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Home , Cicadidae, Orapa (cicada), Tettigarctidae

Eur. J. Entomol 95: 593-598, 1998 ISSN 1210-5759

Liassotettigarcta africana sp. n. (: Cicadoidea: ), the first Mesozoic from Tunisia

A ndré NEL1, Mohamed ZARBOUT2, G eorges BARALE' and M arc PHILIPPE '

'Laboratoire d’Entomologie générale et appliquée. Muséum National d’Histoire Naturelle, 45 rue Buffon, F-75005 Paris, France; e-mail: [email protected] 2Office National des Mines, 24 rue 8601, La Charguia, 1080 Tunis Cedex, Tunisia 'Laboratoire de Biodiversité et Evolution des Végétaux, Université Claude Bernard, Bat. 401 A, 43 Boulevard du 11 Novembre 1918, F-69622 Villeurbanne Cedex, France

Auchenorrhyncha, , Cicadoidea, Tettigarctidae,Liassotettigarcta, new species, fossil, Lower , Tunisia

Abstract. A new Lower Cretaceous , Liassotettigarcta africana sp. n. (Tettigarctidae), is described from Tunisia, based on forewing impression. A brief overview of African fossil is presented.

INTRODUCTION The fossil record of African and Cretaceous insects is very poor, relative to the size of this continent. A few Coleoptera are known from the Upper Jurassic of Madagascar (Paulian, 1969). A few Algerian and Egyptian insects have been described and are listed in Schliiter (1990). A coleopterous larva has been described from the Lower Cretaceous of Angola (Teixeira, 1975). A number of Lower Cretaceous insects have been recorded from the mine, Botswana (Rayner, 1987; Waters, 1989a,b; Rayner et al., 1991; Rayner & Waters, 1989, 1991; Brothers, 1992). Nevertheless, no fossil insect is known from the Mesozoic of Tunisia. Thus, the present discovery of a well preserved wing of a cicada is of considerable interest because it suggests the possibility of a rich Low Cretaceous ento- mofauna in this country. Along the 200 km of the Dahar cliff in SE Tunisia (Fig. 1), a detritic series, the “Groupe Asfer” (Zarbout, 1994) crops out, ranging in age from the Oxfordian up to the Aptian. It has been studied by, among others, Solignac & Berkaloff (1934), Barnaba (1965), Busson (1967), Peybernes et al. (1985), Bouaziz et al. (1989) and Ben Ismail (1991). The literature mentions fossil woods from only two levels in this region. Recently we have described the first results of the discovery of a new flora with Pteridophyta and gymnosperms (Barale et ah, 1997) at the base of the Douiret formation, as well as several wood levels in all the se­ ries. Within these levels, the richest assemblage of leafy remains had been observed in the clay at the base of the “Argiles de Douiret”. We noted one decimetric fish and two insects (this cicada and a small beetle). This is the first record of insect remain from the Mesozoic in Tunisia in a level with numerous autochtonous accumulations of mainly leafy remains. These remains (Barale et ah, 1997) correspond to an accumulation after a minimal trans­ port of a low herbaceous riparian vegetation.

593 There is no consensus on the interpretation of wing venation in Tettigarctidae between Shcherbakov (1981), Dworakowska (1988), and Kukalova-Peck (1991) (see Table 1). We follow Kukalova-Peck (1991) for the present study.

I f f im Sandstone

[ i | 1 1 Limestone [rf-JfJ] Silty clay

|y ‘ / | Dolomite | W A| Gypsum Slump bedding m i Clay and marl ■ & - 54* 4?0 kVv$od Conglomerate level ^ Stipes of Pterldophytes

5 4 0 - ^ t Coniferous trunks

-€?■ Leaves S 4 * 4 k 4' $ Insect 5 4 4 k

■— '4# - e ^ E l 5 4 & z r 5 4 > ^ 5 4 4 k

5 4 4 k

Fig. 1. Synthetic geological section for the Jurassic and Cretaceous of Tataouine area. In the insert, the asterisk corresponds to the locality Merbah el Asfer.

