外来种与本地种西番莲花外蜜腺引起的生物防御特征的比较
|
摘要
花外蜜腺植物是植物中一个比较特殊的类群,这些植物具有着生于花外、般不参与传粉的蜜腺即花外蜜腺。花外蜜腺分泌糖分及微量氨基酸,吸引蚂蚁或寄生蜂等一群捕食者前来取食,蚂蚁取食花外蜜腺分泌物的同时捕食或干扰取食植物的昆虫,形成了蚂蚁与植物的互惠关系。本研究以长叶西番莲Passiflora siamica Craib.(本地种)和红花西番莲Passiflora coccinea Aubl.(外引种)为材料,探讨了在长叶西番莲的自然生境中,两种植物与蚂蚁建立互惠关系的状况及防御效果,两种植物对去蚂蚁、去取食者、去花外蜜腺以及去蚂蚁+去取食者几种处理的反应。主要研究结果如下：
1、在两种西番莲植株上共观察到24种蚂蚁取食花外蜜液,主要包括：立毛蚁一种Paratrechina sp.1、黑可可臭蚁Dolichoderus thoracicus (Smith)、黑头酸臭蚁Tapinoma melanocephalum、伊大头蚁Pheidole yeensis等,分别隶属于猛蚁亚科(1属1种),伪切叶蚁亚科(1属1种),切叶蚁亚科(5属9种)、臭蚁亚科(2属2种)和蚁亚科(4属11种)。
2、利用两种西番莲的蚂蚁数量没有显著差异(t=1.30,P=0.20),但主要的四种蚂蚁的种类组成比例存在显著差异(χ2=14.76,df=4,P<0.01)。长叶西番莲单片叶片泌蜜量显著高于红花西番莲(Willcox,z=-3.87,P<0.01)。去除蚂蚁处理对两种西番莲植物的平均叶面积损失均未造成显著影响。两种西番莲去蚂蚁、去取食者、去蚂蚁及取食者处理之间叶片泌蜜量均没有显著差异。
3、长叶西番莲叶片表皮毛密度显著低于红花西番莲(叶子上表面,t=3.28,P<0.01：叶子下表面,t=26.84,P<0.01)。长叶西番莲的叶片下表面表皮毛密度和植株平均叶面积损失呈显著负相关关系(Pearson correlation, r=-0.38, P=0.04,n=30),而红花西番莲的叶片表皮毛密度和植株平均叶面积损失不显著相关。
4、长叶西番莲植株上蚂蚁数量和平均叶面积损失之间呈显著的负相关关系(Pearson correlation, r=-0.45,P< 0.05, n=20)。利用两种西番莲植株花外蜜腺的蚂蚁群体中不同种类的蚂蚁的行为优势度指数BDI(表示一种蚂蚁在和该群体中其它所有蚂蚁相遇时占优势的比例的指数)和其数量呈显著的正相关关系(Pearson correlation,r=0.86,P=0.01,n=60).
综上所述,蚂蚁和花外蜜腺去除并未显著影响植株被广适性昆虫取食程度；但长叶西番莲植株上蚂蚁数量还是与植株叶片被广适性昆虫取食的程度呈显著负相关关系。而该系统对专食性昆虫取食的反应如何则有待进一步研究。
EFN (extrafloral nectary) plant is a special group of plants. They have nectaries on the other part of the plant more than just in the flower. Those nectars seldom involved in pollination. Most EFN secretions contain sucrose, glucose, fructose, and some amino acids. These sugars attract ants, parasitoids and many predators. Ants come for extrafloral nectar may prey on or disturb herbivores, thus formed the mutualism between plant and ant. We studied the mutualism relationship between ant and plant in the natural habitat of Passiflora siamica with two EFN plants Passiflora siamica (endemic) and Passifora coccinea (introduced). The main results are as follow:
1. At least 24 ant species were recorded utilizing the EFN resource. Include Paratrechina sp.1, Dolichoderus thoracicus (Smith), Tapinoma melanocephalum, Pheidole yeensis. One species from Ponerinae, one from Pseudomyrmecinae, nine from Myrmicinae, two from Dolichoderinae, eleven from Formicinae.
2. No significant difference was found between the number of ants visit two species of plants (t=1.30, P=0.20), while the compositing proportions of the main four ant species were different in the two Passiflora plants (x2=14.76, df=4, P< 0.01).The quantity of nectar secreted by a single leaf of Passiflora siamica is significantly higher than that of Passiflora coccinea plants (Willcox, z=-3.87, P< 0.01).Ant-excluding in this experiment do not significantly affected the average leaf loss per plant. For each species of plant, no significant difference of leaf nectar quantity was found among treatments.
3. The leaf trichome density of both side of Passiflora siamica leaf were significant lower than Passiflora coccinea, (upside leaf, t=-3.28, P< 0.01; underside leaf, t=-26.84, P< 0.01). In Passiflora siamica, the average leaf loss per plant was significant correlated to leaf trichome density of the undersider leaf (Pearson correlation, r=-0.38, P=0.04, n=30), but in Passiflora coccinea wasn't.
4. The average leaf loss (Log(x+1) transformed) of P. siamica was significantly correlated with the number of ants per plant (square root transformed) (Pearson correlation, r=-0.45, P< 0.05, n= 20). Ant behavioral dominance index (BDI, an index calculate as the proportion of dominance times to total encounter times of a species of ant to all other ants species in the same community) was significantly correlated with species total abundance on plants (Pearson correlation, r=0.86, P =0.01).
As a whole, the plant-ant-generalist herbivore in the two Passiflora species did not show a significant differences in the response to the change, while the interactive system with specialist herbivore request further investigation.
1、在两种西番莲植株上共观察到24种蚂蚁取食花外蜜液,主要包括：立毛蚁一种Paratrechina sp.1、黑可可臭蚁Dolichoderus thoracicus (Smith)、黑头酸臭蚁Tapinoma melanocephalum、伊大头蚁Pheidole yeensis等,分别隶属于猛蚁亚科(1属1种),伪切叶蚁亚科(1属1种),切叶蚁亚科(5属9种)、臭蚁亚科(2属2种)和蚁亚科(4属11种)。
2、利用两种西番莲的蚂蚁数量没有显著差异(t=1.30,P=0.20),但主要的四种蚂蚁的种类组成比例存在显著差异(χ2=14.76,df=4,P<0.01)。长叶西番莲单片叶片泌蜜量显著高于红花西番莲(Willcox,z=-3.87,P<0.01)。去除蚂蚁处理对两种西番莲植物的平均叶面积损失均未造成显著影响。两种西番莲去蚂蚁、去取食者、去蚂蚁及取食者处理之间叶片泌蜜量均没有显著差异。
3、长叶西番莲叶片表皮毛密度显著低于红花西番莲(叶子上表面,t=3.28,P<0.01：叶子下表面,t=26.84,P<0.01)。长叶西番莲的叶片下表面表皮毛密度和植株平均叶面积损失呈显著负相关关系(Pearson correlation, r=-0.38, P=0.04,n=30),而红花西番莲的叶片表皮毛密度和植株平均叶面积损失不显著相关。
4、长叶西番莲植株上蚂蚁数量和平均叶面积损失之间呈显著的负相关关系(Pearson correlation, r=-0.45,P< 0.05, n=20)。利用两种西番莲植株花外蜜腺的蚂蚁群体中不同种类的蚂蚁的行为优势度指数BDI(表示一种蚂蚁在和该群体中其它所有蚂蚁相遇时占优势的比例的指数)和其数量呈显著的正相关关系(Pearson correlation,r=0.86,P=0.01,n=60).
