ex gall Aylax minor Hartig on Papaver spp., Spain, Guadalajara: Valdenoches, 31.VII.01, J. L. Nieves leg (n = 1); Madrid: Monte Pajares, 7.IX.03, J. L. Nieves leg (n = 11); Madrid: Rivas-Vaciamadrid, 14.V.03, J. L. Nieves leg (n = 1); Madrid: Valdemorillo, 13.VI.04, J. L. Nieves leg (n = 2); Valladolid: Cabezón-San Martín de Valveni, 22.VI.02, J. L. Nieves leg (n = 7).

n = 22; Body length: 1.68 ± 0.33 mm (min-max: 1.13-2.20 mm), width: 0.92 ± 0.16 mm (min-max: 0.67-1.20 mm). Body fusiform, relatively short and wide, slightly wider at the level of ABS2-ABS3, but not tapering abruptly towards ANS (Figs 3A, 5A) (Table 2); adp present from the second thoracic to fifth abdominal segment, not protruding beyond the dorsal margin of body in lateral view (Fig. 5A); integument of the body smooth; thoracic setae longer than abdominal setae but shorter than length of a thoracic segment Head 1.14 broader than long (Fig. 7A); vertex concave; distance among vam longer than SA; am situated clearly above the antennae (Table 2). On clypeus lcs as long as cs (Fig. 9A), being both situated at the same level; lll not clearly differentiated and almost merged with medial labrum lobe; posterior margin of labrum straight. Mandibles one-toothed with apex of tooth clearly sharp (Table 2).

This species is a common parasitoid in poppy galls of Aylax minor Hartig, 1840 (Hym., Cynipidae) (Fig. 12A and B). The species has also been reared from galls of Aylax papaveris and Barbotinia oraniensis on the heads of Papaver ssp. (Askew et al. 2006). The parasitoid behaviour of Ormyrus capsalis is very similar to the related species Ormyrus papaveris, an idiobiont ectoparasitoid of cynipid larvae (Askew et al. 2006) (Fig. 12C).

Notably, in some cases, we observed terminal-instar larvae of Ormyrus inside cells of Aylax minor, which apparently were not consumed (Fig. 12D).

ex gall Eurytoma gallephedrae Askew on Ephedra nebrodensis, Spain, Madrid: Monte Pajares, 24.I.04, J. L. Nieves leg (n=1).

n = 1; Body length: 1.5 mm, width: 0.61 mm. Body fusiform, broader at the level of abdominal segments ABS2-ABS3 and tapering posteriorly towards ANS; ANS broader than long (Figs 3B, 5B) (Table 2); adp present from second thoracic to fifth abdominal segment, protruding conspicuously beyond the dorsal margin of body in lateral view, but only at the level of abdominal area (Fig. 5B); thoracic and abdominal segments with blister-like sculpture; thoracic setae relatively long, as long as the length of a thoracic segment. Head 1.1 broader than high (Fig. 7B); blister-like sculpture extended over the head; vertex concave in the middle; distance among vam as distance SA; am situated clearly above the antennae (Table 2). The lcs situated above cs; cs separated from lcs 2.5 as distance between cs (Fig. 9B). Lateral lobes of labrum almost fused with the medial lobe; posterior margin of medial piece of medial lobe straight. Maxilary palps indistinct. Mandibles unidentated with apex of tooth acute (Table 2).

The larva of O. cupreus was described as a specific parasitoid of galls induced by Eurytoma gallephedrae Askew (Chalcidoidea, Eurytomidae) on Ephedra nebrodensis stems (Fig. 12E). Additionally, from this host, we also reared larvae and adults of O. cupreus from galls on subterranean runners of Ephedra nebrodensis (Fig. 12F), most likely induced by Eurytoma flaveola (Zerova 1796), which is a species recorded inducing galls on Ephedra roots in Asia (Zerova 1995). Askew and Blasco-Zumeta (1998) performed detailed observations on the biology of this species and found O. cupreus is a primary, solitary idiobiont ectoparasitoid of the larva of E. gallephedrae, also attacking the adult Eurytoma at times or as a hyperparasitoid attacking larvae of the eupelmids Brasema and Eupelmus. These authors also reported cannibalistic behaviour. Because the remains of an adult Eurytoma were found jointly with a larva of O. cupreus, we confirmed the observations of Askew (Fig. 12G and H).

