Type genus.—Megalyra Westwood, 1832, p. 419, pl. 66,4; pl. 106,4a–f.

Diagnosis.—Antennae filiform, inserted below level of ventral margin of eyes, with 14 segments, with scape short and globular; subantennal groove present, with or without dorsal carina; eyes large, with posterior orbit smooth or margined by a groove and/or carina; mandibles symmetrical, with 3 teeth, rarely with 4 or 5. Pronotum reduced medially, barely visible dorsally; median mesoscutal sulcus present; notauli absent; mesoscutum with parapsides exceptionally present; large triangular axillae separated anteriorly by a foveate groove; mesothoracic spiracle generally completely surrounded by pronotal cuticle, rarely exposed posteriorly. Wing venation reduced; fore wing veins 2r-m, 3r-m, and 2m-cu absent; M+Cu absent or spectral, rarely tubular; hind wing without enclosed cell, with Rs usually not extending beyond middle of wing. Legs with metacoxa large and metafemur swollen; first metasomal segment inserted low on propodeum, very close to metacoxal foramina; ovipositor external, at least half as long as metasoma.

Type species.—Prodinapsis succinalis Brues, 1923, p. 32.

Diagnosis.—Head hypognathous, slightly broader than thorax, obliquely narrowed behind eyes; compound eyes large, widely separated; posterior orbital groove narrow or absent, without foveae; lateral ocelli closer to eye margin than to each other; vertex with a distinct longitudinal median sulcus originating behind central ocellus, crenulate or not; occipital margin raised by a small foveate carina; face with a distinct longitudinal median sulcus anterior to central ocellus; antennae inserted below ventral margin of eyes, closer to each other than to eye margin; subantennal grooves deeply excavated, usually with a dorsal carina. Pronotum short, not visible dorsally; mesoscutum with anterolateral carina along vertical portion, with median mesoscutal sulcus usually crenulate; large triangular axillae not meeting at inner angles, connected by a foveate groove, separated from scutellum by grooves that are usually crenulate; propodeum with a transverse row of small, shallow to deep foveae anteriorly, irregularly areolate-rugose in other parts; anterior thoracic spiracle small, entirely surrounded by pronotal cuticle; posterior margin of mesopleuron foveolate; fore metapleural margin nearly straight. Forewing with pterostigma short, only swelling at junction with r-rs; vein r-rs curved toward wing tip, not branching; distal segment of Rs, veins M+Cu, 1m-cu, Cu1, and distal part of A spectral or absent. Hind wing venation reduced to Sc+R, R1, and Rs, with Rs reaching middle of wing. Anterior and median legs slender; hind legs stout, with coxae large, femora swollen, and tibiae enlarged apically; pro tibia with one long curved spur, meso- and metatibiae with one and two short straight spurs, respectively. Abdomen longer than thorax in females, of variable length in males, with 7 nearly equal segments, spiracles not visible; small cerci spatulate, trilobed and with very fine setae at apex; male genitalia symmetrical, projecting downward or forward, with two simple, spinelike aedeagus valves between parameres; female ovipositor barely longer than half body length.

Comment.—The original diagnosis of Brues is modified according to observations of the new material and also to accommodate the three new species described herein. With specimens occurring from the upper Eocene to the upper Oligocene, P. succinalis is now known to have existed for at least 12 m.y. As a whole, the genus Prodinapsis persisted for at least 30 m.y. and was obviously widely distributed from western to Central Europe.

Prodinapsis succinalis Brues, 1923, p. 32, fig. 1 (holotype male, in former Königsberg collection, presumed destroyed).

Prodinapsis succinalis Brues, 1933, p. 21, pl. 2,6 (type female, in former Königsberg collection, presumed destroyed).

Type material.—Neo type male (designated herein; the sex was chosen in order to match as closely as possible to the original holotype, in this case a male), specimen KU-NHM-ENT B-113, in Baltic amber from Samland Peninsula, upper Eocene.

Other material examined.—Bitterfeld amber: specimen MfN-MB.I-5446, one female; MfN-MB.I-5440, one male; MfN-MB.I-5650, one male; MfN-MB.I 5651, one male; GZG-BST-26981a,b, two males. Baltic amber: specimen GZG-BST-6498, one male; JVC-248, one male (Weitschat & Wichard, 2002, pl. 67a); JVC-307, one female; KU-NHM-ENT B-123, B-136, B-138, B-139, B-140, five females; KU-NHM-ENT B-114, B-115, B-116a, B-116b, B-117, B-119, B-121, B-122, B-124, B-127, B-134, eleven males. Rovno amber: SIZK-K-7092a, one male. Possible additional material; specimen RMRS-Ip253/13614, one male missing dorsal part of head and mesosoma; and SIZK-UA-881, one male. Both preserved in Rovno amber.

Diagnosis.—Longitudinal median sulcus on vertex extending from median ocellus to occipital carina; vertex minutely punctured posteriorly to lateral ocelli in females (Fig. 4.5); posterior orbits with a narrow but distinct groove and a sharp carina (Fig. 2.1, 2.4, 3.5, 4.4); antennae long, filiform; flagellomeres very elongate in females, moderately elongate in males; face with deep median depression separated from subantennal grooves by a sharp longitudinal carina extending from clypeus to torulus (Fig. 2.2); subantennal grooves with a sharp dorsal carina (Fig. 3.5, 4.4); clypeus with anterior margin bilobed. Median mesoscutal sulcus and axillar grooves crenulate; axillae widely separated at inner angles by foveate groove (Fig. 3.6); propodeum in females with a longitudinal row of four large, rectangular foveae medially on dorsum. Forewing (Fig. 1.1) with apical segment of Rs and basal segment of Cul present, spectral; veins M+Cu and Cul aligned; basal section of M nearly equal in length to 1cu-a. Metasoma elongate, subcylindrical. Female ovipositor barely longer than half body length, male genitalia directed downward.

Description.—Female. Body minutely shagreened except posterior part of vertex minutely punctate and propodeum rugose; moderately to densely pubescent, with pale short setae, except antennae nearly bare. Measurements as given in Table 1.

Head. Eyes oval, one-third higher than long, with short dispersed setae; posterior orbits with narrow groove and sharp carina; small punctures posteriorly on vertex gradually enlarging toward occipital margin; antennae more than two-thirds body length, not extending beyond apex of metasoma, inserted at a distance of clypeus; scape about two-thirds as broad as long, more than twice as long as pedicel; pedicel scarcely twice as long as broad; flagellomeres I–IV subequal in length, about six times as long as broad, V–VIII and apical one about five times as long as broad, IX–XI about three times as long as broad; mandibles small, with three teeth increasing in size from base to apex.

Mesosoma. Mesoscutum about twice as broad as long; propodeum with small, irregular areolae on each side of the median longitudinal foveae; pleural margins foveolate.

