Type material.—Holotype female IGR.GAR-72.1 (ex coll. Graves), originally preserved with a scelionine wasp and an earwig nymph (the latter described by Engel & Perrichot, 2014: 10J in this volume) in Late Cretaceous (Middle Cenomanian to Early Santonian, 97–85 Ma) Vendean amber, France; deposited in the Geological Department and Museum of the University Rennes 1, France.

Type locality.—La Robinière, departmental road D32, about 2.5 km south-west of La Garnache, Vendée, France.

Etymology.—The specific epithet is a patronym for Didier Graves, the specimens collector.

Diagnosis.—Female. Antenna with 12 flagellomeres, flagellomeres 2–5 transverse, flagellomeres 6–11 spherical, terminal flagellomere as long as the three preceding ones combined; wing with costa prolonged beyond R3; tarsal claw with small denticle at its base and no setae; cerci much elongate, not triangular, with two long apical setae; ratio body length /wing length = 1.38.

Description.—Body stout, total length (without cerci) 0.68 mm. Head with seven setae on vertex but no pair of setae on frons; eyes bare, separated dorsomedially and large, largest diameter (D) 110 µm, small diameter (d) 55 µm; antenna with flagellum 223 µm long; flagellomeres 2–5 transverse, 6–11 spherical, each 16.7 µm in diameter; terminal flagellomere 36.2 µm long, 2.2× as long as flagellomeres 11 or 10 alone, and as long as preceding three flagellomeres combined; palpus four-segmented; third palpal segment 30.6 µm long, 12.5 µm wide; fourth palpal segment slender, 27.8 µm long.

Thorax: 0.21 mm µm long, 0.17 mm µm wide; scutum not prolonged beyond head with around 10 setae; scutellum with rounded margin and two median setae.

Wing 0.49 mm long, 0.17 mm wide; radial cells short, R1 and R3 joining costa in a thickened pterostigma; costa prolonged beyond R3; crossvein r-m absent; vein R4+5 reaching wing apex; median branches reaching wing margin; intercal vein not running close to M.

Legs slender, hairy; length of profemur 0.12 mm, protibia 0.097 mm, protarsi 0.21 mm; mesofemur 0.13 mm, mesotibia 0.097 mm, mesotarsi 0.167 mm; metafemur 0.14 mm, metatibia 0.167 mm, metatarsi 0.194 mm; protibial spur thickened; apex of metatibia with comb of four major spines and a row of smaller spines; metatibial spur not very thick; all basitarsomeres with row of dense setae; fourth tarsomere cylindrical; fifth tarsomere more elongate than fourth; claws sharp, of medium size, equal on all legs, with a small basal tooth and a bristle on each claw; empodia present, small.

Abdomen 0.39 mm long (without cerci), 0.22 mm wide; cerci lamellar, elongate, 0.10 mm long, 41.75 µm wide, with dorsal and ventral margins parallel within anterior half of length, tapering to rounded apex for posterior half of length, dorsally and ventrally with a series of small setae, and apically with two longer setae.

Comments.—According to the phylogenetic analysis proposed by Borkent and Craig (2004), Leptoconops gravesi n. sp. has two synapomorphies supporting its placement within the clade (Leptoconops + Minyohelea), i.e. ‘radial cells very short’ and ‘female wing with R1 and R3 joining costa in a thickened pterostigma’. It also has the following synapomorphies of Leptoconops: 1) ‘wing without r-m’, a character that Borkent and Craig (2004) indicated as ‘unique within at least the Culicomorpha’; 2) ‘female cerci elongate and laterally compressed’.

