钝叶榕果实内繁殖的两种榕小蜂与寄主榕树间的协同进化

doi:10.3773/j.issn.1005-264x.2008.04.005

摘要/Abstract

摘要：

榕树(Ficus)及其传粉榕小蜂(Agaonidae)构成了高度专一的互惠共生体系。榕树的果实(以下简称榕果)内也寄生着一些非传粉小蜂。绝大多数非传粉小蜂在榕果外把产卵器刺入果壁产卵到果腔内, 只有极少数种类能够进入果腔内产卵。在西双版纳地区, 钝叶榕(Ficus curtipes)上的杨氏榕树金小蜂(Diaziella yangi)类似于传粉者钝叶榕小蜂(Eupristina sp.), 它也是进入榕果内产卵繁殖后代的, 这就为比较研究榕果内产卵小蜂与寄主榕树间的关系提供了材料。该文从形态学、行为学和生态学角度比较研究了这两种进入榕果内产卵的小蜂与寄主钝叶榕之间的作用关系, 研究结果显示: 1)杨氏榕树金小蜂与钝叶榕小蜂的雌蜂头部形状存在趋同进化; 2)两种小蜂的产卵器的平均长度都比雌花花柱长, 因而能把卵产在子房里; 3)钝叶榕小蜂从瘿花出来需要3~5 h, 交配需要17~19 min, 杨氏榕树金小蜂从瘿花出来只需18~20 min, 交配时间为20~30 s; 4)在自然群落中, 大约90%的雌花期榕果里都只进一只杨氏榕树金小蜂和一只钝叶榕小蜂, 杨氏榕树金小蜂能通过传粉来增加榕树种子数量, 但对钝叶榕小蜂种群的繁衍造成了极显著的负面影响; 5)两种小蜂于同一时期进入榕果内繁殖, 子代同期成熟羽化, 发育期与榕树雄花的发育期同步。研究表明: 进入榕果内繁殖的两种小蜂与寄主榕树之间存在着协同进化关系, 杨氏榕树金小蜂为榕树有效地传粉, 这可能是一个由寄生者向互惠方向进化的实例。

关键词: 钝叶榕, 钝叶榕小蜂, 杨氏榕树金小蜂, 协同进化

Abstract:

AimsFig trees (Ficus) and their pollinating wasps (Chalcidoidea, Agaonidae) form a highly obligated mutualism. In addition, Ficus species also host some non-pollinating fig wasps, most of which oviposit from the outside of figs, and only a few have evolved fig-entry behavior. Ficus curtipes hosts an internal ovipositing non-ollinating fig wasp, Diaziella yangi, in Xishuangbanna, and it is similar to the pollinating fig wasp (Eupristina sp.). Our objective was to investigate whether there was coevolution between D. yangi and host F. curtipes in aspects of morphology, behavior and ecology.

Methods Head length and width of the fig wasps, along with length of the ovipositors and styles, were measured using dissecting microscopy. In the field, the behavior of wasp’s entry in female floral phase was observed, and the mating and emerging behavior in male floral phase were observed in the laboratory. Moreover, we recorded a number of foundress in the fig female floral phase, and counted the number of wasps and seeds in the male floral phase.

Important findings The head shape (length:width) of female D. yangi was strongly correlated with that of the female pollinator Eupristina sp., while there was no corresponding correlation in male head shape. The style-length of figs was in the reach ofEupristinasp. and D.yangi. The emergence time for each Eupristina sp. from a gall was usually for 3-5 h, and the mating time lasted 17-19 min. The emergence time for each D. yangi was usually for 18-20 min, and the mating time lasted 20-30 s. In nature, almost 90% of figs at female floral phase averaged only one D. yangi and one Eupristina sp. In the natural community, D. yangi has a significant negative effect on the pollinator, but a positive effect on the number of viable seeds. Both internal ovipositing species entered fig in the same day of female floral phase and left the fig at the same time (figs’ male floral phase). Results suggested thatD.yangi can pollinate the figs, similar to the pollinator Eupristinasp. It has high degree of co-adaptation with the hostF. curtipes. This is possibly an example of evolving from parasitism to mutualism.

