The mechanism of pollinator specificity between two sympatric fig varieties: a combination of olfactory signals and contact cues

Background and Aims

Pollinator specificity facilitates reproductive isolation among plants, and mechanisms that generate specificity influence species boundaries. Long-range volatile attractants, in combination with morphological co-adaptations, are generally regarded as being responsible for maintaining extreme host specificity among the fig wasps that pollinate fig trees, but increasing evidence for breakdowns in specificity is accumulating. The basis of host specificity was examined among two host-specific Ceratosolen fig wasps that pollinate two sympatric varieties of Ficus semicordata, together with the consequences for the plants when pollinators entered the alternative host variety.

Methods

The compositions of floral scents from receptive figs of the two varieties and responses of their pollinators to these volatiles were compared. The behaviour of the wasps once on the surface of the figs was also recorded, together with the reproductive success of figs entered by the two Ceratosolen species.

Key Results

The receptive-phase floral scents of the two varieties had different chemical compositions, but only one Ceratosolen species displayed a preference between them in Y-tube trials. Specificity was reinforced at a later stage, once pollinators were walking on the figs, because both species preferred to enter figs of their normal hosts. Both pollinators could enter figs of both varieties and pollinate them, but figs with extra-varietal pollen were more likely to abort and contained fewer seeds. Hybrid seeds germinated at normal rates.

Conclusions

Contact cues on the surface of figs have been largely ignored in previous studies of fig wasp host preferences, but together with floral scents they maintain host specificity among the pollinators of sympatric F. semicordata varieties. When pollinators enter atypical hosts, post-zygotic factors reduce but do not prevent the production of hybrid offspring, suggesting there may be gene flow between these varieties.     

Figs,pollinators, and parasites: A longitudinal study of the effects of nematode infection on fig wasp fitness

Mutualisms are interactions between two species in which the fitnesses of both symbionts benefit from the relationship. Although examples of mutualism are ubiquitous in nature, the ecology, evolution, and stability of mutualism has rarely been studied in the broader, multi-species community context in which they occur. The pollination mutualism between figs and fig wasps provides an excellent model system for investigating interactions between obligate mutualists and antagonists. Compared to the community of non-pollinating fig wasps that develop within fig inflorescences at the expense of fig seeds and pollinators, consequences of interactions between female pollinating wasps and their host-specialist nematode parasites is much less well understood. Here we focus on a tri-partite system comprised of a fig (Ficus petiolaris), pollinating wasp (Pegoscapus sp.), and nematode (Parasitodiplogaster sp.), investigating geographical variation in the incidence of attack and mechanisms through which nematodes may limit the fitness of their wasp hosts at successive life history stages. Observational data reveals that nematodes are ubiquitous across their host range in Baja California, Mexico; that the incidence of nematode infection varies across seasons within- and between locations, and that infected pollinators are sometimes associated with fitness declines through reduced offspring production. We find that moderate levels of infection (1–9 juvenile nematodes per host) are well tolerated by pollinator wasps whereas higher infection levels (≥10 nematodes per host) are correlated with a significant reduction in wasp lifespan and dispersal success. This overexploitation, however, is estimated to occur in only 2.8% of wasps in each generation. The result that nematode infection appears to be largely benign – and the unexpected finding that nematodes frequently infect non-pollinating wasps – highlight gaps in our knowledge of pollinator-Parasitodiplogaster interactions and suggest previously unappreciated ways in which this nematode may influence fig and pollinator fitness, mutualism persistence, and non-pollinator community dynamics.     

Deep mtDNA divergences indicate cryptic species in a fig-pollinating wasp

Background  

Figs and fig-pollinating wasps are obligate mutualists that have coevolved for ca 90 million years. They have radiated together, but do not show strict cospeciation. In particular, it is now clear that many fig species host two wasp species, so there is more wasp speciation than fig speciation. However, little is known about how fig wasps speciate.     