Within the genus Leptoconops, the female cerci elongate and laterally compressed is the synapomorphy of the group of subgenera (Holoconops Kieffer + (Megaconops Wirth & Ashley + Leptoconops s. str. + Proleptoconops Glas trier)) as defined by Borkent (1995), thus excluding affinities of L. gravesi n. sp. with the other subgenera Styloconops Kieffer and Brachyconops Wirth & Ashley, which have short female cerci, a reversal to the plesiomorphic condition according to Borkent and Craig (2004). Following the key of Sontag and Szadziewski (2011), L. gravesi would fall in the extinct subgenus Palaeoconops Borkent (type species L. amplificatus Borkent), because of its costa prolonged beyond R3. Species of this last subgenus have also elongate cerci, as our fossil. Nevertheless, Palaeoconops differs from our fossil by having 13 flagellomeres instead of 12 and thick spines on first tarsomere of all legs. Affinities with Holoconops can be also excluded by the presence of 12 flagellomeres in our fossil, as opposed to 10–11 flagellomeres in Holoconops (Wirth & Atchley, 1973; Szadziewski & Arillo, 2003). Following the key of Downes and Wirth (1981), L. gravesi falls in the subgenus Leptoconops rather than Megaconops Wirth & Atchley on the basis of the absence of a pair of setae on frons and the presence of a basal tooth and bristle on the female claw.

We compare L. gravesi with other fossil Leptoconops described by Szadziewski (1988, 1996, 2004), Kalugina (1991), Borkent (1995, 1996, 1997), Szadziewski and Arillo (2003), Szadziewski and Poinar (2005), Poinar (2008), and Choufani and others (2011). Unfortunately, the Leptoconops species described by Szadziewski & Schluter (1992) from the Cenomanian French amber is too incompletely described for an accurate comparison. The presence of the inner tooth on claws of L. gravesi n. sp. excludes affinities with L. succineus Szadziewski, L. burmiticus Szadziewski, L. rossi Szadziewski, L. zherikhini Szadziewski & Arillo, L. nosopheris Poinar, L. subrossicus Szadziewski & Poinar and L. boreus Kalugina. Leptoconops gravesi has flagellomeres 2–5 transverse and 6–11 spherical, which excludes affinities with L. daugeroni Choufani & others (flagellomeres 2–11 spherical), L. myanmaricus Szadziewski (2–11 cylindrical) and L. rovnensis Son tag & Szadziewski (2–3 transverse, 4–11 spherical). Leptoconops sibiricus Szadwiewski has the costal vein very short, not extending beyond R3, the terminal flagellomere 4× as long as the preceding one (as opposed to 2.2× in L. gravesi), and the upper surface of claws with setae, unlike L. gravesi. Leptoconops primaevus Borkent and L. copiosus Borkent, do not have any long setae at apex of cerci, unlike L. gravesi. The small inner tooth at the base of claw of L. gravesi excludes affinities with L. curvachelus Borkent that has a large thick inner tooth. Finally, L. gravesi differs from L. clava Borkent by the absence of small empodia and absence of tibial spur on midleg.

Material examined.—Female IGR.GAR-31 (ex coll. Dupé), in Late Cretaceous (Middle Cenomanian to Early Santonian, 97–85 Ma) Vendean amber, France; deposited in the Geological Department and Museum of the University Rennes 1, France.

Type locality.—La Robinière, departmental road D32, about 2.5 km south-west of La Garnache, Vendée, France.

Description.—Female. As this specimen is much damaged from head to thorax, several structures could not be examined. Total body length unknown. Head only partly visible, one eye partly visible, 117 µm in diameter, bare; at least three setae on vertex behind compound eye; antenna with 12 flagellomeres, flagellomeres 2–11 more or less spherical, all 23 µm in diameter; terminal flagellomere 53 µm long, 2.3× as long as preceding one; proboscis elongate; lacinia with at least 10 denticles; palpus four-segmented, third segment ovoid, fourth segment slender with three apical setae.

Thorax damaged. Wing 0.35 mm long, 0.16 mm wide; surface covered by microtrichia, venation identical to that of Leptoconops gravesi (n° IGR.GAR-72.1).

Legs; length of profemur 0.26 mm, protibia 0.22 mm, protarsomeres 0.27 mm; mesofemur and metafemur 0.36 mm, slender; fourth tarsomere cylindrical, fifth tarsomere longer than fourth; foreleg tibial spur thickened; hind leg tibial comb composed of a row of five long and thick setae intermixed with smaller ones and a row of short, strong and straight setae; hind leg basitarsus with thick spines on one side and with long setae on other side; tarsal claws with a strong inner tooth in basal third; empodia absent.

