榕小蜂食性分化与榕树―榕小蜂系统稳定性

doi:10.3724/SP.J.1003.2008.08216

摘要/Abstract

摘要：

在榕树与其传粉小蜂组成的互利共生系统中, 理解传粉小蜂与各种非传粉小蜂如何共存是解决这一系统稳定性维持机制问题的关键之一。生态位分化被普遍认为是传粉小蜂与各种非传粉小蜂共存的最主要动力。而作为生态位分化中最基础的食性分化在这一系统中如何具体实现尚不清楚。2006年12月至2007年6月, 我们以聚果榕(Ficus racemosa)为材料, 通过对果内6种榕小蜂进行独立放蜂及两两组合定量放蜂, 并对传粉小蜂分别进行不携带花粉和不能产卵的技术处理, 研究了寄生在聚果榕果内的5种非传粉小蜂的食性及相互关系, 分析了在不同季节下寄生蜂与寄主间的相关系数。研究结果表明: 在5种非传粉小蜂中, Platyneura testacea和P. mayri是造瘿者, 能独立刺激子房发育成瘿花, 并使果实发育成熟; 而Apocryptasp.、A. westwoodi和P. agraensis只能寄生于某些已发育的虫瘿, 为拟寄生者, 它们各自分别与P. testacea、P. mayri和传粉小蜂Ceratosolen fusciceps存在着一对一的寄生关系。拟寄生者与寄主间的相关性在不同季节下会显示出不同的结果, 这表明过去文献中用物种间的相关系数推理而确定的食性关系可能是不可靠的。对自然采集榕果内的小蜂群落分析表明, 传粉小蜂处于优势地位, 这说明在自然情况下非传粉小蜂的种群维持在一个较低水平, 对榕树—传粉小蜂系统稳定性影响较小, 故能与之长期共存。

关键词: 榕树, 非传粉小蜂, 互利共生, 食性, 生态位分化, 西双版纳

Abstract

In the fig-fig wasp reciprocal mutualism, understanding mechanisms of coexistence between pollinating wasps and non-pollinating wasps is an important and relevant topic. Niche partitioning has widely been considered as the most important mechanism in the coexistence of pollinating and non-pollinating wasps. In this study, we experimentally examined the diet of five species of non-pollinators and the relationship among fig wasps in Ficus racemosain Xishuangbanna, southern China, from Dec. 2006 to Jun. 2007. Platyneura testacea and P. mayri are gall-makers, but oviposit sequentially, utilizing different female flowers at different developmental stages; Apocryptasp., A. westwoodi and P. agraensis are parasitoids of P. testacea, P. mayri and Ceratosolen fusciceps respectively, presenting species-specific relationships with the hosted species. Species correlation coefficients differed greatly among seasons and conditions, suggesting that the use of correlation analysis to deduce or identify relationships between species in previous studies may be of limited value. Pollinators were the dominant species at our study sites. In these conditions, non-pollinating wasps exist at relatively low population density, and therefore may have a weak impact on the stability of the mutualism, potentially enabling the non-pollinating wasps coexist with the mutualism between figs and fig wasps.

Key words: mutualism, fig, non-pollinator fig wasps, diet, niche separation, Xishuangbanna

孙宝发, 王瑞武, 胡忠 (2008) 榕小蜂食性分化与榕树―榕小蜂系统稳定性. 生物多样性, 16, 525-532. DOI: 10.3724/SP.J.1003.2008.08216.

Baofa Sun, Ruiwu Wang, Zhong Hu (2008) Diet segregation of fig wasps and the stability of fig-fig wasp mutualism. Biodiversity Science, 16, 525-532. DOI: 10.3724/SP.J.1003.2008.08216.

