Evidence Of Competition Between Two Canopy Ant Species: Is Aggressive Behavior Innate Or Shaped By A Competitive Environment?

Competition occurs in all ecological communities, although it has not always been experimentally tested as a structuring force in the distribution of species. We tested the hypothesis that the aggressiveness exhibited by Camponotus rufipes changes according to the pressures of a competitive environment. This is a dominant species in the montane forest of the Itacolomi State Park, Brazil, where Camponotus sericeiventris does not occur. Using bait traps in a field site where both species occur, (Juiz de Fora site) we showed that C. sericeiventris was able to remove C. rufipes workers at the same bait. In the laboratory, we used dyadic encounters to test workers from both species taken from colonies found in areas where both occur and where only C. rufipes was found. Camponotus rufipes from Itacolomi fought significantly less and was killed during the first few minutes in 60% of the events. On the other hand, the workers that co-existed with C. sericeiventris in the field were more aggressive, but less efficient fighters than the latter. This investigation demonstrated existence of competition between C. rufipes and C. sericeiventris, and also the lower aggressiveness of C. rufipes' individuals that did not co-exist in the field with C. sericeiventris. © 2012 Ndia Barbosa do Esprito Santo et al.Hlldobler, B., Wilson, E.O., (1990) The Ants, , Cambridge, Mass, USA The Belknap Press of Harvard University PressMacArthur, R.H., Levins, R., The limiting similarity, convergence, and divergence of coexisting species (1967) American Naturalist, 101, pp. 377-385Gotelli, N.J., Ellison, A.M., Assembly rules for New England ant assemblages (2002) Oikos, 99 (3), pp. 591-599. , DOI 10.1034/j.1600-0706.2002.11734.xWilson, E.O., (1971) The Insect Societies, , Cambridge, Mass, USA Harvard University PressDavidson, D.W., Resource discovery versus resource domination in ants: A functional mechanism for breaking the trade-off (1998) Ecological Entomology, 23 (4), pp. 484-490. , DOI 10.1046/j.1365-2311.1998.00145.xRoom, P.M., The relative distribution of ant species in Ghanas cocoa farms (1971) Journal of Animal Ecolog, 40, pp. 735-751Leston, D., The ant mosaic, tropical tree crops and the limiting of pests and diseases (1973) Pest Articles and News Summaries, 19, pp. 311-341Leston, D., A neotropical ant mosaic (1978) Annals of the Entomological Society of America, 71, pp. 649-653Majer, J.D., Delabie, J.H.C., Smith, M.R.B., Arboreal ant community patterns in Brazilian cocoa farms (1994) Biotropica, 26 (1), pp. 73-83Holldobler, B., Lumsden, C.J., Territorial strategies in ants (1980) Science, 210 (4471), pp. 732-739Savolainen, R., Vepsalainen, K., A competition hierarchy among boreal ants: Impact on resource partitioning and community structure (1988) Oikos, 51 (2), pp. 135-155Del-Claro, K., Oliveira, P.S., Ant-homoptera interactions in a neotropical savanna: The honeydew-producing treehopper, Guayaquila xiphias (Membracidae), and its associated ant fauna on Didymopanax vinosum (Araliaceae) (1999) Biotropica, 31 (1), pp. 135-144Orr, M.R., Dahlsten, D.L., Benson, W.W., Ecological interactions among ants in the genus Linepithema, their phorid parasitoids, and ant competitors (2003) Ecological Entomology, 28 (2), pp. 203-210. , DOI 10.1046/j.1365-2311.2003.00506.xFonseca, R.C., Diehl, E., Riqueza de formigas (Hymenoptera: Formicidae) epigéicas em povoamentos de Eucalyptus spp. (Myrtaceae) de diferentes idades no Rio Grande do Sul, Brasil (2004) Revista Brasileira de Entomologia, 48, pp. 95-100Peternelli, E.F.O., Della Lucia, T.M.C., Martins, S.V., Espécies de formigas que interagem com sementes de Mabea fistulifera Mart. (Euphorbiaceae) (2004) Revista Rvore, 28, pp. 733-738Lutinski, J.A., Garcia, F.R.M., Anlise faunstica de Formicidae (Hymenoptera: Apocrita) em ecossistema degradado no municpio de Chapec, Santa Catarina (2005) Biotemas, 18, pp. 73-86Santos, M.S., Louzada, J.N.C., Dias, N., Zanetti, R., Delabie, J.H.C., Nascimento, I.C., Litter ants richness (Hymenoptera, Formicidae) in remnants of a semi-deciduous forest in the Atlantic rain forest, Alto do Rio Grande region, Minas Gerais, Brazil (2006) Iheringia - Serie Zoologia, 96 (1), pp. 95-101. , http://www.scielo.br/pdf/isz/v96n1/a17v96n1.pdf, DOI 10.1590/S0073-47212006000100017Esprito Santo, N.B., (2008) Assembléia de Formigas Do Parque Estadual Do Itacolomi (MG) e Relaes Intra- E Interespecficas Entre Espécies Dominantes, M.S. Thesis, , Ps-Graduao em Comportamento e Biologia Animal/Departamento de Biologia Animal/Universidade Federal de Juiz de ForaYamamoto, M., Del-Claro, K., Natural history and foraging behavior of the carpenter ant Camponotus sericeiventris Guérin, 1838 (Formicinae, Campotonini) in the Brazilian tropical savanna (2008) Acta Ethologica, 11 (2), pp. 55-65Campos, R.I., Scares, J.P., Martins, R.P., Ribeiro, S.P., Effect of habitat structure on ant assemblages (Hymenoptera: Formicidae) associated to two pioneer tree species (2006) Sociobiology, 47 (3), pp. 721-738Fujaco, M.A.G., (2007) Influncia Dos Diferentes Tipos de Substrato e Geomorfologia Na Distribuio Espacial e Arquitetnica Do Gnero Eremanthus No Parque Estadual Do Itacolomi, Ouro Preto/MG, M.S. Thesis, , Ps-Graduao em Evoluo Crustal e Recursos Naturais/Departamento de Geologia/Universidade Federal de Ouro PretoBanco de Dados Climticos do BrasilEmbrapa Monitoramento por Satélite e EsalqUSP, , 2007, http://www.bdclima.cnpm.embrapa.br/resultados/balanco.php?UF=spCOD= 450Altmann, J., Observational study of behavior: Sampling methods (1974) Behaviour, 49 (34), pp. 227-267Ribeiro, S.P., Soares, J.P., Campos, R.I., Martins, R.P., Insect herbivores species associated to pioneer tree species: Contrasting within forest and ecotone canopy habitats (2008) Revista Brasileira de Zoocincias, 10, pp. 237-248Lopes, J.F.S., Hughes, W.O.H., Camargo, R.S., Forti, L.C., Larval isolation and brood care in Acromyrmex leaf-cutting ants (2005) Insectes Sociaux, 52 (4), pp. 333-338. , DOI 10.1007/s00040-005-0816-yErrard, C., Hefetz, A., Label familiarity and discriminatory ability of ants reared in mixed groups (1997) Insectes Sociaux, 44 (3), pp. 189-198. , DOI 10.1007/s000400050040Santos, J.C., Del-Claro, K., Ecology and behaviour of the weaver ant Camponotus (Myrmobrachys) senex (2009) Journal of Natural History, 43 (2324), pp. 1423-1435Ferreira Brando, C.R., Sequential ethograms along colony development of Odontomachus affinis Guérin (Hymenoptera, Formicidae, Ponerinae) (1983) Insectes Sociaux, 30 (2), pp. 193-203Santos, J.C., Yamamoto, M., Oliveira, F.R., Del-Claro, K., Behavioral repertory of the weaver ant Camponotus (Myrmobrachys) senex (Hymenoptera: Formicidae) (2005) Sociobiology, 46 (1), pp. 27-37Wilson, E.O., (1975) Sociobiology, , Cambridge, Mass, USA Belknap Press of Harvard University PressLangen, T.A., Tripet, F., Nonacs, P., The red and the black: Habituation and the dear-enemy phenomenon in two desert Pheidole ants (2000) Behavioral Ecology and Sociobiology, 48 (4), pp. 285-292Delsinne, T., Roisin, Y., Leponce, M., Spatial and temporal foraging overlaps in a Chacoan ground-foraging ant assemblage (2007) Journal of Arid Environments, 71 (1), pp. 29-44. , DOI 10.1016/j.jaridenv.2007.02.007, PII S0140196307000705Carroll, C.R., Janzen, D.H., Ecology of foraging by ants (1973) Annual Review of Ecology and Systematics, 4, pp. 231-257Byk, J., Del-Claro, K., Nectar- and pollen-gathering Cephalotes ants provide no protection against herbivory: A new manipulative experiment to test ant protective capabilities (2010) Acta Ethologica, 13 (1), pp. 33-38Jaffé, K., Snchez, C., On the nestmate-recognition system and territorial marking behaviour