综上所述,蚂蚁和花外蜜腺去除并未显著影响植株被广适性昆虫取食程度；但长叶西番莲植株上蚂蚁数量还是与植株叶片被广适性昆虫取食的程度呈显著负相关关系。而该系统对专食性昆虫取食的反应如何则有待进一步研究。
EFN (extrafloral nectary) plant is a special group of plants. They have nectaries on the other part of the plant more than just in the flower. Those nectars seldom involved in pollination. Most EFN secretions contain sucrose, glucose, fructose, and some amino acids. These sugars attract ants, parasitoids and many predators. Ants come for extrafloral nectar may prey on or disturb herbivores, thus formed the mutualism between plant and ant. We studied the mutualism relationship between ant and plant in the natural habitat of Passiflora siamica with two EFN plants Passiflora siamica (endemic) and Passifora coccinea (introduced). The main results are as follow:
1. At least 24 ant species were recorded utilizing the EFN resource. Include Paratrechina sp.1, Dolichoderus thoracicus (Smith), Tapinoma melanocephalum, Pheidole yeensis. One species from Ponerinae, one from Pseudomyrmecinae, nine from Myrmicinae, two from Dolichoderinae, eleven from Formicinae.
2. No significant difference was found between the number of ants visit two species of plants (t=1.30, P=0.20), while the compositing proportions of the main four ant species were different in the two Passiflora plants (x2=14.76, df=4, P< 0.01).The quantity of nectar secreted by a single leaf of Passiflora siamica is significantly higher than that of Passiflora coccinea plants (Willcox, z=-3.87, P< 0.01).Ant-excluding in this experiment do not significantly affected the average leaf loss per plant. For each species of plant, no significant difference of leaf nectar quantity was found among treatments.
3. The leaf trichome density of both side of Passiflora siamica leaf were significant lower than Passiflora coccinea, (upside leaf, t=-3.28, P< 0.01; underside leaf, t=-26.84, P< 0.01). In Passiflora siamica, the average leaf loss per plant was significant correlated to leaf trichome density of the undersider leaf (Pearson correlation, r=-0.38, P=0.04, n=30), but in Passiflora coccinea wasn't.
4. The average leaf loss (Log(x+1) transformed) of P. siamica was significantly correlated with the number of ants per plant (square root transformed) (Pearson correlation, r=-0.45, P< 0.05, n= 20). Ant behavioral dominance index (BDI, an index calculate as the proportion of dominance times to total encounter times of a species of ant to all other ants species in the same community) was significantly correlated with species total abundance on plants (Pearson correlation, r=0.86, P =0.01).
As a whole, the plant-ant-generalist herbivore in the two Passiflora species did not show a significant differences in the response to the change, while the interactive system with specialist herbivore request further investigation.
引文
Andersen, A. N.1995. A classification of Australian ant communities, based on functional groups which parallel plant life-forms in relation to stress and disturbance. Journal of Biogeography 22:15-29.
Andres, M. R., and E. F. Connor.2003. The community-wide and guild-specific effects of pubescence on the folivorous insects of manzanitas. Ecological Entomology 28:383-396.
Apple, J. L., and D. H. Feener, Jr.2001. Ant visitation of extrafloral nectaries of Passiflora:The effects of nectary attributes and ant behavior on patterns in facultative ant-plant mutualisms. Oecologia (Berlin) 127:409-416.
Baker, D. A., J. L. Hall, and J. R. Thorpe.1978. A study of the extrafloral nectaries of Ricinus communis New Phytologist 81:129-137.
Baur, R., S. Binder, and G. Benz.1991. Nonglandular leaf trichomes as short-term inducible defense of the grey alder, Alnus incana (L.), against the chrysomelid beetle, Agelastica alni L. Oecologia 87:219-226.
Becerra, J., and D. Venable.1989. Extrafloral nectaries:a defense against ant-Homoptera mutualisms?. Oikos 55:276-280.
Benson, W. W., K. S. Brown Jr, and L. E. Gilbert.1975. Coevolution of Plants and Herbivores:Passion Flower Butterflies. Evolution 29:659-680.
Bentley, B. L.1976. Plants bearing extrafloral nectaries and the associated ant community:Interhabitat differences in the reduction of herbivore damage. Ecology 57:815-820.
Bentley, B. L.1977a. Extrafloral nectaries and protection by pugnacious bodyguards. Annual Review of Ecology and Systematics 8:407-427.
Bentley, B. L.1977b. Extrafloral nectaries and protection by pugnacious bodyguards. Annual Review of Ecology and Systematics 8:407-427.
Billington, H. L., C. D. Thomas, and L. E. Gilbert.1990. Variation in Stage-Specific Mortality Patterns of a Specialist Herbivore on Different Host Plant Clones. Functional Ecology 4:721-726.
Bluethgen, N., and K. Fiedler.2004a. Competition for composition:lessons from nectar-feeding ant communities. Ecology (Washington D C) 85:1479-1485.
Bluethgen, N., and K. Fiedler.2004b. Preferences for sugars and amino acids and their conditionality in a diverse nectar-feeding ant community. Journal of Animal Ecology 73:155-166.
Buono, R. A., A. B. de Oliveira, and E. A. S. Paiva.2008. Anatomy, Ultrastructure and Chemical Composition of Food Bodies of Hovenia dulcis (Rhamnaceae). Annals of Botany 101:1341.
Cardoso, M. Z.2008. Herbivore handling of a Plant's trichome:the case of Heliconius charithonia (L.)(Lepidoptera:Nymphalidae) and Passiflora lobata (Killip) Hutch.(Passifloraceae). Neotrop. entomol 37:247-252.
Cuautle, M., and V. Rico-Gray.2003. The effect of wasps and ants on the reproductive success of the extrafloral nectaried plant Turnera ulmifolia (Turneraceae). Functional Ecology 17:417-423.
Davidson, D. W.1997. The role of resource imbalances in the evolutionary ecology of tropical arboreal ants. Biological Journal of the Linnean Society 61:153-181.
de la Fuente, M. A. S., and R. J. Marquis.1999a. The role of ant-tended extrafloral nectaries in the protection and benefit of a Neotropical rainforest tree. Oecologia 118:192-202.
de la Fuente, M. A. S., and R. J. Marquis.1999b. The role of ant-tended extrafloral nectaries in the protection and benefit of a Neotropical rainforest tree. Oecologia (Berlin) 118:192-202.
Dejean, A., and B. Corbara.2003. A review of mosaics of dominant ants in rainforests and plantations. In:Arthropods of Tropical Forests:Spatio-Temporal Dynamics and Resource Use in the Canopy (ed. Y. Basset, V. Novotny, S. E. Miller & R. L. Kitching), pp.341-347. Cambrige University Press, Cambridge.