Wings. Hyaline, without infumate banding pattern. Forewing broadly rounded at apex, scarcely extending beyond metasoma; vein R1 long; r-rs with apex nearly aligned with that of tubular segment of M; apical segment of Rs spectral, reaching wing margin; M+Cu, Cu1, 1m-cu, and A distally of 1 cu-a spectral, Cu nebulous to spectral, almost reaching wing margin. Hind wing venation reduced to Sc+R, R1, and Rs, with Rs reaching middle of wing.

Legs. Metacoxa with anterodorsal part rugose, medial part often with a strigate patch laterally (also in males, see Fig. 2.1); metatibial setae prone, metabasitarsomere with a preening brush of stiff setae along inner margin, its length subequal or slightly longer than combined length of tarsomeres II–V; pretarsal claws curved, simple, as long as arolium.

Metasoma. Elongate, about half body length; cerci bearing six to ten very fine setae; hypopygium moderately large, extending backward for a short distance; ovipositor exserted for less than half body length.

Comment.—Males are very similar but usually smaller, and they differ by the following characters: vertex not punctate posteriorly; antennae with flagellomeres distinctly setose, shorter (I–IV about twice as long as broad, V–IX about two-thirds as broad as long, apical one the longest, more than twice as long as broad) ; propodeum without median line of large foveae. The genital appendages are rather large, with parameres bearing a fringe of fine setae at apex and along inner margin (Fig. 2.3).

Prodinapsis minor Brues, 1933, p. 23, pl. 3,9 (holotype, female, in former Königsberg collection, without reference number provided and presumed destroyed).

Type material.—Neotype (designated herein) female, specimen GZG-BST-6480, in Baltic amber from Samland Peninsula, upper Eocene, with old collection number Z2853 engraved on the amber piece.

Other material examined.—Specimen KU-NHM-ENT B-111, one male; specimen JVC-282, one male; JVC-294, one male; in Eocene Baltic amber.

Diagnosis.—Very similar to P. succinalis but female without postorbital groove, with only a small carina; antennae much shorter, less than half body length, with flagellum compact and noticeably thickened toward tip; toruli in contact with clypeus; face without depression between subantennal grooves; a small carina dorsally of subantennal grooves; clypeus with anterior margin not bilobed in female, convex; metasoma cylindrical, moderately elongate.

Description.—Female. Body minutely shagreened except propodeum, entirely bare; ventral and right lateral views mostly obscured by milky coat. Measurements as given in Table 1.

Head. Eyes oval, one-third higher than long, bare; posterior orbit of eyes raised by a weak carina, without groove (Fig. 6.2); well-impressed median line on vertex extending from median ocellus to occipital carina. Antennae inserted close to each other just above clypeus; scape globular, excavated at apex for reception of pedicel, about twice as long as pedicel; flagellomeres compact, subcylindrical (Fig. 5), nearly equal in length to pedicel; FI–FVII slightly longer than thick, FVIII–FXI scarcely longer than thick, FXII the longest, about twice as long as thick; mandibles small, with three teeth, other mouthparts hidden by milky occlusion.

Mesosoma (Fig. 5, 6.1). Median mesoscutal sulcus and inner axillar grooves crenulate; axillae not meeting at inner angle, separated by four shallow foveae; metanotum not foveate medially; propodeum minutely areolate-rugose; mesopleuron foveolate along anterior, dorsal, and posterior margins.

Wings (Fig. 1.2). Membrane hyaline, without infumate banding pattern. Forewing not extending beyond apex of metasoma; vein R1 moderately long; apical segment of Rs absent; veins M+Cu, Cu1, 1m-cu, and distal part of A spectral; basal section of M nearly equal to 1eu-a; M+Cu and Cul aligned; Cu nebulous, almost reaching wing margin. Hind wing vein Rs barely approaching middle of wing.

Legs. Protibia without visible apical stout spines; metacoxa with anterodorsal surface conspicuously rugose; length of metabasitarsomere subequal to combined length of tarsomeres II–V; tarsomeres V of all legs with two median stiff setae apically; pretarsal claws curved, simple, as long as arolium.

Metasoma. Short, less than half body length; ovipositor only partly preserved.

Comment.—Brues (1933) mentioned that he could not find any rationale for distinguishing between males of P. succinalis and P. minor. Indeed, during the present study, and despite a careful morphometric comparison of 25 specimens (Table 1), I failed to find features that would allow the confident distinction of the two species. No male has antennae with flagellomeres compact and thickened toward apex, nor posterior orbits with groove absent and carina reduced, as in females of P. minor. Three specimens are tentatively assigned herein to P. minor on the basis of the forewing length (FWL) relative to the head length (HL), the only apparent morphometric character that allows the discrimination of two groups and matches that of the females (see Table 1); a succinalis-group with 3.5<FWL/HL<5; and a minor-group with FWL/HL≥5.

Type material.—Holotype female, specimen SIZK-K-3258, in upper Eocene Rovno amber.

Type locality.—Klesov, Rovno region, Ukraine.

Etymology.—From the Latin pumilio meaning dwarf, or pygmy, in reference to the very small size of the species.

Diagnosis.—Antennae with flagellum broadened toward tip; posterior orbits without groove, with a very weak carina diverging from eye posteroventrally (Fig. 6.6, 7.2); longitudinal median sulcus on vertex finely impressed. Mesoscutum very short (Fig. 7.1); mesoscutal groove and inner axillar grooves finely crenulate; axillae scarcely meeting at inner angles; metanotum not foveate medially; propodeum with a transverse row of foveae along anterior margin, remaining part irregularly areolate-rugose. Forewing extending only slightly beyond metasoma (Fig. 7.1); apical segment of Rs absent; basal section of M very short, branching from M+Cu proximad to Rs+M; basal segment of Cul absent (Fig. 1.3). Metasoma longer than mesosoma; ovipositor short.

Description.—Female. Body minutely shagreened except propodeum; mostly bare, in ventral view largely hidden by bubbles and milky coat. Measurements as given in Table 1.

Head. Eyes oval, higher than long, bare; longitudinal median sulcus on vertex extending from median ocellus to occipital margin. Antennae two-thirds body length, with flagellum gradually thickened toward tip; pedicel the shortest segment, less than twice as long as thick; FI–FIV subequal in length, at least twice as long as thick; FV–FXI shorter, less than twice as long as thick; FXII the longest. Toruli, face, and clypeus hidden by air bubble and milky occlusion; mandible small; maxillary palps 5-segmented.

Mesosoma. Mesoscutum three times wider than long, with weak lateral carina along vertical anterior portion; axillae meeting at inner angles anteriorly to axillar grooves for a very short distance; metanotum not foveate medially.

Wings. No visible infumate pattern. Forewing with pterostigma very short; vein R1 short, not extending beyond apices of r-rs and M; 1m-cu, distal segment of Cu1, and A distally of 1cu-a spectral; Cu nebulous for a short distance. Hind wing vein Rs reaching middle of wing (Fig. 1.3).

Legs. Protibia with short apical stout spines; metabasitarsomere as long as combined length of following tarsomeres, with a preening brush of very short, stiff setae along inner margin.

Metasoma. Moderately elongate oval, dilated dorsoventrally by bubbles; about half body length; segments I–IV subequal in length, V–VII decreasing in length; ovipositor exserted for a short distance, about one-third body length (Fig. 7.2).

