Revision of Eudorylas Aczél, 1940 (Diptera, Pipunculidae) in the Middle East, with the description of four new species

Abstract Background The Middle Eastern species of Eudorylas Aczél, 1940 are revised through an integrative taxonomic approach by combining morphological and sequence data from the mitochondrial COI barcoding gene. Four new species of the genus Eudorylas are described, males and females of three species are associated, DNA sequence data of 11 Middle Eastern Eudorylas species are provided and 15 additional species are discussed. To facilitate their recognition, we provide diagnoses, descriptions, an identification key and distributional maps for all species. New information The following new species are described from the Middle East: E.avis Motamedinia & Skevington sp. n., E.bihamatus Motamedinia & Skevington sp. n., E.corniculans Motamedinia & Skevington sp. n., E.nasicus Motamedinia & Skevington sp. n.


Introduction
Eudorylas belongs to the tribe Eudorylini (Diptera, Pipunculidae) and is one of the most species-rich and cosmopolitan genera of Pipunculidae in the world with 416 valid species recognised (Skevington 2020). The first Eudorylas species was described in the 19th century as Pipunculus fuscipes (Zetterstedt, 1844). Aczél (1940) established Eudorylas and designated Cephalops opacus Fallén, 1816 as the type species. Without studying type material, he transferred what he believed to be the relevant species from Becker (1897), Cresson (1911) and Sack (1935) to Eudorylas. The absence of propleural setae is one of the diagnostic characters of Eudorylas and overlooking or misinterpreting this caused serious instability in early attempts to place species in the genus. The original type species actually had a propleural fan of hairs and should never have been included in Eudorylas. Kuznetzov (1995) was the first to report on this and complicated things by synonymising Eudorylas with Microcephalops De Meyer, 1989 and introducing Neodorylas as a replacement name for Eudorylas (designating Pipunculus fuscipes Zetterstedt, 1844 as the type species). This action changed the generic name for over 30% of the world's pipunculids. De Meyer and Skevington (2001) appealed to the Commission of Zoological Nomenclature and their proposal to conserve the name Eudorylas by designating a new type species (Pipunculus fuscipes Zetterstedt, 1844) was accepted. Neodorylas became a junior objective synonym of Eudorylas following this and no generic upheaval occurred. The phylogenetic relationships of Eudorylini were studied by Skevington and Yeates (2001), who re-diagnosed the genus Eudorylas and proposed Metadorylas Rafael, 1987 as a synonym. Most recently, revision and description of several new species are provided in Australian (Skevington 2002a), Palaearctic and Oriental , Kehlmaier 2005b), Afrotropical (Földvári 2003, Földvári 2013 regions. The Eudorylas fauna in the Middle East (here defined to include Bahrain, Cyprus, Iran, Iraq, Israel, Jordan, Kuwait, Lebanon, Oman, Palestine, Qatar, Saudi Arabia, Syria, Turkey, United Arab Emirates and Yemen) currently comprises 15 species and now four are added, two of which (E. bipertitus Kehlmaier, 2005 and E. flavicrus De Meyer, 1995) are endemic (De Meyer 1995;Kehlmaier 2005b).
The aim of this study is to shed light on the species of Eudorylas occurring in the Middle East. To achieve this, we mainly focused on characters of male genitalia and sequence data obtained from the mitochondrial COI barcoding region. This paper presents information on taxonomy and includes distributional maps, diagnoses, photo illustrations of important morphological characters and an identification key for species occurring in the Middle East.

