The introduction ofMimorista pulchellalis [Lepidoptera: Pyraustidae] into South Africa for the biological control of jointed cactus,Opuntia aurantiaca 1. Biology and mass-rearing techniques

A laboratory investigation was conducted on a pyraustid moth imported from South America for jointed cactus control. It was found that the moths were crepuscular, with females laying an average of 48±3.5 eggs over a period of 4–8 days. Larvae hatched out at night and usually tunnelled into the terminal ends of cactus cladodes at a site where young cactus growth occurred. The lifecycle was completed inside the cactus, pupation typically occurring in the dry husk of the cladode on which the egg was laid. Mass-rearing of the moth was easy if plants with young cactus growth were used.      

The introduction ofMimorista pulchellalis [Lepidoptera: Pyraustidae] into South Africa for the biological control of jointed cactus,Opuntia aurantiaca. 2. Field evaluation

The mothMimorista pulchellalis was monitored over 2 years after liberation in a jointed cactus (Opuntia aurantiaca) infestation in South Africa. Moth and cactus densities were estimated using a system of randomly-assigned quadrats and the impact of the moth on the cactus population quantified. Moths appeared adapted to survive on the etiolated form of jointed cactus plants, killing approximately 1% of the increment in small plants annually. Large plants were also attacked but damage was negligible. The moths occurred in low numbers throughout the study period and generally went through 3 generations in a year.

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Résumé La pyraleMimorista pulchellalis a été surveillée durant les deux années qui suivirent sa libération dans une infestation de figuier de Barbarie en Afrique du Sud. Les densités de pyrale et de cactus ont été estimées en utilisant un système de quadrats distribués au hasard et l'impact de la pyrale sur la population de cactus a été quantifiée. Les pyrales semblaient adaptées à survivre sur la forme étiolée des plants de cactus tuant approximativement 1% de l'accroissement annuel des petits plants. Les gros plants étaient aussi attaqués, mais les dégats étaient négligeables. La pyrale existait en faibles nombres durant toute la période d'étude et elle présentait généralement 3 générations par an.

     

Trichilogaster sp. [Hymenoptera: Pteromalidae], a potential biocontrol agent for the weedAcacia pycnantha [Fabaceae]

The gall wasp,Trichilogaster sp., was imported from Australia to assess its potential as an agent for the control of the invasive shrub/treeAcacia pycnantha Benth. in South Africa. Host specificity tests indicate safety for release; of 19 tree/shrub species tested, including 16 species closely related toA. pycnantha, galls developed only onA. pycnantha. However, galling intensity remained consistently low on the host plant; only 21–29% of the branches exposed to the wasp were galled during 3 years of rearing. Neither the prolonged presence of males in test cages (someTrichilogaster species are thelytokous) nor the stage of maturity of reproductive buds exposed to oviposition affected the percentage of branches galled. It is not recommended thatTrichilogaster sp. be released before the possibility of insect-plant homeostasis or mis-matching of wasp and host plant populations/strains/subspecies is investigated, especially since galling intensities of 30% were ineffective in reducing seed production of a relatedTrichilogaster species/Acacia association.      

Stamen movements in flowers ofOpuntia (Cactaceae) favour oligolectic pollinators

Opuntia brunneogemmia andO. viridirubra occur sympatrically in the Serra do Sudeste, Rio Grande do Sul, Brazil. Their flowers have 450–600 thigmonastic stamens and provide large amounts of pollen and nectar for bees. Bees of 41 species were registered at the flowers ofO. brunneogemmia and 30 at the flowers ofO. viridirubra. Females of three oligolectic species are the only effective pollinators:Ptilothrix fructifera (Anthophoridae),Lithurgus rufiventris (Megachilidae), andCephalocolletes rugata (Colletidae). During their visits inOpuntia-flowers, bees touch the filaments and stimulate the movement of the stamens to the centre of the flower. At the end of this movement, the anthers are densely packed around the style. As a consequence the pollen is presented in an easily accessible upper layer of anthers and various, nearly inaccessible lower layers. The lower layers contain about 80% of the pollen reward. Only females of the three oligolectic pollinators exploit the pollen from the lower layers and reach the nectar furrow. Therefore, through their stamen movements,Opuntia flowers hide most of their pollen from flower visitors but favour effectively pollinating, oligolectic bees.     

ACactoblastis [Lep.: Phycitidae] for the biological control ofEriocereus martinii [Cactaceae] in Australia

ACactoblastis sp. was found on Harrisia cactus (Eriocereus martinii Lab.) in north-eastern Argentina. In biology and life history it is similar toCactoblastis cactorum (Berg.) though attacking different cacti. Laboratory tests and field observations demonstrate that it will only damage plants in the sub-tribeCereanae of theCactaceae. It was approved for liberation in Queensland, Australia in August 1975 for the control ofEriocereus spp., and field releases were made in October 1978.     

