Genus Ecrizotes Förster, 1861

Figs 1–13

Ecrizotes Förster, 1861: 33; type species by original monotypy Ecrizotes monticola Förster, 1861 .

Henicetrus Thomson, 1876: 188, 190; type species by subsequent designation Henicetrus annellus Thomson, 1876 in Gahan & Fagan 1923: 70; subjective synonym of Ecrizotes in Ashmead 1904: 377.

Spathopus Ashmead, 1904: 272; type species by original designation or monotypy Spathopus anomalipes Ashmead, 1904; syn. nov.

Ecrizotomorpha Mani, 1939b: 537; type species by original monotypy Ecrizotomorpha taskhiri Mani, 1939; syn. nov.

Liaoella Xiao & Huang, 1999; type species by original designation or monotypy Liaoella alternativa Xiao & Huang, 1999; subjective synonym of Ecrizotomorpha in Huang & Xiao 2005: 215–216; syn. nov.

Diagnosis

Body dark, with at most faint metallic reflections, mostly on head (Figs 1–13). Female antenna with 2 microscopic, 1–2 anelliform, and 3–5 large flagellomeres before the 4-segmented clava (with 3 large clavomeres plus a ‘terminal button’) (Figs 1C, 2E, 3E, 4E, 5E, 6C, 7E, 8C, 9D, 10D, 11C, 12F, 13F). Male antenna with 1 microscopic and 6 large flagellomeres before the 3-segmented clava (with 2 large clavomeres and a ‘terminal button’) (Figs 4F, 7F, 9E). Propodeum without plicae or median carina (e.g., Figs 1E, 10E, 13E). Fore wing with parastigmal hyaline break (Figs 1F, 3H, 4H, 5E, 6G, 7F, 8G, 9F, 10C, 11E, 12F, 13C). Female with hypopygium from almost reaching to surpassing the apical tergite of gaster (e.g., Figs 5H, 10F, 11F, 12E). Male sometimes with enlarged tibiae (Figs 5B, 9B).

Etymology

The name Ecrizotes has a Greek origin and comes from the words ‘εκ’ meaning ‘from’ or ‘out’, and ‘ρίζα’ meaning root. It can be translated as ‘the one that takes the root out’ or metaphorically ‘the one that destroys something’; it is of masculine gender.

Description

Female

Body with faint metallic reflections, these most obvious on the head (Figs 1–13). Body sculpture always delicate, alutaceous (e.g., Figs 1E, 3F, 6D, 8E, 10E, 11E, 12C). Setation sparse, setae rather long, piliferous punctures occasionally visible on the upper face (e.g., Figs 10B, 11B, 12B).

Head in frontal view subcircular (e.g., Figs 1B, 2C, 3B, 4C, 5C, 8B, 10B, 11B, 12B). Vertex sometimes conspicuously protruding between posterior ocelli (Figs 5C, 12B). Gena evenly round (Figs 8B, 10B) to conspicuously buccate (Figs 1B, 2C, 3B, 4C, 5C, 6B, 7C, 9C, 12B), not hollowed at mouth corner (e.g., Fig. 6B). Clypeal margin from weakly convex (Figs 4C, 6B, 7D, 8D, 9C, 13D) to strongly convex (Figs 2D, 3D, 5D, 10C, 11D) or acute (Figs 1D, 12D). Tentorial pits invisible. Scrobal depression short and shallow, with weak inter-torular ridge (e.g., Figs 8D, 10B, 12D). Malar sulcus present (e.g., Figs 10A, 12F). Eyes in frontal view slightly diverging in lower part (e.g., Figs 1B, 5C, 8B, 10B, 11B). Temples short and strongly converging in dorsal view of the head (Figs 5G, 8E). Occiput without carina. Antennal inserted much lower than LOL (e.g., Figs 1B, 3B, 4C, 5C 7C, 8B, 10B, 11B, 12B, 13B), with 2 microscopic, 1–2 anelliform (one often intercalated between two larger ones), and 3–5 large flagellomeres before the 4-segmented clava (with 3 large clavomeres plus a ‘terminal button’); antennal clava symmetric, without conspicuous area of microsetation, distal end rounded (Figs 1C, 2E, 3E, 4E, 5E, 6C, 7E, 8C, 9D, 10D, 11C, 12F, 13F). Mandibles usually at least slightly falcate (Figs 1A, 7D, 10C, 12F, 13D), in the few observable cases with the formula 4:3.