Genome‐wide sequence data suggest the possibility of pollinator sharing by host shift in dioecious figs (Moraceae,Ficus)

The obligate mutualism of figs and fig‐pollinating wasps has been one of the classic models used for testing theories of co‐evolution and cospeciation due to the high species‐specificity of these relationships. To investigate the species‐specificity between figs and fig pollinators and to further understand the speciation process in obligate mutualisms, we examined the genetic differentiation and phylogenetic relationships of four closely related fig‐pollinating wasp species (Blastophaga nipponica, Blastophaga taiwanensis, Blastophaga tannoensis and Blastophaga yeni) in Japan and Taiwan using genome‐wide sequence data, including mitochondrial DNA sequences. In addition, population structure was analysed for the fig wasps and their host species using microsatellite data. The results suggest that the three Taiwanese fig wasp species are a single panmictic population that pollinates three dioecious fig species, which are sympatrically distributed, have large differences in morphology and ecology and are also genetically differentiated. Our results illustrate the first case of pollinator sharing by host shift in the subgenus Ficus. On the other hand, there are strict genetic codivergences between allopatric populations of the two host–pollinator pairs. The possible processes that produce these pollinator‐sharing events are discussed based on the level and pattern of genetic differentiation in these figs and fig wasps.     

Expression and evolutionary divergence of the non-conventional olfactory receptor in four species of fig wasp associated with one species of fig

Background  

The interactions of fig wasps and their host figs provide a model for investigating co-evolution. Fig wasps have specialized morphological characters and lifestyles thought to be adaptations to living in the fig's syconium. Although these aspects of natural history are well documented, the genetic mechanism(s) underlying these changes remain(s) unknown. Fig wasp olfaction is the key to host-specificity. The Or83b gene class, an unusual member of olfactory receptor family, plays a critical role in enabling the function of conventional olfactory receptors. Four Or83b orthologous genes from one pollinator (PFW) (Ceratosolen solmsi) and three non-pollinator fig wasps (NPFWs) (Apocrypta bakeri, Philotrypesis pilosa and Philotrypesis sp.) associated with one species of fig (Ficus hispida) can be used to better understand the molecular mechanism underlying the fig wasp's adaptation to its host. We made a comparison of spatial tissue-specific expression patterns and substitution rates of one orthologous gene in these fig wasps and sought evidence for selection pressures.     

Has Pollination Mode Shaped the Evolution of Ficus Pollen?

Background

The extent to which co-evolutionary processes shape morphological traits is one of the most fascinating topics in evolutionary biology. Both passive and active pollination modes coexist in the fig tree (Ficus, Moraceae) and fig wasp (Agaonidae, Hymenoptera) mutualism. This classic obligate relationship that is about 75 million years old provides an ideal system to consider the role of pollination mode shifts on pollen evolution.

Methods and Main Findings

Twenty-five fig species, which cover all six Ficus subgenera, and are native to the Xishuangbanna region of southwest China, were used to investigate pollen morphology with scanning electron microscope (SEM). Pollination mode was identified by the Anther/Ovule ratio in each species. Phylogenetic free regression and a correlated evolution test between binary traits were conducted based on a strong phylogenetic tree. Seventeen of the 25 fig species were actively pollinated and eight species were passively pollinated. Three pollen shape types and three kinds of exine ornamentation were recognized among these species. Pollen grains with ellipsoid shape and rugulate ornamentation were dominant. Ellipsoid pollen occurred in all 17 species of actively pollinated figs, while for the passively pollinated species, two obtuse end shapes were identified: cylinder and sphere shapes were identified in six of the eight species. All passively pollinated figs presented rugulate ornamentation, while for actively pollinated species, the smoother types - psilate and granulate-rugulate ornamentations - accounted for just five and two among the 17 species, respectively. The relationship between pollen shape and pollination mode was shown by both the phylogenetic free regression and the correlated evolution tests.

Conclusions

Three pollen shape and ornamentation types were found in Ficus, which show characteristics related to passive or active pollination mode. Thus, the pollen shape is very likely shaped by pollination mode in this unique obligate mutualism.     

