Proceedings of the XI International Symposium on Biological Control of Weeds
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Politics and ecology in the management of alien invasive woody trees: the pivotal role of biological control agents that diminish seed production V.C. Moran,1 J.H. Hoffmann1 and T. Olckers2 Summary 1. Biological control agents have been established on a total of 22 species of alien, invasive woody trees in Australia (and via Australia in Thailand, Vietnam and Malaysia), South Africa, the United States of America and Hawaii. Fifteen of these tree species (mostly Australian natives) are in South Africa. 2. Of the 39 agent species, 38 are insects (mostly weevils) and one is a pathogen. About 50% of these agents have been released in South Africa. Approximately two-thirds of these agents reduce seed production, directly or indirectly. 3. Usually, pre-dispersal seed mortality will not bring about a reduction in the density of invasive plants because weed populations can maintain near-saturation densities in spite of impressive depri- vations of their seeds by biocontrol agents. However, when humans intervene to clear adult plants, and where recruited seedlings are systematically and repeatedly destroyed, even modest levels of pre-dispersal seed mortality can translate into substantial savings for weed-control managers. Fewer seeds mean easier management through lower recruitment rates, fewer seedlings, and slower dispersal. 4. In the context of the politics and ecology of alien invasive tree control, agents that reduce seed production can also prove to be decisive in resolving conflicts of interest, and can become key elements in rehabilitation and restoration processes. They have become the first-line of defence in the successful management of alien invasive tree species. 5. We illustrate these points with reference to three very different cases, namely biological control against Sesbania punicea, Acacia cyclops and Acacia pycnantha in South Africa. In each case, agents that reduce seeding have played a pivotal role in the biological control campaigns. Keywords: biological control, invasive trees, management, seed-attacking agents. Introduction South Africa). Thirty-eight insect species (and one rust species) have become established as biological control Only about 22 species of alien invasive woody trees agents on trees. About 50% of these agents are estab- have been targeted for biological control. Of these, 3 lished in South Africa, mostly deployed against alien are in the United States of America, 4 in Australia (or Acacia species from Australia (Table 1). About two- via Australia) and 15 in South Africa (Table 1; though thirds of these agents reduce seeding of their host plants an exact species count is not possible because Prosopis either directly through feeding or laying on, or devel- has formed hybrid communities in Australia and in oping in, the seeds, or indirectly by damaging the repro- ductive parts of the plant, or variously debilitate the plants, and thus inhibit seed formation. 1 Zoology Department, University of Cape Town, Rondebosch, 7701, Much has been published on seed dynamics (Harper South Africa. 1977, Crawley 1997) and on the dynamics of 2 Plant Protection Research Institute, Cedara, Hilton, 3245, South Africa. plant–herbivore interactions (Crawley 1983). In the Corresponding author: V.C. Moran <[email protected]>. 434 Seed-attacking agents in biological control of trees Table 1. Alien invasive tree species (and their countries of origin) that have been deliberately subjected to biological control. (Shrubs that less commonly become small trees, and large cactus species, are excluded.) The agents that have become established on each of the target tree species, and the countries of introduction, are listed. The data are mostly extracted from Julien and Griffiths (1998), but modified according to more recent information, indi- cated by superscript numerals in the table, as follows: (1) van Klinken, R.D. et al. (2002) and van Klinken, R.D. (2003, pers. comm.); (2) Adair, R. J. (3, 4, 5) Impson, F.A.C.; (6) Olckers, T. (2003, Report of the Plant Protection Research Institute, Pretoria); (7) Center, T.D. et al. (2000); (8) Wheeler, G.S. et al. (2001). Tree species Agents established Countries of introduction Acacia nilotica ssp. indica Bruchidius sahlbergi Australia Prickly acacia (India) (Coleoptera: Bruchidae) Mimosa pigra Acanthoscelides puniceus Australia; to Thailand Giant sensitive plant (tropical America) (Coleoptera: Bruchidae) Acanthoscelides quadridentatus Via Australia (not established); (Coleoptera: Bruchidae) to Thailand, Vietnam Carmenta mimosa Australia; to Vietnam, (Lepidoptera: Sesiidae) Malaysia Chlamisus mimosae Australia (Coleoptera: Chrysomelidae) Coelocephalapion aculeatum Australia (Coleoptera: Apionidae) Coelocephalapion