Hispanocader lisae Golub, Popov et Arillo, 2012

Golub, V. B., Popov, Yu. A. & Arillo, A., 2012, Hispanocaderidae n. fam. (Hemiptera: Heteroptera: Tingoidea), one of the oldest lace bugs from the Lower Cretaceous Álava amber (Spain), Zootaxa 3270 (1), pp. 41-50 : 43-49

Hispanocader lisae Golub, Popov et Arillo , n. sp.

Figs. 1–9.

Holotype male MCNA-10656 stored at the Alava Museum of Natural Science ( Vitoria , Spain). Specimen was embedded in epoxy resin as described in Corral et al. (1999). Body dorsally brownish-black, head and pronotal disk black. Body elongate, 2.52 times as long as wide, flattened in the dorso-ventral direction. Flattened lateral areas of pronotum (paranota) and elytra small areolated.

Type locality and Stratigraphy. Specimen collected at Peñacerrada I-Moraza outcrop, near the village of Peñacerrada, Álava Province, Spain. Escucha Formation, Lower Cretaceous (Albian).

Etymology. The species is named in honor of Dr. Barbara Lis, well-known Polish researcher of Tingidae .

Description. Head black, long due to extended preorbital part, the ratio of its length, considering from the clypeus apex to posterior margin of eyes, to head width is 0.93, the ratio of its length, considering from apex of clypeus to anterior margin of pronotum, to head width is 1.12. Head depressed dorsally. Eyes large, strongly prominent on either side of the head, cover a significant part of its ventral surface. Rostrum rather long, with short and slightly thickened segment I, the latter is thicker then other segments; third segment thickened at its base. Bucculae short, located on each side of clypeus, slightly diverging towards the apex of head and broadly opened anteriorly, do not extends back more than a quarter length of the ventral side of head. Clypeus seen dorsally and anteriorly. Frons with medial longitudinal ridge in the anterior third, back to the base of clypeus. Head strongly flattened dorso-ventrally and slightly concave in the form of grooves on each side of this ridge in the postclypeal part. Lateral margins of head from the base of clypeus and approximately to the level of ocelli flattened and form a very horizontal narrow carinate plate, which bear barely visible single tiny pores. Rather deep excavation situated on each side of head between of anterior ends of these carinate plates and base of bucculae. Vertex with rather broad longitudinal groove between eyes, behind eyes with vast low calli on either side of median line. A couple of simple ocelli situated near of inner margins of eyes, approximately in the middle of their length, being raised above the head surface. The sides of head behind eyes with pair of strongly prominent angulate outgrowths, apices of that outgrowths almost reach outer margin of eyes. Herewith between eye and lateral occipital angulate outgrowth is pretty deep cut.

Head with one pair of conical frontal short tubercles, located anteriorly of eyes, directed obliquely anteriorly and slightly dorsally, with divergent apices. Antennal tubercles short, conical. Antennae 4-segmented, blackish, long and very thin, with thickened the segment I. Ratio of antennal segments length: I <II> III> IV, segment II is longest. Segment IV with thin, long hairs, that are longer than the thickness of that segment.

Pronotum trapezoidal, with concave anterior and posterior margins, posterior margins being concave in almost all its length. Pronotum posteriorly of its anterior margin is slightly raised in the form of ring collar (vesicula), with hardly visible, very fine areolae, placed in 3-4 irregular transverse rows. Anterior angles of pronotum distinctly, but not strongly prominent anteriorly, with rounded apex. Disc of pronotum is flat in general, depressed posteriorly of collar and slightly raised on sides of midline forming very low calli. Low medial keel runs in the hole length of pronotum and bears one row of small rectangular areolae. Paranota (flattened lateral margins of pronotum) is relatively narrow, with two rows of very small areolae in there widest parts and also with 1-2 areolae of the third row. Outer margin of paranota seated by exclusively small tubercles, has a wavy shape: anteriorly behind of anterior corners slightly is concave, more posteriorly is convex, near of the base of pronotum is slightly concave. Posterior margin of pronotum concave, flattened as a chitinous narrow plate carrying one row of extremely small rectangular areolae. Scutellum large, flat, triangular, with pointed apex and coarse punctation.