in the ant Camponotus rufipes (1984) Insectes Sociaux, 31 (3), pp. 302-315Lucas, C., (2002) Tude des Bases Chimiques et Comportamentales de la Formation du Visa Colonial Chez les Ponérines du Genre Pachycondyla, Thse Docteur, , Paris, France Biologie du Comportement, Universit Paris XIHlldobler, B., Wilson, E.O., The multiple recruitment systems of the african weaver ant Oecophylla longinoda (Latreille) (Hymenoptera: Formicidae) (1978) Behavioral Ecology and Sociobiology, 3 (1), pp. 19-60Hlldobler, B., Territoriality among Oecophylla (1979) National Geographic Society Research Reports, 1977, pp. 369-372Hlldobler, B., Territorial behavior in the green tree ant (Oecophylla smaragdina) (1983) Biotropica, 15, pp. 241-250De Vita, J., Mechanisms of interference and foraging among colonies of the harvester ant Pogonomyrmex californcus in the Mojave Desert (1979) Ecology, 60, pp. 729-73

Competition, Resources And The Ant (hymenoptera: Formicidae) Mosaic: A Comparison Of Upper And Lower Canopy

A canopy crane was used to assess ant defensive behaviour and recruitment at baits in the Parque Natural Metropolitano, Panama. Sardine-honey baits were set within a grid of 25 paired upper and lower canopy points, for which coordinates and height were recorded. We tested the hypothesis that interactions in the ant mosaic become stronger as one moves from the lower to the upper canopy. We sampled 23 ant species, with Azteca (A. trigona, A. velox, Azteca nr. chartifex, and A. snellingi) being by far the most abundant genus, recruiting to 63% of baits and excluding all other ant genera. Camponotus (Myrmobrachys) sp. 1 also showed a statistically significant exclusion of other ant species over 95% of its occurrence. Cephalotes umbraculatus and Dolichoderus bispinosus had exclusive occurrences in smaller areas. Exclusion between dominant or subdominant species was more frequent in the upper than lower canopy. Permeable borders and territory-free spaces are important for ant species diversity, and were more frequent in the lower canopy. Here, a combination of more costly patrolling conditions and less profitable resources, such as extra-floral nectaries and trophobionts, may be the most likely cause of this pattern. The findings presented here could account for the viewpoint of some that ant mosaics exist in plantations but not necessarily in tropical forest canopies.18113120Adams, E.S., Territory defense by the ant Azteca trigona: Maintenance of an arboreal ant mosaic (1994) Oecologia, 97 (2), pp. 202-208Basset, Y., Aberlenc, H.P., Delvare, G., Abundance and stratification of foliage arthropods in a lowland rain forest of Cameroon (1992) Ecological Entomology, 17 (4), pp. 310-318Beugnon, G., Dejean, A., Adaptative properties of the chemical trail system of the African weaver ant Oecophylla longinoda, LATREILLE (Hymenoptera: Formicinae) (1992) Insectes Sociaux, 39, pp. 341-346Blüthgen, N., Feldhaar, H., Food and shelter: How resources influence ant ecology (2009) Ant Ecology, pp. 115-136. , LACH, L., PARR, C. L. & ABBOTT, K.L. (Eds.) Oxford University Press, OxfordBluthgen, N., Stork, N.E., Ant mosaics in a tropical rainforest in Australia and elsewhere: A critical review (2007) Austral Ecology, 32 (1), pp. 93-104. , DOI 10.1111/j.1442-9993.2007.01744.xBluthgen, N., Stork, N.E., Fiedler, K., Bottom-up control and co-occurrence in complex communities: Honeydew and nectar determine a rainforest ant mosaic (2004) Oikos, 106 (2), pp. 344-358. , DOI 10.1111/j.0030-1299.2004.12687.xBlüthgen, N., Verhaagh, M., Goitía, W., Jaffé, K., Morawetz, W., Barthlott, W., How plants shape the ant community in the Amazonian rainforest canopy: The key role of extrafloral nectaries and homopteran honeydew (2000) Oecologia, 125, pp. 229-240Bolton, B., (2012) An Online Catalog of the Ants of the World, , http://www.antcat.org, retrieved on 15 December 