Os grupos tróficos em Coleoptera

Marinoni, Renato C.

doi:10.1590/S0101-81752001000100024

Resumo

The beetles are a useful group for studies on trophic structure of communities, mainly in forested areas. These kind of studies are based on food habits of species groups. The different terms applied to nomminated these groups (trophic category, ecologic group, trophic group, guild, trophic guild) are discussed. The term trophic group, a natural unity, is proposed to form a group of species with the same food habits, not considering the trophic level. The guild, an artificial unity, is recognized as an important tool to group organisms group that use the same resources as defined by the investigator (Hawkins & MacMaiion, 1989; Simberloff & Dayan 1991; Wilson 1999). The known alimentary habits of Coleoptera are classified in five trophic groups - herbivores, algivores, fungívores, detrivores and carnivores. Within each trophic group, subgroups are recognized. The subgroup are in a lower hierarchic level and are defined by morphological (herbivores), taxonomic (fungivores) atributes of the food, by the food origin (detrivores) and by the way that the food is obtained (carnivores). A food resource diagram showing connections among the trophic groups, based on consumer-consumed (predator-prey) relations, is presented.

Coleoptera; trophic groups; guilds; communities; trophic structure

Os grupos tróficos em Coleoptera¹ 1 Contribuição número 1265 do Departamento de Zoologia, Universidade Federal do Paraná.

The trophic groups in Coleoptera

Renato C. Marinoni

Departamento de Zoologia, Universidade Federal do Paraná. Caixa Postal 19020, 81531-990 Curitiba, Paraná, Brasil. Bolsista do CNPq

ABSTRACT

The beetles are a useful group for studies on trophic structure of communities, mainly in forested areas. These kind of studies are based on food habits of species groups. The different terms applied to nomminated these groups (trophic category, ecologic group, trophic group, guild, trophic guild) are discussed. The term trophic group, a natural unity, is proposed to form a group of species with the same food habits, not considering the trophic level. The guild, an artificial unity, is recognized as an important tool to group organisms group that use the same resources as defined by the investigator (Hawkins & MacMaiion, 1989; Simberloff & Dayan 1991; Wilson 1999). The known alimentary habits of Coleoptera are classified in five trophic groups - herbivores, algivores, fungívores, detrivores and carnivores. Within each trophic group, subgroups are recognized. The subgroup are in a lower hierarchic level and are defined by morphological (herbivores), taxonomic (fungivores) atributes of the food, by the food origin (detrivores) and by the way that the food is obtained (carnivores). A food resource diagram showing connections among the trophic groups, based on consumer-consumed (predator-prey) relations, is presented.

Key words: Coleoptera, trophic groups, guilds, communities, trophic structure

Texto completo disponível apenas em PDF.

Full text available only in PDF format.

Recebido em 20.XII.1999, aceito em 09.I.2001.

