Mulleroconis hyalina gen. et. sp. n.

Forewing hyaline; one straight crossvein in proximal part of costal area; ScP2 diverges obliquely from ScP1; crossvein ra-rp absent; crossvein rp-ma undulated; M without macrosetae reaching posterior wing margin with two branches; crossvein cua-cup straight and aligned with 2cup-a1.

The generic name is a combination collector’s surname (Müller) and the Greek ‘conis’ meaning dust. The generic name is feminine in gender.

BUB 2907; lowermost Cenomanian amber (Shi et al. 2012); Myanmar, Kachin, Hukawng Valley; preserved in a polished, transparent yellow piece of amber (5.68 × 5.00 × 1.09 mm), deposited in the Museum für Naturkunde, Berlin (ex. coll. Patrick Müller). The holotype is in amber syninclusion with one representative of the Auchenorrhyncha.

The specific epithet is after the hyaline forewing membrane.

As for the genus (vide supra).

Male. Body length ca. 1.17 mm (measured from tip of head to tip of genitalia). Head poorly preserved, only the last three segments of one maxillary palp are visible and the terminal segment is distinctly broader and longer than two remaining. Length of terminal palpomere ca. 0.09 mm. Thorax length ca. 0.28 mm.

Forewing ca. 1.46 mm long, ca. 0.65 mm wide; ScP1 long and parallel to costal margin; Scp2 present; division of R into RA and RP at about one-third wing length; RA simple distally connected to ScP2, parallel to ScP1; RP forked; rp-ma markedly sinuate, near the fork RP1 and RP2, connected to MA; stem of M running close to stem of R, briefly connecting each other, M distally branched into MA and MP, without stiff setae; crossveins 1m–cua and 2m-cua present, 1m–cua near the base of wing, between M and CuA, 2m-cua slightly sinuate, between M and CuA, situated slightly behind midwing; CuP separated from CuA near the base of wing; cua-cup present, located at level of division veins RA and RP; crossveins 1cup-a1 and 2cup-a1 present, 1cup-a1 connected to CuP near to fork of CuA and CuP, 2cup-a1 almost aligned with cua-cup; A1 and A2 clearly connected near the base of wing, a1-a2 present. Hindwing ca. 1.25 mm long, ca. 0.55 mm wide with venation very similar to forewing, differing in position of sinuate crossvein rp2-ma; crossvein m-cua partially preserved and basal part of wing with hardly recognizable venation pattern. Legs slender; fore femora (only left femur is visible) a bit shorter and wider than femora of second and third pair of legs; tibias covered with setae; tarsi five-segmented; first tarsomere distinctly longer than remaining tarsomeres; fifth tarsomere elongated with two apical claws. Abdomen large, length 0.64 mm, width 0.22 mm, including genitalia, with widest part approximately in middle of its length, greatly tapering to narrow apical segments; abdominal plicatures absent. Genital structures hardly discernible, presumably below projection of gonarcus, ultimate apices of parameres visible from dorsal view.

Figure 1. 

Mulleroconis hyalina gen. et. sp. n., holotype BUB 2907. A Drawing of left fore wing B drawing of left hindwing. Scale bar: 100 µm.

Figure 2. 

Mulleroconis hyalina gen. et. sp. n. holotype BUB 2907. A Habitus dorsally B wing venation – detail. Scale bar: 100 µm.

Mulleroconis gen. n. can be attributed to the Coniopteryginae based on the following combination of forewing characters: M is bifurcate (a trait occurring in almost all members of this subfamily) (Makarkin and Perkovsky 2019), the absence of two macrosetae on the media, the presence of only one crossvein between RP and M (in our species between RP and MA), and the absence of the abdominal plicatures. The Mesozoic record of Coniopteryginae currently comprises four genera, i.e. Jurasiatypus Kaddumi, 2005; Libanosemidalis Azar et al. 2000; Paranimboa Engel, 2016 and Phthanoconis Engel, 2004 (see list below). All four genera share the presence of two basal crossveins in the costal space, unlike Mulleroconis gen. n. In addition, Libanosemidalis, Paranimboa and Jurasiatypus differ from Mulleroconis gen. n. in the presence of one crossvein between RA (ScP2) and RP. Libanosemidalis, described from the Lower Cretaceous amber of Lebanon, differs from Mulleroconis gen. n. in the presence of rp2-ma while in Mulleroconis gen. n. the crossvein rp-ma is shifted proximally, followed by a single crossvein between M and CuA and the presence of crossvein a2 connecting A2 and the hind margin of the wing (Azar et al. 2000). Paranimboa and Phthanoconis are both known from the Cenomanian amber of Myanmar (Engel 2016; Sziráki 2016, 2017). Paranimboa has a simple RP unlike in Mulleroconis gen. n. where the RP has two terminal branches. Furthermore, it differs in the position of the crossvein between RP and M and in the position of crossvein 1cup-a1 which is branching off from point where Cu is divided into CuA and CuP and in the presence of crossvein a2 (Engel 2016; Sziráki 2016, 2017).

