Synergies between research organisations and the wider community in enhancing weed biological control in South Africa

Biological control offers a cost effective and ecologically sustainable tool for the management of invasive alien plants. Its implementation, however, has historically been slow and poorly co-ordinated. In South Africa, as in many other countries, most aspects of biological control programmes were done by researchers, but from 1995 onwards, with the advent of the Working for Water Programme, a more inclusive approach to biological control has been adopted. In this paper, we report on the development of community-based biological control implementation programmes in South Africa, after 1995, and highlight a number of initiatives, including employing persons with disabilities at mass-rearing facilities and in particular, we outline a suite of educational and outreach programmes for the general public and for schools, which have increased capacity, education and employment in the field of weed biological control.     

Ecological studies to assess the efficacy of biological control on populations of alligator weed and lippia

Abstract  The risk of non-target effects from biological control agents can be reduced if we can better identify effective agents prior to release. Introducing only those agents with high potential for effective control will reduce the number of agents released and reduce the probability of both direct and indirect non-target impacts. Identifying effective agents requires understanding the roles that resources, disturbances and herbivory play in regulating plant populations under natural field conditions. Here we propose a series of experiments that will contrast the mechanisms of population regulation of two invasive wetland plants, alligator weed ( Alternanthera philoxeroides ) and lippia ( Phyla canescens ), with native congener species, Alternanthera denticulata and Phyla nodiflora . We hypothesise that the native plants will have greater growth rates when nutrients are low and herbivores are present, whereas the introduced plants will exhibit greater growth rates when nutrients are high and herbivores are absent.     

Climate constrains the establishment and proliferation of Anthonomus santacruzi,a flower-feeding biological control agent of the invasive weed Solanum mauritianum in South Africa

The flowerbud-feeding weevil Anthonomus santacruzi Hustache (Coleoptera: Curculionidae) was released in South Africa in 2008 for the biological control of the invasive tree Solanum mauritianum Scopoli (Solanaceae). The weevil was widely deployed throughout KwaZulu-Natal province, which supports large S. mauritianum infestations, and has become well established in its warmer coastal regions. The aim of this study was to provide field evidence that climate is constraining the weevil’s distribution in South Africa. Solanum mauritianum populations were sampled at 23 sites across an altitudinal gradient in KwaZulu-Natal to determine A. santacruzi densities in relation to food availability and climatic variables. Despite significantly higher amounts of floral material on S. mauritianum at the higher altitude inland sites, A. santacruzi numbers were significantly higher at the lower altitude coastal sites. There was thus a significant negative relationship between A. santacruzi numbers and altitude and significant positive relationships between A. santacruzi numbers and both temperature and humidity. Neither rainfall nor food availability influenced A. santacruzi numbers, although lower amounts of floral material at the coastal sites may well have been caused by higher weevil densities at these sites. Anthonomus santacruzi was absent at only three sites, all at higher altitudes, further demonstrating that conditions in coastal or low-altitude regions are favourable for establishment and population proliferation. Future release efforts in KwaZulu-Natal, but also in other South African provinces, should thus be focused on coastal regions and inland regions that are below 1000 m above sea level.     

Predictable risk to native plants in weed biological control

Data on field host use of 112 insects, 3 fungi, 1 mite, and 1 nematode established for biological control of weeds in Hawaii, the continental United States, and the Caribbean indicate that the risk to native flora can be judged reliably before introduction. Virtually all risk is borne by native plant species that are closely related to target weeds. Fifteen species of insects introduced for biological control use 41 native plant species; 36 of which are congeneric with target weeds, while 4 others belong to two closely allied genera. Only 1 of 117 established biological organisms uses a native plant unrelated to the target weed. Thus the elements of protection for the native flora are the selection of weed targets that have few or no native congeners and the introduction of biological control organisms with suitably narrow diets.     

Suitability of a leaf-mining fly,Hydrellia sp., for biological control of the invasive aquatic weed,Hydrilla verticillata in South Africa

South Africa developed its first biological control programme for a submerged aquatic weed following the discovery of Hydrilla verticillata (L.f.) Royle (Hydrocharitaceae) in a major tourism dam in 2006. A leaf-mining fly, Hydrellia sp. (Diptera: Ephydridae) from Singapore, originating on a closely related biotype of the weed invading South Africa, was considered a priority candidate agent. Host range was investigated through no-choice and paired-choice larval development trials. “Host suitability” trials (multiple generations and performance) were used to further evaluate the potential for non-target host use by the fly. Hydrellia sp. developed on a few closely related species in the Hydrocharitaceae and one species in the Potamogetonaceae. However, paired-choice tests demonstrated a strong preference for its host plant and the host suitability trials indicated that Hydrellia sp. has very limited potential to establish permanent populations on native aquatic plant species. Hydrellia sp. should therefore be considered safe for release in South Africa.     