Beley, J.R.; T.M. Ditswort; S.M. Butt & CD. Johnson. 1982. Arthropods, plants and transmission lines in Arizona: community dynamics during secondary succession in a pinyon-juniper Chaparral habitat. The Southwestern Naturalist 27(3):325-333.
Brown, V.K. 1985. Insect herbivores and plant succession. Oikos 44(1):17-22.
Brown, V.K. & P.S. Hyman. 1986. Successional communities of plants and phytophagous Coleoptera. Jour. Ecol. 74(4):963-975.
Brown, V.K. & T.R.E. Southwood. 1983. Trophic diversity, niche breadth and generation times of exopterigote insects in a secondary succession. Oecologia 56:220-225.
Burns, T.P. 1989. Lindeman's contradiction and the trophic structure of ecosystems. Ecology 70(5):1355-1362.
Butt, S.M.; J.R. Beley; T.M. Ditsworth; C.D. Johnson & R.P. Balda. 1980. Arthropods, plants and transmission lines in Arizona: community dynamics during secondary succession in a desert grassland. Jour. Environ. Management 11:267-284.
Cohen, J.E. 1978. Food webs and niche space. Princeton, Princeton University Press.
Carlton, CE. & H.W. Robinson. 1998. Diversity of litter-dwelling beetles in the Ouachita Highlands of Arkansas, USA (Insecta: Coleoptera). Biodiversity and Conservation 7:1589-1605.
Costa, C; S.A. Vanin; S.A. Casari-Chen. 1988. Larvas de Coleoptera do Brasil. São Paulo, Ed. Museu de Zoologia da Universidade de São Paulo.
Coulson, R.N.; R.O. Flamm; P.E. Pulley; T.L. Payne; E.J. Rykiei. & T.L. Wagner. 1986. Response of the southern pine bark beetle guild (Coleoptera, Scolytidae) to host disturbance. Environ. Entomol. 15(4):850-858.
Crowson, R.A. 1981. The biology of the Coleoptera. London, Academic Press.
Cummins, K.W. 1974. Structure and function of stream ecosystems. Bioscience 24(11):631-641.
Deoca, E.M. & G. Halffter. 1995. Daily and seasonal activities of a guild of the coprophagous, burrowing beetle (Coleoptera, Scarabaeidae, Scarabaeinae) in tropical grassland. Trop. Zool. 8(1):159-180.
Didham, R.K.; J.H. Lawton; P.M. Hammond & P. Eggleton. 1998. Trophic structure stability and extinction dynamics of beetles (Coleoptera) in tropical forest fragments. Philos. Trans. Royal Soc. London 353:437-451.
Ditsworth, T.M.; S.M. Butt; J.R. Beley; CD. Johnson & R.P. Balda. 1982. Arthropods, plants, and transmission lines in Arizona: community dynamics during secondary succession in a pinyon-juniper woodland. The Southwestern Naturalist 27(2):167-181.
Evans, F.C & W.W. Murdoch. 1968. Taxonomic composition, trophic structure and seasonal occurrence in a grassland insect community. Jour. Animal Ecol. 37:259-273.
Finn, J.A., T. Gittings & P.S. Giller. 1999. Spatial and temporal variation in species composition of dung beetle assemblages in southern Ireland. Ecol. Entomol. 24(1):24-36.
Flamm, R.O.; P.E. Pulley & R.N. Coulson. 1993. Colonization of disturbed trees by the southern pine bark beetle guild (Coleoptera, Scolytidae). Environ. Entomol. 22(1):62-70.
Flechtmann, C.A.H.; H.T.Z. Couto; CL. Gaspareto & E. Berti-Filho. 1995. Manual de pragas em florestas. Scolytidae em reflorestamento com pinheiros tropicais (4). Piracicaba, IPEF.
Gaston, K.J.; P.H. Warren & P.M. Hammond. 1992. Predator: non-predator ratios in beetle assemblages. Oecologia 90:417-421.