Submacropterous form. Hemelytra is extending posteriorly to tip of abdomen. Hemelytra is totally sclerotized and areolate including membrane. Areolae of hemelytra are small. Venation well expressed and is richer than in fossil and recent Tingidae . Longitudinal and transversal veins divide the surface of hemelytra on areas and large areolae that include small ones.

Costal area is placed at the outer margin of hemelytron. It separated from the rest of hemelytra by vein C. This area is rather narrow and composed of 2 rows of very small, punctiform, areolae in most of its length, at the basis it is considerably enlarged and with few areolae of the third row there; in addition this area with 1–2 tiny areolae of the 4-th row in the very basis; in its narrowest part, i.e. in the middle of hemelytra length, with one row of areolae only.

Subcostal area wide, bordered by veins C and R+M, with 5–6 rows in the most of its length, with 7–8 irregular rows of areolae having 5–6-angular and rounded forms in the widest part, i.e. in apical sinus.

Discoidal area is delimited from subcostal area by vein R +M and from sutural area by vein Cu, at the holotype on the right elytron is divided by two transverse veins into 3 large areolae. They are named here as basal, radial and medial large areolae, by analogy with the names of large areolae used by Yu. Popov (1989) while describing the Cretaceous Tingidae . Basal large areola composed of 4 rows of small areolae in its widest place; radial large areola composed of 9 rows of small areolae in its widest point, in the inner corner; medial large areola composed of 10–11 rows of small areolae in its widest point, near the apex. On the left elytron of holotype discoidal area includes only two large areolae .

Closed sutural area is disposed medially of discoidal one and is limited by veins Cu and A (A1?). It is divided by two transverse veins in 3 large areolae, which are called conventionally here as the basal, medial and apical ones. These large areolae at their widest places composed accordingly of 7, 5 and 5–6 rows of small areolae. Veins Cu, and A (A1?) unite near of apex of elytron, and CuA terminates at the apex of elytron, without association with vein R+M, medial of the latter.

Membrane narrow, delimited from suture areas by veins A (A1?) and CuA, with 4 rows of small areolae in most of its length and with single areolae of the 5-th row in addition.

Clavus clearly separated from the corium by break, with 7–8 rows of small areolae in its widest place, sizes of clavus areolae does not differ from small areolae into large basal areola of the sutural area. The most outer row of small areolae is isolated from the rest of clavus by the salient veins.

Hypocostal lamella (see below) formed by vein C, has the form of thickened carina.

Scent-gland opening has peritreme in form of small-depth area and the branched off rather long and very strait groove that direct forward along of mesepimerum and comes to the base of elytron. Small semi-ring ridge adjacent to the expanded base of peritreme crosses the ventral surface of costal area, close to its base. Medial part of costal area, surrounded by described above rib, bent down in the form of small lamella. The bottom of vertical carinaform hypocostal lamella adjacents caudal to this lamella but not touch it. Thus, deflected small lamella of costal area and depressed base of hypocosta form a very short and very strait transverse groove passing from extended area of scent gland peritreme to the outer margin of elytron. Herewith hemelytral stenocostal area is not separated.

Body ventrally brown. Pro-, meso-and metasternal plate, limiting the sides of rostrum, absent. Abdominal laterotergites broad and separated from mediotergites by suture dorsally and ventrally.

Legs are long and thin, anterior and posterior coxae are close located but not in contact, middle coxae are rather apart, metacoxae are rotatory type. Anterior and middle femora thicker than posterior ones. The apices of anterior legs with rather thick short bristly hairs. Tarsi 2-segmented.

Measurements (mm). Body length—2.9 body width (elytra folded in rest)—1.15, head length from apex of clypeus to posterior margin of eye—0.35, up to anterior corners of pronotum—0.42, head width—0.375, length of antennal segments (I, II, III, IV)—0.12, 0.42, 0.28, 0.22, length of rostral segments (I, II, III, IV)—0.11, 0.28, 0.58, 0.15, pronotum length—0.45, width of pronotum between anterior corners—0.28, its width between posterior corners—0.79, scutellum length—0.25, its width—0.37; anterior femur length—0.48, anterior tibia length—0.7, anterior tarsus length—0.14, middle femur length—0.52, middle tibia length—0.7, middle tarsus length—0.15, posterior femur length—0.68, posterior tibia length—0.78, posterior tarsus length—0.15.

