Maxillolabial complex in scelionids (Hymenoptera: Platygastroidea): morphology and phylogenetic implications

in Insect Systematics & Evolution
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The Platygastroidea is a highly diverse group of small to minute parasitoid wasps. Despite the reduced size, the morphology of the maxillolabial complex of scelionids is very diverse and phylogenetically informative. 81 characters are scored for 129 genera (representing 75% of the total number of known extant genera of scelionids), as well as for seven outgroup taxa. All taxa examined are illustrated with images, SEM micrographs and/or line drawings. Phylogenetic trees resulting from analyses conducted in TNT under various settings were not fully resolved, but some relationships were repeatedly retrieved. The Platygastroidea are usually corroborated. Nixoniini, Sparasionini, Plaumannion and Huddlestonium are frequently retrieved as the most basal platygastroid lineages. Psix and Paratelenomus often form a monophyletic group close to Gryonini. The Scelioninae, Teleasinae and Telenominae are not supported as monophyletic. However, some major scelionid clades are frequently retrieved, but these are not recognized in the current classification of Platygastroidea. The evolution of the palpal formula, highly variably in scelionids, and previously used in platygastroid systematics, is explored. The number of maxillary palpomeres in the ground plan of platygatroids is probably five, that of labial palpomeres two or three, given the variation among basal scelionids. Our study provides relevant information for resolving the phylogeny of the Platygastroidea, but additional character sources have to be explored to obtain a robust phylogenetic hypothesis.

Maxillolabial complex in scelionids (Hymenoptera: Platygastroidea): morphology and phylogenetic implications

in Insect Systematics & Evolution

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Figures

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    Diversity of the shape of cardo among the genera of scelionids: (13) Sparasion; (14) Sceliomorpha; (15) Nixonia; (16) Neuroscelio; (17) Apegus; (18) Platyscelio; (19) Cremastobaeus; (20) Maruzza; (21) Gryon. This figure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/journals/1876312X.

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    Stipes in (22) Sparasion; (23) Sceliomorpha; (24) Archaeoteleia; (25) Baryconus; (26) Mantibaria; (27) Odontoscelio; (28) Platyscelio; (29) Neuroscelio; (30) Endecascelio. This figure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/journals/1876312X.

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    Stipes in (31) Scelio; (32) Nixonia; (33) Maruzza; (34) Acanthoscelio; (35) Breviscelio; (36) Doddiella. This figure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/journals/1876312X.

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    MLC in (155) Archaeoteleia (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (156) Plaumannion (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (157) Huddlestonium (a,b, galeolacinial complex: a, dorsal view; b, ventral view). Scale bar = 0.1 mm.

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    MLC in (158) Acanthoscelio (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (159) Axea (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (160) Heptascelio (a,b, galeolacinial complex: a, dorsal view; b, ventral view). Scale bar = 0.1 mm.

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    MLC in (170) Chromoteleia (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (171) Oxyscelio (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (172) Anthonyon (a,b, galeolacinial complex: a, dorsal view; b, ventral view). Scale bar = 0.1 mm.

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    MLC in (227) Mantibaria (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (228) Platyscelio (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (229) Doddiella (a,b, galeolacinial complex: a, dorsal view; b, ventral view). Scale bar = 0.1 mm. This figure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/journals/1876312X.

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    MLC in (152) Nixonia (a,b, galeolacinial complex: a, dorsal view; b, ventral view); 153. Sparasion (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (154) Sceliomorpha (a,b, galeolacinial complex: a, dorsal view; b, ventral view). Scale bar: 152, 154 = 0.1 mm, 153 = 0.5 mm.

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    MLC in (257) Microthoron (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (258) Pseudanteris (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (259) Tanaodytes (a,b, galeolacinial complex: a, dorsal view; b, ventral view). Scale bar = 0.1 mm.

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    MLC in (167) Apegus (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (168) Baryconus (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (169) Bracalba (a,b, galeolacinial complex: a, dorsal view; b, ventral view). Scale bar = 0.1 mm.

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    MLC in (173–Amblyscelio sp. 1 (a,b, galeolacinial complex: a, dorsal view; b, ventral view), 174–Amblyscelio sp. 2 (a,b, galeolacinial complex: a, dorsal view; b, ventral view), 175–Calliscelio (a,b, galeolacinial complex: a, dorsal view; b, ventral view). Scale bar = 0.1 mm.

