which averaged 97, 74 and 69 san, A. (1999). Sugarcane press mud 2008)’. (Department of Infrastructure, plants, respectively (Bebawi et al. 2004). cake: accelerator of biogas production Planning and Environment, Northern Furthermore, followup foliar treatments in various weed biomass. Advances in Territory, Australia). implemented one and two years later re- Plant Sciences 12, 73-8. APB Infonote (1994). Agriculture Protec- sulted in only six and three new plants be- Adewunmi, C.O. and Marquis, V.O. (1980). tion Board of Western Australia. In- ing recruited respectively, for every plant Molluscicidal evaluation of some fonote 11/94 AGDEX 647. Plant Protection Quarterly Vol.22(2) 2007 53 Ashley, M. (1995). The biology and man- Bebawi, F.F., Davis, K. and Lockett, C. Association of Societies for Growing agement of Jatropha gossypiifolia. Tech- (2005a). Bellyache bush (Jatropha gos- Australian Plants (ASGAP), http://far- nical Bulletin No. 234. (Department of sypiifolia) biocontrol potential – pre- rer.riv.csu.edu.au. Primary Industry and Fisheries, North- liminary results. Proceedings of the 8th Campbell, S.D. and Grice, A.C. (2000). ern Territory). Queensland Weed Symposium, Weed Weed biology: a foundation for weed Backer, C.A. and Bakhuizen van den Management – Making a Difference, ed. management. Tropical Grasslands 34, Brink, R.C. (1963). ‘Flora of Java. Vol- W.D. Vogler, pp. 78. (Weed Society of 271-9. ume 1’. (N.V.P. Noordhoff-Groningen, Queensland, Brisbane, Queensland). 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Livingstone, Gunasekara, S.P., Bevelle, C.A., Cord- Ltd., London). ell, G.A., Farnsworth, N.R., Kupchan, Wheeler, J.R. (1992). ‘Flora of the Kimber- S.M., Uchida, H., Branfman, A.R., Dai- ley Region’. (Department of Conserva- ley, R.G. and Sneden, A.T. (1983). New tion and Land Management, Como, anti leukemic derivatives from Jatropha Western Australia). gossypiifolia: structural and stereochem- Wild, K.L. (2003). Understanding the ical assignment through nuclear mag- flowering and seeding behaviour of netic resonance spectroscopy. Journal bellyache bush (Jatropha gossypiifolia). 56 Plant Protection Quarterly Vol.22(2) 2007 Appendix 1. Summary of insects, vertebrates, and fungi associated with Jatropha gossypiifolia and locations where collected. All Australian collections were made in Queensland. Source of overseas insects was obtained from Reddi and Reddi (1983) and CSIRO (1998). Source of Queensland insects was identifi ed by Mr. Geoff Monteith, Curator Entomology, Queensland Museum, Brisbane. Family Scientifi c Name Common name Country Insects Acrididae Austracris sp. Spur-throated locust Australia Acrida conica Fabricius Giant green slantface grasshopper Australia Chortoicetes terminifera Walker Australian plague locust Australia Phaulacridium vittatum Sjostedt Wingless grasshopper Australia Alydidae Daclera rufescens Stal. Bug Australia Melanacanthus sp. Brown bean bug Australia Apidae Apis cerana indica Fabricius Asian honey bee India Apis fl orae Fabricius Dwarf honey bee India Apis mellifera Lepeltier Honey bee Trigona carbonaria Smith Native bee Trigona sp. India Bombyliidae Eucahrimia sp. India Geron sp. Bee fl ies India Bruchidae Algarobius bottimeri Kingsolver Mesquite bruchid beetle Australia Buprestidae Chalcophorotaenia australasiae Saunders Jewel beetle Australia Conognatha sp. Australia Calliphoridae Rhyncomya viridaurea Wiedemann Fly India Cerambycidae Lagocheirus araneiformis Linnaeus Stem boring beetle Venezuela Lagocheirus sp. Stem boring beetle Mexico, Venezuela Leptostylus sp. Venezuela Lepturges sp. Venezuela Ozineus sp. Beetle Venezuela Parmenonta sp. Mexico Styloleptus laticollis Drury Broad-necked root borer Dominican Republic Trachyderes mandibularis Audinet-Serville Long-horned beetle Mexico Chrysomelidae Altica sp. Metallic green fl ea beetle Mexico Colaspis sp. Leaf beetle Venezuela Cryptocephalus sp. Cylindrical leaf beetle Venezuela Desmogramma sp. Beetle Venezuela Monomacra sp. Orange beetle Mexico Monoxia sp. Venezuela Paria sp. Sap feeding beetle Venezuela Synbrotica sp. Mexico Coccinellidae Coccinella transversalis Fabricius Transverse ladybird Australia Coccinellidae sp. Mexico Coenagrionidae Pseudagrion aureofrons Tillyard Goldfront sprite damselfl y Australia Coreidae Amorbus alternatus Dallas Squash bug Australia Amorbus obscuricornis Westwood Gumtree bug Australia Mictis profana Fabricius Crusader bug Australia Cosmopterygida Cosmopterygidae sp. Leaf miner Venezuela Eumenidae Rhynchium metallicum Saussure Wasp India Flatidae Siphantha acuta Walker Green plant hopper Australia Formicidae Calomyrmex similis Mayr Beauty ants Australia Camponotus novaehollandiae Northern sugar ants Australia Camponotus sp. Sugar ants Australia Crematogaster sp. Cocktail ants India Iridomyrmex anceps Roger Tropical tyrant ants Australia Iridomyrmex spadius Schattuck Meat ant Australia Iridomyrmex sp. Tyrant ants Australia Paratrechina sp. Parrot ants Australia continued/… Plant Protection Quarterly Vol.22(2) 2007 57