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链接本文: https://www.biodiversity-science.net/CN/10.3724/SP.J.1003.2008.08216

https://www.biodiversity-science.net/CN/Y2008/V16/I6/525

图/表 6

参考文献 43

[1]	Berg CC (1989) Classification and distribution of Ficus. Experientia, 45,605-611.
[2]	Bronstein JL (1988) Predators of fig wasps. Biotropica, 20,215-219.
[3]	Bronstein JL (1991) The non-pollinating wasp fauna of Ficus pertusa: exploitation of a mutualism? Oikos, 61,175-186.
[4]	Chesson P (1991) A need for niches? Trends in Ecology & Evolution, 6,26-28. DOI URL PMID
[5]	Compton SG, Hawkins BA (1992) Determinants of species richness in southern African fig wasp assemblages. Oecologia, 91,68-74.
[6]	Compton SG, Robertson HG (1988) Complex interactions between mutualisms: ants tending homopterans protect fig seeds and pollinators. Ecology, 69,1302-1305.
[7]	Connell JH (1978) Diversity in tropical rain forest and coral reefs. Science, 199,1302-1309.
[8]	Cook JM, Power SA (1996) Effect of within-tree flowering asynchrony on the dynamics of seed and wasp production in an Australian fig species. Journal of Biogeography, 23,487-493.
[9]	Cushman GH, Compton SG, Zachariades C, Ware AB, Nefdt RJC, Rashbrook VK (1998) Geographic and taxonomic distribution of a positive interaction: ant-tended homopterans indirectly benefit figs across southern Africa. Oecologia, 116,373-380. URL PMID
[10]	Galil J, Dulberger R, Rosen D (1970) The effect of Sycophaga sycomori L. on the structure and development of the syconia in Ficus sycomorus L. New Phytologist, 69,103-111.
[11]	Galil J, Eisikowitch D (1968) On the pollination ecology of Ficus sycomorus in east Africa. Ecology, 49,259-269.
[12]	Gause GF (1934) The Struggle for Existence. Williams and Wilkins, London.
[13]	Hardin JW (1960) Studies in the Hippocastanaceae. V. Species of the Old World. Brittonia, 12,26-38.
[14]	Hassell MP, Comins HN, May RM (1991) Spatial structure and chaos in insect population dynamics. Nature, 353,255-258.
[15]	Hawkins BA, Compton SG (1992) African fig wasp communities: undersaturation and latitudinal gradients in species richness. Journal of Animal Ecology, 61,361-372.
[16]	Holling CS (1973) Resilience and stability of ecological systems. Annual Review of Ecology and Systematics, 4,1-23.
[17]	Jazen DH (1979) How to be a fig? Annual Review of Ecology and Systematics, 10,13-51.
[18]	Kerdelhué C, Rasplus JY (1996) The evolution of dioecy among Ficus (Moraceae): an alternative hypothesis involving nonpollinating fig wasp pressure on the fig-pollinator mutualism. Oikos, 77,163-166.
[19]	Kerdelhué C, Rossi JP, Rasplus JY (2000) Comparative community ecology studies on Old World figs and fig wasps. Ecology, 81,2832-2849.
[20]	Ma WL (马炜梁), Chen Y (陈勇), Li HQ (李宏庆) (1997) A summarize of the study on fig trees and its pollinators. Acta Ecologica Sinica (生态学报), 17,209-215. (in Chinese with English abstract)
[21]	May RM (1981) Theoretical Ecology: Principles and Applications. Blackwell Scientific Publishers, Oxford, UK.
[22]	Molles MC (2002) Ecology: Concepts and Applications. McGraw-Hill Companies, Boston.
[23]	Ramirez BW (1970) Host specificity of fig wasps (Agaonidae). Evolution, 24,680-691.
[24]	Schatz B, Hossaert-Mckey M (2003) Interactions of the ant Crematogaster scutellaris with the fig-fig wasp mutualism. Ecological Entomology, 28,359-368.
[25]	Schatz B, Profift M, Rakhi BV, Borges RM, Hossaert-Mckey M (2006) Complex interactions on fig trees: ants capturing parasitic wasps as possible indirect mutualists of the fig-fig wasp interaction. Oikos, 113,344-352.