Taxonomic description
Male genitalia of Eudorylas provide the only absolute characters for secure species identification. Females have only been included in the type material when the DNA data match or geographic overlap with males is unequivocal. Genitalia preparations were made by separating the apical portion of the abdomen and heating in lactic acid (85%) at 100°C for 60-240 minutes and then moving the genitalia into a drop of glycerine on a microscope slide. Potassium hydroxide (KOH) was used for terminalia that were very darkly pigmented or that were to be used for photography. For this, terminalia were treated with 10% KOH at 100°C for 10-30 minutes then immersed in glacial acetic acid for 5 minutes to buffer the reaction and stop the clearing. Terminalia were then washed in ethanol before being placed in glycerine. Following clearing, dissection involved separating syntergosternite 8 and the epandrium from the remainder of the abdomen. After examination of the genitalia in glycerine, the dissections were transferred to microvials filled by glycerine and pinned below the specimen.
External characters were imaged using a Leica DFC450 module fitted on a Leica M205C stereomicroscope using a 0.6x lens. Final images were merged using the image-stacking software Zerene Stacker (Littlefield 2018). Images of the genitalia were taken using a Leica DM5500B microscope, equipped with a Leica DMC4500 module connected to a personal computer running the Leica Application Suite software (https://www.leicamicrosystems.com), which includes an Auto-Montage module that combines multiple layers of photographs into a single fully-focused image. All photos were subsequently modified using Adobe Photoshop CS3 imaging software for mounting photos in the plates.
Specimens examined are based on material deposited in the following collections: CNC = Canadian National Collection of Insects, Arachnids and Nematodes, Ottawa, Canada; HMIM = the Hayk Mirzayans Insect Museum, Tehran, Iran; TAU = Tel Aviv University, Israel. All specimens are labelled with a unique reference number from the CNC database (e.g. Jeff_Skevington_Specimen12345 or CNC_Diptera12345, abbreviated as JSS12345 and CNCD12345, respectively) and can be accessed at https://cnc.agr.gc.ca/. Species are presented in alphabetical order. SimpleMappr (Shorthouse 2010) was used to create the species distribution map. Morphological terminology used is based on Skevington (2002b) and . LDP:LPP = ratio of length of distal part of piercer to length of its proximal part (viewed laterally).

DNA extraction, PCR amplification and sequencing
Total genomic DNA was extracted either from two legs or from whole specimens (dried or in alcohol) using the DNeasy Blood and Tissue Kit (Qiagen Inc., Santa Clara, CA, USA) following the manufacturer's protocol. Following extraction, specimens were critical-point dried and deposited as vouchers in the CNC.
For DNA barcoding, a 658 bp fragment of the 5' end of the mitochondrial coding gene cytochrome oxidase subunit I (COI) was amplified using the primer pair LCO1490 and COI-Dipt-2183R (Gibson et al. 2011). In some cases, initial attempts to amplify the full COI barcode failed, presumably due to the degradation of the DNA. In these cases, a COI minibarcode protocol was employed (Motamedinia et al. 2019) in order to amplify a 214 bp fragment (COI-Fx-C), located at the 3'-end of the COI barcode region, for species identification. In the case of putative new species, efforts were made to amplify the 5' and middle COI mini-barcode fragments (COI-Fx-A and COI-Fx-B, respectively) that, when combined, provide a complete COI barcode sequence. Oligonucleotides (primers) used in this study, are listed in Table 1 Cytochrome c oxidase subunit I mitochondrial gene primers.

Notes
Our single DNA barcode of Eudorylas auctus from Germany (JSS15405) overlaps with barcodes of E. obscurus from France (CNC464954) and E. longifrons from Iran (GB: L T671752). The genitalia of these species are different, so this is likely just a case of incomplete lineage sorting due to ancestral hybridisation or the fact that these are young species whose barcodes have not yet diverged, as seen in many other taxa (e.g. Skevington 2005, Skevington et al. 2007, Young et al. 2016. It does raise the possibility that these three taxa are part of a single variable species with polymorphic genitalia. Future work should explore population genetics within this cluster of species, perhaps using the rapidly evolving marker, ITS2.  Eudorylas species distribution in the Middle East.

Description
Male ( Fig. 2A 3A). Surstyli asymmetrical, right larger than left one. Left surstylus rather rectangular-shaped. Base of left surstylus wider than the right one. Right surstylus with an inner finger-like projection curved towards left surstylus, left surstylus with a projection at apex (Fig. 3A). Genital capsule in ventral view: gonopods unequal, right is longer than the left one (Fig. 3B). Genital capsule in lateral view: both surstyli with a finger-like projection at apices, right surstylus with shorter finger-like projection than left one, base of right surstylus broader than left one (Fig. 3D, E). Phallus trifid; phallic guide strongly broadened, bent shortly before apex with two ventrally feather-like projections and apically with a small projecting hook (Fig. 3D, E); hypandrial apodeme extended (Fig. 3D, E). Ejaculatory apodeme spade-shaped (Fig.  3C).

