Eurytoma asphodeli Hedqvist, 1976

Delvare, G., Escolà, A. Ribes, Stojanova, A. M., Benoit, L., Lecomte, J. & Askew, R. R., 2019, Exploring insect biodiversity: the parasitic Hymenoptera, chiefly Chalcidoidea, associated with seeds of asphodels (Xanthorrhoeaceae), with the description of nine new species belonging to Eurytomidae and Torymidae, Zootaxa 4597 (1), pp. 1-90 : 50-56

Eurytoma asphodeli Hedqvist, 1976

( Figs 22 A–G, 23A–J)

Type material. Paratype ♀ labelled: ‘ Ibiza, San Antonio 7.5.1975. B. Gustafsson’ / ‘ex seed-case of Asphodelus microcarpus’ / ‘Paratypus Eurytoma asphodeli ♀ sp. n. K-J Hedqvist det.’ ; paratype ♂, same data as holotype [in bad condition] (in BMNH)

Other material. Another two males and two females in BMNH are labelled ‘standing over Eurytoma asphodeli in Hedqvist coll. BMNH (E) 2011-27 ’ and, in Hedqvist’s handwriting, ‘Portugal, Cruz , 10/5/[?]1988 K. J. Hedqvist’ . FRANCE: Alpes-Maritimes, Saint-Dalmas-le-Selvage , GR 5-56 Bousieyas-point 44a, N Combe Male, 1930–2000 m, 44.31833°N 6.85833°E, collected on flowers of 20.vi.2012, A. albus delphinensis (G. Delvare) (vouchers GDEL1399 ♀, GDEL1400 ♀, GDEL1401 ♀, GDEL1402 ♀, GDEL1491 ♀, GDEL1492 ♀, GDEL1493 ♀, GDEL1494 ♀, in CIRAD) ; same locality, asphodel and collector, ex seeds of the plant, adults emerged on 01–15.v.2013 (48 ♀ 2 ♂, in GDPC) ; Aveyron, Lapanouse-de-Cernon, Causse Viel , 810 m, 43.98222°N 3.57417°E, collected on flowers of A. cerasiferus , 01.vi.2009 (G. Delvare) (6 ♀ 2 ♂, in GDPC) ; same locality, asphodel and collector, collected on flowers, 23.v.2011 (vouchers GDEL1459 ♀, GDEL1460 ♀, GDEL1461 ♀, GDEL1462 ♀, in CIRAD) ; Bouches-du-Rhône, La Ciotat , ex seeds A. cerasiferus , 12.vi.2010 and 28.v.2011 (H. Dumas) (3 ♀ 6 ♂, donated to A. Ribes, now probably in BMNH) ; Corse, Aléria , Vaccaja , 20 m, 42.12861°N 9.46556°E, ex seeds of A. ramosus , 21.ix.2011 adults emerged 15–30.iv.2012 (J. Balajas) (vouchers GDEL1414 ♀, GDEL1415 ♀, GDEL1416 ♀, GDEL1417 ♀, GDEL1418 ♀, GDEL1419 ♀, GDEL1420 ♀, GDEL1421 ♀, GDEL1422 ♀, GDEL1423 ♀, GDEL1424 ♀, GDEL1425 ♀, GDEL1426 ♀, GDEL1427 ♀, in CIRAD; 18 ♀ 2 ♂, in GDPC) ; Galéria, Piana di l'Olmu , 33 m, 42.41.590°N 8.65796°E, collected on flowers of A. ramosus , 29.iv.2002 (G. Delvare) (8 ♀, in GDPC) ; Linguizetta, 50 m, 42.28194°N 9.51722°E, ex seeds of A. ramosus , 21.ix.2011, adults emerged 15–30.iv.2012 (J. Balajas) (vouchers GDEL1438 ♀, GDEL1439 ♀, GDEL1440 ♀, GDEL1441 ♀, GDEL1442 ♀, in CIRAD; 5 ♀ 46 ♂, in GDPC ) ; Piedicorte-di-Gaggio , N 200, pont génois, 183 m, 42.22167°N 9.26528°E, ex seeds of A. ramosus , 21.ix.2011, adults emerged 15–30.iv.2012 (J. Balajas) (vouchers GDEL1433 ♀, GDEL1434 ♀, GDEL1435 ♀, GDEL1436 ♀, GDEL1437 ♀, in CIRAD; 26 ♀ 8 ♂, in GDPC ) ; San Pietro di Venaco , N 200, ancient hippodrome, 218 m, 42.23806°N 9.22833°E, ex seeds of A. ramosus , 21.ix.2011, adults emerged 15–30.iv.2012 (J. Balajas) (vouchers GDEL1428 ♀, GDEL1429 ♀, GDEL1430 ♀, GDEL1431 ♀, GDEL1432 ♀, in CIRAD; 16 ♀ 18 ♂, in GDPC ) ; Dordogne, Belvès, Forêt de la Bessède , 232 m, 44.79361°N 0.90555°E, ex seeds of A. albus albus , 24.vi.2011 & 23.vi.2013 (R.R. Askew) (15 ♀ 9 ♂, in RAPC) ; La Douze, Forêt Domaniale de la Barade , 235 m, 44.87777 °N 0.86028°E, 19.vi.2011, adults on green fruits of A. albus albus (R.R. Askew) (4 ♀ 2 ♂, 2 in RAPC), 18.v.2011, 14 & 19.vi.2011 and 23.vi.2012, ex seeds, same asphodel and collector (262 ♀ 275 ♂, 21 ♀ in RAPC; 1 ♀ in GDPC) ; Mouleydier, Forêt de Liorac , 127 m, 44.87778°N 0.62861°E, 6.v.2011, 1♀ ovipositing in green fruit of A. a. albus (R.R. Askew) ; 18.vi.2006, 03.vi.2007, 26.vi.2009, same asphodel and collector (3 ♀, 2 ♂ in RAPC) ; Hautes-Pyrénées, Gavarnie-Gèdre , Vallée d'Ossoue , 1655 m, 42.75111°N 0.12166°W, ex seeds of A. albus delphinensis , 24.vii.2007, 9.vii.2009 and 14.vii.2010 (R.R. Askew) (3 ♀ 4 ♂, 4 in RAPC ; same locality, asphodel and collector but adults on fruits (33 ♀ 1 ♂, 3 in GDPC) ; Hérault, Grabels , 07–13.v.1989 (H. Tussac) (1 ♀, in GDPC) ; Mireval , ex seed of A. fistulosus , 11.vi.1988 (G. Delvare) (1 ♀, in GDPC) ; Pyrénées Orientales, Banyuls-sur-Mer, E Col de la Créu , 240 m, 42.46278°N 3.14222°E, ex seeds of A. ramosus , 06.vi.2011, adults emerged 15–30.iv.2012, (J. Lecomte) (vouchers GDEL 1292 ♀, GDEL 1293 ♂, GDEL 1294 ♂, GDEL 1443 ♀, GDEL 1444 ♀, GDEL 1445 ♀, GDEL 1446 ♀, GDEL 1447 ♀, GDEL 1448 ♀, GDEL 1449 ♀, GDEL 1450 ♀, GDEL 1451 ♀, GDEL 1452 ♀, GDEL 1453 ♀, GDEL 1454 ♀, GDEL 1455 ♀, GDEL 1456 ♀, GDEL 1457 ♀, GDEL 1458 ♀, in CIRAD; 179 ♀ 91 ♂, in GDPC) ; Vienne, Châtellerault , Aire de Chagnats and Aire de Meumiers on A10, 131 m, 48.81778°N 0.54611°E, ex seeds of A. a. albus , 28.vii.2007, 22.vi.2008 and 2.vii.2010 (R.R. Askew) (9 ♀ 9 ♂, 13 in RAPC & 1 ♀ in GDPC) ; ITALY: Sicilia, Belpasso, Contrada Segreta , S slope Etna , 900 m, 37.63818°N 14.98592°E, collected on flowers of A. ramosus , 25.v.2014 (F. Turrisi) (vouchers GDEL 1639 ♀, GDEL 1640 ♀, GDEL 1641 ♀, GDEL 1642 ♀, in Cirad; 9 ♀ 1 ♂, in