The occurrence of fig wasps in the fruits of female gynodioecious fig trees

Fig trees are pollinated by wasp mutualists, whose larvae consume some of the plant's ovaries. Many fig species (350+) are gynodioecious, whereby pollinators generally develop in the figs of ‘male’ trees and seeds generally in the ‘females.’ Pollinators usually cannot reproduce in ‘female’ figs at all because their ovipositors cannot penetrate the long flower styles to gall the ovaries. Many non-pollinating fig wasp (NPFW) species also only reproduce in figs. These wasps can be either phytophagous gallers or parasites of other wasps. The lack of pollinators in female figs may thus constrain or benefit different NPFWs through host absence or relaxed competition. To determine the rates of wasp occurrence and abundance we surveyed 11 dioecious fig species on Hainan Island, China, and performed subsequent experiments with Ficus tinctoria subsp. gibbosa to identify the trophic relationships between NPFWs that enable development in female syconia. We found NPFWs naturally occurring in the females of Ficus auriculata, Ficus hainanensis and F. tinctoria subsp. gibbosa. Because pollinators occurred only in male syconia, when NPFWs also occurred in female syconia, overall there were more wasps in male than in female figs. Species occurrence concurred with experimental data, which showed that at least one phytophagous galler NPFW is essential to enable multiple wasp species to coexist within a female fig. Individuals of galler NPFW species present in both male and female figs of the same fig species were more abundant in females than in males, consistent with relaxed competition due to the absence of pollinator. However, these wasps replaced pollinators on a fewer than one-to-one basis, inferring that other unknown mechanisms prevent the widespread exploitation by wasps of female figs. Because some NPFW species may use the holes chewed by pollinator males to escape from their natal fig, we suggest that dispersal factors could be involved.     

鸡嗉子榕隐头果雌花期特征对传粉榕小蜂选择的影响

为了探讨榕树隐头果的发育期、性别、大小等外部特征对传粉榕小蜂选择的影响,采取人为控制雌花期的方法,对鸡嗉子榕(Ficus sermicordata)及其传粉榕小蜂(Ceratosolen gravelyi)的选择行为进行研究。结果表明,在隐头花序发育到雌花期后,如果阻止传粉小蜂进入,隐头果会继续生长。直径较小的雌果和雄果的进蜂量较多,且在雌雄果同时存在时,小蜂仍然会选择进入雌果,但进蜂量显著低于雄果。小蜂优先选择进入雌花期前期的隐头花序,雌雄果皆有此特点。对于相同发育期的隐头果,果径和进蜂量呈正相关关系,说明对于相同发育期的隐头果,小蜂更倾向于进入较大的隐头果。因此,真正控制小蜂行为的是隐头花序所处的发育期,以及不同发育期所产生的化学挥发物,而非隐头果直径大小。这为进一步研究榕-蜂系统的稳定机制提供依据。     

Finding hidden females in a crowd: Mate recognition in fig wasps

Multi-species mating aggregations are crowded environments within which mate recognition must occur. Mating aggregations of fig wasps can consist of thousands of individuals of many species that attain sexual maturity simultaneously and mate in the same microenvironment, i.e, in syntopy, within the close confines of an enclosed globular inflorescence called a syconium – a system that has many signalling constraints such as darkness and crowding. All wasps develop within individual galled flowers. Since mating mostly occurs when females are still confined within their galls, male wasps have the additional burden of detecting conspecific females that are “hidden” behind barriers consisting of gall walls. In Ficus racemosa, we investigated signals used by pollinating fig wasp males to differentiate conspecific females from females of other syntopic fig wasp species. Male Ceratosolen fusciceps could detect conspecific females using cues from galls containing females, empty galls, as well as cues from gall volatiles and gall surface hydrocarbons.In many figs, syconia are pollinated by single foundress wasps, leading to high levels of wasp inbreeding due to sibmating. In F. racemosa, as most syconia contain many foundresses, we expected male pollinators to prefer non-sib females to female siblings to reduce inbreeding. We used galls containing females from non-natal figs as a proxy for non-sibs and those from natal figs as a proxy for sibling females. We found that males preferred galls of female pollinators from natal figs. However, males were undecided when given a choice between galls containing non-pollinator females from natal syconia and pollinator females from non-natal syconia, suggesting olfactory imprinting by the natal syconial environment.     

