Abstract
Old-growth tropical forests are widely believed to be immune to the establishment of alien species. Collections from tropical regions throughout the world, however, have established that this generalization does not apply to inbreeding host generalist bark and ambrosia beetles. Scolytine saproxylophages are readily spread by shipping, inbreeders can easily establish new populations, and host generalists readily find new breeding material, apparently regardless of stage of forest succession. Consequently, many inbreeding scolytines are globally distributed and abundant in all forest types, often being among the dominant species in their wood-borer communities. We report the recent introductions to lower Central America of two Old World inbreeding ambrosia beetles: Xylosandrus crassiusculus, which breeds primarily in smaller diameter trunks, small branches, and twigs, and Xyleborinus exiguus, which is apparently not size selective. We also document the establishment of Euwallacea fornicatus in the region, known previously from a single collection in Panama. Xylosandrus crassiusculus and E. fornicatus are notorious agricultural and forestry pests, as are several previously established alien species in the region. Studying the spread of species such as these three new arrivals into millions of years-old faunas could help us to understand if the saproxylic communities of old-growth tropical forests are peculiarly vulnerable to invasion.References
Atkinson, T.H. & Equihua, A. (1986) Biology of bark and ambrosia beetles (Coleoptera: Scolytidae and Platypodidae) of a tropical rain forest in southeastern Mexico with an annotated checklist of species. Annals of the Entomological Society of America, 79, 414–423.
Atkinson, T.H., Foltz, J.L. & Wilkinson, R.C. (1988) Xylosandrus crassiusculus (Moltschultsky), an Asian ambrosia beetle recently introduced into Florida (Coleoptera: Scolytidae). Florida Department of Agriculture and Consumer Services, Division of Plant Industry, Entomology Circular, 310, 1–4.
Atkinson, T.H., Foltz, J.L., Wilkinson, R.C. & Mizell, R.F. (2005) Featured creature: granulate or Asian ambrosia beetle Xylosandrus crassiusculus (Motschulsky). University of Florida Institute of Food and Agricultural Sciences, and Florida Department of Agriculture and Consumer Services. http://creatures.ifas.ufl.edu/trees/asian_ambrosia_beetle.htm (accessed 6 June 2007).
Basset, Y., Aberlenc, H.-P., Barrios, H., Curletti, G., BJrenger, J.-M., Vesco, J.-P., Causse, P., Haug, A., Hennion, A.-S., Lesobre, L., Marques, F. & O'Meara, R. (2001) Stratification and diel activity of arthropods in a lowland rain forest in Gabon. Biological Journal of the Linnean Society, 72, 585–607.
Beaver, R.A. (1979) Host specificity of temperate and tropical animals. Nature, 281, 139–141.
Beaver, R.A. (1988) Insect-fungus relationships in the bark and ambrosia beetles. In: Wilding, N., Collins, N.M., Hammond, P.M. & Webber, J.F. (Eds.) Insect-Fungus Interactions. Academic Press, London, pp. 121–143.
Beaver, R.A. (2005) A remarkable new species of Cyclorhipidion Hagedorn, and new records of bark and ambrosia beetles from Gabon (Coleoptera: Curculionidae, Scolytinae and Platypodinae). Entomologists Monthly Magazine, 141, 113–119.
Bright, D.E. & Skidmore, R.E. (2002) A Catalog of Scolytidae and Platypodidae (Coleoptera), Supplement 2 (1995–1999). NRC Research Press, Ottawa, 523 pp.
Brockerhoff, E.G., Bain, J., Kimberley, M. & KníÒek, M. (2006) Interception frequency of exotic bark and ambrosia beetles (Coleoptera : Scolytinae) and relationship with establishment in New Zealand and worldwide. Canadian Journal of Forest Research, 36, 289–298.
Browne, F.G. (1961) The biology of Malayan Scolytidae and Platypodidae. Malayan Forest Record, 22, 1–255.
Browne, F.G. (1963) Notes on the habits and distribution of some Ghanaian bark beetles and ambrosia beetles (Coleoptera: Scolytidae and Platypodidae). Bulletin Of Entomological Research, 54, 229–266.
CAB International. (2005a) Xyleborus fornicatus (Eichhoff, 1868) [original text by R.A. Beaver]. CAB International – Crop Pest Compendium. www.cabicompendium.org/cpc (commercial, electronic version not seen).
