Prioritisation of potential agents for the biological control of the invasive alien weed,Pereskia aculeata (Cactaceae), in South Africa

Pereskia aculeata Miller (Cactaceae) is an invasive alien species in South Africa that is native in Central and South America. In South Africa, P. aculeata outcompetes native plant species leading to a reduction in biodiversity at infested sites. Herbicidal and mechanical control of the plant is ineffective and unsustainable, so biological control is considered the only potential solution. Climatic matching and genotype matching indicated that the most appropriate regions in which to collect biological control agents were Santa Catarina and Rio de Janeiro provinces in Southern Brazil. Surveys throughout the native distribution resulted in 15 natural enemy species that were associated with the plant. Field host range data, as well as previous host plant records, were used to prioritise which of the species were most likely to be suitably host specific for release in South Africa. The mode of damage was used to determine which species were most likely to be damaging and effective if released. The most promising species prioritised for further study, including host specificity and impact studies, were the stem-wilter Catorhintha schaffneri Brailovsky & Garcia (Coreidae); the stem boring species Acanthodoxus machacalis Martins & Monné (Cerambycidae), Cryptorhynchus sp. (Curculionidae) and Maracayia chlorisalis (Walker) (Crambidae) and the fruit galler Asphondylia sp. (Cecidomyiidae). By prioritising the potential biological control agents that are most likely to be host-specific and damaging, the risk of conducting host specificity testing on unsuitable or ineffective biological control agents is reduced.     

Interactions between two biological control agents and their target weed: a beetle,a bug and a cactus weed

ABSTRACT

Pereskia aculeata Miller (Cactaceae) is an invasive alien shrub introduced into South Africa from Brazil. The leaf-feeding beetle, Phenrica guerini Bechyne (Chrysomelidae), was released as a biological control agent in South Africa in 1991 followed by the stem-wilting bug, Catorhintha schaffneri Brailovsky & Garcia (Coreidae), in 2014. This study investigated the interactions between the two agents under laboratory conditions. Potted plants were exposed to one of four treatments: control (no agents), P. guerini only, C. schaffneri only and both species together. Four densities, ranging from 2 to 12 insects per plant were used. Cathorhitha schaffneri alone at low to moderate densities resulted in the same reduction in number of leaves and shoot length as when combine with P. guerini. At the highest density, C. schaffneri reduced the number of leaves significantly more than any treatment. Mortality of P. guerini was significantly higher than C. schaffneri at the highest density when in combination. The antagonistic interaction between P. guerini and C. schaffneri suggests that these agents should not be released together because this would impact negatively on the overall biocontrol programme against P. aculeata. It is recommended that C. schaffneri should be released at sites where P. guerini is not present. Extrapolation of laboratory-based studies into the field is often challenging, so mass-rearing and releases of P. guerini should continue until there is convincing proof that C. schaffneri alone is more effective than P. guerini in the field.     

Pereskiophaga brasiliensis,a natural enemy of the invasive alien cactus Pereskia aculeata,is not suitably host specific for biological control in South Africa

The stem-mining weevil, Pereskiophaga brasiliensis, was a candidate biological control agent for the invasive cactus Pereskia aculeata in South Africa. In host specificity trials, it developed on two indigenous test plant species under choice and no-choice conditions. Pereskiophaga brasiliensis is therefore not suitably host specific for release in South Africa.     

The effect of host plant intraspecific genetic variation on the fitness of a monophagous biological control agent

Variation in insect life-history traits when developing on different host plant genotypes with known genetic relationships was tested using the monophagous biological control agent Phenrica guérini Bechyné (Chrysomelidae) and its host plant, Pereskia aculeata Miller (Cactaceae). Differences in insect fitness among tested host plant genotypes were expected because of geographic isolation and large genetic distances between plant genotypes, and because P. guérini has been associated with some of the plant accessions in the field while others have never been exposed to herbivory by the insect. There was little variation in insect life-history traits among plant genotypes and no differences in the insect's ability to utilise different plant genotypes. The results of the study suggest that, in some cases, biological control agents collected from the native genotype most closely related to the target weed population will not always be more effective than those collected from more distantly related genotypes.     

