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. (C) 2013 Elsevier Masson SAS. All rights reserved.

A coat of many scents: Cuticular hydrocarbons in multitrophic interactions of fig wasps with ants

The fig fig wasp system of Ficus racemosa constitutes an assemblage of galler and parasitoid wasps in which tritrophic interactions occur. Since predatory ants (Oecophylla smaragdina and Technomyrmex albipes) or mostly trophobiont-tending ants (Myrmicaria brunnea) were previously shown to differentially use volatile organic compounds (VOCs) from figs as proximal cues for predation on fig wasps, we examined the response of these ants to the cuticular hydrocarbons (CHCs) of the wasps. CHC signatures of gallers were distinguished from those of parasitoids by the methyl-branched alkanes 5-methylpentacosane and 13-methylnonacosane which characterised trophic group membership. CHC profiles of wasp predator and wasp prey were congruent suggesting that parasitoids acquire CHCs from their prey; the CHC composition of the parasitoid Apocrypta sp 2 clustered with that of its galler host Apocryptophagus fusca, while the CHC profile of the parasitoid Apocryptophagus agraensis clustered with its galler prey, the fig pollinator Ceratosolen fusciceps. In behavioural assays with ants, parasitoid CHC extracts evoked greater response in all ant species compared to galler extracts, suggesting that parasitoid CHC extracts contain more elicitors of ant behaviour than those of plant feeders. CHCs of some wasp species did not elicit significant responses even in predatory ants, suggesting chemical camouflage. Contrary to earlier studies which demonstrated that predatory ants learned to associate wasp prey with specific fig VOCs, prior exposure to fig wasp CHCs did not affect the reaction of any ant species to these CHCs. (C) 2015 Elsevier Masson SAS. All rights reserved.

榕小蜂的产卵模式及其对子代性比的影响

于2007年4月-2007年7月在中国科学院西双版纳热带植物园,通过干预控制榕小蜂产卵的方法研究了聚果榕的传粉榕小蜂Ceratosolen fusciceps和非传粉小蜂Platyneura mayri及垂叶榕的非传粉小蜂Acophila sp.1和Wakerella benjamini等的产卵顺序.对传粉榕小蜂,在完成放蜂后不同间隔时间向榕果内注入乙醚杀死小蜂:对非传粉榕小蜂,在放蜂后不同间隔时间将其从纱网袋内全部放出,从而控制了各种榕小蜂的产卵时间.到榕果成熟后,收集了榕果内的小蜂,并分析比较各种榕小蜂在不同产卵时间下的子代性比.结果表明:Ceratosolen fusciceps、P.mayri和Acophila sp.1在产卵的最初时间内倾向于产下更多的雄性后代,而随后的时间内则产下更多的雌性伴随少量的雄性后代,这样的产卵顺序导致子代性比随着母代产卵时间的延长而下降,榕小蜂后代雌性比例显著高于雄性.同时,子代榕小蜂数晕随母代产卵时间的延长而增加,这在一定程度上解释了单头繁殖雌蜂的子代性比随子代数量的增加而减少的现象.而Wakerella benjamini在产卵顺序上是随机的,在其开始产卵后的不同时间段内子代性比都接近于50%.这一结果表明榕小蜂的产卵顺序与母代产卵时间的长短对子代性比有极为重要的影响.

聚果榕两种非传粉小蜂产卵与果实脱落的关系

下载PDF阅读器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.

榕小蜂食性分化与榕树-榕小蜂系统稳定性

在榕树与其传粉小蜂组成的互利阿共生系统中,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小清楚.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最放蜂,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 Apocrypta sp、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.

