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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... more
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 Bou£ek, 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.
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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... more
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 reappearance of winged morphs.
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Fig trees and their pollinating fig wasps arose about 75 million years ago in the Cretaceous period. Several other groups of chal- cid wasps also utilize figs for larval development, including syco- phagines, the putative sister group to... more
Fig trees and their pollinating fig wasps arose about 75 million years ago in the Cretaceous period. Several other groups of chal- cid wasps also utilize figs for larval development, including syco- phagines, the putative sister group to pollinating fig wasps. Whereas stone and amber fossil pollinators are known, no fossils representing non-pollinating fig wasp groups have been con- firmed previously. Here, we describe the first Sycophaginae from the c.15–20 Ma Dominican amber, Idarnes thanatos sp. nov. Farache, Rasplus, Pereira and Compton, and discuss its relation- ships within the Idarnes carme species group. Additionally, we use linear regression to compare body size, ovipositor sheaths length, and host fig size data from extant Idarnes species to estimate the size of its host figs. Idarnes thanatos was most likely associated with small to medium sized figs (diameter ≤1.0 cm), that were likely to have been dispersed by birds and primates. The discovery of this close relative of extant non-pollinating fig wasps suggests that early Miocene and modern fig wasp communities may share similar ecological and functional features.
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Mutualism is the name given to interspecific interactions which result in benefit for all partners involved. However, such cooperation is open to opportunistic strategies: individuals that extract the benefit from the partner, but do not... more
Mutualism is the name given to interspecific interactions which result in benefit for all partners involved. However, such cooperation is open to opportunistic strategies: individuals that extract the benefit from the partner, but do not offer any benefit in exchange. The fig-fig wasp interaction is an appropriate case to study mutualistic interactions and opportunistic strategies (parasites of mutualism). Plants of the genus Ficus maintain a mutualistic interaction with tiny pollinating wasps (Agaonidae) and are exploited by other non-pollinating fig wasp species. This study aimed to assess the effect of non-pollinating wasps over the mutualistic relation between Ficus citrifolia (Moraceae) and its pollinator Pegoscapus tonduzi Grandi, 1919 (Hymenoptera, Agaonidae). We compared both seed (female plant function) and pollinating female (male plant function) production rates among samples of high- and low-infested figs by non-pollinating wasps, sampled in three Brazilian cities, Londrina (State of Paraná), Campinas and Ribeirão Preto (state of São Paulo), Brazil. Our results have shown a negative impact over both female and male floral reproductive components. This effect was higher on the male plant component (production of pollinating females). Pollinator production was approximately seven times lower in infested figs, whereas seed production was 1.5 times lower in those figs. We discuss hypotheses about mutualism stability with the occurrence of opportunistic species.
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Ficus microcarpa is an Asian g tree that is ornamentally cultivated. The specic pollinator, Eupristina verticillata Waterston, and the non-pollinators Walkerella microcarpae Bou ek and Philotrypesis emeryi Grandi, have been reported... more
Ficus microcarpa is an Asian g tree that is ornamentally cultivated. The specic pollinator, Eupristina verticillata Waterston, and the non-pollinators Walkerella microcarpae Bou ek and Philotrypesis emeryi Grandi, have been reported associated to F. microcarpa in Brazil. In here we report for the rst 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 inux of these wasps into Brazil. Considering that ~20 g wasp species are associated with F. microcarpa in its native area, novel occurrences can be reported in the future in Brazil.