Phthanoconis and Jurasiatypus both differ from Mulleroconis gen. n. in the presence of a single crossvein between M and CuA and in the complete absence of crossveins between CuA and CuP, CuP and A1, as well as between A1 and A2. Moreover, Phthanoconis differs from Mulleroconis gen. n. in the absence of a crossvein between RP and M (Engel 2004). Jurasiatypus was based on single found specimen (holotype) from the Lower Cretaceous amber of the Kurnub in Jordan (Kaddumi 2005). It differs from Mulleroconis gen. n. in the position of the crossvein between RP2 and MA (Kaddumi 2005).

Palaeoconis azari gen. et. sp. n.

Antennae with 19 flagellomeres. Forewing with three pigmented spots; two crossveins present in apical part of costal area. Crossvein rp2-ma approximately as long as basal abscissa of RP2. Media with one macroseta, ending with three terminal branches. Crossveins 1m-cua, 2mp2-cua and a1 present. Abdomen with discernible plicatures on sternites II-IV.

The generic name is a combination of Palaeo and the suffix conis meaning dust.

BUB 2914; lowermost Cenomanian amber (Shi et al. 2012); Myanmar, Kachin, Hukawng Valley; preserved in a polished, transparent yellow piece of amber (9.18 × 6.77 × 2.18 mm), deposited in the Museum für Naturkunde, Berlin (ex. coll. Patrick Müller). The amber piece contains also one syninclusion of an imago of Diptera.

The specific epithet honors Prof. Dany Azar (Lebanese University, Fanar, Lebanon), friend and colleague of AN and JP and worldwide known palaeoentomologist.

As for the genus (vide supra).

Male. Body length ca. 1.93 mm (measured from tip of head to tip of genitalia). Head hypognathous, ca. 0.34 mm. Compound eyes well developed, 0.22 mm x 0.14 mm (from lateral view). Antennae 21-segmented (with 19 flagellomeres), scape and pedicel stouter, longer and broader than flagellomeres, first flagellomere longer and wider than remaining flagellomeres, flagellomeres subquadrate, nearly as long as wide, terminal flagellomere conical. Maxillary palps five-segmented, fifth segment distinctly larger than other palpomeres, length of fifth segment ca. 0.12 mm. Labial palps three-segmented, third segment larger than remaining. Thorax well developed. Prothorax narrower and overall smaller than meso- and metathorax.

Forewing ca. 2.19 mm long, ca. 0.93 mm wide; two distinct apical crossveins in costal area; division of ScP1 and ScP2 0.31 mm from wing apex; R branching into RA and RP 0.48 mm from wing base; RA simple, distally ending connecting with branch of ScP2; RP branched into RP1 and RP2; ra-rp1 present; first spot on basal abscissa of RP2 near RP1 (near place where is RP forked); crossvein rp-m oblique crossing dark spot about mid-length; M with three branches (MA, MP1 and MP2), M with one stiff seta near crossvein rp-m (Figs 3A, 4C); crossvein rp2-ma relatively long, approximately as long as basal abscissa of RP2; crossvein 1m–cua near rp-m, but not aligned; crossvein 2mp2-cua near to wing apex, with third spot over it; crossveins cua-cup and a1 present. Hindwing ca. 2.12 mm long, ca. 0.92 mm wide; venation pattern very similar to forewing with exception of only single crossvein between C and ScP, wing membrane in hindwing hyaline without any pigmented spots; crossveins ra-rp1 and rp2-ma present; basal part of hindwing without clearly recognizable venation.

Figure 3. 

Palaeoconis azari gen. et sp. n., holotype BUB 2914. A Drawing of right fore wing B drawing of left hindwing. Scale bars: 100 µm.

Figure 4. 

Palaeoconis azari gen. et sp. n., holotype BUB 2914. A Habitus lateral view B wing venation and spots C detail of forewing venation with stiff seta on vein M D head from dorso-lateral view showing antennae, terminal segment of maxillary palp and eye. Scale bars: 500µm (A, B); not in scale: (C); 50 µm (D).