Diffusive coevolution and mutualism maintenance mechanisms in a fig-fig wasp system

In reciprocal mutualism systems, the exploitation events by exploiters might disrupt the reciprocal mutualism, wherein one exploiter species might even exclude other coexisting exploiter species over an evolutionary time frame. What remains unclear is how such a community is maintained. Niche partitioning, or spatial heterogeneity among the mutualists and exploiters, is generally believed to enable stability within a mutualistic system. However, our examination of a reciprocal mutualism between a fig species (Ficus racemosa) and its pollinator wasp (Ceratosolen fusciceps) shows that spatial niche partitioning does not sufficiently prevent exploiters from overexploiting the common resource (i.e., the female flowers), because of the considerable niche overlap between the mutualists and exploiters. In response to an exploiter, our experiment shows that the fig can (1) abort syconia-containing flowers that have been galled by the exploiter, Apocryptophagus testacea, which oviposits before the pollinators do; and (2) retain syconia-containing flowers galled by Apocryptophagus mayri, which oviposit later than pollinators. However, as a result of (2), there is decreased development of adult non-pollinators or pollinator species in syconia that have not been sufficiently pollinated, but not aborted. Such discriminative abortion of figs or reduction in offspring development of exploiters while rewarding cooperative individuals with higher offspring development by the fig will increase the fitness of cooperative pollinating wasps, but decrease the fitness of exploiters. The fig fig wasp interactions are diffusively coevolved, a case in which fig wasps diversify their genotype, phenotype, or behavior as a result of competition between wasps, while figs diverge their strategies to facilitate the evolution of cooperative fig waps or lessen the detrimental behavior by associated fig wasps. In habitats or syconia that suffer overexploitation, discriminative abortion of figs or reduction in the offspring development of exploiters in syconia that are not or not sufficiently pollinated will decrease exploiter fitness and perhaps even drive the population of exploiters to local extinction, enabling the evolution and maintenance of cooperative pollinators through the movement between habitats or syconia (i.e., the metapopulations).

Biased oviposition and biased survival together help resolve a fig-wasp conflict

We report evidence that helps resolve two competing explanations for stability in the mutualism between Ficus racemosa fig trees and the Ceratosolen fusciceps wasps that pollinate them. The wasps lay eggs in the tree's ovules, with each wasp larva developing at the expense of a fig seed. Upon maturity, the female wasps collect pollen and disperse to a new tree, continuing the cycle. Fig fitness is increased by producing both seeds and female wasps, whereas short-term wasp fitness increases only with more wasps, thereby resulting in a conflict of interests. We show experimentally that wasps exploit the inner layers of ovules first (the biased oviposition explanation), which is consistent with optimal-foraging theory. As oviposition increases, seeds in the middle layer are replaced on a one-to-one basis by pollinator offspring, which is also consistent with biased oviposition. Finally, in the outer layer of ovules, seeds disappear but are only partially replaced by pollinator offspring, which suggests high wasp mortality (the biased survival or ‘unbeatable seeds’ explanation). Our results therefore suggest that both biased oviposition and biased survival ensure seed production, thereby stabilizing the mutualism. We further argue that biased oviposition can maintain biased survival by selecting against wasp traits to overcome fig defenses. Finally, we report evidence suggesting that F. racemosa balances seed and wasp production at the level of the tree. Because figs are probably selected to allocate equally to male and female function, a 1:1 seed:wasp ratio suggests that fig trees are in control of the mutualism.

Evaluación de la palatabilidad de Eugenia myrtifolia, justicia carnea y monstera deliciosa con potencial para la alimentación de marimonda (Ateles fusciceps robustus), en la fundación zoológico Santacruz, Cundinamarca

Tesis (Zootecnista). -- Universidad de La Salle. Facultad de Ciencias Agropecuarias. Programa de Zootecnia, 2013

西双版纳热带雨林聚果榕小蜂的传粉生态学

对聚果榕小蜂 (Ceratosolensp )传粉生态学进行了首次研究。结果表明 ,聚果榕小蜂的雄蜂比雌蜂早羽化数小时 ;雌蜂羽化不能自行打开瘿花和果肉出蜂口 ,两个出蜂口均需雄蜂开凿。而聚果榕的成熟花粉 ,不能自行地从开裂处散发出来 ,必须经榕小蜂的繁殖性雌蜂采集才能散到表面。羽化后的雌蜂在开裂的雄花中不停地用触角柄节、口器上颚和足推动和采集花粉。雌蜂飞出熟榕果寻找嫩隐头花果 ,一般在外飞翔 5～ 80min。雌蜂进入嫩聚果榕的隐头花果内后 ,立即把粘附在足、头、触角和身上的花粉不停地推动到长柱头雌花中 ,授粉行为长达 4～ 9h。然后 ,才把卵产在短柱头雌花中

