Persistence of the flowerbud weevil Anthonomus santacruzi in optimal versus marginal areas: implications for the biological control of the invasive tree Solanum mauritianum in South Africa

Anthonomus santacruzi Hustache (Coleoptera: Curculionidae) was released in South Africa to offset the extensive reproductive output of the invasive tree Solanum mauritianum Scopoli (Solanaceae). Widespread establishment has occurred predominantly in the coastal areas of KwaZulu-Natal province, with limited success in higher-altitude inland areas. Irrespective of location, populations exhibit peaks in the austral autumn and decline during winter. In this study, we evaluated the persistence of A. santacruzi populations in climatically-optimal coastal areas versus climatically-marginal inland areas. The weevil’s pre- and post-winter abundance was surveyed at six coastal and six inland sites during 2018, and compared between areas and seasons. The 2018 pre-winter data were also compared to 2016 pre-winter data collected at the same sites. During 2018, pre- and post-winter numbers were six times and 22 times higher, respectively, at optimal sites than at marginal sites, with substantial winter declines at all sites. Post-winter weevil numbers at optimal sites were significantly higher than pre-winter numbers at marginal sites. Pre-winter numbers at optimal sites were not significantly different between 2016 and 2018, but at marginal sites were significantly lower in 2018. Inflorescences of S. mauritianum at marginal sites contained significantly more floral material and fruit than those at optimal sites, during both seasons in 2018. Significant negative correlations between A. santacruzi numbers and floral/fruit production suggest some impact on the reproductive output of S. mauritianum. Since A. santacruzi populations are barely persisting in marginal areas, releases in other South African provinces should target locations that are below 300?m in altitude.     

Inflorescence-inhabiting spiders do not threaten the survival and fecundity of Anthonomus santacruzi (Curculionidae), a florivorous biocontrol agent of Solanum mauritianum (Solanaceae) in South Africa

The florivorous weevil Anthonomus santacruzi is being released in South Africa to reduce the reproductive capacity of the invasive tree, Solanum mauritianum. Initiated to allay concerns about biotic interference, our study suggests that inflorescence-inhabiting spiders pose no risk to the survival and proliferation of A. santacruzi populations in the field.     

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.     

Should the flower bud weevil Anthonomus santacruzi (Coleoptera: Curculionidae) be considered for release against the invasive tree Solanum mauritianum (Solanaceae) in New Zealand?

The invasive tree Solanum mauritianum Scop. has been targeted for biological control in South Africa and New Zealand, by deploying insect agents that could constrain its excessive reproductive output. The flower-feeding weevil Anthonomus santacruzi (Curculionidae) was approved for release in South Africa in 2007 but following the loss of the original culture in quarantine, new stocks were introduced from Argentina in 2008–2009. This study was initiated to confirm that the host range of the new culture was the same as that of the previous one, but also to assess the risks associated with the weevil's release in New Zealand. Different testing procedures, including no-choice tests and multi-choice tests in different arenas, produced inconsistent and ambiguous results. During no-choice tests, oviposition and larval development to adulthood occurred on three non-target species including two native South African and one native New Zealand Solanum species. However, subsequent multi-choice tests and a risk assessment suggested that the risks of anything more than collateral damage to non-target Solanum species are low. Overall, these data do not deviate substantially from the results of the original quarantine tests which facilitated the release of A. santacruzi in South Africa in 2009. Although we argue that none of the New Zealand native and cultivated species are at risk, stronger evidence from open-field trials and chemical ecology studies may be required to convince the regulatory authorities that A. santacruzi is suitable for release in New Zealand.     

A review of <Emphasis Type="Italic">Solanum mauritianum</Emphasis> biocontrol: prospects,promise and problems: a way forward for South Africa and globally

The invasive tree Solanum mauritianum Scopoli remains one of the world’s most widespread environmental weeds. Despite biocontrol providing one of the few viable long-term solutions to tackling S. mauritianum invasions globally, only South Africa and, more recently, New Zealand, have programmes in place. Ongoing biocontrol efforts against S. mauritianum are reviewed here with particular reference to South Africa. The South African programme has suffered a troubled history, with considerable research efforts culminating in the eventual release and establishment of only two insect agents, Gargaphia decoris Drake and Anthonomus santacruzi Hustache. The difficulties experienced have hindered research into new agents, causing apprehension in using biocontrol internationally. However, recent studies have demonstrated that biocontrol may be deserving of renewed investment, particularly within an integrated management context. In this review, we advocate for the revival of the S. mauritianum biocontrol programme in South Africa, and discuss possible avenues for future research internationally.     