Mesosoma dorsally convex (Figs 3A, 4A, 5A, 6A, 8A, 10A, 11A) or almost flat (Figs 1A, 2A, 7A, 9A, 12A, 13A). Pronotum often long, with large diverging shoulders (Figs 1E, 2G, 5G, 10E, 11E, 12C, 13E), occasionally shorter, with smaller shoulders (Figs 3F, 4G, 6D, 7H, 8E, 9F, 10F). Pronotum evenly sloping, without transverse carina (e.g., Fig. 10A). Mesoscutum shorter than mesoscutellum; notauli complete and deep (Figs 1E, 2G, 3F, 4G, 5G, 6D, 7H, 8E, 9F, 10E, 11E, 12C, 13E). Axillae only slightly advanced. Mesoscutellum from convex to almost flat; frenum mostly indistinct. Dorsellum subhorizontal, smooth, semicircular (e.g., Figs 1E, 3F, 4G, 5G, 7H, 8E, 10E, 11E, 12C, 13E). Propodeum much shorter than mesoscutellum, smooth or uniformly and superficially sculptured; plicae and median carina absent; hind corners not prominent and not sharp; spiracles almost touching posterior margin of metanotum (e.g., Figs 1E, 4G, 6D, 10E, 13E). Hind coxa dorsally bare. Fore wing hyaline (Figs 1F, 2H, 3C, 4H, 5H, 6E, 7G, 8F, 9G, 10F, 11F, 12E, 13C), extensively setose, fringe present; veins slender or slightly thickened; parastigma with hyaline break; stigmal vein much shorter than marginal vein, stigma moderately capitate; postmarginal vein much shorter than marginal vein and only slightly longer than stigmal vein.

Gaster at least slightly compressed laterally, occasionally strongly so (Figs 3A, 4A, 6A, 7A, 8A, 9A). Petiole inconspicuous. Gastral tergites subequal in length, their posterior margin straight. Hypopygium from almost reaching to surpassing the apical tergite of gaster, its tip with (e.g., Figs 3G, 6F) or without a small incision (e.g., Fig. 7I).

Male

Similar to female (Figs 2B, 4B, 5B, 7B, 9B), except mainly for the differential features given in the diagnosis.

Distribution

The genus Ecrizotes is newly recorded in Africa (six species). Various species are present on all continents except for South America, Australia and Antarctica: Europe (seven species), Asia (five species) and North America (one species).

Hosts

Bouček (1964: 258) considered that Ecrizotes hofferi (Bouček, 1964) comb. nov. “probably develops as a parasite of some Cecidomyids”. Ecrizotes taskhiri (Mani, 1939) comb. nov. was cited as a hyperparasitoid of Dasineura lini (Barnes, 1936) ( Diptera: Cecidomyiidae) in India (Pruthi et al. 1940).

Taxonomic comments

Of the four described species of Spathopus, males are known only for S. anomalipes and S. hofferi (Fig. 4B). In both species all tibiae are strongly swollen; this feature only characterizes the males and makes them easily discernible from both conspecific females and males of Ecrizotes, which have normal tibiae.According to Mani (1939:538), the male of Ecrizotomorpha taskhiri has the hind tibiae “[…]broad, compressed laterally”. The males of the other two species classified in Ecrizotomorpha are unknown. This feature was used by both Graham (1969) and Bouček & Rasplus (1991) to separate the Ecrizotes males from the Spathopus males. However, in our examined material from Africa, we found several males that have slender fore and mid tibiae (as in Ecrizotes) and inflated hind tibiae (as in Spathopus) (Fig. 6B). Furthermore, the strong sexual dimorphism present in some species and not in others is not a reliable indication of their generic distinctiveness, as shown in the related genus Macroglenes Westwood, 1832 ( Pirenidae: Pireninae): the males of some species (e.g., M. gibsoni Mitroiu, 2010) display abnormally large eyes, the males of other species have a strongly inflated antennal scape (e.g., M. bouceki (Graham, 1969)), while the males of other species (e.g., M. paludum (Graham, 1969)) do not have any of these unusual characters.

Regarding the separation of the females of Ecrizotes and Spathopus, according to Graham (1969) and Bouček & Rasplus (1991), the main differences are summarized in Table 1.

