Daily Activity Schedule, Gregariousness, And Defensive Behaviour In The Neotropical Harvestman Goniosoma Longipes (opiliones: Gonyleptidae)

In this paper we provide a field account of some aspects of the behavioural biology of Goniosoma longipes (Roewer), a harvestman which commonly occurs in caves in South-east Brazil. During daytime, solitary and aggregated individuals can be found resting on the cave walls. Just after sunset, however, many individuals leave the cave to forage for live and dead arthropods. Foraging individuals return to the cave before dawn. Aggregations of G. longipes contain on average 34 individuals (range 7-200), and the groups are usually found close to the water source and away from the cave entrance. The main predators of G. longipes are the spider Ctenus fasciatus Mello-Leitao (Ctenidae) and the opossum Philander opossum (L.) (Didelphidae). Upon disturbance solitary and aggregated individuals may either flee, or drop from the cave wall or vegetation. The harvestmen can also release a repugnatory liquid upon manipulation, and aggregated individuals collectively discharge this secretion toward the aggressor before fleeing. The activity schedule of G. longipes shows that individuals need to leave the cave periodically to forage, and therefore the population can be considered trogloxene. Data on the food items collected by G. longipes indicate that the harvestman is a generalist predator which also feeds on dead animal matter. Gregarious behaviour is considered relatively common among harvestmen and has been interpreted in several ways. We suggest that gregarious behaviour in G. longipes may be related with the choice of more suitable microconditions in the cave habitat and/or with group chemical defence.344587596Acosta, L.E., Poretti, T.I., Mascarelli, P.E., The defensive secretions of Pachyloidellus goliath (Opiliones, Laniatores, Gonyleptidae) (1993) Bonner Zoologishche Beiträge, 44, pp. 19-31Anuradha, K., Parthasarathy, M.D., Field studies on the ecology of gagrellula saddlana roewer (Palpatores, Opiliones, Arachnida) and its behaviour in the laboratory condition (1976) Bulletin of the Ethological Society of India, 1, pp. 68-71Barr T.C., Jr., Holsinger, J.R., Speciation in cave faunas (1985) Annual Review of Ecology and Systematics, 16, pp. 313-337Berland, L., Ordre des opilions (1949) Traité de Zoologie, Anatomie, Systematique, Biologie, pp. 761-793. , P. Grassé (ed.) (Paris: Masson et Cie)Bristowe, W.S., Notes on habits of insects and spiders in Brazil (1925) Transactions of the Royal Entomological Society of London, 1924, pp. 475-504Bristowe, W.S., The distribution of harvestmen (Phalangida) in Great Britain and ireland, with notes on their names, enemies and food (1949) Journal of Animal Ecology, 18, pp. 100-114Capocasale, R., Bruno-Trezza, L.B., Biologia de acanthopachylus aculeatus (Kirby, 1819), (Opiliones: Pachylinae) (1964) Revista de la Sociedad Uruguay a de Entomologia, 6, pp. 19-32Cloudsley-Thompson, J.L., (1958) Spiders, Scorpions, Centipedes and Mites, , (London: Pergamon Press), 227 ppCockerill, J.J., Notes on aggregations of Leiobunum (Opiliones) in the southern U.S.A. (1988) Journal of Arachnology, 16, pp. 123-126Coddington, J., Horner, M., Soderstrom, E.A., Mass aggregations in tropical harvestmen (Opiliones, Gagrellidae: Prionostemma sp.) (1990) Revue Arachnologique, 8, pp. 213-219Cokendolpher, J.C., Observations on the defensive behaviors of a neotropical Gonyleptidae (Arachnida: Opiliones) (1987) Revue Arachnologique, 7, pp. 59-63Cokendolpher, J.C., Pathogens and parasites of Opiliones (Arthropoda: Arachnida) (1993) Journal of Arachnology, 21, pp. 120-146Duffield, R.M., Olubajo, O., Wheeler, J.W., Shear, W.A., Alkyphenols in the