Comment.—Brues (1933) distinguished P. succinalis from P. minor based, in females, on larger body size and antennae and a sharp posterior orbital carina separated from compound eyes by a distinct groove. This new species is closely similar to P. minor in the proportions of antennae relative to the body and the weak ocular carina. It is less than 1.40 mm in length, however, while females of P. minor range between 1.60 and 2.00 mm (2.30 to 4.00 mm for P. succinalis). It differs from both species by its forewing venation with the basal segment of M being very short and the basal part of vein Cul being absent and by the axillae meeting at inner angles.

Type material.—Holotype male specimen KU-NHM-ENT B-128; paratypes male specimens KU-NHM-ENT B-126, B-133, in upper Eocene Baltic amber from Samland Peninsula; paratype male specimen SIZK-K-27, in upper Eocene Rovno amber from Klesov, Ukraine.

Etymology.—The species name is in honor of Jens-Wilhelm Janzen, former owner of the Baltic amber type material.

Diagnosis.—Posterior orbits of eyes without groove, only raised by a small carina; inner orbits finely crenulate; antennae as long as or longer than body, with flagellum filiform; subantennal grooves with a faint dorsal carina, separated by shallow depression medially on face; clypeus with anterior margin bilobed; forewing extending beyond apex of metasoma, with apical segment of Rs and basal segment of Cu1 absent; basal section of M very short, branching from spectral M+Cu proximad to fork of Rs and Rs+M; metasoma compact; genitalia small, projected downward.

Description.—Male. Body minutely shagreened except pronotum; pubescence barely visible, on frons; vertex, eyes, and mesosoma with scattered, very short pale setae. Measurements as given in Table 1.

Head. Eyes oval, about one-third higher than long; posterior orbits raised by small carina; inner orbits irregularly and finely crenulate (Fig. 8); longitudinal median sulcus on vertex distinct, extending from median ocellus to occipital margin; antennae extending slightly beyond apex of metasoma, filiform (Fig. 8, 9.1–9.6); scape and pedicel less than twice as long as broad; pedicel the shortest segment; flagellomeres cylindrical, scarcely one-third as broad as long, densely covered by short setae, with multiporous plates elliptical, prominent; FIV and FXII the longest, nearly four times as long as broad; other flagellomeres subequal in length; toruli each connected to clypeus by a short, sharp carina, forming a shallow median depression above clypeus; subantennal grooves deep, with a faint dorsal carina; clypeus about one-third wider than long; mandibles with three small teeth increasing in size from base to apex; maxillary palps with segments greatly increasing in length from base to apex.

Mesosoma. Mesoscutum short, 0.4 times as long as wide; median mesoscutal sulcus and inner axillar grooves crenulate; triangular axillae separated at inner angles by small foveae; propodeum areolate-rugose (Fig. 8, 9.1).

Wings (Fig. 1.4). Hyaline, without infumate banding pattern. Forewing broadly rounded at apex; pterostigma reduced; R1 short; vein r-rs not conspicuously curved toward wing tip, with apex nearly aligned with that of tubular segment of M; M+Cu, 1m-cu, apical segment of Cu1, and A distally of 1cu-a spectral; Cu nebulous for a short distance. Hind wing with vein Rs straight, reaching middle of wing.

Legs. Protibia without apical stout spines; anterodorsal surface of metacoxa minutely rugose; metabasitarsomere subequal in length to combined length of tarsomeres II–V; metabasitarsomere with a preening brush of stiff setae along inner margin; pretarsal claws simple, curved, as long as arolium.

Metasoma. Compact, less than half body length, nearly as broad as long; genitalia with small parameres and spinelike valves of aedegus projecting downward (Fig. 9.3).

Comment.—It is observed in P. succinalis that most males are smaller than females (Table 1). Therefore, the smaller size of the present specimens, together with a small posterior ocular carina, could indicate males of P. minor or even P. pumilio. But they differ from females of P. minor by the forewing with basal section of M being very short and a single medial cell, and from both species by very long antennae. They are thus attributed to a new, distinct species.

Type material.—Holo type male, specimen MNHN-PA-48, in lower Eocene (Ypresian) French amber.

Type locality.—Farm Le Quesnoy, Chevrière, near Creil, Oise department, France.

Stratigraphic horizon.—Lowermost Eocene, Ypresian, ca. 53 Ma, level MP7 of the mammal fauna of Dormaal (Nel & others, 1999).

Etymology.—The specific epithet refers to Oesia, the Latin name of the Oise river, which gave its name to the French department from where the fossil specimen originates.

Diagnosis.—Easily distinguished from other species by the absence of median depression on face, the clypeus not being bilobed, the absence of crenulation on median mesoscutal sulcus and inner axillar grooves, a bifoveate groove crossed by small longitudinal ribs between axillae, the anterior margin of propodeum with a transversal row of deep foveae, and the male genitalia with large parameres strongly deflected forward.

Description.—Male. Body minutely shagreened except pronotum and propodeum; head and mesoscutum covered by sparsely scattered short setae, antennae and legs densely covered by short setae; metasoma bare. Measurements as given in Table 1.

Head. Eyes oval, higher than long, apparently bare; posterior orbits raised by a weak carina, without groove; vertex with a short, only finely impressed longitudinal median sulcus extending from median ocellus slightly posterior to lateral ocelli (Fig. 10, 11.3, 11.4); antennae filiform, about two-thirds body length; scape globular, about twice as long as pedicel (Fig. 11.3); flagellomeres cylindrical, with short setae and elliptical multiporous plates, FII–FIV and FXII twice as long as thick, subequal to scape length, FI and FV–FXI nearly equal in length, less than twice as long as thick; face without deep median depression between subantennal grooves; only a weak dorsal carina visible dorsally of subantennal grooves; clypeus with anterior margin convex, bearing a fringe of erect setae; mouthparts not visible, hidden by milky occlusion.

Mesosoma (Fig. 10, 11.4). Mesoscutum nearly half as long as wide; median mesoscutal sulcus and inner axillar grooves not crenulate; inner angles of axillae separated by a large bifoveate groove crossed by small longitudinal ribs; metanotum in dorsal view entirely foveate, with posterior edge straight; propodeum with a transverse row of deep rectangular foveae along anterior margin, otherwise densely areolate-rugose, with scattered stiff setae on declivity.

Wings (Fig. 1.5). No visible infumate pattern. Forewing broadly rounded at apex, not extending beyond metasoma; vein R1 long; r-rs distinctly curved toward wing tip, apical segment of Rs absent; apices of veins Rs and M nearly aligned; veins M+Cu, Cu1, 1m-cu, and distal part of A spectral; basal section of M long, subequal to 1cu-a; M+Cu and Cu1 aligned. Hind wing with three visible hamuli.

Legs. Protibia with a fringe of apical stout spines and long curved spur; meso- and metatibial spurs short and straight; metacoxa large, minutely rugose; metatibia as long as combined length of tarsomeres, covered by prone setae; metabasitarsomere with a preening brush of short, stiff setae along inner margin; pretarsal claws simple, curved.