CAB International. (2005b) Xylosandrus crassiusculus (Motschulsky) [original text by R.A. Beaver]. CAB International – Crop Pest Compendium. www.cabicompendium.org/cpc (commercial, electronic version not seen).
Chornesky, E.A., Bartuska, A.M., Aplet, G.H., Britton, K.O., Cummings-Carlson, J., Davis, F.W., Eskow, J., Gordon, D.R., Gottschalk, K.W., Haack, R.A., Hansen, A.J., Mack, R.N., Rahel, F.J., Shannon, M.A., Wainger, L.A. & Wigley, T.B. (2005) Science priorities for reducing the threat of invasive species to sustainable forestry. Bioscience, 55, 335–348.
Colwell, R. & Longino, J.T. (2006) Project ALAS Arthropods of La Selva. http://viceroy.eeb.uconn.edu/ALAS/ALAS.html (accessed 6 June 2007).
Connell, J.H. (1978) Diversity in tropical rain forests and coral reefs. Science, 199, 1302–1310.
Elton, C.S. (1958) The ecology of invasions by animals and plants. Methuen, London, UK,pp.
Haack, R.A. (2001) Intercepted Scolytidae (Coleoptera) at U.S. ports of entry: 1985–2000. Integrated Pest Management Reviews, 6, 253–282.
Haack, R.A. (2006) Exotic bark- and wood-boring Coleoptera in the United States: recent establishments and interceptions. Canadian Journal of Forest Research, 36, 269–288.
Hanson, P.E. (1992) The Costa Rican Malaise Trap Network. http://www.phorid.net/phoridae/malaise.html (accessed 6 June 2007).
Hanson, P.E. & Gauld, I.D. (1995) The Hymenoptera of Costa Rica. Oxford, Oxford University Press, 893 pp.
Herbold, B. & Moyle, P.B. (1986) Introduced species and vacant niches. The American Naturalist, 128, 751–760.
Hooper, D.U., Chapin, F.S., III, Ewel, J.J., Hector, A., Inchausti, P., Lavorel, S., Lawton, J.H., Lodge, D.M., Loreau, M., Naeem, S., Schmid, B., Setälä, H., Symstad, A.J., Vandermeer, J. & Wardle, D.A. (2005b) Effects of biodiversity on ecosystem functioning: a consensus of current knowledge. Ecological Monographs, 75, 3–35.
Humble, L.M. & Allen, E.A. (2006) Forest biosecurity: alien invasive species and vectored organisms. Canadian Journal of Plant Pathology, 28, S256–S269.
Huston, M. (1979) A general hypothesis of species diversity. The American Naturalist, 113, 81–101.
Huston, M.A. (1994) Biological Diversity. The coexistence of species on changing landscapes. Cambridge University Press, Cambridge, 681 pp.
Jordal, B.H. (2002) Elongation Factor 1 alpha resolves the monophyly of the haplodiploid ambrosia beetles Xyleborini (Coleoptera : Curculionidae). Insect Molecular Biology, 11, 453–465.
Jordal, B.H., Beaver, R.A. & Kirkendall, L.R. (2001) Breaking taboos in the tropics: inbreeding promotes colonization by wood-boring beetles. Global Ecology and Biogeography, 10, 345–357.
Jordal, B.H., Normark, B.B. & Farrell, B.D. (2000) Evolutionary radiation of an inbreeding haplodiploid beetle lineage (Curculionidae, Scolytinae). Biological Journal of the Linnean Society, 71, 483–499.
Kalshoven, L.G.E. (1924) Een geval van aantasting van kina door boeboek-keertjes (Xyleborus sp. div). [An attack of Cinchona by shothole borers (Xyleborus spp.)]. Mededeelingen van het Instituut voor Plantenziekten, 65, 25.
Kalshoven, L.G.E. (1958) Studies on the biology of Indonesian Scolytoidea. I. Xyleborus fornicatus Eichh. as a primary and secondary shot-hole borer in Java and Sumatra. Entomologische Berichten, 18, 147–160.
Kirkendall, L.R. (1983) The evolution of mating systems in bark and ambrosia beetles (Coleoptera: Scolytidae and Platypodidae). Zoological Journal of the Linnean Society, 77, 293–352.
Kirkendall, L.R. (1993) Ecology and evolution of biased sex ratios in bark and ambrosia beetles (Scolytidae). In: Wrensch, D.L. & Ebbert, M.A. (Eds.) Evolution and Diversity of Sex Ratio: Insects and Mites. Chapman and Hall, New York, pp. 235–345.