ex gall Liposthenes kerneri (Wachtl) on Nepeta hispanica Spain, Madrid: Casa Eulogio, 01.VI.03, J. L. Nieves leg (n = 8); Madrid: Rivas-Vaciamadrid, 01.VI.03, J. L. Nieves leg (n = 5); Madrid: Rivas-Vaciamadrid, 13.VI.03, J. L. Nieves leg (n = 1); Madrid: Rivas-Vaciamadrid, 17.V.03, J. L. Nieves leg (n = 22)

n = 36; Body length: 1.56 ± 0.34 mm (min-max: 1.00-2.40 mm), width: 0.83 ± 0.12 mm (min-max: 0.60-1.00 mm). Body fusiform, not tapering abruptly towards anal segment (Figs 3C, 5C) (Table 2); adp present from second thoracic to fifth abdominal segment protruding beyond the dorsal margin of body in lateral view (Fig. 5C); integument smooth; thoracic setae relatively long, abdominal setae shorter. Head 1.12 times wider than high (Fig. 7C); integument of genal area with blister-like sculpture; vam more separated than distance SA; am situated slightly above antennae (Table 2).

On clypeus lcs situated at the same level of cs; lateral lobes of labrum strongly remarked and incompletely fused with the medial lobe; posterior margin of medial piece of labrum convex (Fig. 9C); mandible unidentate with the apex slightly visible under lb; tooth acute (Fig. 11B).

The species is a common parasitoid reared from cynipid galls of Liposthenes kerneri (Wachtl) on fruits of Nepeta ssp. (Lamiaceae; Fig. 12I) (Askew et al. 2006). Full-growth larva of O. diffinis occupied the entire primary cell of a parasitized gall after the host larvae was devoured (Fig. 12J and K). We observed that larvae of O. diffinis inside galls apparently entered prolonged periods of diapause, without causing normal pupation and adult emergence after the winter diapause period. Moreover, live terminal-instar larvae of O. diffinis were found in galls dissected two years after collection. The data indicate that O. diffinis is an idiobiont ectoparasitoid with a univoltine life cycle that is synchronized with the emergence and growth of their host galls on species of Nepeta. The insects emerge in the second year when the new galls are available again on the host plant. This species has also been reared from galls of Neaylax salviae and N. nemorosae on different species of Salvia (Lamiaceae) and from those of Rhodus cyprius on Salvia triloba (Lamiaceae) (Askew et al. 2006).

ex gall Isocolus scabiosae (Giraud) on Centaurea scabiosa, Spain, Guadalajara: Pozo de Guadalajara, 31.VII.02, J. L. Nieves leg (n = 4); Pozo de Guadalajara, 03.X.04, J. L. Nieves leg (n = 7).

n = 11; Body length: 2.22 ± 0.63 mm (min-max: 1.40-3.60 mm), width: 1.28 ± 0.31 mm (min-max: 0.67-1.87 mm). The species differs from O. capsalis in the following characters: head 1.1 times as wide as high; genal area, vertex and first thoracic segment with blister-like sculpture; antennae mid-situated in anterior view of the head; antennal setae 0.35 as long as distance between antennae; lcs situated above lc (Figs 3D, 5D, 7D; Table 2).

Larvae of O. gratiosus are oligophagous idiobiont ectoparasitoids of species of Isocolus that induce galls on flower heads of Centaurea and Serratula species (Asteraceae) (Askew et al. 2006). Additionally, the species has been reared from the galls of Diastrophus mayri on Potentilla argentea (Rosaceae). Our examined material was from dissected galled achenes of the flower heads of Centaurea scabiosa in Spain (Fig. 13A, B, and C).

ex gall Andricus hispanicus on Quercus canariensis, Spain, Málaga: Algatocín, 19.VIII.02, J. L. Nieves leg (n = 1); ex gall Andricus hispanicus on Quercus faginea, Spain, Salamanca: Laguna de San Marcos, 26.VIII.03, J. L. Nieves leg (n = 1)

n = 2; Body length: 4.28 ± 0.87 mm (min-max: 3.67-4.90 mm), width: 2.13 ± 0.18 mm (min-max: 2.00-2.25 mm).