N:P stoichiometry in Ficus racemosa and its mutualistic pollinator

Aims Nitrogen (N) and phosphorus (P) are limiting nutrients to life across a variety of ecosystems. N:P stoichiometry, concerning the balance of these two elements, has recently received great attention. However, little is known about the nature of N:P stoichiometry in obligate mutualism.Methods N:P stoichiometry of Ficus racemosa and its pollinating wasp Ceratosolen fusciceps, an example of coevolving obligate mutualism, was investigated, and the N:P stoichiometric traits of male versus female wasps were compared.Important findings Nutrient concentrations in C. fusciceps were much higher than in its host. N enrichment in fig wasp was evidently stronger than phosphorus. N concentrations of male fig wasps were significantly higher than those of females, while P concentrations of female fig wasps were remarkably higher than those of male ones. Therefore, N:P ratios in male fig wasps were significantly greater than in female fig wasps. N:P ratio in fig-pollinating wasp displayed linear functions to fig N contents, suggesting that N limitation in fig wasps may dominate the nutritional relationship between fig pollinator and its host. Fig wasp population size had significant influences on N concentrations in host fig and female wasp per se. Driven by the nutritional stress of pollinating and parasite insects, fig fruit preferred increasing its diameter first but not nutrient richness. Values for N and P contents of fig pollinators showed seasonal differences with greater N:P ratios in dry season than in rainy season. The observations suggest that tropical climate change would result in more severe N limitation to fig-pollinating wasp and may further influence the stability of fig–fig wasp mutualism.     

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.     

New insights into the fungal community from the raw genomic sequence data of fig wasp Ceratosolen solmsi

Background

To date, biologists have discovered a large amount of valuable information from assembled genomes, but the abundant microbial data that is hidden in the raw genomic sequence data of plants and animals is usually ignored. In this study, the richness and composition of fungal community were determined in the raw genomic sequence data of Ceratosolen solmsi (RGSD-CS).

Results

To avoid the interference from sequences of C. solmsi, the unmapped raw data (about 17.1%) was obtained by excluding the assembled genome of C. solmsi from RGSD-CS. Comparing two fungal reference datasets, internal transcribed spacer (ITS) and large ribosomal subunit (LSU) of rRNA, the ITS dataset discovered a more diverse fungal community and was therefore selected as the reference dataset for evaluating the fungal community based on the unmapped raw data. The threshold of 95% sequence identity revealed many more matched fungal reads and fungal richness in the unmapped raw data than those by identities above 95%. Based on the threshold of 95% sequence identity, the fungal community of RGSD-CS was primarily composed of Saccharomycetes (88.4%) and two other classes (Agaricomycetes and Sordariomycetes, 8.3% in total). Compared with the fungal community of other reported fig wasps, Agaricomycetes and Eurotiomycetes were found to be unique to C. solmsi. In addition, the ratio of total fungal reads to RGSD-CS was estimated to be at least 4.8 × 10⁻³, which indicated that a large amount of fungal data was contained in RGSD-CS. However, rarefaction measure indicated that a deeper sequencing coverage with RGSD-CS was required to discover the entire fungal community of C. solmsi.

Conclusion

This study investigated the richness and composition of fungal community in RGSD-CS and provided new insights into the efficient study of microbial diversity using raw genomic sequence data.

Electronic supplementary material

The online version of this article (doi:10.1186/s12866-015-0370-3) contains supplementary material, which is available to authorized users.     

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.     

High-Efficiency Thermal Asymmetric Interlaced PCR (hiTAIL-PCR) for Determination of a Highly Degenerated Prophage WO Genome in a Wolbachia Strain Infecting a Fig Wasp Species

Temperate bacteriophage WO is a model system for studying tripartite interactions among viruses, bacteria, and eukaryotes, especially investigations of the genomic stability of obligate intracellular bacteria. Few WO genomes exist because of the difficulty in isolating viral DNA from eukaryotic hosts, and most reports are by-products of Wolbachia sequencing. Only one partial genome of a WO phage has been determined directly from isolated particles. We determine the complete genome sequence of prophage WO (WOSol) in Wolbachia strain wSol, which infects the fig wasp Ceratosolen solmsi (Hymenoptera: Chalcidoidea), by high-efficiency thermal asymmetric interlaced PCR. The genome of WOSol is highly degenerated and disrupted by a large region (14,267 bp) from Wolbachia. Consistent with previous molecular studies of multiple WO genomes, the genome of WOSol appears to have evolved by single nucleotide mutations and recombinations.     

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.     

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.     

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.     

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.     

