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4. Habitus of selected eulophids. 1. Closterocerus tau (Entedoninae: Entedonini). 2. Astichus sp. (Entiinae). 3. Ophelimus maskelli (Opheliminae). 4. Aprostocetus sp. (Tetrastichinae).
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AbstractA new combined molecular and morphological phylogeny of the Eulophidae is presented with special reference to the subfamily Entedoninae. We examined 28S D2–D5 and CO1 gene regions with parsimony and partitioned Bayesian analyses, and examined the impact of a small set of historically recognized morphological characters on combined analyses....
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... Sulcus across vertex between median and lateral ocelli: 0 = absent (Fig. 14); 1 = present ( Fig. 15: ...
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... External completeness of notauli posteriorly: 0 = reaching trans-scutal articulation ( Figs 23 and 26); 1 = not reaching trans-scutal articulation, essentially absent (Figs 24, 25, 28 and 33). ...
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... Advancement of axillae: 0 = dorsal axillar surface not completely advanced beyond anterior margin of scutellar disc (Figs 23, 24, 26-28 and 33: ax); 1 = dorsal axillar surface completely advanced beyond anterior margin of scutellar disc (Fig. 25: ax). ...
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... axillae are advanced entirely beyond the scutellar disc (Fig. 24) in some genera of Euderomphalini (LaS- alle and Schauff, 1994), causing them to be interpreted as the side lobes of the mesoscutum (Gumovsky, 2002). They are similarly advanced in the outgroup taxa Ceratogramma and Colotrechnus. In other eulophids, the dorsal surface of the axilla extends alongside the scutellar ...
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... Epipygium (Mt 9 ) in females: 0 = separate from Mt 8 (Fig. 39: Mt 9 ); 1 = fused with Mt 8 , forming a syntergum ( Fig. 40: Mt 8+9 ...
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... and Bayesian analyses agree on the higher classification of Eulophidae in all respects except on Entedoninae and Ophelimus (Figs 43-46). Monophyly of Eulophidae, excluding Trisecodes agromyzae, is supported in all analyses. ...
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... results (Figs 45 and 46) support a monophyletic Entiinae with Ophelimus as its sister group. Parsimony (Figs 43 and 44) instead indicates either a clade of Ophelimus + Entiinae with Ophelimus arising within Entiinae, or an unresolved grouping of Ophelimus and Entiinae with parts of Entedoninae. ...
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... results (Figs 45 and 46) support a monophyletic Entiinae with Ophelimus as its sister group. Parsimony (Figs 43 and 44) instead indicates either a clade of Ophelimus + Entiinae with Ophelimus arising within Entiinae, or an unresolved grouping of Ophelimus and Entiinae with parts of Entedoninae. The clade of Ophelimus plus or within Entiinae has not been recognized by any previous author. ...
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... lack all three of the characters specified by Coote (1994) as helpful in recognizing Entiinae: a bare area under the forewing marginal vein exposing ventral admarginal setae, scutellum overhanging the reduced and concave axillulae, and the separated Mt 9 (character 30:0, Fig. 39). These characterize all Entiinae except Beornia and Hubbardiella (Fig. 40). While these three characters are neither unique to, nor universally found in, Entiinae, there have been no shared characters found for ophelimines and entiines that are not also found in all other eulophid subfamilies. Combining the two would therefore result in a group that is more difficult to characterize morphologically than ...
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... raised doubts over its family and subfamily affinities. All molecular and combined analyses (Figs 43-46) place it with Entiinae. ...
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... only consistently supported clade within Entiinae was the grouping of Beornia + Euderus + Hubbardiella. Although Beornia and Hubbardiella are the only entiines with a fused Mt 8+9 (character 30:1, Fig. 40), Hubbardiella was consistently the sister group to Euderus in molecular and combined analyses. ...
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... raised some doubts concerning placement of the two exceptions. Colpoclypeus was not available for sequencing, but Dicladocerus westwoodi was consistently part of a monophyletic Eulophini in all molecular and combined analyses, between the clades Eulophus + Pnigalio and Elachertus + Euplectrus (Figs 43-46). This suggests that a reversal in propleural form has occurred at least once within Eulophini. ...
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... from Entiinae, the only other eulophid subfamily that was paraphyletic in any molecular analysis was Entedoninae. Likewise, its tribe Euderomphalini was paraphyletic in parsimony analyses (Figs 43 and 44). Entedonini was paraphyletic in all but the two combined analyses (Figs 44 and 46). ...