Hawkins, B.A. 1988. Species-diversity in the third and fourth trophic levels: patterns and mechanisms. Jour. Animal Ecol. 57(1):137-162.
Hawkins, CP. & J.A. MacMahon. 1989. Guilds: the multiple meanings of a concept. Ann. Rev. Entomol. 34:423-451.
Heatwole, H. & R. Levins. 1972. Trophic structure stability and faunal change during recolonization. Ecology 53:531-534.
Hengeveld, R. 1980. Polyphagy, oligophagy and food specialization in ground beetles (Coleoptera, Carabidae). Netherlands Jour. Zool. 30(4):564-584.
Hering, D. 1998. Riparian beetles (Coleoptera) along a small stream in the Oregon Coast Range and their interactions with the aquatic environment. Coleop. Bull. 52(2):161-170.
Hodek, I. 1996. Food relationships. 6, p. 143-234. In: I. Hodek & A. Honek (Eds). Ecology of Coccinellidae. London, Kluwer Academic Publishers.
Holloway, B.A. 1982. Anthribidae (Insecta: Coleoptera). Fauna of New Zealand nº 3. Wellington, Science Information Division, DSIR.
Hutcheson, J. 1990. Characterization of terrestrial insect communities using quantified, Malaise-trapped Coleoptera. Ecol. Entomol. 15:143-151.
Hutcheson, J. & D. Junes. 1999. Spatial variability of insect communities in a homogenous system: measuring biodiversity using Malaise trapped beetles in a Pinus radiata plantation in New Zealand. Forest Ecol. Management 118:93-105.
Johnson, CD. & D.H. Siemens. 1995. Bruchid guilds, host preferences, and new host records from Latin America and Texas for the genus Stator Bridwell (Coleoptera, Bruchidae). Coleop. Bull. 49(2):133-142.
______. 1997. Distribution, oviposition guilds, behavior and new host records from Latin America for Algarobius Bridwell, Scutobruchus Kingsolver and Pseudopachymerina spinipes (Erichson) (Coleoptera: Bruchidae). Coleop. Bull. 51(1):37-42.
Johnson, CD.; J.R. Beley; T.M. Ditsworth & S.M. Butt. 1983. Secondary succession of arthropods and plants in the Arizona Sonoran Desert in response to transmission line construction. Jour. Environ. Management 16:125-137.
Kaila, L.; P. Martikainen; P. Punttila & E. Yakovlev. 1994. Saproxylic beetles (Coleoptera) on dead birch trunks decayed by different polypore species. Ann. Zool. Fennici 31:97-107.
Kistler, R.A. 1995. Influence of temperature on populations within a guild of mesquite bruchids (Coleoptera, Bruchidae). Environ. Entomol. 24(3):663-672.
Lawrence J.F. & E.B.Britton. 1991. Coleoptera. In: CSIRO (Ed.). The Insects of Australia. New York, Cornell University Press, Vol. 2.
Lima, A.M. da C. 1952-56. Insetos do Brasil, Coleópteros. Rio de Janeiro, Escola Nacional de Agronomia, Vol. 7-10.
Lovelady, C.N.; P.E. Pulley; R.N. Coulson & R.O. Flamm. 1991. Relation of lightning to herbivory by the southern pine bark beetle guild (Coleoptera, Scolytidae). Environ. Entomol. 20(5):1279-1284.
Marinoni, R.C & R.R.C. Dutra. 1997. Famílias de Coleoptera capturadas com armadilha malaise em oito localidades do Estado do Paraná, Brasil. Diversidades alfa e beta. Revta bras. Zool. 14(3):751-770.
Marinoni, R.C; N.G. Ganho; M. Monné & J.R. Mermudes. (no prelo). Hábitos alimentares em Coleoptera. Ribeirão Preto, Ed. Holos.
Mills, N.J. 1994. Parasitoid guilds: defining the structure of the parasitoid communities of endopterygote insect host. Environ. Entomol. 23(5):1066-1083.