Pollination ofOpuntia lindheimeri and related species

The large, yellow, bowl-shaped flowers ofOpuntia lindheimeri, O. discata, O. phaeacantha major, andO. compressa in Texas are visited by various species of beetles and bees. The beetles and small bees (Perdita, Dialictus) are pollen thieves. The pollinators are the medium-sized and larger bees (Melissodes, Diadasia, Lithurge, Megachile, Agapostemon, etc.). Different species of theOpuntia lindheimeri group have the same pollination system and there is no evidence of any floral isolation between them. The pollination system of these species ofOpuntia in Texas is essentially the same as that ofEchinocereus fasciculatus andFerocactus wislizeni in Arizona.Pollination of North American Cacti, II.—SeeGrant & Grant (1979).     

The biology and host specificity of the stem-boring weevilListronotus setosipennis [Col.: Curculionidae] A potential biocontrol agent forParthenium hysterophorus [Asteraceae]

The stem-boring weevilListronotus setosipennis (Hustache) is widespread and damaging to its hostParthenium hysterophorus L. in northern Argentina and southern Brazil. In detailed host-testing it was shown to have a restricted host-range and, despite some feeding and development on sunflower in tests, to be a safe biological control agent againstP. hysterophorus. Field releases in Queensland, Australia took place from 1982 to 1986 and the weevil has established over several thousand hectares at numerous sites. Spread is however slow and the effect on the plant still negligible.      

Investigations on the parasitism ofOtiorrhynchus salicicola andO. Sulcatus [Col.: Curculionidae] byHeterorhabditis sp. [Nematoda]

Experiments were conducted to study the efficacy of the insect parasitic nematodeHeterorhabditis sp. (HW79) as a biological control agent ofOtiorrhynchus salicicola. This weevil species is reported as a pest of ornamental plants in Switzerland and Italy. Dipping plastic boxes containing heavily infested cuttings of laurel (Prunus laurocerasus) in a nematode suspension resulted in approximately 100% parasitisation of full-grown larvae, pupae and non-emerged young adults. The average dose resulting from dipping varied between 56,000 and 62,000 nematodes per liter soil. This experiment was run under natural outdoor conditions. In a further outdoor experiment, pottedLigustrum plants were inoculated with eggs ofO. salicicola and later 20,000 infective juvenile nematodes per liter soil were added to the soil surface. The resulting weevil mortality in the treated pots was 78%. In seven greenhouse tests using the same nematode dose in pots with horticultural soil to which weevil larvae had been added, weevil mortality varied between 76% and 100%, the arithmetic average being 90%. These results indicate that Heterorhabditid nematodes may provide an effective means of controllingO. salicicola. In an other experiment usingO. sulcatus larvae, the influence of application time on nematode efficacy was investigated. When nematodes were added a few days before weevil larvae had hatched from the eggs, no parasitic effect was obtained. Nematode applications done shortly after larval hatching however, resulted in complete weevil control. These results are of significance in timing nematode applications in practice.      

Pre-release impact assessment of two stem-boring weevils proposed as biological control agents for Alliaria petiolata

Experimental studies can be useful tools to test plant responses to herbivory and to quantify the impact of potential biological control agents prior to their release. We evaluated the per-capita effect of Ceutorhynchus alliariae and C. roberti, two stem-boring weevils currently being investigated as potential biological control agents for garlic mustard, Alliaria petiolata, in North America. Weevils were released at three different densities in individual and mixed-species treatments onto potted plants of A. petiolata. Damage by C. roberti alone and by both weevils combined caused an increase in the numbers of inflorescences produced per plant. Although plants could compensate for low levels of damage, moderate to high levels of damage by both C. alliariae and C. roberti, individually and in combination, caused a decrease in plant height and a reduction in seed output per plant. The damage inflicted by both weevil species is similar so the overall impact of both species combined can be predicted by summing the impact of each species alone. Provided they are sufficiently host specific, both weevils could be released as biocontrol agents. Because reduced seed production is necessary to suppress A. petiolata populations, both species have the potential to contribute to control of A. petiolata in North America.     

Chamaesphecia doryliformis [Lep.: Sesiidae], a second root borer for the control ofRumex spp. [Polygonaceae] in Australia

The host-specificity and biology ofChamaesphecia doryliformis (Ochsenheimer) [Lep.: Sesiidae] are described and the insect is assessed for its potential as a biological control agent for weeds of the genusRumex (Polygonaceae) in Australia. The insect is found in the western Mediterranean region, chiefly in north Africa. Adults emerge from pupation in late spring to summer when they lay eggs. The larvae feed inside the roots of post-reproductive plants belonging to the subgeneraRumex andAcetosa. In host-specificity tests 1^st instar larvae attacked only plants of thePolygonaceae. The insect was judged safe to release in Australia after assessing its level of host-specificity, and attributes of its biology which indicate that native AustralianPolygonaceae will not be endangered.      