defensive secretion of the Nearctic opilionid, Stygnomma spinifera (Arachnida: Opiliones) (1981) Journal of Chemical Ecology, 7, pp. 445-452Edgar, A.E., Studies on the biology and ecology of Michigan Phalangida (Opiliones) (1971) Miscellaneous Publications Museum of Zoology University of Michigan, 144, pp. 1-64Edgar, A.E., Opiliones (Phalangida) (1990) Soil Biology Guide, pp. 529-581. , D. L. Dindal (ed.) (New York: John Wiley and Sons)Eisner, T., Alsop, D., Meinwald, J., Secretions of opilionids, whip scorpions and pseudoscorpions (1978) Handbook of Experimental Pharmacology, 48, pp. 87-99. , S. Bettini (ed.) (Arthropod venoms) (Berlin: Springer-Verlag)Estable, C., Ardao, M.I., Brasil, H.P., Fieser, L.F., Gonyleptidine (1955) Journal of the American Chemical Society, 77, p. 4942Gautier, J.Y., Delaporte, P., Rivault, C., Relationships between ecology and social behavior in cockroaches (1988) The Ecology of Social Behavior, pp. 335-351. , C. N. Slobodchikoff (ed.) (New York: Academic Press)Gnaspini, P., Reproduction and postembryonic development of Goniosoma spelaeum, a cavernicolous harvestman from southeastern Brazil (Arachnida: Opiliones: Gonyleptidae) (1995) Invertebrate Reproduction and Development, 28, pp. 137-151Gnaspini, P., Population ecology of goniosoma spelaeum, a cavernicolous harvestman from southeastern Brazil (Arachnida: Opiliones: Gonyleptidae) (1996) Journal of Zoology, 239, pp. 417-435Gnaspini, P., Cavalheiro, A.J., Chemical and behavioral defences of a Neotropical cavernicolous harvestman: Goniosoma spelaeum (Opiliones: Laniatores: Gonyleptidae) (1998) Journal of Arachnology, 26, pp. 81-90Gnaspini, P., Trajano, E., Brazilian cave invertebrates, with a checklist of troglomorphic taxa (1994) Revista Brasileira de Entomologia, 38, pp. 549-584Holmberg, R.G., The scent glands of opiliones: A review of their function (1983) Proceedings of the 5th Congress International of Arachnology, 1983, pp. 131-133Holmberg, R.G., Angerilli, N.P.D., Lacasse, J.L., Overwintering aggregation of Leiobunum paessleri in caves and mines (Arachnida, Opiliones) (1984) Journal of Arachnology, 12, pp. 195-204Holsinger, J.R., Culver, D.C., The invertebrate cave fauna of Virginia and a part of Eastern Tennessee: Zoogeography and ecology (1988) Brimleyana, 14, pp. 1-162Kaestner, A., Arachnids and myriapods (1980) Invertebrate Zoology, 2. , trans. and adapted by H. W. Levi and E. R. Levi (Hutington: Robert E. Krieger Publications Co.), 472 ppKrebs, J.R., Davies, N.B., (1987) An Introduction to Behavioural Ecology, , (Oxford: Blackwell), 389 ppKunz, T., (1982) Ecology of Bats, , New York: Plenum Press, 425 ppMachado, G., Oliveira, P.S., Reproductive biology of the neotropical harvestman goniosoma longipes (Arachnida, Opiliones, Gonyleptidae): Mating and oviposition behaviour, brood mortality, and parental care (1998) Journal of Zoology, 246, pp. 359-367Machado, G., Vasconcelos, C.H.F., Multi-species aggregations in Neotropical harvestmen (Opiliones: Gonyleptidae) (1998) Journal of Arachnology, 26, pp. 389-391Martin, P., Bateson, P., (1986) Measuring Behaviour: An Introductory Guide, , (New York: Cambridge University Press), 200 ppPellegati-Franco, F.P., Gnaspini, P., Use of caves by philander opossum (Mammalia: Didelphidae) in Southeastern Brazil (1996) Papéis Avulsos de Zoologia, 39, pp. 351-364Phillipson, J., A contribution to the feeding biology of mitopus morio (F) (Phalangida) (1960) Journal of Animal Ecology, 29, pp. 35-43Pinto-Da-Rocha, R., Invertebrados cavernícolas da porção meridional da provincia espeleológica do Vale do Ribeira, Sul do Brasil (1993) Revista Brasileira de Zoologia, 10, pp. 229-255Ramires, E.N., Giaretta, A.A., Maternal care in a Neotropical harvestman, acutisoma