Death of the cereal root cortex: its relevance to biological control of take-all

Cell death in the root cortex of cereals was assessed by an inability to detect nuclei, using acridine orangelfluorescence microscopy after fixation and mild acid hydrolysis. Seminal roots were scanned at x 100 magnification and their cortices were considered dead when nuclei were absent from all cell layers except the innermost one, adjacent to the endodermis; this cell layer remains alive long after the rest of the cortex has died. Cortical death of wheat and barley roots occurred in the absence of major pathogens. Cell death started behind the root hair zone of the main root axis, initially in the outermost cell layer of the cortex and then progressively inwards towards the endodermis; however, the cortex remained alive for a distance of c. 800 μm around emerging root laterals. The rate of cortical death was more rapid in wheat than in barley, both under field conditions and in the glasshouse at 20 °C. Thus, field-grown spring wheat (Sicca) showed 50% death of the root cortex in the top 6 cm of first seminal roots after 35 days (growth stage 1–2), whereas spring barley (Julia) showed 50% death of the root cortex after 67 days (growth stage 8). In the glasshouse, the top 9 cm of first seminal roots on 16-day plants showed 55% cortical death in wheat (Cappelle-Desprez) but only 2.5% cortical death in barley (Igri). The same rates of death were found in all subsequent seminal roots. The wheat root cortex died at the same rate in sterile and unsterile conditions, and at the same rate in the presence/absence of Phialophora radicicola Cain var. graminicola Deacon or Aureobasidium bolleyi (Sprague) von Arx. Hence, although P. radicicola and other soil microorganisms may benefit from root cortex death they do not exert biological control of take-all by enhancing or retarding the rate of this process. To study the effects of cortical death on take-all, Gaeumannomyces graminis (Sacc.) Arx & Olivier var. tritici Walker was point-inoculated at the tips and on older (5 and 15 day) regions of wheat seminal roots. After 17 days at 20 °C the fungus had grown to the same extent as runner-hyphae in all cases, but the severity of disease decreased with increasing age of the root cortex prior to inoculation; thus, G. graminis caused most extensive vascular discoloration and most intense vascular blockage in roots inoculated at their tips. Similar experiments on wheat and barley roots inoculated separately with P. radicicola and G. graminis suggest that at least three factors associated with cortical death influence infection by these fungi: (1) initially, cell death may enhance infection because nutrients are made available to the parasites and host resistance within the cortex is reduced; (2) weak parasites and soil saprophytes may colonise dead and dying cortices in competition with G. graminis and P. radicicola and thereby reduce infection by these fungi; (3) changes in the endodermis and adjacent cell layers may be associated with cortical death and may retard invasion of the stele. Future work will seek to establish the relative importance of these factors and extend this study to other cereal host-fungus combinations.     

European origin of Bradyrhizobium populations infecting lupins and serradella in soils of Western Australia and South Africa

We applied a multilocus phylogenetic approach to elucidate the origin of serradella and lupin Bradyrhizobium strains that persist in soils of Western Australia and South Africa. The selected strains belonged to different randomly amplified polymorphic DNA (RAPD)-PCR clusters that were distinct from RAPD clusters of applied inoculant strains. Phylogenetic analyses were performed with nodulation genes (nodA, nodZ, nolL, noeI), housekeeping genes (dnaK, recA, glnII, atpD), and 16S-23S rRNA intergenic transcribed spacer sequences. Housekeeping gene phylogenies revealed that all serradella and Lupinus cosentinii isolates from Western Australia and three of five South African narrow-leaf lupin strains were intermingled with the strains of Bradyrhizobium canariense, forming a well supported branch on each of the trees. All nodA gene sequences of the lupin and serradella bradyrhizobia formed a single branch, referred to as clade II, together with the sequences of other lupin and serradella strains. Similar patterns were detected in nodZ and nolL trees. In contrast, nodA sequences of the strains isolated from native Australian legumes formed either a new branch called clade IV or belonged to clade I or III, whereas their nonsymbiotic genes grouped outside the B. canariense branch. These data suggest that the lupin and serradella strains, including the strains from uncultivated L. cosentinii plants, are descendants of strains that most likely were brought from Europe accidentally with lupin and serradella seeds. The observed dominance of B. canariense strains may be related to this species' adaptation to acid soils common in Western Australia and South Africa and, presumably, to their intrinsic ability to compete for nodulation of lupins and serradella.     