Diagnosis
This species can be distinguished by the shape of the surstyli in dorsal view; base of left surstylus broader than the right one, right surstylus apically with inner long fingerlike projection curved towards left one and small outer finger-like projection (

Etymology
The specific epithet is derived from the Latin avis which means bird and refers to the similarity between the shape of the phallic guide apically in lateral view to that of a bird.

Notes
Based on DNA barcodes, this species is closest to one or more species from South Africa (2.09-2.45% pairwise divergence). Unidentified female specimens from Yemen (CNCD6818) and Angola (CNC395962) are sufficiently different that they are not likely the same species (4.21% and 3.14%, respectively).  Fig. 4 A, B). Body length (excluding antennae): 3.8-4.1 mm (n = 6). Head. Scape black, pedicel and arista dark brown, pedicel with a pair of short upper bristles, flagellum light brown, tapering and grey pruinose (LF:WF = 3.0); arista with thickened base. Eyes meeting for a distance of 12-13 facets. Frons dark silver-grey pruinose. Vertex black, bearing an elevated slightly ocellar triangle. Occiput dark and grey pruinose. Thorax. Postpronotal lobe brown, grey pruinose with 2-3 short bristles along upper margin. Prescutum and scutum black with scattered long setae at anterior supraalar area. Scutellum black with eight thin short setae along posterior margin (up to 0.04 mm). Subscutellum black, grey pruinose. Pleura brown. Wing. Length: 3.5-3.6 mm. LW:MWW = 3.1. Wing almost entirely covered with microtrichia. Pterostigma brown and complete. LS:LTC = 1.0. LTC:LFC = 1.0. Cross-vein r-m reaches dm shortly after onethird of the cell's length. Halter length: 0.5 mm, base dark brown stem and knob light brown. Legs. dark brown, grey pruinose. Mid coxa with three brown anterior bristles. Trochanters partly grey pruinose. Femora dark brown with pale apices, grey pruinose. Mid and hind femora bearing two rows of dark anteroventral small spines in apical half. Tibiae grey pruinose, with two rows of short setae on anterior side and three rows on posterior. Hind tibia with three wrinkled indentations in middle without erect anteromedial setae. Tarsi yellowish with light brown bristles at posterior margin and dark brown scattered setae at anterior margin. Pulvilli light brown, slightly large. Claws light brown with black tips. Abdomen. Ground colour dark brown, tergite 1 silver-grey pruinose, with three dark lateral bristles (up to 0.1 mm). Tergites 2-5 posterolaterally grey pruinose, slightly extending on to dorsal surface along posterior margin, extending on to dorsal surface, otherwise brown pruinose. Tergite 5 slightly wider than other tergites and almost asymmetrical in dorsal view (LT35:WT5 = 1.2, WT5:LT5 = 0.4, T5R:T5L = 1.2). Sternites brown, lighter than tergites, grey pruinose. Syntergosternite 8 enlarged, dark brown and grey pruinose. Viewed laterally, longer than high (LS8:HS8 = 1.8). Membranous area large and triangular-shaped caudally. Genitalia. Genital capsule in dorsal view: epandrium and surstyli brown. Epandrium longer than wide (MLE:MWE = 1.1). Surstyli rather asymmetrical, wider than long, both surstyli with a small inner finger-like projections, tip of both finger-like projections curved towards inner side, left surstylus with a broad projection at the base (Fig. 5A). Genital capsule in ventral view: subepandrial sclerite wide without setae (Fig. 5B); gonopods unequal, right is higher and broader than left one (Fig. 5B); phallic guide strong and straight, pointed apically, lateraly with two downwards sclerotised hook-like projections shortly before the apex, right hook is longer than left one (Fig. 5C). Phallus trifid with circular ejaculatory ducts (Fig. 5B, C). Genital capsule in lateral view: both surstyli wide at the base with a small finger-like projection at apex (Fig. 5E, F). Ejaculatory apodeme small, spade-shaped (Fig. 5D).

Diagnosis
This species can be recognised by the shape of surstyli in dorsal view, wider than long, both surstyli with a small inner finger-like projection apically, left surstylus with a broad projection at outer side (Fig. 5A); phallic guide strong and straight with two laterally hook-like projections shortly before the apex (Fig. 5C).

Etymology
The specific name is derived from the Latin bihamatus which means "with two hooks" and references the two lateral hooks on its phallic guide.