Djieto-Lordon, C., A. Dejean, M. Gibernau et al.2004. Symbiotic mutualism with a community of opportunistic ants:protection, competition, and ant occupancy of the myrmecophyte Barteria nigritana (Passifloraceae). Acta Oecologica 26:109-116.
Doak, P., D. Wagner, and A. Watson.2007. Variable extrafloral nectary expression and its consequences in quaking aspen. Canadian Journal of Botany 85:1-9.
Elias, T. S.1983. Extrafloral nectaries:their structure and distribution. In:The Biology of Nectaries, (ed. B Bentley, TS Elias), pp.174-203. Columbia Univ. Press, New York
Federle, W., U. Maschwitz, B. Fiala, M. Riederer, and B. Hoelldobler.1997. Slippery ant-plants and skillful climbers:Selection and protection of specific ant partners by epicuticular wax blooms in Macaranga (Euphorbiaceae). Oecologia (Berlin) 112:217-224.
Federle, W., U. Maschwitz, and B. Holldobler.2002. Pruning of host plant neighbours as defence against enemy ant invasions:Crematogaster ant partners of Macaranga protected by "wax barriers" prune less than their congeners. Oecologia (Berlin) 132:264-270.
Fiala, B., H. Grunsky, U. Maschwitz, and K. E. Linsenmair.1994. Diversity of ant-plant interactions: Protective efficacy in Macaranga species with different degrees of ant association. Oecologia (Heidelberg) 97:186-192.
Fischer, E., and I. R. Leal.2006. Effect of nectar secretion Tate on pollination success of Passiflora coccinea (Passifloraceae) in the Central Amazon. Brazilian Journal of Biology 66:747-754.
Freitas, L., L. Galetto, G. Bernardello, and A. A. S. Paoli.2000. Ant exclusion and reproduction of Croton sarcopetalus (Euphorbiaceae). Flora (Jena) 195:398-402.
Galen, C.2005. Catching ants with honey:an experimental test of distraction and satiation as alternative modes of escape from flower-damaging ants. Oecologia 144:80-87.
Gaume, L., and D. McKey.1999. An ant-plant mutualism and its host-specific parasite:activity rhythms, young leaf patrolling, and effects on herbivores of two specialist plant-ants inhabiting the same myrmecophyte. Oikos 84:130-144.
Gaume, L., D. McKey, and M. C. Anstett.1997. Benefits conferred by" timid" ants:active anti-herbivore protection of the rainforest tree Leonardoxa africana by the minute ant Petalomyrmex phylax. Oecologia 112:209-216.
Gaume, L., D. McKey, and S. Terrin.1998. Ant-plant-homopteran mutualism:How the third partner affects the interaction between a plant-specialist ant and its myrmecophyte host. Proceedings of the Royal Society of London Series B Biological Sciences 265:569-575.
Gaume, L., M. Zacharias, and R. M. Borges.2005. Ant-plant conflicts and a novel case of castration parasitism in a myrmecophyte. Evolutionary Ecology Research 7:435-452.
Gentry, G.2003. Multiple parasitoid visitors to the extrafloral nectaries of Solanum'adherens. Is S. adherens an insectary plant? Basic and Applied Ecology 4:405-411.
Gill, F. B.1988. Effects of nectar removal on nectar accumulation in flowers of Heliconia imbricata (Heliconiaceae). Biotropica 20(2):169-171.
Gruner, D. S., A. D. Taylor, and R. E. Forkner.2005. The effects of foliar pubescence and nutrient enrichment on arthropod communities of Metrosideros polymorpha (Myrtaceae). Ecological Entomology 30:428-443.
Guimaraes, P. R., Jr., R. L. G Raimundo, C. Bottcher, R. R. Silva, and J. R. Trigo.2006. Extrafloral nectaries as a deterrent mechanism against seed predators in the chemically protected weed Crotalaria pallida (Leguminosae). Austral Ecology 31:776-782.
Heil, M.2004. Induction of two indirect defences benefits Lima bean (Phaseolus lunatus, Fabaceae) in nature. Journal of Ecology 92:527-536.
Heil, M., S. Bueno, and J. Carlos.2007. Within-plant signaling by volatiles leads to induction and priming of an indirect plant defense in nature. Proceedings of the National Academy of Sciences 104:5467-5472.
Heil, M., B. Fiala, B. Baumann, and K. E. Linsenmair.2000. Temporal, spatial and biotic variations in extrafloral nectar secretion by Macaranga tanarius. Functional Ecology 14:749-757.
Heil, M., A. Hilpert, R. Kruger, and K. E. Linsenmair.2004. Competition among visitors to extrafloral nectaries as a source of ecological costs of an indirect defence. Journal of Tropical Ecology 20:201-208.
Heil, M., T. Koch, A. Hilpert, B. Fiala, W. Boland, and K. E. Linsenmair.2001. Extrafloral nectar production of the ant-associated plant, Macaranga tanarius, is an induced, indirect, defensive response elicited by jasmonic acid. Proceedings of the National Academy of Sciences of the United States of America 98:1083-1088.
Heil, M., and C. Kost.2006. Priming of indirect defences. Ecology Letters 9:813-817.
Heil, M., and D. McKey.2003a. Protective ant-plant interactions as model systems in ecological and evolutionary research. Annual Review of Ecology Evolution and Systematics 34:425-453.
Heil, M., and D. McKey.2003b. Protective ant-plant interactions as model systems in ecological and evolutionary research. Annual Review of Ecology Evolution and Systematics.34:425-553.
Heil, M., J. Rattke, and W. Boland.2005. Postsecretory hydrolysis of nectar sucrose and specialization in ant/plant mutualism. Science (Washington D C) 308:560-563.
Holldobler, B., and C. J. Lumsden.1980. Territorial Strategies in Ants. Science 210:732-739.
Horvitz, C. C., and D. W. Schemske.1984. Effects of ants and an ant-tended herbivore on seed production of a Neotropical herb. Ecology 65:1369-1378.
Hossaert-McKey, M., J. Orivel, E. Labeyrie, et al..2001. Differential associations with ants of three co-occurring extrafloral nectary-bearing plants. Ecoscience 8:325-335.
Inouye, D. W., and O. R. Taylor, Jr.1979. A temperate region plant-ant-seed predator system: Consequences of extra floral nectar secretion by Helianthella quinquenervis. Ecology 60:1-7.
Itioka, T., M. Nomura, Y. Inui, T. Itino, and T. Inoue.2000. Difference in intensity of ant defense among three species of Macaranga myrmecophytes in a southeast Asian dipterocarp forest. Biotropica 32:318-326.
Janzen, D. H.1966. Coevolution of mutualism between ants and acacias in Central America. Evolution 20:249-275.
Katayama, N., and N. Suzuki.2003. Changes in the use of extrafloral nectaries of Vicia faba (Leguminosae) and honeydew of aphids by ants with increasing aphid density. Annals of the Entomological Society of America 96:579-584.
Keeler, K. H.1981. Function of Mentzelia nuda (Loasaceae) postfloral nectaries in seed defense. American Journal of Botany 68:295-299.
Kelly, C. A.1986. Extrafloral nectaries:ants, herbivores and fecundity in Cassia fasciculate Oecologia 69:600-605.