Key words: Ficus curtipes, Eupristina sp., Diaziella yangi, coevolution

张凤萍, 彭艳琼, 杨大荣. 钝叶榕果实内繁殖的两种榕小蜂与寄主榕树间的协同进化. 植物生态学报, 2008, 32(4): 768-775. DOI: 10.3773/j.issn.1005-264x.2008.04.005

ZHANG Feng-Ping, PENG Yan-Qiong, YANG Da-Rong. COEVOLUTION BETWEEN TWO INTERNAL OVIPOSITING FIG WASPS AND HOST FICUS CURTIPES. Chinese Journal of Plant Ecology, 2008, 32(4): 768-775. DOI: 10.3773/j.issn.1005-264x.2008.04.005

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链接本文: https://www.plant-ecology.com/CN/10.3773/j.issn.1005-264x.2008.04.005

https://www.plant-ecology.com/CN/Y2008/V32/I4/768

图/表 6

表1 钝叶榕小蜂和杨氏榕树金小蜂头部形态特征(平均值±标准偏差)

Table 1 Head feature of female and male of Eupristina sp. and Diaziella yangi(mean ± SD)

种类 Species	雌蜂 Female (n=37)			雄蜂 Male (n=35)
种类 Species	长 Length (mm)	宽 Width (mm)	长/宽 Length /width	长 Length (mm)	宽 Width (mm)	长/宽 Length /width
钝叶榕小蜂 Eupristina sp.	0.41±0.03	0.39±0.01	1.03±0.06	0.41±0.03	0.39±0.02	1.04±0.07
杨氏榕树金小蜂 Diaziella yangi	0.40±0.03	0.49±0.04	0.83±0.22	0.47±0.04	0.53±0.04	0.88±0.03

图1 钝叶榕小蜂和杨氏榕树金小蜂雌蜂头部形状之间的相关性 ♀: 雌蜂

Fig. 1 Relationship between the head shapes of female Diaziella yangi andEupristina sp. Female wasps

图2 钝叶榕小蜂和杨氏榕树金小蜂雄蜂头部形状之间的相关性 ♂: 雄蜂

Fig. 2 Relationship between the head shapes of maleDiaziella yangi andEupristina sp. Male wasps

图3 钝叶榕小蜂和杨氏榕树金小蜂产卵器长度之间的相关性

Fig. 3 Relationship between the ovipositor lengths of female Diaziella yangi andEupristina sp.

图4 钝叶榕榕果果腔内传粉小蜂和进入果腔产卵的非传粉小蜂的进蜂量

Fig. 4 Number of the foundresses of Eupristina sp. and Diaziella yangi trapped in the fig cavity

表2 进入果腔产卵的非传粉小蜂杨氏榕树金小蜂对钝叶榕小蜂种群数量和种子数量的影响(平均值±标准偏差)

Table 2 Effects of Diaziella yangi on pollinators Eupristina sp. and seeds (mean ± SD)

模式 Model	雌花 No. of female flowers	种子数量 No. of seeds	传粉小蜂数量 No. of pollinators	杨氏榕树金小蜂数量 No. of Diaziella yangi
传粉榕小蜂 Eupristina sp. (n=46)	281.26±42.39	5.54±8.53	108.46±22.01	0
传粉榕小蜂+杨氏榕树金小蜂 Eupristina sp.+ Diaziella yangi (n=23)	240.61±50.51	10.00±13.80	10.52±5.17	80.69±21.25

