J.H. Hoffmann

The population dynamics of an introduced tree, Sesbania punicea, in South Africa, in response to long-term damage caused by different combinations of three species of biological control agents.

Abstract This paper contributes to the relatively sparse literature on the effects of insect herbivory on the population dynamics of plants and is probably unique in that it reports the long-term effects of combinations of three insect herbivore species on the population densities of a moderately long-lived tree species. The tree is Sesbania punicea, a leguminous perennial from South America that has been the target of a biological control programme in South Africa for almost 20 years. Sixteen infestations of the weed have been monitored for periods of up to 10 years to determine changes in the density of the mature, reproductive plants under the influence of different combinations of three biological control agents (i.e. with one, two or three of the agent species present in the weed infestation). The three biological control agents, all weevil species, include Trichapion lativentre, which primarily destroys the flower-buds, Rhyssomatus marginatus, which destroys the developing seeds, and Neodiplogrammus quadrivittatus, whose larvae bore into the trunk and stems of the plants. While T. lativentre occurs throughout the range of the weed in South Africa, the other two species are less mobile, more recent introductions and are largely confined to the vicinity of selected release sites. There has been a significant decline in the density of mature S. punicea in areas where two or more of the agents are established. The decline of the weed has been most evident where N. quadrivittatus is active and particularly so where both of the other two weevil species are also present.

Biological control of invasive alien plants in South Africa: necessity, circumspection, and success

The negative impacts of invasive alien plants on the biodiversity, water supplies, and economy of South Africa are severe. Thousands of people are employed to clear the landscape of these invasive alien species, particularly trees, from conservation and riparian areas. The long-term suppression of the most virulent invasive plants in South Africa will never be possible without the intervention of biological control. Furthermore, the use of specially selected and carefully tested plant-feeding insects, mites, and pathogens as biological control agents is very safe, albeit never completely risk free. We present examples and data from South Africa to support these assertions and to show that biological control is often extremely successful and highly cost effective.

Seed ecology of an invasive alien species, Acacia longifolia (Fabaceae), in Portuguese dune ecosystems.

UNLABELLED PREMISE OF THE STUDY Worldwide, invasive plants threaten biodiversity, by disrupting habitats and ecosystem processes, and cause major economic losses. Invasiveness in plants is frequently associated with prolific production of seeds that accumulate in the soil. Knowledge of the extent and persistence of invasive seed banks helps explain invasion processes and enables management planning. A study of Acacia longifolia, an invasive species in Portuguese dune ecosystems, provides an informative example. • METHODS Seed rain and dispersal (seed traps), the persistence of seeds in the soil (burial), and the extent of seed banks were measured and analyzed. • KEY RESULTS Seed rain is concentrated under the canopy with about 12000 seeds · m(-2) falling annually. The number of seeds in the soil declined with time, with only 30% surviving after 75 mo. Losses were lowest at greater depths. Seed germinability was low (<12%), but viability was high (>85%) for surviving seeds. The seed bank under the canopy was approximately 1500 and 500 seeds · m(-2) in long- and recently invaded stands, respectively. Some seeds were found up to 7 m from the edge of stands, indicating that outside agencies facilitate dispersal. • CONCLUSIONS Acacia longifolia produces large numbers of seeds, some of which are lost through germination, decay, and granivory. The remainder form vast and persistent seed banks that serve as a source of replenishment and make it difficult to control the invader once it is established. Control costs escalate as the duration of an invasion increases, highlighting the urgency of initiating and persevering with control efforts.

