Host-specificity and coevolution among pollinating and nonpollinating New World fig wasps

Figs (Ficus spp., Moraceae) and their pollinating wasps (Hymenoptera, Agaonidae, Chalcidoidea) constitute a classic example of an obligate plant-pollinator mutualism, and have become an ideal system for addressing questions on coevolution, speciation, and the maintenance of mutualisms. In addition to pollinating wasps, figs host several types of nonpollinating, parasitic wasps from a diverse array of Chalcid subfamilies with varied natural histories and ecological strategies (e.g. competitors, gallers, and parasitoids). Although a few recent studies have addressed the question of codivergence between specific genera of pollinating and nonpollinating fig wasps, no study has addressed the history of divergence of a fig wasp community comprised of multiple genera of wasps associated with a large number of sympatric fig hosts. Here, we conduct phylogenetic analyses of mitochondrial DNA sequences (COI) using 411 individuals from 69 pollinating and nonpollinating fig wasp species to assess relationships within and between five genera of fig wasps (Pegoscapus, Idarnes, Heterandrium, Aepocerus, Physothorax) associated with 17 species of New World Urostigma figs from section Americana. We show that host-switching and multiple wasp species per host are ubiquitous across Neotropical nonpollinating wasp genera. In spite of these findings, cophylogenetic analyses (TREEMAP 1.0, TREEMAP 2.02beta, and parafit) reveal evidence of codivergence among fig wasps from different ecological guilds. Our findings further challenge the classical notion of strict-sense coevolution between figs and their associated wasps, and mirror conclusions from detailed molecular studies of other mutualisms that have revealed common patterns of diffuse coevolution and asymmetric specialization among the participants.     

Contrasting patterns of fig wasp communities along Mt. Wilhelm,Papua New Guinea

The fig (Moraceae) and pollinating fig wasp (Agaonidae) mutualism is best known as a model system for the study of coevolution in plant–pollinator interactions and its central role in shaping vertebrate communities in tropical forests. Figs also host myriad antagonistic parasitic fig wasps which impose costs on both partners threatening mutualism stability. Spatiotemporal variation in parasitic wasp abundance is a key factor in mitigating these effects. Because fig wasps are temperature sensitive and likely vary in their ability to traverse environmental gradients, we expect community assemblages and abundance of both pollinating and non-pollinating fig wasps to respond to changes along an elevational gradient. In the present study, we compare the fig wasp communities and abundance of three fig species growing along the slopes of the Mount Wilhelm altitudinal gradient in Papua New Guinea. We quantified wasps from over 100 male fig trees and calculated seed set for 55 female trees along each of the species’ distribution on the transect. Our results show that the abundance of both pollinating and non-pollinating fig wasps follow a mid-elevation peak, consistent with fig species richness found in the same transect. The patterns, however, are different according to the host's species distribution. Seed set remained relatively constant along the gradient for all species with some decrease along higher elevations, potentially affecting connectivity along the gradient. As suggested for insects in general, temperature and habitat diversity appear to play a fundamental role in the species richness and abundance of fig wasps.     

非传粉小蜂对榕-蜂共生系统的影响

榕-蜂共生系统是桑科榕属(Ficus)植物与传粉榕小蜂专一互惠形成的生态学关系。但是,也有一些非传粉的小蜂出现在这个系统中,对榕-蜂共生系统可能产生较大的影响。西双版纳的聚果榕(Ficus racemosa)树上主要有5种非传粉小蜂,分别在榕果发育的不同阶段从果外向果内产卵。在传粉榕小蜂进果之前的花前期,Platyneura testace、Apocrypta sp.和P. mayri这3种非传粉小蜂先后到果外产卵繁殖后代,对榕-蜂共生系统造成显著影响,尤其是影响传粉榕小蜂的繁殖。在传粉榕小蜂进果之后的间花期,P. mayri、A. westwoodi和P. agraensis这3种非传粉小蜂相继到果外产卵,它们虽然能减少种子形成和传粉榕小蜂繁殖的数量,但最终没有对榕-蜂共生系统造成显著的影响。造瘿类的P. mayri可在花前期和间花期产卵繁殖,在花前期产卵时它主要是影响传粉榕小蜂的繁殖,而在间花期产卵时它则更多地是影响种子的生产。     

