RESUMEN
BACKGROUND: Non-pollinating Sycophaginae (Hymenoptera, Chalcidoidea) form small communities within Urostigma and Sycomorus fig trees. The species show differences in galling habits and exhibit apterous, winged or dimorphic males. The large gall inducers oviposit early in syconium development and lay few eggs; the small gall inducers lay more eggs soon after pollination; the ostiolar gall-inducers enter the syconium to oviposit and the cleptoparasites oviposit in galls induced by other fig wasps. The systematics of the group remains unclear and only one phylogeny based on limited sampling has been published to date. Here we present an expanded phylogeny for sycophagine fig wasps including about 1.5 times the number of described species. We sequenced mitochondrial and nuclear markers (4.2 kb) on 73 species and 145 individuals and conducted maximum likelihood and Bayesian phylogenetic analyses. We then used this phylogeny to reconstruct the evolution of Sycophaginae life-history strategies and test if the presence of winged males and small brood size may be correlated. RESULTS: The resulting trees are well resolved and strongly supported. With the exception of Apocrytophagus, which is paraphyletic with respect to Sycophaga, all genera are monophyletic. The Sycophaginae are divided into three clades: (i) Eukoebelea; (ii) Pseudidarnes, Anidarnes and Conidarnes and (iii) Apocryptophagus, Sycophaga and Idarnes. The ancestral states for galling habits and male morphology remain ambiguous and our reconstructions show that the two traits are evolutionary labile. CONCLUSIONS: The three main clades could be considered as tribes and we list some morphological characters that define them. The same biologies re-evolved several times independently, which make Sycophaginae an interesting model to test predictions on what factors will canalize the evolution of a particular biology. The ostiolar gall-inducers are the only monophyletic group. In 15 Myr, they evolved several morphological adaptations to enter the syconia that make them strongly divergent from their sister taxa. Sycophaginae appears to be another example where sexual selection on male mating opportunities favored winged males in species with small broods and wingless males in species with large broods. However, some species are exceptional in that they lay few eggs but exhibit apterous males, which we hypothesize could be due to other selective pressures selecting against the re-appearance of winged morphs.
Asunto(s)
Evolución Biológica , Ficus/parasitología , Filogenia , Enfermedades de las Plantas/parasitología , Avispas/clasificación , Avispas/genética , Animales , Femenino , Masculino , Datos de Secuencia Molecular , Avispas/anatomía & histologíaRESUMEN
Sycophaginae is a group of non-pollinating fig wasps considered closely related to the fig pollinators (Agaoninae, Tetrapusiinae, and Kradibiinae) in the most recent phylogenetic analyses. They occur in all tropical regions and are associated with Ficus subgenera Urostigma and Sycomorus. There are six described genera of Sycophaginae, and two are native and confined to the Neotropics, namely Idarnes Walker, 1843 and Anidarnes Boucek, 1993. Genus Idarnes is divided into three morphologically distinct groups that were proven to be monophyletic by recent molecular phylogenetic analyses. In this paper we reviewed the Idarnes incertus species-group and provide detailed morphological descriptions and illustrations for the species belonging to this group. Three previously described species were redescribed: I. brasiliensis (Mayr, 1906) comb. nov., I. hansoni Boucek, 1993, and I. incertus (Ashmead, 1900). Seventeen new species are described by Farache and Rasplus: I. amacayacuensissp. n., I. amazonicussp. n., I. americanaesp. n., I. badiovertexsp. n., I. brevissp. n., I. brunneussp. n., I. comptonisp. n., I. cremersiaesp. n., I. dimorphicussp. n., I. flavicrussp. n., I. flaviventrissp. n., I. gibberosussp. n., I. gordhisp. n., I. maximussp. n., I. nigriventrissp. n., I. pseudoflavussp. n. and I. ramirezisp. n. We provided keys for the identification of the species as well as for recognising the different species-groups of Idarnes and a closely related genus (Sycophaga Westwood, 1840). Additionally, phylogenetic relationships among 13 species of the I. incertus species-group were inferred using four molecular markers and discussed in the light of Ficus taxonomy and host specificity.
RESUMEN
Ficus microcarpa is an Asian fig tree that is ornamentally cultivated. The specific pollinator, Eupristina verticillata Waterston, and the non-pollinators Walkerella microcarpae Boucek and Philotrypesis emeryi Grandi, have been reported associated to F. microcarpa in Brazil. In here we report for the first time the occurrence of Odontofroggatia ishii Wiebes and Philotrypesis taiwanensis Chen et al in F. microcarpa in Brazil. Our results suggest that P. taiwanensis and O. ishii represent a recent influx of these wasps into Brazil. Considering that approximately 20 fig wasp species are associated with F. microcarpa in its native area, novel occurrences can be reported in the future in Brazil.
Asunto(s)
Ficus , Avispas , Animales , BrasilRESUMEN
Ficus microcarpa is an Asian fig tree that is ornamentally cultivated. The specific pollinator, Eupristina verticillata Waterston, and the non-pollinators Walkerella microcarpae Bouček and Philotrypesis emeryi Grandi, have been reported associated to F. microcarpa in Brazil. In here we report for the first time the occurrence of Odontofroggatia ishii Wiebes and Philotrypesis taiwanensis Chen et al in F. microcarpa in Brazil. Our results suggest that P. taiwanensis and O. ishii represent a recent influx of these wasps into Brazil. Considering that ~20 fig wasp species are associated with F. microcarpa in its native area, novel occurrences can be reported in the future in Brazil.
Ficus microcarpa é uma figueira asiática cultivada com finalidades ornamentais. A espécie polinizadora específica, Eupristina verticillata Waterston, e as não-polinizadoras Walkerella microcarpae Bouček e Philotrypesis emeryi Grandi, foram relatadas associadas a F. microcarpa no Brasil. No presente estudo, relatamos pela primeira vez a ocorrência de Odontofroggatia ishii Wiebes e Philotrypesis taiwanensis Chen et al em F. microcarpa no Brasil. Nossos resultados sugerem que tais vespas foram introduzidas recentemente no país. Considerando que ~20 espécies de vespas são associadas a F. microcarpa em sua região de origem, novas ocorrências poderão ser registradas no futuro no Brasil.