Kenzo, T., R. Yoneda, M. A. Azani, and N. M. Majid.2008. Changes in leaf water use after removal of leaf lower surface hairs on Mallotus macrostachyus (Euphorbiaceae) in a tropical secondary forest in Malaysia. Journal of Forest Research 13:137-142.
Kersch, M. F., and C. R. Fonseca.2005. Abiotic factors and the conditional outcome of an ant-plant mutualism. Ecology 86:2117-2126.
Knox, R. B., J. Kenrick, P. Bernhardt, et al.1985. Extrafloral nectaries as adaptations for bird pollination in Acacia terminalis. American Journal of Botany 72:1185-1196.
Koptur, S.1979. Facultative mutualism between weedy vetches bearing extrafloral nectaries and weedy ants in California. American Journal of Botany 66:1016-1020.
Koptur, S.1983. Flowering phenology and floral biology of Inga (Fabaceae:Mimosoideae). Systematic botany 8(4):354-368.
Koptur, S.1984. Experimental evidence for defense of Inga (Mimosoideae) saplings by ants. Ecology 65:1787-1793.
Koptur, S.1985. Alternative defenses against herbivores in Inga (Fabaceae:Mimosoideae) over an elevational gradient. Ecology 66:1639-1650.
Koptur, S.1991. Extrafloral nectaries of herbs and trees:modelling the interaction with ants and parasitoids. In:Ant-plant interactions.(ed. C. R. H. a. D. F. Cutler), pp.213-230. Oxford University Press, Oxford.
Koptur, S.1994. Floral and extrafloral nectars of Costa Rican Inga trees:A comparison of their constituents and composition. Biotropica 26:276-284.
Koptur, S.2005. Nectar as fuel for plant protectors. In:Plant-Provided Food for Carnivorous Insects:A Protective Mutualism and Its Applications.(ed. Wackers FL, Van Rijn PCJ, Bruin J) pp.75-108. Cambridge University Press, Cambridge
Kost, C., and M. Heil.2005. Increased availability of Extrafloral nectar reduces herbivory in Lima bean plants (Phaseolus lunatus., Fabaceae). Basic Appl. Ecol.6:237-248.
Kost, C., and M. Heil.2006. Herbivore-induced plant volatiles induce an indirect defence in neighbouring plants. Journal of Ecology 94:619-628.
Lanza, J.1988. Ant Preferences for Passiflora Nectar Mimics That Contain Amino Acids. Biotropica 20(4):341-344.
Leal, I. R., E. Fischer, C. Kost, M. Tabarelli, and R. Wirth.2006. Ant protection against herbivores and nectar thieves in Passiflora coccinea flowers. Ecoscience 13:431-438.
Levin, D. A.1973. The role of trichomes in plant defense. The quarterly review of biology 48:3-15.
Lorch, J.1978. The discovery of nectar and nectaries and its relation to views on flowers and insects. Isis 69:514-533.
MacDougal, J. M.1994. Revision of passiflora subgenus decaloba section pseudodysosmia (passifloraceae). Systematic Botany Monographs 41:1-146.
McKey, D.1974. Adaptive Patterns in Alkaloid Physiology. American Naturalist 108:305-320.
却舍不得让你难过啊McLain, D. K.1983. Ants, Extrafloral Nectaries and Herbivory on the Passion Vine, Passiflora incarnata. American Midland Naturalist 110:433-439.
Medeiros, L., and G. R. P. Moreira.2005. Larval feeding behavior of Gratiana spadicea (Klug)(Coleoptera:Chrysomelidae:Cassidinae) on its host plant, Solanum sisymbriifolium Lamarck (Solanaceae):interaction with trichomes. The Coleopterists Bulletin 59:339-350.
Meunier, L., A. Dalecky, C. Berticat, L. Gaume, and D. McKey.1999. Worker size variation and the evolution of an ant-plant mutualism:Comparative morphometrics of workers of two closely related plant-ants, Petalomyrmex phylax and Aphomomyrmex afer (Formicinae). Insectes Sociaux 46:171-178.
Miller, T. E. X.2007. Does having multiple partners weaken the benefits of facultative mutualism? A test with cacti and cactus-tending ants. Oikos 116:500-512.
Mondor, E. B., and J. F. Addicott.2003. Conspicuous extra-floral nectaries are inducible in Vicia faba. Ecology Letters 6:495-497.
Moya-Raygoza, G. 2005. Relationships between the ant Brachymyrmex obscurlor (Hymenoptera, Formicidae) and Acacia pennatula (Fabaceae). Insectes Sociaux 52:105-107.
Ness, J. H., W. F. Morris, and J. L. Bronstein.2006. Integrating quality and quantity of mutualistic service to contrast ant species protecting Ferocactus wislizeni. Ecology(Durham) 87:912-921.
O'Dowd, D. J.1979. Foliar nectar production and ant activity on a neotropical tree, Ochroma pyramidale. Oecologia 43:233-248.
Offenberg, J.2000. Correlated evolution of the association between aphids and ants and the associalion between aphids and plants with extrafloral nectaries. Oikos 91:146-152.
Oliveira, P. S., V. Rico-Gray, C. Diaz-Castelazo, and C. Castillo-Guevara.1999. Interaction between ants, extrafloral nectaries and insect herbivores in Neotropical coastal sand dunes:herbivore deterrence by visiting ants increases fruit set in Opuntia stricta (Cactaceae). Functional Ecology 13:623-631.
Palmer, T. M., and A. K. Brody.2007. Mutualism as reciprocal exploitation:African plant-ants defend foliar but not reproductive structures. Ecology (Washington D C) 88:3004-3011.
Palmer, T. M., T. P. Young, and M. L. Stanton.2002. Burning bridges:Priority effects and the persistence of a competitively subordinate acacia-ant in Laikipia, Kenya. Oecologia (Berlin) 133:372-379.
Palmer, T. M., T. P. Young, M. L. Stanton, and E. Wenk.2000. Short-term dynamics of an acacia ant community in Laikipia, Kenya. Oecologia (Berlin) 123:425-435.
Pemberton, R. W., and J. H. Lee.1996. The influence of extrafloral nectaries on parasitism of an insect herbivore. American Journal of Botany 83:1187-1194.
Pickett, C. H., and W. D. Clark.1979. The function of extrafloral nectaries in Opuntia acanthocarpa (Cactaceae). American Journal of Botany 66:618-625.
Pulice, C. E., and A. A. Packer.2008. Simulated herbivory induces extrafloral nectary production in Prunus avium. Functional Ecology 22:801-807.
Pyke, G. H.1991. What does it cost a plant to produce floral nectar? Nature 350:58-59.
Queiroz, J. M., and P. S. Oliveira.2001. Tending ants protect honeydew-producing whiteflies (Homoptera:Aleyrodidae). Environmental Entomology 30:295-297.
Raine, N. E., N. Gammans, I. J. Macfadyen, G. K. Scrivner, and G. N. Stone.2004. Guards and thieves: antagonistic interactions between two ant species coexisting on the same ant-plant. Ecological Entomology 29:345-352.
Raine, N. E., P. Willmer, and G. N. Stone.2002. Spatial structuring and floral avoidance behavior prevent ant-pollinator conflict in a Mexican ant-acacia. Ecology 83:3086-3096.
Romero, G. Q., and J. Vasconcellos-Neto.2004. Beneficial effects of flower-dwelling predators on their host plant. Ecology 85:446-457.