Basal cactus phylogeny: implications of Pereskia (Cactaceae) paraphyly for the transition to the cactus life form

The cacti are well-known desert plants, widely recognized by their specialized growth form and essentially leafless condition. Pereskia, a group of 17 species with regular leaf development and function, is generally viewed as representing the "ancestral cactus," although its placement within Cactaceae has remained uncertain. Here we present a new hypothesis of phylogenetic relationships at the base of the Cactaceae, inferred from DNA sequence data from five gene regions representing all three plant genomes. Our data support a basal split in Cactaceae between a clade of eight Pereskia species, centered around the Caribbean basin, and all other cacti. Two other Pereskia clades, distributed mostly in the southern half of South America, are part of a major clade comprising Maihuenia plus Cactoideae, and Opuntioideae. This result highlights several events in the early evolution of the cacti. First, during the transition to stem-based photosynthesis, the evolution of stem stomata and delayed bark formation preceded the evolution of the stem cortex into a specialized photosynthetic tissue system. Second, the basal split in cacti separates a northern from an initially southern cactus clade, and the major cactus lineages probably originated in southern or west-central South America.     

Pathogenicity and Host Specificity of Mycovellosiella lantanae var. lantanae,a Potential Biocontrol Agent for Lantana camara in South Africa

Lantana camara is a poisonous, bushy shrub from South and Central America that has invaded much of the moist, warm sub-tropical areas of South Africa. In the last decade, evidence of a conspicuous and damaging mycobiota on lantana in the Neotropics has persuaded biocontrol researchers to consider fungi as potential biocontrol agents for this plant. Samples of diseased L. camara leaves were collected during field surveys in South, North and Central America from 1987 to 1997. Pathogenicity tests showed the leaf spot fungus, Mycovellosiella lantanae var. lantanae to be a promising biocontrol pathogen. Inoculations onto South African biotypes of L. camara and a number of closely related species within the family Verbenaceae were made under quarantine conditions to determine the pathogenicity and host specificity of the fungus. Inoculation of L. camara biotypes resulted in necrotic, grey lesions (20-60 per leaf), necrosis of flower buds and stalks, as well as defoliation of certain biotypes after three weeks. None of the other plant species tested showed any disease symptoms. This indicates a very restricted host range, making this pathogen a suitable candidate for application as a biological control agent. It is expected that the fungus will reduce the vigour and reproductive potential of L. camara. Due to the variation in virulence of M. lantanae var. lantanae when tested on South African lantana biotypes, release strategies may include the use of combinations of the isolates to target a wider range of lantana biotypes in the field. Permission to release M. lantanae var. lantanae in South Africa was granted in September 2001. The pathogen will be released as a classical biocontrol agent in South Africa.     

The tortoise beetle Physonota maculiventris (Chrysomelidae: Cassidinae) is suitable for release against the weedy Mexican sunflower Tithonia diversifolia (Asteraceae) in South Africa

The biology and host range of the defoliating beetle Physonota maculiventris Boheman (Chrysomelidae: Cassidinae) were studied in quarantine to determine its suitability for release as a biological control agent against Mexican sunflower, Tithonia diversifolia (Hemsl.) A. Gray (Asteraceae), in South Africa. Females laid 5.25?±?0.25 (mean?±?SE) egg batches during their lifetimes, with each batch consisting of approximately 33 eggs. Larvae were highly gregarious as early instars and both larvae and adults fed voraciously, often defoliating the plants completely. The life cycle of the beetle was completed in 67.5?±?7.5 days under quarantine conditions. Among the 58 test plant species subjected to no-choice tests, P. maculiventris developed successfully on T. diversifolia and some sunflower (Helianthus annuus L.) cultivars. However, only minor damage was recorded on non-target species, notably the exotic weed Xanthium strumarium L. and some H. annuus cultivars. Also, survival to adulthood was considerably lower on sunflower cultivars than on the target weed during these tests. During choice tests, P. maculiventris oviposited and developed successfully on T. diversifolia only, with only minor feeding damage on some H. annuus cultivars, suggesting that the beetle’s field host range will be confined to the target weed. Risk analysis also showed that P. maculiventris presents an extremely low risk to non-target plant species (e.g. within the tribe Heliantheae and other close relatives). The study concluded that P. maculiventris is safe for release in South Africa and an application for its release is being considered by the relevant South African regulatory authorities.     