Moeed, A. & M.J. Meads. 1985. Seasonality of pitfall trapped invertebrates in three types of native forest, Orongorongo Valley, New Zealand. New Zealand Jour. Zool. 12:17-53.
Moran, V.C & T.R.E. Southwood. 1982. The guild composition of arthropod communities in trees. Jour. Animal Ecol. 51:289-306.
Morris, M.G. 1980. Insects and the environment in the United Kingdom. Atti XII Congresso Nazionale Italiano di Entomologia, Roma, p. 203-235.
Muona, J. & I. Rutanen. 1994. The short-term impact of fire on the beetle fauna in boreal coniferous forest. Ann. Zool. Fennici 31:109-121.
Murrin, F. 1996. Fungi and insects, p. 365-388. In: D.H. Haward & J.D. Miller (Ed.). The Mycota. Human and Animal Relationships. Berlin, Springer, Vol. 6.
Parmenter, R.R. & J.A. MacMahon. 1987. Early successional patterns of arthropod recolonization on reclaimed strip mines in southwestern Wyoming: The ground-dwelling beetles fauna (Coleoptera). Environ. Entomol. 16(1):168-175.
Pellmyr, O. 1985. Flower constancy in individuals of an anthophilous beetle, Byturus ochraceus (Coleoptera: Byturidae). The Coleop. Bull. 39(4):341-345.
Penny , N.D. & J.R. Arias . 1982. Insects of an amazon forest. New York, Columbia University Press.
Ricklefs, R.E. 1990. Ecology. New York, W.H. Freeman and Company, 3^th ed.
Root, R.B. 1967. The niche exploitation pattern of the blue-gray gnatcatcher. Ecol. Monogr. 37:317-350.
Shtonen, J. 1994. Decaying wood and saproxylic Coleoptera in two old spruce forests: a comparison based on two sampling methods. Ann. Zool. Fennici 31:89-95.
Simandi., J. 1993. The spatial pattern, diversity niche partitioning in xylophagous beetles (Coleoptera) associated with Frangula alnus Mill. Acta Oecol. Internat. Jour. Ecol. 14(2):161-171.
Simberloff, D. & T. Dayan. 1991. The guild concept and the structure of ecological communities. Ann. Rev. Ecol. Syst. 22:115-143.
Southwood, T.R.E.; V.K. Brown & P.M. Reader. 1979. The relationship of plant and insect diversities in succession. Biol. Jour. Linnean Soc. 12:327-348.
Southwood, T.R.E.; V.C. Moran & C.E.J. Kennedy. 1982. The richness, abundance and biomass of the arthropod communities on trees. Jour. Animal Ecol. 51:635-649.
Stork, N.E. 1987. Guild structure of arthropods from Bornean rain forest trees. Ecol. Entomol. 12:69-80.
Stribling, J.B. & R.L. Seymour. 1988. Evidence of mycophagy in Ptilodactylidae (Coleoptera: Dryopoidea). Coleop. Bull. 42(2):152-154.
Tanaka, L.K.& S.K. Tanaka. 1982. Rainfall and seasonal changes in arthropod abundance on a tropical oceanic island. Biotropica 14(2):114-123.
Torre-Bueno, J.R. 1978. A glossary of entomology. New York, Ed. G.S. Tulloch, New York Entomological Society, 335p.
White, I.M. 1977. Larvae of some british species of Gyrophaena Mannerheim (Coleoptera-Saphylinidae) with notes on taxonomy and biology of the genus. Zool. Jour. Linnean Soc. 60(4):297-317.
Wilson , J.B. 1999. Guilds, functional types and ecological groups. Oikos 86(3):507-522.
Yodzys, P. 1982. The compartmentation of real and assembled ecosystems. Amer. Naturalist 120:551-570.
Yodzys, P. 1993. Environment and trophodiversity, p. 26-38. In: R.E. Ricklefs & D. Schluter (Ed.). Species diversity in ecological communities. Chicago, The University of Chigago Press.