Rudgers, J. A.2004a. Enemies of herbivores can shape plant traits:Selection in a facultative ant-plant mutualism. Ecology (Washington D C) 85:192-205.
Rudgers, J. A.2004b. Enemies of herbivores can shape plant traits:Selection in a facultative ant-plant mutualism. Ecology 85:192-205.
Rudgers, J. A., and M. C. Gardener.2004. Extrafloral nectar as a resource mediating multispecies infractions. Ecology 85:1495-1502.
Rudgers, J. A., and S. Y. Strauss.2004. A selection mosaic in the facultative mutualism between ants and wild cotton. Proceedings of the Royal Society Biological Sciences Series B 271:2481-2488.
Ruhren, S., and S. N. Handel.1999. Jumping spiders (Salticidae) enhance the seed production of a plant with extrafloral nectaries. Oecologia (Berlin) 119:227-230.
Santini, G., L. Tucci, L. Ottonetti, and F. Frizzi.2007. Competition trade-offs in the organisation of a Mediterranean ant assemblage. Ecological Entomology 32:319-326.
Savolainen, R., and K. Vepsalainen.1988. A competition hierarchy among boreal ants:impact on resource partitioning and community structure. Oikos 51:135-155.
Schappert, P. J., and J. S. Shore.1999. Effects of cyanogenesis polymorphism in Turnera ulmifolia on Euptoieta hegesia and potential Anolis predators. Journal of Chemical Ecology 25:1455-1479.
Schmid, R.1988. Reproductive versus extra-reproductive nectaries:historical perspectives and terminological recommendations. The Botanical Review 54:179-232.
Smiley, J.1986. Ant Constancy at Passiflora Extrafloral Nectaries:Effects on Caterpillar Survival. Ecology 67(2):516-521.
Stanton, M. L., T. M. Palmer, and T. P. Young.2002. Competition-colonization trade-offs in a guild of African acacia-ants. Ecological Monographs 72:347-363.
Stapel, J. O., A. M. Cortesero, C. M. De Moraes, J. H. Tumlinson, and W. J. Lewis.1997. Extrafloral nectar, honeydew, and sucrose effects on searching behavior and efficiency of Microplitis croceipes (Hymenoptera:Braconidae) in cotton. Environ. Entomol.26:617-623.
Storti, E. F.2002. Pollination biology and breeding system of Passiflora coccinea Aubl. in Manaus, Amazonas, Brazil. Acta Amazonica 32:421-429.
Suarez, A. V., C. Moraes, and A. Ippolito.1998. Defense of Acacia collinsii by an Obligate and Nonobligate Ant Species:the Significance of Encroaching Vegetation 1. Biotropica 30:480-482.
Tilman, D.1978. Cherries, Ants and Tent Caterpillars:Timing of Nectar Production in Relation in Relation to Susceptibility of Catepillars to Ant Predation. Ecology 59:686-692.
Van Mele, P., and N. T. T. Cuc.2001. Farmers'Perceptions and Practices in Use of Dolichoderus thoracicus (Smith)(Hymenoptera:Formicidae) for Biological Control of Pests of Sapodilla. Biological Control 20:23-29.
Vanderplank, J.1996. Passion Flowers. The MIT, Cambridge
Wackers, F. L., and C. Bonifay.2004. How to be sweet? Extrafloral nectar allocation by Gossypium hirsutum fits optimal defense theory predictions. Ecology 85:1512-1518.
Wagner, D.1997. The influence of ant nests on Acacia seed production, herbivory and soil nutrients. Journal of Ecology 85:83-93.
Wagner, D., and A. Kay.2002. Do extrafloral nectaries distract ants from visiting flowers? An experimental test of an overlooked hypothesis. Evol. Ecol. Res.4:293-305.
Way, M. J., and K. C. Khoo.1991. Colony dispersion and nesting habits of the ants, Dolichoderus thoracicus and Oecophylla smaragdina(Hymenoptera:Formicidae), in relation to their success as biological control agents on cocoa. Bulletin of Entomological Research 81:341-350.
Wheeler, H. C., and C. C. Bennington.2001. Effects of light availability on herbivory and defense chemistry in Passiflora incarnata. Florida Scientist 64:237-245.
Willmer, P. G., and G. N. Stone.1997. How aggressive ant-guards assist seed-set in Acacia flowers. Nature (London) 388:165-167.
Wirth, R., and I. R. Leal.2001. Does rainfall affect temporal variability of ant protection in Passiflora coccinea? Ecoscience 8:450-453.
Young, T. P., C. H. Stubblefield, and L. A. Isbell.1997. Ants on swollen-thorn acacias:species coexistence in a simple system. Oecologia 109:98-107.
方文培,张泽荣.1999《中国植物志》,第52(1)卷.北京：科学出版社,1983.pp 97-101
李绍文.1991.生态生物化学：(八)植食性昆虫对食物的选择.生态学杂志(Chinese journal of ecology)10:65-70.
张丽霞.2002.五种杀虫剂防治西番莲大斑芫菁的药效试验.植物保护(Plant protection) 28:51-53
杨萍,邓合黎,漆波,and刘琼.2006.红锯蛱蝶Cethosia biblis (Drury)的生物学研究.西南农业大学学报28：549-552.
Andres, M. R., and E. F. Connor.2003. The community-wide and guild-specific effects of pubescence on the folivorous insects of manzanitas. Ecological Entomology 28:383-396.
Apple, J. L., and D. H. Feener, Jr.2001. Ant visitation of extrafloral nectaries of Passiflora:The effects of nectary attributes and ant behavior on patterns in facultative ant-plant mutualisms. Oecologia (Berlin) 127:409-416.
Baker, D. A., J. L. Hall, and J. R. Thorpe.1978. A study of the extrafloral nectaries of Ricinus communis New Phytologist 81:129-137.
Baur, R., S. Binder, and G. Benz.1991. Nonglandular leaf trichomes as short-term inducible defense of the grey alder, Alnus incana (L.), against the chrysomelid beetle, Agelastica alni L. Oecologia 87:219-226.
Becerra, J., and D. Venable.1989. Extrafloral nectaries:a defense against ant-Homoptera mutualisms?. Oikos 55:276-280.
Benson, W. W., K. S. Brown Jr, and L. E. Gilbert.1975. Coevolution of Plants and Herbivores:Passion Flower Butterflies. Evolution 29:659-680.
Bentley, B. L.1976. Plants bearing extrafloral nectaries and the associated ant community:Interhabitat differences in the reduction of herbivore damage. Ecology 57:815-820.
Bentley, B. L.1977a. Extrafloral nectaries and protection by pugnacious bodyguards. Annual Review of Ecology and Systematics 8:407-427.
Bentley, B. L.1977b. Extrafloral nectaries and protection by pugnacious bodyguards. Annual Review of Ecology and Systematics 8:407-427.
Billington, H. L., C. D. Thomas, and L. E. Gilbert.1990. Variation in Stage-Specific Mortality Patterns of a Specialist Herbivore on Different Host Plant Clones. Functional Ecology 4:721-726.
Bluethgen, N., and K. Fiedler.2004a. Competition for composition:lessons from nectar-feeding ant communities. Ecology (Washington D C) 85:1479-1485.