Temperature requirements for seed germination of Pereskia aculeata and Pereskia grandifolia

Pereskia aculeata and Pereskia grandifolia have been studied widely due to their high nutritional and therapeutic values. However, little is known about the biological requirements of their seeds for the various germination factors. Thus, this experiment aimed to evaluate the thermal effects on the germination of these species at the temperatures of 24 °C, 27 °C, 30 °C, 33 °C and 36 °C. After verification of the existence of differences in the performance of germination, a non-linear regression was carried out, relating the germination to temperature and identifying its point of maximum efficiency. We found that the lowest synchronization indexes of germination were observed close to 30 °C. The best germination response of the P. aculeata and P. grandifolia was observed at 30 °C and 33 °C, respectively, with greater germination strength and fewer days to attain 63.21% of germinations. The results obtained from the germination of P. aculeata and P. grandifolia can be described by the Weindull distribution model with three parameters, as proposed by Carneiro and Guedes (1992).     

Using molecular methods to determine the origin of weed populations of Pereskia aculeata in South Africa and its relevance to biological control

New biological control agents are required in order to reach and sustain an adequate level of control of the declared environmental weed Pereskia aculeata Miller (Cactaceae) in South Africa. Identifying the origin of weed genotypes has been important in a number of biological control programmes and is likely to be of importance for the control of P. aculeata due to its disjunct native distribution and morphological polymorphisms between plants from different regions of the native and introduced distribution. DNA sequencing of the trnL chloroplastic intron and the phyC nuclear gene indicate that the South African weed population’s origin was in the southern region of native distribution. Inter-Simple Sequence Repeats (ISSRs) confirmed this result and added resolution to the analysis indicating that the native plants with the closest genetic distance to the South African weed population were found in Rio de Janeiro Province, Brazil. The relationship between the South African weed population and garden variety plants as well as the large genetic distance between the South African plants and the native plants suggests that the South African population may be the progeny of escaped garden variety plants that have been cultivated and possibly hybridized. The low levels of genetic variation within the South African population and the monophyly of the South African plants indicates that these plants are the progeny of a single introduction or multiple introductions from the same source. Rio de Janeiro Province in Brazil is the most appropriate region in which to survey for new biological control agents.     

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.     

Biology and host range of Mada polluta,a potential biological control agent of Tecoma stans in South Africa

The host range of Mada polluta Mulsant (Coleoptera:Coccinnellidae) was studied to assess its suitability as a biological control agent of Tecoma stans (L.) Juss ex Kunth var stans (Bignoniaceae), an invasive weed in South Africa. Biology of M. polluta and its host range were determined in the laboratory using no-choice and multi-choice feeding, oviposition and larval survival tests. Out of 36 plant species from 12 plant families (Bignoniaceae, Acanthaceae, Asteraceae, Verbenaceae, Lamiaceae, Oleaceae, Cucurbitaceae, Fabaceae, Scrophulariaceae, Solanaceae, Apiaceae, Chenopodiaceae and Poaceae) within the order Lamiales that were tested during the host specificity testing, M. polluta showed a very strong preference for T. stans, depositing its eggs on T. stans and none on non-target plant species. In no-choice tests, an average of 246 eggs was laid on T. stans, and from these, 133 larvae developed to adulthood. The beetle also showed very promising biological attributes that will contribute to its success as a biological control agent of T. stans. These attributes include highly damaging larvae and adults, high fecundity (532 eggs/female) and a short life cycle (four weeks). The short life cycle will enable multiple generations per year and rapid population increase in the field. It is concluded that M. polluta is sufficiently host-specific to be released against T. stans in South Africa. It is, therefore, strongly recommended that permission be granted to release this beetle from quarantine for biological control of T. stans in South Africa.     

