Spiloconis oediloma Engel and Grimaldi

Grimaldi, David, Engel, Michael S., Nascimbene, Paul c. & Singh, Hukam, 2013, Coniopterygidae (Neuroptera: Aleuropteryginae) in amber from the Eocene of India and the Miocene of Hispaniola, American Museum Novitates 2013 (3770), pp. 1-20 : 4-14

Spiloconis oediloma Engel and Grimaldi

Figures 1C ; 2A ; 3 ; 4B, C ; 5A

Spiloconis oediloma Engel and Grimaldi, 2007: 19 .

DIAGNOSIS: Distinguished from S. glaesaria by S. oediloma being slightly smaller species; oral margin in frontal view narrow (0.4× width of distance between compound eyes); antenna with 20 flagellomeres, basal flagellomere significantly larger than distal ones; much of frontal portion of head membranous/lightly sclerotized, but with very sparse, fine setulae on only lower third of face; dorsal margin of head greatly protruding; wing with r-rs distal to fork of R 2+3 –R 4+5 (i.e., r-rs connected to R 2+3), length of basal discal cell 3.0–3.2× of its greatest width.

DESCRIPTIVE NOTES: The forewing of DR15-421 (fig. 4C) is distinctive, having the base of Rs and the basal rs-m crossvein equal in length (i.e., the long stem of Rs is midway between R and M, forming a symmetrical V); also, sc-r and r-rs are in line and diagonal, not perpendicular to the longitudinal veins, and m-cu is diagonal to the adjoining longitudinal veins, not perpendicular. In DR14-1427 (fig. 5A), the wing has areas of infuscation that are darker and more extensive than in the other specimens, including a diffuse, longitudinal band in the distal discal cell. Also, the tips of the longitudinal veins are distinctly tapered before contacting the wing margin, and the anterior seta on M lies at the junction of the basal rs-m and m-cu (base of the forewing venation is unfortunately obscured). Other interesting details include the abdomen and appendages; abdominal segments 2–6 each with pair of plicaturae; plicature roughly tridentate in shape (fig. 3A); male genitalia (fig. 3B) relatively simple externally, with broad gonocoxae apparently fused to epandrium, aedeagus large and broad in lateral view, apically pointed; antenna with stout setae having blunt apices; tarsomere 4 much broader than other tarsomeres, tarsomere 5 inserted into proximal end of tarsomere 4; tibiae with 4 longitudinal rows of stiff setae on dorsal and lateral surfaces, lengths of setae slightly less than width of tibia.

TYPE AND OTHER MATERIAL: Known only from Dominican amber. Holotype AMNH DR14-1097 , in a clear, dark yellow amber 2 × 8 × 10 mm, filled with bubbles. Specimen is complete but partially obscured by some bubbles. Nontypes: Female , AMNH DR14-1427 , a very well-preserved female in a small square (4 × 3 × 1.5 mm) of clear yellow amber. Male , AMNH DR15-421 , well preserved but amber split in half, each half ca. 8 × 2 × 2 mm, one piece contains the body and basal portion of the wings, the other piece contains the apical half of the wings .

Spiloconis eominuta Grimaldi and Engel , new species

Figures 1E, F ; 2C ; 5C ; 6

DIAGNOSIS: Known only from Eocene Cambay amber; very small species, forewing length 1.50 mm; front of head with cuticle weakly sclerotized, frons centrally with fine wrinkles (but not collapsed), oval membranous area just dorsal to antennal insertion; oral margin broad; scape large, antenna with 15 flagellomeres. Wing evenly fuscous, crossveins not clouded; with short basal cells (e.g., L/W of basal discal cell = 1.8); M with thickened, setigerous spot on each side of 1rs-m and m-cu (aMs, pMs setae). Male genitalia as described and figured below.

