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A multilocus phylogeny of the world Sycoecinae fig wasps (Chalcidoidea: Pteromalidae).

Cruaud A, Underhill JG, Huguin M, Genson G, Jabbour-Zahab R, Tolley KA, Rasplus JY, van Noort S - PLoS ONE (2013)

Bottom Line: We therefore proposed a new classification for the subfamily.Comparisons of our results with fig phylogenies showed that, despite sycoecines being internally ovipositing wasps host-switches are common incidents in their evolutionary history.Finally, by studying the evolutionary properties of the markers we used and profiling their phylogenetic informativeness, we predicted their utility for resolving phylogenetic relationships of Chalcidoidea at various taxonomic levels.

View Article: PubMed Central - PubMed

Affiliation: INRA, UMR1062 CBGP Centre de Biologie pour la Gestion des Populations, Montferrier-sur-Lez, France.

ABSTRACT
The Sycoecinae is one of five chalcid subfamilies of fig wasps that are mostly dependent on Ficus inflorescences for reproduction. Here, we analysed two mitochondrial (COI, Cytb) and four nuclear genes (ITS2, EF-1α, RpL27a, mago nashi) from a worldwide sample of 56 sycoecine species. Various alignment and partitioning strategies were used to test the stability of major clades. All topologies estimated using maximum likelihood and Bayesian methods were similar and well resolved but did not support the existing classification. A high degree of morphological convergence was highlighted and several species appeared best described as species complexes. We therefore proposed a new classification for the subfamily. Our analyses revealed several cases of probable speciation on the same host trees (up to 8 closely related species on one single tree of F. sumatrana), which raises the question of how resource partitioning occurs to avoid competitive exclusion. Comparisons of our results with fig phylogenies showed that, despite sycoecines being internally ovipositing wasps host-switches are common incidents in their evolutionary history. Finally, by studying the evolutionary properties of the markers we used and profiling their phylogenetic informativeness, we predicted their utility for resolving phylogenetic relationships of Chalcidoidea at various taxonomic levels.

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Related in: MedlinePlus

Phylogram of relationships among the Sycoecinae obtained from the analysis of the MAFFT alignment (combined dataset, without Gblocks cleaning, 6 partitions: mtDNAcodon1&2, mtDNAcodon3, EF-1α, ITS2, RpL27a, magonashi).Uppercase letters refer to clades discussed in the text. The new classification is indicated by colored bars on the right (yellow = oriental species, blue = afrotropical species). Nodes with likelihood bootstrap (BP) values < 70 have been collapsed. BP (> 70) and Bayesian posterior probabilities (> 0.90) are indicated at nodes. Illustrations of female habitus for the main clades are provided on the right. Host fig tree subsections are indicated between parentheses. Black boxes at nodes show cases of probable speciation on a single host Ficus species.
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pone-0079291-g001: Phylogram of relationships among the Sycoecinae obtained from the analysis of the MAFFT alignment (combined dataset, without Gblocks cleaning, 6 partitions: mtDNAcodon1&2, mtDNAcodon3, EF-1α, ITS2, RpL27a, magonashi).Uppercase letters refer to clades discussed in the text. The new classification is indicated by colored bars on the right (yellow = oriental species, blue = afrotropical species). Nodes with likelihood bootstrap (BP) values < 70 have been collapsed. BP (> 70) and Bayesian posterior probabilities (> 0.90) are indicated at nodes. Illustrations of female habitus for the main clades are provided on the right. Host fig tree subsections are indicated between parentheses. Black boxes at nodes show cases of probable speciation on a single host Ficus species.

