Pollinating Fig Wasps Research Articles

Specialist fig-consuming lepidopterans can inflict costs to plant reproductive success that are mitigated by ant bodyguards

Insect herbivores can inflict substantial costs on plant reproductive success. Seed herbivory impacts directly by reducing the number of seeds and therefore the dispersal and reproductive potential of the plant. Fig trees, Ficus, provide keystone resources for tropical forests. The pollinating fig wasps develop inside figs, so consumption of unripe figs results in trees not only losing seeds but also their pollen dispersers. Selection to defend figs should therefore be strong. Seed herbivory is understudied in tropical forests and most data has been collected from fallen fruits. Here we use canopy sampling to identify fig-consuming larvae in central Panama and quantify both their consequences for the fig trees’ reproductive success and the defensive value that ants provide against these larvae. Field surveys of 46 crops from nine fig species revealed that larvae could destroy up to 80% of figs on a tree. From seven Ficus species we barcoded (using COI) 51 individual fig consuming larvae (mainly Lepidoptera) that grouped into seven molecular operational taxonomic units. Lepidopteran larvae formed two feeding strategies, either stationary within a fig or tunneling between figs. Within the context of our study, stationary larvae were specialists whereas tunneling larvae were either specialists or generalists found on different Ficus species. Trees with ants had significantly fewer figs consumed by larvae (9% ± 17% (mean ± SD) for trees with azteca ants and 16% ± 24% for trees with other ants) than did trees without ants (51% ± 27%). Our results corroborate earlier findings that hosting ants can be an effective defensive mechanism for trees against seed herbivores or other antagonistic insects. Our study contributes to a wider body of research around the networks of insects associated with figs that highlights the importance of a multitrophic approach for understanding mutualism stability and persistence in the face of antagonism.

Who holds the reins? Context-dependent resource allocation in the mutualism between fig trees and their fig wasp pollinators.

Mutualisms are consumer-resource interactions, in which goods and services are exchanged. Biological market theory states that exchanges should be regulated by both partners. However, most studies on mutualisms are one-sided, focusing on the control exercised by host organisms on their symbionts. In the brood-site pollination mutualism between fig trees and their symbiont wasp pollinators, galled flowers are development sites for pollinator larvae and are exchanged for pollination services. We determined if pollinator galls influenced resource allocation to fig inflorescences called syconia and considered feedbacks from the host tree. We experimentally produced syconia containing only seeds (S), only pollinator galls (G) or seeds and galls (SG) with varying number of introduced female pollinator wasps, i.e., foundress wasps. Biomass allocation to syconia was affected by foundress numbers and treatment groups; SG treatments received highest biomass allocation at low foundress numbers, and both G and SG treatments at high foundress numbers. Seeds are important determinants of allocation at low foundress numbers; galls are likely more influential at high foundress numbers. Most allocation in the G and SG treatment was to the syconium wall, likely as protection from parasitoids and temperature/humidity fluctuations. Dry mass of individual seeds and wasps (except at low foundress numbers) was unchanged between treatment groups, indicating seeds and wasps regulate resource flow into them, with lower flow into galls containing the smaller males compared to females commensurate with sexual dimorphism. We demonstrate the importance of considering the direct role of symbionts in accessing resources and controlling exchanges within mutualisms.

A new player in the Panamanian fig tree – fig wasp mutualism; a study on the effect of gall midges on Ficus citrifolia