Biology, host specificity and risk assessment of the leaf-mining flea beetle, Acallepitrix sp. nov., a candidate agent for the biological control of the invasive tree Solanum mauritianum in South Africa

The South American tree Solanum mauritianum, a major environmentalweed in the high-rainfall regions of SouthAfrica, has proved to be a difficult target forbiological control. Artificially expanded hostranges of candidate agents during quarantinehost-specificity tests, have resulted in therelease of only one agent species which has sofar had a negligible impact on the weed. Thenecessity for additional agents resulted in theimportation of a leaf-mining flea beetle, anunidentified species of Acallepitrix,from Brazil in 1997. No-choice tests inquarantine showed that potential host plantsare confined to the genus Solanum.Although several non-target plants, includingcultivated eggplant (aubergine) and some nativeSouth African Solanum species, sustainedfeeding, oviposition and the development oflarval leaf mines, most of these were inferiorhosts. These results were confirmed by pairedchoice tests, where Acallepitrix sp. nov.displayed significant feeding and ovipositionpreferences for S. mauritianum and where,with few exceptions, more larval leaf mineswere initiated on S. mauritianum.Interpretation of the host-specificity testswas facilitated by a risk assessment matrixwhich suggested that the risk of feeding andoviposition on non-target Solanum plants,including eggplant cultivations, was relativelylow. These and other considerations, such asthe lack of evidence of damage inflicted oncultivated Solanaceae in South America, suggestthat Acallepitrix sp. nov. could beconsidered for release against S. mauritianum in South Africa. However, the results of the host-specificity tests remain ambiguous and until more compelling evidence is obtained from field surveys and open-field trials in Brazil, an application for permission to release the flea beetle in South Africa will not be considered.     

Comparing the effects of the exotic cactus-feeding moth, Cactoblastis cactorum (Berg) (Lepidoptera: Pyralidae) and the native cactus-feeding moth, Melitara prodenialis (Walker) (Lepidoptera: Pyralidae) on two species of Florida Opuntia

This study examined the effects of the native cactus moth borer, Melitara prodenialis, and the invasive cactus moth borer, Cactoblastis cactorum, on two common cactus species, Opuntia stricta and O. humifusa at coastal and inland locations in central Florida. Opuntia stricta were present only at coastal sites and O. humifusa were present at coastal and inland sites. Throughout the duration of the study, coastal plants were subject to damage solely by C. cactorum and inland plants solely by M. prodenialis. Results showed marginally significantly higher numbers of eggsticks on O. stricta than O. humifusa and significantly higher numbers at coastal sites than at inland sites. There was also significantly higher moth damage on O. stricta than O. humifusa and at coastal sites than inland sites, but not significantly so. However, there was a higher level of plant mortality for O. humifusa than for O. stricta and a significantly higher level of cactus mortality at inland sites when compared to coastal sites. This increased mortality may be due to increased attack by true bugs, Chelinidea vittiger, and by Dactylopius sp., combined with attack by M. prodenialis. Inland plants also tended to be smaller than coastal plants and could be more susceptible to the combined effects of all insects. Further long-term research on coastal cactus survival when attacked and unattacked by Cactoblastis is necessary to fully determine the effects of this moth on Opuntia survival.     

Distribution and habitats of Bulinus natalensis and its role as intermediate host of economically important helminth parasites in South Africa

This article focusses on the geographical distribution and habitats of Bulinus natalensis, as reflected by the samples taken from 244 collection sites on record in the database of the National Freshwater Snail Collection (NFSC) at the Potchefstroom campus of the North-West University. This snail species is experimentally susceptible to Schistosoma haematobium and is incriminated in the natural transmission of S. margrebowiei, S. bovis and Calicophoron microbothrium elsewhere in Africa. The 78 different loci (1/16 degree squares) on record reflect a geographical distribution that is largely limited to the KwaZulu-Natal Province. Details of each habitat as described by collectors during surveys, as well as altitude and mean annual air temperature and rainfall for each locality, were processed and chi-squared and effect size values calculated. A decision tree constructed from all the available data indicated that temperature and altitude, followed by the type of water body, seemed to be the more important factors that significantly influenced the distribution of this species in South Africa. The possible role of this species as intermediate host of economically important helminth species is briefly examined. The need to clarify its relationships with B. tropicus and B. depressus, and to update the geographical distribution in South Africa, is also emphasised.     