2010Campos, R.I., Vasconcelos, H.L., Ribeiro, S.P., Neves, F.S., Soares, J.P., Relationship between tree size and insect assemblages associated with Anadenanthera macrocarpa (2006) Ecography, 29 (3), pp. 442-450. , DOI 10.1111/j.2006.0906-7590.04520.xDavidson, D.W., Ecological studies of Neotropical ant gardens (1998) Ecology, 69, pp. 1138-1153Dejean, A., Beugnon, G., Persistent intercolonial trunk route marking in the African weaver ant Oecophylla longinoda, LATREILLE (Hymenoptera: Formicinae) - Tom Thumb's versus Ariadne's orienting strategies (1991) Ethology, 88, pp. 89-98Dejean, A., Corbara, B., Fernández, F., Delabie, J.H.C., Mosaicos de hormigas arbóreas en bosques y plantaciones tropicales (2003) Introducción a Las Hormigas de la Región Neotropical, pp. 149-158. , FERNÁNDEZ, F. (Ed.) Instituto de Investigación de Recursos Biológicos Alexander von Humboldt, BogotáDejean, A., Corbara, B., Orivel, J., Leponce, M., Rainforest canopy ants: The implications of territoriality and predatory behavior (2007) Functional Ecosystems and Communities, 1, pp. 105-120Delabie, J.H.C., Trophobiosis between formicidae and hemiptera (sternorrhyncha and auchenorrhyncha): An overview (2001) Neotropical Entomology, 30 (4), pp. 501-516Del Claro, K., Oliveira, P.S., Ant-Homoptera interaction: Do alternative sugar sources distract tending ants? (1993) Oikos, 68, pp. 202-206Del Claro, K., Oliveira, P.S., Honeydew flicking by treehoppers provides cues to potential tending ants (1996) Animal Behaviour, 51, pp. 1071-1075Del-Claro, K., Oliveira, P.S., Conditional outcomes in a neotropical treehopper-ant association: Temporal and species-specific variation in ant protection and homopteran fecundity (2000) Oecologia, 124 (2), pp. 156-165Fagundes, F., Del-Claro, K., Ribeiro, S.P., Effects of the trophobiont herbivore Calloconophora pugionata (Hemiptera) on ant fauna associated with Myrcia obovata (Myrtaceae) in a montane tropical forest (2012) Psyche, 2012. , Article ID 783945Fitzjarrald, D.R., Moore, K.E., Physical methods of heat and mass exchange between forests and the atmosphere (1995) Forest Canopies, pp. 45-72. , LOWMAN, M.D. & NADKARNI, N.M. (Eds.) Academic Press, San Diego, CAFloren, A., Linsenmair, K.E., Do ant mosaics exist in pristine lowland rain forests? (2000) Oecologia, 123 (1), pp. 129-137Floren, A., Linsenmair, K.E., The importance of primary tropical rain forest for species diversity: An investigation using arboreal ants as an example (2005) Ecosystems, 8 (5), pp. 559-567. , DOI 10.1007/s10021-002-0272-8Goitía, W., Jaffé, K., Ant-plant associations in different forests in Venezuela (2009) Neotropical Entomology, 38, pp. 1-41Guerrero, R.J., Delabie, J.H.C., Dejean, A., Taxonomic contribution to the Aurita group of the ant genus Azteca (Formicidae: Dolichoderinae) (2010) Journal of Hymenopteran Research, 19, pp. 51-65Hahn, D.A., Wheeler, D.E., Seasonal foraging activity and bait preferences of ants on Barro Colorado Island, Panama (2002) Biotropica, 34 (3), pp. 348-356Holbrook, N.M., Lund, C.P., Photosynthesis in forest canopies (1995) Forest Canopies, pp. 411-430. , LOWMAN, M.D. & NADKARNI, N.M. (Eds.) Academic Press, San Diego, CAHölldobler, B., Territoriality among Oecophylla (1979) National Geographic Society Research Reports, 1977, pp. 369-372Holldobler, B., Lumsden, C.J., Territorial strategies in ants (1980) Science, 210 (4471), pp. 732-739Hölldobler, B., Wilson, E.O., (1990) The Ants, 732p. , The Belknap Press of Harvard University Press, Cambridgȩ MAJaffé, K., Hernandez, J.V., Goitía, W., Osio, A., Osborn, F., Cerda, H., Arab, A., Lopez, H., Flower ecology in the Neotropics: A flower-ant love-hate relationship (2003) Arthropods of Tropical Forests: Spatio-temporal Dynamics and Resource use in the Canopy, pp. 213-219. , BASSET, Y., NOVOTNY, V., MILLER, S. & KITCHING, R. (Eds.) Cambridge University Press, Cambridge, UKLeston, D., A Neotropical ant