The larva of this species is the largest among all the European species. Is quite similar in most diagnostic characters to the larvae of the related species O. pomaceus, being differentiated by its large size and the blister like sculpture much less conspicuous. Other diagnostic characters are as follows: body short and wide, not tapering towards the anal segment. Setae of thoracic segments shorter than ½ length of a thoracic segment; ratio AC/AV 0.77, the shortest among all the studied species (Table 2); anteromedial seatae of antennal area short, 0.3 as long as distance among antennae; lcs separated from cs 0.7 times the distance between cs; maxillary palps conspicuous (Figs 3E, 5E, 7E; Table 2).

The species O. nitidulus is a member, with the closely allied O. pomaceus, of the parasitoid community associated with oak gall wasps (Hymenoptera, Cynipini). The two species were reared from more than 50 different species of cynipids associated with Quercus species in the west Palaearctic (Askew et al. 2013); however, O. nitidulus is not as common and is less polyphagous than O. pomaceus. In contrast to the closely related species O. pomaceus, O. nitidulus prefers to attack the large galls of asexual generations of heteroecic species of Andricus. On the Iberian Peninsula, O. nitidulus was reared primarily from galls of Andricus hispanicus (Fig. 13D, E, and F) and the asexual generation of Andricus grossulariae. Our observations of dissected galls showed the larva of O. nitidulus was a primary ectoparasitoid of the galling inducer. In the host galls of Andricus hispanicus, the larvae always occupied the host central larval chamber, not the secondary cells occupied by inquilines.

ex gall of an undetermined Tephritidae (Diptera) on Microlonchus salmanticus, Spain, Salamanca: La Flecha (23/X/02), J. L. Nieves leg (n = 1).

n = 1; Body length: 2.35 mm, width: 1.35 mm

Body fusiform, short and wide, slightly wider at the level of ABS2-ABS3, but not tapering abruptly towards ANS (Fig. 3F) (Table 2); body segments with conspicuous blister-like sculpture; thoracic setae relatively long, clearly shorter than abdominal setae. Head 1.14 as wide as high (Fig. 7F); face integument smooth; medial area of vertex regularly convex; antennae situated at mid distance among vertex and ventral margin of clypeus; ratio AC/AV 1.22; am situated at the same level of antennae; antennal setae 0.4 as long as distance among antennae (Table 2).

On clypeus lcs situated at the same level of cs, both equal in length (Fig. 9F); lateral lobes of labrum inconspicuous and almost fused with the medial lobe; posterior margin of the medial piece of labrum straight; mandibles unidentated with the apex of tooth acute (Table 2).

In contrast to most European species of Ormyrus, the larvae of O. orientalis attack dipteran galls induced by tephritids (Diptera, Tephritidae) in the heads of different species of Asteraceae. On the Iberian Peninsula, tephritid galls containing O. orientalis were found on Microlonchus salmanticus (Asteraceae) (Fig. 13I), and the species was also reared from galls of Myopites limbardae Schiner (Tephritidae) on Inula viscosa (Asteraceae) (Fig. 13G and H). Based on our unpublished data from Malaise traps and sweep net samples, O. orientalis was one of the most abundant ormyrid species in many habitats on the Iberian Peninsula; consequently, the list of hosts could be wider than that reported in the literature and in the data of the authors.

ex gall Aylax papaveris on Papaver spp., Spain, Guadalajara: El Cardoso de la Sierra, 30.VI.02, J. L. Nieves leg. (n = 4); Soria: San Andrés, 14.VII.05, J. L. Nieves & J. F. Gómez leg. (n = 1); ex gall Barbotinia oraniensis on Papaver spp., Spain, Madrid: Rivas-Vaciamadrid, 25.V.02, J. L. Nieves leg. (n = 2); Madrid: Rivas-Vaciamadrid, 13.VI.04, J. L. Nieves leg. (n = 1).

n = 8; Body length: 1.88 ± 0.24 mm (min-max: 1.53-2.13 mm), width: 0.94 ± 0.19 mm (min-max: 0.67-1.20 mm). This species is similar to Ormyrus capsalis from which may be distinguished in the body fusiform, slightly wider at the level of body segments ABS2-ABS3, tapering towards the ANS (Figs 4A, 5F) and the anteromedial setae of antennal area being relatively short, <0.3 the distance among antennae. Other descriptive diagnostic characters as follows: thoracic setae short; head 1.07 times wider than high (Fig. 8A); face integument smooth; antennae situated at mid position in the face; am short and situated above antennae (Table 2). On clypeus lcs situated at the same level of cs, both equal in length but short and inconspicuous (Fig. 9G); lateral lobes of labrum conspicuous and clearly separated from the medial lobe; mandibles with a single tooth with acute apex (Fig. 11E).