Ecology of a fig ant–plant

Mutualistic interactions are embedded in networks of interactions that affect the benefits accruing to the mutualistic partners. Figs and their pollinating wasps are engaged in an obligate mutualism in which the fig is dependent on the fig pollinator for pollination services and the pollinator is dependent on fig ovules for brood sites. This mutualism is exploited by non-pollinating fig wasps that utilise the same ovules, but do not provide a pollination service. Most non-pollinating wasps oviposit from outside the inflorescence (syconium), where they are vulnerable to ant predation. Ficus schwarzii is exposed to high densities of non-pollinating wasps, but Philidris sp. ants patrolling the syconia prevent them from ovipositing. Philidris rarely catch wasps, but the fig encourages the patrolling by providing a reward through extra-floral nectaries on the surface of syconia. Moreover, the reward is apparently only produced during the phase when parasitoids are ovipositing. An ant-exclusion experiment demonstrated that, in the absence of ants, syconia were heavily attacked and many aborted as a consequence. Philidris was normally rare on the figs during the receptive phase or at the time of day when wasp offspring are emerging, so predation on pollinators was limited. However, Myrmicaria sp. ants, which only occurred on three trees, preyed substantially on pollinating as well as non-pollinating wasps. F. schwarzii occurs in small clusters of trees and has an exceptionally rapid crop turnover. These factors appear to promote high densities of non-pollinating wasps and, as a consequence, may have led to both a high incidence of ants on trees and increased selective pressure on fig traits that increase the payoffs of the fig–ant interaction for the fig. The fig receives no direct benefit from the reward it provides, but protects pollinating wasps that will disperse its pollen.     

Restructuring of a mutualism following introduction of Australian fig trees and pollinating wasps to Europe and the USA

Figs and fig-pollinating wasps are obligate mutualists that require each other to complete sexual reproduction. However, landscapers can establish populations of fig trees outside their native ranges by propagation through exported seeds, seedlings or cuttings. Once mature, these trees could be colonized by pollinating wasps and/or various non-pollinating wasps that also develop in figs. In recent decades, the Australian endemic Ficus rubiginosa has been planted widely in the Mediterranean region and in parts of the USA. Observation of ripe fruit production suggested that a pollination mutualism has been re-established by pollinating wasps colonizing trees in the plant’s introduced range. We therefore used sampling of pollinators from mainland Spain, Tenerife and California (USA) and molecular studies to characterize the restructured mutualism and compare it with the native range. In the native range, the plant is pollinated by five wasp species that form the Pleistodontes imperialis complex. However, all wasps in the introduced ranges belonged to just one of these species (P. imperialis sp. 1). Moreover, their mtDNA diversity was close to zero and the sequences clearly link them with the native southern population of this species. None of the?>?20 non-pollinating wasp species from the native range were found in the introduced ranges. In summary, the restructured mutualism has been dramatically simplified, lacking all non-pollinating wasps and all but one pollinator species from the native range. Moreover, the one pollinator species to establish successfully shows a drastic reduction in genetic diversity relative to its source population.     

Secondary galling: a novel feeding strategy among ‘non‐pollinating’ fig wasps from Ficus curtipes

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Detecting the elusive cost of parasites on fig seed production

Mutualisms provide essential ecosystem functions such as pollination and contribute considerably to global biodiversity. However, they are also exploited by parasites that remove resources and thus impose costs on one or both of the mutualistic partners. The fig/pollinator interaction is a classic obligate mutualism; it is pantropical and involves >750 Ficus species and their host-specific pollinating wasps (family Agaonidae). Figs also host parasites of the mutualism that should consume pollinators or seeds, depending on their larval ecology. We collected data from a large crop of figs on Ficus glandifera var. brachysyce in a Sulawesi rainforest with an unusually high number of Eukoebelea sp. parasites. We found that these parasites have a significant negative correlation with fig seed production as well as with pollinator offspring production. Eukoebelea wasps form the basal genus in subfamily Sycophaginae (Chalcidoidea) and their larval biology is considered unknown. Our analysis suggests that they feed as flower gallers and impose direct costs on the fig tree, but a strategy including the consumption of pollinator larvae cannot be ruled out. We also present baseline data on the composition of the fig wasp community associated with F. glandifera var brachysyce and light trap catch data.