Adaptation of the externally feeding bug <Emphasis Type="Italic">Elasmucha necopinata</Emphasis> (Hemiptera: Acanthosomatidae) to its fig host

Figs (Ficus) are keystone resources that maintain tropical biodiversity. Pollinators, non-pollinating fig wasps, and some insects that feed internally in syconia (the fruits of figs) synchronize their developmental stages with syconia. Other insects feed and develop externally on the syconia but little is known about adaptations in these insects. We investigated the life cycle of Elasmucha necopinata, a bug that develops externally on the syconia of Ficus hispida, a functionally dioecious fig. The bug oviposits about a week after fig receptivity, the nymphs feed externally on the syconia and eclosion occurs when the syconia mature. Thus, nymphs synchronize their developmental stages with the male syconia of F. hispida. Although the relationship does not appear to be mutualistic, we suggest that E. necopinata, feeding externally, has adapted to figs, in a similar way to Agaonid fig wasps that live internally. We believe that this is the first direct evidence of adaptation of an externally feeding insect to figs.     

Pollinators entering female dioecious figs: why commit suicide?

In the dioecious fig/pollinator mutualism, the female wasps that pollinate figs on female trees die without reproducing, whereas wasps that pollinate figs on male trees produce offspring. Selection should strongly favour wasps that avoid female figs and enter only male figs. Consequently, fig trees would not be pollinated and fig seed production would ultimately cease, leading to extinction of both wasp and fig. We experimentally presented pollinators in the wild (southern India) with a choice between male and female figs of a dioecious fig species, Ficus hispida L. Our results show that wasps do not systematically discriminate between sexes of F. hispida. We propose four hypotheses to explain why wasp choice has not evolved, and how a mutualism is thus maintained in which all wasps that pollinate female figs have zero fitness.     

The dominant exploiters of the fig/pollinator mutualism vary across continents,but their costs fall consistently on the male reproductive function of figs

1. Fig trees (Moraceae: Ficus) are keystone species, whose ecosystem function relies on an obligate mutualism with wasps (Chalcidoidea: Agaonidae) that enter fig syconia to pollinate. Each female flower produces one seed (fig female reproductive function), unless it also receives a wasp egg, in which case it supports a wasp. Fig male reproductive function requires both male flowers and pollinator offspring, which are the only vectors of fig pollen. 2. The mutualism is exploited by other wasps that lay eggs but provide no pollination service. Most of these non‐pollinating fig wasps (NPFWs) do not enter syconia, but lay eggs through the wall with long ovipositors. Some are gall‐makers, while others are parasitoids or lethal inquilines of other wasps. 3. Ficus is pan‐tropical and contains >750 fig species. However, NPFW communities vary across fig lineages and continents and their effects on the mutualism may also vary. This provides a series of natural experiments to investigate how the costs to a keystone mutualism vary geographically. 4. We made the first detailed study of the costs of NPFWs in a fig (Ficus obliqua G. Forst) from the endemic Australasian section Malvanthera. In contrast to the communities associated with section Americana in the New World, wasps from the subfamily Sycoryctinae (Chalcidoidea: Pteromalidae) dominated this community. 5. These sycoryctine wasps have a negative impact on pollinator offspring numbers, but not on seed production. Consequently, while the NPFW fauna varies greatly at high taxonomic levels across continents, we show that the consistent main effect of locally dominant exploiters of the mutualism is to reduce fig male reproductive function.     