proximum (Opiliones, Gonyleptidae) (1994) Journal of Arachnology, 22, pp. 179-180Roth, V.D., Roth, B.M., A review of appendotomy in spiders and other arachnids (1984) Bulletin of British Arachnological Society, 6, pp. 137-146Sabino, J., Gnaspini, P., Harvestman (Opiliones, Gonyleptidae) takes prey from a spider (Araneae, Ctenidae) (1999) Journal of Arachnology, , in pressSavory, T.H., Notes on the biology of harvestman (1938) Journal of the Queckett Microscopical Club, 1, pp. 89-94Wagner, H.O., Massenansammlungen von weberknechten in Mexico (1954) Zeitschrift Tierpsychologie, 11, pp. 348-35

Reciprocal Interference Between Ants And Stingless Bees Attending The Honeydew-producing Homopteran Aetalion Reticulatum (homoptera: Aetalionidae)

We evaluated the reciprocal effects between foragers of the ants Camponotus crassus and of the stingless bees Trigona hyalinata on aggregations of the honeydew-producing treehopper Aetalion reticulatum. The interactions were observed in Bauhinia variegata (Caesalpiniaceae) and Mangifera indica (Anacardiaceae) trees. We recorded the presence/absence of each attendant species in homopteran aggregations to test if the observed co-occurrence is lower than that expected by chance. An exclusion experiment was performed in which each attendant species was excluded from aggregations in order to test if an attendant species is more likely to occupy aggregations where the other attendant is not present. We also recorded the number of individuals of each attendant species in homopteran aggregations to search for any correlation between homopteran and attendant abundances. Additionally, we performed experiments using termites (Termitidae, Isoptera) as models to verify if the attendant species have the potential to defend A. reticulatum against natural enemies. The co-occurrence of attendant species was lower than that expected by chance. Homopteran aggregations without stingless bees were more visited by ants than those in which T. hyalinata was present, and vice-versa. The abundance of stingless bees was proportional to homopteran abundance, while ants abundance was not correlated to homopteran abundance. Both attendant species attacked the natural enemies models when we glued the termites ca. 1 cm away from homopteran aggregations, but only ants removed termites glued 5-7 cm away from aggregations. We suggest that the effects of non-formicid attendants should be included as another factor influencing the costs and benefits of ant-homopteran interactions, since honeydew availability for ants also depends on the presence and behavior of interspecific attendants.422369380Addicott, J.F., Multispecies aphid ant association - Density dependence and species-specific effects (1979) Canadian Journal of Zoology - Revue Canadienne de Zoologie, 57, pp. 558-569Barrows, E.M., Polistes wasps (Hymenoptera, Vespidae) show interference competition with other insects for Kermes scale insect (Homoptera, Kermesidae) secretions (1979) Proceedings of the Entomological Society of Washington, 81, pp. 570-575Bentley, B.L., Extra-floral nectaries and protection by pugnacious bodyguards (1977) Annual Review of Ecology and Systematics, 8, pp. 407-427Bentley, B.L., Benson, W.W., The influence of ant foraging patterns on the behavior of herbivores (1988) Advances in Myrmecology, pp. 297-306. , J.C. Trager (ed.). E.J.Brill, New YorkBlank, S., Setter, C., Bruce, P., (2001) Resampling Stats in Excel - Version 2, , Arlington, VA: Resampling Stats, IncBreton, L.M., Addicott, J.F., Does host-plant quality mediate aphid-ant mutualism? (1992) Oikos, 63, pp. 253-259Bristow, C., Differential benefits from ant attendance to two tropical species of homopterans of New York ironweed (1984) Journal of