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... its tribe Euderomphalini was paraphyletic in parsimony analyses (Figs 43 and 44). Entedonini was paraphyletic in all but the two combined analyses (Figs 44 and 46). While the placement of Trisecodes agromyzae had admittedly been controversial (Delvare and LaSalle, 2000), there had never been any doubt concerning the subfamily placement of Closterocerus Westwood. ...
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... analyses (Figs 43 and 44). Entedonini was paraphyletic in all but the two combined analyses (Figs 44 and 46). While the placement of Trisecodes agromyzae had admittedly been controversial (Delvare and LaSalle, 2000), there had never been any doubt concerning the subfamily placement of Closterocerus Westwood. The molecular Bayesian analysis (Fig. 45) placed Closterocerus sensu stricto as the sister group to other Entedoninae + Opheliminae + Entiinae. This is in agreement with the previous analysis by Gauthier et al. (2000), suggesting that these findings are unlikely to be due to sequencing error. Combined analyses (Figs 44 and 46) instead indicated a monophyletic Entedonini with ...
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... is in agreement with the previous analysis by Gauthier et al. (2000), suggesting that these findings are unlikely to be due to sequencing error. Combined analyses (Figs 44 and 46) instead indicated a monophyletic Entedonini with Closterocerus s.s. arising within it. ...
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... et al. (2000), suggesting that these findings are unlikely to be due to sequencing error. Combined analyses (Figs 44 and 46) instead indicated a monophyletic Entedonini with Closterocerus s.s. arising within it. The molecular-only parsimony analysis differed in that Closterocerus s.s. was sister group to all other Entedoninae except Neopomphale (Fig. ...
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... bases, the 3e¢ subregion in the 28S D2 rDNA, from all sequences in the analysis resulted in the placement of Closterocerus s.s. within Entedonini as the sister group to Chrysonotomyia in both molecular-only analyses (Table 3). The 3e¢ subregion in both sampled species of Closterocerus s.s. is very different from that of other entedonines ( Fig. 47) and could be both the defining trait of the subgenus and the reason why molecular results consistently place it far from morphologically similar ...
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... if one is not convinced by the condition in Euderomphale, it is even clearer that Entedononecremnus (Fig. 24), another euderomphaline genus, has no externally indicated notauli. Its more typically-shaped axillae are only weakly advanced anteriorly and extend posteriorly as a long slope towards the metanotum, as in most other chalcidoids. Gumovsky (2002) acknowledged this, n n n 68 n n Closterocerus s.s. + Chrysonotomyia n n n n 71 n Mol = ...
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... anteriorly and extend posteriorly as a long slope towards the metanotum, as in most other chalcidoids. Gumovsky (2002) acknowledged this, n n n 68 n n Closterocerus s.s. + Chrysonotomyia n n n n 71 n Mol = Molecular only (= ⁄ -, with and without 3e¢ region); Comb = combined analysis; y = clade present but without support; n = clade absent. Fig. 47. The 3e and 3e¢ subregions for Closterocerus compared with that of other selected eulophids as aligned by the secondary structure model provided by Gillespie et al. (2005). Species Other Entedonini have the same sequence as Chrysocharis sp. in these subregions. Intervening bases between the two subregions omitted. but maintained that ...
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... our analyses, this grouping does not occur. Instead, Euderomphalini is either sister group to Entedonini (Figs 44 and 46), sister group to Entedonini minus Closterocerus s.s. (Fig. 45), or part of an unresolved clade including Entedonini, Ophliminae, and Entiinae (Fig. 43). ...
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... change in interpretation leaves only clypeal form and 28S D2 data supporting a grouping of Euderomphalini + Entiinae. In our analyses, this grouping does not occur. Instead, Euderomphalini is either sister group to Entedonini (Figs 44 and 46), sister group to Entedonini minus Closterocerus s.s. (Fig. 45), or part of an unresolved clade including Entedonini, Ophliminae, and Entiinae (Fig. 43). While clypeal form in Entiinae and Euderomphalini may be similar in some taxa, the clypeus is not indicated in some species of both groups. This leaves no unambiguous support for Euderomphalini + Entiinae, and therefore it seems best to return ...
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... clypeal form and 28S D2 data supporting a grouping of Euderomphalini + Entiinae. In our analyses, this grouping does not occur. Instead, Euderomphalini is either sister group to Entedonini (Figs 44 and 46), sister group to Entedonini minus Closterocerus s.s. (Fig. 45), or part of an unresolved clade including Entedonini, Ophliminae, and Entiinae (Fig. 43). While clypeal form in Entiinae and Euderomphalini may be similar in some taxa, the clypeus is not indicated in some species of both groups. This leaves no unambiguous support for Euderomphalini + Entiinae, and therefore it seems best to return Euderomphalini to Entedoninae stat. ...