The number and development of fore tibial spines (character 1) is difficult to assess and it is virtually impossible to separate the two instances. Characters 2 and 3 are also variable inside many chalcid genera. Character 4 was considered relevant by Bouček & Rasplus (1991), but not Graham (1969) and indeed Ecrizotes monticola and E. caudatus have a posteriorly constricted pronotum, although less long (Fig. 10F). Character 5 refers to the head thickness in anteroposterior axis, which is a variable character inside many chalcid genera. The ventral clypeal margin is more or less convex in both Spathopus and Ecrizotes and has a continuous degree of projection, ranging from only slightly convex to sharp (Figs 1D, 2D, 3D, 4D, 5B, 6D, 7D, 9C, 10D, 11D, 12D, 13D).

According to Mani (1939: 537), Ecrizotomorpha has affinities with both Ecrizotes (“in venation and moniliform antennae of male”) and Spathopus (“in its stout and compressed hind tibiae and short stouter fore tibiae”), but differs from Ecrizotes in “the pubescent eyes, absence of ring-joints and triarticulate maxillary palpi” and from Spathopus in “the longer hind tibiae, pubescent eyes, absence of ring-joints and clypeus produced obtusely but not triangularly”. All of the above differences do not hold upon a closer examination of the three genera involved: (1) the “ring-joints” refer to the microscopic flagellomeres, which are not absent in Ecrizotomorpha but can be seen (at least the second) if enough magnification is used; (2) all species of these genera have some eye pubescence; (3) the ventral clypeal margin shows various degrees of convexity, ranging from broadly convex to narrowly pointed (see above); (4) the maxillary palpus is triarticulate at least in Spathopus hofferi; (5) the length of hind tibia is irrelevant as it can be variable inside a given genus.

Furthermore, the only character that separates females of Ecrizotes from those of Gastrancistrus Westwood, 1833 is the position of the hypopygium, which is situated at the same level with the distal tip of the apical tergite or slightly beyond it in Ecrizotes and clearly anterior of it (mostly anterior to middle of gaster) in Gastrancistrus . Males of Ecrizotes are indistinguishable from those of Gastrancistrus, except for those having enlarged hind tibiae. However, Gastrancistrus is a hugely diverse genus and until its revision other taxonomic changes are postponed. Melancistrus Graham, 1969, which also has the hypopygium near the tip of gaster, differs in having a translucid median projection (the mucro) extending posteriorly from the hypopygium, and a transverse propodeal carina; it could also belong here but unavailability of material prevented further investigations. Afrothopus Mitroiu, 2024 also has similarities with Ecrizotes, Gastrancistrus and Spathopus but differs in several key characters (see Discussion and Mitroiu et al. 2024).

In conclusion, there are no reliable characters for the separation of Ecrizotes, Ecrizotomorpha and Spathopus and consequently Ecrizotomorpha syn. nov., Liaoella syn. nov. (previously synonymized with Ecrizotomorpha), and Spathopus syn. nov. are regarded as junior synonyms of Ecrizotes, with the following new combinations: Ecrizotes alternativa (Xiao & Huang, 1999) comb. nov.; E. anomalipes (Ashmead, 1904) comb. nov.; E. hofferi (Bouček, 1964) comb. nov.; E. montanus (Huggert, 1976) comb. nov.; E. nasalis (Springate & Noyes, 1990) comb. nov.; E. taskhiri (Mani, 1939) comb. nov.; and E. tenkasiensis (Jamal Ahmad & Shafee, 1993) comb. nov.

Key to Afrotropical and West-Palaearctic species of Ecrizotes (females)

1. Antenna with at least proximal funiculars longer than wide or quadrate and none anelliform (Figs 4E, 8C); Palaearctic species .................................................................................................................... 2

– Antenna with all funiculars wider than long, usually at least some anelliform (Figs 1C, 2E, 3E, 5E, 6C, 7E, 9D, 10D, 11C, 12F, 13F); Afrotropical and Palaearctic species .......................................... 3

2. Funiculars distinctly longer than wide, Fu1 length about twice width (Fig. 8C); clava at most as long as combined length of the three preceding funiculars; scape in lateral view about 6–7 × as long as wide ..................................................................................................... E. longicornis (Walker, 1848)

– Funiculars at most slightly longer than wide or quadrate, Fu1 length about 1.1× width (Fig. 4E); clava length 1.25 × combined length of the three preceding funiculars; scape in lateral view at most about 5.5× as long as wide ................................................................. E. filicornis (Thomson, 1876)

