Yue-Guan Fu

Obligate mutualism within a host drives the extreme specialization of a fig wasp genome

BackgroundFig pollinating wasps form obligate symbioses with their fig hosts. This mutualism arose approximately 75 million years ago. Unlike many other intimate symbioses, which involve vertical transmission of symbionts to host offspring, female fig wasps fly great distances to transfer horizontally between hosts. In contrast, male wasps are wingless and cannot disperse. Symbionts that keep intimate contact with their hosts often show genome reduction, but it is not clear if the wide dispersal of female fig wasps will counteract this general tendency. We sequenced the genome of the fig wasp Ceratosolen solmsi to address this question.ResultsThe genome size of the fig wasp C. solmsi is typical of insects, but has undergone dramatic reductions of gene families involved in environmental sensing and detoxification. The streamlined chemosensory ability reflects the overwhelming importance of females finding trees of their only host species, Ficus hispida, during their fleeting adult lives. Despite long-distance dispersal, little need exists for detoxification or environmental protection because fig wasps spend nearly all of their lives inside a largely benign host. Analyses of transcriptomes in females and males at four key life stages reveal that the extreme anatomical sexual dimorphism of fig wasps may result from a strong bias in sex-differential gene expression.ConclusionsOur comparison of the C. solmsi genome with other insects provides new insights into the evolution of obligate mutualism. The draft genome of the fig wasp, and transcriptomic comparisons between both sexes at four different life stages, provide insights into the molecular basis for the extreme anatomical sexual dimorphism of this species.

Molecular approaches to identify cryptic species and polymorphic species within a complex community of fig wasps.

Cryptic and polymorphic species can complicate traditional taxonomic research and both of these concerns are common in fig wasp communities. Species identification is very difficult, despite great effort and the ecological importance of fig wasps. Herein, we try to identify all chalcidoid wasp species hosted by one species of fig, using both morphological and molecular methods. We compare the efficiency of four different DNA regions and find that ITS2 is highly effective for species identification, while mitochondrial COI and Cytb regions appear less reliable, possibly due to the interference signals from either nuclear copies of mtDNA, i.e. NUMTs, or the effects of Wolbachia infections. The analyses suggest that combining multiple markers is the best choice for inferring species identifications as any one marker may be unsuitable in a given case.

Rampant host switching and multiple female body colour transitions in Philotrypesis (Hymenoptera: Chalcidoidea: Agaonidae)

Figs (Ficus, Moraceae) and their associated fig waSPS (Hymenoptera, Chalcidoidea and Agaonidae) have attracted much attention and have been used as a model system for many studies. Fig waSPS belonging to the genus Philotrypesis are very common in most figs but their taxonomy, ecology and biology are currently poorly explored. A previous study on African Philotrypesis showed that their host association is phylogenetically conserved at subsection level. We reconstructed a molecular phylogeny with extended sampling from seven sections of figs. Our study suggested that the diversification of Philotrypesis is less constrained by host figs. Host switching is rampant between figs at species level and even at section level. We also investigated the evolution of the body colour forms in female Philotrypesis. Our study first suggested that female body colour is not evolutionarily stable and that there have been multiple transitions. Possible mechanisms for multiple colour transitions are expected to be determined in the near future.

Chaos of Wolbachia sequences inside the compact fig syconia of Ficus benjamina (Ficus: Moraceae)

Figs and fig wasps form a peculiar closed community in which the Ficus tree provides a compact syconium (inflorescence) habitat for the lives of a complex assemblage of Chalcidoid insects. These diverse fig wasp species have intimate ecological relationships within the closed world of the fig syconia. Previous surveys of Wolbachia, maternally inherited endosymbiotic bacteria that infect vast numbers of arthropod hosts, showed that fig wasps have some of the highest known incidences of Wolbachia amongst all insects. We ask whether the evolutionary patterns of Wolbachia sequences in this closed syconium community are different from those in the outside world. In the present study, we sampled all 17 fig wasp species living on Ficus benjamina, covering 4 families, 6 subfamilies, and 8 genera of wasps. We made a thorough survey of Wolbachia infection patterns and studied evolutionary patterns in wsp (Wolbachia Surface Protein) sequences. We find evidence for high infection incidences, frequent recombination between Wolbachia strains, and considerable horizontal transfer, suggesting rapid evolution of Wolbachia sequences within the syconium community. Though the fig wasps have relatively limited contact with outside world, Wolbachia may be introduced to the syconium community via horizontal transmission by fig wasps species that have winged males and visit the syconia earlier.