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... tribes of Entedoninae were monophyletic in the combined analyses (Figs 44 and 46). In the molecular parsimony analysis (Fig. 43), Neopomphale was part of an unresolved clade with Entiinae and Opheliminae, and the rest of Euderomphalini rendered Entedonini paraphyletic. ...
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... tribes of Entedoninae were monophyletic in the combined analyses (Figs 44 and 46). In the molecular parsimony analysis (Fig. 43), Neopomphale was part of an unresolved clade with Entiinae and Opheliminae, and the rest of Euderomphalini rendered Entedonini paraphyletic. The molecular-only Bayesian analysis (Fig. 45) indicated a poorly supported but monophyletic Euderomphalini, but Entedonini was paraphyletic because Closterocerus s.s. was sister group to other ...
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... tribes of Entedoninae were monophyletic in the combined analyses (Figs 44 and 46). In the molecular parsimony analysis (Fig. 43), Neopomphale was part of an unresolved clade with Entiinae and Opheliminae, and the rest of Euderomphalini rendered Entedonini paraphyletic. The molecular-only Bayesian analysis (Fig. 45) indicated a poorly supported but monophyletic Euderomphalini, but Entedonini was paraphyletic because Closterocerus s.s. was sister group to other Entedonini + Entiinae + ...
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... combined parsimony and Bayesian analyses (Figs 44 and 46) contained a monophyletic group of Cabeza, Euderomphale and Neopomphale, all of which have a transverse sulcus or sharp carina extending across the vertex between the median and lateral ocelli (character 6:1, Fig. 15). Molecular-only analyses did not reflect this group (Figs 43 and 45). ...
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... Chrysonotomyia within Entedonini (Table 3). The combined analyses (Figs 44-46) also bring Closterocerus s.s. back within Entedonini, but not as sister group to Chrysonotomyia. ...
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... in a sister-group relationship of Closterocerus s.s. + Chrysonotomyia within Entedonini (Table 3). The combined analyses (Figs 44-46) also bring Closterocerus s.s. back within Entedonini, but not as sister group to Chrysonotomyia. No analysis supports inclusion of Asecodes and Neochrysocharis with Closterocerus. Instead, combined Bayesian results (Fig. 46) suggest that they are closely related to a clade of Pediobomyia + Pediobius + Rhynchentedon. A similar relationship between Asecodes, Neochrysocharis, and Pediobius was found independently by Gumovsky (2002) using 28S D2 ...
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... is paraphyletic with respect to Asecodes and some species of Pediobius or other ''core'' entedonines in the molecular-only analyses and the combined parsimony analysis (Figs 43-45); it is monophyletic in the combined Bayesian analysis (Fig. 46). There is no known morphological reason to expect Neochrysocharis to be paraphyletic with respect to Pediobius. ...
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... is paraphyletic with respect to Asecodes and some species of Pediobius or other ''core'' entedonines in the molecular-only analyses and the combined parsimony analysis (Figs 43-45); it is monophyletic in the combined Bayesian analysis (Fig. 46). There is no known morphological reason to expect Neochrysocharis to be paraphyletic with respect to Pediobius. While Neochrysocharis is morphologically similar to Asecodes, they differ in some characters and do not consistently form a clade in molecular analyses. While Neochrysocharis and Asecodes may eventually be combined, they ...
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... (2004) suggested a novel set of characters defining Chrysonotomyia, most importantly the presence of a single spine on the volsellar digitus (character 31:1, Fig. 42) and an at least partially delimited clypeus (character 10:0, Fig. 20). He reclassified some Neotropical and Nearctic species from the subgenus Closterocerus (Achrysocharis) into Chrysonotomyia, based on this character, but other members of the subgenus were not discussed. The European species Closterocerus (Achrysocharis) germanicus ...
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... similar to Omphale in possessing a delimited clypeus (character 10:0, Figs 20 and 21). The combined set of genera from these two lists included in this study are Astichomyiia, Chrysonotomyia, Parzaommomyia, and Tropicharis. None of the analyses in the current study produced a monophyletic group of these genera, but the combined Bayesian analysis (Fig. 46) presents an unsupported clade of Chrysonotomyia + Omphale + Parzaommomyia. The only supported monophyletic relationship between any genera with a delimited clypeus was Omphale + Parzaommomyia, in Bayesian analyses (Figs 45 and 46). Parsimony results (Figs 43 and 44) do not support any monophyletic grouping of entedonine genera with an ...