DESCRIPTION: Male. Head: Narrow in lateral view, with long gena. Compound eye relatively large, length 0.55× that of head; compound eye L/W (in lateral view) ca. 1.8; posterior margin of compound eye slightly concave. Frons apparently not membranous but very weakly sclerotized, wrinkled medially; with sparse, fine setulae on ventral half of face and labrum; ovoid membranous areas dorsal to and contiguous with bases of scape (diameter of membranous areas slightly greater than that of scape). Scape large, length 3× greatest width, length ca. 0.9× depth of compound eye, expanded in width apicad (fig. 6A). Pedicel slightly larger than basal flagellomere. Antenna with 17 antennomeres (15 flagellomeres [only 13 visible in Tad- 316D]); flagellomeres setulose, but without whorls; length and width of each flagellomere approximately equal. Maxillary palp with five palpomeres, apical palpomere large, slightly more than twice length of penultimate palpomere. Labial palps with three palpomeres, apical palpomere enlarged. Mesal surface of each apical palpomere with dense, short, fine mat of setulae. Labrum broad, 0.70× width between compound eyes (vs. ca. 0.50).

Thorax: Short, length in lateral view 0.35 mm, dorsally (with cervical and pronotal extension) 0.40 mm; mesopleuron largest segment, slightly thicker than pro- and metapleura. Legs: Femur and tibia of each leg approximately equal in length. Foreleg with tibia apically swollen (fig. 6B), having comb of fine, stiff, erect setae on ventral surface (some setae recurved); tarsomeres dorsally with dense, long setae (setal lengths approximately equal to diameter of tarsi); distal three tarsomeres with ventral mat of fine, dense setae; preapical tarsomere with expanded lateral lobes, cordate in shape.

Forewing: Length ca. 1.50 mm, width 0.60 mm (Tad-316B, fig. 5C), entirely fuscous, without any apparent clouds over crossveins. Crossvein sc-r slightly distal to r-rs; base of Rs and 1rs-m approximately equal in length (fork symmetrical); basal discal cell very short, length 2.0× greatest width (vs. 3.0–5.2), basal medial cell concomitantly short, basal radial cell long (nearly equal in length to basal discal cell). Basal fork of Sc-R very distal to h crossvein, by 5.5× length of h (vs. 1–2x); m-cu slightly proximal to 1rs-m (vs. distal or opposite each other in other Tertiary and in Recent species); both M setae approximately equidistant from 1rs-m; anterior cubital cell rather large, approximately equal in size to basal medial cell (vs. 0.3–0.4× the size). Hindwing: Bases of veins M and Cu very close together and parallel, separated by distance approximately equal to diameter of either vein.

Abdomen: Number of pairs of abdominal plicaturae not entirely visible. Male terminalia well preserved, almost fully visible (fig. 6C, D). Male terminalia: Ventrally with very narrow, U-shaped hypandrium visible, posteriorly connected to pair of long, digitate parameres. Parameres with distinctive wartlike structures (possibly campaniform sensilla), each paramere with faint transverse suture near midlength or distal third. Gonostyli fully articulated (not fused) to gonocoxae; gonostylus scimitar shaped, apically pointed; thin, pointed accessory lobe dorsal to each gonostylus, base of which articulates with gonocoxa. Aedeagus short, pointed, central structure (sclerotization difficult to discern), flanked by pair of clavate, sclerotized lobes approximately equal in length to aedeagus (the gonarcus, or styli, sensu Meinander, 1972).

Female. Unknown.

TYPE MATERIAL: Holotype, male, Tad-316D, in BSIP . Paratype, male, Tad-316B, in AMNH. Both specimens were in the same piece of turbid amber, but were embedded in EpoTek and the inclusions then separated. Tad-316D also contains a small beetle and midge, and occurs in an irregular piece of amber 1 × 2 × 3 mm; wings are mostly obscured, right side of the head is collapsed into the head capsule, and most of the abdomen is compressed; however, the thorax and terminalia are very well preserved, and the latter with its minute structures very well displayed (figs. 6B, C). Tad-316B is in a piece of amber 1 × 3 × 6 mm, with the wings and legs well displayed (fig. 1F, 5C), though basal portion of the venation required slight reconstruction; head and much of the body is compressed, and most of the genitalia is obscured .

ETYMOLOGY: The specific epithet is a combination of eo -, from “Eocene,” the geological epoch from which the amber derived, itself derived from the Greek eos, for “dawn” or “early,” and Latin minutus, in reference to the small body size.