Assessing the risks of releasing a sap-sucking lace bug, <Emphasis Type="Italic">Gargaphia decoris</Emphasis>, against the invasive tree <Emphasis Type="Italic">Solanum mauritianum</Emphasis> in New Zealand

The South American tree Solanum mauritianum Scopoli (Solanaceae), a major environmental weed in South Africa and New Zealand, has been targeted for biological control, with releases of agents restricted to South Africa. The leaf-sucking lace bug, Gargaphia decoris Drake (Tingidae), so far the only agent released, has become established in South Africa with recent reports of severe damage at a few field sites. To evaluate the insect’s suitability for release in New Zealand, host-specificity testing was carried out in South Africa in laboratory and open-field trials, with selected cultivated and native species of Solanum from New Zealand. No-choice tests confirmed the results of earlier trials that none of the three native New Zealand Solanum species are acceptable as hosts. Although the cultivated Solanum muricatum Aiton and S. quitoense Lam. also proved unacceptable as hosts, some cultivars of S. melongena L. (eggplant) supported feeding, development and oviposition in the no-choice tests. Although eggplant was routinely accepted under laboratory no-choice conditions in this and previous studies, observations in the native and introduced range of G. decoris, open-field trials and risk assessment based on multiple measures of insect performance indicate that the insect has a host range restricted to S. mauritianum. These results strongly support the proposed release of G. decoris in New Zealand because risks to non-target native and cultivated Solanum species appear to be negligible. An application for permission to release G. decoris in New Zealand will be submitted to the regulatory authority. Handling editor: John Scott.     

The introduction and release of Chiasmia inconspicua and C. assimilis (Lepidoptera: Geometridae) for the biological control of Acacia nilotica in Australia

Two geometrid moths Chiasmia inconspicua and Chiasmia assimilis, identified as potential biological control agents for prickly acacia Acacia nilotica subsp. indica, were collected in Kenya and imported into quarantine facilities in Australia where laboratory cultures were established. Aspects of the biologies of both insects were studied and CLIMEX® models indicating the climatically favourable areas of Australia were developed. Host range tests were conducted using an approved test list of 74 plant species and no-choice tests of neonate larvae placed on both cut foliage and potted plants. C. inconspicua developed through to adult on prickly acacia and, in small numbers, Acacia pulchella. C. assimilis developed through to adult on prickly acacia and also in very small numbers on A. pulchella, A. deanei, A. decurrens, and A. mearnsii. In all experiments, the response on prickly acacia could be clearly differentiated from the responses on the non-target species. Both insects were approved for release in Australia. Over a three-year period releases were made at multiple sites in north Queensland, almost all in inland areas. There was no evidence of either insect’s establishment and both colonies were terminated. A new colony of C. assimilis was subsequently established from insects collected in South Africa and releases of C. assimilis from this new colony were made into coastal and inland infestations of prickly acacia. Establishment was rapid at one coastal site and the insect quickly spread to other infestations. Establishment at one inland area was also confirmed in early 2006. The establishment in coastal areas supported a CLIMEX model that indicated that the climate of coastal areas was more suitable than inland areas.     

The viability and germination characteristics of exhumed Solanum mauritianum seeds buried for different periods of time

The site, depth and duration of burial significantly influenced the viability and state of dormancy of Solanum mauritianum seeds. Burial at a depth of 15 cm was most effective in reducing the level of conditional dormancy. Secondary dormancy was not induced at any of the environmental (burial) sites when seeds were maintained at 15 cm, where light and temperature fluctuations were minimal. When buried at 4 cm or maintained on the soil surface secondary dormancy was induced, particularly at the inland sites where environmental conditions such as temperature and moisture were more extreme. Conditional dormancy could generally be overcome by incubating seeds at 15/30 °C in the light, even after prolonged burial at unfavourable germination conditions. Gibberellic acid (500 mg l^–1) was very effective in breaking secondary dormancy of seeds induced by storage under unfavourable conditions after burial. These results have important implications for the control of this week in commercial forests.     