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... of the analyses in the current study produced a monophyletic group of these genera, but the combined Bayesian analysis (Fig. 46) presents an unsupported clade of Chrysonotomyia + Omphale + Parzaommomyia. The only supported monophyletic relationship between any genera with a delimited clypeus was Omphale + Parzaommomyia, in Bayesian analyses (Figs 45 and 46). Parsimony results (Figs 43 and 44) do not support any monophyletic grouping of entedonine genera with an indicated clypeus. ...
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... only supported monophyletic relationship between any genera with a delimited clypeus was Omphale + Parzaommomyia, in Bayesian analyses (Figs 45 and 46). Parsimony results (Figs 43 and 44) do not support any monophyletic grouping of entedonine genera with an indicated clypeus. ...
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... was consistently placed near the genus Emersonella, forming a clade with it and Ceranisus in the combined Bayesian analysis (Fig. 46) or with Neochrysocharis formosa in the molecular-only parsimony analysis (Fig. 44). It does not actually have a delimited clypeus (Hansson, 2002: Fig. 404). Given that Astichomyiia also possesses a pronotal collar carina (character 12:1), which is absent in Omphale, we suggest that there is no evidence of any close relationship between ...
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... was consistently placed near the genus Emersonella, forming a clade with it and Ceranisus in the combined Bayesian analysis (Fig. 46) or with Neochrysocharis formosa in the molecular-only parsimony analysis (Fig. 44). It does not actually have a delimited clypeus (Hansson, 2002: Fig. 404). Given that Astichomyiia also possesses a pronotal collar carina (character 12:1), which is absent in Omphale, we suggest that there is no evidence of any close relationship between it and ...
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... was consistently placed near the genus Emersonella, forming a clade with it and Ceranisus in the combined Bayesian analysis (Fig. 46) or with Neochrysocharis formosa in the molecular-only parsimony analysis (Fig. 44). It does not actually have a delimited clypeus (Hansson, 2002: Fig. 404). Given that Astichomyiia also possesses a pronotal collar carina (character 12:1), which is absent in Omphale, we suggest that there is no evidence of any close relationship between it and ...
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... was sister group either to most other Entedonini (Figs 44-46), or to most Euderomphalini (Fig. 43). Combined analyses (Figs 44 and 46) suggest it is at the base of a grade including Chrysonotomyia, Closterocerus s.s., Omphale, and Parzaommomyia. ...
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... was sister group either to most other Entedonini (Figs 44-46), or to most Euderomphalini (Fig. 43). Combined analyses (Figs 44 and 46) suggest it is at the base of a grade including Chrysonotomyia, Closterocerus s.s., Omphale, and Parzaommomyia. This seems plausible if one includes a delimited clypeus in the groundplan state for Entedonini, the character being lost multiple times in entedonine evolution. This scenario is supported ...
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... was sister group either to most other Entedonini (Figs 44-46), or to most Euderomphalini (Fig. 43). Combined analyses (Figs 44 and 46) suggest it is at the base of a grade including Chrysonotomyia, Closterocerus s.s., Omphale, and Parzaommomyia. This seems plausible if one includes a delimited clypeus in the groundplan state for Entedonini, the character being lost multiple times in entedonine evolution. ...
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... (2002) No species from the former genus Edovum were included in this analysis, but the others formed a monophyletic and supported group in the Bayesian molecular-only analysis (Fig. 45). They did not form a monophyletic group in parsimony results or in combined Bayesian analysis ( Figs 43, 44 and 46). Parsimony consistently indicated a sister-group relationship between Paracrias and Ceranisus, while Paracrias was in an unresolved clade in combined Bayesian results. Horismenus was monophyletic in both combined ...
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... all analyses, Pediobomyia and Rhynchentedon were sister groups forming a clade with at least one species of Pediobius. The three genera formed a monophyletic group in all analyses (Figs 43-46). Both parsimony analyses indicated a monophyletic Pediobius as sister group to Pediobomyia + Rhynchentedon, but Pediobius was paraphyletic in both Bayesian analyses. ...
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... (2007) listed a set of genera possessing a longitudinal carina on the lateral surface of the pronotum (character 13:1, Fig. 22). Three of these genera were included in this analysis: Achrysocharoides, Entedon, and Pleurotroppopsis. These genera formed a monophyletic group in the combined Bayesian analysis (Fig. 46), but not in any other ...
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... Bayesian results (Figs 45 and 46), these three genera formed a moderately supported clade. With parsimony, Emersonella and Astichomyiia formed a clade in molecular-only results (Fig. 43), but not in combined results (Fig. 44). Ceranisus was consistently the sister group of Paracrias in parsimony results (Figs 43 and 44). Hansson (2002) recognized the morphological similarity between Astichomyiia and Emersonella, but also listed some similarities between Astichomyiia and Closterocerus, a grouping that is not supported ...