榕树–榕小蜂协同进化中的非对称性相互作用及其集合种群效应

在协同进化研究中,传统观点认为物种之间的相互作用是对称性的,在进化过程中将形成一个稳定的均衡状态或进化稳定策略。然而,近来的观测和实验数据表明,物种间的协同进化可能存在非对称性相互作用,而且这种非对称性可能造成集合种群效应(metapopulation effect)或形成非均衡状态(例如混沌,chaos)。本文利用"榕树–榕小蜂"这一经典的协同进化模式系统来介绍协同进化过程中的非对称性相互作用,以及这种非对称性如何产生集合种群效应。在榕–蜂共生系统中,栖身于榕果中的榕小蜂除了传粉小蜂之外,还有一些"投机"的非传粉小蜂。非传粉小蜂比传粉小蜂具有更强的竞争力,而榕树则可惩罚不合作的非传粉蜂,同时奖励合作的传粉小蜂,从而形成了复杂的非对称性种间相互作用。在某个斑块生境中,非传粉小蜂通过竞争作用排斥传粉小蜂,然而随着非传粉小蜂在蜂群中的比例不断升高,榕树惩罚作用(如落果等)常常会导致整个非传粉蜂群的数量急剧下降,甚至局域性灭绝。随后,其他斑块的传粉小蜂则迁移过来填补空白。然而,随着传粉小蜂种群的数量增长,该斑块又会集聚更多的投机性非传粉小蜂,进而诱发榕树再次进行惩罚。这样的非对称性相互作用导致了非传粉小蜂、传粉小蜂以及榕树种群的集合效应,其种群大小呈现周期性的"此消彼长"式的循环。     

Divvying up an incubator: How parasitic and mutualistic fig wasps use space within their nursery microcosm

Differential occupancy of space can lead to species coexistence. The fig–fig wasp pollination system hosts species-specific pollinating and parasitic wasps that develop within galls in a nursery comprising a closed inflorescence, the syconium. This microcosm affords excellent opportunities for investigating spatial partitioning since it harbours a closed community in which all wasp species are dependent on securing safe sites inside the syconium for their developing offspring while differing in life history, egg deposition strategies and oviposition times relative to nursery development. We determined ontogenetic changes in oviposition sites available to the seven-member fig wasp community of Ficus racemosa comprising pollinators, gallers and parasitoids. We used species distribution models (SDMs) for the first time at a microcosm scale to predict patterns of spatial occurrence of nursery occupants. SDMs gave high true-positive and low false-positive site occupancy rates for most occupants indicating species specificity in oviposition sites. The nursery microcosm itself changed with syconium development and sequential egg-laying by different wasp species. The number of sites occupied by offspring of the different wasp species was negatively related to the risk of syconium abortion by the plant host following oviposition. Since unpollinated syconia are usually aborted, parasitic wasps ovipositing into nurseries at the same time as the pollinator targeted many sites, suggesting response to lower risk of syconium abortion owing to reduced risk of pollination failure compared to those species ovipositing before pollination. Wasp life history and oviposition time relative to nursery development contributed to the co-existence of nursery occupants.     

福州大叶榕隐头果内榕小蜂的分类

通过对福州两个样地大叶榕雄花期隐头果进行定时、定点采集,观察与鉴定,从546个花序果内共收集到榕小蜂33641头,隶属于小蜂总科中的7个科(亚科)7个属的11个种,其中传粉小蜂为榕小蜂科的Platyscapa coronata,其余10种非传粉小蜂分别隶属于隐针榕小蜂亚科Epichrysomallinae,金小蜂科Pteromalidae的锥尾榕小蜂亚科Otitesellinae和延腹榕小蜂亚科Sycoryctinae,广肩小蜂科Eurytomidae,刻腹小蜂科Ormyridae和姬小蜂科Eulophidae。传粉小蜂和非传粉小蜂的种类和数量呈现明显的季节性。冬-春季(12-翌年5月)榕果内小蜂的种类和数量较多,传粉榕小蜂是优势种;夏-秋季(6-11月间)小蜂种类和数量略少,Camarothorax bismasculinus和Sycophila sp.1是优势种。在非传粉小蜂中,Camarothorax、Sycophila、Otitesella、Sycoscapter等4个属的榕小蜂为常见种,而Ormyrus和Aprostocetus等2个属的榕小蜂为偶见种。根据小蜂的形态特征制定大叶榕隐头果中榕小蜂种类检索表,本实验为榕小蜂分类、榕-蜂协同进化研究,以及非传粉小蜂对榕-蜂共生体系的影响等研究提供科学依据。     