Mentions: Species identification was based on morphological characteristics according to the current literature and was conducted by SvN and JYR. Morphological observations were performed on 10 to 20 representatives of each species. A total of 81 sycoecine specimens representing all the known genera and 56 species, of which 50% are undescribed, were included in the molecular study (Table 2). Seventy percent of the species were represented by sequences from one specimen only. The type-species of the genera Seres (S. armipes) and Philocaenus (P. barbarus) were also included in our analyses. All necessary permits were obtained for the collection of specimens in nature reserves and national parks (Ministère des Eaux et Forêts et du Reboisement, Libreville, Gabon permit granted by Emile Mamfoumbi Kombila, Directeur de la Faune et de la Chasse; Uganda Wildlife Authority UWA/TDO/33/02; UNCST NS 214; Namibia Ministry of Environment and Tourism permit 1289/2008; Ezemvelo KZN Wildlife permits 2985/1999; 24139/2000; 29820/2002; 4502/2005; 4345/2005; 4346/2005;1958/2007; Kenyan Wildlife Service KWS/RP/5001; Cape Nature permits 288/1999; AAA004-00092-0035; AAA007-00324-0035; Northern Cape Province permits Fauna 131/2010, Fauna 132/2010; Eastern Cape Province permits CRO 101/11CR and CRO 102/11CR; Permits for field work in Borneo were obtained from the Sarawak Forestry Corporation and the Forest Research Institute Malaysia; Permits for specimen collection in China were obtained from the Chinese Natural Science Foundation (30670358, 30571507), KSCX2-YW-Z-003; Permits for the collection of specimen in Sulawesi were obtained from the Research Center for Biology LIPI (No : 3180/SU.3/KS/2007); Specimen collection in Taiwan was funded by the ANR project BioFigs/National Science Council, Taiwan, R. O. C. code: 98WFA0100291). Wasps were collected by sampling figs containing either adults or developing wasp larvae. The figs, containing wasps no more than a few days short of their emergence, were placed in handmade wasp-rearing chambers. Once emerged, adult wasps were killed and preserved in 95% ethanol. While the relationships between the chalcidoid subfamilies remain controversial, closer and more distant relatives were included as outgroups [9,10,35]. Six species belonging to the genera Haltichella (Haltichellinae, Chalcididae), Bruchophagus (Eurytominae, Eurytomidae), Grandiana (Otitesellinae, Pteromalidae), Micranisa (Otitesellinae, Pteromalidae), Walkerella (Otitesellinae, Pteromalidae), and Megastigmus (Megastigminae, Torymidae) were used (Table 2). Each time destructive extraction was used, vouchers were selected among specimens sampled from the same fig tree and the same fig after careful identification. Vouchers are deposited at CBGP, Montferrier-sur-Lez, France and Iziko South African Museum, Cape Town (SAMC). A high definition image library of vouchers was also constructed, using the EntoVision Premium Portable Imaging System, to allow future identification of specific taxa and traceability of our results (see Figure 1 for examples).


A multilocus phylogeny of the world Sycoecinae fig wasps (Chalcidoidea: Pteromalidae).

Cruaud A, Underhill JG, Huguin M, Genson G, Jabbour-Zahab R, Tolley KA, Rasplus JY, van Noort S - PLoS ONE (2013)

Phylogram of relationships among the Sycoecinae obtained from the analysis of the MAFFT alignment (combined dataset, without Gblocks cleaning, 6 partitions: mtDNAcodon1&2, mtDNAcodon3, EF-1α, ITS2, RpL27a, magonashi).Uppercase letters refer to clades discussed in the text. The new classification is indicated by colored bars on the right (yellow = oriental species, blue = afrotropical species). Nodes with likelihood bootstrap (BP) values < 70 have been collapsed. BP (> 70) and Bayesian posterior probabilities (> 0.90) are indicated at nodes. Illustrations of female habitus for the main clades are provided on the right. Host fig tree subsections are indicated between parentheses. Black boxes at nodes show cases of probable speciation on a single host Ficus species.
© Copyright Policy
Related In: Results  -  Collection