Legs long and slender; femora covered with fine and sparse setae; tibiae covered with stiff and dense setae; tarsi five-segmented, first tarsomere distinctly longer than remaining tarsomeres, terminal tarsomere elongated ending with two claws, tarsi covered with fine dense setae. Abdomen large and broad, tapered to the end (visible from lateral view only); plicatures present on sternites II-IV; length of abdomen including genitalia 1.22 mm, width 0.54 mm. Structures of external genitalia hardly discernible with exception of domed and well-sclerotized ectoproct in lateral view and the caudal projection of gonarcus below.

Palaeoconis gen. n. can be attributed to the Aleuropteryginae based on the presence of two crossveins between RP and M and also the presence of plicatures on abdominal sternites II-IV. Nevertheless, Zimmermann et al. (2009) considered this last character as a symplesiomorphy of the Coniopterygidae. However, Palaeoconis does not possess in the forewings two long stiff setae on M, which is the diagnostic character for Aleuropteryginae. In contrast to this, the absence of these setae on the media and the presence of three-branched media are considered as plesiomorphic characters typical for Mesozoic Coniopterygidae (Engel 2002). In addition, Palaeoconis possesses in forewing crossvein a1 connecting A1 to posterior wing margin. According to Engel (2016) the subfamily Aleuropteryginae contains 14 genera (Mesozoic and Cenozoic), i.e., Achlyoconis Engel, 2016; Alboconis Nel et al., 2005; Apoglaesoconis Grimaldi, 2000; Archiconiocompsa Enderlein, 1910; Archiconis Enderlein, 1930; Garnaconis Perrichot and Nel in Perrichot et al., 2014; Geroconiocompsa Engel, 2010; Glaesoconis Meinander, 1975; Hemisemidalis Meinander, 1972; Juraconiopteryx Meinander, 1975; Libanoconis Engel, 2002; Neoconis Enderlein, 1930; Pararchiconis Nel, 1991, and Spiloconis Enderlein, 1907.

Achlyoconis, Alboconis, Apoglaesoconis, Glaesoconis, and Libanoconis share the presence of two distinct basal crossveins in the costal area, and in addition Achlyoconis, Alboconis and Libanoconis share the presence of crossvein a1–a2, vein A2 and crossvein a2. These traits separate these genera from Palaeoconis gen. n. In addition, Glaesoconis described from the Lower Cretaceous Myanmar amber differs from Palaeoconis gen. n. by a distinctly shorter crossvein rp2-ma , single crossvein m–cua, presence of A2 and four pigmented spots on wing membrane instead of three in Palaeoconis (Engel 2004). Libanoconis, known from the Lower Cretaceous amber of Lebanon, differs from Palaeoconis gen. n. in the positions of crossveins between M and CuA (1m–cua is near to wing base; 2m–cua approximately at mid-length of wing), and lack pigmented spots on membrane (Engel 2002; Nel et al. 2004). Achlyoconis described from the Upper Cretaceous amber of northern Myanmar differs from Palaeoconis gen. n. in the presence of seven thickenings with specialized setae on M, positions of crossveins 1m-cua and 2m-cua, presence of a single crossvein between CuP and A1, and four pigmented spots (Engel 2016). Alboconis, described from the Lower Cretaceous French amber, differs from Palaeoconis gen. n. by a two branched media bearing two long setae, crossvein between CuA and R+M, and two crossveins between CuP and A1 (Nel et al. 2004). Apoglaesoconis and Garnaconis share the following features such as M with two long setae, presence of crossveins cup–a1; a1–a2, vein A2 and absence of pigmented spots on wing membrane. Apoglaesoconis, from the Late Cretaceous amber of New Jersey, differs from Palaeoconis gen. n. by one crossvein between RP and M, branches RP2 and MA forming a cross, and by a single crossvein between M and CuA (Engel 2002). Garnaconis, from the Late Cretaceous amber of Vendée in northwestern France, differs from Palaeoconis gen. n. in the remote positions of separated transverse ScP2 and crossvein between RA+ScP2 and RP1 (see Fig. 3), unlike both being aligned in Garnaconis. Palaeoconis gen. n. bears two crossveins beween RP and M, a two-branched media with setae instead of only one crossvein between RP2 and MA and a three-branched media without setae in Garnaconis (Perrichot et al. 2014). Juraconiopteryx, Upper Jurassic of Kazakhstan, represents the oldest record of Coniopterygidae. Wing venation of Juraconiopteryx is very poorly preserved (Meinander 1975), therefore we are unable to make a reliable comparison with Palaeoconis.