Metasoma. Elongate, about half body length, subcylindrical; genitalia with large parameres and spinelike valves of aedegus strongly curved forward (Fig. 11.2); inner margin of parameres with short setae.

Comment.—The new fossil is identical to all Prodinapsis species in most characters, including the longitudinal median sulcus on vertex, a weak ocular carina, wing venation, a straight fore metapleural margin, and large axillae not meeting at inner angles. The most apparent difference is the absence of crenulation on the median mesoscutal sulcus and axillar grooves. Among species of Prodinapsis, it differs from P. succinalis by the anterior margin of clypeus not being bilobed, the absence of median depression on face between subantennal grooves, and the absence of posterior orbital groove. It is more similar to P. minor in these characters, but it differs by having longer antennae, a shorter median line on vertex, and entirely foveate metanotum.

Type species.—Ukrainosa prolata n. sp., by designation and monotypy herein.

Etymology.—The new genus-group name is a combination of Ukraine, the country from which the specimen originates, and the Ukrainian word osa meaning wasp.

Diagnosis.—Head scarcely wider than mesosoma, with posterior orbits smooth, without groove or carina; vertex with a faint longitudinal median sulcus; occipital carina minutely foveate; antennae with flagellomeres much longer than broad, cylindrical, flagellomeres II–IV being longest; apical flagellomere obtusely pointed at apex; subantennal grooves deep, with a broad dorsal carina; clypeus bilobed; maxillary palps 5-segmented, labial palps 3-segmented. Mesoscutum moderately elongate, with anterolateral carina present; median mesoscutal sulcus and inner axillar grooves deeply crenulate; axillae widely separated at inner angles by foveae; metanotum entirely foveate medially; propodeum with a transverse row of small, deep foveae anteriorly, remaining part areolate-rugose; mesopleuron with anterior, dorsal, and posterior margins foveate; fore metapleural margin straight. Forewing extending moderately beyond apex of metasoma; wing venation identical to that of P. succinalis except Cu long and tubular on forewing. Hind legs stout; metacoxa large, smooth; protibia with a fringe of apical stout spines and a long, curved spur; meso- and metatibiae with one and two short straight spurs, respectively; basitarsomeres of all legs with a brush of stiff setae along inner margin. Metasoma cylindrical, elongate; hypopygium large, protruding backward; ovipositor long, well exserted.

Comment.—The new genus is very similar to Prodinapsis in most features, but differs by the absence of postocular groove and carina and the forewing vein Cu being tubular for a long distance.

Type material.—Holotype female, specimen SIZK-K-7046, in upper Eocene Rovno amber from Klesov, Ukraine.

Etymology.—The specific epithet is based on the Latin prolatus meaning elongate and refers to the general aspect of the specimen with an elongate metasoma and ovipositor.

Diagnosis.—As for genus.

Description.—Female. Body minutely shagreened; head (including eyes) and mesosoma uniformly covered by short, pale setae. Measurements as given in Table 1.

Head (Fig. 11.6). Hypognathous, about half as long as high; eyes oval, less than half longer than high; ocelli in a large triangle; longitudinal median sulcus on vertex faintly impressed, barely extending beyond lateral ocelli. Antennae about three-quarters body length, with very short setae; scape one-third longer than broad, twice as broad as pedicel; pedicel the shortest segment, about twice as long as broad; flagellomeres I, V–IX, and XII three times longer than broad, II–IV four times longer than broad, X–XI twice as long as broad; torulus separated from clypeus by a short longitudinal carina; clypeus three times wider than long; mouthparts not visible by preservation.

Mesosoma. Mesoscutum about one-third mesosomal length; mesothoracic spiracle small, entirely surrounded by pronotal cuticle. Fore metapleural margin straight (Fig. 11.6).

Wings (Fig. 1.6). No visible infumate pattern. Forewing with apical segment of Rs, M+Cu, 1m-cu, Cu1, and A distally of 1cu-a spectral; Cu tubular, long, approaching wing margin. Hind wing venation reduced to Sc+R, R1, and Rs, with Rs reaching middle of wing.

Legs. Metabasitarsomere as long as combined length of following tarsomeres; pretarsal claws simple, curved.

Metasoma. First tergum the longest, following ones gradually decreasing in length; cerci small, spatulate, with a tuft of long, fine setae; ovipositor as long as body length.

Type species.—Rubes bruesi n. sp., by designation and monotypy herein.

Etymology.—The new genus-group name is an anagram of Brues (Charles T.), as an acknowledgement of his contribution to the study of fossil hymenopterans.

Diagnosis.—Head distinctly wider than mesosoma, with large prominent oval eyes covering length of head, transversely narrowed immediately behind eyes; inner orbits irregularly crenulate, posterior orbits with only a small carina; toruli almost touching; antennae gradually broadened toward apex, with flagellomeres compact; ocelli in a very large triangle; distinct longitudinal median sulcus on vertex posterior to central ocellus, and on face anterior to central ocellus; clypeus projecting downward, not bilobed. Mesoscutum short, with anterolateral carina present; median mesoscutal sulcus and inner axillar grooves punctate; axillae separated anteromedially by faint foveae; propodeum areolate-rugose. Forewing with veins r-rs and M short; apical segment of Rs absent; veins M+Cu, 1m-cu, Cu1, and A distal of 1cu-a spectral, others tubular; a prebasal (proximal) segment of Cu1 present, aligned with 1cu-a, resulting in a pentagonal medial cell. Legs with metacoxa dorsally rugose; tibial spur formula 1-1-2. Metasoma compact, nearly as wide as long; ovipositor less than half body length.

Comment.—This fossil has some general similarities with Maimetsha, but it differs by its reduced number of antennomeres and its forewing venation lacking the basal and apical segments of Rs (thus excluding it from the tribe Dinapsini). It is tentatively placed in the tribe Cretodinapsini on the basis of the longitudinal median sulcus on vertex (possible synapomorphy of the tribe, according to Shaw, 1990b). The postorbital carina is present as in Dinapsini, however without an orbital groove, thus it is more similar to that observed in Prodinapsis.

Type material.—Holotype female, specimen KU-NHM-ENT B-129. Paratype females, specimens KU-NHM-ENT B-112, B-118, B-120, B-125, B-135, GZG-BST-3132; all preserved in upper Eocene Baltic amber from Samland Peninsula.

Etymology.—The species name is in honor of Charles T. Brues for his contribution to the knowledge of Megalyridae.

Diagnosis.—As for genus.

Description.—Female. Body densely shagreened except propodeum and tarsomeres, sparsely covered by very short, pale setae except on metasoma, and longer pale setae on legs and propodeum. Measurements as given in Table 1.