Kovach, J. & Gorsuch, C.S. (1985) Survey of the ambrosia beetle species infesting South Carolina peach orchards and a taxonomic key for the most common species. Journal of Agricultural Entomology, 2, 238–247.
Kühnholz, S., Borden, J.H. & Uzunovic, A. (2001) Secondary ambrosia beetles in apparently healthy trees: adaptations, potential causes and suggested research. Integrated Pest Management Reviews, 6, 209–219.
LaBonte, J.R., Mudge, A.D. & Johnson, K.J.R. (2005) Nonindigenous woodboring coleoptera (Cerambycidae, Curculionidae : Scolytinae) new to Oregon and Washington, 1999–2002: Consequences of the intracontinental movement of raw wood products and solid wood packing materials. Proceedings of the Entomological Society of Washington, 107, 554–564.
Leponce, M. & Basset, Y. (2006) IBISCA Panama 2003–2004: Spatio-temporal distribution of arthropods in a tropical rainforest. Royal Belgian Institute of Natural Sciences. http://www.naturalsciences.be/cb/ants/projects/ibisca-panama2003-2004.htm (accessed 6 June 2007).
Maeto, K., Fukuyama, K. & Kirkon, L.K. (1999) Edge effects on ambrosia beetle assemblages in a lowland rain forest, bordering oil palm plantations, in Peninsula Malaysia. Journal of Tropical Forest Science, 11, 537–547.
Noguera-Martinez, F.A. & Atkinson, T.H. (1990) Biogeography and biology of bark and ambrosia beetles (Coleoptera: Scolytidae and Platypodidae) of a mesic montane forest in Mexico, with an annotated checklist of species. Annals of the Entomological Society of America, 83, 453-466.
Rabaglia, R.J. (2003) Annotated list of the bark and ambrosia beetles (Coleoptera : Scolytidae) of Maryland, with new distributional records. Proceedings of the Entomological Society of Washington, 105, 373–379.
Rabaglia, R.J., Dole, S.A. & Cognato, A.I. (2006) Review of American Xyleborina (Coleoptera : Curculionidae : Scolytinae) occurring North of Mexico, with an illustrated key. Annals of the Entomological Society of America, 99, 1034–1056.
Roberts, H. (1969) Forest insects of Nigeria with notes on their biology and distribution. Commonwealth Forestry Institute Paper, 44, 1–206.
Samuelson, G.A. (1981) A synopsis of Hawaiian Xyleborini (Coleoptera: Scolytidae). Pacific Insects, 23, 50–92.
Schedl, K.E. (1962) Scolytidae und Platypodidae Afrikas. Band 2. Familie Scolytidae. Rev. Entomol. Moambique, 5, 1–594.
Simberloff, D. (1981) Community effects of introduced species. In: Nitecki, T.H. (Ed.) Biotic crises in ecological and evolutionary time. Academic Press, New York, pp. 53–81.
Springate, N.D. & Basset, Y. (2004) IBISCA: 2003–2005, Panama: progress report. Bulletin of the British Ecological Society, 35, 21–23.
Stachowicz, J.J. & Byrnes, J.E. (2006) Species diversity, invasion success, and ecosystem functioning: disentangling the influence of resource competition, facilitation, and extrinsic factors. Marine Ecology-Progress Series, 311, 251–262.
Thomas, M.C. (2005) Two Asian ambrosia beetles recently established in Florida (Curculionidae: Scolytinae). Florida Department of Agriculture and Consumer Services, Division of Plant Industry, Pest Alert. http://www.doacs.state.fl.us/pi/enpp/ento/twonewxyleborines.html (accessed 6 June 2007).
Wood, S.L. (1982) The bark and ambrosia beetles of North and Central America (Coleoptera: Scolytidae), a taxonomic monograph. Great Basin Naturalist Memoirs, 6, 1–1359.
Wood, S.L. (2007) The Bark and Ambrosia Beetles of South America (Coleoptera, Scolytidae). Monte L. Bean Life Science Museum, Brigham Young University, Provo, Utah, USA, 900 pp + 230 plates pp.
Wood, S.L. & Bright, D.E. (1992) A Catalog of Scolytidae and Platypodidae (Coleoptera), Part 2: Taxonomic Index. Great Basin Naturalist Memoirs, 13, 1–1553.