The mutualism between the fig tree and the pollinating fig wasps is a keystone interaction in tropical forests. However, many antagonistic interactions also occur in the system, taking advantage of the fig trees and the pollinator. One such example is an antagonistic gall midge (Cecidomyiidae) that develops inside figs. Gall midges inside figs have been documented in a few Ficus species around the world, but to our knowledge they have not previously been observed in Panama. In this study the newly observed Panamanian fig gall midge is documented, together with its parasitoid wasp. The fig gall midge was only found in Ficus citrifolia figs. We investigated the effect of fig gall midge presence on the number of seeds and the number of pollinating wasps (Pegoscapus tonduzi) in a fig and aimed to identify the species based on morphology and barcoding of the COI region. We found that the fig gall midge had no, or a negligible effect, on the reproduction of the fig tree - fig wasp mutualism. The fig gall midge most likely belongs to the genus Ficiomyia, close to Ficiomyia perarticulata. The parasitoid belongs to the genus Physothorax, close to Physothorax russelli. This study suggests that the potentially newly arrived fig gall midge currently has no major effect on the fig tree - fig wasp mutualism. However, should infestation rates increase, it is likely that the fig gall midge would affect the mutualism negatively as it has in other parts of the world. More studies on the fig gall midge species distributions in this region would be valuable and would connect these newly observed species to a larger community, adding yet another species to this complex but classic example of a mutualism.

Multiple mating does not benefit females of a polyandrous pollinating fig wasp

In some species, females vary in the numbers of times they mate. While polyandry will always be beneficial to a male that mate with a previously mated female, the effect on female fitness is unclear. From females' perspective, variation in matedness can reflect adaptive differences in females’ requirements for mating or non-adaptive chance factors. Pollinating fig wasps have been considered to be mostly monandrous although polyandry has been confirmed in a number of species. Here we first show that the pollinating fig wasp, Platyscapa awekei, is polyandrous. Second, we show that intraspecific variation in female matedness may be explained best by chance encounters between males and receptive females. The mean number of offspring does not increase with polyandry. Nor is there evidence of sperm limitation. These observations rule out direct benefits to females. Despite evidence for multiply-mated females having mated with less compatible males, multiple mating is not combined with selective preference for more compatible males' sperm, ruling out indirect benefits. Therefore variation in female matedness seems to have no fitness benefit to females and from the females' perspective may be best explained by chance variation in encounter rates between males and receptive females.

Field sampling of fig pollinator wasps across host species and host developmental phase: Implications for host recognition and specificity.

Previous genetic studies of pollinator wasps associated with a community of strangler figs (Ficus subgenus Urostigma, section Americana) in Central Panama suggest that the wasp species exhibit a range in host specificity across their host figs. To better understand factors that might contribute to this observed range of specificity, we used sticky traps to capture fig-pollinating wasp individuals at 13 Ficus species, sampling at different phases of the reproductive cycle of the host figs (e.g., trees with receptive inflorescences, or vegetative trees, bearing only leaves). We also sampled at other tree species, using them as non-Ficus controls. DNA barcoding allowed us to identify the wasps to species and therefore assign their presence and abundance to host fig species and the developmental phase of that individual tree. We found: (1) wasps were only very rarely captured at non-Ficus trees; (2) nonetheless, pollinators were captured often at vegetative individuals of some host species; (3) overwhelmingly, wasp individuals were captured at receptive host fig trees representing the fig species from which they usually emerge. Our results indicate that wasp occurrence is not random either spatially or temporally within the forest and across these hosts, and that wasp specificity is generally high, both at receptive and vegetative host trees. Therefore, in addition to studies that show chemicals produced by receptive fig inflorescences attract pollinator wasps, we suggest that other cues (e.g., chemicals produced by the leaves) can also play a role in host recognition. We discuss our results in the context of recent findings on the role of host shifts in diversification processes in the Ficus genus.

Pollinator attraction in the Ficus deltoidea complex: Varietal specificity in a fig wasp that likes to stay close to home