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    MLC in (218) Styloteleia (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (219) genus T (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (220) Trichoteleia (a,b, galeolacinial complex: a, dorsal view; b, ventral view). Scale bar = 0.1 mm.

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    MLC in (221) Opisthacantha (Trissoscelio) (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (222) Opisthacantha (Vardhana) (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (223) Tyrannoscelio (a,b, galeolacinial complex: a, dorsal view; b, ventral view; c, prementum; d,e, stipes and maxillary palp). Scale bar = 0.1 mm.

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    MLC in (269) Baeus (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (270) Neobaeus (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (271) Ceratoteleas (a,b, galeolacinial complex: a, dorsal view; b, ventral view). Scale bar = 0.1 mm.

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    MLC in (161) Oreiscelio (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (162) Pseudoheptascelio (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (163) Scelio, Neotropical species (a,b, galeolacinial complex: a, dorsal view; b, ventral view). Scale bar = 0.1 mm.

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    MLC in (164) Scelio (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (165) Scelio (Lepidoscelio) (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (166) Synoditella (a,b, galeolacinial complex: a, dorsal view; b, ventral view). Scale bar = 0.1 mm.

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    MLC in (224) Aradophagus (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (225) Janzenella (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (226) Cremastobaeus (a,b, galeolacinial complex: a, dorsal view; b, ventral view). Scale bar = 0.1mm. This figure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/journals/1876312X.

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    Galeolacinial complex and base of maxillary palp in (37) Glyphomerus; (38) Pteromalus; (39) Pelecinus; (40) Ropronia; (41) Vanhornia; (42) Andricus; (43) Ibalia; (44) Belyta; (45) Sparasion.

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    Maxillary palp in (46) Sparasion sp.1; (47) Sparasion sp.2; (48) Sceliomorpha; (49) Dichoteleas; (50) Neoscelio; (51) Nyleta; (52) Doddiella; (53) Acolomorpha; (54) Opisthacantha; (55) Baeus; (56) Neuroscelio; (57) Scelio sp.1; (58) Scelio sp.2; (59) Teleas. This figure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/journals/1876312X.

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    MLC in (197) Calotelea (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (198) Chakra (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (199) Duarina (a,b, galeolacinial complex: a, dorsal view; b, ventral view). Scale bar = 0.1 mm.

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    MLC in (212) Nyleta (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (213) Opisthacantha (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (214) Oxyteleia (a,b, galeolacinial complex: a, dorsal view; b, ventral view). Scale bar = 0.1 mm.

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    MLC in (230) Neoscelio (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (231) Parascelio (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (232) Sceliacanthella (a,b, galeolacinial complex: a, dorsal view; b, ventral view). Scale bar = 0.1 mm.

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    MLC in (254) Palaeogryon (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (255) Acolomorpha (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (256) genus Q (a,b, galeolacinial complex: a, dorsal view; b, ventral view). Scale bar = 0.1 mm.

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    MLC in (281) Gryonoides 1 (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (282) Xenomerus (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (283) Eumicrosoma (a,b, galeolacinial complex: a, dorsal view; b, ventral view). Scale bar = 0.1 mm.

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    Galeolacinial complex in scelionids: (69) Nixonia; (70) Archaeoteleia; (71) Sceliomorpha; (72) Lapitha; (73) Dichoteleas; (74) genus G.; (75) Sparasion; (76) Trissoscelio; (77) Doddiela; (78) Orwellium; (79) Dicroscelio; (80) Gryonoides; (81) Baeus; (82) Trimorus; (83) Xenomerus; (84) Teleas. This figure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/journals/1876312X.

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    Galeolacinial complex in outgroup taxa: (85) Andricus; (86) Ibalia; (87) Belyta; (88) Glyphomerus; (89) Ropronia; (90) Proctotrupes; (91) Helorus; (92) Pelecinus; (93) Vanhornia; (94) Pteromalus.

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    MLC in (239) Gryon (Eremioscelio) (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (240) Gryon (Exon) (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (241) Gryon 1 (a,b, galeolacinial complex: a, dorsal view; b, ventral view). Scale bar = 0.1 mm.

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    MLC in (284) Paratelenomus (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (285) Phanuromyia (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (286) Phanuropsis (a,b, galeolacinial complex: a, dorsal view; b, ventral view). Scale bar = 0.1 mm.