photosynthesis and sink activity of wasp-induced galls in acacia pycnantha

Although insect galls are widely known to influence source-sink relationships in plants, the relationship between photosynthesis and gall activity has not been extensively studied. In this study we used 14CO2, photosynthesis, and respiration measurements to examine the capacity of bud galls induced by the wasp Trichilogaster signiventris (Pteromalidae) as carbon sinks in Acacia pycnantha. Galls of this species develop either in vegetative or reproductive buds, depending on the availability of tissues at different times of the year, and effectively eliminate seed production by the plant. Photosynthetic rates in phyllodes subtending clusters of galls were greater than rates in control phyllodes, a result we attributed to photosynthesis compensating for increased carbon demand by the galls. Contrary to previous studies, we found that photosynthesis within galls contributed substantially to the carbon budgets of the galls, particularly in large, mature galls, which exhibited lower specific respiration rates allowing for a net carbon gain in the light. To determine the sink capacity and competitive potential of galls, we measured the proportion of specific radioactivity in galls originating from either vegetative or reproductive buds and found no difference between them. The proportion of the total amount of phyllode-derived 14C accumulated in both clustered and solitary galls was less than that in fruits. Galls and fruits were predominantly reliant on subtending rather than on distant phyllodes for photosynthate. Solitary galls that developed in vegetative buds constituted considerably stronger sinks than galls in clusters on inflorescences where there was competition between galls or fruits for resources from the subtending phyllode. Wasps developing in solitary vegetative galls were correspondingly significantly larger than those from clustered galls. We conclude that, in the absence of inflorescence buds during summer and fall, the ability of the wasps to cause gall formation in vegetative tissues tempers intraspecific competition and substantially increases the availability of plant resources for the development of wasps in such galls.

Biological Control of Australian Acacia Species and Paraserianthes lophantha (Willd.) Nielsen (Mimosaceae) in South Africa

In total, ten agent species have been released in South Africa for the biological control of ten invasive Australian Acacia species and Paraserianthes lophantha (Willd.) Nielsen (Mimosaceae). Besides a single fungal pathogen species which affects both reproductive and vegetative growth of its host plant, Acacia saligna (Labill.) H.L.Wendl., there are nine herbivorous insect species which predominantly suppress the reproductive output of their host plants. These include five seed-feeding weevil species, two flower-galling fly species and two bud-galling wasp species. An indigenous basidiomycete fungus, which causes die-back disease of Acacia cyclops A. Cunn. ex G. Don, has also been investigated. During the last ten years, considerable effort has been directed at searching for new agents in Australia and in collecting additional material to bolster populations of recently-established agents in South Africa. Concurrently, ongoing evaluation studies in South Africa have measured the dynamics of the introduced agents as well as their impact on the vigour and fecundity of their host plants and the extent to which their damage is reducing the density, distribution and invasiveness of the Acacia species. Progress with all of these projects is reviewed.

Biological Control of Cactaceae in South Africa

This review is a summary of developments that have contributed to the success of several biological control programmes against invasive cactus species (Cactaceae) that have been worked on in South Africa over the last 12 years. Six potential biological control agents have been identified for the control of Pereskia aculeata Mil. and molecular studies have identified the origin of the South African P. aculeata population. Host-specificity testing is now required for the three most promising of these agents. The successful biological control programme against Opuntia stricta (Haw.) Haw. has resulted in a change in management strategies against this weed in the Kruger National Park and the control of O. stricta is now almost entirely reliant on biological control. Taxonomic problems associated with the identification of Cylindropuntia fulgida var. fulgida (Engelm.) F.M.Knuth var. fulgida have been resolved and an appropriate cochineal insect (Hemiptera: Dactylopiidae) biotype has been released, resulting in substantial declines in Cyl. fulgida var. fulgida populations. A long-term monitoring programme has been initiated to evaluate the progress of this new cochineal insect biotype. The Harrisia mealybug, Hypogeococcus pungens Granara de Willink (Hemiptera: Pseudococcidae), which was originally released on Harrisia martinii (Labour.) Britton & Rose has been collected and redistributed onto Cereus jamacaru DC., where it reduces fruit production and leads to the death of both seedlings and large plants. Resolving taxonomic problems to ensure the correct identification of plant species and the most appropriate biological control biotypes have been key issues that have led to the successful control of several cactaceous weed species in South Africa.

Some Perspectives on the Risks and Benefits of Biological Control of Invasive Alien Plants in the Management of Natural Ecosystems

Globally, invasions by alien plants are rapidly increasing in extent and severity, leading to large-scale ecosystem degradation. Weed biological control offers opportunities to arrest or even reverse these trends and, although it is not always effective or appropriate as a management strategy, this practice has an excellent record of safety and many notable successes over two centuries. In recent years, growing concerns about the potential for unintended, non-target damage by biological control agents, and fears about other unpredictable effects on ecosystems, have created an increasingly demanding risk-averse regulatory environment. This development may be counter-productive because it tends to overemphasize potential problems and ignores or underestimates the benefits of weed biological control; it offers no viable alternatives; and it overlooks the inherent risks of a decision not to use biological control. The restoration of badly degraded ecosystems to a former pristine condition is not a realistic objective, but the protection of un-invaded or partial restoration of invaded ecosystems can be achieved safely, at low cost and sustainably through the informed and responsible application of biological control. This practice should therefore be given due consideration when management of invasive alien plants is being planned. This discussion paper provides a perspective on the risks and benefits of classical weed biological control, and it is aimed at assisting environmental managers in their deliberations on whether or not to use this strategy in preference, or as a supplement to other alien invasive plant control practices.

