Table 1.
Classification, biology and collection site for the cynipoid species included in the study.
Table 2.
Data matrix based on the characters listed in the Materials and methods.
Figure 1.
Phylogenetic relationships among the studied species of Cynipoidea, as depicted from recent studies [59]–[64] and unpublished data (see Methods for details).
The main life-history trait for each species is mapped on the tree. For non-gall parasitoids, U = unconcealed host and C = concealed host. “?” denotes that biology is unknown for Araucocynips queulensis.
Figure 2.
Variability in the general aspect of the antennae of Cynipoidea.
A) Aulacidea freesei (filiform (3-0) with 10 flagellomeres (1-0) and F1 about as long as F2 (2-0)), B) Andricus crispator (sexual) (filiform (3-0) with 10 flagellomeres (1-0) and F1 1.2–1.5 times longer than F2 (2-1)), C) Eschatocerus acaciae (filiform (3-0) with 11 flagellomeres (1-1) and F1 1.2–1.5 times longer than F2 (2-1)), D) Cecinothofagus gallaelenga (clavate (3-0) with 10 flagellomeres (1-0) and F1 about as long as F2 (2-0)), E) Trigonaspis mendesi (asexual) (filiform (3-0) with 11 flagellomeres (1-1) and F1>1.5 times longer than F2 (2-2)), F) Roophilus loewi (filiform (3-0) with 12 flagellomeres (1–2) and F1 1.2–1.5 times longer than F2 (2-1)), G) Phanacis centaureae (filiform (3-0) with 12 flagellomeres (1–2) and F1>1.5 times longer than F2 (2-2)), H) Qwaqwaia scolopiae (filiform (3-0) with 12 flagellomeres (1–2) and F1 about as long as F2 (2-0)), I) Synergus hayneanus (filiform (3-0) with 12 flagellomeres (1–2) and F1 about as long as F2 (2-0)), J) Pediaspis aceris (asexual) (filiform (3-0) with 13 flagellomeres (1–3) and F1 1.2–1.5 times longer than F2 (2-1)), K) Ganaspis sp. (moniliform (3-0; 4-1) with 11 flagellomeres (1-1) and F1 about as long as F2 (2-0)), L) Parnips nigripes (filiform (3-0) with 11 flagellomeres (1-1) and F1 about as long as F2 (2-0)).
Figure 3.
Dendrogram depicted by the Hierarchical Cluster Analysis (Ward's method) based on the matrix of presence/absence of the 12 different types of sensilla for each species.
The dashed line represents the most probable truncation that segregates different clusters. The main life-history trait for each species is mapped on the dendrogram, as well as the taxonomic position of each species. For non-gall parasitoids, U = unconcealed host and C = concealed host. “?” denotes that biology is unknown for Araucocynips queulensis. Note that one cluster is exclusively composed of Cynipini and in particular of species in the genus Andricus, while the other two groups include a less defined mixture of species.
Figure 4.
Variability in number, relative size and arrangement of sensilla placoidea (SP) in the flagellomere Fn of Cynipoidea.
A) Andricus corarius (asexual) (arranged in one row (11-0), present only dorsally (12-0), 6–8 SP per row (13-1), widely separated in a row (14-0), almost flat (15-0), with surface constantly plane (16-0), more or less overlapping the distal margin of Fn (17-1), linear, with parallel margins (18-0)), B) Acanthaegilips sp. (arranged in two rows (11-1), present dorsally and ventrally (12-1), 6–8 SP per row (13-1), widely separated in a row (14-0), raising on Fn (15-0), with surface constantly plane (16-0), only reaching the distal margin of Fn (17-0), linear, with parallel margins (18-0)), C) Andricus burgundus (sexual) (arranged in one row (11-0), present only dorsally (12-0), 3–5 SP per row (13-0), widely separated in a row (14-0), almost flat (15-0), with surface constantly plane (16-0), more or less overlapping the distal margin of Fn (17-1), linear, with parallel margins (18-0)), D) Apocharips sp. (arranged in one row (11-0), present dorsally and ventrally (12-1), 3–5 SP per row (13-0), widely separated in a row (14-0), almost flat (15-0), with surface constantly plane (16-0), more or less overlapping the distal margin of Fn (17-1), linear, with parallel margins (18-0)), E) Callaspidia notata (arranged in two rows (11-1), present dorsally and ventrally (12-1), 6-8 SP per row (13-1), widely separated in a row (14-0), almost flat (15-0), with surface constantly plane (16-0), more or less overlapping the distal margin of Fn (17-1), more or less sinuate (18-1)), F) Cecinothofagus gallaelenga (arranged in one row (11-0), present dorsally and ventrally (12-1), 6–8 SP per row (13-1), widely separated in a row (14-0), almost flat (15-0), with a longitudinal groove (16-1), only reaching the distal margin of Fn (17-0), more or less sinuate (18-1)), G) Diastrophus rubi (arranged in one row (11-0), present dorsally and ventrally (12-1), 6–8 SP per row (13-1), widely separated in a row (14-0), almost flat (15-0), with surface constantly plane (16-0), more or less overlapping the distal margin of Fn (17-1), more or less sinuate (18-1)), H) Hedickiana levantina (arranged in three-four rows (11-2), present dorsally and ventrally (12-1), >8 SP per row (13-2), narrowly separated in a row (14-1), almost flat (15-0), with surface constantly plane (16-0), only reaching the distal margin of Fn (17-0)), I) Eschatocerus acaciae (arranged in one rows (11-0), present dorsally and ventrally (12-1), 6–8 SP per row (13-1), narrowly separated in a row (14-1), almost flat (15-0), with surface constantly plane (16-0), only reaching the distal margin of Fn (17-0), linear, with parallel margins (18-0)), J) Ganapsis sp. (arranged in one rows (11-0), present dorsally and ventrally (12-1), 3–5 SP per row (13-0), widely separated in a row (14-0), almost flat (15-0), with surface constantly plane (16-0), only reaching the distal margin of Fn (17-0), linear, with parallel margins (18-0)), K) Ibalia rufipes (arranged in three-four rows (11-2), present dorsally and ventrally (12-1), 6–8 SP per row (13-1), narrowly separated in a row (14-1), almost flat (15-0), with surface constantly plane (16-0), only reaching the distal margin of Fn (17-0), more or less sinuate (18-1)), L) Oberthuerella sp. (arranged in three-four rows (11-2), present dorsally and ventrally (12-1), >8 SP per row (13-2), closely spaced in a row (14-2), almost flat (15-0), with surface constantly plane (16-0), only reaching the distal margin of Fn (17-0), with parallel margins (18-0)).