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    MLC in (266) Hickmanella (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (267) Idris (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (268) Odontacolus (a,b, galeolacinial complex: a, dorsal view; b, ventral view). Scale bar = 0.1 mm.

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    MLC in (275) Odontoscelio (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (276) Teleas (Proteleas) (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (277) Teleas (a,b, galeolacinial complex: a, dorsal view; b, ventral view). Scale bar = 0.1 mm.

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    MLC in (278) Trimorus 1 (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (279) Trimorus 2 (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (280) Trissacantha (a,b, galeolacinial complex: a, dorsal view; b, ventral view). Scale bar = 0.1 mm.

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    Galeolacinial complex, laterodorsal view in (95) Archaeoteleia; (96) Sceliomorpha; (97) Axea; (98) genus G (with white dots, visible through transparency); (99). Scelio; (100) Bracalba; (101) Cremastobaeus; (102) Doddiella; (103) Gryon. This figure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/journals/1876312X.

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    Galeolacinial complex, latero-dorsal view in (104) Baeus; (105) Gryonella; (106) Gryonoides; (107) Teleas; (108) Xenomerus; (109) Mantibaria; (110) Orwellium; (111) Psix.

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    Prementum in (121) Archaeoteleia; (122) Nixonia; (123) Sparasion; (124) Sceliomorpha; (125) Parascelio; 126; (127) Habroteleia; (128) Chromoteleia; (129) Doddiella; (130) Acolomorpha; (131) Encyrtoscelio. This figure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/journals/1876312X.

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    MLC in (179) Habroteleia (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (180) Holoteleia (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (181) Macroteleia (a,b, galeolacinial complex: a, dorsal view; b, ventral view). Scale bar = 0.1 mm.

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    MLC in (182) Oethecoctonus (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (183) Palpoteleia (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (184) Paridris (a,b, galeolacinial complex: a, dorsal view; b, ventral view). Scale bar = 0.1 mm.

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    MLC in (188) Triteleia (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (189) Triteleia (Parapegus) (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (190) genus A (a,b, galeolacinial complex: a, dorsal view; b, ventral view). Scale bar = 0.1 mm.

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    MLC in (191) genus B (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (192) Crama (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (193) Lispoteleia (a,b, galeolacinial complex: a, dorsal view; b, ventral view). Scale bar = 0.1 mm.

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    MLC in (215) Psilanteris (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (216) Shreemana (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (217) Spiniteleia (a,b, galeolacinial complex: a, dorsal view; b, ventral view). Scale bar = 0.1 mm.

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    MLC in (194) Xentor (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (195) Yunkara (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (196) Anteris (a,b, galeolacinial complex: a, dorsal view; b, ventral view). Scale bar = 0.1 mm.

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    MLC in (206) Leptoteleia sp. 2 (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (207) Lidgbirdius (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (208) Mallateleia (a,b, galeolacinial complex: a, dorsal view; b, ventral view). Scale bar = 0.1 mm.

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    MLC in (209) Masnerella (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (210) Mecix (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (211) Merriwa (a,b, galeolacinial complex: a, dorsal view; b, ventral view). Scale bar = 0.1 mm.

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    Prementum in (132) Cremastobaeus; (133) Mantibaria; (134) Dyscritobaeus; (135) Trimorus; (136) Odontacolus; (137) Platyscelio; (138) Eumicrosoma. This figure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/journals/1876312X.

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    Prementum in (139, 140) Andricus; (141) Belyta; (142, 143) Ibalia; (144) Glyphomerus; (145) Helorus; (146) Pelecinus; (147) Ropronia. This figure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/journals/1876312X.

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    Cardo and stipes in (2) Andricus (Cynipoidea); (3) Brachymeria (Chalcididae: Chalcidoidea); (4) Codrus (Proctotrupidae: Proctotrupoidea); (5) Helorus (Heloridae: Proctotrupoidea); (6) Ibalia (Cynipoidea); (7) Pelecinus (Pelecinidae: Proctotrupoidea); (8) Proctotrupes (Proctotrupidae: Proctotrupoidea); (9) Ropronia (Roproniidae: Proctotrupoidea); (10) Vanhornia (Vanhorniidae: Proctotrupoidea). This figure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/journals/1876312X.

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    The cardo/stipes ratio in scelionids.

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    The correlation between length of cardo and length of stipes. This figure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/journals/1876312X.