594 T able 1. Interpretations of the wing venation of the Tettigarctidae. Shcherbakov (1981) Dworakowska (1988) Kukalova-Peck (1991) C PC + C PC + (C = CA + CP) ScP ScP + R (ScP) + R R + M M MP R1 ScP + RA RA Rs RP + MA RP M MP MP CuA CuA CuA CuP CuP CuP Pcu AA AA1 +2 A1 AP' AA2 + 3 A2 AP" AP

TAXONOMIC ACCOUNT OF THE NEW FOSSIL

Superfamily Cicadoidea Latreille, 1802 Family Tettigarctidae Distant, 1905 Genus Liassotettigarcta Nel, 1996

Liassotettigarcta ofricana sp. n.

Description . Impression of a nearly complete forewing, with the apex and the extreme base missing (Figs 2, 3). It is broken along the nodal line and vein CuP. There are some weak traces of colouration along the posterior margin of the clavus (anal area) but the rest of surface seems to have been hyaline. Some more or less dark spots are visible along the costal margin but they continue outside of the wing, thus they are not part of the original colouration. The nodus is in a rather distal position. Veins C and PC cannot be distin­ guished from each other. Veins R and MP are basally separated but very closely parallel. MP separates from R+MP just distal of the point of fusion between R and MP, then MP is strongly curved and strongly approximates CuA, with a point of contact between the two veins; there after distally diverge again. There are two short crossveins between RA and the costal margin in the nodus, basal and distal of the nodal furrow (interpreted by Dworakowska, 1988 as branches of ScP). There is a rather poorly preserved crossvein be­ tween RP and RA below the more basal of these crossveins. RA has two costal branches reaching the costal margin and a posterior branch directed towards the wing apex. RP is a long and straight vein. The nodal line is well defined. There is a crossvein between MP1+2 and RP, closely parallel with the nodal line. The second apical cell a2 and the cell a3 are closed by distal oblique crossveins respectively between RA and RP and between RP and MP1+2. Five other apical cells a4 to a8 are present between the branches of MP and CuA. Vein Cu is divided into CuA and CuP 3.4 mm distal of the wing base, CuA and CuP are well separated. CuP is nearly straight. The cubital triangle (Ct) is narrow. CuA is curved and has a strong angle at the level of the nodal line, there after it becomes parallel with it. AA1+2 is weakly curved. AA3+4 is basally nearly fused with AP but these veins are dis­ tally separated again. AA3+4 is finally fused with AP about 9.0 mm from its base. The ju­ gal area is not preserved.

595 Fig. 2. Forewing venation of Liassotettigarcta africana sp. n., holotype specimen No. 103.

D imensions . The wing is about 38 mm long and 12 mm wide. Distance from the base to the nodus, 23.0 mm; from the nodus to the apex, about 15.0 mm. Width of the costal area between PC + C and R, 3.0 mm. Length of the first postnodal cell defined by RA and the costal margin, 8.2 mm, width, 0.4 mm; veins R and MP fused 8.2 mm from the wing base; maximal width between them, 0.1 mm; vein RP diverges from RA 2.5 mm distal of the base of MP; length of aO, 7.6 mm, width, 0.6 mm; width of al, 0.6 mm; width of a2, 0.8 mm; width of a3, 1.3 mm; width of a4, 1.4 mm; width of a5, 1.4 mm; width of a6, 1.5 mm; width of a7, about 2.0 mm; width of a8, 2.4 mm (the lengths of these cells are unknown). Length of the cubital triangle, 1.8 mm, width, 0.4 mm. Maximal distance between CuA and CuP, 2.6 mm. Width of the cell between AA1+2 and CuP, 1.8 mm; between AAI+2 and AA3+4, 3.4 mm; between AA3+4 and AP, 0.9 mm.

Fig. 3. Photograph of Liassotettigarcta africana sp. n., holotype specimen No. 103. Scale: 10 mm. 596 Diagnosis . This species differs from L. mueckei Nel, 1996, the only other species in the genus, in the following points: Its wing is nearly completely hyaline, unlike that of L. mueckei which is covered with dark spots; it has two costal branches of RA distal of the nodus, instead of one in L. mueckei.