中国榕小蜂触角感受器形态特征及进化适应性分析

【目的】榕树与传粉榕小蜂体系是协同进化、专性传粉的经典系统,每种榕树上一般还生活有种类丰富、数目众多的非传粉榕小蜂。在选择压力下,榕小蜂为在榕果内生存产生了明显的适应性形态。触角感受器是昆虫通讯系统的单元,其形态是反映生态和进化适应性的最佳特征之一。本文旨在对中国部分传粉和非传粉榕小蜂的触角感受器的形态多样性和进化适应性进行系统研究。【方法】对来源于海南和云南15种榕树上24种榕小蜂54个型的触角感受器形态进行了扫描电镜观察,基于现有的分子系统发育树,对传粉榕小蜂触角感受器的形态特征进行了性状演化分析,并对榕小蜂的形态特征进行了进化适应性分析。【结果】榕小蜂触角感受器普遍存在雌雄二型现象。雌蜂触角感受器种类有毛状、锥状、板状、刺状、钟形、腔锥状和栓锥状等,数目丰富,并且进果产卵的传粉雌蜂和果外产卵的非传粉雌蜂之间、进果产卵的传粉雌蜂和进果产卵的非传粉雌蜂之间在形态上存在差异。传粉雄蜂和不具有雄性多型现象的非传粉雄蜂触角感受器极为退化,具有雄性多型的非传粉雄蜂触角感受器形态在种内不具有显著差异。性状演化分析表明进化路径相当复杂,可能存在多次独立进化过程。触角感受器的形态类型与其进化适应性相关。【结论】榕小蜂触角感受器类型多样,形态变化丰富,并为适应榕果内的生存而产生了进化适应性特点。雌蜂和雄蜂在榕果内受到了完全不同的选择压力,行使不同的生态功能,从而产生了不同的适应性形态。不同的适应性形态可能与雌蜂不同的产卵行为、雄蜂不同的交配策略具有一定联系。该文是首次对中国榕小蜂触角感受器形态进行系统研究的报道,有助于更好地理解榕小蜂的形态特征、进化路线、行为策略和生态关系。     

榕小蜂温度耐受性及其对种间共存关系的影响

全球已知有约800种榕属(Ficus)植物, 主要分布在热带, 部分种类延伸至亚热带地区。温度是限制榕‒蜂共生系统分布北界的主要因素, 也显著影响榕树及其榕小蜂的繁殖成功, 其中榕小蜂对温度的响应更加敏感。榕小蜂只有在一定的温度范围内才能保持正常的生理机能, 其对温度耐受能力直接影响榕果内小蜂种群数量和群落内种间关系。然而目前对榕小蜂温度耐受性的研究尚少, 榕小蜂的温度耐受能力如何影响榕果内小蜂的共存关系还未见报道。本文研究了分布于西双版纳的2种雌雄同株和1种雌雄异株榕树果内传粉榕小蜂和非传粉小蜂的温度耐受能力。结果表明: 3种传粉榕小蜂对高温的耐受性极差, 相对于雌雄同株的高榕(F. altissima)和聚果榕(F. racemosa)传粉榕小蜂, 雌雄异株的鸡嗉子榕(F. semicordata)传粉榕小蜂对低温有增强的耐受趋势。聚果榕小蜂群落结构显示: 在适宜其生长的西双版纳地区, 传粉榕小蜂的数量占绝对优势, 在温度较低的季节其数量显著减少; 而非传粉榕小蜂呈相反模式, 较强的温度耐受能力使其在低温的雾凉季维持了较高的种群数量。鸡嗉子榕果内非传粉小蜂Sycoscapter trifemmensis相对于Philotrypesis dunia有更强的温度耐受能力, 在种群数量和种间关系上有更多的竞争优势及数量。榕小蜂的温度耐受性差异在物种分布、种间关系的维持和共存上起了重要作用, 本研究结果为阐明榕小蜂种间共存的维持机制提供了科学依据。     

Interactions of the ant Crematogaster scutellaris with the fig/fig wasp mutualism