Head. Wider than long, as wide as high, with anterior and posterior margins nearly parallel as viewed dorsally (Fig. 14.1, 14.2, 14.4); eyes very large, covering all the sides of head, sparsely covered by very short, pale setae; inner orbits minutely crenulate, posterior orbits raised by a small carina but without groove (Fig. 13.1); subantennal groove present, without dorsal carina (Fig. 13.2); toruli closer to each other than to eye margins; antennae less than two-thirds body length, with antennomeres cylindrical, compact (Fig. 13.2, 14.1, 14.3), covered by very short, barely visible setae; scape less than twice as long as broad; pedicel the shortest segment, half as long as scape, nearly as long as broad; apical flagellomere the longest, not acute apically, about twice as long as broad; other flagellomeres gradually shortened and broadened from base to apex, with small, multiporous plates rounded; ocelli widely separated, lateral ones closer to eye margins than to each other; vertex with a distinct longitudinal median sulcus extending from central ocellus to occipital margin (Fig. 13.1, 14.4); occipital carina bordered by a row of small to large foveae; clypeus and mouthparts conspicuously protruding downward; anterior margin of clypeus not bilobed; mandibles small, largely hidden by clypeus when closed, with one small basal and two larger apical teeth; labial palps short, with three nearly equal segments, maxillary palps long, with five segments gradually increasing in length.

Mesosoma. Mesoscutum short, about three times wider than long, with anterolateral carina along vertical anterior portion, median mesoscutal sulcus punctate; large triangular axillae connected anteriorly, separated from scutellum by punctate grooves (Fig. 13.1, 14.4); metanotum narrow, with a transverse row of small foveae medially and larger foveae laterally, with posterior edge nearly straight; propodeum areolate-rugose; mesothoracic spiracle very small, entirely surrounded by pronotal cuticle; fore metapleural margin slightly sinuate, anteriorly bordered by a row of small foveae.

Wings. Membrane hyaline. Forewing with only very short, fine marginal setae; pterostigma much reduced, only swelling at r-rs junction; apices of veins r-rs and M nearly aligned; apical segment of Rs absent; basal segment of M short, about half length of 1cu-a; veins M+Cu, 1m-cu, Cu1, and A distal of 1 cu-a spectral; a prebasal (proximal) segment of Cul present, aligned with 1cu-a, resulting in a pentagonal medial cell. Hind wing venation reduced to Sc+R, R1, and Rs, with Rs reaching middle of wing (Fig. 1.7).

Legs. Slender, sparsely setose, with femora swollen (Fig. 14.3). Protibia with small, apical, stout spines and one long, curved spur; meso- and metatibiae with one and two straight spurs, respectively; metacoxa large, with dorsal surface areolate-rugose; metabasitarsomere longer than combined length of those that follow, with a ventral row of short, comblike spines; pretarsal claws small, simple.

Metasoma. Compact, shorter than mesosoma, nearly as wide as long; first tergum the longest; following one gradually decreasing in size; cerci very small, with long apical setae; ovipositor less than half body length, mostly straight (Fig. 13.1, 14.3, 14.5, 14.6).

Type genus.—Megazar n. gen., by designation herein.

Diagnosis.—Head hypognathous, globular; eyes large, oval, posterior orbits without groove or carina; vertex with or without longitudinal sulcus. Antennae long, with 14 segments, flagellum filiform; scape moderately long, pedicel very short, flagellomeres cylindrical, elongate; subantennal groove shallow, without dorsal carina; mandibles broad and short, with four teeth; occipital carina present, foveate or not. Pronotum with anteromedial part moderately high as viewed from side, weakly visible dorsally; mesoscutum very large, nearly half length of mesosoma, with parapsides present; median mesoscutal sulcus not crenulate; axillae large, for a short distance connected anteriorly at inner angles; inner axillar grooves deep, crenulate or not; propodeum carinate, with sharp angle between dorsum and declivity to form a nearly vertical posterior surface; mesothoracic spiracle not surrounded by pronotal cuticle posteriorly; fore metapleural margin sinuate. Forewing with marginal and submarginal cells closed by tubular veins, medial cell small, basal segment of Rs very long, vein M+Cu tubular, and apical segments of M and Cu nebulous or tubular, nearly reaching wing margin. Hind wing venation reduced to Sc+R. Hind leg much stouter than fore and midlegs; metacoxa very large, not rugose; metatrochanter, metafemur, and metatibia with row of comblike spines along inner margin; first two metatarsomeres with preening brush of stiff setae along inner margin; tibial spurs thick. Metasoma elongate, cylindrical, with large tergum 7 and sternum 6 (hypopygium); small spatulate cerci; ovipositor well exserted, at least half body length.

Comment.—The new tribe is immediately distinctive from all other megalyrid tribes by its mandibular form, the mesothoracic spiracle being free posteriorly, the large mesoscutum having parapsides, the forewing vein M+Cu being tubular, the basal segment of Rs very long, the marginal cell enclosed and narrow, the hind wing venation reduced to Sc+R, and the hind leg with comblike spines on trochanter, femur, and tibia. This tribe comprises the genera Megazar n. gen. and Megalava n. gen. and is known from the mid-Cretaceous (Albian) of France and Spain.

Type species.—Megazar elegans n. sp., by designation and monotypy herein.

Etymology.—Combination of the suffix Mega-, in reference to the family Megalyridae, and the name of the Lebanese paleoentomologist Dany Azar, in acknowledgement of his contribution to the study of French amber insects.

Diagnosis.—Head globular, without longitudinal sulcus on vertex; eyes very large, posterior orbits without groove or carina; antennae long, with scape globular and flagellomeres elongate; mandibles broad and short, rectangular, with four teeth. Mesoscutum very large, more than half length of mesosoma dorsally, with faint parapsides present; median mesoscutal sulcus and inner axillar grooves smooth; axillae broader than long, shortly connecting at inner angles; propodeum carinate, with posterior portion nearly vertical; mesothoracic spiracle not surrounded by pronotal cuticle posteriorly; fore metapleural margin distinctly sinuate. Forewing with vein M+Cu tubular; basal segment of M very short; vein Rs present between r-rs and Rs+M to form a large submarginal cell; apical segment of Rs tubular, without elbow indicating former insertion of vein r-m; marginal cell narrow; presence of a short prebasal segment Cu1 between M+Cu and 1cu-a, forming a small pentagonal medial cell; apical segments of M and Cu nebulous, reaching wing margin. Hind wing venation reduced to Sc+R. Hind leg very stout; protibia with a fringe of apical stout spines and one long and curved spur, meso- and metatibiae with two straight long spurs; metacoxa very large, not rugose; hind leg with row of comblike spines along inner margin of trochanter, femur, and tibia, a preening brush of stiff setae along inner margin of tarsomeres I–II. Metasoma elongate, with large hypopygium protruding backward; ovipositor well exserted, at least half body length.