Abdomen broad, 0.73 mm long, 0.4 mm wide; two visible spermathecae visible in segment 8, maybe moved from their original place by damages caused to the specimen; cerci elongate, 0.083 mm long, 33.4 µm wide, with dorsal and ventral margins converging for entire length to broadly acute apex, dorsally and ventrally with a series of small setae, and apically with two longer setae.

Comments.—The wing venation and long cerci are typical of Leptoconops. Similarly to L. gravesi n. sp. described above, the presence of inner tooth on claws of L. gravesi n. sp. excludes affinities with L. succineus, L. burmiticus, L. rossi, L. zherikhini, L. nosopheris, L. subrossicus, and L. boreus. Our fossil has flagellomeres 2–11 spherical, which excludes affinities with L. myanmaricus (2–11 cylindrical) and L. rovnensis (2–3 transverse, 4–11 spherical). The extend of costal vein beyond R3 and terminal flagellomere only 2.3× as long as preceding one, instead of 4×, exclude affinities with L. sibiricus. Leptoconops primaevus and L. copiosus differ from IGR.GAR-31 in the absence of the two long setae at apex of cerci. Also affinities with L. clava can be excluded because IGR.GAR-31 lacks the midleg tibial spur. Affinities with L. daugeroni can be excluded because IGR.GAR-31 has only three apical setae on fourth palpomere, instead of six. L. curvachelus has thick setae on first and fifth tarsomeres of its legs and has not a pair of apical setae on cerci, unlike IGR.GAR-31, which has thick setae only on first tarsomere of hind leg and two elongated setae on apex of cerci.

Affinities with L. gravesi can be excluded on the basis of the shorter cerci, different proportions in leg structures, all flagellomeres rounded, not transverse, and presence of a stronger basal tooth on claws in specimen IGR.GAR-31. Therefore, it corresponds to a different, probably new species. Nevertheless we prefer not to give it a formal name because of its incomplete state of preservation, and maintain it as a Leptoconops species of uncertain subgenus attribution.

Type material.—Holotype male IGR.GAR-127.2 (ex coll. Doyen), originally preserved with a Berothidae (Neuroptera), a Thysanoptera, and a Coleoptera indeterminate; in Late Cretaceous (Middle Cenomanian to Early Santonian, 97–85 Ma) Vendean amber, France; deposited in the Geological Department and Museum of the University Rennes 1, France.

Type locality.—La Robinière, departmental road D32, about 2.5 km south-west of La Garnache, Vendée, France.

Etymology.—The specific epithet is a patronym for Dominique Doyen, the specimens collector.

Diagnosis.—Wing covered with macrotrichia; costa ending at R3; vein R4+5 absent; cells r1 and r2 small to moderately large, distinctly longer than broad, touching but separated; second radial cell square ended; a single row of hairs on wing fringe; eyes narrowly separate above antennae; palpus five-segmented; antenna with 13 flagellomeres, first one with sensilia coeloconica; flagellomeres 11–13 elongate; terminal flagellomere without apical stylet-like prolongation but rounded apex; all legs having nearly similar tarsal ratios; mesobasitarsus with only slender setae, without small spines; metabasitarsus with palisade-like setae, without subbasal spine; gonostylus with a thick base tapering to pointed apex, not divided apically nor with subapical lobe.

Description.—Head: eyes bare, separated dorsomedially by two ommatidial distance; vertex with more than 12 setae; antenna with plumes directed towards apex of flagellum; pedicel 100 µm long, 140 µm large; flagellum 416 µm long; only last three flagellomeres elongate, nearly equal in length, 62 µm long, the last two flagellomeres with sensilla basiconica; first flagellomere with sensilla coeloconica, with two whorls of long setae; palpus five-segmented, third palpal segment ovoid, 24 µm long, 14 µm wide, with three long sensillae medially; fourth and fifth palpal segments slender, each 18 µm long, 12 µm wide.

Thorax: 0.4 mm long, 0.388 mm wide; scutellum rounded, with four setae; scutum very hairy; katepisternum lacking setae.

Wing 0.608 mm long, 0.24 mm wide; membrane and veins covered with short macrotrichia; radial cells small and nearly equal, almost touching, both distinctly longer than broad; second radial cell square-ended; costal vein not extending beyond R3; CR 0.65; r-m oblique; medial bifurcation not visible but most likely distal to r-m; M2 partially reduced; no patterns of dark and light spots.