Species boundaries are maintained by limitations on gene flow between taxa. In flowering plants pollinator specificity can strongly influence gene flow patterns and facilitate speciation. Fig trees are species rich and ecologically significant plants. It was once believed that each species had its own unique pollinator fig wasp, but numerous exceptions are now known. The dioecious Ficus deltoidea complex is distributed across southern South-East Asia with 13 recognised varieties of uncertain status, seven of which are found in Peninsular Malaysia. We investigated the host preferences of one of their Blastophaga pollinators, reared from the epiphytic F. deltoidea var. angustifolia. Our series of experimental manipulations using free-flying fig wasps and figs on host trees showed that male figs were preferred, that only their natal variety was attractive and that pollinators only dispersed from their natal trees if no suitable figs were present there. Fig wasps can only reproduce in male hosts, so preferred entry to male figs has clear advantages for them, as does their reluctance to disperse from natal trees. This behaviour is likely to reduce seed set on female plants, with potential consequences for host population dynamics. The extreme specificity of the Blastophaga sp. suggests gene flow between varieties of F. deltoidea is low or absent and that different varieties may be different biological species.

How to be a fig nematode

Fig (Moraceae: Ficus) species host vast communities of organisms that are bound together by complicated ecological networks that have influenced community structure and dynamics over evolutionary timescales. Much attention has been paid to the mutualism between figs and their specialized pollinating fig wasps, as well as with often antagonistic non-pollinating fig wasps. Equally ubiquitous to fig systems, but much less understood are the multitude of nematode groups that have independently evolved obligate associations with pollinating fig wasps and proliferate inside fig syconia. In this review we describe what is currently known (and unknown) about these numerous and increasingly studied nematode taxa and how they interact with the fig systems they inhabit. We identify the groups that are currently understood to associate with fig pollinators and outline their known species distributions and evolutionary history, where possible. Special attention is paid to the life history of these nematode groups, especially which features of nematode biology are generalizable across groups and what idiosyncratic peculiarities exist within individual genera. We outline key biological features including host choice, dispersal, disembarkation, diet, mating, and proliferation within figs. We address biological conundrums that have been raised following observational work such as, why do nematodes sometimes infect the wrong host? What adaptations were required for nematodes to successfully adapt and coexist with figs and their pollinators for millions of years? How do nematodes overcome the constraints of low mating group size? Finally, we outline key considerations, gaps in the knowledge, and limitations to expand this field forward towards promising areas of future research. Through better understanding of fig nematodes, we stand to not only know more about Ficus communities, but also more about the evolution and maintenance of interspecific interactions, development, adaptation, and co-evolution in general.

A NEW SPECIES OF <i>CERATOSOLEN</i> (AGAONIDAE) ASSOCIATED WITH <i>FICUS HISPIDA</i> L. AND PROPOSED CHANGES IN STATUS OF TWO OTHER POLLINATOR FIG WASPS

An increasing number of fig trees are known to support more than one species of pollinator fig wasp (Agaonidae). This brings into question the relative importance of co-evolution, radiation and host switching in the development of this mutualistic association. The functionally dioecious Ficus hispida has a wide distribution in Asia. It supports at least three morphologically distinct Ceratosolen pollinators, one of which we describe as C. abscondus Wong & Yu sp. n., based on collections in China and Thailand. The other two taxa were originally described as separate species, but subsequently downgraded to sub-species. The morphological differences between the three taxa are similar in extent to those recorded between other Ceratosolen species and we therefore re-instate them as the distinct species, C. solmsi (Mayr) and C. marchali (Mayr). An identification key is provided to distinguish between them and relationships between Ficus species and their Ceratosolen pollinators of this and related species are discussed.

The coexistence of two related fig wasp species sharing the same host fig species across a broad geographical area

Although more and more cases of breakdown of the 1: 1 partner specificity are being documented, figs and their pollinating fig wasps constitute perhaps the most tightly integrated pollination mutualism known. However, there are rare occasions where the pollinating fig wasp evolves cheating in this obligated system. The pollinator loses the ability to carry pollens but still lays eggs in female flowers. It has been reported that the figs of Ficus altissima, a functionally monoecious fig species, are occupied by two Eupristina species, the effective pollinator Eupristina altissima and the cheater Eupristina sp. in Xishuangbanna, SW China. However, little is known about whether the two Eupristina species entering figs can coexist widely in the nursery pollination system. Here, we used molecular methods to investigate the genetic diversity of Eupristina species in the widespread Asian fig species. Standard barcoding genes support two wasp species. Both fig wasp species were co-occurring in most distribution regions, raising the question of how two species can coexist within similar or identical resource niches. Our study offers a striking example of two closely related fig wasps that share a host can co-exist across a broad geographic area in a natural setting, but the pollinating wasps are more abundant.