The larvae of O. papaveris are common ectoparasitoids in poppy galls of different Aylacini (Cynipidae) species, primarily Aylax papaveris and Barbotinia oraniensis (Figs 13J and K; 14A), and attack the host in the early stages of development (Askew et al. 2006). We observed the remains of the host larva on the body of a mature ormyrid larva (Fig. 14C). The host larval chamber of Barbotinia oraniensis was normally spherical and regular (Fig. 14B); however, when O. papaveris attacked the host, the chamber was shorter and irregular (Fig. 14C). The larva of O. papaveris moved inside the host gall larval cell touching the gall chamber walls with their mandibles, which suggested that during the terminal larval stage, O. papaveris might exhibit a similar phytophagous behaviour to that of Eurytoma species inhabiting galls (Askew and Blasco Zumeta 1998, La Salle 2005). In the galls of Aylax papaveris, the host larval cells were regularly ellipsoidal and were coated with a thin scum, whereas the cells attacked by ormyrid larvae were larger, more irregular and lacked the thin scum. Because we observed “in vivo” in dissected galls, the phytophagous behaviour of the Ormyrus larvae during their final larval stage caused the change in gall morphology.

ex gall Andricus grossulariae asex. on Quercus faginea, Spain, Cádiz: La Suara-Jérez, 16.X.04, J. L. Nieves leg. (n = 1); ex gall Plagiotrochus fusifex on Quercus coccifera, Spain, Madrid: Arganda, 01.VI.03, J. L. Nieves leg. (n = 1); ex gall Plagiotrochus razeti on Quercus ilex, Spain, Madrid: Villanueva del Pardillo, 07/X/02, J. L. Nieves leg. (n = 11). ex gall Trigonaspis mendesi on Quercus faginea, Spain, Madrid: Boadilla del Monte, 23/IX/02, J. L. Nieves leg. (n = 1).

Ex gall Andricus grossulariae asex., on Quercus faginea, n = 1; Body length: 2.73 mm, width: 1.67 mm; ex gall Plagiotrochus fusifex on Quercus coccifera, n = 1; Body length: 1.13 mm, width: 0.53 mm; ex gall Plagiotrochus razeti on Quercus ilex, n = 11; Body length: 2.21 ± 0.46 mm (min-max: 1.80–3.40 mm), width: 1.10 ± 0.10 mm (min-max: 0.93–1.27 mm); ex gall Trigonaspis mendesi on Quercus faginea, n = 1; Body length: 1.55 mm, width: 0.80 mm.

The morphology of the terminal larva of this species is very similar to that of the Ormyrus nitidulus larva. The larvae of O. pomaceus from galls of Andricus and Trigonaspis species were distinguished from those of O. nitidulus by the following characters: integument of thoracic and abdominal segments with conspicuous blister-like sculpture; distance among vertex setae longer than the distance between antennae; am 0.47 as long as the distance between antennae; lcs situated above the level of cs, being separated from cs by 1.2-fold the distance between cs; and maxillary palps not visible (Table 2).

For the O. pomaceus larvae that inhabited Plagiotrochus galls (Figs 4C, 6B, 8C, and 10A), the differences between O. pomaceus ex Andricus and ex other host genera, such as Trigonaspis (Figs 4B, 6A, 8B, and 9H), were the following: am shorter in length than the separation between antennae (Table 2) and maxillary palps conspicuous.