Comment.—Shaw (1988) proposed the recent Megalyridae to be a monophyletic group on the basis of two synapomorphies, viz. a reduced hind wing venation and the mesothoracic spiracle being completely surrounded by pronotal cuticle (see also Gibson, 1985, for this last character). However, the new fossil is the first megalyrid ever known to possess a spiracle not surrounded by pronotal cuticle posteriorly. It also displays unusual mandibles with four teeth, while most megalyrids have only three, except Carminator, which has five teeth. In addition, the new fossil has parapsides, which are absent in all other megalyrids. Nevertheless, the new genus can be included in Megalyridae as it has 14-segmented antennae inserted below the level of the eyes, subantennal grooves present, median mesoscutal sulcus present, a swollen metafemora, and wing venation rather similar to that of the Dinapsini. Unlike the three dinapsine genera Dinapsis, Neodinapsis, and Ettchellsia, however, the new genus has no postorbital carina and dorsal carina of subantennal grooves, and its forewing has a tubular vein M+Cu and a narrow marginal cell closed by a tubular vein Rs without elbow, reminiscent of the lost vein r-m. These unique characters preclude an assignment to any of the existing tribes and justify the new tribe.

Type material.—Holotype female, specimen MNHN-CDL-2.11 (coll. Arnaud), originally in a piece of amber with 45 various arthropod inclusions, now isolated and embedded in Canada balsam (see Néraudeau and others, 2008, fig. 3A).

Type locality.—Cadeuil, Charente-Maritime, southwestern France.

Stratigraphic horizon.—Lower Cretaceous, uppermost Albian, level Alb (sensu Néraudeau & others, 2008).

Etymology.—The specific epithet is the Latin adjective elegans and refers to the elegant position of preservation of the specimen.

Diagnosis.—As for genus.

Description.—Female. Body moderately setose, with setae short and fine, except stouter on antennae and longer on pronotum; surface sculpturing not visible.

Head. One-third broader and higher than long, slightly broader than mesosoma. Eyes oval, very high (Fig. 15.1, 16.1), sparsely setose; posterior orbit of eyes without groove or carina; ocelli equidistant, forming a large triangle; no longitudinal median sulcus on vertex; occipital carina weak, not margined by foveae. Antennae long, more than half body length, inserted below ventral margin of eyes; scape short, globular; pedicel the shortest segment, a quarter longer than thick; flagellomeres elongate, cylindrical (Fig. 16.3), densely covered by very short, stout setae, with rounded multiporous plates; FI, FIII–FIV, FXII subequal in length, about three times as long as thick; FII, FV-FXI shorter, about twice as long as thick. Distance between toruli equivalent to distance between torulus and eye; subantennal grooves shallow, without dorsal carina; clypeus with anterior margin convex, simple; mandibles rectangular, endodont, with 4 teeth increasing in size toward posterior margin (Fig. 16.1); maxillary palps 5-segmented, labial palps 3-segmented.

Mesosoma. Pronotum weakly visible dorsally, medially moderately high as viewed laterally (Fig. 16.1, 16.3); hind lateral margin of pronotum with circular notch around spiracle, making it free posteriorly (Fig. 16.3); mesoscutum only slightly broader than long, more than half mesosomal length dorsally, with anterior and lateral margins foveate, without anterolateral carina; parapsides very faintly impressed, widely separated from median mesoscutal sulcus, long but not reaching anterior mesoscutal margin (Fig. 16.2); median mesoscutal sulcus and axillar grooves smooth; axillae broader than long, meeting at inner angles anteriorly; propodeum carinate, with dorsum short, declivity nearly vertical; mesopleuron apparently not foveate posteriorly; fore metapleural margin sinuate (Fig. 15.1).

Wings. Membrane hyaline. Forewing not extending beyond sixth metasomal segment; pterostigma small, vein R1 long, marginal cell narrow (Fig. 15.2); Rs present between r-rs and Rs+M, straight and tubular; apical segment of Rs tubular, reaching wing margin; basal segment of M very short, apical part of M reaching wing margin, nebulous; presence of a short prebasal segment of Cul between M+Cu and 1cu-a; medial cell small, pentagonal; 1m-cu, second and third portions of Cu1, distal portion of M, and A distally of 1cu-a nebulous; Cu reaching wing margin, tubular. Hind wing venation reduced to Sc+R diverging from wing margin for a short distance, other veins absent. Five distal hamuli, the two most proximal ones somewhat distant from other three.

Legs. All tibiae distinctly thickened apically, tip about twice as broad as base, with short apical stout spines; all but apical tarsomeres with two or three apical stiff setae; metacoxa without sculpturing on dorsal surface; one or two rows of comblike spines on inner margin of metatrochanter, metafemur, and metatibia (Fig. 15.3, 16.4); preening brush of stiff setae on tarsomeres shorter than comblike spines, not aligned in rows; metafemur almost half as broad as long in its middle part; metatarsomeres II–IV gradually decreasing in length, with IV only slightly longer than thick; apical tarsomere about as long as second one; pretarsal claws long and curved, simple.

Metasoma. Cylindrical elongate, half body length; first segment the broadest; first and sixth tergites the longest, subequal in length, other tergites gradually decreasing in length (Fig. 15.1, 16.1); cerci spatulate; ovipositor with only right sheath completely preserved, about half body length; sheath with microscopic transverse striations, with small perforations at apex; hypopygium well developed.

Measurements (in mm). Body length 3.50; head length 0.62, width 0.92, height 0.67; antennae length ∼1.90; scape length and width 0.12, pedicel length 0.10, width 0.08; mesosoma length 1.25; forewing length 2.10; metasoma length 1.63, height 0.65, maximal width 0.87; ovipositor sheath external length ∼1.70.

Type species.—Megalava truncata n. sp., by designation and monotypy herein.

Etymology.—Combination of the suffix Mega-, in reference to the family Megalyridae, and Álava, name of the Spanish province from which the amber containing the specimen originates.

Diagnosis.—Distinctive from Megazar in having a longitudinal median sulcus on vertex, occipital carina being foveate, the inner axillar grooves broadly crenulate, and the forewing with the medial cell being rectangular instead of pentagonal, with M and Cu entirely tubular. The eyes are also slightly smaller and the antennal scape thinner than in Megazar. The mesothoracic spiracle is considered herein to be free posteriorly, although it is not possible to ascertain this character very confidently.

Material.—Specimen MCNA-9416, in Cretaceous Spanish amber.

Type locality.—Amber geological site Peñacerrada 1, near Moraza, Basque-Cantabrian province, Spain.

Stratigraphic horizon.—Escucha Formation, Lower Cretaceous, lower Albian.

Etymology.—The specific epithet is from the Latin truncare (truncated), in reference to the incomplete preservation of the specimen.

Diagnosis.—As for genus.

Description.—Sex unknown. Metasoma, left fore and mid legs not preserved. Ventral structures barely visible, obscured by detritus.

Head. Shorter than wide and high, broader than mesosoma. Eyes large, higher than long, not pubescent; postorbital margin without groove, foveae, or carina (Fig. 17.1, 17.2). Lateral ocelli closer to inner eye margin than to each other; vertex with a longitudinal median sulcus extending from median ocellus to level of posterior margin of eyes; occipital margin raised by a small carina, with a row of small foveae. Antennae long, with 14 segments, inserted below level of ventral margin of eyes; toruli slightly closer to each other than to eyes; pedicel half as long as scape, scarcely longer than broad; flagellomeres cylindrical, first and apical ones subequal in length, more than twice as long as broad, intermediate ones shorter, nearly equal to scape in length. Clypeus and mouthparts poorly visible, except maxillary palps 5-segmented.