Conservation of the fynbos biome in the Cape Floral Region: the role of biological control in the management of invasive alien trees

Fynbos is a vegetation type in the Cape Floral Region (CFR), at the southern tip of Africa. Portions of the CFR are recognised as a ‘serial’ World Heritage site and acclaimed by UNESCO as the world’s ‘hottest hot spot’ for plant species richness and endemism. Habitat degradation and species losses through human intrusion in the CFR include transformations brought about by introduced invasive alien tree species. Since 1970, ten invasive tree species in the fynbos biome have been subjected to biological control, namely: six Acacia species and Paraserianthes lophantha (Mimosaceae), Hakea sericea (Proteaceae) and Leptospermum laevigatum (Myrtaceae), all from Australia, and Sesbania punicea (Fabaceae) from South America. A total of 19 species have been deployed as biological control agents, including nine weevil species (eight Curculionidae and one species in the family Brentidae: Apioninae), a seed-feeding moth species (Lepidoptera: Carposinidae), two species of bud-gallers (Hymenoptera: Pteromalidae), two species of flower-gallers (Diptera: Cecidomyiidae), and a gall-forming rust fungus (Uredinales: Pileolariaceae). Most of these agents primarily reduce seed production, directly or indirectly, but some also cause die-back and mortality of their host plants. The overall result, often in combination with mechanical clearing and herbicide applications, has been a substantial decline in the abundance and/or aggressiveness of most of the targeted host-plants. In this review, four representative but contrasting case studies are used to show that biological control is an indispensible supplement to other management practices for long-term conservation of the remnants of the fynbos biome.

Biocontrol of a perennial legume, Sesbania punicea, using a florivorous weevil, Trichapion lativentre: weed population dynamics with a scarcity of seeds

SummaryThe establishment in South Africa of a florivorous, apionid weevil, Trichapion lativentre, on Sesbania punicea, a leguminous weed of South American origin, has reduced seed production of the plants by >98%. Surveys of the age structure and density of plants in infestations of S. punicea throughout South Africa have shown that the rate of recruitment of seedlings has drastically declined within a few years in many areas, due to the weevils. However, there has unexpectedly not been a corresponding decline in the density of mature plants in extant infestations of S. punicea. In spite of this, T. lativentre has curtailed the rate of spread of the weed into uninvaded habitats and has impeded reinvasion into areas cleared of infestations by mechanical means or by another complimentary biocontrol agent.

Temporal changes in the impacts on plant communities of an invasive alien tree, Acacia longifolia

Studies on the impacts of invasive plants are common but most are short term and fail to consider the temporal context of invasion. The present work investigated particular invasions on two different time scales and asked: (1) Do the impacts of an invasive tree on plant communities change after decades of invasion? (2) Are patterns of impacts dynamic over a period of five years? (3) Can multiple parameters reveal impacts that are unnoticed when single-value parameters are measured alone? Contrasting plant communities (long invaded, recently invaded and non-invaded) of a Portuguese coastal dune were compared during a 5-year period to assess the impacts of the invasive Acacia longifolia. Plant diversity, richness, cover, plant traits, indicator species analysis, similarity between areas and species turnover were analysed to reveal spatial and temporal patterns of change. Native species richness declined in both invaded areas. As invasion time increased, species shared with natural habitats decreased along with native plant cover, diversity and species turnover. Many species typical of dunes were replaced by generalists and exotics species, and richness of species with some level of nitrophily increased, particularly in recently invaded areas. Life form spectrum was radically transformed in invaded areas with several small-sized life forms being replaced by one single microphanerophyte. Germination of A. longifolia was greatest in long invaded areas. Findings show that modification of plant communities intensifies with invasion time and that invasion promotes a diversity of structural and functional changes which are dynamic over a few years (medium-term scale) but which tend to stabilize after several decades (long-term scale). An analysis of parameters in combination revealed changes that were not apparent when the same parameters were considered separately.