Abstract. 1. A classical example of specialised pollination mutualisms is the relationship between fig trees and their pollinating wasps, in which each partner depends completely on the other for its reproduction; however the fig/fig wasp association is also the target of a great diversity of other species, ranging from specialised parasites to opportunistic foragers, among them ants.
2. The ant community and the sources of ant attraction observed on the Mediterranean fig tree Ficus carica were characterised.
3. A guild of ants attracted by homopterans tended on the plant was distinguished from a second guild composed of two co-dominant ant species ( Crematogaster scutellaris and Pheidole pallidula ) that prey mostly on pollinating wasps, abundant during certain parts of the fig cycle.
4. Foraging workers of C. scutellaris search for prey on the fig inflorescence (syconium), capturing pollinating wasps mostly at the peak of wasp emergence and at a rate estimated to reach 600 prey per day for an entire tree.
5. Detailed study of the predatory sequences displayed under experimental conditions showed that ant workers captured 100% of the pollinating wasps offered, while they captured only 5.5% of the parasitoid wasp specific to the pollinator. The respective impacts of the interaction on ants and on the figs are discussed, as well as several behavioural traits of predation by the dominant ant on pollinators.     

Relative investment in egg load and poison sac in fig wasps: Implications for physiological mechanisms underlying seed and wasp production in figs

Fig pollinating wasps and most non-pollinator wasps apply secretions from their poison sacs into oviposited flowers that appear necessary to the formation of the galls that their developing offspring consume. Thus, both eggs and poison sac secretions appear to be essential for wasp reproduction, but the relative investment in each is unknown. We measured relative investment in poison sac and egg production in pollinating and non-pollinating wasps associated with seven species of monoecious Panamanian figs representing both active and passive pollination syndromes. We then collected similar data for four fig hosts in China, where some wasp species in the genus Eupristina have lost the ability to pollinate (“cheaters”). All wasps examined possessed large poison sacs, and we found a strong positive correlation between poison sac size and absolute egg production. In the Panamanian species, the relative poison sac to egg investment was highest in the externally ovipositing non-pollinator wasps, followed by active pollinators, then by passive pollinators. Further, pollinator wasps of fig species with demonstrated host sanctions against “cheating” wasps showed higher investment in the poison sac than wasps of species without sanctions. In the Chinese samples, relative investment in the poison sac was indistinguishable between pollinators and “cheaters” associated with the same fig species. We suggest that higher relative investment in poison sac across fig wasp species reflects higher relative difficulty in initiating formation of galls and subsequently obtaining resources from the fig. We discuss the implications for the stability of the fig–wasp mutualism, and for the ability of non-pollinators to exploit this mutualism.     

Measuring the discrepancy between fecundity and lifetime reproductive success in a pollinating fig wasp

Lifetime reproductive success in female insects is often egg‐ or time‐limited. For instance in pro‐ovigenic species, when oviposition sites are abundant, females may quickly become devoid of eggs. Conversely, in the absence of suitable oviposition sites, females may die before laying all of their eggs. In pollinating fig wasps (Hymenoptera: Agaonidae), each species has an obligate mutualism with its host fig tree species [Ficus spp. (Moraceae)]. These pro‐ovigenic wasps oviposit in individual ovaries within the inflorescences of monoecious Ficus (syconia, or ‘figs’), which contain many flowers. Each female flower can thus become a seed or be converted into a wasp gall. The mystery is that the wasps never oviposit in all fig ovaries, even when a fig contains enough wasp females with enough eggs to do so. The failure of all wasps to translate all of their eggs into offspring clearly contributes to mutualism persistence, but the underlying causal mechanisms are unclear. We found in an undescribed Brazilian Pegoscapus wasp population that the lifetime reproductive success of lone foundresses was relatively unaffected by constraints on oviposition. The number of offspring produced by lone foundresses experimentally introduced into receptive figs was generally lower than the numbers of eggs carried, despite the fact that the wasps were able to lay all or most of their eggs. Because we excluded any effects of intraspecific competitors and parasitic non‐pollinating wasps, our data suggest that some pollinators produce few offspring because some of their eggs or larvae are unviable or are victims of plant defences.     