GDPC) ; San Fratello, Monte Nebrodi , N Casello Volpe , 1150 m, 37.94402°N 14.62579°E, on flowers of A. ramosus , 26.vi.2014 (G. Delvare) (vouchers GDEL 1643 ♀, GDEL 1644 ♀, GDEL 1645 ♀, GDEL 1646 ♀, in CIRAD; 24 ♀, in GDPC) ; San Fratello, Monte Nebrodi , Passo dei Tre , 775 m, 37.97803°N 14.61925°E, on flowers of A. ramosus , 26.vi.2014 (G. Delvare) (1 ♀, in GDPC) ; Vizzini, Monte Iblei , SE Lago Dirillo , contrado Rubalé , 420 m, 37.10778°N 14.72578°E, ex seeds of A. ramosus , 19.vi.2014, adults emerged 07–15.iv.2015 (G. Delvare) (179 ♀ 170 ♂, in GDPC) ; same locality, asphodel and collector, 350 m, 37.12103°N 14.72166°E, ex seeds of A. ramosus , 22.vi.2014, (192 ♀ 250 ♂, in GDPC) ; Sicilia, Palermo, Zingaro Reserve , ex seeds of A. ramosus , 9.v.2005 (R.R. Askew, M.C. Rizzo) (22 ♀ 10 ♂, 9 in RAPC, 2 ♀ in GDPC) ; SPAIN: Huesca, Fraga , 125 m, 41.41892°N 0.10911°E, ex seeds of A. cerasiferus , 15.iii.2010 & 29.iii.2011 (A. Ribes) (34 ♀ 21 ♂ including vouchers GDEL 1486 ♀, GDEL 1488 ♀, in CIRAD; 6 ♀ 3 ♂, in GDPC, 15 in RAPC) ; Bielsa, Col de Larri , Val de Pineta , 1200 m, 42.64507°N 0.212893°E, ex seeds of A. a. delphinensis 6.vii.2007 (R.R. Askew) (9 ♀ 20 ♂, 24 in RAPC) ; on flower spikes of A. a. delphinensis, same date & collector, 19 ♀ 5 ♂, (22 in RAPC) ; Lleida, Aitona , 150 m, 41.48333°N 0.46667°E, ex seeds of A. cerasiferus (A. Ribes) 28.v.2009 (10 ♀ 3 ♂ emerged 30.v–3.vi.2009), 17.vi.2009 (1 ♀ emerged 22.vi.2009, 1 ♂ emerged 19.vi.2009, 6 ♀ emerged about 6.v.2011 (1 ♀, in GDPC), 13.iii.2010 (1 ♀ emerged 23.iv.2010), 15.vi.2010 (2 ♀ 1 ♂ emerged 16–17.vi.2010), 23.vii.2010 (1 ♀ emerged 9.viii.2010), 19.iii.2011 (4 ♀ 4 ♂ emerged 8–24.iv.2011) (voucher GDEL 1472 ♀, in CIRAD; 7 ♀ 1 ♂, in GDPC) ; Tunel Vielha , near northern entrance, 1390 m, 42.67278°N 0.77333°E, ex seeds of A. a. delphinensis, 18.viii.2008, adults emerged 22–29.iv.2010 (A. Ribes) (71 ♀ 59 ♂) (vouchers GDEL 1299 ♀, GDEL 1495 ♀, in CIRAD; 2 ♀, in GDPC; 15 ♀ 6 ♂, 9 in RAPC; 1 ♀ in GDPC) ; same locality and asphodel, 14.vii.2011 (R.R. Askew) 2 ♀ 7 ♂ emerged vii.2011, 3 ♀ 15 ♂ emerged 2012 ; 30.v.2012 same locality, collected on flowers of the plant (vouchers GDEL 1473 ♀, GDEL 1474 ♀, GDEL 1475 ♀, GDEL 1476 ♀, GDEL 1477 ♀, GDEL 1478 ♀, GDEL 1479 ♀, GDEL 1480 ♀, GDEL 1481 ♀, in CIRAD) ; TURKEY: Aegean sea Gökçe Ada Island , Şirinköy village , 62 m, 40.12972°N 25.73806°E, ex seeds of Asphodelus sp., 8.vi.2011 (A. Stojanova) (vouchers GDEL 1489 ♀, GDEL 1490 ♀, in CIRAD; 2 ♂, in GDPC) .