Family Scientifi c Name Common name Country Insects continued/… Geometridae Geometridae sp. Moth Venezuela Gracillariidae Epicephala gracillariidae Leaf miner, moth Australia Epicephala sp. Moth Australia Halictidae Pseudapis oxybeloides Smith Bee India Hemiptera Auchenorrhyncha sp. Leaf hopper Mexico Histeridae Histeridae sp. Clown beetles, Hister beetles Mexico Lampyridae Lampyridae sp. Fire fl ies Mexico Lepidoptera Gelechioidea sp. Moth Mexico Papilionoidae sp. Venezuela Libellulidae Diplacodes haematodes Burmeister Common percher dragonfl y Australia Orthetrum caledonicum Brauer Blue skimmer dragonfl y Australia Lycaenidae Nacaduba biocellata Felder & Felder Two-spotted line blue butterfl y Australia Lycarnidae Zizina labradus Godart Common grass blue butterfl y Australia Mantidae Austrovates sp. Blackbarrel mantid Australia Tenodera australasiae Leach Purple-winged mantid Australia Meloidae Meloidae sp. Blister beetles Mexico Milichidae Milichiella sp. Fly India Myrmicinae Myrmecia pilosula Smith Jumping jacks Australia Nitidulidae Carpophilus marginellus Motschulsky Australia Carpophilus obsoletus Erichson Australia Noctuidae Achaea janata Linnaeus Castor oil looper Australia Bulia confi rmans Walker Venezuela Chrysodeixis argentifera Guenee Tobacco looper butterfl y Australia Ischyja neocherina Butler Sugarcane moth looper Australia Metaponpneumata rogenhoferi Moschler Venezuela Ponometia exigua Fabricius Moth Venezuela Spodoptera latifascia Walker Moth Mexico Nyphalidae Euploea core-corinna Cramer Common crow butterfl y Australia Pieridae Belonis java Linnaeus Caper white butterfl y Australia Catopsilia crocale Cramer Butterfl y India Catopsilia pomona Fabricius Lemon migrant butterfl y Australia Eurema hecabe Linnaeus Common grass yellow butterfl y Australia Pompilidae Ctenostegus sp. Spider wasp Australia Ponerinae Odontomachus turneri Forel Giant snappy ants Australia Rhytidoponera convexa Mayr Convex pony ants Australia Rhytidoponera metallica Smith Metallic pony ants Australia Pyralidae Herpetogramma licarsisalis Walker Grass webworm butterfl y Australia Reduviidae Apiomerus vexillarius Champion Venezuela Gminatus Australia Stal Orange assassin bug Australia Poccilobdallus formosus Stal Assassin bug Australia Poecilosphadrus sp. Assassin bug Australia Pristhesancus plagipennis Walker Common assassin bug Australia Sarcophagidae Sarcophaga orchidea Asahina Housefl y India Sarcophaga sp. India Scarabaeidae Glycyphana sp. Harvest beetle Australia Protaetia sp. Beetle Australia Scoliidae Campsomeris radula Fabricius Wasp Australia Scolia cruenta Klug Wasp India Scolia sp. Wasp Australia Scutelleridae Agonosoma trilineatum Fabricius Jewel bug, sucking bug Venezuela Agonosoma sp. Venezuela Coleotichus artensis Montroonzier Bug Australia Coleotichus costatus Fabricius Shield bug Australia Lampromicra senator Fabricius Australia continued/… 58 Plant Protection Quarterly Vol.22(2) 2007

Family Scientifi c Name Common name Country Insects continued/… Sphecidae Sphex cognatus Smith Sand wasp Australia Sphingidae Hyles lineata Fabricius White-lined sphinx moth Australia Stratiomyidae Exaireta spinigera Weidemann Garden soldier fl y Australia Tenebrionidae Blapstinus sp. Darkling beetle Venezuela Branchus opatroides Champion Mexico Epitragus sp. Defoliator Venezuela Hylocrinus sp. Mexico Lyphia australis Blackburn Australia Phegoneus sp. Mexico Platycotylus nitidus Macleay Australia Xystropus sp. Venezuela Tephritidae Dacus tryoni Froggatt Queensland fruitfl y Australia Teragnathidae Nephila edulis Koch Golden orb web spider Australia Theridiidae Latrodectus hasseltii Thorell Redback spider Australia Thyreocoridae Galgupha sp. Venezuela Tineidae Tineidae sp. Moth Venezuela Xylesthia sp. Stem mining moth Mexico, Venezuela Tortricidae Amorbia sp. Totricid moth Mexico Platynota rostrana Walker Peel-feeding totricid caterpillar Mexico Platynota sp. Leafrollers caterpillars Mexico Vespidae Epiodynerus nigrocinctus Saussure Potter wasp Australia Ropalidia romandi Le Guillon Australian paper wasp Australia Ropalidia spatulate Van der Vecht Wasp India Xylocopidae Xylocopa latipes Drury Carpenter bee India Fungi Uredinales Phakopsora jatrophicola Cummins Rust fungus Mexico Scytalidium dimidiatum Sutton & Dyko Soil-borne fungus Australia