聚果榕两种非传粉小蜂产卵与果实脱落的关系

在聚果榕与其传粉榕小蜂Ceratosolen fusciceps组成的互利共生系统中,与传粉榕小蜂共存的还有榕树的寄生性小蜂Platyneura testacea和Platyneura mayri.这些寄生性小蜂由于不能给榕树带来任何收益而只是利用榕树的种子或与传粉小蜂竞争植物的雌花资源,因而可能导致榕树与榕小蜂之间合作系统的崩溃.植物果实的脱落机制普遍被认为是维持系统稳定的关键因素之一.然而,定量实验和野外观测发现P. mayri产卵并不会引起果实脱落,只有P. testacea产卵会使果实大量脱落.通过对3株样树进行比较发现:当产卵时的P. testacea数量越多时,它产生的瘿花数就越多,榕果发生脱落的比例越高.P. testacea的过度产卵是导致榕果选择性脱落的主要原因.结果表明:脱落机制并不能完全阻止非传粉小蜂的寄生,榕树只能选择性地脱落掉先于传粉小蜂产卵的榕果.这也同时表明维持榕树与榕小蜂互利共存的机制不仅仅只有榕果的脱落机制,可能还存在其它未被发现的机制.     

Egg deposition patterns of fig pollinating wasps: implications for studies on the stability of the mutualism

1. Fig pollinating wasps (Agaonidae) enter Ficus inflorescences (figs), oviposit in some of the flowers, and pollinate in the process. Each larva completes its development within a single flower. In most cases, an inflorescence entered by a wasp will represent its only egg‐laying site. The mechanisms that prevent pollinating wasps from exploiting all the flowers inside a fig are not understood. In this study, hypotheses about flower use by pollinating fig wasps were tested by investigating egg deposition patterns in three species. 2. Either one or three wasps were introduced into figs. The figs were then harvested. Serial sections allowed assessment of the presence or absence of a wasp egg in a sample of flowers in each fig. The overall proportion of flowers with eggs and the spatial distribution of eggs were then compared in single wasp figs and three foundress figs. 3. In all species, the proportion of flowers with a wasp egg increased with foundress number but less than three‐fold. 4. In all species, at least in single foundress figs, flowers near the fig cavity were more likely to receive a wasp egg than were flowers near the fig wall. 5. In two species, when the number of foundresses was multiplied by three, there was an increase in the use of flowers near the fig wall, while in the third species, the increase was spread evenly among flowers. 6. Factors affecting wasp egg deposition patterns and the potential of investigating such patterns for studying the stability of the mutualism are discussed.     

聚果榕榕果小蜂种群分布格局及其生境和季节差异比较

采用多个指数对西双版纳地区聚果榕榕果小蜂的种群分布格局进行了检测,并重点运用基于方差/均值比率法的t值测度比较了其生境和季节差异.结果表明,6种聚果榕榕果小蜂的种群分布格局皆呈聚集分布;传粉小蜂聚集强度在生境序列上,原始雨林≈严重干扰地段中度干扰地段,在季节序列上,雨季>干热季>雾凉季;非传粉小蜂不同种类不同性别随着生境和季节变化皆表现出明显的聚集强度差异.对各种聚集强度变化的发生机理与其对聚果榕传粉者互惠共生体进化稳定性进行了讨论,认为保护榕树、保护榕树和传粉昆虫互惠共生体的重点应放在保护其原始森林生境方面.     

聚果榕榕果小蜂种群分布格局及其生境和季节差异比较

采用多个指数对西双版纳地区聚果榕榕果小蜂的种群分布格局进行了检测,并重点运用基于方差／均值比率法的t值测度比较了其生境和季节差异．结果表明,6种聚果榕榕果小蜂的种群分布格局皆呈聚集分布;传粉小蜂聚集强度在生境序列上,原始雨林≈严重干扰地段>>中度干扰地段。在季节序列上,雨季>干热季>雾凉季;非传粉小蜂不同种类不同性别随着生境和季节变化皆表现出明显的聚集强度差异。对各种聚集强度变化的发生机理与其对聚果榕-传粉者互惠共生体进化稳定性进行了讨论,认为保护榕树、保护榕树和传粉昆虫互惠共生体的重点应放在保护其原始森林生境方面。     

Geographic structuring into vicariant species-pairs in a wide-ranging,high-dispersal plant–insect mutualism: the case of <Emphasis Type="Italic">Ficus racemosa</Emphasis> and its pollinating wasps