Matching the origin of an invasive weed for selection of a herbivore haplotype for a biological control programme

The Florida Everglades have been invaded by an exotic weed fern, Lygodium microphyllum. Across its native distribution in the Old World tropics from Africa to Australasia it was found to have multiple location-specific haplotypes. Within this distribution, the climbing fern is attacked by a phytophagous mite, Floracarus perrepae, also with multiple haplotypes. The genetic relationship between mite and fern haplotypes was matched by an overarching geographical relationship between the two. Further, mites that occur in the same location as a particular fern haplotype were better able to utilize the fern than mites from more distant locations. From a biological control context, we are able to show that the weed fern in the Everglades most likely originated in northern Queensland, Australia/Papua New Guinea and that the mite from northern Queensland offers the greatest prospect for control.     

Removal of post-dispersed seeds in Acacia cyclops thickets under biological control in South Africa

Biological control programmes have been mounted against all invasive Australian acacias with two agent species, Melanterius servulus (seed feeding weevil), and Dasineura dielsi (flower galling midge), being released on Acacia cyclops (rooikrans) in 1991 and 2002 respectively. Both of these agents are prolific and are causing high levels of damage, resulting in reduced seeding capacity of A. cyclops which in turn is expected to limit the invasiveness of the species. As part of an ongoing study to determine the long-term effectiveness of the biological control programme, we measured seed removal rates of A. cyclops by invertebrates and vertebrates, and the composition of granivorous species, to determine how these compare with earlier studies when there was no biological control. Results show that in A. cyclops thickets under biological control, 13% of seeds were removed by invertebrates 59% by rodents and 15% by ground-foraging birds and large mammals within 24 h. The removal rates of seeds with arils intact were double than those of seeds without arils. Camera traps captured 10 vertebrate species comprising six birds and four mammals including Mellivora capensis Storr (Cape ratel) and Raphicerus melanotis Thunberg (Cape grysbok) consuming seeds from stashes. The most frequent visitors were Rhabdomys pumilio (striped mouse), Streptopelia capicola (Cape turtledove) and Cossypha caffra (Cape robin-chat) (33%, 27% and 20% of visits respectively). We conclude that levels of granivory and the composition have not changed substantially with biological agents in the system.     

Best of both worlds: The thermal physiology of Hydrellia egeriae,a biological control agent for the submerged aquatic weed,Egeria densa in South Africa

BioControl - The submerged aquatic weed, Egeria densa Planch. (Hydrocharitaceae) or Brazilian waterweed, is a secondary invader of eutrophic freshwater systems in South Africa, following the...     

石榴巾夜蛾生物学特性及防治试验

描述了石榴巾夜蛾Paralleliastuposa的形态特征、生物学特性,建议用95%敌敌畏乳油500～800倍液防治石榴巾夜蛾,防治效果在938%以上。     

Flightlessness in mayflies and its relevance to hypotheses on the origin of insect flight

Until now, only fully winged mayflies have been known. It has been proposed recently that brachyptery could be a missing link in the development of insect flight, via sailing or skimming aquatic insects. To our knowledge, we report here the first documented case of brachyptery in mayflies. The flightless genus Cheirogenesia is endemic to Madagascar, and the adults skim the water surface. This loss of the flight function has induced important physiological changes, such as a shift from lipids to carbohydrates in the energy reserves used during their adult life. Comparison of wing area of living mayflies with fossil species indicates that brachyptery could have already occurred in early flying insects (in the Permian). We argue that flight loss in Cheirogenesia has been made possible by the lack of fish predation in its natural habitats.     

Using molecular gut content analysis to identify key predators in a classical weed biological control system: a study with Neomusotima conspurcatalis (Lepidoptera: Crambidae)