Diagnosis
This species can be recognised by the divided phallic guide in ventral view, separated directly from gonopods, left is stronger and wider than right one (

Diagnosis
The male of this species can be recognised by a large membranous area, as wide as long; unequal gonopods (

Notes
Eudorylas blascoi is part of the E. mutillatus species complex. Based on uncorrected pairwise genetic distances (p-distance), E. blascoi differs from E. mutillatus by 3.58%.

Eudorylas chvalai Kozánek, 1988 Diagnosis
This species can be recognised by asymmetrical surstyli, left surstylus slightly triangular-shaped, right surstylus quadratic-shaped at the base with long inner fingerlike projection and a short outer one in dorsal view; gonopods unequal, right larger than left one in ventral view; phallic guide straight, but at dorsal margin, concave in apical two thirds in lateral view (for illustration, see Kehlmaier 2005a: Figure 69a, k).

Distribution
Greece, Iran (Fig. 1), Turkmenia , Motamedinia et al. 2017, Skevington 2020 (Fig. 9A, B). Body length ( Coxae dark brown, grey pruinose. Trochanters somewhat light brown partly grey pruinose. Femora brown with pale apices, grey pruinose. Mid and hind femora bearing two rows of dark anteroventral small spines in apical half. Tibiae light brown, grey pruinose, with two rows of short setae on anterior side and three rows on posterior. Hind tibia with one or two wrinkled indentations in middle without erect anteromedial setae. Tarsi yellowish with scattered dark setae at anterior margin. Pulvilli yellow. Claws brown with black tips. Abdomen. Ground colour dark brown, tergite 1 silver-grey pruinose, with three long (up to 0.2 mm) and two short (up to 0.08 mm) dark lateral bristles. Tergite 2 silver-grey pruinose. Tergites 3-5 brown pruinose with scattered brown setae. Sternites white-yellow laterally with dark mid-line centrally, grey pruinose. Syntergosternite 8 enlarged, dark brown and grey pruinose. Membranous area small. Genitalia. Genital capsule in dorsal view: epandrium dark brown and surstyli light brown, grey pruinose. Epandrium longer than wide (MLE:MWE = 1.59) (Fig. 10A). Surstyli perpendicular to the epandrium. Right surstylus broader than left surstylus at the base, with two finger-like projections at the apex, inner fingerlike projection longer than the outer one and slightly curved towards left surstylus (Fig.  10B). Left surstylus with an inner finger-like projection at the apex which is slightly broader and longer than the inner finger-like projection of right surstylus (Fig. 10B). Genital capsule in ventral view: subepandrial sclerite wide (Fig. 10C). Gonopods unequal, right gonopod slightly higher than the left one (Fig. 10C), left gonopod with a long finger-like projection towards surstyli in ventrolateral view (Fig. 10D). Genital capsule in lateral view: phallus trifid, long and circular, with strong membranous sheath; phallic guide strong, divided at base, right phallic guide with a projection in the middle which is divided by two branches, a downward branch being longer than upward one (Fig. 10E, G); ejaculatory apodeme spade-shaped (Fig. 10F).    (Fig. 11A, B).

Diagnosis
This species can be distinguished by the specific shape of the phallic guide, divided at the base, right phallic guide with two branched projections in the middle (Fig. 10D); strong membranous sheath (Fig. 10E, G), left gonopod with a long upward finger-like projection (Fig. 10D).

Etymology
The specific epithet is derived from the Latin corniculans, the diminutive form of cornuatus which means horned and references the shape of the phallic guide.

Diagnosis
This species can be recognised by the triangular-shaped left surstylus and broad base of right surstylus with inner finger-like projection in dorsal view; gonopods unequal, right larger than left one in ventral view; phallic guide bent with two small lobes bearing some short setae in lateral view (for illustration, see Kehlmaier 2005a: Figure 30a, h).

Notes
No sequence data exist for this species.

Diagnosis
This species can be recognised by asymmetrical surstyli, right surstylus quadraticshaped at the base with a small finger-like projection apically towards left surstylus, left surstylus slightly rectangular-shape at the base with a triangular projection in dorsal view (Fig. 13A); phallic guide short, apically with downwards hook-like projection in lateral view (Fig. 13B); gonopods slightly unequal, right higher than left one in ventral view (Fig. 13B). The genitalia of this species are similar to E. fluviatilis (Becker, 1900). It differs by the shape of left surtylus in dorsal view and shape of right gonopod in ventral view. In E. flavicrus, the left surstylus has a rather rectangular base followed by a triangular projection in dorsal view (Fig. 13a) and the right gonopod has a distinct finger-like projection (Fig. 13B).