参考文献 25

[1]	Boucek Z (1988). Australasian Chalcidoidea (Hymenoptera): errors and omissions. Entomologist’s Monthly Magazine, 125,85-86.
[2]	Bronstein JL (1987). Maintenance of species-specificity in a neotropical fig-pollinator wasp mutualism. Oikos, 48,39-46.
[3]	Bronstein JL (1988). Mutualism, antagonism, and the fig- pollinator interaction. Ecology, 69,1298-1302.
[4]	Compton SG (1993). One way to be a fig. African Entomology, 1,151-158.
[5]	Compton SG, Rasplus JY, Ware AB (1994). African fig wasp parasitoid communities. In: Hawkins B, Sheehan W eds. Parasitoid Community Ecology. Oxford University Press, Oxford,343-368.
[6]	Galil J, Neeman G (1977). Pollen transfer and pollination in the common fig ( Ficus carica L. ). New Phytologist, 79,163-171.
[7]	Galil J, Zerroni M, Shalom DB (1973). Carbon dioxide and ethylene effects in the coordination between the pollinator Blastophaga quadraticeps and the syconium in Ficus religiosa . New Phytologist, 72,1113-1127.
[8]	Ganeshaiah KN, Katyuria P, Shaanker RU (1999). Does optimal packing of flowers in syconia shape style length variation in monoecious figs? Biotropica, 31,312-320.
[9]	Greeff JM (1999). Mating ecology of the non-pollinating fig wasps of Ficus ingens. Animal Behaviour, 57,215-222. URL PMID
[10]	Hossaert-Mckey M, Gibernau M, Frey JE (1994). Chemosensory attraction of fig wasps to substances produced by receptive figs. Entomologia Experimentalis et Applicata, 70,185-192.
[11]	Janzen DH (1979). How to be a fig? Review of Ecology and Systematics, 10,13-51.
[12]	Jousselin E, Kjellberg F (2001). The functional implications of active and passive pollination in dioecious figs. Ecology Letters, 4,151-158.
[13]	Jousselin E, Rasplus JY, Kjellberg F (2001). Shift to mutualism in parasitic lineages of the fig/fig wasp interaction. Oikos, 94,287-294.
[14]	Kathuria P (1999). Pollination Biology of Monoecious Fig Trees in India. PhD dissertation, University of Leeds, Leeds, UK.
[15]	Kathuria P, Ganeshaiah KN, Shaanker RU, Vasudeva R (1995). Is there dimorphism for style lengths in monoecious figs? Current Science, 68,1047-1049.
[16]	Machado CA, Herre EA, McCafferty S, Bermingham E (1996). Molecular phylogenies of fig pollinating and non-pollinating wasps and the implications for the origin and evolution of the fig-fig wasp evolution. Journal of Biogeography, 23,531-542.
[17]	Michaloud G, Michaloud-Pelletier S, Wiebes JT, Berg CC (1986). The co-occurrence of two pollinating species of fig wasp and one species of fig. Proceedings of the Koninklijke Nederlandse Akademie Van Wetenschappen Series C-Biological and Medical Sciences, 88,93-119.
[18]	Nefdt RJC, Compton SG (1996). Regulation of seed and pollinator production in the fig-fig wasp mutualism. Journal of Animal Ecology, 65,170-182.
[19]	Ramirez WB (1970). Host specificity of fig wasps (Agaonidae). Evolution, 24,680-691. DOI URL PMID
[20]	Ramirez WB (1974). Coevolution of Ficus and Agaonidae . Annals of the Missouri Botanical Garden, 61,770-780.
[21]	van Noort S, Compton SG (1996). Convergent evolution of agaonine and sycoecine (Agaonidae, Chalcidoidea) head shape in response to the constraints of host fig morphology. Journal of Biogeography, 23,415-424.
[22]	van Noort S, Peng YQ, Rasplus JY (2006). First record of the fig wasp genus Diaziella Grandi (Hymenoptera: Chalcidoidea: Pteromalidae: Sycoecinae) from the Asian mainland with description of two new species from China . Zootaxa, 1337,39-59.
[23]	Weiblen GD (2002). How to be a fig wasp? Annual Review of Entomology, 47,299-330.
[24]	West SA, Herre EA (1994). The ecology of the New World fig-parasitizing wasps Idarnes and implications for the evolution of the fig-pollinator mutualism . Proceedings of the Royal Society of London, Series B: Biological Sciences, 258,67-72.
[25]	Wiebes JT (1979). Coevolution of figs and their insect pollinators. Annual Review of Zoology and Systematics, 10,1-12.