Show All Figures
getmorefigures.php?uid=PMC3818460&req=5

pone-0079291-g001: Phylogram of relationships among the Sycoecinae obtained from the analysis of the MAFFT alignment (combined dataset, without Gblocks cleaning, 6 partitions: mtDNAcodon1&2, mtDNAcodon3, EF-1α, ITS2, RpL27a, magonashi).Uppercase letters refer to clades discussed in the text. The new classification is indicated by colored bars on the right (yellow = oriental species, blue = afrotropical species). Nodes with likelihood bootstrap (BP) values < 70 have been collapsed. BP (> 70) and Bayesian posterior probabilities (> 0.90) are indicated at nodes. Illustrations of female habitus for the main clades are provided on the right. Host fig tree subsections are indicated between parentheses. Black boxes at nodes show cases of probable speciation on a single host Ficus species.
Mentions: Species identification was based on morphological characteristics according to the current literature and was conducted by SvN and JYR. Morphological observations were performed on 10 to 20 representatives of each species. A total of 81 sycoecine specimens representing all the known genera and 56 species, of which 50% are undescribed, were included in the molecular study (Table 2). Seventy percent of the species were represented by sequences from one specimen only. The type-species of the genera Seres (S. armipes) and Philocaenus (P. barbarus) were also included in our analyses. All necessary permits were obtained for the collection of specimens in nature reserves and national parks (Ministère des Eaux et Forêts et du Reboisement, Libreville, Gabon permit granted by Emile Mamfoumbi Kombila, Directeur de la Faune et de la Chasse; Uganda Wildlife Authority UWA/TDO/33/02; UNCST NS 214; Namibia Ministry of Environment and Tourism permit 1289/2008; Ezemvelo KZN Wildlife permits 2985/1999; 24139/2000; 29820/2002; 4502/2005; 4345/2005; 4346/2005;1958/2007; Kenyan Wildlife Service KWS/RP/5001; Cape Nature permits 288/1999; AAA004-00092-0035; AAA007-00324-0035; Northern Cape Province permits Fauna 131/2010, Fauna 132/2010; Eastern Cape Province permits CRO 101/11CR and CRO 102/11CR; Permits for field work in Borneo were obtained from the Sarawak Forestry Corporation and the Forest Research Institute Malaysia; Permits for specimen collection in China were obtained from the Chinese Natural Science Foundation (30670358, 30571507), KSCX2-YW-Z-003; Permits for the collection of specimen in Sulawesi were obtained from the Research Center for Biology LIPI (No : 3180/SU.3/KS/2007); Specimen collection in Taiwan was funded by the ANR project BioFigs/National Science Council, Taiwan, R. O. C. code: 98WFA0100291). Wasps were collected by sampling figs containing either adults or developing wasp larvae. The figs, containing wasps no more than a few days short of their emergence, were placed in handmade wasp-rearing chambers. Once emerged, adult wasps were killed and preserved in 95% ethanol. While the relationships between the chalcidoid subfamilies remain controversial, closer and more distant relatives were included as outgroups [9,10,35]. Six species belonging to the genera Haltichella (Haltichellinae, Chalcididae), Bruchophagus (Eurytominae, Eurytomidae), Grandiana (Otitesellinae, Pteromalidae), Micranisa (Otitesellinae, Pteromalidae), Walkerella (Otitesellinae, Pteromalidae), and Megastigmus (Megastigminae, Torymidae) were used (Table 2). Each time destructive extraction was used, vouchers were selected among specimens sampled from the same fig tree and the same fig after careful identification. Vouchers are deposited at CBGP, Montferrier-sur-Lez, France and Iziko South African Museum, Cape Town (SAMC). A high definition image library of vouchers was also constructed, using the EntoVision Premium Portable Imaging System, to allow future identification of specific taxa and traceability of our results (see Figure 1 for examples).

Bottom Line: We therefore proposed a new classification for the subfamily.Comparisons of our results with fig phylogenies showed that, despite sycoecines being internally ovipositing wasps host-switches are common incidents in their evolutionary history.Finally, by studying the evolutionary properties of the markers we used and profiling their phylogenetic informativeness, we predicted their utility for resolving phylogenetic relationships of Chalcidoidea at various taxonomic levels.

View Article: PubMed Central - PubMed

Affiliation: INRA, UMR1062 CBGP Centre de Biologie pour la Gestion des Populations, Montferrier-sur-Lez, France.

ABSTRACT
The Sycoecinae is one of five chalcid subfamilies of fig wasps that are mostly dependent on Ficus inflorescences for reproduction. Here, we analysed two mitochondrial (COI, Cytb) and four nuclear genes (ITS2, EF-1α, RpL27a, mago nashi) from a worldwide sample of 56 sycoecine species. Various alignment and partitioning strategies were used to test the stability of major clades. All topologies estimated using maximum likelihood and Bayesian methods were similar and well resolved but did not support the existing classification. A high degree of morphological convergence was highlighted and several species appeared best described as species complexes. We therefore proposed a new classification for the subfamily. Our analyses revealed several cases of probable speciation on the same host trees (up to 8 closely related species on one single tree of F. sumatrana), which raises the question of how resource partitioning occurs to avoid competitive exclusion. Comparisons of our results with fig phylogenies showed that, despite sycoecines being internally ovipositing wasps host-switches are common incidents in their evolutionary history. Finally, by studying the evolutionary properties of the markers we used and profiling their phylogenetic informativeness, we predicted their utility for resolving phylogenetic relationships of Chalcidoidea at various taxonomic levels.

Show MeSH
Related in: MedlinePlus