Notes
No sequence data exist for this species.

Diagnosis
This species can be recognised by the very large membranous area; rather symmetrical surstyli in dorsal view, left surstylus with squared base, right one with base longer than wide, both surstyli with a rather inner finger-like process in dorsal view; gonopods small, left is higher than right one; phallic guide broad and straight with inner lateral margin distinctly convex, outer lateral margin slightly convex to straight and pointed apex in ventral view (for illustration, see Kehlmaier 2005a: Figure 37a, k).

Diagnosis
The male of this species can be recognised by the small membranous area; unequal gonopods in ventral view (Fig. 14A); right surstylus apically with two short projection in lateral view (Fig. 14E); left surstylus apically with long projection in lateral view (Fig.  14D); phallic guide with hook pointing downwards in ventral view (Fig. 14B).

Notes
Eudorylas fluviatilis is in the E. mutillatus species complex. It is genetically close to specimens from Indonesia, Bangladesh, China, Vietnam, Australia, Israel, Egypt, Yemen and Pakistan, differring by 0.77-4.02%. The genitalia are very similar between E. fluviatilis and E. mutillatus (Loew, 1858), raising the possibility that the two taxa are conspecific. Eudorylas mutillatus was illustrated and re-described by Skevington (2002a) and Földvári (2013), but this species must be re-assessed across its entire range. A second genetic marker should be used in conjunction with COI to test the species concept (likely ITS2) and to examine closely-related and potentiallysynonymous species like E. fluviatilis.

Diagnosis
This species can be recognised by a long phallic guide in ventral view (Fig. 15B); epandrium longer than wide (Fig. 15A); surstyli with small finger-like projection apically in dorsal view (Fig. 15A); gonopods asymmetrical and small with a small-sized humplike projections on each side (Fig. 15B); subepandrial sclerite wide with scattered setae in ventral view (Fig. 15B).

Diagnosis
This species can be recognised by the size of the right surstylus in dorsal view, wider than long with inner finger-like projection; left surstylus triangular-shaped in dorsal view and dorsal margin of left surstylus humped in lateral view; gonopods small and equal in height; phallic guide short and straight with two triangular projection dorsomedially in lateral view (for illustration, see Kehlmaier, 2005: Fig. 31a, n).

Notes
DNA barcodes of Eudorylas jenkinsoni overlap with those of E. obliquus (0.62-1.63% pairwise divergence). The genitalia of these species differ by the size of the right surstylus in dorsal view, wider than long in E. jenkinsoni, so this is likely another case of recently-diverged species or ancestral hybridisation. There is always a possibility that it is a single species with polymorphic genitalia, so future genetic work is warranted.

Diagnosis
This species can be recognised by asymmetrical surstyli with short inner finger-like projection, right longer than left one in dorsal view; gonopods small, right slightly higher than left one in ventral view; phallic guide straight and broad in lateral view (for illustration, see Kehlmaier 2005a: Fig. 50a, l).

Distribution
Belgium, Croatia, Czech Republic, Denmark, France, Germany, England, Hungary, Iran, Israel, Italy, Latvia, Macedonia, Romania, Slovakia, Switzerland , Skevington 2020 (Fig. 6).  Coxae dark brown, grey pruinose. Fore and hind coxae with four to five short brown setae and mid coxa with two long dark setae and three brown setae on inner apical corner. Trochanters somewhat light brown partly grey pruinose. Fore femur dark brown with pale apices, grey pruinose bearing two rows of dark anteroventral small spines in apical half. Tibiae dark brown, grey pruinose, with two rows of short setae on anterior side and three rows on posterior. Tarsi light brown with scattered dark setae at anterior margin. Pulvilli yellow. Claws light brown with black tips. Abdomen. Ground colour dark brown, tergite 1 brown pruinose, with three to four long dark (up to 0.16 mm) and five to six short (up to 0.08 mm) brown lateral bristles. Tergites 2-5 brown pruinose with scattered brown setae. Sternites brown with some scattered dark setae. Membranous area ovate. Genitalia. Genital capsule in dorsal view: epandrium brown, grey pruinose. Epandrium longer than wide (MLE:MWE = 1.2) (Fig. 17A). Both surstyli rather rectangular-shaped at base. Left surstylus triangular-shaped in apical one third. Right surstylus broader than left one, with a small projection at outer margin in middle and with a longer finger-like projection at inner margin shortly before its apex, pointing towards left surstylus (Fig. 17A). Genital capsule in ventral view: subepandrial sclerite wide (Fig. 17B). Gonopods unequal, right gonopod higher than the left one. Phallus trifid; phallic guide short, broad and straight, with a long lateral projection towards right gonopod horizontally and bent upwards at its apex (Fig. 17B). Genital capsule in lateral view: phallic guide straight with a small dorsolateral projection before its apex (Fig.  17D). Left surstylus rounded with a finger-like projection apically and with a ventral margin distinctly concave in apical one third (Fig. 17D). Right surstylus broadened at base with two finger-like projection apically, the longer one is situated ventrally before the apex and the shorter one arising from the apex (Fig. 17E). Ejaculatory apodeme spade-shaped (Fig. 17C).