1

Contribuição número 1265 do Departamento de Zoologia, Universidade Federal do Paraná.

Datas de Publicação

Publicação nesta coleção
30 Abr 2009
Data do Fascículo
Mar 2001

Histórico

Recebido
20 Dez 1999
Aceito
09 Jan 2001

This work is licensed under a Creative Commons Attribution-NonCommercial 4.0 International License.

[1] Beley, J.R.; T.M. Ditswort; S.M. Butt & CD. Johnson. 1982. Arthropods, plants and transmission lines in Arizona: community dynamics during secondary succession in a pinyon-juniper Chaparral habitat. The Southwestern Naturalist 27(3):325-333.

[2] Brown, V.K. 1985. Insect herbivores and plant succession. Oikos 44(1):17-22.

[3] Brown, V.K. & P.S. Hyman. 1986. Successional communities of plants and phytophagous Coleoptera. Jour. Ecol. 74(4):963-975.

[4] Brown, V.K. & T.R.E. Southwood. 1983. Trophic diversity, niche breadth and generation times of exopterigote insects in a secondary succession. Oecologia 56:220-225.

[5] Burns, T.P. 1989. Lindeman's contradiction and the trophic structure of ecosystems. Ecology 70(5):1355-1362.

[6] Butt, S.M.; J.R. Beley; T.M. Ditsworth; C.D. Johnson & R.P. Balda. 1980. Arthropods, plants and transmission lines in Arizona: community dynamics during secondary succession in a desert grassland. Jour. Environ. Management 11:267-284.

[7] Cohen, J.E. 1978. Food webs and niche space. Princeton, Princeton University Press.

[8] Carlton, CE. & H.W. Robinson. 1998. Diversity of litter-dwelling beetles in the Ouachita Highlands of Arkansas, USA (Insecta: Coleoptera). Biodiversity and Conservation 7:1589-1605.

[9] Costa, C; S.A. Vanin; S.A. Casari-Chen. 1988. Larvas de Coleoptera do Brasil. São Paulo, Ed. Museu de Zoologia da Universidade de São Paulo.

[10] Coulson, R.N.; R.O. Flamm; P.E. Pulley; T.L. Payne; E.J. Rykiei. & T.L. Wagner. 1986. Response of the southern pine bark beetle guild (Coleoptera, Scolytidae) to host disturbance. Environ. Entomol. 15(4):850-858.

[11] Crowson, R.A. 1981. The biology of the Coleoptera. London, Academic Press.

[12] Cummins, K.W. 1974. Structure and function of stream ecosystems. Bioscience 24(11):631-641.

[13] Deoca, E.M. & G. Halffter. 1995. Daily and seasonal activities of a guild of the coprophagous, burrowing beetle (Coleoptera, Scarabaeidae, Scarabaeinae) in tropical grassland. Trop. Zool. 8(1):159-180.

[14] Didham, R.K.; J.H. Lawton; P.M. Hammond & P. Eggleton. 1998. Trophic structure stability and extinction dynamics of beetles (Coleoptera) in tropical forest fragments. Philos. Trans. Royal Soc. London 353:437-451.

[15] Ditsworth, T.M.; S.M. Butt; J.R. Beley; CD. Johnson & R.P. Balda. 1982. Arthropods, plants, and transmission lines in Arizona: community dynamics during secondary succession in a pinyon-juniper woodland. The Southwestern Naturalist 27(2):167-181.

[16] Evans, F.C & W.W. Murdoch. 1968. Taxonomic composition, trophic structure and seasonal occurrence in a grassland insect community. Jour. Animal Ecol. 37:259-273.

[17] Finn, J.A., T. Gittings & P.S. Giller. 1999. Spatial and temporal variation in species composition of dung beetle assemblages in southern Ireland. Ecol. Entomol. 24(1):24-36.

[18] Flamm, R.O.; P.E. Pulley & R.N. Coulson. 1993. Colonization of disturbed trees by the southern pine bark beetle guild (Coleoptera, Scolytidae). Environ. Entomol. 22(1):62-70.

[19] Flechtmann, C.A.H.; H.T.Z. Couto; CL. Gaspareto & E. Berti-Filho. 1995. Manual de pragas em florestas. Scolytidae em reflorestamento com pinheiros tropicais (4). Piracicaba, IPEF.

[20] Gaston, K.J.; P.H. Warren & P.M. Hammond. 1992. Predator: non-predator ratios in beetle assemblages. Oecologia 90:417-421.

[21] Hawkins, B.A. 1988. Species-diversity in the third and fourth trophic levels: patterns and mechanisms. Jour. Animal Ecol. 57(1):137-162.

[22] Hawkins, CP. & J.A. MacMahon. 1989. Guilds: the multiple meanings of a concept. Ann. Rev. Entomol. 34:423-451.

[23] Heatwole, H. & R. Levins. 1972. Trophic structure stability and faunal change during recolonization. Ecology 53:531-534.