[26]	Sun RY (孙儒泳) (1992) Fundamentals of Animal Ecology (动物生态学原理). Beijing Normal University Publishing House, Beijing. (in Chinese)
[27]	Tilman D (1982) Resource Competition and Community Structure. Princeton University Press, Princeton.
[28]	Wang RW, Shi L, Ai SM, Zheng Q (2008) Trade-off between the reciprocal mutualists: local resource availability oriented interaction in fig/fig wasp mutualism. Journal of Animal Ecology, 77,616-623.
[29]	Wang RW, Yang CY, Zhao GF, Yang JX (2005) Fragmentation effects diversity of wasp community and its impact on fig/fig wasp interaction in Ficus racemosa. Journal of Integrative Plant Biology, 47,144-152.
[30]	Weiblen GD (2002) How to be fig wasp? Annual Review of Entomology, 47,299-330. DOI URL PMID
[31]	Weiblen GD, Bush GL (2002) Speciation in fig pollinators and parasites. Molecular Ecology, 11,1573-1578. DOI URL PMID
[32]	Weiblen GD, Yu DW, West SA (2001) Pollination and parasitism in functionally dioecious figs. Proceedings of the Royal Society of London B, Biological Sciences, 268,651-659.
[33]	West SA, Herre EA (1994) The ecology of the New World fig parasitizing wasps Idarens and the implications for the evolution of the fig-pollinator mutualism. Proceedings of the Royal Society of London B, Biological Sciences, 258,67-72.
[34]	West SA, Herre EA, Windsor DM, Green PRS (1996) The ecology and evolution of the New World non-pollinating fig wasp communities. Journal of Biogeography, 23,447-458.
[35]	Whittaker RH (1965) Dominance and diversity in land plant communities. Science, 147,250-260. DOI URL PMID
[36]	Wiebes JT (1979) Co-evolution of figs and their insect pollinators. Annual Review of Ecology and Systematics, 10,1-12.
[37]	Wilson JB (1990) Mechanisms of species coexistence: twelve explanations for Hutchinson’s ‘Paradox of the Plankton’: evidence from New Zealand plant communities. New Zealand Journal of Ecology, 13,17-42.
[38]	Yang CY (杨成云), Wang RW (王瑞武), Zhao GF (赵桂仿), Yang DR (杨大荣) (2005) Diet of non-pollinating wasps and their impact on the stability of fig-pollinator wasp mutualism. Zoological Research (动物学研究), 26,379-385. (in Chinese with English abstract)
[39]	Yang DR (杨大荣), Peng YQ (彭艳琼), Zhang GM (张光明), Song QS (宋启示), Gu HY (谷海燕), Wang QY (王秋艳) (2003) Structure and biodiversity of insect community on syconia fruits of Ficus racemosa in the tropical rainforest of Xishuangbanna, China. Chinese Journal of Applied Ecology (应用生态学报), 14,1710-1714. (in Chinese with English abstract)
[40]	Yang DR (杨大荣), Wang RW (王瑞武), Song QS (宋启示), Zhang GM (张光明) (2000) Rule of seasonal changes of the Ceratosolen sp. in the tropical rainforest of Xishuangbanna, China. Forest Research (林业科学研究), 13,477-484. (in Chinese with English abstract)
[41]	Yang DR (杨大荣), Zhao TZ (赵庭周), Wang RW (王瑞武), Zhang GM (张光明), Song QS (宋启示) (2001) Study on pollination ecology of fig wasp (Ceratosolen sp.) in the tropical rainforest of Xishuangbanna, China. Zoological Research (动物学研究), 22,125-130. (in Chinese with English abstract)
[42]	Zhang GM (张光明), Yang DR (杨大荣), Xu L (徐磊), Peng YQ (彭艳琼), Lu Y (卢耀) (2003) The study on the relationships of the species fig wasps in the figs of Ficus racemosa in Xishuangbanna. Chinese Journal of Ecology (生态学杂志), 22(4),20-26. (in Chinese with English abstract)
[43]	Zhao TZ (赵庭周), Yang DR (杨大荣), Xu JH (许继宏) (2001) The role and comprehensive value of fig trees in tropical rainforest of Xishuangbanna. Forest Research (林业科学研究), 14,441-445. (in Chinese with English abstract)