Weed biological control: applying science to solve seemingly intractable problems

Abstract   Exotic weeds pose a problem of considerable economic and environmental importance to Australia. As a consequence, Australia has developed into a leading centre of research on weed biological control, with over 60 weeds the targets of past and current projects. Using primarily entomological examples, this review highlights the contributions made by Australian scientists to the development of theory and the improvement of practice in weed biological control. It also shows how biological control practitioners have made use of, and contributed to, broader theory and knowledge of plant-herbivore relationships. Finally, it concludes with some reflections on the future direction of biological control in Australia.     

Pollination biology ofOpuntia polyacantha andOpuntia phaeacantha (Cactaceae) in southern Colorado

The phenology, compatibility system, and pollinator fauna ofOpuntia polyacantha Haw. andO. phaeacantha Engelm. in southern Colorado were studied and compared. The total blooming periods overlap, but the peak of blooming differs between the species withO. polyacantha blooming first and for longer. Neither species is apomictic,O. polyacantha is largely self-incompatible andO. phaeacantha is self-compatible but may not automatically fully self-pollinate. Examination of flowering structures showed that the former has larger flowers with more perianth parts and heavier (but fewer) anthers. Further the flowers of the allogamous species are visited by greater diversity (but similar abundance) of pollinators. Medium to large bees of the generaDiadasia, Lithurge, Melissodes, Bombus, Agapostemon andMegachile were found to be effective pollinators of the Opuntias studied. The two species were compared as to their floral attributes, breeding systems, and pollinators, and the possible role of competition for pollinators is discussed with respect to its role in their evolutionary paths.     

Pre-release evaluation of Heikertingerella sp. as a potential biocontrol agent for Tecoma stans in South Africa

Native to Central America, Tecoma stans (L.) Juss ex Kunth var. stans (Bignoniaceae) is a small tree that is invasive in South Africa and neighbouring countries. The plant was targeted for biological control in South Africa in 2003, with two insect agents released and established so far. The root-feeding flea beetle, Heikertingerella sp. (Coleoptera: Galerucinae: Alticini), was imported from Mexico as an additional biocontrol agent and its biology and host specificity was assessed under quarantine conditions. The beetle displayed a generation time (i.e. from adult to adult) of 49 to 67 days, ensuring four annual generations under laboratory conditions. The beetle's larval and adult stages inflicted high levels of damage on the root system and the leaves of T. stans, respectively. No-choice tests with 40 test-plant species revealed adult feeding on only two non-target species, Tecoma × alata and T. capensis (Thunb.) Spach, with feeding four times higher on T. stans. Larvae developed to adulthood on T. stans only. Multi-choice tests involving the three Tecoma species confirmed these trends, demonstrating that Heikertingerella sp. is host specific. Since T. × alata is a hybrid of T. stans with invasive tendencies, any unlikely attacks by Heikertingerella sp. would be inconsequential in South Africa. The native T. capensis, which suffered little leaf damage and produced no F1 adults, is also at minimal risk of attack. We conclude that Heikertingerella sp. is a suitable biocontrol agent for T. stans and that permission for its release in South Africa be sought.     

Parasitoids acquired byColeophora parthenica [Lepidoptera: Coleophoridae] ten years after its introduction into southern California for the biological control of Russian thistle

Parasitism of the stem and branch-boring mothColeophora parthenica Meyrick [Lepidoptera: Coleophoridae], introduced into California for the biological control of Russian thistle,Salsola australis R. Brown [Chenopodiaceae] was studied in the Coachella Valley of southern California during 1985–1986. Eight parasitoid species were reared from overwintering larvae ofC. parthenica, but none from the F₁ larvae, and just 2 individuals of one species from the F₂ summer generation. The level of parasitism of overwintering larvae was positively correlated with branch diameter, and ranged from 2% in the primary (main) branches to 45% in the tertiary branches in the spring 1985 sample, and from 2% to 19% in the spring 1986 sample, respectively. Rates of parasitism>20% were only found at sites with higher plant cover and chenopod diversity, but no other plant source or alternate hosts of the parasitoids ofC. parthenica were found. The 2 dominant species, the solitary, hymenopterous ectoparasitoids,Norbanus perplexus (Ashmead) [Pteromalidae] andEurytoma strigosa Bugbee [Eurytomidae], are both congeners of native parasitoids ofC. parthenica in Pakistan. The 2 other species of parasitoids ofC. parthenica in southern California for which other hosts are known are polyphagous and external on the larvae. No specialized endoparasitoid Braconidae, like those which dominate the native parasitoid complex in Pakistan and the U.S.S.R., have transferred toC. parthenica during its first 10 years in southern California.      