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Besides the pollinators (Agaonidae), several chalcidoid lineages of nonpollinating fig wasps are strictly dependent on Ficus (Moraceae) for reproduction. Overall, nonpollinating fig wasps have not received much consideration. Reliable... more
Besides the pollinators (Agaonidae), several chalcidoid lineages of nonpollinating fig wasps are strictly dependent on Ficus (Moraceae) for reproduction. Overall, nonpollinating fig wasps have not received much consideration. Reliable phylogenetic and taxonomic frameworks are lacking for most groups, which hampers progress in our understanding of the evolution and functioning of fig wasp communities. Here we focus on Anidarnes Boucˇek (Sycophaginae), a member of the Neotropical fig wasp community. We present a detailed morphological analysis as well as the first molecular phylogeny of the genus inferred from two nuclear and two mitochondrial genes (Cytb, COI, EF-1α and 28S rRNA, 3937 bp). Twelve species are recognised, illustrated and a key to their identification is provided. Of those, only three were previously described and are redescribed here: A. bicolor (Ashmead); A. globiceps (Mayr) and A. brevicauda Boucˇek. In addition, nine new species are described: Anidarnes brevior sp.n. Farache & Rasplus (Costa Rica); A. dissidens sp.n. Farache & Rasplus (Brazil); A. gracilis sp.n. Farache & Rasplus (Costa Rica); A. isophlebiae sp.n. Farache & Rasplus (Costa Rica); A. jimeneziae sp.n. Farache & Rasplus (Costa Rica); A. longiscutellum sp.n. Farache & Rasplus (Brazil); A. martinae sp.n. Farache & Rasplus (U.S.A., Florida); A. nigrus sp.n. Farache & Rasplus (Colombia) and A. rugosus sp.n. Farache & Rasplus (Brazil). Finally, phylogenetic relationships inferred using parsimony, bayesian and maximum likelihood methods are discussed in the light of our morphological observations and the host fig tree taxonomy.
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Paracrias strii Schauff, 1985 and P. ceratophaga Palmieri & Hansson sp. nov. are rst record in Brazil and both are associated with Ceratopus Schoenherr larvae (Coleoptera, Curculionidae) reared from syconia of two species of g-trees.... more
Paracrias strii Schauff, 1985 and P. ceratophaga Palmieri & Hansson sp. nov. are  rst record in Brazil and both are associated with Ceratopus Schoenherr larvae (Coleoptera, Curculionidae) reared from syconia of two species of  g-trees. Both Paracrias species are diagnosed and illustrated. Males of P. ceratophaga sp. nov. are described. The association of Paracrias with weevil larvae is brie y discussed.
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e species of Pseudidarnes are revised, and six species are described: P. acaudus Farache & Rasplus, sp. n.; P. astridae Farache & Rasplus, sp. n.; P. badiogeminus Farache & Rasplus, sp. n.; P. cooki Farache & Rasplus, sp. n.; P.... more
e species of Pseudidarnes are revised, and six species are described: P. acaudus Farache & Rasplus, sp. n.; P. astridae Farache & Rasplus, sp. n.; P. badiogeminus Farache & Rasplus, sp. n.; P. cooki Farache & Rasplus, sp. n.; P. kjellbergi Farache & Rasplus, sp. n.; P. laevis Farache & Rasplus, sp. n. Pseudidarnes minerva Girault, 1927 and P.  avicollis Bouček, 1988 are redescribed. A key to the species is provided as well as illustrations for all females and all known males (except the wingless male of P. minerva). We also provided further discussion on ecology, morphological patterns, and host taxonomy. Online dichotomous and multi-access interactive LUCID keys to all Pseudidarnes species are available at http://www. gweb.org/.
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e sycophagines are strictly associated with two subgenera of Ficus L. (Moraceae), namely Sycomorus and Urostigma. ey mostly oviposit through the g wall and lay their eggs within the g owers, being either gall-makers or parasitoids of... more
e sycophagines are strictly associated with two subgenera of Ficus L. (Moraceae), namely Sycomorus and Urostigma.  ey mostly oviposit through the  g wall and lay their eggs within the  g  owers, being either gall-makers or parasitoids of other  g wasps. In this contribution, a new genus of Sycophaginae, Conidarnes Farache & Rasplus, gen. n., is described with seven new species: Conidarnes achterbergi Far- ache & Rasplus, sp. n.; Conidarnes bergi Farache & Rasplus, sp. n.; Conidarnes laevis Farache & Rasplus, sp. n.; Conidarnes santineloi Farache & Rasplus, sp. n.; Conidarnes subtectae Farache & Rasplus, sp. n.; Conidarnes sulcata Farache & Rasplus, sp. n.; and Conidarnes sumatranae Farache & Rasplus, sp. n. Illus- trations, morphological diagnoses, dichotomous keys and multi-entry online keys to species are provided. Conidarnes species strictly occur in the oriental region, and their distribution does not overlap with the distribution of the two other genera belonging to the same clade. Due to their relative rarity, we encourage extensive sampling of Conosycea  gs to improve our knowledge of the genus.
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