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    The correlation between length of cardo and length of maxillary palp. This figure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/journals/1876312X.

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    MLC in (242) Gryon 2 (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (243) Gryon (Plesiobaeus) (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (244) Maruzza (a,b, galeolacinial complex: a, dorsal view; b, ventral view). Scale bar = 0.1 mm.

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    MLC in (233) Apteroscelio (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (234) Breviscelio (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (235) Encyrtoscelio 1 (a,b, galeolacinial complex: a, dorsal view; b, ventral view). Scale bar = 0.1 mm.

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    MLC in (236) Encyrtoscelio 2 (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (237) Encyrtoscelio 3 (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (238) Epigryon (a,b, galeolacinial complex: a, dorsal view; b, ventral view). Scale bar = 0.1 mm.

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    MLC in (251) Endecascelio (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (252) Mirobaeoides (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (253) Mirobaeus (a,b, galeolacinial complex: a, dorsal view; b, ventral view). Scale bar = 0.1 mm.

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    MLC in (245) Neuroscelio (a,b: galeolacinial complex: a, dorsal view; b, ventral view); (246) Orwellium (a,b: galeolacinial complex: a, dorsal view; b, ventral view); (247) Titta (a,b, galeolacinial complex: a, dorsal view; b, ventral view). Scale bar = 0.1 mm.

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    MLC in (185) Phoenoteleia (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (186) Probaryconus (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (187) Romilius (a,b, galeolacinial complex: a, dorsal view; b, ventral view). Scale bar = 0.1 mm.

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    Maxillary palp in (60) Pelecinus; (61) Proctotrupes; (62) Ropronia; (63) Vanhornia; (64) Paramesius; (65) Dendrocerus; (66) Belyta; (67) Andricus; (68) Ibalia. This figure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/journals/1876312X.

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    MLC in (272) Dvivarnus (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (273) Echinoteleas (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (274) Gryonella (a,b, galeolacinial complex: a, dorsal view; b, ventral view). Scale bar = 0.1 mm.

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    MLC in (176) Dichoteleas (a,b, galeolacinial complex: a, dorsal view; b, ventral view), 177–Dicroscelio sp. 1 (a,b, galeolacinial complex: a, dorsal view; b, ventral view), 178–Dicroscelio sp. 2 (a,b, galeolacinial complex: a, dorsal view; b, ventral view). Scale bar = 0.1 mm.

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    MLC in (248) Echthrodesis (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (249) Embidobia (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (250) Embioctonus (a,b, galeolacinial complex: a, dorsal view; b, ventral view). Scale bar = 0.1 mm.

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    Galeolacinial complex, latero-dorsal view in (112) Andricus; (113) Ibalia; (114) Glyphomerus; (115) Belyta; (116) Helorus; (117) Proctotrupes; (118) Pelecinus; (119) Ropronia; (120) Vanhornia.

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    Prementum in (148) Proctotrupes; (149) Vanhornia; (150) Pteromalus.

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    MP tree of 1068 steps based on MLC data under NT with all characters unordered and equally weighted. Palpal formula (numbers at nodes) mapped on tree.

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    MLC in (290) Telenomus (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (291) Trissolcus (a,b, galeolacinial complex: a, dorsal view; b, ventral view). Scale bar = 0.1 mm.

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    MLC in (263) Tiphodytes (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (264) genus X (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (265) Dyscritobaeus (a,b, galeolacinial complex: a, dorsal view; b, ventral view). Scale bar = 0.1 mm.

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    MLC in (260) Thoron (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (261) Thoronella (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (262) Thoronidea (a,b, galeolacinial complex: a, dorsal view; b, ventral view). Scale bar = 0.1 mm.

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    MLC in (287) Telenomus (Platytelenomus) (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (288) Protelenomus (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (289) Psix (a,b, galeolacinial complex: a, dorsal view; b, ventral view). Scale bar = 0.1 mm.

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    MLC in (200) Duta (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (201) Fusicornia (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (202) Genatropis (a,b, galeolacinial complex: a, dorsal view; b, ventral view). Scale bar = 0.1 mm.

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    MLC in (203) genus G (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (204) Opisthacantha (Lapitha) (a,b, galeolacinial complex: a, dorsal view; b, ventral view); (205) Leptoteleia sp. 1 (a,b, galeolacinial complex: a, dorsal view; b, ventral view). Scale bar = 0.1 mm.

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