Material : Holotype specimen No. 103, collection of the Musée de Paléontologie de l’Office des Mines, Tunis, Tunisia. Locality deposit . Merbah el Asfer, Tataouine region, Tunisia, Africa. G eological age . Lower Cretaceous, Aptian, Douiret Formation, Asfer Group. Etymology . After Africa.

DISCUSSION The venation of this forewing is nearly identical to those of the extant spp. The only differences are as follows: There is a crossvein between MP and RP along the nodal line; there is no crossvein (arculus) between CuA and MP because these veins are in contact; CuA has a strong curve along the nodal line before its division into two branches CuA 1+2 and CuA3+4; the cubital triangle is narrow. All these characters are present in Li- assotettigarcta mueckei Nel, 1996 from the Lower Jurassic of Germany and were consid­ ered by Nel (1996) as diagnostic of this last genus. Nel also gave an annotated list of the genera and species which have been attributed to the Tettigarctidae. Except Liassotettigarcta, only the genera Cicadoprosbole Becker- Migdisova, 1947 (Liassic of Kirghizia), Turutanovia Becker-Migdisova, 1949 (Jurassic of Kazakhstan), Shuraboprosbole Becker-Migdisova, 1949 (Lower or Middle Jurassic of Ta­ jikistan), Elkinda Shcherbakov, 1988 (Lower Cretaceous, Transbaikalia, Siberia) and Hy- laeoneura Lameere & Severin, 1897 (Lower Cretaceous, Belgium) are known from the Mesozoic. Elkinda and Hylaeoneura differ from the two species of Liassotettigarcta in the presence of numerous branches and crossveins in the apical third of the forewing (Shcher­ bakov, 1988; Lameere & Severin, 1897).Shuraboprosbole differs from Liassotettigarcta in its veins MP and CuA fused on a long distance (Becker-Migdisova, 1949). Turutanovia is based on the apical third of a forewing. It has no clear diagnostic character preserved and cannot be compared with the other Tettigarctidae. Cicadoprosbole differs from Lias­ sotettigarcta in its distinctly wider costal area between CP + C and ScP + R and in its shorter distal branches of RA (Becker-Migdisova, 1947). Liassotettigarcta africana sp. n. is the first record of the Tettigarctidae in Africa. Ar- chitettix Hamilton, 1990 from the Lower Cretaceous of Brazil is attributed to the Cicado- prosbolidae (Hamilton, 1990). The two extant species of this family are now restricted to Australia and Tasmania (Moulds, 1990). This group is also known in the Mesozoic and Cenozoic of Laurasia. The present discovery suggests that it was more widespread. Ne­ vertheless, further discoveries of fossil taxa would be necessary to analyse the palaeobio- geography of the Tettigarctidae. Fossil cicada-like insects are uncommon almost everywhere, thus the present first record suggests that “Merbah el Asfer” could become an important locality for Lower Cretaceous palaeoentomology.

REFERENCES

Barale G., Philippe M., Tayech -M annai B .& Z arbout M. 1997: Découverte d’ une flore à Ptéridophytes et Gymnospermes dans le Crétacé inférieur de la région de Tataouine (sud-tunisien). C. R. Acad. Sci. (Paris) (II) 325: 221-224. Barnaba P.F. 1965:Studio stratigraphico sul Cretaceo della Tunisia méridionale. Rev. Ital. Paleont. Stratigr. 71: 883-922.