Invaded habitat incompatibility affects the suitability of the potential biological control agent Listronotus sordidus for Sagittaria platyphylla in South Africa

Sagittaria platyphylla (Engelmann) J.G. Smith (Alismataceae) was first recorded in South Africa in 2008 and is considered to be an emerging weed with naturalised populations occurring throughout the country. A biological control programme was initiated in Australia and surveys conducted between 2010 and 2012 yielded potential agents, including the crown feeding weevil, Listronotus sordidus Gyllenhal (Coleoptera: Curculionidae). The potential of L. sordidus as a candidate biological control agent against S. platyphylla in South Africa was examined. Although adult feeding was recorded on a number of plant species, oviposition and larval development indicated a narrow host range restricted to the Alismataceae. In South Africa, S. platyphylla populations are primarily found in inundated systems. However, laboratory studies showed that L. sordidus did not oviposit on inundated plants, potentially nullifying the impact of the insect on South African populations. It is suggested that even though L. sordidus is a damaging, specific agent, its limited impact on inundated plant populations in South Africa does not justify the inherent risk associated with the release of a biological control agent.     

Megamelus scutellaris (Hemiptera: Delphacidae), a biocontrol agent of water hyacinth,is not sufficiently specific for release in Australia

Water hyacinth Eichhornia crassipes (Pontederiaceae) is one of the world's worst invasive species, responsible for damaging aquatic systems in many warmer parts of the globe including north America, Africa, Asia and Australia. The planthopper Megamelus scutellaris Berg (Delphacidae) has been released in USA and approved for release in South Africa for biocontrol of water hyacinth. We assessed this agent for suitability for release in Australia and found that a related native aquatic plant, Monochoria cyanea (Pontederiaceae) is within the fundamental host range of this insect. Adult survival, oviposition and development of nymphs to adult was equally high on M. cyanea as on the target species, although the quality of these next generation adults was lower than those reared on the target species. This demonstrates that M. scutellaris is not sufficiently specific for release in Australia. Nymphal development to adults occurred only in very low numbers on the three other Australian species of Monochoria. M. cyanea only occurs in Australia so M. scutellaris is still a possible water hyacinth biocontrol candidate for other regions depending on the results of assessment of the risk to local species of Monochoria. This study demonstrates the effectiveness of modern biocontrol agent assessment and reinforces the importance of testing of local non-target species.     

Life History and Laboratory Host Range ofEccritotarsus catarinensis(Carvalho) (Heteroptera: Miridae), a New Natural Enemy Released on Water Hyacinth (Eichhornia crassipes(Mart.) Solms-Laub.) (Pontederiaceae) in South Africa

A mirid,Eccritotarsus catarinensis(Carvalho), was studied as a potentially damaging natural enemy for water hyacinth, (Eichhornia crassipes(Mart.) Solms-Laub.), in South Africa. In the laboratory, eggs were inserted into the leaf tissue parallel to the leaf surface. The four nymphal instars fed gregariously with the adults mainly on the undersurface of the leaves, causing severe chlorosis at high population levels. The duration of immature stages (egg and nymphs) was approximately 23 days, while the adults survived for approximately 50 days. Favorable biological characteristics ofE. catarinensisincluded a high rate of increase, gregarious habits, long-lived and mobile adults, and several generations per year. Laboratory host range of the mirid was determined by adult choice trials on 67 plant species in 36 families and adult no-choice trials on five species in the Pontederiaceae. Feeding was recorded on all Pontederiaceae tested and oviposition on four of the five species. However, these plant species proved to be inferior hosts forE. catarinensisin comparison to water hyacinth, suggesting thatE. catarinensiswould be an acceptable natural enemy for water hyacinth in South Africa.     