Multi-omics analysis the differences of VOCs terpenoid synthesis pathway in maintaining obligate mutualism between Ficus hirta Vahl and its pollinators.

Volatile organic compounds (VOCs) emitted by the receptive syconia of Ficus species is a key trait to attract their obligate pollinating fig wasps. Ficus hirta Vahl is a dioecious shrub, which is pollinated by a highly specialized symbiotic pollinator in southern China. Terpenoids are the main components of VOCs in F. hirta and play ecological roles in pollinator attraction, allelopathy, and plant defense. However, it remains unclear that what molecular mechanism difference in terpenoid synthesis pathways between pre-receptive stage (A-phase) and receptive stage (B-phase) of F. hirta syconia. Transcriptome, proteome and Gas Chromatography-Mass Spectrometer (GC-MS) were applied here to analyze these difference. Compared to A-phase syconia, the genes (ACAT2, HMGR3, GGPS2, HDR, GPS2, TPS2, TPS4, TPS10-4, TPS14) related to the terpenoid synthesis pathway had higher expression level in receptive syconia (B-phase) according to transcriptome sequencing. Seven differentially expressed transcription factors were screened, namely bHLH7, MYB1R1, PRE6, AIL1, RF2b, ANT, VRN1. Specifically, bHLH7 was only specifically expressed in B-phase. 235 differentially expressed proteins (DEPs) were mainly located in the cytoplasm and chloroplasts. Kyoto Encyclopedia of Genes and Genomes (KEGG) analysis showed that the DEPs were mainly enriched in the metabolic process. A total of 9 terpenoid synthesis proteins were identified in the proteome. Among them, 4 proteins in methylerythritol phosphate (MEP) pathway were all down-regulated. Results suggested the synthesis of terpenoids precursors in B-phase bracts were mainly accomplished through the mevalonic acid (MVA) pathway in cytoplasm. Correlation analysis between the transcriptome and proteome, we detected a total of 1082 transcripts/proteins, three of which are related to stress. From the VOCs analysis, the average percent of monoterpenoids emitted by A-phase and B-phase syconia were 8.29% and 37.08%, while those of sesquiterpenes were 88.43% and 55.02% respectively. Monoterpenes (camphene, myrcene, camphor, menthol) were only detected in VOCs of B-phase syconia. To attract pollinators, B-phase syconia of F. hirta need more monoterpenoids and less sesquiterpenes. We speculate that transcription factor bHLH7 may regulate the terpenoid synthesis pathway between A- and B-phase syconia. Our research provided the first global analysis of mechanism differences of terpenoid synthesis pathways between A and B phases in F. hirta syconia.

Composition Diversity and Expression Specificity of the TPS Gene Family among 24 Ficus Species

Volatile organic compounds (VOCs) released by the receptive syconia of Ficus species play a vital role in attracting highly species-specific pollinating fig wasps. The components of VOCs vary considerably among Ficus species, but are generally dominated by a few common terpenoid compounds or specific proportions of several compounds. Terpene synthase (TPS) is the main source of specific and diverse terpenoids, but the evolution of the TPS gene family in Ficus and the potential functions of the TPS genes in species-specific pollination remain largely unelucidated. In this study, using transcriptomes of ostiole bracts of receptive male figs from 24 Ficus species collected from South China and Southeast Asia, we comprehensively scanned and investigated the composition and evolutionary characteristics of all TPS genes in all 24 species. We identified 248 TPS genes, including 33 orthologous genes and six singletons. Sequence and phylogenetic analysis showed that a majority of the 248 TPSs contained the DDXXD and DTE motifs, rather than the DXDD motif, and involved all subfamilies (TPS-a,b,c,e/f and g) known in other angiosperm genomes, suggesting a very diverse and complex composition of class I TPSs during the receptive phase. In addition, compared to TPS-a, which is generally the largest subfamily in some plants, the TPS-b subfamily contained the highest number of genes in Ficus species. Expression profile comparison showed that the distribution and expression levels of different TPSs among different Ficus species differed considerably, but a few TPS genes were common across most species. Positive selection analysis showed that the Ficus TPS genes were mainly under purifying selection, with only four genes having positive selection signals and two genes having positive selection sites, and two genes having relatively fast-evolving rates. The present study demonstrates the basic evolutionary characteristics of TPS genes in Ficus and reveals the roles of TPSs in shaping the diversity and specificity of the fig–fig wasp symbiotic relationship.