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... Bayesian results (Figs 45 and 46), these three genera formed a moderately supported clade. With parsimony, Emersonella and Astichomyiia formed a clade in molecular-only results (Fig. 43), but not in combined results (Fig. 44). Ceranisus was consistently the sister group of Paracrias in parsimony results (Figs 43 and 44). Hansson (2002) recognized the morphological similarity between Astichomyiia and Emersonella, but also listed some similarities between Astichomyiia and Closterocerus, a grouping that is not supported by molecular ...
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... parsimony, Emersonella and Astichomyiia formed a clade in molecular-only results (Fig. 43), but not in combined results (Fig. 44). Ceranisus was consistently the sister group of Paracrias in parsimony results (Figs 43 and 44). Hansson (2002) recognized the morphological similarity between Astichomyiia and Emersonella, but also listed some similarities between Astichomyiia and Closterocerus, a grouping that is not supported by molecular data. ...
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... results present the first published phylogenetic analysis of Eulophidae where Entedoninae has been supported as monophyletic. The phylogenetic hypothesis presented in the combined Bayesian analysis (Fig. 46) presents strongly supported nodes, suggesting answers to some controversies concerning eulophid morphology. The combined parsimony analysis (Fig. 43) suggests an alternative hypothesis differing in important ways that call for further ...
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... results present the first published phylogenetic analysis of Eulophidae where Entedoninae has been supported as monophyletic. The phylogenetic hypothesis presented in the combined Bayesian analysis (Fig. 46) presents strongly supported nodes, suggesting answers to some controversies concerning eulophid morphology. The combined parsimony analysis (Fig. 43) suggests an alternative hypothesis differing in important ways that call for further ...
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... The pronotal shoulder of Entedon and allied groups is bordered by a semicircular ridge (or plica) below (Gumovsky 2002(Gumovsky , 2011Burks et al. 2011;Fig. 2, lpl), whereas the upper sector of the lateral panel of the pronotum bears no plica in most other entedonine genera. ...
... The V-shaped or transverse frontal sutures (frontal sulcus: Graham 1959Graham , 1971; frontal grooves/sutures: Schauff 1988Schauff , 1991; the transverse facial sulcus: Burks et al. 2011;Fig. 5) is found in most of the Entedonini, and is likely one of the morphological synapomorphies for Entedoninae (e.g., Schauff 1991). ...
... The non-sulcate scrobal grooves are conserved across genera and shared by Chrysocharis (Fig. 5D), Achrysocharoides ( Fig. 5E-F), Entedon ( Fig. 6A-D), and Derostenus (Schauff 1988). Entedon and Achrysocharoides have been shown to be closely related (Gumovsky 2002;Burks et al. 2011). The genera Colpixys and Xiphentedon also possess these non-sulcate scrobal grooves ( Fig. 6E-F), which may indicate their affinity to Entedon and Achrysocharoides. ...
The genera Colpixys and Xiphentedon are revised. Both genera are of Afrotropical distribution and are very similar to Entedon, but distinguished by a median strip or furrow on the propodeum replacing the median carina of Entedon. The genus Xiphentedon is characterized by the complete lateral propodeal sulci delimiting convex submedian areas, prosternum with a flange, axillula with indentate projection, and densely hairy central mesopectus. The subgenus Cederholmia Gumovsky, 1997 of Entedon is considered a junior synonym of Xiphentedon, so two of its species are moved to the genus as X. halli (Gumovsky, 1997) comb. nov. and X. danielssoni (Gumovsky, 1997) comb. nov. The genus Colpixys is characterized by the lack of characters of Xiphentedon, but also by the broadly sculptured propodeum with the deep median furrow delimited laterally by two sinuous margins. Three (one new) species are recognized in Colpixys and sixteen (thirteen new) species allocated to three groups (kayovei, danielssoni and forceps), are assigned to Xiphentedon. The new species are: Colpixys eburnus sp. nov. from Ivory Coast and the Republic of South Africa (RSA), Xiphentedon neserorum sp. nov. from RSA, X. simoni sp. nov. from Tanzania, X. dewittei sp. nov. and X. musimba sp. nov. from the Democratic Republic of the Congo (DRC), X. jeanyvesi sp. nov. from Tanzania, the Central African Republic (CAR) and RSA, X. wieringai sp. nov. from Gabon and Ivory Coast, X. kivuensis sp. nov. from DRC, X. palabora sp. nov. from RSA, X. sangha sp. nov. from CAR, X. nimba sp. nov. from Guinea, X. forceps sp. nov. from Ivory Coast, Benin and RSA, X. gerardi sp. nov. from Benin, DRC and RSA, and X. acutigena sp. nov. from Ivory Coast, DRC, Tanzania and RSA. Morphological peculiarities and possible relationships of Colpixys, Xiphentedon, Entedon and some other genera, are discussed.