Predators can severely limit or prevent the establishment of newly introduced biological control agents. Identifying key predators and quantifying their impacts, however, has often proved difficult, especially in cases where the biological control agent has low establishment or detectability. We used molecular gut content analysis to identify predators feeding on Neomusotima conspurcatalis Warren (Lepidoptera: Crambidae), a biological control agent for Lygodium microphyllum (Cavanilles) R Brown (Lygodiaceae), an invasive weed in Florida, USA. We developed DNA primers specific to N. conspurcatalis, and conducted laboratory feeding trials to confirm that agent DNA could be detected in predator guts through PCR amplification. We then released N. conspurcatalis at three L. microphyllum infested sites and one week later conducted field surveys for predators. Our experimental field survey indicated that predatory arthropods were attracted to the immediate locations of N. conspurcatalis releases. A total of 351 predatory arthropods were collected from L. microphyllum infested sites, representing a broad taxonomic range. At least 33% of individuals from all 13 taxa analyzed had fed on N. conspurcatalis. Of the four predator functional groups encountered, spiders were the most numerous, and appeared to be the most important group based on predation rate. Our results can inform the N. conspurcatalis release program, providing guidance on avoiding generating predation hotspots, and facilitating predictions about where and when N. conspurcatalis releases are most likely to result in establishment. Additionally, we highlight the potential of molecular gut content analysis to improve our understanding of the effects of predators on newly introduced classical biological control agents.

     

The stability of host-pathogen interactions of plant disease in relation to biological weed control

A theoretical analysis was conducted to investigate the dynamics of plant-pathogen interactions for biological weed control. Computer simulation showed that the dynamics of plant-pathogen interactions can be determined by the properties of the pathogen. Pathogens with high levels of virulence may exist in nature in low frequencies due to high extinction rates. Pathogens of this type are suitable for the mycoherbicide strategy. Pathogens with a low level of virulence are frequent and may coexist stably with their host. Good candidates for the classical strategy may be the pathogens with intermediate pathogenicity, which maintain a stable interaction and a high control efficiency. The probability of extinction of a pathogen increases when pathogenicity is greater than a critical value at the intermediate range. The regulation of plant populations through reducing host reproductivity and increasing host mortality has similar results.     

Scientific advances in the analysis of direct risks of weed biological control agents to nontarget plants

Research on host specificity testing protocols over the last 10 years has been considerable. Traditional experimental designs have been refined and interpretation of the results is benefiting from an improved understanding of agent behavior. The strengths, weaknesses, and best practice for the different test types are now quite clearly understood. Understanding the concept of fundamental host range (the genetically determined limits to preference and performance) and using this to maximize reliability in predicting field host specificity following release (behavioral expression of the fundamental host range under particular conditions) are still inconsistently understood or adopted despite having been identified as the critical steps in analyzing the threats posed by biological control agents to the agriculture and biodiversity of novel environments. This needs to be consistently understood and applied so the process of testing can follow a recognized process of risk analysis from hazard identification (identifying life stages of the agent that pose a threat and defining their fundamental host range) to uncertainty analysis based on the magnitude (predicted field host specificity following release) and likelihood of threats (predicted actual damage and impact) to nontargets. Modern molecular techniques are answering questions associated with subspecific variation in biological control agents with respect to host use and the chance of host shifts of agents following release. Guidelines for assessment of nontarget impacts need to recognize and adopt such recent developments and emphasize a general increased understanding of the evolution of host choice and the phylogenetic constraints to shifts in host use. This review covers all these recent advances for the first time in one document, highlighting how inconsistent interpretation by biological control practitioners can be avoided.     

Phylogeography of the invasive weed Hypochaeris radicata (Asteraceae): from Moroccan origin to worldwide introduced populations

In an attempt to delineate the area of origin and migratory expansion of the highly successful invasive weedy species Hypochaeris radicata, we analysed amplified fragment length polymorphisms from samples taken from 44 populations. Population sampling focused on the central and western Mediterranean area, but also included sites from Northern Spain, Western and Central Europe, Southeast Asia and South America. The six primer combinations applied to 213 individuals generated a total of 517 fragments of which 513 (99.2%) were polymorphic. The neighbour-joining tree presented five clusters and these divisions were supported by the results of Bayesian analyses: plants in the Moroccan, Betic Sierras (Southern Spain), and central Mediterranean clusters are all heterocarpic. The north and central Spanish, southwestern Sierra Morena, and Central European, Asian and South American cluster contain both heterocarpic (southwestern Sierra Morena) and homocarpic populations (all other populations). The Doñana cluster includes two homocarpic populations. Analyses of fragment parameters indicate that the oldest populations of H. radicata are located in Morocco and that the species expanded from this area in the Late Quaternary via at least three migratory routes, the earliest of which seems to have been to the southwestern Iberian Peninsula, with subsequent colonizations to the central Mediterranean area and the Betic Sierras. Homocarpic populations originated in the southwestern Iberian Peninsula and subsequently spread across north and central Spain, Central Europe and worldwide, where they became a highly successful weed.