title: Phylogeny and evolution of life-history strategies in the Sycophaginae non-pollinating fig wasps (Hymenoptera, Chalcidoidea).
abstract: 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.
authors.affiliation: INRA-UMR Centre de Biologie et de Gestion des Populations, CBGP, (INRA/IRD/CIRAD/Montpellier SupAgro), Campus international de Baillarguet, CS 30016, 34988 Montferrier-sur Lez, France. cruaud@supagro.inra.fr
authors.forename: Astrid
authors.initials: A
authors.lastname: Cruaud
authors.forename: Roula
authors.initials: R
authors.lastname: Jabbour-Zahab
authors.forename: Gwenaëlle
authors.initials: G
authors.lastname: Genson
authors.forename: Finn
authors.initials: F
authors.lastname: Kjellberg
authors.forename: Noppol
authors.initials: N
authors.lastname: Kobmoo
authors.forename: Simon
authors.initials: S
authors.lastname: van Noort
authors.forename: Yang
authors.initials: Y
authors.lastname: Da-Rong
authors.forename: Peng
authors.initials: P
authors.lastname: Yan-Qiong
authors.forename: Rosichon
authors.initials: R
authors.lastname: Ubaidillah
authors.forename: Paul E
authors.initials: PE
authors.lastname: Hanson
authors.forename: Otilene
authors.initials: O
authors.lastname: Santos-Mattos
authors.forename: Fernando H A
authors.initials: FH
authors.lastname: Farache
authors.forename: Rodrigo A S
authors.initials: RA
authors.lastname: Pereira
authors.forename: Carole
authors.initials: C
authors.lastname: Kerdelhué
authors.forename: Jean-Yves
authors.initials: JY
authors.lastname: Rasplus
mesh_terms: D000818:Animals; D005075:Biological Evolution; D005260:Female; D030681:Ficus / Q000469:parasitology*; D008297:Male; D008969:Molecular Sequence Data; D010802:Phylogeny; D010935:Plant Diseases / Q000469:parasitology*; D014863:Wasps / Q000033:anatomy & histology / Q000145:classification* / Q000235:genetics*