Ficus and their mutualistic pollinating wasps provide a unique model to investigate joint diversification in a high dispersal system. We investigate genetic structuring in an extremely wide-ranging Ficus species, Ficus racemosa, and its pollinating wasp throughout their range, which extends from India to Australia. Our samples were structured into four large, vicariant populations of figs and wasps which may correspond to distinct (sub)species, located in India, China-Thailand, Borneo, and Australia. However, the genetically most divergent group was the Indian population for the figs and the China-Thailand population for the wasps, suggesting different evolutionary histories of populations. Molecular dating for the wasps shows that diversification of the pollinator clade is surprisingly old, beginning about 13.6 Ma. Data on both the host fig species and its pollinating wasps suggest that strong genetic flow within biogeographic groups over several hundreds of kilometers has limited genetic and morphological differentiation and, potentially, local adaptation. This is probably due to long-distance dispersal of pollinating wasps. The genetic clustering into large geographic units observed in F. racemosa and its pollinators is reminiscent of what can be observed in some other high-dispersal organisms characterized by morphology that varies little over huge distances. The implications of strong gene flow for diversification processes and adaptation to different ecological conditions in Ficus and their pollinating wasps are just beginning to emerge.     

Non‐pollinating Fig Wasps Decrease Pollinator and Seed Production in Ficus andicola (Moraceae)

Fig trees ( Ficus spp.) and Agaonine fig‐wasps participate in an obligate mutualism. Fig wasps can only develop within fig inflorescences (syconia) and they are the only organisms capable of pollinating fig flowers. Other non‐pollinating wasps that lay eggs by inserting their ovipositors from the outside can also develop in syconia. These parasitic wasps may be parasitoids of either pollinating or other non‐pollinating wasps, or form galls in fig flowers or other tissues. Depending on this interaction, parasitic wasps may have various effects on the production of pollinating wasps and seeds. Wasps in the genus Idarnes, which parasitize New World figs (subgenus Urostigma), have an effect on wasp production but not on seed production. Heterandrium spp., which have short ovipositors and lay on external flowers, are infrequent and no effect on seed production has been documented. In the Colombian Andes, Idarnes spp. and Heterandrium spp. are the most frequent parasites of the Ficus andicola — Pegoscapus sp. mutualism, affecting 62 and 43 percent of syconia, respectively. Controlling for other factors that influence wasp and seed production, such as number of foundresses, syconium size and tree, we found that Idarnes reduced pollinator production by almost half but did not reduce seed production, whereas Heterandrium reduced seed production by 40 percent, and marginally affected pollinator production. Our results provide the first clear documentation of Heterandrium spp. impact on fig seed production. Whether the relative abundance of this genus is a generalized phenomenon in montane forest remains to be determined.     

高榕雌花期传粉榕小蜂和欺骗性小蜂的繁殖特点

榕树及其专一性传粉榕小蜂组成了动植物界最为经典的协同进化关系,传粉榕小蜂演化出欺骗性是非常罕见的。在雌雄同株的高榕隐头果内,共存着一种传粉榕小蜂Eupristina altissima和一种欺骗性的小蜂Eupristina sp.,两种小蜂在雌花期进入隐头果内繁殖,但有不同的繁殖特点。对比研究了两种小蜂从成虫羽化到产卵和传粉这个阶段的雌蜂个体大小、孕卵量及繁殖差异,结果表明:羽化期两种雌蜂的平均个体小,经飞行小个体的雌蜂易死亡,大个体雌蜂到达接受树,但通过苞片通道,一些个体较大的传粉榕小蜂被夹死导致进入果腔的雌蜂相对小,而欺骗性小蜂易通过苞片以至进入果腔的雌蜂个体较大。两种未产卵雌蜂均表现为个体大者孕卵量较多,但两种雌蜂的平均孕卵量没有差异。即使有充足雌花资源产卵,两种雌蜂均未产完所有卵,产卵后两种雌蜂卵巢中的卵量均显著减少,遗留下的卵量两种小蜂间没有差异。传粉榕小蜂只有部分个体传完所携带花粉,并表现为传粉越成功的雌蜂,产卵越多。存在种内竞争时,两种小蜂的产卵量均减少,传粉榕小蜂的传粉效率也降低。在种间竞争背景下,欺骗性小蜂产卵更成功,传粉榕小蜂的产卵和传粉量均受到极大抑制。研究结果说明雌花期隐头果内传粉榕小蜂只适量利用雌花资源产卵繁殖后代,更有效地传粉繁殖榕树种子,这可能是维持榕-蜂互惠系统稳定共存的重要机制之一;欺骗者稳定存在需降低与传粉者的直接竞争,而欺骗者和传粉者分散在不同果内,甚至是不同的树上繁殖是理想的繁殖策略。     