Genus Neoconis Enderlein

Neoconis Enderlein, 1930: 112 . Type species: Helicoconis pistrix Enderlein, 1906 , by original designation.

COMMENTS: The genus comprises 16 modern species occurring in the southern Nearctic southward to southern Brazil, Bolivia, and Paraguay and west to the Virgin Islands ( Sziráki, 2011). The species geographically closest to Hispaniola are Neoconis cubana ( Banks, 1938) , which is recorded from Cuba and Dominica; N. insulana ( Meinander, 1974) from Jamaica; and Neoconis bispina Meinander, 1972 , from the Virgin Islands.

Neoconis paleocaribis Grimaldi and Engel , new species

Figures 1D , 5B , 7

DIAGNOSIS: Larger than the other two Dominican amber aleuropterygines (forewing length 2.4 mm); distinguished by short scape and wing with long basal discal cell (L/W = 5.0, vs. 3.2–4.1). Distinguished from Recent species of the genus (all New World) based on following: 20 flagellomeres, basal 7–8 ones distinctly longer and thinner; dorsal half of frons membranous, ventral half with pair of lateral membranous lines only; forewing with two M setae on each side of basal rs-m and m-cu (vs. pMs on junction of rs-m and M); hind wing with bases of CuA and M separated by distance of 3–4× diameter of veins.

DESCRIPTION: Female. Largest fossil species in amber, wing length 2.4 mm. Head: Compound eye large and deep, depth more than 2× width, 0.5× depth of head. Frons entirely covered with fine, light setulae; depressed in large circular area between and surrounding antennae (apparently membranous, without any sclerotized spots in center); ventral to this area with pair of weak, membranous lines laterally, but area between lines not depressed or collapsed (either membranous or lightly sclerotized, not forming “tongue” of membrane contacting clypeus). Membranous lines on frons do not extend to clypeus; anterior end of each line with well-defined anterior tentorial pit. Portion of head ventral to compound eyes deep, nearly equal in length to depth of compound eyes. Antenna with short, stout scape, length slightly greater than width; scape slightly wider than pedicel, lengths of both approximately equal and 1.3× longer than longest flagellomere. Antenna with 22 segments (20 flagellomeres), basal 7–8 flagellomeres distinctly longer and thinner than apical ones; preapical 11 flagellomeres approximately square in shape, apical flagellomere with tapered apex. Flagellomeres with very fine setulae only, not arranged in whorls. Maxillary palp with five segments; basal four segments small, apical segment large (length approximately equal to that of segments 2–4). Labrum narrow.

Thorax: Pronotum cordate, with slightly emarginate posterior margin; median line of pronotum membranous. Pleuron with sutures between thoracic segments arrayed at angle distinctly oblique to dorsal margin and parallel to each other. Legs slender, tibiae not swollen apically; apex of each tibia dorsally with thin, blackish, cuticular extension. Femora virtually bare; tibiae with faint longitudinal rows of light, short, sparse setae (setal lengths no more than 0.6–0.7× width of tibia). Tarsus: Basitarsomere slender, approximately equal in length to tarsomeres 2–5; distitarsomere (t-5) inserted into base of preapical tarsomere (t-4), not into apex; preapical tarsomere with two large lobes.

Forewing: Length 2.40 mm; nearly entirely hyaline, with extremely faint, small areas of infuscation over crossveins sc-r, r-rs, and m-cu. Crossvein sc-r slightly distal to r-rs; base of Rs slightly longer than 1rs-m (= proximal rs-m), 1rs-m nearly perpendicular to Rs (fork asymmetrical); basal discal cell long and slender, length 4.6× greatest width; basal medial cell extremely long and slender, length 7.0× greatest width, basal radial cell short (0.3× length of basal discal cell). Basal fork of Sc-R slightly distal to level of h (humeral, or c-sc, crossvein); m-cu slightly distal to 1rs-m (vs. distal or opposite each other); aMs (= apical M seta) approximately half distance from 1rs-m than is pMs (= proximal M seta); anterior cubital cell narrow, 0.3× length of basal medial cell. Hindwing: Length of forked segments of Rs approx. 0.3× total length of Rs; bases of veins M and Cu not very close, separated by distance 3–4× diameter of either vein.

