Ebboa areolata, Perrichot, Vincent, Nel, André, Guilbert, Éric & Didier, 2006

Perrichot, Vincent, Nel, André, Guilbert, Éric & Didier, 2006, Fossil Tingoidea (Heteroptera: Cimicomorpha) from French Cretaceous amber, including Tingidae and a new family, Ebboidae, Zootaxa 1203, pp. 57-68 : 62-66

Ebboa areolata , sp. nov.

( Figs 2­3 )

Material. Holotype MNHN Arc 236.4, paratypes MNHN Arc 312 and MNHN Sis 1.1; all are housed in the Laboratoire de Paléontologie, Muséum National d’Histoire Naturelle, Paris, France.

Etymology. In reference to the entire areolate hemelytra and corium shown by the specimens.

Stratigraphic horizon. Uppermost Albian to Cenomanian.

Localities. Archingeay­Les Nouillers, Charente­Maritime, and Salignac, near Sisteron, Alpes­de­Haute­Provence, France.

Description. Head narrow and elongate, with compound eyes reduced and in a rather median position; neck long; no visible ocelli; antennae four­segmented, first segment elongate but relatively short, second longer than first and third, fourth the longest; rostrum elongate, reaching level of base of abdomen.

Pronotum. Trapezoidal, with anterior margin narrower than posterior margin, covered with small areolae and with a median carina (better preserved in Arc 312 than in other specimens); scutellum large, covered with very small dark punctures; peritrema of metapleural scent gland opening straight (quite visible in holotype Arc 236.4 and paratype Sis 1.1).

Hemelytra. Macropterous, entirely areolate; membrane absent; clavus well defined; corium subdivided into large ‘cells’ by elevated veins; discoidal and subcostal areas subdivided into seven large cells and costal area subdivided into five slightly smaller cells. Hind wing not visible.

Legs. Elongate, pulvilli absent; tarsi three­segmented, with first segment very short and the distal two of equal length.

Abdomen. Globular, broad; no abdominal trichobotria on sternites.

Measurements (in mm): length of body 2.85 (from head to apex of abdomen); length/ width of head 0.75/0.2; length of eye 0.2; length of antennal segment I 0.16, II 0.45, III 0.2, IV 0.5; length of rostrum ca. 1.00; length of pronotum 0.25, min/max width 0.35/0.5; length/width of scutellum 0.28/0.5; length of hemelytra 1.38; length of femur/tibia/tarsi 0.7/0.9/0.23, 0.85/1.05/0.23, 1.0/1.42/ 0.23 in fore, mid and hind legs respectively; length/ width of abdomen 1.85/1.35.

Discussion. These fossils, although in ambers from different outcrops, can be attributed to the same taxon because of their identical habitus. According to Schuh and Slater (1995), Ebboa gen. nov. would share with some Leptopodomorpha: Leptopodoidea the following characters: body shape rounded ( Omaniidae ), convex hemelytra, long antennal segment II, and a rather similar pattern of corium venation. In this group however, only the Leptopodidae have at least the clavus, and often much of the dorsum, heavily punctured. Nevertheless, we exclude affinities of Ebboa with Leptopodoidea because of its distinctly smaller eyes, and the absence of trichobotria on the dorsal surface of the head. The Dipsocoromorpha: Hypsipterygidae , the majority of Tingoidea, some Thaumastocoridae , and Piesmatidae have the dorsal surface of body and hemelytra areolate, as in the new genus Ebboa . This genus can be attributed to the Tingoidea because of the presence of an open peritrema of the metapleural scent­gland, but it is not Y­shaped (plesiomorphy) as in the Vianaididae . Furthermore, Ebboa lacks abdominal trichobotria and pulvilli, as in the recent Tingoidea ( Péricart 1983). According to Golub and Popov (2000a, 2003) and Schuh et al. (2006), the hemelytra of the Vianaididae can or cannot be divided into a clavus, corium, and membrane, but are not areolate as they are in all other Tingidae . However, it is not clear when punctuation becomes areolae, even in the discussion by Golub and Popov (2000a). According to these authors, the punctuation or tiny areolae is a symplesiomorphy of Vianaididae and Cantacaderinae. The hemelytra of Ebboa is highly divided in the main area, as in Tingidae , and in other secondary areas, as in Cantacaderinae, suggesting affinities with this subfamily. But the vianaidid Vianagramma goldmani Golub & Popov has a corium divided into large secondary areas. Thus the hemelytral pattern varies greatly within both the Tingidae and the Vianaididae . The cantacaderine­like hemelytra of Ebboa does not exclude affinities with the latter. Also, Ebboa has a considerably elongate second antennal segment, a ‘secondary’ synapomorphy of the Vianaididae ( Golub & Popov 2000a, 2003). The length of an antennal segment can vary within a family, even if it is not the case in the known recent and fossil Tingidae ; thus this last character does not exclude affinities with the Tingidae . Ebboa has a large scutellum, unlike the known recent and fossil Tingidae , but this is probably a plesiomorphy. Lastly, Ebboa has three­segmented tarsi, unlike the Tingidae and Vianaididae , which could correspond to a plesiomorphy, as the tarsi are two­ or three­segmented in the tingoid sister­group Miridae .

Ebboa shares with Ignotingis Zhang, Golub, Popov & Shcherbakov, type species of the extinct family Ignotingidae ( Zhang et al. 2005), the presence of a rather long second antennal segment (longer than first) and three­segmented tarsi. But these two fossils have very different hemelytral pattern of venation, head and pronotum shape, and the longest antennal segment of Ignotingis is the third, unlike in Ebboa .

Ebboa strongly differs from all the modern and fossil vianaidid genera in its elongate head and very long second antennal segment, distinctly longer than the third. It differs from the two Cretaceous genera Vianagramma and Vianathauma in the lack of hemelytral membrane, the elongate head, and distinctly smaller eyes. The two modern genera Anommatocoris China and Thaumamannia Drake & Davis also lack a hemelytral membrane and have very small eyes, even smaller than those of Ebboa ( Kormilev 1955; Drake & Davis 1960). Both these reductions of the modern taxa are probably correlated to their biology, owing that they inhabit ant nests. In conclusion, Ebboa fits neither into the Tingidae nor the Vianaididae , and its phylogenetic affinities remain uncertain within the Tingoidea. Nevertheless, this new genus shows sufficient peculiar characters to justify its attribution to a distinct, new tingoid family.

This and the recent finding of the extinct family Ignotingidae provide evidence for a larger diversity of the Tingoidea during the Cretaceous, the various taxa having more diversified chorologies. Indeed, the morphological disparity of the recent and fossil Tingoidea is very important, with species having areolate hemelytra and others not, species with a coleopteroid habitus and others not. Interestingly, the Tingidae , which is now the more diverse tingoid family, is still underrepresented in the Mesozoic record comparatively to its sister families. A similar situation occurs between the two neuropteran sister families Rhachiberothidae and Mantispidae . The former is well represented in the Cretaceous and relictual in the present time ( Nel et al. 2005), while the latter is rare in the Mesozoic but very diverse now.

Figures

FIGURE 2. Ebboa areolata new genus and species, in mid­Cretaceous amber of France. A. Holotype MNHN Arc 236.4. B. Paratype MNHN Arc 312. C. Paratype MNHN Sis 1.1. Scale bars represent 1 mm.

FIGURE 3. Ebboa areolata new genus and species, holotype MNHN Arc 236.4, general habitus.