THE STABILITY OF THE SYMBIOSIS BETWEEN DIOECIOUS FIGS AND THEIR POLLINATORS: A STUDY OF FICUS CARICA L. AND BLASTOPHAGA PSENES L.

Each Ficus species depends on a specific mutualistic wasp for pollination. The wasp breeds on the fig, each larva destroying a female flower. It is, however, not known why the wasps have not evolved the ability to use all female flowers. In “dioecious” figs, the wasp can only breed in the female flowers of the “male” trees, so that pollination of a female tree is always lethal. The wasps should therefore be selected to avoid female trees. Field data is presented showing that the fruiting phenology of the dioecious fig Ficus carica is such that this selection does not occur: syconia are not receptive at the same time on “male” and female trees. Most wasps are forced to emerge from the syconia of “male” trees at a time when they will not be able to reproduce, whether they avoid female trees or not. This aspect of the life cycle of the wasp, although noticed, has been obscured in most previous studies. It is shown that the fruiting phenology of Ficus carica, which stabilizes the symbiosis, is the result of short-term selective pressures on the male function of the trees. Such selective pressures suggest a possible pathway from monoecy to dioecy in Ficus under seasonal climates.     

Sexual specialization in two tropical dioecious figs

Ficus species (figs) and their species-specific pollinator wasps are involved in an intimate mutualism in which wasps lay eggs in some ovaries of the closed inflorescences (syconia), and mature, inseminated offspring carry pollen from mature syconia to fertilize receptive inflorescences. In monoecious species, each syconium produces seeds and wasps. In functionally dioecious fig species, making up approximately half the figs worldwide, male and female functions are separated; hermaphrodite (functionally male) trees produce wasps and pollen only, while female trees produce seeds only. This sexual separation allows selection to act independently on the reproductive biology of each sex. Examining sexual specialization in a tight mutualism allows us to determine aspects of the mutualism that are flexible and those that are canalized. In this study, we quantified the phenology of two species of dioecious figs, F. exasperata and F. hispida, for 2 years by following the fates of several thousand syconia over time. In studying each of these species in a dry and a wet site in south India, we tested specific predictions of how dioecious figs might optimize sexual function. On female trees of both species, more inflorescences matured during the wet (monsoon) season than in any other season; this fruiting period enabled seeds to be produced during the season most suitable for germination. In F. exasperata, functionally male trees released most wasps from mature syconia in the dry season, during peak production of receptive female syconia, and thus maximized successful pollination. In F. hispida, “male” trees produced more syconia in the dry and monsoon seasons than in the post-monsoon season. In both species, male and female trees abscised more unpollinated, young inflorescences than pollinated inflorescences, but abscission appeared to be more likely due to resource- rather than pollinator- limitation. The phenology of F. exasperata requires that male inflorescences wait in receptive phase for scarce pollinators to arrive. As expected, male inflorescences of this species had a longer receptive phase than female inflorescences. In F. hispida, where pollinators are rarely scarce, duration of receptive phase was the same for both sexes. Duration of developing phase was longer in female syconia of both species than in male syconia, most likely because they need a longer period of investment in a fleshy fruit. Variation in developing phase of female syconia in one species (F. exasperata) was also greater than that in male syconia, and enabled female trees to sample a variety of germination environments in time. The strong sexual differences in both fig species support the hypothesis that selection for sexual specialization has strongly influenced the reproductive biology of these species. Received: 28 May 1997 / Accepted: 2 February 1998     