[24] Hengeveld, R. 1980. Polyphagy, oligophagy and food specialization in ground beetles (Coleoptera, Carabidae). Netherlands Jour. Zool. 30(4):564-584.

[25] Hering, D. 1998. Riparian beetles (Coleoptera) along a small stream in the Oregon Coast Range and their interactions with the aquatic environment. Coleop. Bull. 52(2):161-170.

[26] Hodek, I. 1996. Food relationships. 6, p. 143-234. In: I. Hodek & A. Honek (Eds). Ecology of Coccinellidae. London, Kluwer Academic Publishers.

[27] Holloway, B.A. 1982. Anthribidae (Insecta: Coleoptera). Fauna of New Zealand nº 3. Wellington, Science Information Division, DSIR.

[28] Hutcheson, J. 1990. Characterization of terrestrial insect communities using quantified, Malaise-trapped Coleoptera. Ecol. Entomol. 15:143-151.

[29] Hutcheson, J. & D. Junes. 1999. Spatial variability of insect communities in a homogenous system: measuring biodiversity using Malaise trapped beetles in a Pinus radiata plantation in New Zealand. Forest Ecol. Management 118:93-105.

[30] Johnson, CD. & D.H. Siemens. 1995. Bruchid guilds, host preferences, and new host records from Latin America and Texas for the genus Stator Bridwell (Coleoptera, Bruchidae). Coleop. Bull. 49(2):133-142.

[31] ______. 1997. Distribution, oviposition guilds, behavior and new host records from Latin America for Algarobius Bridwell, Scutobruchus Kingsolver and Pseudopachymerina spinipes (Erichson) (Coleoptera: Bruchidae). Coleop. Bull. 51(1):37-42.

[32] Johnson, CD.; J.R. Beley; T.M. Ditsworth & S.M. Butt. 1983. Secondary succession of arthropods and plants in the Arizona Sonoran Desert in response to transmission line construction. Jour. Environ. Management 16:125-137.

[33] Kaila, L.; P. Martikainen; P. Punttila & E. Yakovlev. 1994. Saproxylic beetles (Coleoptera) on dead birch trunks decayed by different polypore species. Ann. Zool. Fennici 31:97-107.

[34] Kistler, R.A. 1995. Influence of temperature on populations within a guild of mesquite bruchids (Coleoptera, Bruchidae). Environ. Entomol. 24(3):663-672.

[35] Lawrence J.F. & E.B.Britton. 1991. Coleoptera. In: CSIRO (Ed.). The Insects of Australia. New York, Cornell University Press, Vol. 2.

[36] Lima, A.M. da C. 1952-56. Insetos do Brasil, Coleópteros. Rio de Janeiro, Escola Nacional de Agronomia, Vol. 7-10.

[37] Lovelady, C.N.; P.E. Pulley; R.N. Coulson & R.O. Flamm. 1991. Relation of lightning to herbivory by the southern pine bark beetle guild (Coleoptera, Scolytidae). Environ. Entomol. 20(5):1279-1284.

[38] Marinoni, R.C & R.R.C. Dutra. 1997. Famílias de Coleoptera capturadas com armadilha malaise em oito localidades do Estado do Paraná, Brasil. Diversidades alfa e beta. Revta bras. Zool. 14(3):751-770.

[39] Marinoni, R.C; N.G. Ganho; M. Monné & J.R. Mermudes. (no prelo). Hábitos alimentares em Coleoptera. Ribeirão Preto, Ed. Holos.

[40] Mills, N.J. 1994. Parasitoid guilds: defining the structure of the parasitoid communities of endopterygote insect host. Environ. Entomol. 23(5):1066-1083.

[41] Moeed, A. & M.J. Meads. 1985. Seasonality of pitfall trapped invertebrates in three types of native forest, Orongorongo Valley, New Zealand. New Zealand Jour. Zool. 12:17-53.