597 Becker-M igdisova E.F. 1947: Cicadoprosbole sogutensis g. n., sp. n. - A transitional form between the Prosbolidae and the Recent . Dokl. Akad. Nauk SSSR 55: 441-444 (in Russian). Becker-M igdisova E.F. 1949: Mesozoic Homoptera from Central Asia. Trudy Paleozool. Inst. (Lenin­ grad) 40: 1-68 (in Russian). Ben Ismail M.H. 1991:Les bassins mésozoïques (Trias et Aptien) du sud de la Tunisie: Stratigraphie inté­ grée: caractéristiques géophysiques et évolution géodynamique. Thesis, Université de Tunis II, Tunis, 446 pp. Bouaziz S., Donze P., G hanmi M. & Zarbout M. 1989: La série à dominante continentale (Oxfordien à Cé­ nomanien) de la falaise du Dahar (Sud tunisien): son évolution du Tebaga de Médenine à la frontière tripolitaine. Géol. Méditerran. 15: 67-76. Brothers D.J. 1992: The first Mesozoic Vespidae (Hymenoptera) from the southern hemisphere, Bot­ swana. J. Hymenopterist Res. 1: 119-124. Busson G. 1967:Le Mésozoïque saharien. I. L'extrême-sud tunisien. Géologie 8. Centre Recherche sur les Zones arides, CNRS, Paris, 194 pp. Dworakowska I. 1988: Main veins of the wings of Auchenorrhyncha. Entomol. Abh. Staatl. Mus. Tierkde Dresden 52: 63-108. H amilton K.G.A. 1990: Homoptera. In Grimaldi D.A. (ed.): Insects from the Santana Formation, Lower Cretaceous, of Brazil. Bull. Am. Mus. Nat. Hist. 195: 191 pp. Kukalová -Peck J. 1991: Fossil history and the evolution of hexapod structures. In Naumann I.D., Crane P.B., Lawrence J.F., Nielsen E.S., Spradbery J.P., Taylor R.W., Whitten M.J. & Littlejohn M.J. (eds): The Insects o f Australia. A Textbook for Students and Research Workers. 2nd Melbourne ed. Vol. I. University Press, Melbourne, pp. 141—179. Lameere A. & Severin G. 1897: Les insectes de Bernissart. Ann. Soc. Entomol. Belg. 41: 35-38. M oulds M.S. 1990:Australian . New South Wales University Press, Kensington, 217 pp. N el A. 1996: Un Tettigarctidae fossile du Lias européen (Cicadomorpha, Cicadoidea, Tettigarctidae). Ecole Pratique des Hautes Etudes, Biologie et Evolution des Insectes 9: 83-94. Paulian R. 1969: Sur quatre empreintes fossiles de Coléoptères du Jurassique supérieur de Madagascar. Mém. Acad. Malgache 43: 127-129. Peybernes B., A lmeras Y., Ben Y oussef M., Kamoun F., M ello J., Rey J. & Zargouni F. 1985: Nouveaux éléments de datation dans le Jurassique du Sud tunisien (plate-forme saharienne). C. R. Acad. Sci. (Paris) (II) 300: 113-118. Rayner R.J. 1987: March flies from an African Cretaceous springtime. Lethaia 20: 123-127. Rayner R.J. & W aters S.B. 1989: A new from the Cretaceous of Botswana. Palaeontology 32: 669-673. Rayner R.J. & W aters S.B. 1991: Floral sex and the fossil record. Naturwissenschaften 78: 280-282. Rayner R.J., W aters S.B., M cKay I.J., Dobbs P.N. & Shaw A.L. 1991: The Mid-Cretaceous palaeoenvi- ronment of central Southern Africa (Orapa, Botswana). Palaeogeogr., Palaeoclimatol., Palaeoecol. 88: 147-156. Schlüter T. 1990: Fossil insect localities in Gondwanaland. Entomol. Gen. 15: 61-76. Shcherbakov D.Y e. 1981: Diagnostics of the families of the Auchenorrhyncha (Homoptera) on the basis of the wings. I. Fore wing. Entomol. Rev. 60: 64-81. S hcherbakov D.Y e. 1988: New cicadas (Cicadina) from the Later Mesozoic of Transbaikalia. Paleont. J. 1988: 52-63. Solignac M. & B erkaloff E. 1934: Carte géologique provisoire de la Tunisie au 1/200000: Medenine. Publication de la Direction générale des Travaux publics, Tunis. T eixeira C. 1975: Sur une larve d’insecte fossile (Coléoptère) du “Karroo” de l’Angola. Bolm Soc. Geol. Port. 19: 131-133. W aters S.B. 1989a: A Cretaceous dance fly (Díptera: Empididae) from Botswana. Syst. Entomol. 14: 233-241. W aters S.B. 1989b: A new hybotine dipteran from the Cretaceous of Botswana. Palaeontology 32: 657-667. Zarbout M. 1994: In Zarbout M. & Ghanmi M.: Carte géologique de Bir Touila i/100000. Service Géologique de la Tunisie (Office Nationale des Mines).

Received March 13, 1998; accepted July 22, 1998

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