pigrae Australia (Coleoptera: Apionidae) Malacorhinus irregularis Australia (Coleoptera: Chrysomelidae) Neurostrota gunniella Australia (Lepidoptera: Gracillariidae) Parkinsonia aculeata Mimosestes ulkei Australia Palo verde (tropical America) (Coleoptera: Bruchidae) Penthobruchus germaini Australia (Coleoptera: Bruchidae) Rhinacloa callicrates Australia (Hemiptera: Miridae) Prosopis spp. (various hybrids including pallida and velutina) Algarobius prosopis Australia Mesquite (North and Central America) (Coleoptera: Bruchidae) Evippe sp. #11 Australia (Lepidoptera: Gelechiidae) Prosopidopsylla flava 1 Australia (Hemiptera: Psyllidae) Acacia cyclops Dasineura dielsi 2 South Africa Rooikrans (Australia) (Diptera: Cecidomyiidae) Melanterius servulus South Africa (Coleoptera: Curculionidae) Acacia dealbata Melanterius maculatus 3 South Africa Silver wattle (Australia) (Coleoptera: Curculionidae) Acacia decurrens Melanterius maculatus 4 South Africa Green wattle (Australia) (Coleoptera: Curculionidae) Acacia longifolia Melanterius ventralis South Africa Long leaved wattle (Australia) (Coleoptera: Curculionidae) Trichilogaster acaciaelongifoliae South Africa (Hymenoptera: Pteromalidae) Acacia mearnsii Melanterius maculatus South Africa Black wattle (Australia) (Coleoptera: Curculionidae) Acacia melanoxylon Melanterius acaciae South Africa Australian blackwood (Australia) (Coleoptera: Curculionidae) Acacia pycnantha Trichilogaster sp. South Africa Golden wattle (Australia) (Hymenoptera: Pteromalidae) Acacia saligna Melanterius compactus 5 South Africa Port Jackson willow (Australia) (Coleoptera: Curculionidae) Uromycladium tepperianum South Africa (Fungus: Uredinales) Hakea gibbosa Erytenna consputa South Africa Rock hakea (Australia) (Coleoptera: Curculionidae) 435 Proceedings of the XI International Symposium on Biological Control of Weeds Table 1. (Continued) Alien invasive tree species (and their countries of origin) that have been deliberately subjected to biological control. (Shrubs that less commonly become small trees, and large cactus species, are excluded.) The agents that have become established on each of the target tree species, and the countries of introduction, are listed. The data are mostly extracted from Julien and Griffiths (1998), but modified according to more recent information, indicated by superscript numerals in the table, as follows: (1) van Klinken, R.D. et al. (2002) and van Klinken, R.D. (2003, pers. comm.); (2) Adair, R. J. (3, 4, 5) Impson, F.A.C.; (6) Olckers, T. (2003, Report of the Plant Protection Research Institute, Pretoria); (7) Center, T.D. et al. (2000); (8) Wheeler, G.S. et al. (2001). Tree species Agents established Countries of introduction Hakea sericea Carposina autologa South Africa Silky hakea (Australia) (Lepidoptera: Carposinidae) Cydmaea binotata South Africa (Coleoptera: Curculionidae) Erytenna consputa South Africa (Coleoptera: Curculionidae) Leptospermum laevigatum Dasineura sp. South Africa Australian myrtle (Australia) (Diptera: Cecidomyiidae) Parectopa thalassias South Africa (Lepidoptera: Gracillariidae) Paraserianthes lophantha Melanterius servulus South Africa Crested wattle (tropics and subtropics) (Coleoptera: Curculionidae) Prosopis spp. (various hybrids including glandulosa and velutina) Algarobius prosopis South Africa Mesquite (North America) (Coleoptera: Bruchidae) Neltumius arizonensis South Africa (Coleoptera: Bruchidae) Sesbania punicea Neodiplogrammus quadrivittatus South Africa Red sesbania (Argentina, Brazil, Uruguay) (Coleoptera: Curculionidae) Rhyssomatus marginatus South Africa (Coleoptera: Curculionidae) Trichapion lativentre South Africa (Coleoptera: Apionidae) Solanum mauritianum Gargaphia decoris 6 South Africa Bugweed (South America) (Hemiptera: Tingidae) Melaleuca quinquenervia Oxyops vitiosa 7 Florida, U.S.A Paper-bark tree (Australia) (Coleoptera: Curculionidae) Myrica faya Caloptilia nr schinella Hawaii , U.S.A Fire tree (Azores, Madeira, Canary Islands) (Lepidoptera: Gracillariidae) Schinus terebinthifolius Episimus utilus Hawaii, U.S.A. Brazilian pepper tree (South America) (Lepidoptera: Tortricidae) Lithraeus atronotatus Hawaii, U.S.A (Coleoptera: Bruchidae) Metastigmus transvaalensis 8 Florida, U.S.A (Hymenoptera: Torymidae) context of biological control of weeds, the majority of of the seedlings they generated. Weed populations can studies quantify the actual damage inflicted on the often persist in spite of the destruction of high propor- target plants by the biological control agents (e.g. the tions of their seeds by biological control agents. number of shoots destroyed, or the number of feeding The situation is very different where humans inter- holes in the leaves, and so on). Relatively few studies vene to harvest (see Crawley 1997) or to clear parent quantify the impact of the agents