Delimitation and description of the immature stages of a pollinating fig wasp, Ceratosolen solmsi marchali Mayr (Hymenoptera: Agaonidae)

The mutualism between fig trees and their wasp pollinators is a model system for many ecological and evolutionary studies. However, the immature stages of pollinating fig wasps have rarely been studied. We monitored developing fig wasps of known ages and performed a series of dissections at 24 h intervals to identify key developmental traits of Ceratosolen solmsi marchali Mayr (Hymenoptera: Agaonidae), a pollinator of Ficus hispida L. (Moraceae). We identified where in the Ficus ovary eggs were deposited and time to hatch. We were also able to identify the timing and key underlying characters of five larval instars, three sub-pupal stages, and a single prepupal stage. We provide detailed morphological descriptions for the key stages and report some behavioral observations of the wasps in the several developmental stages we recorded. Scanning electron microscope images were taken.

Adaptation of the externally feeding bug Elasmucha necopinata (Hemiptera: Acanthosomatidae) to its fig host

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

Brood size: a major factor influencing male dimorphism in the non‐pollinating fig wasp Sycobia sp.

1 Male polymorphisms have been described in some non‐pollinating fig wasps, as well as in other animals. The proximal basis and the maintenance of alternative male reproductive strategies are either genetic or environmental. 2 Here we studied male dimorphism in the non‐pollinating fig wasp Sycobia sp. We conducted experimental manipulations to study the factors influencing offspring male morph allocations and explore a possible basis for the determination and maintenance of male dimorphism in Sycobia sp. 3 The results showed that brood size was the major and underlying factor influencing the male morph ratio. When the brood size increases, the wingless male ratio also increases. 4 Also, our results indicated that there was no direct maternal control on offspring male morph allocation. 5 Male dimorphism in Sycobia sp. probably represents an environmentally determined conditional strategy, which responded to offspring population density at the level of the individual fig.

Pollinating fig wasp Ceratosolen solmsi adjusts the offspring sex ratio to other foundresses

Abstract  Local mate competition theory predicts that offspring sex ratio in pollinating fig wasps is female‐biased when there is only one foundress, and increased foundress density results in increased offspring sex ratio. Information of other foundresses and clutch size have been suggested to be the main proximate explanations for sex ratio adjustment under local mate competition. Our focus was to show the mechanism of sex ratio adjustment in a pollinating fig wasp, Ceratosolen solmsi Mayr, an obligate pollinator of the functionally dioecious fig, Ficus hispida Linn., with controlled experiments in the field. First, we obtained offspring from one pollinator and offspring at different oviposition sequences, and found that offspring sex ratio decreased with clutch size, and pollinators produced most of their male offspring at the start of bouts, followed by mostly females. Second, we found that offspring sex ratio increased with foundress density, and pollinators did adjust their offspring sex ratio to other females in the oviposition patches. We suggest that when oviposition sites are not limited, pollinators will mainly adjust their offspring sex ratio to other foundresses independent of clutch size changes, whereas adjusting clutch size may be used to adjust sex ratio when oviposition sites are limited.

A NEW SPECIES OF METAPHYCUS MERCET (HYMENOPTERA: ENCYRTIDAE) FROM CHINA, PARASITOID OF PARASAISSETIA NIGRA (NIETNER) (HOMOPTERA: COCCOIDEA)

ABSTRACT Metaphycus parasaissetiae sp. n., belonging to M. zebratus species group, is described from China. Photomicrographs are provided to illustrate morphological characters of the species. Metaphycus parasaissetiae is an important parasitoid of the Nigra Scale, Parasaissetia nigra (Nietner).

Nymphal antennae and antennal sensilla in Aleurodicus dispersus (Hemiptera: Aleyrodidae).

Whiteflies have distinct nymphal stages: their first stage is mobile, whereas the later immature stages are sessile. The developmental and structural changes of antennae and antennal sensilla in whiteflies during these stages have rarely been investigated. This paper describes the morphology of antennae and antennal sensilla in four nymphal stages of Aleurodicus dispersus based on scanning electron microscopy. There were significant differences found in shape and length of the antennae, and differences in type, number, morphological structure and distributional pattern of antennal sensilla in the four nymphal stages of A. dispersus. We found two types of sensilla on the antennae of first-instar nymph, three types on the third-instar nymphal antennae, four types on the second-instar and seven types on the fourth-instar nymphal antennae. Sensilla trichoidea (ST) and elevated sensilla placodea were found on the antennae of each nymphal stage, sensilla chaetica only occurred on the antennae of fourth-instar nymph. Sensilla furcatea occurred on the antennae of second- and third-instar nymphs, and sensilla basiconica were found on the antennae of second- and fourth-instar nymphs. In addition, there were sensilla campaniform and sensilla coeloconica found only on the antennae of fourth-instar nymph, whereas the ST of fourth-instar nymphs included sensilla trichoidea 1 and sensilla trichoidea 2. The possible functions of antennal sensilla are discussed. Our results contribute to a better understanding of the development of the olfactory system of whitefly nymphal stages, and provide a basis for further exploration of chemical communication mechanisms between whiteflies and host plants.