福州大叶榕隐头果内的小蜂群落结构与多样性

对福州2个样地10株大叶榕果内小蜂群落组成和物种多样性进行研究。全年在两个样地267个隐头果内共收集到小蜂13458只。发现大叶榕隐头果内有7种小蜂,隶属小蜂总科Chalcidoidae中的榕小蜂科Agaonidae、金小蜂科Pteromalidae的隐针榕小蜂亚科Epichrysomallinae、锥尾榕小蜂亚科Otitesellinae和延腹榕小蜂亚科Sycoryctinae;广肩小蜂科Eurytomidae、刻腹小蜂科Ormyridae、姬小蜂科Eulophidae,其中榕小蜂科的Platyscapa coronata是大叶榕唯一的传粉者,传粉方式为主动传粉;非传粉榕小蜂的雄性多型现象普遍。传粉与非传粉小蜂的性比明显具有偏雌现象。传粉小蜂性比为0.19±0.07,非传粉小蜂中Camarothorax bismasculinus小蜂性比为0.36±0.10;Walkerella sp.小蜂性比为0.36±0.22,Sycoscapter sp.小蜂性比为0.31±0.22,Sycophila sp.小蜂性比为0.35±0.13。雄性多型现象可能是导致非传粉小蜂性比提高的原因之一。根据各种榕小蜂发生数量及连续性,可将大叶榕隐头果中的榕小蜂分为常见种和偶见种,Platyscapa coronata、Camarothorax bismasculinus、Walkerella sp.、Sycoscapter sp.和Sycophila sp.为常见种,Omyrus sp.和Aprostocetus sp.为偶见种。偶见种的存在对常见种的数量几乎没有影响,偶见种利用的是榕果内未饱和的一部分资源,也可能是榕果为偶见种的发生预留了空间和资源。首次发现姬小蜂科的昆虫寄生在榕果内,且有一定的种群数量,为姬小蜂科昆虫分类及其生物学、生态学特性研究提供了基础资料。在大叶榕小蜂群落结构中,传粉小蜂和非传粉小蜂的种类和数量呈现明显的季节性。冬-春季(12-翌年5月)榕果内小蜂的种类和数量较多,传粉榕小蜂是优势种,其重要值达到0.42,榕果种子结实率高;夏-秋季(6-11月间)小蜂种类和数量略少,Camarothorax bismasculinus和Sycophila sp.是优势种,其重要值分别为0.56和0.28,而传粉小蜂的重要值仅为0.025,此期大叶榕榕果中几乎找不到传粉小蜂,榕果结实率极低,对大叶榕的繁殖利益有较大的负面影响。雨水和高温等不良气候,可能是导致夏-秋季雄花期榕果内的传粉小蜂数量骤减的主要原因。两个实验样地的小蜂群落结构组成没有明显差异,年变化趋势相似。研究结果为城市绿化和热带雨林生物多样性保护提供科学依据。     

Phylogeography and virulence structure of the powdery mildew population on its 'new' host triticale

Background

The interaction between insects and plants takes myriad forms in the generation of spectacular diversity. In this association a species host range is fundamental and often measured using an estimate of phylogenetic concordance between species. Pollinating fig wasps display extreme host species specificity, but the intraspecific variation in empirical accounts of host affiliation has previously been underestimated. In this investigation, lineage delimitation and codiversification tests are used to generate and discuss hypotheses elucidating on pollinating fig wasp associations with Ficus.

Results

Statistical parsimony and AMOVA revealed deep divergences at the COI locus within several pollinating fig wasp species that persist on the same host Ficus species. Changes in branching patterns estimated using the generalized mixed Yule coalescent test indicated lineage duplication on the same Ficus species. Conversely, Elisabethiella and Alfonsiella fig wasp species are able to reproduce on multiple, but closely related host fig species. Tree reconciliation tests indicate significant codiversification as well as significant incongruence between fig wasp and Ficus phylogenies.

Conclusions

The findings demonstrate more relaxed pollinating fig wasp host specificity than previously appreciated. Evolutionarily conservative host associations have been tempered by horizontal transfer and lineage duplication among closely related Ficus species. Independent and asynchronistic diversification of pollinating fig wasps is best explained by a combination of both sympatric and allopatric models of speciation. Pollinator host preference constraints permit reproduction on closely related Ficus species, but uncertainty of the frequency and duration of these associations requires better resolution.