[42] Moran, V.C & T.R.E. Southwood. 1982. The guild composition of arthropod communities in trees. Jour. Animal Ecol. 51:289-306.

[43] Morris, M.G. 1980. Insects and the environment in the United Kingdom. Atti XII Congresso Nazionale Italiano di Entomologia, Roma, p. 203-235.

[44] Muona, J. & I. Rutanen. 1994. The short-term impact of fire on the beetle fauna in boreal coniferous forest. Ann. Zool. Fennici 31:109-121.

[45] Murrin, F. 1996. Fungi and insects, p. 365-388. In: D.H. Haward & J.D. Miller (Ed.). The Mycota. Human and Animal Relationships. Berlin, Springer, Vol. 6.

[46] Parmenter, R.R. & J.A. MacMahon. 1987. Early successional patterns of arthropod recolonization on reclaimed strip mines in southwestern Wyoming: The ground-dwelling beetles fauna (Coleoptera). Environ. Entomol. 16(1):168-175.

[47] Pellmyr, O. 1985. Flower constancy in individuals of an anthophilous beetle, Byturus ochraceus (Coleoptera: Byturidae). The Coleop. Bull. 39(4):341-345.

[48] Penny , N.D. & J.R. Arias . 1982. Insects of an amazon forest. New York, Columbia University Press.

[49] Ricklefs, R.E. 1990. Ecology. New York, W.H. Freeman and Company, 3^th ed.

[50] Root, R.B. 1967. The niche exploitation pattern of the blue-gray gnatcatcher. Ecol. Monogr. 37:317-350.

[51] Shtonen, J. 1994. Decaying wood and saproxylic Coleoptera in two old spruce forests: a comparison based on two sampling methods. Ann. Zool. Fennici 31:89-95.

[52] Simandi., J. 1993. The spatial pattern, diversity niche partitioning in xylophagous beetles (Coleoptera) associated with Frangula alnus Mill. Acta Oecol. Internat. Jour. Ecol. 14(2):161-171.

[53] Simberloff, D. & T. Dayan. 1991. The guild concept and the structure of ecological communities. Ann. Rev. Ecol. Syst. 22:115-143.

[54] Southwood, T.R.E.; V.K. Brown & P.M. Reader. 1979. The relationship of plant and insect diversities in succession. Biol. Jour. Linnean Soc. 12:327-348.

[55] Southwood, T.R.E.; V.C. Moran & C.E.J. Kennedy. 1982. The richness, abundance and biomass of the arthropod communities on trees. Jour. Animal Ecol. 51:635-649.

[56] Stork, N.E. 1987. Guild structure of arthropods from Bornean rain forest trees. Ecol. Entomol. 12:69-80.

[57] Stribling, J.B. & R.L. Seymour. 1988. Evidence of mycophagy in Ptilodactylidae (Coleoptera: Dryopoidea). Coleop. Bull. 42(2):152-154.

[58] Tanaka, L.K.& S.K. Tanaka. 1982. Rainfall and seasonal changes in arthropod abundance on a tropical oceanic island. Biotropica 14(2):114-123.

[59] Torre-Bueno, J.R. 1978. A glossary of entomology. New York, Ed. G.S. Tulloch, New York Entomological Society, 335p.

[60] White, I.M. 1977. Larvae of some british species of Gyrophaena Mannerheim (Coleoptera-Saphylinidae) with notes on taxonomy and biology of the genus. Zool. Jour. Linnean Soc. 60(4):297-317.

[61] Wilson , J.B. 1999. Guilds, functional types and ecological groups. Oikos 86(3):507-522.

[62] Yodzys, P. 1982. The compartmentation of real and assembled ecosystems. Amer. Naturalist 120:551-570.

[63] Yodzys, P. 1993. Environment and trophodiversity, p. 26-38. In: R.E. Ricklefs & D. Schluter (Ed.). Species diversity in ecological communities. Chicago, The University of Chigago Press.