Animal Ecology, 53, pp. 15-726Buckley, R.C., Interactions between ants and membracid bugs decreases growth and seed set of host plant bearing extrafloral nectaries (1983) Oecologia, 58, pp. 132-136Buckley, R.C., Interactions involving plants, Homoptera, and ants (1987) Annual Review of Ecology and Systematics, 18, pp. 111-138Castro, P.R., Mutualismo entre Trigona spinipes (Fabricius, 1973) e Aetalion reticulatum (L. 1767) em Cajanus indicus Spreng. na presença de Camponotus spp (1975) Ciência & Cultura, 27, pp. 537-539Cortopassi-Laurino, M., Notas sobre associações de Trigona (Oxitrigona) taitara (Apidae, Meliponinae) (1977) Boletim de Zoologia da Universidade de São Paulo, 2, pp. 183-187Dejean, A., Turillazzi, S., Territoriality during trophobiosis between wasps and homopterans (1992) Tropical Zoology, 5, pp. 237-247Del-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, pp. 156-165Figueiredo, R.D.A., Interactions between stingless meliponine bees, honeydew-producing homopterans, ants and figs in a cerrado area (1996) Naturalia, 2, pp. 159-164Flatt, T., Weisser, W.W., The effects of mutualistic ants on aphid life history traits (2000) Ecology, 81, pp. 3522-3529Fowler, H.G., Aethalionidae: Functional equivalents of extrafloral nectaries in Bauhinia (Caesalpinoidea) (1992) Anales de Biologia, 18, pp. 155-159Hubbell, S.P., Johnson, L.K., Comparative foraging behavior of six stingless bee species exploiting a standardized resource (1978) Ecology, 59, pp. 1123-1136Johnson, L.K., Hubbell, S.P., Aggression and competition among stingless bees - Field studies (1974) Ecology, 55, pp. 120-127Krebs, J., Optimal foraging - Theory and experiment (1977) Nature, 268, pp. 583-584Laroca, S., Mutualism between Trigona fuscipennis (Hymenoptera, Apoidea) and Anchistrotus amitteraglobus, an amazonian membracid that suffers mutilation of the pronotum (1997) Acta Biológica Paranaense, 26, pp. 1-18Laroca, S., Sakakibara, A.M., Mutualismo entre Trigona hyalinata branneri (Apidae) e Aconophora flavipes (Membracidae) (1976) Revista Brasileira de Entomologia, 20, pp. 71-72Letourneau, D.K., Choe, J.C., Homopteran attendance by wasps and ants: The stochastic nature of interactions (1987) Psyche, 94, pp. 81-91Messina, F.J., Plant-protection as a consequence of an ant-membracid mutualism - Interactions on goldenrod (Solidago sp) (1981) Ecology, 62, pp. 1433-1440Morales, M.A., Mechanisms and density dependence of benefit in an ant-membracid mutualism (2000) Ecology, 81, pp. 482-489Oliveira, P.S., Silva, A.F., Martins, A.B., Ant foraging on extrafloral nectaries of Qualea grandiflora (Vochysiaceae) in cerrado vegetation: Ants as potential antiherbivore agents (1987) Oecologia, 74, pp. 228-230Roubik, D.W., (1989) Ecology and Natural History of Tropical Bees, , Cambridge Tropical Biology series, New YorkSilva, A.G.A., Gonçalves, C.R., Galvão, D.M., Gonçalves, A.J.L., Silva, M.N., Simoni, L., (1968) Quarto Catálogo dos Insetos que vivem nas Plantas do Brasil: Seus Parasitos e Predadores. Parte II - 1 Tomo. Insetos, Hospedeiros e Inimigos Naturais, , Ministério da Agricultura. Rio de Janeiro, BrasilSokal, R.R., Rohlf, F.J., (1995) Biometry. 3rd Edition, , W.H. Freeman, New YorkSudd, J.H., Sudd, M.E., Seasonal changes in the response of wood-ants (Formica lugubris) to sucrose baits (1983) Ecological Entomology, 10, pp. 89-97Völkl, W., Woodring, J., Fischer, M., Lorenz, M.W., Hoffmann, K.H., Ant-aphid mutualisms: The impact of honeydew production and honeydew sugar composition on ant preferences (1999) Oecologia, 118, pp. 483-491Way, M.J., Mutualism between ants and honeydew-producing Homoptera (1963) Annual Review of Entomology, 8, pp. 307-344Wilson, E.O., (1971) The Insects Societies, , Belknap, Cambridge, Mas