Bluethgen, N., and K. Fiedler.2004b. Preferences for sugars and amino acids and their conditionality in a diverse nectar-feeding ant community. Journal of Animal Ecology 73:155-166.
Buono, R. A., A. B. de Oliveira, and E. A. S. Paiva.2008. Anatomy, Ultrastructure and Chemical Composition of Food Bodies of Hovenia dulcis (Rhamnaceae). Annals of Botany 101:1341.
Cardoso, M. Z.2008. Herbivore handling of a Plant's trichome:the case of Heliconius charithonia (L.)(Lepidoptera:Nymphalidae) and Passiflora lobata (Killip) Hutch.(Passifloraceae). Neotrop. entomol 37:247-252.
Cuautle, M., and V. Rico-Gray.2003. The effect of wasps and ants on the reproductive success of the extrafloral nectaried plant Turnera ulmifolia (Turneraceae). Functional Ecology 17:417-423.
Davidson, D. W.1997. The role of resource imbalances in the evolutionary ecology of tropical arboreal ants. Biological Journal of the Linnean Society 61:153-181.
de la Fuente, M. A. S., and R. J. Marquis.1999a. The role of ant-tended extrafloral nectaries in the protection and benefit of a Neotropical rainforest tree. Oecologia 118:192-202.
de la Fuente, M. A. S., and R. J. Marquis.1999b. The role of ant-tended extrafloral nectaries in the protection and benefit of a Neotropical rainforest tree. Oecologia (Berlin) 118:192-202.
Dejean, A., and B. Corbara.2003. A review of mosaics of dominant ants in rainforests and plantations. In:Arthropods of Tropical Forests:Spatio-Temporal Dynamics and Resource Use in the Canopy (ed. Y. Basset, V. Novotny, S. E. Miller & R. L. Kitching), pp.341-347. Cambrige University Press, Cambridge.
Djieto-Lordon, C., A. Dejean, M. Gibernau et al.2004. Symbiotic mutualism with a community of opportunistic ants:protection, competition, and ant occupancy of the myrmecophyte Barteria nigritana (Passifloraceae). Acta Oecologica 26:109-116.
Doak, P., D. Wagner, and A. Watson.2007. Variable extrafloral nectary expression and its consequences in quaking aspen. Canadian Journal of Botany 85:1-9.
Elias, T. S.1983. Extrafloral nectaries:their structure and distribution. In:The Biology of Nectaries, (ed. B Bentley, TS Elias), pp.174-203. Columbia Univ. Press, New York
Federle, W., U. Maschwitz, B. Fiala, M. Riederer, and B. Hoelldobler.1997. Slippery ant-plants and skillful climbers:Selection and protection of specific ant partners by epicuticular wax blooms in Macaranga (Euphorbiaceae). Oecologia (Berlin) 112:217-224.
Federle, W., U. Maschwitz, and B. Holldobler.2002. Pruning of host plant neighbours as defence against enemy ant invasions:Crematogaster ant partners of Macaranga protected by "wax barriers" prune less than their congeners. Oecologia (Berlin) 132:264-270.
Fiala, B., H. Grunsky, U. Maschwitz, and K. E. Linsenmair.1994. Diversity of ant-plant interactions: Protective efficacy in Macaranga species with different degrees of ant association. Oecologia (Heidelberg) 97:186-192.
Fischer, E., and I. R. Leal.2006. Effect of nectar secretion Tate on pollination success of Passiflora coccinea (Passifloraceae) in the Central Amazon. Brazilian Journal of Biology 66:747-754.
Freitas, L., L. Galetto, G. Bernardello, and A. A. S. Paoli.2000. Ant exclusion and reproduction of Croton sarcopetalus (Euphorbiaceae). Flora (Jena) 195:398-402.
Galen, C.2005. Catching ants with honey:an experimental test of distraction and satiation as alternative modes of escape from flower-damaging ants. Oecologia 144:80-87.
Gaume, L., and D. McKey.1999. An ant-plant mutualism and its host-specific parasite:activity rhythms, young leaf patrolling, and effects on herbivores of two specialist plant-ants inhabiting the same myrmecophyte. Oikos 84:130-144.
Gaume, L., D. McKey, and M. C. Anstett.1997. Benefits conferred by" timid" ants:active anti-herbivore protection of the rainforest tree Leonardoxa africana by the minute ant Petalomyrmex phylax. Oecologia 112:209-216.
Gaume, L., D. McKey, and S. Terrin.1998. Ant-plant-homopteran mutualism:How the third partner affects the interaction between a plant-specialist ant and its myrmecophyte host. Proceedings of the Royal Society of London Series B Biological Sciences 265:569-575.
Gaume, L., M. Zacharias, and R. M. Borges.2005. Ant-plant conflicts and a novel case of castration parasitism in a myrmecophyte. Evolutionary Ecology Research 7:435-452.
Gentry, G.2003. Multiple parasitoid visitors to the extrafloral nectaries of Solanum'adherens. Is S. adherens an insectary plant? Basic and Applied Ecology 4:405-411.
Gill, F. B.1988. Effects of nectar removal on nectar accumulation in flowers of Heliconia imbricata (Heliconiaceae). Biotropica 20(2):169-171.
Gruner, D. S., A. D. Taylor, and R. E. Forkner.2005. The effects of foliar pubescence and nutrient enrichment on arthropod communities of Metrosideros polymorpha (Myrtaceae). Ecological Entomology 30:428-443.
Guimaraes, P. R., Jr., R. L. G Raimundo, C. Bottcher, R. R. Silva, and J. R. Trigo.2006. Extrafloral nectaries as a deterrent mechanism against seed predators in the chemically protected weed Crotalaria pallida (Leguminosae). Austral Ecology 31:776-782.
Heil, M.2004. Induction of two indirect defences benefits Lima bean (Phaseolus lunatus, Fabaceae) in nature. Journal of Ecology 92:527-536.
Heil, M., S. Bueno, and J. Carlos.2007. Within-plant signaling by volatiles leads to induction and priming of an indirect plant defense in nature. Proceedings of the National Academy of Sciences 104:5467-5472.
Heil, M., B. Fiala, B. Baumann, and K. E. Linsenmair.2000. Temporal, spatial and biotic variations in extrafloral nectar secretion by Macaranga tanarius. Functional Ecology 14:749-757.
Heil, M., A. Hilpert, R. Kruger, and K. E. Linsenmair.2004. Competition among visitors to extrafloral nectaries as a source of ecological costs of an indirect defence. Journal of Tropical Ecology 20:201-208.
Heil, M., T. Koch, A. Hilpert, B. Fiala, W. Boland, and K. E. Linsenmair.2001. Extrafloral nectar production of the ant-associated plant, Macaranga tanarius, is an induced, indirect, defensive response elicited by jasmonic acid. Proceedings of the National Academy of Sciences of the United States of America 98:1083-1088.
Heil, M., and C. Kost.2006. Priming of indirect defences. Ecology Letters 9:813-817.
Heil, M., and D. McKey.2003a. Protective ant-plant interactions as model systems in ecological and evolutionary research. Annual Review of Ecology Evolution and Systematics 34:425-453.