Condition of examined paratypes. Male badly damaged, in three fragments mounted on one card: 1) head with antennae enclosed in pupal cuticle, 2) mesosoma with legs and propodeum missing, 3) petiole and body of gaster. The female paratype from Ibiza that was examined is in good condition.

Description of female paratype examined (figures from other specimens). Length 3.0 mm. Body black except for 2 faint pale spots on anterior face of pronotum, pilosity white; scape (as in Fig. 23D ) blackish; legs with coxae black, knees narrowly testaceous, tibiae with testaceous apices and protibia pale on inner face, tarsi testaceous with last segments infuscate; wing pilosity white, venation testaceous; ovipositor sheath with pale apex.

Head in dorsal view 2.0× as broad as long with frons projecting in an even curve in front of eyes; temples about 0.37× eye length; POL 2.2× OOL, OOL 1.6× OD. Head in front view ( Fig. 22A ) 1.4× as broad as high; height of eye 1.3× malar space; mouth 1.3× as broad as malar space; toruli very slightly nearer anterior ocellus than anterior margin of clypeus; clypeus with a small median depression on anterior margin, radiating carinulae extending almost to level of toruli; gena buccate with strong reticulate sculpture, a seta in each areole, extending to malar sulcus; head in occipital view ( Fig. 22B ) with genal carina strong and postgenal groove complete.

Antenna ( Figs 23 C–D) 11153; scape almost 4× as long as broad, reaching level of middle of anterior ocellus; pedicel plus flagellum 0.9× as long as breadth of head; F1 as broad as pedicel, about 1.5× as long as broad and as long as pedicel plus anellus; following funicle segments of about equal breadth; F5 very slightly longer than broad; clava 2.2× as long as broad, slightly longer than F4 plus F5; sensilla in 2 slightly overlapping transverse rows on F1 and in 2 much overlapping rows (or 1 irregular row) on F2–F5.

Mesosoma in dorsal view 1.6× as long as broad, narrower than head (20:23); pronotum medially shorter than mesoscutum (10:11) and 0.8× length of mesoscutellum; mesoscutellum with base narrower than length of scutelloaxillar suture. Mesosoma in profile 1.3× as long as depth at mesosternal shelf, the mesoscutellum dorsally strongly convex and propodeum sloping at an angle of 80° to the scutello-mesoscutal plane. Ventral shelf of mesepisternum ( Fig. 23D ) developed, about twice as broad as its median length; mesosternum, between depressions that receive the procoxae, elevated as a rounded longitudinal ridge which in profile appears very slightly concave and is untoothed, median spine on transverse carina at anterior margin of mesosternum well-developed. Procoxa ( Fig. 22F ) with an oblique oval scrobe on anterior face. Metatibia with longest spines on dorsal edge about half breadth of tibia. Propodeum with a broad, shallow median depression with fine reticulate sculpture and irregular, carinulae radiating from above the petiolar foramen.

Fore wing lengths of costal cell: marginal vein: stigmal vein: postmarginal vein as 110:20:23:29; stigmal vein ( Figs 23 G–H) weakly curved, expanding gradually into an elongated stigma; basal vein pilose, basal cell closed below in apical half.

Metasoma. Gaster 1.3× as long as rest of body, laterally compressed, in dorsal view 3.4× as long as broad and 0.8× breadth of mesoscutum; petiole transverse. Gaster in lateral view about 2× as long (including ovipositor sheath) as deep, as deep as mesosoma at level of ventral shelf of mesepisternum, ascending at only 45° from petiole to apex of GT2; exposed part of GT4 dorsally almost twice as long as GT3; syntergum not upturned, in dorsal view 1.4× as long as breadth at level of apex of basal emargination; syntergum plus exposed ovipositor sheaths about half as long as metatibia; ovipositor sheaths extending beyond apex of syntergum by about 0.7× length of syntergum.

Description of male paratype examined. Length estimated 1.8 mm.

Head in front view 1.4× as broad as high; radiatingly strigose sculpture on clypeus and lower face weak.

Antenna ( Fig. 22G ) 11152; scape 3.2× as long as broad, expanding gradually from base into a ventral boss, broadest at about three-quarters of length thence narrowing rather abruptly to an apical neck; pedicel plus flagellum 1.3× breadth of head; funicle segments petiolate, bodies of F1 1.5×, F2–F5 about 1.3× as long as deep; clava 3.2× as long as broad; setae on F1 at least as long as body of segment.