Mesosoma. Pronotum short medially although anteromedial part slightly visible dorsally, with posteromedial part moderately high as viewed laterally (Fig. 17.2). Mesoscutum slightly longer than broad, not abruptly truncate anteriorly, without anterolateral carina, with posterior margin not straight dorsally; median mesoscutal sulcus deep, not crenulate (Fig. 17.1); faint parapsides present, widely separated from median mesoscutal sulcus. Axillae large, anteriorly slightly meeting at inner angles, with inner grooves broadly crenulate. Propodeum carinate, with sharp angle between dorsum and declivity to form an almost vertical posterior surface (Fig. 17.2). Pronotal spiracle hardly visible by preservation, possibly free posteriorly, as the pronotal cuticle seems to make a notch along posterior margin (Fig. 17.2); mesopleuron with a row of small foveolae along posterodorsal margin. Fore metapleural margin apparently sinuate.

Forewings hyaline; pterostigma, vein r-rs and apical part of Rs not preserved; M+Cu tubular, aligned with Rs+M (Fig. 17.1); medial cell subrectangular, very small, entirely closed by tubular veins, with M and 1m-cu very short; distal segments of M and Cu tubular, reaching wing margin. Hind wings not visible.

Legs. Fore legs poorly preserved; protibia with two short apical spurs; hind leg stout, with large coxa; metafemur swollen, with at least two rows of short, comblike spines along inner margin (Fig. 17.2); metatibia with two short apical spurs, with a few visible stout spines along inner margin; pretarsal claws small, simple.

Measurements (in mm). Head length 0.30, width and height 0.40; antennae length 0.80; mesosomal length 0.72, maximal width 0.36; forewing length ∼1.10.

Comment.—Although incomplete, this fossil shows characteristic features of the Megalyridae with the 14-segmented antennae inserted below eyes, the mesoscutum with a deep median mesoscutal sulcus, and the hind legs with swollen femora. Critical characters like the complete wing venation are missing, but it is very similar to Megazar in its overall aspect. Indeed, the two fossil specimens are the only ones with such a preening or grooming device on the hind legs.

Type species.—Valaa delclosi n. sp., by designation and monotypy herein.

Etymology.—The genus-group name is an anagram of Álava, name of the Spanish province from which the amber containing the specimen originates.

Diagnosis.—Head hypognathous, wider than long, without longitudinal median sulcus on vertex; eyes large, oval; posterior orbits without groove or carina; antennae filiform, with 14 segments, flagellomeres elongate, cylindrical; subantennal groove shallow, without dorsal carina; clypeus transverse, much wider than high, anterior margin convex, simple; mandibles short, endodont, with three large teeth increasing in size from base to apex. Mesoscutum moderately large, with a small anterolateral carina; median mesoscutal sulcus minutely crenulate (Fig. 18.1); axillae not connected at inner angles, separated by two shallow triangular foveae, with inner edges separated from scutellum by smooth grooves; propodeum irregularly carinate; mesopleuron with posterior margin foveate; fore metapleural margin straight. Forewing extending beyond metasoma, with venation identical to that of Neodinapsis. Fore and midlegs slender, hind legs moderately stouter; all legs with tibiae shorter than combined length of tarsomeres; protibia with fringe of apical stout spines and one long and curved spur, meso- and metatibiae with one and two straight, short spurs, respectively; metacoxa not rugose; metabasitarsomere with short, stiff setae along inner margin. Metasoma compact, oval, with acute tip.

Comment.—The absence of longitudinal sulcus on the vertex and the forewing venation with tubular basal and apical segments of Rs strongly indicate affinities with the Dinapsini. The forewing venation is remarkably similar to that of Neodinapsis but, as for Megazar n. gen. described above, the new genus differs from it and all other extant genera of this tribe by the absence of a foveate-reticulate sculpture on the head and mesosoma, the absence of postorbital groove and carina, and the absence of dorsal carina on subantennal grooves.

Type material.—Holotype male, specimen MCNA-12578a, in a piece of Spanish Cretaceous amber with two Hymenoptera: Falsiformicidae (larger, 12578b) and Chalcidoidea (smaller, 12578c).

Type locality.—Amber geological site Peñacerrada 1, near Moraza, Basque-Cantabrian province, Spain.

Stratigraphic horizon.—Escucha Formation, Lower Cretaceous, lower Albian.

Etymology.—The species name is in honor of the Spanish paleoentomologist Xavier Delclòs who facilitated access to the material, and for his contribution to the knowledge of Spanish amber.

Diagnosis.—As for genus.

Description.—Male. Body without apparent pubescence.

Head. Shorter than broad and high, slightly broader than mesosoma; eyes bare, higher than long; ocelli in a large triangle; small occipital carina present, finely foveolate (Fig. 18.1); antennae moderately elongate (Fig. 18.3), about 0.75 times body length; scape less than twice as long as broad; pedicel the shortest segment, about twice as long as broad; all flagellomeres nearly subequal in length, 2.5 times longer than broad; apical flagellomere acute at tip; distance between toruli subequal to distance between torulus and eye; clypeus about four times wider than long.

Mesosoma. Pronotum not visible dorsally. Mesoscutum less than twice as broad as long, with a short carina along each anterolateral corner; propodeum with fine carinae arranged irregularly (Fig. 18.1); lateral parts of mesosoma barely visible.

Wings. No visible banding pattern. Forewing broadly rounded at apex (Fig. 18.2); pterostigma broad and short, with marginal and submarginal cells closed by tubular veins; veins M+Cu, Cu1, 1m-cu, and A distally of 1cu-a absent; vein R1 long, extending beyond Rs; r-rs short, branching into Rs proximad to level of apical midlength of pterostigma; Rs tubular between Rs+M and r-rs, straight; apical segment of Rs entirely tubular, with a distinct elbow, curving toward anterior margin of wing; M long distal of Rs+M. Hind wing vein Rs short, not reaching middle of wing (Fig. 18.1).

Legs. Metabasitarsomere the longest, combined length of metatarsomeres II–IV about 1.7 times length of apical tarsomere.

Metasoma. Compact, slightly shorter than mesosoma, less than half body length; genitalia barely visible by preservation, apparently reduced.

Measurements (in mm). Body length 1.13; head length 0.31, width 0.44, height 0.38; antennae length 0.87; mesosoma length 0.59, maximal width 0.38; mesocutum length 0.20, width 0.34; forewing length 1.12; metasoma length 0.50, maximal width 0.25.

Type species.—Megallica parva n. sp., by designation and monotypy herein.

Etymology.—Combination of the suffix Mega- in reference to the family Megalyridae, and the Latin Gallica, meaning from France, country from which the specimen originates.