Heil, M., and D. McKey.2003b. Protective ant-plant interactions as model systems in ecological and evolutionary research. Annual Review of Ecology Evolution and Systematics.34:425-553.
Heil, M., J. Rattke, and W. Boland.2005. Postsecretory hydrolysis of nectar sucrose and specialization in ant/plant mutualism. Science (Washington D C) 308:560-563.
Holldobler, B., and C. J. Lumsden.1980. Territorial Strategies in Ants. Science 210:732-739.
Horvitz, C. C., and D. W. Schemske.1984. Effects of ants and an ant-tended herbivore on seed production of a Neotropical herb. Ecology 65:1369-1378.
Hossaert-McKey, M., J. Orivel, E. Labeyrie, et al..2001. Differential associations with ants of three co-occurring extrafloral nectary-bearing plants. Ecoscience 8:325-335.
Inouye, D. W., and O. R. Taylor, Jr.1979. A temperate region plant-ant-seed predator system: Consequences of extra floral nectar secretion by Helianthella quinquenervis. Ecology 60:1-7.
Itioka, T., M. Nomura, Y. Inui, T. Itino, and T. Inoue.2000. Difference in intensity of ant defense among three species of Macaranga myrmecophytes in a southeast Asian dipterocarp forest. Biotropica 32:318-326.
Janzen, D. H.1966. Coevolution of mutualism between ants and acacias in Central America. Evolution 20:249-275.
Katayama, N., and N. Suzuki.2003. Changes in the use of extrafloral nectaries of Vicia faba (Leguminosae) and honeydew of aphids by ants with increasing aphid density. Annals of the Entomological Society of America 96:579-584.
Keeler, K. H.1981. Function of Mentzelia nuda (Loasaceae) postfloral nectaries in seed defense. American Journal of Botany 68:295-299.
Kelly, C. A.1986. Extrafloral nectaries:ants, herbivores and fecundity in Cassia fasciculate Oecologia 69:600-605.
Kenzo, T., R. Yoneda, M. A. Azani, and N. M. Majid.2008. Changes in leaf water use after removal of leaf lower surface hairs on Mallotus macrostachyus (Euphorbiaceae) in a tropical secondary forest in Malaysia. Journal of Forest Research 13:137-142.
Kersch, M. F., and C. R. Fonseca.2005. Abiotic factors and the conditional outcome of an ant-plant mutualism. Ecology 86:2117-2126.
Knox, R. B., J. Kenrick, P. Bernhardt, et al.1985. Extrafloral nectaries as adaptations for bird pollination in Acacia terminalis. American Journal of Botany 72:1185-1196.
Koptur, S.1979. Facultative mutualism between weedy vetches bearing extrafloral nectaries and weedy ants in California. American Journal of Botany 66:1016-1020.
Koptur, S.1983. Flowering phenology and floral biology of Inga (Fabaceae:Mimosoideae). Systematic botany 8(4):354-368.
Koptur, S.1984. Experimental evidence for defense of Inga (Mimosoideae) saplings by ants. Ecology 65:1787-1793.
Koptur, S.1985. Alternative defenses against herbivores in Inga (Fabaceae:Mimosoideae) over an elevational gradient. Ecology 66:1639-1650.
Koptur, S.1991. Extrafloral nectaries of herbs and trees:modelling the interaction with ants and parasitoids. In:Ant-plant interactions.(ed. C. R. H. a. D. F. Cutler), pp.213-230. Oxford University Press, Oxford.
Koptur, S.1994. Floral and extrafloral nectars of Costa Rican Inga trees:A comparison of their constituents and composition. Biotropica 26:276-284.
Koptur, S.2005. Nectar as fuel for plant protectors. In:Plant-Provided Food for Carnivorous Insects:A Protective Mutualism and Its Applications.(ed. Wackers FL, Van Rijn PCJ, Bruin J) pp.75-108. Cambridge University Press, Cambridge
Kost, C., and M. Heil.2005. Increased availability of Extrafloral nectar reduces herbivory in Lima bean plants (Phaseolus lunatus., Fabaceae). Basic Appl. Ecol.6:237-248.
Kost, C., and M. Heil.2006. Herbivore-induced plant volatiles induce an indirect defence in neighbouring plants. Journal of Ecology 94:619-628.
Lanza, J.1988. Ant Preferences for Passiflora Nectar Mimics That Contain Amino Acids. Biotropica 20(4):341-344.
Leal, I. R., E. Fischer, C. Kost, M. Tabarelli, and R. Wirth.2006. Ant protection against herbivores and nectar thieves in Passiflora coccinea flowers. Ecoscience 13:431-438.
Levin, D. A.1973. The role of trichomes in plant defense. The quarterly review of biology 48:3-15.
Lorch, J.1978. The discovery of nectar and nectaries and its relation to views on flowers and insects. Isis 69:514-533.
MacDougal, J. M.1994. Revision of passiflora subgenus decaloba section pseudodysosmia (passifloraceae). Systematic Botany Monographs 41:1-146.
McKey, D.1974. Adaptive Patterns in Alkaloid Physiology. American Naturalist 108:305-320.
却舍不得让你难过啊McLain, D. K.1983. Ants, Extrafloral Nectaries and Herbivory on the Passion Vine, Passiflora incarnata. American Midland Naturalist 110:433-439.
Medeiros, L., and G. R. P. Moreira.2005. Larval feeding behavior of Gratiana spadicea (Klug)(Coleoptera:Chrysomelidae:Cassidinae) on its host plant, Solanum sisymbriifolium Lamarck (Solanaceae):interaction with trichomes. The Coleopterists Bulletin 59:339-350.
Meunier, L., A. Dalecky, C. Berticat, L. Gaume, and D. McKey.1999. Worker size variation and the evolution of an ant-plant mutualism:Comparative morphometrics of workers of two closely related plant-ants, Petalomyrmex phylax and Aphomomyrmex afer (Formicinae). Insectes Sociaux 46:171-178.
Miller, T. E. X.2007. Does having multiple partners weaken the benefits of facultative mutualism? A test with cacti and cactus-tending ants. Oikos 116:500-512.
Mondor, E. B., and J. F. Addicott.2003. Conspicuous extra-floral nectaries are inducible in Vicia faba. Ecology Letters 6:495-497.
Moya-Raygoza, G. 2005. Relationships between the ant Brachymyrmex obscurlor (Hymenoptera, Formicidae) and Acacia pennatula (Fabaceae). Insectes Sociaux 52:105-107.
Ness, J. H., W. F. Morris, and J. L. Bronstein.2006. Integrating quality and quantity of mutualistic service to contrast ant species protecting Ferocactus wislizeni. Ecology(Durham) 87:912-921.
O'Dowd, D. J.1979. Foliar nectar production and ant activity on a neotropical tree, Ochroma pyramidale. Oecologia 43:233-248.
Offenberg, J.2000. Correlated evolution of the association between aphids and ants and the associalion between aphids and plants with extrafloral nectaries. Oikos 91:146-152.
Oliveira, P. S., V. Rico-Gray, C. Diaz-Castelazo, and C. Castillo-Guevara.1999. Interaction between ants, extrafloral nectaries and insect herbivores in Neotropical coastal sand dunes:herbivore deterrence by visiting ants increases fruit set in Opuntia stricta (Cactaceae). Functional Ecology 13:623-631.