Abdomen: Five pairs of plicaturae in pleural membrane of segments 2–6; plicaturae on segments 3–6 flat and fan shaped, without pointed corners (cf. fig. 3C); plicaturae on segment 2 minute, budlike. Spiracles visible on segments 2–6. Female terminalia: Small, cerci small, ovoid, with apical setae longer than length of cercus; pair of small, scoop-shaped lobes medial to cerci (gonovalves); epiproct and hypoproct small, triangular sclerites.

Male. Unknown.

HOLOTYPE: Female (unique specimen), AMNH DR15-29 , in an irregular piece of clear yellow amber 7 × 4 × 2 mm containing no other inclusions; from the Dominican Republic (specific provenance unknown). Deposited in AMNH.

ETYMOLOGY: The specific epithet is a combination of Greek paleo, for “ancient,” and Spanish caribe, in reference to the Caribbean (Hispaniolan) origin of the fossil.

Figures

FIGURE 1. Photomicrographs of Aleuropteryginae in amber. A–D: In Miocene amber from the Dominican Republic. A. Spiloconis glaesaria Meinander, holotype, AMNH DR10-59. B. S. glaesaria, AMNH DR14-1094. C. Spiloconis oediloma Engel and Grimaldi, AMNH DR14-1427. D. Neoconis paleocaribis Grimaldi and Engel, new species, holotype, AMNH DR15-29. E–F: Spiloconis eominuta Grimaldi and Engel, new species, in Early Eocene Cambay amber from India; both are males. E. Holotype, BSIP Tad 316D. F. AMNH Tad 316B.

FIGURE 2. Frontal views of head of fossil species of Spiloconis. A–B: Dominican amber; C: Cambay amber. A. Spiloconis oediloma Engel and Grimaldi, AMNH DR15-421. B. S. glaesaria Meinander, holotype, AMNH DR10-59. Dashed line above head approximates dorsal margin. C. S. eominuta Grimaldi and Engel, new species, BSIP Tad 316D (reconstructed, based on right half of head). Note differences in scale between both A and B and C.

FIGURE 3. Features of Spiloconis oediloma Engel and Grimaldi in Dominican amber. A. Abdominal segments 2–5, AMNH DR14-1427. B. Male terminalia, AMNH DR15-421; oblique lateral view. C. Terminal five flagellomeres, AMNH DR14-1427. D. Detail of apical protarsomeres, AMNH DR14-1427; dorsal view and oblique lateral view. E. Mesotibia and mesotarsus, AMNH DR14-1427; dorsal view.

FIGURE 4. Forewings of fossil species of Spiloconis in Dominican amber. A. Spiloconis glaesaria Meinander, AMNH DR14-1094, also showing detail of costal setae. B. S. oediloma Engel and Grimaldi, holotype, AMNH DR14-1097. C. S. oediloma, AMNH DR15-421 (slightly reconstructed from fractured specimen). Costal setae omitted in B, C, and in figure 5.

FIGURE 5. Forewings of fossil species of Neoconis and Spiloconis in Dominican (A, B) and Eocene Cambay amber (C). A. Spiloconis oediloma Engel and Grimaldi, AMNH DR14-1427, which shows extensive pigmentation. B. Neoconis paleocaribis Grimaldi and Engel, new species, holotype, AMNH DR15-29. C. S. eominuta Grimaldi and Engel, new species, paratype, AMNH Tad-316B.

FIGURE 6. Spiloconis eominuta Grimaldi and Engel, new species, in Eocene Cambay amber. Holotype, BSIP Tad-316D. A. Left lateral view of anterior half of body. B. Lateral view of protibia and protarsus. C. Ventral view of male terminalia. D. Detail of parameres. E. Ventral view of mesotarsus.

FIGURE 7. Neoconis paleocaribis Grimaldi and Engel, new species, in Dominican amber. Holotype, AMNH DR15-29. A. Head and pronotum, dorsolateral view. B. Female terminalia, oblique terminal view.