薜荔榕小蜂对薜荔和爱玉子雌花期榕果挥发物的行为反应

薜荔和爱玉子均属雌雄异株桑科榕属植物,两者互为原变种与变种的关系,分别与薜荔传粉小蜂和爱玉子传粉小蜂(二者互为隐存种)建立了专性共生关系,榕树榕果挥发物在维系传粉小蜂与其寄主的共生关系上起着重要作用。利用Y型嗅觉仪测定薜荔榕小蜂(薜荔和爱玉子的传粉小蜂)对薜荔和爱玉子雌花期榕果挥发物的行为反应。结果表明:(1)雌花期果型的大小对薜荔榕小蜂行为反应无显著影响,薜荔大、小果型雌花期雌(雄)榕果挥发物对其传粉小蜂均具有强烈的吸引作用;(2)榕果挥发物浓度影响薜荔榕小蜂行为反应,薜荔、爱玉子雌花期雌(雄)榕果挥发物对其传粉小蜂的吸引作用均可能存在阈值反应,即榕果挥发物浓度未超过阈值时,雌花期榕果挥发物对传粉小蜂的吸引作用与挥发物浓度成正相关关系,而一旦超过阈值,榕果挥发物对传粉蜂的吸引作用显著下降,表明寄主榕果挥发物浓度影响传粉小蜂的寄主定位;(3)薜荔传粉小蜂对低浓度爱玉子雌花期雌(雄)榕果挥发物、爱玉子传粉小蜂对低浓度薜荔雌花期雌(雄)榕果挥发物均既无趋向也无驱避行为;薜荔传粉小蜂对高浓度的爱玉子雌花期雌(雄)榕果挥发物表现为显著的驱避行为,而爱玉子传粉小蜂对高浓度薜荔雌(雄)雌花期榕果挥发物表现为显著的趋向行为,因此,薜荔传粉小蜂与爱玉子传粉小蜂存在寄主专一性不对称现象,爱玉子传粉小蜂进入薜荔雌(雄)果内传粉或产卵的可能性较大,而福州地区的薜荔传粉小蜂可能难以进入爱玉子雌(雄)果内传粉或产卵。本研究结果将为榕-蜂共生体系的化学生态学理论研究以及爱玉子栽培提供科学依据。     

影响对叶榕及其传粉者繁殖的生态学因素

在西双版纳,分别统计了对叶榕(Ficus hispida)雌花期雌雄果的进蜂量和花后期雌雄果繁殖的多个特征值,以此来探讨自然条件下,影响对叶榕及其传粉榕小蜂(Ceratosolen solmsi marchali)繁殖的因素。结果表明:单果内有效进蜂数量是影响种子生产和传粉榕小蜂繁殖的首要因素,而雌花期进果的传粉榕小蜂并不是都能全部进入果腔传粉或产卵,大部分蜂还未进到果腔就被夹死在顶生苞片层的通道里,能进入雌果内传粉的榕小蜂为(2.72±2.04)只·果^-1,约占总进蜂量的52%;而在雄果里,能进入果腔的蜂量只有(2.08±1.65)只·果^-1,占35%左右。由于雌果内的雌花显著比雄果内的雌花多,结合单果进蜂量雌多雄少的格局,最终单果生产的种子数量 (1 891.63 ± 471.53)比传粉榕小蜂的数量 (367.20 ± 208.02) 多5倍有余。在雌果里,供给传粉的雌花数量与所生产的种子数量之间呈显著的正相关,而没有接受到花粉或不能正常受精的雌花数量与种子数量呈显著的负相关。雄果不仅生产花粉,也是传粉榕小蜂繁殖的场所,在相关于传粉榕小蜂自身繁殖力的因子中,传粉榕小蜂产卵制造的瘿花数量对其种群数量有最大的影响;影响次之的是发育过程中死亡的个体数量,它可降低30%左右的传粉榕小蜂数量;影响排在第三位的是寄主的雌花数量。此外,3类非传粉者的存在,单果内平均可减少30多只传粉小蜂。     