Palmer, T. M., and A. K. Brody.2007. Mutualism as reciprocal exploitation:African plant-ants defend foliar but not reproductive structures. Ecology (Washington D C) 88:3004-3011.
Palmer, T. M., T. P. Young, and M. L. Stanton.2002. Burning bridges:Priority effects and the persistence of a competitively subordinate acacia-ant in Laikipia, Kenya. Oecologia (Berlin) 133:372-379.
Palmer, T. M., T. P. Young, M. L. Stanton, and E. Wenk.2000. Short-term dynamics of an acacia ant community in Laikipia, Kenya. Oecologia (Berlin) 123:425-435.
Pemberton, R. W., and J. H. Lee.1996. The influence of extrafloral nectaries on parasitism of an insect herbivore. American Journal of Botany 83:1187-1194.
Pickett, C. H., and W. D. Clark.1979. The function of extrafloral nectaries in Opuntia acanthocarpa (Cactaceae). American Journal of Botany 66:618-625.
Pulice, C. E., and A. A. Packer.2008. Simulated herbivory induces extrafloral nectary production in Prunus avium. Functional Ecology 22:801-807.
Pyke, G. H.1991. What does it cost a plant to produce floral nectar? Nature 350:58-59.
Queiroz, J. M., and P. S. Oliveira.2001. Tending ants protect honeydew-producing whiteflies (Homoptera:Aleyrodidae). Environmental Entomology 30:295-297.
Raine, N. E., N. Gammans, I. J. Macfadyen, G. K. Scrivner, and G. N. Stone.2004. Guards and thieves: antagonistic interactions between two ant species coexisting on the same ant-plant. Ecological Entomology 29:345-352.
Raine, N. E., P. Willmer, and G. N. Stone.2002. Spatial structuring and floral avoidance behavior prevent ant-pollinator conflict in a Mexican ant-acacia. Ecology 83:3086-3096.
Romero, G. Q., and J. Vasconcellos-Neto.2004. Beneficial effects of flower-dwelling predators on their host plant. Ecology 85:446-457.
Rudgers, J. A.2004a. Enemies of herbivores can shape plant traits:Selection in a facultative ant-plant mutualism. Ecology (Washington D C) 85:192-205.
Rudgers, J. A.2004b. Enemies of herbivores can shape plant traits:Selection in a facultative ant-plant mutualism. Ecology 85:192-205.
Rudgers, J. A., and M. C. Gardener.2004. Extrafloral nectar as a resource mediating multispecies infractions. Ecology 85:1495-1502.
Rudgers, J. A., and S. Y. Strauss.2004. A selection mosaic in the facultative mutualism between ants and wild cotton. Proceedings of the Royal Society Biological Sciences Series B 271:2481-2488.
Ruhren, S., and S. N. Handel.1999. Jumping spiders (Salticidae) enhance the seed production of a plant with extrafloral nectaries. Oecologia (Berlin) 119:227-230.
Santini, G., L. Tucci, L. Ottonetti, and F. Frizzi.2007. Competition trade-offs in the organisation of a Mediterranean ant assemblage. Ecological Entomology 32:319-326.
Savolainen, R., and K. Vepsalainen.1988. A competition hierarchy among boreal ants:impact on resource partitioning and community structure. Oikos 51:135-155.
Schappert, P. J., and J. S. Shore.1999. Effects of cyanogenesis polymorphism in Turnera ulmifolia on Euptoieta hegesia and potential Anolis predators. Journal of Chemical Ecology 25:1455-1479.
Schmid, R.1988. Reproductive versus extra-reproductive nectaries:historical perspectives and terminological recommendations. The Botanical Review 54:179-232.
Smiley, J.1986. Ant Constancy at Passiflora Extrafloral Nectaries:Effects on Caterpillar Survival. Ecology 67(2):516-521.
Stanton, M. L., T. M. Palmer, and T. P. Young.2002. Competition-colonization trade-offs in a guild of African acacia-ants. Ecological Monographs 72:347-363.
Stapel, J. O., A. M. Cortesero, C. M. De Moraes, J. H. Tumlinson, and W. J. Lewis.1997. Extrafloral nectar, honeydew, and sucrose effects on searching behavior and efficiency of Microplitis croceipes (Hymenoptera:Braconidae) in cotton. Environ. Entomol.26:617-623.
Storti, E. F.2002. Pollination biology and breeding system of Passiflora coccinea Aubl. in Manaus, Amazonas, Brazil. Acta Amazonica 32:421-429.
Suarez, A. V., C. Moraes, and A. Ippolito.1998. Defense of Acacia collinsii by an Obligate and Nonobligate Ant Species:the Significance of Encroaching Vegetation 1. Biotropica 30:480-482.
Tilman, D.1978. Cherries, Ants and Tent Caterpillars:Timing of Nectar Production in Relation in Relation to Susceptibility of Catepillars to Ant Predation. Ecology 59:686-692.
Van Mele, P., and N. T. T. Cuc.2001. Farmers'Perceptions and Practices in Use of Dolichoderus thoracicus (Smith)(Hymenoptera:Formicidae) for Biological Control of Pests of Sapodilla. Biological Control 20:23-29.
Vanderplank, J.1996. Passion Flowers. The MIT, Cambridge
Wackers, F. L., and C. Bonifay.2004. How to be sweet? Extrafloral nectar allocation by Gossypium hirsutum fits optimal defense theory predictions. Ecology 85:1512-1518.
Wagner, D.1997. The influence of ant nests on Acacia seed production, herbivory and soil nutrients. Journal of Ecology 85:83-93.
Wagner, D., and A. Kay.2002. Do extrafloral nectaries distract ants from visiting flowers? An experimental test of an overlooked hypothesis. Evol. Ecol. Res.4:293-305.
Way, M. J., and K. C. Khoo.1991. Colony dispersion and nesting habits of the ants, Dolichoderus thoracicus and Oecophylla smaragdina(Hymenoptera:Formicidae), in relation to their success as biological control agents on cocoa. Bulletin of Entomological Research 81:341-350.
Wheeler, H. C., and C. C. Bennington.2001. Effects of light availability on herbivory and defense chemistry in Passiflora incarnata. Florida Scientist 64:237-245.
Willmer, P. G., and G. N. Stone.1997. How aggressive ant-guards assist seed-set in Acacia flowers. Nature (London) 388:165-167.
Wirth, R., and I. R. Leal.2001. Does rainfall affect temporal variability of ant protection in Passiflora coccinea? Ecoscience 8:450-453.
Young, T. P., C. H. Stubblefield, and L. A. Isbell.1997. Ants on swollen-thorn acacias:species coexistence in a simple system. Oecologia 109:98-107.
方文培,张泽荣.1999《中国植物志》,第52(1)卷.北京：科学出版社,1983.pp 97-101
李绍文.1991.生态生物化学：(八)植食性昆虫对食物的选择.生态学杂志(Chinese journal of ecology)10:65-70.
张丽霞.2002.五种杀虫剂防治西番莲大斑芫菁的药效试验.植物保护(Plant protection) 28:51-53
杨萍,邓合黎,漆波,and刘琼.2006.红锯蛱蝶Cethosia biblis (Drury)的生物学研究.西南农业大学学报28：549-552.