Biology and host range of <Emphasis Type="Italic">Aconophora compressa,</Emphasis> a candidate considered as a biocontrol agent of <Emphasis Type="Italic">Lantana camara</Emphasis> in Africa

The shrub, Lantana camara L. (Verbenaceae), of tropical American origin, is a serious weed in South Africa and is the target of a biological control programme. The stem-sucking membracid, Aconophora compressa Walker (Homoptera: Membracidae), from Mexico was reported to be very damaging and was therefore imported into South Africa for biology and host range studies. The female partially inserts the eggs into the woody portion of actively growing stems, and guards them against potential predators. Nymphs develop through five instars to the adult stage in about 45 days. The adults and nymphs feed on the sap of stems causing the leaves to wilt, the flowers to abort and the gradual dieback of stems. No-choice experiments showed that adult survival, egg production, and nymphal emergence was high on L. camara, and also on related ornamental and indigenous species. The adult reproductive performance was higher on some indigenous Lippia species than on L. camara. Furthermore, the nymphs developed faster on these indigenous species and high rates of reproductive performance on these non-target species were sustained over several generations. In multiple-choice trials, equal or larger numbers of egg batches were recorded from indigenous Lippia species and the ornamental plant, Aloysia citrodora Palau, than on L. camara. The treehopper, A. compressa, poses an unacceptable threat to indigenous Lippia species and has therefore been rejected as a biocontrol agent for L. camara in Africa.     

The fixed plot survey method for determining the host range of the flowerbud-feeding weevil Dicomada rufa, a candidate for the biological control of Hakea sericea in South Africa

The hakea bud weevil, Dicomada rufa (Curculionidae), is a promising candidate for the biological control of the weed Hakea sericea (Proteaceae) in South Africa. Because D. rufa could not be successfully cultured on potted plants in quarantine, most of theconventional methods for host range determination were not suitable. A type of open-field testing method, the fixed plot survey method, was developed to show that D. rufa is host specific to H. sericea. The trial was conducted in three 1–2 ha plots at three localities in New SouthWales, Australia, involving 41 test plantspecies. This result was combined with otherconventional considerations to apply for therelease of D. rufa in South Africa. Theseincluded a multiple choice feeding trial inquarantine in South Africa, during which 10test species from seven genera of South AfricanProteaceae were not accepted for feeding. AllSouth African Proteaceae, except Brabejumstellatifolium, are phylogenetically distinctfrom H. sericea. Also, there are norecords of D. rufa interacting withcommercially important plants in Australia,including commercially cultivated South AfricanProteaceae. Dicomada rufa adult andlarval feeding destroys buds, flowers, smallfruits and succulent shoots. It is expectedthat this damage will supplement that of thetwo seed-feeding biological control agentsalready established in South Africa and furtherreduce the reproductive potential of the weed,particularly that of young plants regeneratingafter fires. Without compromising safety, thefixed plot survey method may also contribute toreducing the time and cost normally associatedwith conventional host specificity testing.     

Identification and genetic diversity of two invasive <Emphasis Type="Italic">Pissodes</Emphasis> spp. Germar (Coleoptera: Curculionidae) in their introduced range in the southern hemisphere

During the first half of the twentieth century, two accidental cases of introduction of Pissodes weevils were recorded from the southern hemisphere. The weevils in South Africa were identified as the deodar weevil (Pissodes nemorensis) and those in South America as the small banded pine weevil (Pissodes castaneus). Wide distribution of the two species in their invasive range, general difficulty in identifying some Pissodes spp., and the varying feeding and breeding behaviours of the species in South Africa has necessitated better evidence of species identity and genetic diversity of both species and population structure of the species in South Africa. Barcoding and the Jerry-to-Pat region of the COI gene were investigated. Morphometric data of the South African species was analysed. Our results confirmed the introduction of only one Pissodes species of North American origin to South Africa. However, this species is not P. nemorensis, but an unrecognized species of the P. strobi complex or a hybrid between P. strobi and P. nemorensis. Only P. castaneus, of European origin, was identified from South America. We identified ten mitochondrial DNA haplotypes from South Africa with evidence of moderate genetic structure among geographic populations. Terminal leader and bole-feeding weevils did not differ at the COI locus. A single haplotype was identified from populations of P. castaneus in South America. Results of the present study will have implications on quarantine, research and management of these insect species.     