Metasoma including petiole estimated at 0.65× combined length of head and mesosoma; petiole about 0.35× length of body of gaster, its dorsal surface flat, 1.6× as long as broad with an anterior transverse carina; body of gaster 2× as long as deep, in profile GT1 rising at an angle of about 45°, dorsal surface after GT1 and all ventral surface relatively flat to apex of GT4, the two surfaces converging only slightly; aedeagus directed posteriorly.

Comment. Two males (and two females) from the Hedqvist collection now in BMNH are labelled ‘standing over Eurytoma asphodeli in Hedqvist coll. BMNH(E) 2011-27 ’ and, in Hedqvist’s handwriting, ‘ Portugal, Cruz, 10/5[?]/1988 K.J. Hedqvist’. The males are quite large, about 3.5 mm long, each with the base of the antennal scape pale and with a relatively strongly protruding swelling, front leg including coxa and mesofemur entirely reddish, F1 with setae hardly half the length of the body of the segment and the gaster short and rounded with the aedeagus directed ventrally. The females also have scapes with pale bases and procoxae partly yellowish. These are characters of the red-legged summer form (see below) of E. asphodeli . It is testimony to Hedqvist’s perception that he correctly recognized this Portuguese material as conspecific with his type series of E. asphodeli from the Balearic Islands, especially considering that its capture was apparently unlinked to Asphodelus ( Hedqvist 1976) .

Variation. Eurytoma asphodeli is subject to considerable intraspecific morphological variation. The relatively small and dark typical form has been reared from seeds of A. a. albus from France (Vienne, Dordogne) and from A. albus delphinensis from France (Hautes-Pyrénées) and Spain (Lleida, Huesca) (see Table 1 for details), as well as from A. ramosus (= microcarpus) from Spain (Ibiza), Corse and Sicilia and A. cerasiferus ( France, Bouches-du- Rhône and Spain, Lleida).

The setation of the fore wing of all phenotypes is white with the exception of the form found in Corse, the females of which have partly or entirely dark setation.

A very distinctive phenotype ( Figs 23A, C, E, G, I ) is characterized by large size (body length of females 4.7–5.2 mm), mainly reddish yellow antennal scape ( Fig. 23C , compare Fig. 23D ), mainly reddish pro- and mesocoxae ( Fig. 22F ), reduced dark markings on femora, gaster of female in lateral view 1.75× as long as deep and 1.25× as deep as mesosoma with combined length of syntergum and exposed ovipositor sheath 0.65× length of metatibia ( Fig. 23I ). This phenotype emerged in early summer from the large first year seeds of A. cerasiferus in Spain (Lleida), the larvae not having overwintered. Adults emerging in spring from overwintering larvae in the same sample were of the typical form although, in comparison to overwintering specimens reared from other species of asphodel, larger and with a deeper and more laterally compressed gaster, more strongly upturned ovipositor sheaths and more transverse head.

A phenotype somewhat intermediate between the typical form and the red-legged variant was reared from A. ramosus in Sicilia and Asphodelus sp. in Turkey (Muğla). It has scape and much of the upper legs yellowish to reddish, and the female gaster is high, but the syntergum plus exposed ovipositor sheaths are not quite half as long as the metatibia.

The different morphological forms described above are all referable to E. asphodeli and not to separate species. This has been clearly demonstrated by DNA analysis in conjunction with a morphometric examination, and will be discussed elsewhere (Château et al. in prep.).

Diagnosis. Because of the extreme variation in the reared specimens, it is quite difficult to find a diagnosis that applies to all.

Recognition. The recognition of E. asphodeli within the robusta species group of Eurytoma in its restricted concept ( Lotfalizadeh et al. 2007; not Zerova & Seryogina 2006 or Zerova 2010) is challenging because of the intraspecific variation. The non-hibernating females reared from A. cerasiferus approach the condition of E. robusta Mayr ; but they differ from that species in having a more transverse head in dorsal view and largely testaceous legs. The other phenotypes approach the condition of E. strigifrons but with different notauli and relatively longer gaster.

Distribution ( Fig. 36 ). Widely distributed in Europe, it was found at almost all sites where asphodels, except A. fistulosus , were present. It is also quoted from Israel ( Zerova et al. 2005).