Diagnosis
This species can be recognised by asymmetrical surstyli in dorsal view, both surstyli rather rectangular-shaped at base, left surstylus triangular-shaped in apical one third, right surstylus with an inner finger-like projection shortly before its apex, pointing towards left surstylus in dorsal view and with small projection at outer margin in middle (Fig. 17A); gonopods unequal, right slightly higher than left one in ventral view (Fig.  17B); phallus trifid; phallic guide short and straight, with a long lateral projection towards right gonopod horizontally and bent upwards at its apex in ventral view (Fig.  17B). Phallic guide with a small but distinct dorsal projection shortly before its apex in lateral view (Fig. 17D, E).

Etymology
The specific epithet is derived from the Latin nasicus (= nose), referring to the long projection of the right surstylus in dorsal view.

Taxon discussion
Male E. nasicus sp. n. can be identified by the shape of surstyli and phallic guide, which place it in close relation to E. unicolor, E. wahisi and E. pannonicus. The right surstylus of all four of these species shows an inner finger-like projection in dorsal view and the phallic guide has a distinct projection in ventral view. Compared to E. unicolor, the right surstylus of E. nasicus sp. n. has a distinct small projection at the outer margin in the middle in dorsal view (Fig. 17A), whereas in E. unicolor, it does not have a distinct small projection at the outer margin in the middle (see Kehlmaier 2005a: Fig. 67j). Meanwhile, in E. nasicus sp. n., the left surstylus in lateral view is rounded (circleshaped), with a finger-like projection apically and with a ventral margin distinctly concave in the apical one third (Fig. 17D), whereas in E. unicolor, the left surstylus is not rounded and has the ventral margin distinctly concave from the base to the apex. In E. wahisi, the finger-like projection of right surstylus is wider and the lateral projection of the phallic guide is long (see Kehlmaier 2005a: Fig. 68a, j). In E. pannonicus, the finger-like projection of right surstylus is longer and the right gonopod has two distinct projections (Fig. 18B).

Diagnosis
This species can be recognised by the asymmetrical surstyli, base of the right surstylus rectangular shape with inner finger-like projection in dorsal view (Fig. 19A); epandrium wider than long (Fig. 19A); phallic guide dorsally with two finger-like projections situated half way up in lateral view (Fig. 19D, E); small and equal gonopods in ventral view (Fig. 19B). The genitalia of this species are similar to E. jenkinsoni Coe, 1966. It differs by smaller size; shorter setae on the abdominal tergite 2-5; right surstylus longer than wide in dorsal view (Fig. 19A).

Notes
DNA barcodes of Eudorylas obliquus overlap with E. jenkinsoni. See the notes under E. jenkinsoni for more details.

Identification keys
Key to males of Eudorylas species in the Middle East 1 Phallic guide divided (Fig. 7C, Fig. 10E) 2 -Phallic guide not divided 3 2 Right phallic guide with distinct projections in ventral view (Fig.  10C)

DNA barcoding
Based on morphology and DNA barcoding, the present paper introduces four new species of Eudorylas, E. avis sp. n., E. bihamatus sp. n., E. corniculans sp. n. and E. nasicus sp. n. and associates or confirms the association of males and females of three species: E. blascoi, E. corniculans sp. n. and E. fluviatilis. DNA sequence data are provided for 11 Middle Eastern Eudorylas species.