Host-specificity of fiveAnacassis species [Col.: Chrysomelidae] introduced into australia for the biological control ofBaccharis halimifolia [Compositae]

Foliage-feeding beetles of the genusAnacassis [A. phaeopoda Buzzi,A. fuscata (Klug),A. fuscata var.unicolor (Burmeister),A. cribrum (Klug) andA. dubia (Boheman)] were collected fromBaccharis spp. andBaccharidastrum spp. in South America. Specificity studies showed that they were restricted to these 2 host genera. Between 1974 and 1976 these species were introduced into Australia for the control ofBaccharis halimifolia. A. phaeopoda andA. fuscata were first released in 1976. One field colony ofA. fuscata persisted for up to 3 years but no recoveries ofA. phaeopoda were made after the 1st field generation. The other species died out in quarantine and were not released.      

Host specificity and biology ofLioplacis elliptica [Col.: Chrysomelidae] introduced into Australia for the biological control ofBaccharis halimifolia [Compositae]

The nocturnal foliage feeding ChrysomelidLioplacis elliptica Stal occurs onBaccharis gaudichaudiana DC. in Brazil. Studies of its biology and host plant specificity showed it was restricted tobaccharis spp. It was introduced into Australia for the control ofB. halimifolia L. in 1976 and first released in 1977. Field colonies persisted for up to 3 years after which no further evidence of survival was found.      

Host-specificity and biology ofMetallactus patagonicus [Col.: Chrysomelidae] introduced into Australia for the biological control ofBaccharis halimifolia [Compositae]

Biology and host-specificity of the foliage-feeding chrysomelidMetallactus patagonicus Suffrian were studied in Brazil to determine its suitability for introduction into Australia to control the shrubBaccharis halimifolia L. Multiple-choice host preference testing of plants related toBaccharis and of desirable plants from a range of plant families, showed thatM. patagonicus was restricted toBaccharis spp. It was released in Australia in 1975, but did not establish.      

Host-specificity of a root borer,Bembecia chrysidiformis [Lep.: Sesiidae], a potential control agent forRumex spp. [Polygonaceae] in Australia

Bembecia chrysidiformis (Esper) [Lep.: Sesiidae] was examined for its natural history and specificity toRumex spp. (Polygonaceae) which are weeds in Australia. Adults of this southern European insect appear in late spring to summer. Eggs are laid on the dried, seed bearing stems of perennialRumex plants. The larvae tunnel inside the root during summer through to the next spring. In nature, the larvae are round inRumex species of the subgeneraRumex andAcetosa. In host-specificity tests with 1st instar larvae, the roots of a number of genera within thePolygonaceae were attacked. Larvae died on a range of plants from other families except inPersea americana Miller(Lauraceae), Helianthemum nummularium (L.) Miller (Cistaceae) andQuercus ilex L. (Fagaceae) where larvae fed on the stems. The insect was judged safe for release in Australia by assessing aspects of its biology, its known host plants, and the lack of reported attack on other plants.      

A leaf feeding moth,Euclasta whalleyi [Lep.: Pyralidae] for the biological control ofCryptostegia grandiflora [Asclepiadaceae] in Queensland,Australia

In Madagascar, the leaf feeding Pyralid,Euclasta whalleyi Popescu-Gorj & Constantinescu, occurs onCryptostegia grandiflora R. Br. and other genera in the subfamilyPeriplocoideae of the familyAsclepiadaceae. Studies on its biology and host plant specificity have shown it is restricted to this subfamily. The Australian native,Gymnanthera nitida R. Br., also belongs to this sub-family and is host to an Australian native moth,Euclasta maceratalis Lederer. Despite the likelihood ofE. whalleyi feeding onG. nitida and competing withE. maceratalis, the moth was approved for release in Australia in October 1987. During 1988 over 22,000 larvae were released; field establishment is not yet proven.      

Host-specificity and virus-vector potential ofAphis chloris [Hemiptera: Aphididae], a biological control agent for St John's wort in Australia

Under laboratory conditionsAphis chloris Koch has been demonstrated to be specific to plants belonging to the genusHypericum. It can effect severe damage to its principal host,H. perforatum L., and shows good potential for contributing to the control of this noxious weed. Other species ofHypericum are less favoured hosts ofA. chloris and would not be endangered by it. A. chloris shows a high level of host-discrimination and does not transmit persistent viruses between non-host plants. Whereas in the laboratory it is capable of transmitting non-persistent viruses, it would contribute only marginally to the risk of virus transfer posed by the Australian aphid fauna as a whole, and its release would not necessitate changes to existing control practices, where these are required to reduce plant virus transmission. As a consequence,A. chloris is considered safe for release againstH. perforatum in Australia.