Ormyrus pomaceus is a polyphagous ectoparasitoid that attacks more than 56 different cynipid galls on Quercus trees (Figs 14F and I; 15D and E) (Askew et al. 2013). Nevertheless, results from ongoing unpublished molecular studies clearly indicate that O. pomaceus includes a complex of sibling or cryptic species that are segregated according to cynipid hosts, host plant species and ecological preferences (Hernandez Nieves et al. unpublished, Stone pers. comm.). On the Iberian Peninsula, among the most regular host species of O. pomaceus, we found the asexual generations of Andricus grossulariae (Fig. 14D and E) and A. pictus (Fig. 14G and H), Trigonaspis mendesi (Fig. 14J and K) and T. brunneicornis (Figs 14L and 15A) on Quercus pyrenaica and Q. faginea and the galls of asexual species of Plagiotrochus on Q. ilex and Q. coccifera (Fig. 15B and C).

ex gall Xestophanes potentillae on Potentilla reptans, Spain, Madrid: Cotos de Monterrey, 24.VI.03, J. L. Nieves leg (n = 2); Madrid: Villalvilla, 26.VIII.05, J. L. Nieves leg (n = 9); Madrid: Villar del Olmo, 03.X.04, J. L. Nieves leg (n = 23); Tarragona: Colldejou, 14.VIII.03, J. L. Nieves leg (n = 7).

n = 41; Body length: 1.69 ± 0.39 mm (min-max: 1.13–2.53 mm), width: 0.84 ± 0.22 mm (min-max: 0.47-1.27). Body fusiform, abdominal segments tapering abruptly towards ANS (Figs 4D, 6C); adp strongly remarked; integument of abdominal segments smooth but with blister-like sculpture extended in part of thoracic segments; setae on thoracic segment long, not longer than length of a thoracic segment, shorter on abdominal region. Head 1.03 as wide as high (Fig. 8D); integument on the face smooth; vertex concave; an situated at mid position in the face; vertex setae equally separated than the distance between antennae; am situated at the same level of an; am short, 0.22 times as long as the separation between antennae (Table 2). On clypeus lcs situated at the same level of cs, both equal in length (Fig. 10B); lateral lobes of labrum slightly differentiated and almost fused with the medial lobe; posterior margin of medial lobe of labrum straight; mandibles unidentated; tooth apex acute (Table 2).

This species is extremely host-specific and is exclusively associated with galls on the runners and roots of Potentilla reptans (Rosaceae) induced by Xestophanes potentillae (Retzius) (Fig. 15H and I) (Askew et al. 2006). On the Iberian Peninsula, two forms of the galls were found. One form was on stems or runners close to or beneath the soil surface that consisted of round swellings (Fig. 15G), and the others formed on subterranean rhizomes (Fig. 15F).

In the first stages, the larva of O. rufimanus and the paralyzed host larva co-occurred; in later stages, the remains of the host larva appeared on the ventral surface of the O. rufimanus larva. In dissected galls, the larvae of O. rufimanus were extracted from irregularly shaped larval gall cells, which indicated that vegetal material was consumed at the terminal larval stage, as observed with other Ormyrus species such as O. papaveris. Based on additional observations, we found larvae of Eupelmus vesicularis (Chalcidoidea, Eupelmidae) were hyperparasitoids of O. rufimanus pupae.

ex gall Neaylax verbenacus on Salvia verbenaca, Spain, Madrid: Dehesa de Arganda, 09.VI.02 J. L. Nieves leg (n = 1).

n = 1; Body length: 1.67 mm, width: 0.80 mm. The larva of this species is similar to the larva of O. diffinis, from which may be distinguissed as follows: body fusiform, wider at the level of segments ABS2-ABS3, tapering progresively towards ANS; anal segment wider tan length; adp absent; integument of the abdominal and thoracic segments blister-like. Head 1.18 as wide as high (Fig. 8E) with blister-like sculpture extended on all the head; antennae situated at mid position in the face; ams situated clearly above the antennae; lateral lobes of labrum almost fused with the medial lobe; ventral margin of medial lobe of labrum straight (Fig. 10C).

The larva of O. wachtli is a solitary ectoparasitoid of larvae of cynipids, inducing galls on fruits of Salvia (Lamiaceae). Along the Iberian Peninsula and in southern Europe, the species is associated with galls of Neaylax salviae (Giraud) on Salvia lavandulifolia (Fig. 15L) and Neaylax verbenacus (Nieves-Aldrey) on Salvia verbenaca (Fig. 15J and K) (Nieves-Aldrey 2001, Nieves-Aldrey and Askew 2002, Askew et al. 2006). The species has a bivoltine life cycle.