Floral constraint resulting from intersexual mimicry in a gynodioecious fig tree

Fig trees (Ficus: Moraceae) are pollinated by female fig wasps (Agaonidae) whose larvae develop inside galled flowers of unusual inflorescences (figs). Most fig trees also support communities of non‐pollinating fig wasps. Figs of different species display great size variation and contain tens to tens of thousands of flowers. Around one‐half the species of fig trees have the gynodioecious breeding system, where female trees have figs that produce seeds and male trees have figs that support development of pollinators. Mutual mimicry between receptive male and female figs ensures that pollinators enter female figs, even though the insects will die without reproducing, but the need to give no sex‐specific cues to the pollinators may constrain differences in size between receptive male and female figs. We compared relationships between inflorescence size and some measures of reproductive success in male and female figs of Ficus montana grown under controlled conditions in the presence of the pollinator Kradibia tentacularis and its main parasitoid Sycoscapter sp. indesc. Female figs that contained more flowers produced more seeds, but male figs did not increase the production of female pollinator K. tentacularis fig wasps in proportion of the flower number. Although more flowers were galled by the pollinators in male figs containing more female flowers, the high larval mortality caused by parasitism and nutritional limitation prevented the increase in the production of adult female offspring. Selection may favor the increase in flower numbers within figs in female plants of F. montana, but contrarily constrain this attribute in male plants.     

两种Eupristina属小蜂对宿主榕果挥发物和合成信息素的行为反应

高榕,雌雄同株,与其共生的形态高度相似的传粉小蜂Eupristina altissima与非传粉小蜂Eupristina sp.均为果内产卵。利用Y型嗅觉仪生物测定法研究E.altissima和Eupristina sp.对不同发育期高榕榕果挥发物以及合成信息素混合物的行为反应差异。结果表明:E.altissima和Eupristina sp.对高榕雌花期榕果挥发物具极显著趋向行为,对高榕雄花期榕果挥发物具极显著驱避行为,表明E.altissima与Eupristina sp.雌蜂能够敏锐辨别高榕雌花期和雄花期榕果的挥发物,快速远离雄花期榕果,并实现对宿主雌花期榕果的定位。但是,E.altissima与Eupristina sp.对其它发育时期榕果挥发物的行为反应具有差异性,前者对雌前期榕果和传粉或产卵后榕果释放的挥发物存在显著的趋避反应,而后者无明显的行为偏向性,表明E.altissima能够识别雌前期榕果和传粉或产卵后榕果,可避免误入其中而无法繁殖后代,Eupristina sp.则不具备这种识别能力。E.altissima和Eupristina sp.小蜂对合成信息素的行为反应也高度相似,表现为芳樟醇、苯乙烯,以及苯乙烯和金合欢醇的交互作用对两种小蜂选择行为影响显著,组合A_1B_2C_2(0.5%Linalool+1%Benzyl ethylene+1%Farnesol)的混合信息素对两种小蜂具有极显著的驱避作用,而组合A_2B_1C_1(1%Linalool+0.5%Benzyl ethylene+0.5%Farnesol),A_2B_1C_2(1%Linalool+0.5%Benzyl ethylene+1%Farnesol),A2B2C1(1%Linalool+1%Benzyl ethylene+0.5%Farnesol)的混合信息素对其具有极显著的吸引作用,但两种小蜂对组合A_1B_1C_1(0.5%Linalool+0.5%Benzyl ethylene+0.5%Farnesol)的行为反应存在差异。两种小蜂对榕果挥发物和合成信息素的反应异同,可能与其触角结构的差异密切相关。研究结果为探究榕-蜂共生体系的化学信号传导机制提供科学依据。     

Seasonal change in the structure of fig-wasp community and its implication for conservation