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New genus Zhijilingia Xu, Liu et He, gen. n. (type species: Zhijilingia tettigonia Xu, Liu et He, sp. n.) is described from Sichuan, South China. This new genus can be distinguished from other genera of Tetrastichinae by following characters: antenna with 3 anelli and 4 funicular segments, only one clava segment, all funicular segments slender and longer than broad, funicular segments almost same in length, mesoscutum without a median line, propodeum with a distinct median carina, the whole rim of propodeal spiracle exposed, postmarginal vein absent. The new species probably parasitize eggs of the katydids Prohimerta dispar (Bey-Bienko, 1951) (Orthoptera: Tettigoniidae).
... Gumovsky (2001 synonymized Closterocerus, Hispinocharis Bouček, 1988, Mangocharis Bouček 1986, Neochrysocharis and Asecodes, based on their morphological similarity, in particular their possession of subtorular grooves. Burks et al. (2011) demonstrated a basal relationship of Closterocerus with the clade comprising Asecodes + Neochrysocharis, and suggested the potential value of the form of the basiconic peg sensilla for informing the phylogenetic position of the genera. Afronympha is close to Asecodes in terms of the general habitus, the subtorular grooves, the occiput with median furrow (though weak), the short postmarginal and stigmal veins, and presence of the anterior groove on the propodeum. ...
... However, A. eminpashai cannot be classified as a derived species of Asecodes due to the possession of the following character states: (1) scrobal grooves meet below the transverse facial sulcus (ending separately in Asecodes); (2) flagellum with distinct subdivision into 3-segmented funicle and 2-segmented clava (unlike Asecodes where the subdivision is ambiguous, since the funicular and claval segments are rather similar in shape and length); (3) flagellum with short and symmetric sensilla ampullacea (type I sensu Hansson 2002, basiconic capitate peg sensilla sensu Yang et al. 2018) unlike the L-shaped (type 2 sensu Hansson 2002) sensilla ampullacea in Asecodes. Similar sensilla ampullacea (type I) are peculiar to the genus Neochrysocharis, which is also a sister group to Asecodes (Burks et al. 2011). Neochrysocharis is clearly different from Afronympha (apart from the autapomorphies of the genus) in having a clavate antenna with 2-segmented funicle and 3-segmented clava, all flagellar segments being generally transverse. ...
Afronympha gen. nov. is described for the single known species A. eminpashai sp. nov. collected in the Democratic Republic of the Congo and Uganda. This new genus is similar to some extent to Asecodes, but differs from this and other genera of Entedoninae in having a produced and medially notched anterior margin of the clypeus, and robust mandibles. Similarities and differences between Afronympha and other genera are discussed. The type species, A. eminpashai sp. nov., was collected in degraded mid-altitude rainforest in Kibale National Park, western Uganda and disturbed areas of lowland rainforest in the Ituri region of eastern Democratic Republic of the Congo, suggesting its ecological plasticity under circumstances of habitat transformation.
... The character matrix can be accessed on Dryad Data Repository (DOI https://doi.org/10.6086/D19098). Some features that could not be assessed on the fossil were chosen to ensure that subfamily monophyly resembled that of recent phylogenies (Gauthier et al. 2000;Burks et al. 2011;Gumovsky 2011;Heraty et al. 2013). ...
A new fossil species of Eulophidae, Kressleinius celans , gen. et sp. nov. , is described from Eocene Baltic amber. It does not place into any extant genera within Eulophidae due to a lack of distinctive synapomorphies. The results of a morphology-based phylogenetic analysis placed K. celans in Tetrastichinae because of features shared with several genera, including having 6 flagellomeres past the anelli, admarginal setae short, propleura diverging, and a newly described feature of the transepimeral sulcus. This represents the oldest known fossil for the largest family of Chalcidoidea in terms of described species.
... Our phylogenetic results support two major clades within Eulophidae: a clade formed by Tetrastichinae + Eulophinae and a clade containing Entedoninae sister to Entiinae + Opheliminae. This result corroborates the hypothesis of relationships among subfamilies proposed by Burks et al. (2011) and Munro et al. (2011). ...
... Within Entedoninae, Euderomphalini (parasitoids of whiteflies) is recovered sister to Entedonini, a placement corroborated by the results from Burks et al. (2011). The only species of the genus Closterocerus subgenus Closterocerus included in this study was recovered sister to all other Entedonini, a result also similar to Burks et al. (2011), yet with higher support. ...