3. Gaster length (without ovipositor sheath) 1.15–1.40 × combined length of head and mesosoma, strongly compressed laterally (Figs 3A, 6A, 7A, 9A); ovipositor sheath length at least 0.5 × length of hind tibia .................................................................................................................................... 4

– Gaster length (without ovipositor sheath) at most equal to combined length of head and mesosoma, at most moderately compressed laterally (Figs 1A, 2A, 5A, 10A, 11A, 12A, 13A); ovipositor sheath at most 0.4 × length of hind tibia ...................................................................................................... 7

4. Tip of hypopygium incised (Figs 3G, 6F); Fu3 smaller than Fu2 and Fu4, but not anelliform (Figs 3E, 6C); Afrotropical and Palaearctic species ....................................................................... 5

– Tip of hypopygium not incised (e.g., Fig. 7I); Fu3 variable (Figs 7E, 9D); Afrotropical species .... 6

5. Ovipositor sheath about 0.9× as long as hind tibia; hind leg slender, tibia length about 8 × width; tibiae extensively yellowish brown (Fig. 6A); Afrotropical species ........ E. incisus Mitroiu sp. nov.

– Ovipositor sheath about 0.5–0.6 × as long as hind tibia; hind leg stouter, tibia length about 5 × width; legs entirely dark brown (Fig. 3A); Palaearctic species ..................... E. caudatus (Thomson, 1876)

6. Ovipositor sheath very long, about 1.2× as long as hind tibia (Fig. 7A); Fu3 only slightly smaller than Fu2 and Fu4 (Fig. 7E) ................................................................ E. longicauda Mitroiu sp. nov.

– Ovipositor sheath shorter, about 0.5–0.6 × as long as hind tibia (Fig. 9A); Fu3 anelliform (Fig. 9D) ................................................................................................................... E. longus Mitroiu sp. nov.

7. Ventral clypeal margin strongly protruding and acute (Fig. 1D); Afrotropical species ...................... ....................................................................................................................... E. acer Mitroiu sp. nov.

– Ventral clypeal margin more or less evenly curved (Figs 2C, 5D, 10C, 11D, 13D), if rarely almost acute ( E. nasalis), then less strongly protruding (Fig. 12D); Afrotropical and Palaearctic species … 8

8. Antenna with Fu3 not anelliform, not or only slightly smaller than Fu2 or Fu4 (Figs 5E, 10D, 12F); Palaearctic species ............................................................................................................................ 9

– Antenna with Fu3 conspicuously smaller than Fu2 or Fu4, usually anelliform (Figs 2E, 11C, 13F); Afrotropical and Palaearctic species ................................................................................................11

9. Ventral clypeal margin acute (Fig. 12D) .............. E. nasalis (Springate & Noyes, 1990) comb. nov.

– Ventral clypeal margin evenly convex (Figs 5D, 10C) ................................................................... 10

10. Head in frontal view with vertex less convex between posterior ocelli and gena evenly rounded (Fig. 10B) .......................................................................... E. montanus (Huggert, 1976) comb. nov.

– Head in frontal view with vertex more strongly convex between posterior ocelli and gena buccate (Fig. 5C) .................................................................................. E. hofferi (Bouček, 1964) comb. nov.

11. Ventral clypeal margin weakly convex (Fig. 13D); MV about 2.4× SV; scape, pedicel, and legs except basal part of femora yellowish (Fig. 13A); Afrotropical species ............................................ ................................................................................................................ E. rovumae Mitroiu sp. nov.

– Ventral clypeal margin strongly convex (Figs 2D, 11D); MV about 2.5–3.0× SV; scape, pedicel and legs more extensively dark, femora completely dark (Figs 2A, 11A); Afrotropical and Palaearctic species ............................................................................................................................................. 12

12. Mesosoma dorsally convex (Fig. 11A); pronotum shorter than mesoscutum (Fig. 11E); Fu3 short but clearly visible (Fig. 11C); hind tibia length 5.0–5.2 × width; MV 2.5–2.9× SV; ovipositor sheath length 0.3–0.4 × length of hind tibia; Palaearctic species ........................ E. monticola Förster, 1861

– Mesosoma dorsally almost flat (Fig. 2A); pronotum about as long as mesoscutum (Fig. 2G); Fu3 hardly distinct (Fig. 2E); hind tibia length almost 7× width; MV about 3 × SV; ovipositor sheath length about 0.01 × length of hind tibia; Afrotropical species ........... E. brevicauda Mitroiu sp. nov.