mosaic (1978) Annals of the Entomological Society of America, 71, pp. 649-653Longino, J.T., (2010) Specializing in Neotropical Myrmecology, , http://academic.evergreen.edu/projects/ants/, John T. Longino retrieved on 15 December 2010Lowman, M.D., Herbivory as a canopy process in rain forest trees (1995) Forest Canopies, pp. 431-455. , LOWMAN, M.D. & NADKARNI, N.M. (Eds.) Academic Press, San Diego, CAMajer, J.D., The ant mosaic in Ghana cocoa farms (1972) Bulletin of Entomological Research, 62, pp. 151-160Majer, J.D., Comparison of the arboreal ant mosaic in Ghana, Brazil, Papua New Guinea and Australia - Its structure and influence on ant diversity (1992) Hymenoptera and Biodiversity, pp. 115-141. , LASALLE, J. & GAULD, I. (Eds.) CAB International, Wallingford, UKMajer, J.D., Camerpesci, P., Ant species on tropical Australian tree crop and native ecosystems - Is there a mosaic? (1991) Biotropica, 23, pp. 173-181Majer, J.D., Delabie, J.H.C., An evaluation of Brazilian cocoa farm ants as potential biological control agents (1993) Journal of Plant Protection in the Tropics, 10, pp. 43-49Majer, J.D., Delabie, J.H.C., Smith, M.R.B., Arboreal ant community patterns in Brazilian cocoa farms (1994) Biotropica, 26, pp. 73-83Parker, G.G., Structure and microclimate of forest canopies (1995) Forest Canopies, pp. 73-106. , LOWMAN, M.D. & NADKARNI, N.M. (Eds.) Academic Press, San Diego, CAParr, C.L., Gibb, H., Competition and the role of dominant ants (2009) Ant Ecology, pp. 115-136. , LACH, L., PARR, C.L. & ABBOTT, K.L. (Eds.) Oxford University Press, OxfordPfeiffer, M., Linsenmair, K.E., Territoriality in the Malaysian giant ant Camponotus gigas (Hymenoptera/Formicidae) (2001) Journal of Ethology, 19 (2), pp. 75-85. , DOI 10.1007/s101640170002Ribas, C.R., Schoereder, J.H., Are all ant mosaics caused by competition? (2002) Oecologia, 131 (4), pp. 606-611. , DOI 10.1007/s00442-002-0912-xRibeiro, S.P., Insect herbivores in the canopies of savannas and rainforests (2003) Arthropods of Tropical Forests: Spatiotemporal Dynamics and Resource use in the Canopy, pp. 348-359. , BASSET, Y., NOVOTNY, V., MILLER, S. & KITCHING, R. (Eds.) Cambridge University Press, Cambridge, UKRichards, P.W., (1952) The Tropical Rainforest, 450p. , Cambridge University Press, Cambridge, UKRoom, P.M., The relative distribution of ant species in Ghana's cocoa farms (1971) Journal of Animal Ecology, 40, pp. 735-751Sanders, N.J., Crutsinger, G.M., Dunn, R.R., Majer, J.D., Delabie, J.H.C., An ant mosaic revisited: Dominant ant species disassemble arboreal ant communities but co-occur randomly (2007) Biotropica, 39 (3), pp. 422-427. , DOI 10.1111/j.1744-7429.2007.00263.xSchupp, E.W., Feener Jr., D.H., Phylogeny, lifeform, and habitat dependence of ant-defended plants in a Panamanian forest (1991) Ant-plant Interactions, pp. 175-197. , HUXLEY, C.R. & CUTLER, D.F. (Eds.) Oxford Science Publications, OxfordStork, N.E., The composition of the arthropod fauna of Bornean lowland rain forest trees (1991) Journal of Tropical Ecology, 7, pp. 161-180Tanaka, H.O., Yamane, S., Itioka, T., Effects of a ferndwelling ant species Crematogaster difformis, on the ant assemblages of emergent trees in a Bornean rainforest (2012) Annals of the Entomological Society of America, 105, pp. 592-598Tobin, J.E., Ecology and diversity of tropical forest canopy ants (1995) Forest Canopies, pp. 129-147. , LOWMAN, M.D. & NADKARNI, N. (Eds.) Academic Press, LondonWright, S.J., Horlyck, V., Basset, Y., Tropical canopy biology program, Republic of Panama (2003) Studying Forest Canopies from Above: The International Canopy Crane Network, pp. 138-158. , BASSET, Y., HORLYCK, V. & WRIGHT, S.J. (Eds.) Smithsonian Tropical Research Institute and UNEP, Panama City, PanamaYanoviak, S.P., Kaspari, M., Community structure and the habitat templet: Ants in the tropical forest canopy and litter (2000) Oikos, 89, pp. 256-26