... Within Entedoninae, Euderomphalini (parasitoids of whiteflies) is recovered sister to Entedonini, a placement corroborated by the results from Burks et al. (2011). The only species of the genus Closterocerus subgenus Closterocerus included in this study was recovered sister to all other Entedonini, a result also similar to Burks et al. (2011), yet with higher support. The second species of Closterocerus, belonging to the subgenus Achrysocharis, was nested within Entedonini and recovered sister to Horismenus with high statistical support in ML analyses. ...
Eulophidae is a hyper-diverse family of chalcidoid wasps with 324 genera, about 5300 described species and probably thousands of others to be described. Until now, the absence of unequivocal morphological apomorphies and the low resolution provided by the handful of Sanger sequenced genes have hampered the reconstruction of phylogenetic relationships within the family. Here we used Ultra-Conserved Elements (UCEs) and their flanking regions to resolve relationships among 84 species of eulophids included in 63 genera representing all subfamilies and most tribes, plus 15 outgroups. Our analyses recover all traditional Eulophidae subfamilies and tribes with high support and globally agree with the traditional classification of the family. Our results confirm that Eulophinae + Tetrastichinae is the sister group of (Opheliminae + Entiinae) + Entedoninae. At the generic level, our analyses provide high support for intergeneric relationships for which morphology and Sanger markers previously failed to provide resolution. Our results also confirm that Trisecodes does not group with Eulophidae and may not belong to this family; however, its correct classification still awaits a large-scale phylogenomic hypothesis for Chalcidoidea. This work opens new avenues towards a better understanding of the evolutionary history, biogeography and evolution of host-parasitoid associations in this hyper-diverse family of chalcidoid wasps.
... The description of the new species is based on adult females and males that emerged from the breeding chamber. Terminology of the morphological structures followed Protasov et al. [6], Burks et al. [10], and Gibson et al. [36] As a measurement and observation instrument, a trinocular flat chromatic stereo mic magnifying glass, (BEL model, Solaris-T-Led) and a Microscope (OPTIKAL B-1000PH) were used (OPTIKA, Microscopes, Via Rigia, 32, 24010, Ponteranica, BG, Italy). For the photographs and measurements, the program OPTIKALS (view version 3.9.0.602) was used. ...
... A comparative table was prepared for a comparison of the morphological characteristics presented by the individuals from the samples collected in the field, and those described for O. maskelli by Burks et al., [10] (Table S1). ...
... Additional DNA sequences from generous Ophelimus and outgroup members were downloaded from GenBank, [40] and BoldSystem. This sampling was complemented with the inclusion of COI sequences belonging to the Entiineae subtribes: Astichus, Bellerus, Beornia and Euderus [10]. Hubbardiella did not have COI sequences at the time of this sampling. ...
In 2003, a new gall-inducing wasp of the genus Ophelimus was detected in the Valparaíso Region (Chile), affecting tree plantations of Eucalyptus globulus Labill and Eucalyptus camaldulensis Dehnh. Since then Ophelimus has been frequently detected in different plantations in Chile, covering a widespread area. A preliminary collaborative study suggests that the micro-wasp detected should be classified as a new Ophelimus species. In this paper, using an integrative approach (including genetic, morphological, and behavioral data), we addressed the delimitation and description of this new species. This study involved the use of brood adult specimens, raised at the laboratory of MIPlagas Ltda., from infested twigs of E. globulus collected in several localities between of Valparaíso and Los Lagos Regions (Chile). Morphological structures were described according to current Eulophidae taxonomic keys, as well as additional traits, such as gall morphology and behavior. Genetic characterization was implemented using a phylogenetic approach, based on a 648 bp specific fragment of the mitochondrial Cytochrome Oxidase I gene (COI 5 region) obtained from collected specimens and available databases (Genbank, NCBI, and BOLDSystem). Specifically, distinctive patterns of variation were detected in traits like gall and antennae morphology, growth habit trends, and a notorious polyphenism in the setae from the sub marginal vein. Overall evidence suggests that this new entity should be considered a new species in Ophelimus, which is henceforth named Ophelimus migdanorum Molina-Mercader.
... 5A-5B) was reared exclusively from wild olives in Paarl (n = 23), and represented the lowest proportion (3.6%) of the total chalcids (Table S4 2 ). A previous phylogeny of Eulophidae based on morphological and molecular markers (COI and 28S rRNA D2-D5) showed that N. formosus and N. clinias were a paraphyletic group with respect to Asecodes sp., although the study included a single sequence for each Neochrysocharis species from Italy (Burks et al. 2011). Due to the poor sequence coverage of the genus Neochrysocharis, public Asecodes sequences with high quality (A. ...