Comparison. The following complex of morphological features confirms that this new taxon belongs to superfamily Tingoidea .

Antennae 4-segmented. Tarsi 2-segmented. Posterior coxae are rotatory type. Pronotum with areolate paranota and slightly raised anterior part—collar (hood). Elytra entirely areolate, including membrana. Corium divided by raised veins into costal, subcostal, discoidal and sutural areas.

All these features are characteristic of the overwhelming majority of tingoid Hemiptera .

The similarity of described taxon with representatives of the Cantacaderinae (Tingidae) or Cantacaderidae sensu Lis (1999) is proven by the following features: a strongly elongated head, open anteriorly bucculae, the absence of pronotal posterior triangular projection and open so scutellum dorsally. Besides, there is a short transverse groove connecting peritreme of scent-ostiolar opening with costal area of elytra (this condition almost corresponds to ostiolar-stenocostal system of recent Australian genus— Carldrakeana Froeschner, 1968 ). Transversal veins in discoidal area of elytra are present in some recent genera such as Cyperobia Bergroth, 1927 ( New Zealand) and Carldrakeana Froeschner.

At the same time, the new taxon has a complex of the following features that distinguish it from all Tingidae and Cantacaderidae sensu Lis.

Segment of antennae II is long and longer than the other antennal segments, eyes large below, concealing significant part of the ventral (gular) surface of head, the presence of simple ocelli, the presence of large lateral outgrowths behind eyes, “probing” type of rostrum with thickened at the base and the longest segment III, absence of pro-, meso- and metasternal lamelli, limiting rostrum in the sides, elytron vein R +M and CuA not connecting with one another and reach the apex of elytra independently .

There are some similarities with the monotypic Mesozoic family Ignotingidae Zhang et al., 2005 in the venation of elytra. In particular, vast discoidal area of Ignotingis mirifica Zhang et al., 2005 including several large areolae, can be represented as consisting of two areas, located parallel to each other, namely discoidal and sutural ones, suture area bordering membrane. In addition I. mirifica , as the species described here, has rather long antennal segment II, although shorter than segment III. Ignotingidae however have some cardinal differences from new the taxon: head is short, antennal segment III is longest, simple ocelli obviously absent, hemyelytral membrane evidently expressed and still without areolae, etc. The main difference of the taxon described here from tingoid Vianaididae is another form of scent gland peritreme (Y-form in Vianaididae ). Besides the whole complex of the above differences of the new taxon from Tingidae is also real for Vianaididae .

Discussion. Complex features of Hispanocaderidae indicates that this family represents one more tingoid link, which in all probability, is the ancestral group of one of the initial recent groups of Cantacadedrinae ( Cantacaderidae sensu Lis ) and, in particular, Cantacaderini . A rudimentary state of ostiolar-stenocostal system in Hispanocader lisae , in which the stenocostal area is still not in isolation, confirms that it is primary. In this aspect, the recent genus Carldrakeana holds an intermediate position between Hispanocaderidae and Cantacadedrinae ( Cantacaderidae sensu Lis ). Ostiolar-stenocostal system has already been developed, although weakly in the representatives of the genus Carldrakeana (Froeshner, 1996) (or may be it is a results from a reduction of the costal area). Hence the author referred it to Cantacaderinae .

There are two possible options for a status of Hispanocader .

1. On the basis of plesiomorphic state of the ostiolar-stenocostal system the new genus should be placed as a separate subfamily in the family Cantacaderidae sensu Lis. In this case Phatnomatinae should also be given the rank of subfamily, as it was done by B. Lis (1996). However she referred it to the family Tingidae , but not to Cantacaderidae .

2. To give Hispanocader the status of family, taking into account all the above features of it and especially the presence of ocelli, rich venation of hemelytra, not merging veins R+M and CuA in the apical part of hemelytron and only short segment I of antennae. The latter feature is also a major one of the Early Cretaceous Ignotingidae .

The whole complex of morphological characteristics of the new taxon described above, distinguishing it from Tingidae , Cantacaderidae sensu Lis , Ignotingidae and Vianaididae , allows us considering it as an independent family Hispanocaderidae within of the superfamily Tingoidea .