Biology. Trophic relationships ( Figs 39–42 ). Hedqvist (1976) implies that E. asphodeli is phytophagous but the available data relative to the biology of other Eurytoma belonging to the robusta group showed that most species are strictly entomophagous ( Claridge 1961; Delvare 1988; Domenichini 2002; Zwölfer et al. 2007). In addition, a few gall dissections provided no evidence that E. strigifrons larvae in seed bract galls of Isocolus scabiosae (Giraud, 1859) (= rogenhoferi Wachtl, 1880) ( Hymenoptera , Cynipidae ) in Centaurea scabiosa L. in England consume any plant tissue. We have found solitary fully grown larvae of E. asphodeli in seeds of A. a. albus and A. a. delphinensis together with the larval remains of B. abscedus and B. ribesi on which they had been feeding externally. E. asphodeli may, however, also feed on seed tissues after the insect host has been consumed. This latter possibility is based on the rather similar sizes of E. asphodeli and the Bruchophagus host species, and on the contemporaneous activity of the adult insects.

Phenology. The time taken by E. asphodeli to complete its development from egg to adult is variable. Some adults emerge from first year asphodel seeds collected as early as May ( Fig. 47 ), evidently the offspring of an early spring emergence. Such adults chew an exit hole from the seed and through the fruit wall when the fruit is still attached to the flower spike; later in the year adult Eurytoma mostly emerge directly from seeds that have fallen to the ground. In contrast to Bruchophagus , there are at least two flight periods in a year. Most individuals, however, pass one or two winters as larvae in the seeds, the emergence of E. asphodeli from our samples being as follows:

Number of winters in seeds 0 1 2 3

A. albus albus 2 478 47 0 A. albus delphinensis 17 38 138 0 A. cerasiferus 29 68 6 0 A. ramosus 28 4 0 0

The number of individuals recorded as not overwintering will partly depend on the date of sampling. The proportion of emergences of insects destined not to overwinter will decrease with the lateness of sample collection. It seems that a large proportion of E. asphodeli in A. cerasiferus seeds, and more especially in A. ramosus seeds, do not overwinter (and are of the red-legged form), whilst those in A. albus delphinensis seeds mostly overwinter twice. The data are distorted, however, because some collections of A. cerasiferus fruits were made early in spring after overwintering once.

E. asphodeli is protandrous, the peak of male emergence preceding that of females by about three days. Considering emergences from a large collection of A. albus albus fruits made in Dordogne, France on 19.vi.2011 and overwintered outdoors, the emergence dates of all E. asphodeli in 2012 were as follows:

Emergence dates 8– 10 April 11–13 April 14– 16 April 17–20 April

Numbers of Males 124 91 2 0

Numbers of Females 0 127 88 7

Both sexes of E. asphodeli were found on 6.vii.2007 walking on fruits and flowers of A. a. delphinensis near Bielsa in the Spanish Pyrénées. A collected sample comprised 18 ♀ and 5 ♂ together with 11 B. ribesi and 3 Pteromalus tethys . Also on A. a. delphinensis, over thirty females were seen on green fruits on 14.vii 2010 in the Val d’Ossoue, French Pyrénées, and several red-legged males were collected on blackening fruits on 31.vii.2013 on the Col du Portillon, Lleida, Spain. Such large aggregations of E. asphodeli have not been found on A. a. albus , although in Dordogne, France females were observed on this plant on 6.v.2011, 2.vi.2012 and 16.vii.2009 with ovipositors penetrating mature green fruits.

The large red-legged summer seasonal form of E. asphodeli associated with A. cerasiferus emerged within days from fresh fruits collected in Spain (Lleida, Aitona) at the end of May and in June, and was found in the field on A. cerasiferus fruits at about this time. On 28.v.2012 totals of 19 females and 21 males were found. These appeared to have developed in basal fruits on the host plant racemes, some of the older blackened fruits having exit holes, and females were on younger green and more apically situated fruits, seemingly ovipositing or in mating pairs, whilst males were mostly on the older fruits, perhaps awaiting the emergence of females. Dry, brown fruits of A. cerasiferus collected at Aitona during winter produced only the typical dark form of A. asphodeli the following spring (April). This population of E. asphodeli is therefore at least partly bivoltine with the first adult generation emerging in spring from overwintered fruits and being typical in appearance. Most of the progeny of this darkcoloured spring generation emerge during summer as the red-legged form, although occasional dark-legged typical individuals may emerge with them. All specimens emerging from A. cerasiferus after overwintering are attributed to typical E. asphodeli .