Overlaps in olfactive signalling coupled with geographic variation may result in localised pollinator sharing between closely related Ficus species

BackgroundIn brood site pollination mutualisms, pollinators are attracted by odours emitted at anthesis. In Ficus, odours of receptive figs differ among species and the specific pollinators generally only enter figs of their host species ensuring a pre-zygotic barrier to plant interspecific hybridisation. However, field observations recorded that, in Guangdong province in China, Valisia javana hilli, the local pollinator of F. hirta, entered and reproduced successfully in the figs of the closely related F. triloba on a regular basis. We propose that closely related Ficus species produce similar receptive fig odours. Under particular contexts of odours locally present, the receptive fig odours of non-host figs of a Ficus species may become attractive to pollinators of closely related Ficus species. We used the headspace technique to collect in situ receptive fig odours of F. triloba in a series of locations in China. Under controlled conditions, we tested the attraction of fig pollinating wasps from F. hirta and F. triloba to host figs and non-host figs in Y tube experiments.ResultsReceptive fig odours of F. triloba though different from those of F. hirta, were mainly composed of a same set of volatile organic compounds. When given the choice between receptive fig odours and air, the pollinating wasps were only attracted by their host’s odours. However, when given a choice between host and non-host figs the pollinators of F. hirta were equally attracted by the two odours while the pollinators of F. triloba tended to be more attracted by their host’s fig odours.ConclusionsReceptive fig odours vary geographically within species and the differentiation of receptive fig odours between closely related Ficus species is often incomplete. This allows localised or occasional pollinator sharing following different modalities. Cross stimulation when wasps are exposed simultaneously to odours of host and non-host species may be important. While occasional pollinator sharing may play a marginal role when wasp populations are robust, it may ensure the provisioning of new pollinators from the closest relative of a Ficus species if its pollinators go extinct.

Maintenance of specificity in sympatric host-specific fig/wasp pollination mutualisms.

BackgroundFig/wasp pollination mutualisms are extreme examples of species-specific plant-insect symbioses, but incomplete specificity occurs, with potentially important evolutionary consequences. Why pollinators enter alternative hosts, and the fates of pollinators and the figs they enter, are unknown.MethodsWe studied the pollinating fig wasp, Ceratosolen emarginatus, which concurrently interacts with its typical host Ficus auriculata and the locally sympatric alternative host F. hainanensis, recording frequencies of the wasp in figs of the alternative hosts. We measured ovipositor lengths of pollinators and style lengths in female and male figs in the two host species. Volatile organic compounds (VOCs) emitted by receptive figs of each species were identified using GC-MS. We tested the attraction of wasps to floral scents in choice experiments, and detected electrophysiologically active compounds by GC-EAD. We introduced C. emarginatus foundresses into figs of both species to reveal the consequences of entry into the alternative host.ResultsC. emarginatus entered a low proportion of figs of the alternative host, and produced offspring in a small proportion of them. Despite differences in the VOC profiles of the two fig species, they included shared semiochemicals. Although C. emarginatus females prefer receptive figs of F. auriculata, they are also attracted to those of F. hainanensis. C. emarginatus that entered male figs of F. hainanensis produced offspring, as their ovipositors were long enough to reach the bottom of the style; however, broods were larger and offspring smaller than in the typical host. Female figs of F. hainanensis failed to produce seeds when visited by C. emarginatus. These findings advance our current understanding of how these species-specific mutualisms usually remain stable and the conditions that allow their diversification.