Figure 5.
Examples of sensilla coeloconica type A (SCo-A) found in the flagellomere Fn of Cynipoidea.
A) Aulacidea tragopogonis, B) Andricus curvator (sexual), C) Periclistus brandtii, D) Neralsia sp., E) Pediaspis aceris (asexual), F) Timaspis phoenixopodos, G) Callaspidia notata, H) Ceroptres cerri, I) Dryocosmus kuriphilus (asexual), J) Eschatocerus acaciae, K) Iraella luteipes, L) Xestophanes potentillae. Note the variability in the diameter of the SCo pit.
Figure 6.
Variability in number, relative size and arrangement of sensilla coeloconica type A (SCo-A) in the flagellomere Fn of Cynipoidea.
A) Acanthaegilips sp. (one per flagellomere (21-1), far from the distal margin (22-1), small (23-0)), B) Hedickiana levantina (≥3 per flagellomere (21-3), far from the distal margin (22-1), small (23-0)), C) Aulacidea tragopogonis (one per flagellomere (21-1), far from the distal margin (22-1), small (23-0)), D) Andricus curvator (sexual) (one per flagellomere (21-1), on or close the distal margin (22-0), large (23-2)), E) Eschatocerus acaciae (two per flagellomere (21-2), far from the distal margin (22-1), large (23-2)), F) Pediaspis aceris (asexual) (≥3 per flagellomere (21-3), on or close the distal margin (22-0), medium size (23-1)).
Figure 7.
Examples of sensilla coeloconica type B (SCo-B) and sensilla campaniformia (SCa) found in the flagellomeres of Cynipoidea.
A) Andricus quercusilicis (sexual) (arrow poiting at the peg of SCa), B) Andricus burgundus (sexual), C) Diastrophus rubi (arrow poiting at the peg of SCo-B), D) Cecinothofagus gallaelenga, E) Qwaqwaia scolopiae, F) Hedickiana levantina, G) Andricus curvator (sexual), H) Isocolus lichtensteini, I) Andricus grossulariae (sexual), J) Andricus multiplicatus (sexual), K) Ceroptres cerri, L) Pediaspis aceris (asexual). Note that these two types of sensilla are overall similar but in SCa the peg a bit smaller and is on the top of a doomed area, while in SCo-B a slightly larger peg visibly arises from a pit in a less doomed and even often depressed, concave area. Note also a rare case of a pair of SCa in E (arrows).
Figure 8.
Examples of sensilla basiconica (SB) found in the flagellomeres of Cynipoidea.
A) Andricus corarius (asexual), B) Periclistus brandtii, C) Roophilus loewi, D) Trigonaspis sinaspis (sexual), E) Andricus multiplicatus (sexual), F) Synergus physocerus, G) Synergus clandestinus, H) Aylax papaveris, I) Callaspidia notata, J) Diastrophus rubi, K) Qwaqwaia scolopiae, L) Oberthuerella sp. Note one small and one large SB in I.
Figure 9.
Examples of sensilla trichoidea (ST-A, ST-B, ST-C, ST-D, ST-E) found in the flagellomeres of Cynipoidea.
A) Andricus curvator (sexual), B) Andricus coriarius (asexual), C) Andricus grossulariae (asexual), D) Andricus grossulariae (asexual), E) Pediaspis aceris (asexual), F) Aulacidea tragopogonis, G) Aulacidea papaveris, H) Synergus hayneanus, I) Oberthuerella sp., J) Cynips quercusfolii (asexual), K) Eschatocerus acaciae, L) Neralsia sp.
Table 3.
Measurements (in µm) taken on the antennae and sensilla of the studied species.
Figure 10.
Unique sensilla types found on the apex of the antennae of Paraulacini (Cynipidae) and Plectocynipinae (Figitidae).
A) last three flagellomeres of Plectocynips pilosus (Plectocynipinae), with arrows pointing the three-disc Large Disc Sensilla (SLD), B) last three flagellomeres of Araucocynips queulensis (Plectocynipinae), with arrows pointing the five-disc Large Disc Sensilla (SLD), C) Detail of the SLD in A. queulensis (Plectocynipinae), D) apical clava of Cecinothofagus gallaelenga (Paraulacini), with arrow poiting the Large Volcano Sensilla (SLV), E) lateral view of the SLV in C. gallaelenga (Paraulacini), F) frontal view of the SLV in C. gallaelenga (Paraulacini), G) detail of the cone entrance of the SLV in C. gallaelenga (Paraulacini).