种类 Species	生态型 Ecotype	产卵特征 Oviposition	产卵时隐头果发育阶段 Developing phase of syconium on oviposition
榕小蜂亚科 Agaonidae
Ceratosolen fusciceps	传粉小蜂 Pollinator	果内产卵 Inside	雌花期 Female phase
长鞘榕小蜂亚科 Sycophaginae
Platyneura testacea	非传粉小蜂 Non-pollinator	果外产卵 Outside	花前期 Prefemale phase
P. mayri	非传粉小蜂 Non-pollinator	果外产卵 Outside	花前期至间花期 Prefemale to interfloral phase
P. agraensis	非传粉小蜂 Non-pollinator	果外产卵 Outside	间花期 Interfloral phase
延腹榕小蜂亚科 Sycoryctinae
Apocryptasp.	非传粉小蜂 Non-pollinator	果外产卵 Outside	花前期 Prefemale phase
A. westwoodi	非传粉小蜂 Non-pollinator	果外产卵 Outside	间花期 Interfloral phase

种类 Species	生态型 Ecotype	产卵特征 Oviposition	产卵时隐头果发育阶段 Developing phase of syconium on oviposition
榕小蜂亚科 Agaonidae
Ceratosolen fusciceps	传粉小蜂 Pollinator	果内产卵 Inside	雌花期 Female phase
长鞘榕小蜂亚科 Sycophaginae
Platyneura testacea	非传粉小蜂 Non-pollinator	果外产卵 Outside	花前期 Prefemale phase
P. mayri	非传粉小蜂 Non-pollinator	果外产卵 Outside	花前期至间花期 Prefemale to interfloral phase
P. agraensis	非传粉小蜂 Non-pollinator	果外产卵 Outside	间花期 Interfloral phase
延腹榕小蜂亚科 Sycoryctinae
Apocryptasp.	非传粉小蜂 Non-pollinator	果外产卵 Outside	花前期 Prefemale phase
A. westwoodi	非传粉小蜂 Non-pollinator	果外产卵 Outside	间花期 Interfloral phase

处理 Treatments	样本量 N	瘿花数 Galls (mean ± SD)	种子数 Seeds (mean ± SD)
正常小蜂 Natural foundresses	20	1,541.25±276.51^a	2,095.20±565.53^a
不带花粉小蜂 Foundresses without pollen	20	1,562.95±220.70^a	0.35±1.02^b
不能产卵小蜂 Foundresses that cannot oviposit	20	0.40±1.23^b	1,996.25±202.42^a

处理 Treatments	样本量 N	瘿花数 Galls (mean ± SD)	种子数 Seeds (mean ± SD)
正常小蜂 Natural foundresses	20	1,541.25±276.51^a	2,095.20±565.53^a
不带花粉小蜂 Foundresses without pollen	20	1,562.95±220.70^a	0.35±1.02^b
不能产卵小蜂 Foundresses that cannot oviposit	20	0.40±1.23^b	1,996.25±202.42^a

	人工放蜂果 Control fruits (n= 30)	自然采集榕果 Natural mature fruits
	人工放蜂果 Control fruits (n= 30)	雾凉季 Fog-cool (n= 36)	干热季 Dry-hot (n= 51)	雨季 Rainy (n= 45)
P. agraensisvs.C. fusciceps	-0.265^ns	0.536^*	0.489^**	-0.291^ns
A. westwoodi vs. P. mayri	-0.591^**	0.344^*	-0.375^*	0.007^ns
Apocryptasp. vs.P. testacea	-0.197^ns	0.183^ns	-0.392^**	-0.104^ns