Breeding distribution,population size and conservation of the Greyheaded Gull Larus cirrocephalus in southern Africa

Brooke, R.K., Allan, D.G., Cooper, J., Cyrus, D.P., Dean, W.R.J., Dyer, B.M., Martin, A.P. & Taylor, R.H. 1999. Breeding distribution, population size and conservation of the Greyheaded Gull Larus cirrocephalus in southern Africa. Ostrich 70 (3&4): 157–163.

The Greyheaded Gull Larus cirrocephalus occurs throughout southern Africa, both coastally and inland, and has bred at one time or another at 67 known localities since the 1860s. Most of these sites have been occupied by small numbers of birds, even single pairs, and for only one or a few years. The two principal breeding areas are the East Rand in Gauteng and Lake St Lucia in KwaZulu-Natal, both in South Africa. The next most important sites are Walvis Bay, Namibia and Lake Ngami, Botswana. The total southern African breeding population is estimated as about 2000 pairs. The Greyheaded Gull is not a threatened species in southern Africa, with 27 breeding sites supporting more than half the breeding population within formally conserved areas.     

Potential spread of the invasive plant Hydrilla verticillata in South Africa based on anthropogenic spread and climate suitability

Hydrilla verticillata (L.f.) Royle is a submerged aquatic plant native to Asia and Australia that is highly invasive in the USA and was first recorded in South Africa in 2006. It is only known from one locality, Pongolapoort Dam in KwaZulu-Natal Province, but there are fears that it might spread to other sites. The primary vector of spread in the USA is recreational boaters and anglers. A survey at a fishing competition on Pongolapoort Dam showed that anglers travel considerable distances around South Africa (73% of water bodies were >200 km, visited by 68% of the respondents). A Threat Index for freshwater bodies throughout South Africa visited by participants of the competition was calculated showing that dams in the vicinity of the infestation were more at risk from invasion. Further, the potential distribution of the weed based on climatic matching with the region of origin showed that most of the country was suitable for establishment, with the exception of the high-lying interior of the country. Recommendations for reducing the potential spread of hydrilla in South Africa are presented.     

Dispersal of the Cotton Boll Weevil (Coleoptera: Curculionidae) in South America: Evidence of RAPD Analysis

RAPD technique provides useful information on the geographic origin and dispersal of the boll weevil Anthonomus grandis in South America. Nine populations from Argentina, Brazil, Paraguay, Mexico and USA were analyzed. Weevils were captured on native plants (Misiones province, Argentina) and on cotton cultures, except the sample from the United States (USDA laboratory-reared colony). A sample of the ‘Peruvian square weevil’, A. vestitus, from Ecuador, was included in the analysis in order to compare interspecific variation. The four primers used in the analysis revealed 41 ‘anonymous loci’. The neighbor-joining tree based on Nei's distances and values of Nm (migrants per generation), indicate that genetic similarity between samples from Tecomán (Mexico) and Puerto Iguazú (Argentina), is higher than among remaining South American populations. This result supports an hypothesis of natural occurrence of the boll weevil in South America, prior to extensive cotton cultivation. Population outbreaks of the species would be associated with increase of agricultural lands. This revised version was published online in August 2006 with corrections to the Cover Date.     

Toward sustainable use of Palm leaves by a rural community in Kwazulu-Natal, South Africa

There is increasing pressure from communities neighbouring protected areas in KwaZulu-Natal, South Africa, to harvest resources from these areas. For example,Hyphaene coriacea Geartn. (vegetable ivory palm) leaves are extensively used by Zulu people for commercial and subsistence craftwork. This study investigates the effect of varying leaf harvest intensities on leaf production ofHyphaene coriacea in the Greater St Lucia Wetland Park in KwaZulu-Natal, South Africa, and aims to provide some management guidelines for sustainable use. Replicate plots subjected to three different harvest intensities were established and numbers and lengths of unopened leaves were monitored over four years. Results suggest that a harvest intensity of one leaf stem^-1 year^-1 would be sustainable. This is equivalent to 34% of calculated annual production. The 650 ha harvest area is estimated to produce 191 750 leaves year^-1 (4793 bundles) which could be harvested sustainably. The effects of leaf harvest, even at maximum harvest rate, on leaf production over the four year study period appear negligible although there was a slight decline over time in leaf production of trees subjected to high harvest intensity. A slight increase in leaf production was observed at moderate levels of harvest intensity.     