Legs slender, very hairy; protibial spur with a small comb; mesotibia apparently with a spur; metatibial comb barely visible; metabasitarsus with stout setae; fourth tarsomere cylindrical; tarsal claws simple, empodia absent.

Abdomen; tergite IX shorter than gonocoxite with small cerci; apicolateral processes not visible; gonocoxites long, with bases not contiguous, and with elongated lateral setae and a tuff of setae near the outer apical angle; gonostylus not forked, single, with thick base, gradually tapering to apex, apex not divided, with no apical tooth nor setae; aedeagus short, with short basal arms, lateral arms longer, arched, bound into slightly longer caudal arm; parameres symmetric, a little extended beyond apex of gonocoxite, with simple rounded apex directed inward.

Comments.—Following the keys of Szadziewski (1996), our fossil is distinguishable from Lebanoculicoides Szadziewski, by the absence of vein R4+5; from Archiculicoides Szadziewski, by the absence of apical stylet-like prolongation of terminal flagellomere but the presence of a rounded apex; from Leptoconopinae on the basis of the eyes narrowly separated above antennae, and the palpus five-segmented; from Austroconops Wirth & Lee, Archiaustroconops Szadziewski, and Minyohelea Borkent (incl. Lebanoconops Szadziewski), on the basis of tarsal ratios of all legs nearly similar; from Brachycretacea Szadziewski, by the presence of five palpal segments instead of four; from Heleageron Borkent, by the wing with cells r1 and r2 feebly separated; from Protoculicoides Boesel, by the costa ending at R3 and radial cells small to moderately large (as opposed to a costa prolonged to wing apex and very large radial cells in Protoculicoides); from Washingtonhelea Wirth & Grogan, Ceratopogon Meigen, Palaeobrachypogon Borkent, and Peronehelea Borkent, by the absence of sub-basal spine on metabasitarsus; and from Atriculicoides Remm, by its flagellomeres 11–13 elongate (as opposed to flagellomeres 10–13 in Atriculicoides), a situation similar to what is seen in Culicoides. Our fossil is also distinguishable from Devalquia Choufani & Nel, by the presence of a sensilia coeloconica on the first flagellomere, the presence of a single row of hairs on the wing fringe, and the second radial cell square-ended (as opposed to narrowed in Devalquia; see Choufani & others, 2013).

It is not possible to compare this fossil to the numerous recent species of Culicoides; we only compare it to other Cretaceous species. Following the keys of males proposed by Borkent (1996), C. doyeni n. sp. is distinguishable from C. flipalpis Remm, C. tyrrelli Boesel, and C. bifidus Borkent, by its gonostylus tapering to pointed apex, not divided apically nor with subbapical lobe; from C. bullus Borkent, and C. succineus Remm, by its gonostylus with a thick base, gradually tapering to apex; from C. sphenostylus Remm, C. obuncus Borkent, C. annosus Borkent, and C. casei Grogan & Szadziewski, by its wing with macrotrichia present all over the wing, and especially basal to fork of M and Cu (as opposed to a wing with macrotrichia restricted to an area distant from M and Cu in these species). It differs from C. canadensis (Boesel, 1937) in the radial cells weakly separated and square-ended (well separated and nearly as long as broad in C. canadensis); from C. agamus Borkent, by the second radial cell relatively thinner compared to its length, and presence of palisade-like setae on metabasitarsomere. According to the keys of Borkent (1995), C. doyeni n. sp. is also distinguishable from C. kaluginae Remm, by the mesobasitarsomere with only slender setae, without small spines.

The comparison with species of Culicoides known only by females is more tentative and limited to characters that are supposedly not dimorphic. Culicoides doyeni n. sp. is distinguishable from C. grandibocus Borkent, by its wing with macrotrichia not restricted to the distal half; from C. yoosti Borkent, by its metabasitarsomere with stout, palisade-like setae (no such setae visible in C. yoosti); and from the very distinctive C. truncatus Borkent, by its antenna with 13 flagellomeres (as opposed to only five flagellomeres in C. truncatus).