Wolbachia infection and dramatic intraspecific mitochondrial dna divergence in a fig wasp

Mitochondria and Wolbachia are maternally inherited genomes that exhibit strong linkage disequilibrium in many organisms. We surveyed Wolbachia infections in 187 specimens of the fig wasp species, Ceratosolen solmsi, and found an infection prevalence of 89.3%. DNA sequencing of 20 individuals each from Wolbachia-infected and -uninfected subpopulations revealed extreme mtDNA divergence (up to 9.2% and 15.3% in CO1 and cytochrome b, respectively) between infected and uninfected wasps. Further, mtDNA diversity was significantly reduced within the infected group. Our sequencing of a large part of the mitochondrial genome from both Wolbachia-infected and -uninfected individuals revealed that high sequence divergence is common throughout the mitochondrial genome. These patterns suggest a partial selective sweep of mitochondria subsequent to the introduction of Wolbachia into C. solsmi, by hybrid introgression from a related species.

A Mechanical Analysis of Suspensory Locomotion in Primates and Other Mammals

For primates, and other arboreal mammals, adopting suspensory locomotion represents one of the strategies an animal can use to prevent toppling off a thin support during arboreal movement and foraging. While numerous studies have reported the incidence of suspensory locomotion in a broad phylogenetic sample of mammals, little research has explored what mechanical transitions must occur in order for an animal to successfully adopt suspensory locomotion. Additionally, many primate species are capable of adopting a highly specialized form of suspensory locomotion referred to as arm-swinging, but few scenarios have been posited to explain how arm-swinging initially evolved. This study takes a comparative experimental approach to explore the mechanics of below branch quadrupedal locomotion in primates and other mammals to determine whether above and below branch quadrupedal locomotion represent neuromuscular mirrors of each other, and whether the patterns below branch quadrupedal locomotion are similar across taxa. Also, this study explores whether the nature of the flexible coupling between the forelimb and hindlimb observed in primates is a uniquely primate feature, and investigates the possibility that this mechanism could be responsible for the evolution of arm-swinging.

To address these research goals, kinetic, kinematic, and spatiotemporal gait variables were collected from five species of primate (Cebus capucinus, Daubentonia madagascariensis, Lemur catta, Propithecus coquereli, and Varecia variegata) walking quadrupedally above and below branches. Data from these primate species were compared to data collected from three species of non-primate mammals (Choloepus didactylus, Pteropus vampyrus, and Desmodus rotundus) and to three species of arm-swinging primate (Hylobates moloch, Ateles fusciceps, and Pygathrix nemaeus) to determine how varying forms of suspensory locomotion relate to each other and across taxa.

From the data collected in this study it is evident the specialized gait characteristics present during above branch quadrupedal locomotion in primates are not observed when walking below branches. Instead, gait mechanics closely replicate the characteristic walking patterns of non-primate mammals, with the exception that primates demonstrate an altered limb loading pattern during below branch quadrupedal locomotion, in which the forelimb becomes the primary propulsive and weight-bearing limb; a pattern similar to what is observed during arm-swinging. It is likely that below branch quadrupedal locomotion represents a “mechanical release” from the challenges of moving on top of thin arboreal supports. Additionally, it is possible, that arm-swinging could have evolved from an anatomically-generalized arboreal primate that began to forage and locomote below branches. During these suspensory bouts, weight would have been shifted away from the hindlimbs towards forelimbs, and as the frequency of these boats increased the reliance of the forelimb as the sole form of weight support would have also increased. This form of functional decoupling may have released the hindlimbs from their weight-bearing role during suspensory locomotion, and eventually arm-swinging would have replaced below branch quadrupedal locomotion as the primary mode of suspensory locomotion observed in some primate species. This study provides the first experimental evidence supporting the hypothetical link between below branch quadrupedal locomotion and arm-swinging in primates.