Genome-Wide Identification and Analysis of Lipases in Fig Wasps (Chalcidoidea, Hymenoptera).

Simple SummaryLipases are a large family of enzymes involved in lipid metabolism. Lipids play diverse roles in insect growth and responses to environmental stimuli. Fig wasps are a polyphyletic assemblage of Chalcidoidea that develop in the inflorescences of fig trees. Based on whether they can pollinate, they are separated into pollinator fig wasp (PFW) and non-pollinating fig wasp (NPFW). In this study, we conducted a genome-wide screening of lipases in the 12 fig wasp genomes using bioinformatics tools, including seven PFWs and five NPFWs. In total, 481 lipase genes were identified with the neutral and acid lipases as the most numerous families. NPFWs had significantly more lipases than PFWs. Tandem duplication accounted for the expansion of the gene family. Phylogenetic analysis indicated that the lipase genes were conserved. This study provided evidence of insect metabolism to understand the obligate mutualism between figs and fig wasps. Our results will facilitate the understanding of the molecular mechanism of how lipase proteins contribute to the distinctions of life histories between PFWs and NPFWs.Lipases are the main enzymes involved in lipid metabolism. However, the characteristics of lipases in insects were scarcely investigated. Here, we screened the recently sequenced genomes of 12 fig wasp species consisting of seven pollinator fig wasps (PFWs) and five non-pollinating fig wasps (NPFWs) for the six major lipase gene families. In total, 481 lipase genes were identified, and the two most numerous families were the neutral and acid lipases. Tandem duplication accounted for the expansion of the gene family. NPFWs had significantly more lipases than PFWs. A significant gene family contraction occurred in the clade of PFWs. The difference of lipases between NPFWs and PFWs might contribute to their distinction in life histories and feeding regimes. Phylogenetic analysis showed that the lipase genes of each fig wasp species was almost equally distributed in each clade, indicating that the lipase genes were conserved. The gene structures were similar within each clade, while they were different among clades. Most of the neutral and acid lipases were signal peptides and located extracellularly. The pathways of lipases involved were predicted. This genome-wide study provides a systematic analysis of lipase gene families in 12 hymenopteran insects and further insights towards understanding the potential functions of lipases.

Predicting distributions of Wolbachia strains through host ecological contact-Who's manipulating whom?

Reproductive isolation in response to divergent selection is often mediated via third‐party interactions. Under these conditions, speciation is inextricably linked to ecological context. We present a novel framework for understanding arthropod speciation as mediated by Wolbachia, a microbial endosymbiont capable of causing host cytoplasmic incompatibility (CI). We predict that sympatric host sister‐species harbor paraphyletic Wolbachia strains that provide CI, while well‐defined congeners in ecological contact and recently diverged noninteracting congeners are uninfected due to Wolbachia redundancy. We argue that Wolbachia provides an adaptive advantage when coupled with reduced hybrid fitness, facilitating assortative mating between co‐occurring divergent phenotypes—the contact contingency hypothesis. To test this, we applied a predictive algorithm to empirical pollinating fig wasp data, achieving up to 91.60% accuracy. We further postulate that observed temporal decay of Wolbachia incidence results from adaptive host purging—adaptive decay hypothesis—but implementation failed to predict systematic patterns. We then account for post‐zygotic offspring mortality during CI mating, modeling fitness clines across developmental resources—the fecundity trade‐off hypothesis. This model regularly favored CI despite fecundity losses. We demonstrate that a rules‐based algorithm accurately predicts Wolbachia infection status. This has implications among other systems where closely related sympatric species encounter adaptive disadvantage through hybridization.