Eurytoma asphodeli attacking A. ramosus seeds in Sicilia have a similar life cycle to that described above for the population in A. cerasiferus seeds, but the form emerging in early summer without overwintering has less bright colouration and is not much larger than insects that have overwintered.

Larval morphology. Larvae of E. asphodeli are easily distinguished from those of B. abscedus and B. ribesi . The fully grown Eurytoma larva is smaller (about 2.6 mm long) than that of Bruchophagus , noticeably more mobile and with considerably longer setae on head and body ( Fig. 6D ). The head of the larva of E. asphodeli ( Fig. 6F ) bears four pairs of long setae, each seta longer than half the distance separating the antennae, and the mandibles ( Fig. 6E ) are bidentate with the principal tooth more than twice as long as the inner tooth.

Figures

FIGURES 6A–F. Morphology of larval instars. A–C. Bruchophagus abscedus. A, fully grown larva. B, mouthparts. C, head in frontal view. D–F. Eurytoma asphodeli. D, fully grown larva. E, mouthparts. F, head in frontal view.

FIGURES 22A–G. Eurytoma asphodeli. A–F, ♀. G, ♂. A, head in frontal view. B, same in posterior view. C, same in anterolateral view. D, mesopleuron in anterolateral view. E, propodeum. F, procoxa in anterior view. G, antenna. Abbreviations. epm, epicnemium; epmc, epicnemial carina; epst2-vf, ventral shelf of mesepisternum; pgl, postgenal lamina; ppt-mvt, medioventral tooth of prepectal belt. A, B & C, ex Asphodelus sp., Şirinköy village, Gökçe Ada Island, Aegean Sea, Turkey. D, on collected on spike of A. cerasiferus, Lapanouse-de-Cernon, Aveyron, France. E & G, ex A. albus delphinensis from Vielha, Lleida, Spain. F, ex A. cerasiferus, Aitona, Lleida, Spain.

FIGURES 23A–J. Eurytoma asphodeli, ♀. A, C, E, G & I, aestivalis phenotype, ex A. cerasiferus, non-overwintering generation, Aitona, Lleida, Spain. B, D, F, H & J, corsicae phenotype, ex A. ramosus, overwintering generation, Aléria, Corse, France. A–B, ventral view of mesepisternum. C–D, antenna. E–F, propodeum. G–H, fore wing. I–J, metasoma in lateral view.

FIGURE 36. Distribution map of Eurytoma asphodeli, E. genale and E. pistaciae (only reared from asphodels)

FIGURE 39. Trophic web of A. albus albus. Horizontal scale bars proportional to the numbers of reared specimens examined that are indicated in the bars. Arrows pointing out the direction of the trophic relationships: from phytophagous Bruchophagus spp. to the asphodel and from parasitoids to chalcid hosts.

FIGURE 40. Trophic web of A. albus delphinensis. Horizontal scale bars proportional to the numbers of reared specimens examined that are indicated in the bars. Arrows pointing out the direction of the trophic relationships: from phytophagous Bruchophagus spp. to the asphodel and from parasitoids to chalcid hosts.

FIGURE 41. Trophic web of A. cerasiferus. Horizontal scale bars proportional to the numbers of reared specimens examined that are indicated in the bars. Arrows pointing out the direction of the trophic relationships: from phytophagous Bruchophagus spp. to the asphodel and from parasitoids to chalcid hosts.The estimated number of Bruchophagus abscedus parasitized by P. depilata was estimated considering that a mean of 10 Puklina emerge from a larva of Bruchophagus.

FIGURE 42. Trophic web of A. ramosus. Horizontal scale bars proportional to the numbers of reared specimens examined that are indicated in the bars. Arrows pointing out the direction of the trophic relationships: from phytophagous Bruchophagus spp. to the asphodel and from parasitoids to chalcid hosts. The estimated number of Bruchophagus abscedus parasitized by P. depilata was estimated considering that a mean of 10 Puklina emerge from a larva of Bruchophagus.

FIGURE 47. Phenology of the two sexes of Eurytoma asphodeli according to the asphodel.