... Neochrysocharis (N. formosus HM365028 and N. clinias HM365038) was previously shown to be, with respect to Asecodes, paraphyletic in molecular analyses, paraphyletic in combined (molecular and morphological) parsimony analysis, and monophyletic in combined Bayesian analysis (Burks et al. 2011). The inclusion of the N. formosus Nf21 and N. formosus Nf18 sequences recovered the paraphyly of the genus Neochrysocharis with regards to Asecodes. ...
Wild and cultivated olives harbor and share a diversity of insects, some of which are considered agricultural pests, such as the olive fruit fly. The assemblage of olive-associated parasitoids and seed wasps is rich and specialized in sub-Saharan Africa, with native species possibly coevolving with their hosts. Although historical entomological surveys reported on the diversity of olive wasp species in the Western Cape Province of South Africa, no comprehensive study has been performed in the region in the molecular era. In this study, a dual approach combining morphological and DNA-based methods was used for the identification of adult specimens reared from olive fruits. Four species of Braconidae and six species of Chalcidoidea were morphologically identified, and DNA barcoding methodologies were used to investigate conspecificity among individuals, based on randomly selected representative specimens. Morphological identifications were congruent with DNA data, as NJ and ML trees correctly placed the sequences for each species either at the genus or species level, depending on the available taxa coverage, and genetic distances strongly support conspecificity. No clear evidence of cryptic diversity was found. Overall seed infestation and parasitism rates were higher in wild olives compared to cultivated olives, and highest for Eupelmus spermophilus and Utetes africanus . These results can be used for early DNA-based detection of wasp larvae in olives, and investigating the biology and ecology of these species.
... The close relationships of Pediobius with Rhynchentedon and Pediobomyia were demonstrated by Burks et al. (2011) using molecular markers. The genera were always closely related in all phylogenetic analyses provided by the authors, and at least in the Bayesian molecular-only results and in the combined morphological and molecular results, Rhynchentedon + Pediobomyia appeared as a derived node within Pediobius with high bootstrap support (Burks et al. 2011). ...
... The close relationships of Pediobius with Rhynchentedon and Pediobomyia were demonstrated by Burks et al. (2011) using molecular markers. The genera were always closely related in all phylogenetic analyses provided by the authors, and at least in the Bayesian molecular-only results and in the combined morphological and molecular results, Rhynchentedon + Pediobomyia appeared as a derived node within Pediobius with high bootstrap support (Burks et al. 2011). So, based on original morphological data and in part on results of molecular research of other authors, the generic names Rhynchentedon Girault and Pediobomyia Girault are hereby considered junior synonyms of Pediobius Walker (syn. ...
Three new species groups and seven species of the genus Pediobius Walker are described from the Afrotropical realm: the marjoriae group, with P. marjoriae Kerrich (described from Uganda), P. rohombaya Gumovsky sp. n. (from the Central African Republic (CAR), the Democratic Republic of the Congo (DRC), Gabon, Uganda) and P. orungu Gumovsky sp. n. (from Gabon); the afroteres group with P. afroteres Gumovsky sp. n. (from South Africa, Zimbabwe, Malawi, Kenya, Uganda, DRC, Cameroon) and P. kafroteres Gumovsky sp. n. (from Cameroon); the askari group with P. askari Gumovsky sp. n. (from South Africa, Zimbabwe, Kenya, Tanzania, DRC, Ivory Coast) and P. maleficus Gumovsky sp. n. (from Kenya, DRC, CAR, Cameroon, Gabon); and P. nganga Gumovsky sp. n. (from DRC) (not assigned to any group). All these species are characterized by the relatively wide and robust head with elongate and/or narrowed lower face and the antennae attached near or below the lower eye margins. Morphological features of the species, as well as their habitat distribution, are discussed. The comparison of the new species with similar taxa of Entedoninae suggested the following synonymies: Rhynchentedon Girault and Pediobomyia Girault under Pediobius (syn. n.); Bomyiabius frontus Narendran, Pediobomyia budaicus Narendran and Pediobomyia lankicus Narendran under Pediobomyia darwini Girault (syn. n.). The following new combinations are proposed: Pediobius maximus (Girault), P. achterbergi (Gumovsky), P. narendrani (Gumovsky), P. brevicaulis (Hansson), P. canaliculatus (Hansson) and P. darwini (Girault) (comb. n.). Ant parasitoids P. marjoriae and Myrmokata diparoides Bouček are recorded from DRC for the first time.