DISTRIBUTION AND NUMBERS OF COASTAL WADERS (CHARADRII) IN THE SOUTHWESTERN CAPE,SOUTH AFRICA,SUMMER 1975–76

Summers, R. W., Cooper, J. & Pringle, J. S. 1970. Distribution and numbers of coastal waders (Charadrii) in the southwestern Cape, South Africa, summer 1975–76. Ostrich 48: 85–97. A survey of the distribution and numbers of waders (suborder Charadrii) in the coastal regions of the southwestern Cape was conducted from November 1975 to February 1976. Forty-four coastal wetlands and 49 sections of coastline were visited between the Olifants and Groot-Brak rivers. The estimated total wader population was 119 008 of which 102 841 (86,4%) were Palaearctic migrants. Langebaan Lagoon and the Berg estuary supported the largest numbers. The Curlew Sandpiper Calidris ferruginea and Sanderling Calidris alba were the most numerous species. The abundance of waders was greater on west coast shores than along the south coast.     

Biological control of Salvinia molesta in some areas of Moremi Game Reserve,Botswana

The Curculionid weevil, Cyrtobagous salviniae Calder & Sands, has been established on Salvinia molesta (salvinia) in Botswana. The study determined the intensity of weevil activity on the weed at the margin and the centre of four selected sites on the Khwai system of the Okavango Delta. Random samples of salvinia were collected from each site at monthly intervals in 1998 to extract weevils and to demonstrate the effect of weevil on the weed. The rate at which the weed was controlled at different sites varied with mat and weevil density. The biological control at Paradise Pools was moderate without much increase in the weed biomass per m², while fluctuations in both weevil and plant populations were noticed in the Khwai streams. Weevil numbers remained low at Dombo for the first nine months of the year, during which time the mat density increased. With the onset of higher temperatures, weevil numbers increased to 155 and 457/kg fresh weight of salvinia at the margin and centre respectively in Dombo Pool in early summer. A significant control in MGR 6 HATAB pool between mat biomass and the weevil number resulted in the disappearance of the weed in three months. This study shows that C. salviniae is an effective biological control agent of S. molesta in semiarid areas.     

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.     

Inferred hybridisation,sympatry and movements of Chorister Robin-Chat Cossypha dichroa and Red-capped Robin-Chat C. natalensis

Inferred hybrid Chorister Robin-Chat Cossypha dichroa and Red-capped Robin-Chat C. natalensis have previously only been recorded from the Eastern Cape province, South Africa. We extend the occurrence of inferred hybrids with ringed and photographed examples from KwaZulu-Natal and Limpopo provinces. Sympatry at a fine scale was investigated at Vernon Crookes Nature Reserve, KwaZulu-Natal, using observational data over 30 years (1978–2008) and intensive mist-netting during two field-trips in June and October 2010. Both robin-chat species were found throughout the year and occurred in the same forest patches. Inferred hybrids are known from five different forest patches in Vernon Crookes, three with photographic evidence. During winter an increase in Chorister Robin-Chat numbers was detectable both from mist-netting and checklist data. An increase of Red-capped Robin-Chats was observed from mist-netting and more equivocally from checklist data in spring. Red-capped and Chorister Robin-Chats are sympatric along the eastern escarpment of South Africa, with co-occurrence in 96 quarter degree squares from Southern African Bird Atlas Project 1 data. During the breeding season, co-occurrence was recorded in 62 quarter degree squares and in nine grid-cells evidence of breeding by both species is available. Hybridisation may be more common along the extensive area of sympatry than currently realised.     

Evaluation of hydrolytic enzyme activities from digestive fluids of Anthonomus grandis (Coleoptera: Curculionidae)

The cotton boll weevil, Anthonomus grandis, is a major pest of cotton crops in South America. In this work, partial biochemical characterizations of (hemi) cellulases and pectinases activities in the digestive system (head- and gut- extracts) of A. grandis were evaluated. Gut extract section from third instar larvae exhibited endoglucanase, xylanase, β-glucosidase, and pectinase activities. The endoglucanase and xylanase activities were localized in the foregut, whereas β-glucosidase activity was mainly detected in the hindgut. In addition, no difference in pectinase activity was observed across the gut sections. Thus, A. grandis digestive system is a potentially interesting reservoir for further lignocellulolytic enzymes research.