Candidates for biocontrol of Macfadyena unguis-cati in South Africa: biology, host ranges and potential impact of Carvalhotingis visenda and Carvalhotingis hollandi under quarantine conditions

Macfadyena unguis-cati (L.) Gentry (Bignoniaceae) was introduced as an ornamental in South Africa, but is fast becoming an important invasive plant in many areas. It is difficult to control the plant chemically and mechanically. The first biocontrol agent, the chrysomelid Charidotis auroguttata (Boheman), has been released. It established at some release sites, but numbers have so far remained low. Additional biocontrol agents were sought to augment C. auroguttata. The potential host ranges of two foliage feeding lace bugs, Carvalhotingis visenda (Drake and Hambleton) and C. hollandi (Drake) (Hemiptera: Tingidae) were evaluated on the basis of nymphal no-choice and adult multi-choice tests involving 23 plant species in 11 families. In no-choice tests, nymphs of both species were able to survive and complete development on M. unguis-cati only, and adults of both species only fed and oviposited on M. unguis-cati during the adult multi-choice tests. Host specificity tests thus confirm that the tingids are highly host specific biocontrol agents, and will not pose risk to any non-target plants in South Africa. A study to determine the potential impact of C. hollandi nymphal feeding on M. unguis-cati showed a significant decrease in the chlorophyll contents of leaves when compared to those of control plants. These studies indicate that, once released, the two lace bug species could contribute significantly to the biological control of M. unguis-cati in South Africa.     

中国新发现外来入侵害虫——南美番茄潜叶蛾(鳞翅目:麦蛾科)

2017年8月,在新疆维吾尔自治区伊犁哈萨克自治州露地鲜食番茄上发现一种鳞翅目害虫,以幼虫潜食叶肉、蛀食果实,经鉴定为南美番茄潜叶蛾。该害虫原产南美洲的秘鲁,2006年入侵欧洲的西班牙,截至2017年5月,已在南美洲、欧洲、非洲、中美洲和亚洲的80多个国家和地区发生,严重危害鲜食番茄、加工番茄和樱桃番茄/圣女果,产量损失最高可达80%～100%。本文提供了南美番茄潜叶蛾的危害情况及主要形态鉴定特征,提出了防范其进一步扩散危害的措施建议以及今后应开展的主要研究方向,包括植物检疫、发生分布调查、生物学生态学特性研究、天敌资源挖掘利用等,以为积极应对南美番茄潜叶蛾对我国农业生产安全的威胁提供参考。     

Assessing the risks associated with the release of a flowerbud weevil, Anthonomus santacruzi, against the invasive tree Solanum mauritianum in South Africa

Biological control of Solanum mauritianum Scopoli, a major environmental weed in the high-rainfall regions of South Africa, is dependent on the establishment of agents that can reduce fruiting and limit seed dispersal. The flowerbud weevil, Anthonomus santacruzi Hustache, is a promising fruit-reducing agent, despite ambiguous results obtained during host-specificity evaluations in quarantine. Adult no-choice tests showed that although feeding is confined to Solanum species, normal feeding and survival occurred on the foliage (devoid of floral material) of cultivated eggplant (aubergine), potato, and several native South African Solanum species. During paired-choice tests, involving floral bouquets in 10-liter containers, A. santacruzi oviposited in the flower buds of 12 of the 17 test species, including potato and eggplant, although significantly more larvae were recovered on S. mauritianum than on eight other species. Larvae survived to adults on all 12 species, with survival significantly lower on only four species than on S. mauritianum. However, during multi-choice tests, involving potted plants in a large walk-in cage, A. santacruzi consistently displayed significant feeding and oviposition preferences for S. mauritianum over all of the 14 Solanum species tested. Analyses of the risk of attack on nontarget Solanum plants suggested that, with the possible exception of two native species, none is likely to be extensively utilized as a host in the field. Also, host records and field surveys in South America have suggested that A. santacruzi has a very narrow host range and that the ambiguous laboratory results are further examples of artificially expanded host ranges. These and other considerations suggest that A. santacruzi should be considered for release against S. mauritianum in South Africa, and an application for permission to release the weevil was submitted in 2003.