Figs (Moraceae) and their pollinating wasps (Agaonidae) constitute a famous reciprocal mutualism in which figs provide some female flowers for the development of fig wasp offspring while the fig wasps pollinate fig flowers. However, figs also host many non-pollinating wasps which are either parasitoids or resource competitors of pollinators, and bring no benefit for figs and are detrimental to fig’ fitness. Our data onFicus racemosa in Xishuangbanna showed that the numbers of non-pollinators and the mature syconia without pollinator wasps increase in rainy season, especially in the highly fragmented forest. This might be because of the longer developing time of the syconia and thereby longer oviposition time to non-pollinators in the dry season. The galled flower and the viable seed percentages in dry seasons are also larger than in rainy seasons in both primary forest and fragmented forest, and the development of non-pollinators is mainly at the expense of pollinator wasps. Our results showed that there exists a discriminative seasonal impact of non-pollinators and fragmentation effects on population size of fig’s pollinators. This implies that fig/fig wasp mutualism is more fragile in dry season, and that the critical population size and breeding units of figs in seasonal area might be larger than previously estimated without considering the seasonal change of pollinator population.     

西双版纳热带雨林对叶榕传粉生物学(英)

研究了西双版纳热带雨林地区雌雄异株植物对叶榕 (FicushispidaL .)的生物学、传粉物候学以及榕小蜂(CeratosolensolmsimarchaliMayr)的传粉和繁殖行为。研究结果表明 :雌雄异株的对叶榕与其他雌雄同株的榕属植物不同 ,它的种子形成与传粉者有着更为复杂的相互关系。对叶榕一年结隐花果 6～ 8次 ,结果高峰期 4～ 5次 ,其中雄性植株仅产生花粉和孕育榕小蜂 ,而雌性植株 (无雄蕊 )则需榕小蜂带花粉进入隐花果内 ,进行传粉授精 ,使之发育成种子。对叶榕的成熟花粉不能从花药开裂处自行散发出来 ,必须由榕小蜂采集才能散落。榕小蜂雌蜂羽化、交配后 ,找到雄花区 ,用足、触角、口器在推动中采集花粉。雌蜂飞出熟榕果找寻雌株或雄株榕树上的幼嫩隐花果 ,一般需 3～ 6 7min ;一部分雌蜂在雄株中寻找幼嫩的隐花果 ,进去产卵繁殖 ,另一部分雌蜂则寻找雌株雌花期嫩隐花果进去传粉。雌蜂在雌株榕树的隐花果内传粉时间为 15～ 2 3h ,在雄株榕树的隐花果内产卵时间为 6～ 9h。对叶榕小蜂在雌株上进入单个隐花果的数量多少关系到雌花结实率 ;观察表明 ,每个隐花果最佳进蜂数为 2头 ;榕小蜂传粉后榕树成熟种子形成率在 5 4 .1%～ 82 .5 %之间 ,平均为 73.8% ;而在雄株上雌蜂进蜂数量则关系到榕小蜂在隐花果内的产卵率 ,     

Between‐species facilitation by male fig wasps in shared figs

1. Facilitation is recorded from diverse plant–insect interactions, including pollination and herbivory. 2. The significance of facilitation resulting from the behavior of males of multiple fig wasp species inside figs was investigated. Female fig wasps emerge from natal figs via exit holes dug by males, especially male pollinators. When no males are present, the females struggle to escape and may die. 3. Ficus microcarpa L. is a widely‐established invasive fig tree from Southeast Asia. Its pollinator is absent in South Africa, so the tree cannot reproduce, but two Asian non‐pollinating fig wasps (NPFW) Walkerella microcarpae and Odontofroggatia galili occupy its figs. Abundance patterns of the two NPFW and the proportion of male‐free figs in South Africa, Spain (where the pollinator is introduced), and in China, where the native fig wasp community is diverse, were compared to determine the consequences of reduced species richness for insect survival. 4. Female fig wasps in male‐free figs were found to be trapped, and small clutch sizes contributed to the absence of males in both species. The presence of pollinators in Spain allowed most NPFW to develop in figs containing males. Far more male‐free figs were present in South Africa, elevating mortality rates among female NPFW. Facilitation of female release by males of other NPFW species nonetheless benefitted the rarer species. 5. Selection pressures in South Africa currently favour greater aggregation of NPFW offspring and/or less female biased sex ratios.     