Diagnosis.—Body small. Head without postocular carina or orbital row of foveae, with a weak longitudinal median sulcus on vertex; occipital carina very weak; antennae about half body length, with all but apical segments nearly equal in length; pedicel and flagellomeres scarcely twice as long as thick, apical flagellomere about twice as long as thick; subantennal grooves shallow; clypeus simple, mandibles with three teeth. Median mesoscutal sulcus and axillar inner grooves crenulate; axillae not connected, separated by two large triangular foveae; pronotum areolate-rugose. Forewing with pterostigma much swollen at junction with r-rs; vein Rs beyond r-rs elbowed, entirely tubular to form a fully closed radial cell [2R1]; Rs segment between Rs+M and r-rs present as a short tubular stub not reaching Rs+M, no medial cell; veins M+Cu and A distal of 1cu-a spectral. Legs slender; protibia without apical stout spines, with one long curved spur; meso- and metatibiae with two short straight spurs; metacoxa scarcely larger than others, metafemur swollen; metabasitarsomere short, with a preening brush of stiff setae along inner margin. Metasoma elongate oval, slightly longer than mesosoma. Male genitalia reduced.

Comment.—This genus is easily distinguishable from all other megalyrids by its unique forewing venation with a broad pterostigma, the radial cell being closed, the short stub of Rs present between Rs+M and r-rs but not reaching Rs+M, and the absence of medial cells (Dinapsini-like venation pattern), combined with the presence of a longitudinal median sulcus on vertex. This last character was a possible synapomorphy of the Cretodinapsini, according to Shaw (1990b), but the monophyly of this tribe has not been demonstrated and a sulcus on vertex is also found in Megalava, which belongs in the new tribe Megazarini. The configuration of the forewing venation, with the radial vein present, even if not complete basally, supports affinities with the Dinapsini. The posterior orbital carina, another putative synapomorphy of this tribe, is not present in the new fossil, but Shaw (1990b) indicated that this character is homoplastic. The new genus strengthens the possibility that the presence of this carina is not a synapomorphy for this tribe. Therefore, the new genus cannot be placed confidently in any existing tribe.

Type material.—Holotype male, specimen MNHN-ARC-362.2, originally in a piece of amber with two Diplopoda belonging to Synxenidae, now isolated and embedded in a polished block of epoxy resin.

Type locality.—Font-Benon quarry, Archingeay-Les Nouillers, in Charente-Maritime, France.

Stratigraphic horizon.—Lower Cretaceous, uppermost Albian, level A1sl2 (sensu Néraudeau & others, 2002).

Etymology. From the Latin parvus meaning small, in reference to the minute size of the species.

Diagnosis.—As for genus.

Description.—Body mostly bare, minutely shagreened except pronotum.

Head. One-third broader than long, slightly broader than maximum width of mesosoma (Fig. 19.1, 20.1, 20.2). Eyes bare, large; orbital margins without groove or raised carina; lateral ocelli widely separated, closer to inner orbit of eyes than to each other; vertex with a fine, impressed median line extending from median ocellus to occipital margin; occipital margin raised by a very weak carina, without foveae. Antennae about half body length, inserted below level of ventral margin of eyes, distance between toruli larger than distance between torulus and eye; scape short, globular; pedicel ovoid, less than twice as long as thick; flagellomeres cylindrical, scarcely half as broad as long, subequal in length to pedicel except apical one longer; sub antennal grooves present but shallow, without dorsal carina; clypeus with anterior margin convex, not bilobed; mandibles short, with three teeth increasing in size from base to apex (Fig. 19.2).

Mesosoma. Pronotum not visible dorsally; mesoscutum less than twice as long as broad, with anterior portion subvertical, without visible anterolateral carina; median mesoscutal sulcus and axillar inner grooves crenulate (Fig. 19.1, 20.2); large triangular axillae not meeting at inner angles, separated by two triangular foveae; metanotum with a transverse row of small foveae dorsomedially; propodeum finely carinate; mesopleuron with posterodorsal margin foveolate.

Wings. No visible infumate pattern. Forewing not extending beyond metasoma (Fig. 19.1); pterostigma broadly swelling at junction with r-rs; R1 long, extending beyond apical segment of Rs; Rs between Rs+M and r-rs present as a short, tubular stub branching from r-rs proximad to midlength of pterostigma, not reaching Rs+M; apical segment of Rs with a slight but distinct elbow, tubular, reaching wing margin to form a closed marginal cell; vein 1cu-a distinctly longer than basal segment of M; veins 1m-cu and Cu1 absent; veins M+Cu and A spectral distal of 1cu-a. Hind wing barely visible, venation apparently reduced to Sc+R, R1, and Rs.

Legs. Hind leg moderately stouter than others; metacoxa not rugose on dorsal surface; metafemur swollen; all tarsomeres with short apical stout setae; metabasitarsomere scarcely longer than combined length of tarsomeres II–IV; apical tarsomere as long as combined length of tarsomeres III–IV; pretarsal claws very small, curved, and simple.

Metasoma. Elongate oval, scarcely longer than mesosoma, in its median part nearly half as broad as long (Fig.20.1); segments nearly equal in length; genitalia barely visible.

Measurements (in mm). Body length 1.06; head length 0.22, width 0.34; antennae length 0.60; scape length/width 0.05/0.05; pedicel length/width 0.04/0.02; mesosoma length 0.40, maximal width 0.26; forewing length ∼0.75; metasoma length 0.44, width 0.22.

Megalyra baltica Poinar and Shaw, 2007, p. 65, fig. 1–4 (holotype female, in Baltic amber).

Material examined.—Female specimen KU-NHM-ENT B-137, in upper Eocene (Priabonian) Baltic amber.

Measurements (in mm). Body length 6.00; head length 0.78, width 1.47; ratio first/second flagellomeres length 0.80; mesosoma length 2.10; forewing length ∼4.80; metasoma length 3.12; ovipositor length ∼5.60.

Comment.—The new fossil matches almost perfectly the original description of M. baltica given recently by Poinar and Shaw (2007), including range sizes. The only visible differences are the forewing veins M+Cu, Cu1, 1m-cu, and distal segments of Rs and M being spectral instead of sclerotized. They also described the first flagellomere as being the longest, but it is not coherent with the ratio of the length of first flagellomere/length of second flagellomere of 0.70 that they give in the measurements section. On the new fossil, the first flagellomere is shorter than the subequal second to fourth flagellomeres, and the following ones are nearly equal to the first one. Finally, in the discussion, Poinar and Shaw (2007, p. 67) mentioned the character “anterolateral sharp spines on mesoscutum,” a synapomorphy for Megalyra according to Shaw (1990a). Rather than “sharp spines,” however, the terminology small projection is preferred (S. Shaw, personal communication, April 2009) as it is sometimes reduced as in M. fasciipennis. Indeed, only a very small anterolateral projection is visible dorsally on our specimen, and laterally this projection merely resembles the anterolateral carina that is visible in Prodinapsis or Ettchellsia. Some photographs (Fig. 20.3–20.6) give a few additional details present but not described nor visible in Poinar and Shaw's (2007) original account, such as the foveate occipital carina, the metacoxae with carinate sculpturing on dorsal surface, and the metafemora and metatibiae with long erect setae. All tarsomeres I–IV have a row of 5–6 apical spines on ventral surface (not figured herein).