EURYTOMIDAE

Lotfalizadeh, Hosseinali, Delvare, Gérard & Rasplus, Jean-Yves, 2007, Phylogenetic analysis of Eurytominae (Chalcidoidea: Eurytomidae) based on morphological characters, Zoological Journal of the Linnean Society 151 (3), pp. 441-510 : 461-464

MONOPHYLY OF EURYTOMIDAE

In all results the family is polyphyletic, as Hockeria (illustrating the Chalcididae ) is the sister group of Heimbra (illustrating the Heimbrinae ) with high bootstrap support (= 91–95). Such a relationship was also obtained in a study dealing with the phylogeny of Chalcididae ( Delvare, 2004) . Altogether this relationship is corroborated by numerous derived states, seven of which are synapomorphies: (1) structure and articulation of the mandible (see Wijesekara, 1997 for details); (2) pronotum with posterolateral projection ( Fig. 143 ); (3) parascutal and axillar carinae forming an arch; (4) mesopleuron with differentiated adscrobal area (homoplastic); (5) epicnemium completely delimited (homoplastic); (6) epicnemium with a median crest; (7) mesopleuron with a ventral shelf (homoplastic); (8) mesofurcal pits reduced or evanescent ( Fig. 143 ; (9) metafurcal pits absent through secondary reduction ( Fig. 143 ); (10) discrimen lamella quite high (a component of the internal skeleton within metapectus; see Krogmann, 2005, for details).

In cladograms 3 and 4, Rileya is placed as the sister group of ( Hockeria + Heimbra ). The support is moderate (76–80) and one putative synapomorphy is the short prepectus (more reduced in Chalcididae ), both on its lateral panel and its ventral part ( Fig. 151 ).

In the same cladograms ( Eurytomidae + Hockeria ) are monophyletic. This weakly supported relationship (bootstrap = 67) is, however, sustained by several synapomorphies: (1) mesothoracic spiracle hidden by a posterolateral flange of the pronotum ( Figs 133–138 ); (2) mesocoxal cavities closed posteriorly ( Figs 143, 144 ); (3) metapleuron at least partially fused with propodeum ( Fig. 142 ); (4) metapleuron with a ventral shelf ( Figs 157–159 ); (5) petiole completely closed and tube-like ( Figs 187–193 ). The first character is also shared by Eulophidae Entedoninae ( Gumovsky, 2002) , and the last ones are probably found within a number of chalcidoid families, e.g. the Podagrionini have a ventral shelf ( Grissell, 1995), and the species with petiolate gaster also share a tube-like petiole. Moreover, the mesothoracic spiracle is exposed in the Cratocentrini ( Chalcididae ): either a possible reversion in that tribe or an independent evolution in both groups? Further evidence for a close relationship between these families is the special structure of the syntergum, shared by the Rileyinae s.l. and several taxa in Chalcididae (Cratocentrini, Brachymeria , and Haltichellinae ): the segment has a transverse carina in front of the cercal plates, which are situated within foveae; this carina might represent a trace of the fused sclerites that constitute the syntergum. The Eurytomidae as presently understood therefore appear paraphyletic relative to the Chalcididae , and may even be polyphyletic. This corroborates recent results reported by Gates (2005, 2007) in which Rileyinae are polyphyletic and consist of two different lineages: (1) the Rileyinae s.s., the limits of which were redefined by the same author; (2) the genera Macrorileya , Archirileya , and Buresium included in what we called the Macrorileya genus group.

The Macrorileya genus group is monophyletic in cladograms 2, 3, and 4 (bootstrap = 63–69). It is corroborated by the very small metafurcal pits that are only visible at strong magnification (> ×1000) ( Fig. 167 ). The sister-group relationship for the Macrorileya genus group + Eurytominae is sustained by another feature of the metafurcal pits that are slightly to distinctly moved backwards ( Figs 160– 164 ), whereas they are placed along the anteroventral margin of the metapleuron in most chalcidoids ( Krogmann, 2005). One of us (JYR) recently found that similar pits occurred in all Sycophaginae he examined (an unplaced family of figwasps, see Rasplus et al., 1998), but they are closer to each other and their orientation is oblique (vertical in Eurytomidae ).

Figures

Figures 139–150. Figures 139, 140: Mesopleuron in lateral view – 139, Bruchophagus squamea; 140, Eurytoma compressa. Figure 141: scutellum in lateral view; Eurytoma Peru 1. Figure 142: propodeum in lateral view; 142, Eurytoma dentata. Figures 143, 144, 148: meso- and metapleuron in ventral view – 143, Heimbra opaca; 144, Aximogastra Guinea; 148, Prodecatomidea Cameroon. Figures 146, 147: mesopleuron in anterior view; 146, Syceurytoma ficus; 147, Bephratelloides pomorum. Figures 145, 149, 150: prepectus in ventral view; 145, Tetramesa romana; 149, Bruchophagus Alicante; 150, Phylloxeroxenus San Alberto 2.

Figures 151–159. Figures 151, 152, 154, 157, 158: meso- and metapleuron in ventral view – 151, Rileya pulchra; 152, Bruchophagus Guyoniana; 154, Bruchophagus phlei; 157, Eurytoma lepidopterae; 158, Tenuipetiolus Guadeloupe. Figures 153 and 156: mesopleuron in ventral view – 153, Ficomila Gabon; 156, Eurytoma Senegal. Figure 155: mesopleuron in lateroventral view; Bruchophagus caucasicus. Figure 159: mesosoma in lateroventral view; Eurytoma San Alberto 1.

Figures 127–138. Figures 127–131: propodeum – 127, Sycophila biguttata; 128, Eurytoma ochraceipes; 129, Bruchophagus roddi; 130, Eurytoma collaris; 131, Conoaxima affinis. Figure 132: mesosoma in lateral view; Bephratoides Shushufindi. Figure 133: Mesosoma, anterior part in lateral view; Aximopsis Colombia 1. Figure 134: Pronotum and prepectus in lateral view; Eurytoma braconidis. Figures 135–138: mesopleuron in lateral view; 135, Eurytoma dentata; 136, Eurytoma Cébazan; 137, Plutarchia bicariniventris; 138, Eurytoma braconidis.

Figures 187–197. Figures 187–192: petiole and base of gaster in lateral view; 187, Rileya pulchra; 188, Eurytominae Ecuador; 189, Tetramesa fulvicollis; 190, Eurytoma cressoni; 191, Phylloxeroxenus San Alberto; 192, Axima brevicornis. Figure 193: petiole in ventral view – Eurytoma nodularis. Figure 194: base of gaster in frontal view – Bruchophagus squamea. Figure 195: petiole and median part of first gastral tergite in frontal view – Eurytoma volkovi. Figure 196: petiole and gaster – Axima brevicornis. Figure 197: syntergum in lateral view – Eurytoma amygdali.

Figures 160–171. Figures 160–164: meso- and metapleuron in ventral view – 160, Prodecatoma maculiventris; 161, Eurytoma ficusgallae; 162, Axima brevicornis; 163, Eurytoma Cébazan; 164, Eurytoma braconidis. Figure 165: mesopleuron in ventral view; Conoaxima affinis. Figures 166, 167, 168, 170: metapleuron in ventral view; 166, Glyphomerus stigma; 167, Buresium rufum; 168, Archirileya Cicada; 170, Bruchophagus Alicante. Figure 169: metapleuron (after dissection) in anterior view (internal skeleton visible) – Archirileya Cicada. Figure 171: hind coxal cavity and metafurcal pit (enlarged) – Rileya pulchra.