西双版纳热带雨林对叶榕传粉生物学

研究了西双版纳热带雨林地区雌雄异株植物对叶榕(Ficus hispida L.)的生物学、传粉物候学以及榕小蜂(Ceratosolen solmsi marchali Mayr)的传粉和繁殖行为.研究结果表明:雌雄异株的对叶榕与其他雌雄同株的榕属植物不同,它的种子形成与传粉者有着更为复杂的相互关系.对叶榕一年结隐花果6～8次,结果高峰期4～5次,其中雄性植株仅产生花粉和孕育榕小蜂,而雌性植株(无雄蕊)则需榕小蜂带花粉进入隐花果内,进行传粉授精,使之发育成种子.对叶榕的成熟花粉不能从花药开裂处自行散发出来,必须由榕小蜂采集才能散落.榕小蜂雌蜂羽化、交配后,找到雄花区,用足、触角、口器在推动中采集花粉.雌蜂飞出熟榕果找寻雌株或雄株榕树上的幼嫩隐花果,一般需3～67 min;一部分雌蜂在雄株中寻找幼嫩的隐花果,进去产卵繁殖,另一部分雌蜂则寻找雌株雌花期嫩隐花果进去传粉.雌蜂在雌株榕树的隐花果内传粉时间为15～23 h,在雄株榕树的隐花果内产卵时间为6～9 h.对叶榕小蜂在雌株上进入单个隐花果的数量多少关系到雌花结实率;观察表明,每个隐花果最佳进蜂数为2头;榕小蜂传粉后榕树成熟种子形成率在54.1%～82.5%之间,平均为73.8%;而在雄株上雌蜂进蜂数量则关系到榕小蜂在隐花果内的产卵率,每个隐花果最佳进蜂数为3～4头,产卵率在72.3%～93.8%之间,平均为84.4%.     

Ability to gall: the ultimate basis of host specificity in fig wasps?

1. Fig trees (Ficus spp.) and their host‐specific pollinator fig wasps (Agaonidae) are partners in an obligate mutualism. Receptive phase figs release specific volatiles to attract their pollinators, and this is generally effective in preventing pollinator species from entering figs of the wrong hosts. 2. If entry is attempted into atypical host figs, then ostiole size and shape and style length may also prevent reproduction. In spite of these barriers, there is increasing evidence that fig wasps enter atypical hosts, and that this can result in hybrid seed and fig wasp offspring. 3. This study examines the basis of pollinator specificity in two dioecious fig species from different geographical areas. Kradibia tentacularis pollinates Ficus montana in Asia. Ficus asperifolia from East Africa is closely related but is pollinated by a different species of Kradibia. 4. In glasshouses, K. tentacularis was attracted to its normal host, F1s and backcrosses, but only rarely entered figs of F. asperifolia. Foundresses were able to lay eggs in hybrids, backcrosses, and F. asperifolia, although flower occupancy was lowest in F. asperifolia figs and intermediate in hybrids. 5. The fig wasp failed to reproduce in female F. montana, male F. asperifolia, and male F1s, and most but not all backcrosses to F. montana. This was a result of the failure to initiate gall production. 6. Host specificity in this fig wasp is strongly influenced by host volatiles, but the ability to gall may be the ultimate determinant of whether it can reproduce.     

Comparison of reproductive strategies in two externally ovipositing non-pollinating fig wasps