Association of Fig Pollinating Wasps and Fig Nematodes inside Male and Female Figs of a Dioecious Fig Tree in Sumatra, Indonesia

Simple SummaryIt has been known that fig pollinating wasps can transfer nematodes when they enter receptive figs to deposit their eggs (male figs in male trees) and pollinate the flowers (female figs in female trees) of a dioecious fig tree. However, the fate of nematodes transferred into female figs was unknown, since the pollinating wasps that enter female figs cannot reproduce. This study examined whether the nematodes transferred by pollinating wasps into female figs of Ficus hispida could develop and reproduce in the same way as in male figs. Three species of nematodes were found to develop within the male and female figs of F. hispida, with populations peaking at about the same time for both sexes of figs. Our findings showed that the female figs could support the growth and reproduction of the three nematodes; however, there was no pollinator offspring to transfer the nematodes out of the figs. The results provide a better understanding of the development of nematodes in male and female figs of a dioecious fig tree, which may also influence the biodiversity of the tropical ecosystem.Nematodes can grow within the inflorescences of many fig trees (Ficus spp., Moraceae); however, the feeding behaviour of most nematodes is not known. Fig pollinating wasps (Hymenoptera: Agaonidae) transfer nematodes into young figs upon the wasps’ entry into the figs to deposit their eggs. Most Asian fig trees, however, are functionally dioecious, and the pollinating wasps that enter female figs are unable to reproduce. They fail to produce the offspring required to carry the new generations of nematodes. We examined whether female figs of F. hispida can nonetheless support the development of phoretic nematode populations. Nematodes were extracted from male and female figs sampled in Sumatra, Indonesia, to compare the growth of their populations within the figs. We found three species of nematodes that grew within figs of male and female trees of F. hispida: Ficophagus cf. centerae (Aphelenchoididae), Martininema baculum (Aphelenchoididae) and Caenorhabditis sp (Rhabditidae). The latter species (Caenorhabditis sp.) has never been reported to be associated with F. hispida before. Nematode populations peaked at around 120–140 individuals in both sexes of figs, at the time when a succeeding generation of adult fig wasps appeared within male figs. The female figs could support the growth and reproduction of the three nematodes species; however, the absence of vectors meant that female figs remained as traps from which there could be no escape.

Asymmetric sharing of pollinator fig wasps between two sympatric dioecious fig trees: a reflection of supply and demand or differences in the size of their figs?

BackgroundHost specificity among pollinator fig wasps (Agaonidae) depends on host plant specific volatile cues, but fig wasps must also pass through a narrow physical barrier (the ostiole) if they are to pollinate and oviposit. Across South East Asia the dioecious shrub Ficus hirta is associated with at least ten pollinator species allied to Valisia javana. Ficus triloba has a single recorded pollinator, Valisia esquirolianae. Receptive figs of F. hirta are usually much smaller than those of F. triloba, but at a mainland site where F. hirta has atypically large figs we identified both V. esquirolianae and V. javana from both Ficus species using COI and ITS2 sequencing. To investigate whether this host overlap was exceptional we reared fig wasps from the two trees elsewhere and recorded features that may facilitate host transfer between them, including attractant volatiles, reproductive phenology and the sizes of their figs and fig wasps.ResultsThe two Ficus species were found to support both Valisia species at several of the sites, suggesting that the differences we detected in volatile profiles, ostiole sizes and pollinator head sizes are not strict barriers to host sharing. Valisia javana colonised F. triloba more frequently than V. esquirolianae colonised F. hirta.ConclusionsThis asymmetric sharing of pollinators may reflect the relative abundance of the two species of fig wasps and differences in host reproductive phenology. Asynchronous flowering of individual F. hirta may favor local retention of pollinators, in contrast to the tree-wide synchrony of F. triloba figs, which can generate local shortages of V. esquirolianae. If the pollinator sharing by male figs of F. triloba and F. hirta also occurs in female figs then this could result in gene flow between them.