The reproductive strategies of Walkerella sp.1 associated with Ficus curtipes and Walkerella sp.2 associated with Ficus benjamina were investigated. Both species oviposited from outside the fig wall. Walkerella sp.1 was the first non-pollinating fig to oviposit on Ficus curtipes and began to do this ten days after figs syconia began to develop. The larvae of Walkerella sp.1 were only found in the most external ovary layer of the fig. Walkerella sp.2 starts ovipositing after several other non-pollinating fig wasps have already laid their eggs in F. benjamina. The progeny of Walkerella sp.2 are distributed in the external ovary layer, the middle ovary layer, and/or the inner ovary layer of the figs. However, more than a quarter of the offspring were found in the most external layer and only a few in the inner layer. Experimental studies proved that the two Walkerella species are gall formers. In both manipulated figs and in natural figs, the sex ratios of Walkerella sp.1 and Walkerella sp.2 were female-biased. In Walkerella sp.2, the overall sex ratio increased with the proportion of figs parasitized in a crop, but this was not the case for Walkerella sp.1. Females of both Walkerella species appear not to have information about the patches on which they oviposit because sex ratios of both species decreased as brood sizes within individual figs increased and foundresses of both species were able to lay clutches containing a single male egg and several female eggs.     

木瓜榕传粉生物学

 木瓜榕（Ficus auriculata Lour.）是雌雄异株植物,它只有通过大果榕小蜂（Ceratosolen emarginatus Mayr）传粉才能获得有性繁殖,大果榕小蜂也仅在木瓜榕雄果内产卵才得以繁衍后代。经两年来的实验和研究结果表明：木瓜榕繁殖能力在不同季节随进蜂量的多少而变化,雌树在4月和11月出现的两次结果高峰期与进蜂量最多的时期相遇;同时雄树与雌树在11月挂果高峰期重叠,导致该期进入雄果产卵的雌蜂量减少,进蜂量高峰推迟到部分雄果成熟、只有稀少接受果的3月下旬和4月上旬。木瓜榕在与大果榕小蜂互惠共生的进化历程中,逐渐完善了自身保证远交的繁殖机制,它们在不同程度上受到雌雄异熟、雌雄异株和短花柱雌花不亲和3种机制的共同作用。温湿度对大果榕小蜂访果行为有明显的影响,在温湿度变化缓和的早晚,有利于榕小蜂出行,雌蜂访果最为频繁。大果榕小蜂活动最佳的温湿度是：温度20～24 ℃之间、湿度85%～93%之间;温度高于29 ℃、湿度低于58% 或风雨天均不利于该蜂出行活动。榕小蜂日活动高峰的出现先后在不同季节略有差异,高温高湿的夏季榕小蜂活动出现早,7∶40达高峰;秋季榕小蜂活动推迟,高峰出现在11∶00;干凉的冬季最晚,13∶00才到达高峰。通过人工套袋、放蜂传粉试验,平均进8头的隐头果传粉效率最高,单果平均种子数量为11 247粒,结实率50.8%,未受精的败育花8 207朵;平均进蜂4头以下或10头以上的隐头果,由于花粉量不足或者雌蜂争夺资源的损伤使种子量下降,平均结实率分别达37.2% 和44.4%;放蜂试验结果与大量调查自然界中每果进蜂量和结实率的结果一致。     

薜荔和爱玉及其传粉昆虫繁殖特性

薜荔(Ficus pumila L．var．pumila)隶属桑科榕属,爱玉(F．pumila L．var．awkeotsanmg Corner)为其变种,它们的花是单性的,雌雄异株。雌花序中着生雌花,雄花序中有瘿花和雄花,每个花序中花的数量极多,达4000～6000朵。薜荔榕小蜂是唯一能进入薜荔和爱玉的隐头花序中产卵或传粉的共生昆虫,自然状态下雌花的结实率分别为82％、83．52％;瘿花的成虫瘿率分别为58．71％、51．32％,因此可形成大量的果实和虫瘿。物候观察表明薜荔和爱玉花期不遇,它们花序中的榕小蜂种群已经生殖隔离。人为的放蜂实验表明,生活于爱玉花序中的榕小蜂,已无法在薜荔花序中繁殖,生殖隔离进一步得到证实;实验同时表明爱玉的花粉亦不能使薜荔雌花结实,宿主两变种间生理上已不亲和。本文从共生双方协同进化的角度出发,探讨了榕树2变种间与传粉昆虫繁殖特性的差异,以及变种产生的主要原因。