The complete chloroplast genome of two related fig species Ficus squamosa and Ficus heterostyla

The Ficus squamosa and Ficus heterostyla share an undescribed pollinating fig wasp Ceratosolen sp. in Xishuangbanna region, which constitutes the most excellent model to study the role of convergent evolution and hybridization in the species-specific fig-wasp mutualism. The plastomes were 160,350 bp for Ficus squamosa and 160,300 bp for F. heterostyla, both in length with the typical quadripartite structure. In the two genomes, the LSC region was 88,615 bp (F. squamosa) and 88,535 bp (F. heterostyla), the SSC region was 20,071 bp (F. squamosa) and 20,101 bp (F. heterostyla), and the IR regions of both figs were 25,832 bp. They contained 113 unique genes, including a set of 78 protein-coding genes, 30 transfer RNA (tRNA) genes, four ribosomal RNA (rRNA) genes, and one pseudogene (infA). Phylogenetic analysis based on the complete chloroplast (cp) genomes within the Ficus genus suggests that they are closely related sister species.

Pollinating and non-pollinating fig wasps (Hymenoptera: Chalcidoidea) from Ficus elastica, the living root bridge tree of Meghalaya in northeast India

The Indian rubber fig tree Ficus elastica Roxb. ex Hornem. Moraceae is the constituent of the iconic living root bridges (LRBs) in Meghalaya, India, and is characterized by a highly specific mutualism between the fig and its pollinating agaonid fig wasp, in which the wasps breed within fig inflorescences. F. elastica is restricted to south and southeast Asia in its distribution. We identified the pollinating fig wasp as Platyscapa clavigera (Mayr 1885) which was first described from F. elastica in Bogor in 1885 and from Singapore in 2017. This is the first record of the pollinator (family Agaonidae) from F. elastica in Meghalaya, northeast India, in the westernmost portion of the fig’s range. We also discovered and identified in F. elastica, a non-pollinating fig wasp of the genus Micranisa which appears close to Micranisa ralianga Mathew and Balakrishnan 1981 (Pteromalidae). This fig wasp has not been earlier reported anywhere from the closed urn-shaped inflorescences (i.e. syconia) of F. elastica and was only described from the syconia of Ficus altissima Blume in 1981 from Meghalaya. Notes on the morphology of both fig wasps are provided and illustrated. The phenology and developmental cycle of F. elastica syconia are documented. Evidence of passive pollination was confirmed in F. elastica which sheds light on the evolution of character traits in figs and their wasps

More examples of breakdown the 1:1 partner specificity between figs and fig wasps

BackgroundThe obligate mutualism between fig trees (Ficus, Moraceae) and pollinating fig wasps (Agaonidae) is a model system for studying co-evolution due to its perceived extreme specificity, but recent studies have reported a number of examples of trees pollinated by more than one fig wasp or sharing pollinators with other trees. This will make the potential of pollen flow between species and hybridization more likely though only few fig hybrids in nature have been found. We reared pollinator fig wasps from figs of 13 Chinese fig tree species and established their identity using genetic methods in order to investigate the extent to which they were supporting more than one species of pollinator (co-pollinator).ResultsOur results showed (1) pollinator sharing was frequent among closely-related dioecious species (where pollinator offspring and seeds develop on different trees); (2) that where two pollinator species were developing in figs of one host species there was usually one fig wasp with prominent rate than the other. An exception was F. triloba, where its two pollinators were equally abundant; (3) the extent of co-pollinator within one fig species is related to the dispersal ability of them which is stronger in dioecious figs, especially in small species.ConclusionsOur results gave more examples to the breakdown of extreme specificity, which suggest that host expansion events where pollinators reproduce in figs other than those of their usual hosts are not uncommon among fig wasps associated with dioecious hosts. Because closely related trees typically have closely related pollinators that have a very similar appearance, the extent of pollinator-sharing has probably been underestimated. Any pollinators that enter female figs carrying heterospecific pollen could potentially generate hybrid seed, and the extent of hybridization and its significance may also have been underestimated.