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Ent17 4 343 348 (Shcherbakov).Pmd Russian Entomol. J. 17(4): 343348 © RUSSIAN ENTOMOLOGICAL JOURNAL, 2008 Review of the fossil and extant genera of the cicada family Tettigarctidae (Hemiptera: Cicadoidea) Îáçîð âûìåðøèõ è ñîâðåìåííûõ ðîäîâ öèêàä ñåìåéñòâà Tettigarctidae (Hemiptera: Cicadoidea) D.E. Shcherbakov Ä.Å. Ùåðáàêîâ Paleontological Institute, Russian Academy of Sciences, Profsoyuznaya ul. 123, 117647 Moscow, Russia. E-mail: [email protected] Ïàëåîíòîëîãè÷åñêèé èíñòèòóò ÐÀÍ, Ïðîôñîþçíàÿ óë. 123, 117647 Ìîñêâà, Ðîññèÿ. KEY WORDS: Hemiptera, Cicadomorpha, Cicadoidea, Tettigarctidae, fossil, phylogeny, venation, Mesozoic, Jurassic, Cretaceous, Paleogene. ÊËÞ×ÅÂÛÅ ÑËÎÂÀ: Hemiptera, Cicadomorpha, Cicadoidea, Tettigarctidae, èñêîïàåìûå, ôèëîãåíèÿ, æèëêîâàíèå, ìåçîçîé, þðà, ìåë, ïàëåîãåí. ABSTRACT. Tettigarctidae, the most primitive fam- Nel, 1996, syn.n. Ïðåäëîæåíî íàçâàíèå Shaanxiarcta ily of Cicadoidea, recorded since terminal Triassic (ca. nom.n. äëÿ Involuta Zhang, 1993, non Cox, 1931.Çàäíåå 200 Myr) and ancestral to singing cicadas (Cicadidae), êðûëî èç ñðåäíåãî ýîöåíà Ýêôåëüäà (Ãåðìàíèÿ) is briefly characterized. Seventeen described fossil tet- ïðåäïîëîæèòåëüíî îòíåñåíî ê ñîâðåìåííîé òðèáå tigarctid genera are keyed and compared to the only Tettigarctini. Îäèííàäöàòü âûìåðøèõ ðîäîâ èñêëþ- extant genus (hairy cicadas); some fossil taxa are com- ÷åíû èç ñîñòàâà ñåìåéñòâà. mented. Based on tegminal venation, the family is di- vided into two subfamilies, Cicadoprosbolinae with tribes Introduction Cicadoprosbolini stat.n., Architettigini trib.n. and Tu- rutanoviini trib.n., and Tettigarctinae with tribes Prota- Tettigarctidae is a relict family, comprising the only banini stat.n., Meunierini and Tettigarctini. Sunotettig- extant genus of hairy cicadas with two species restricted arcta kudryashevae sp.n. (Upper Jurassic of Karatau) to the mountains of Tasmania and South-East Australia. and Magrebarcta gen.n. for Liassotettigarcta africana Tettigarcta is the most primitive living member of the Nel, Zarbout, Barale et Philippe are described. Liassoci- superfamily Cicadoidea, autplesiomorphic in the prono- cada Bode, 1953 = Liassotettigarcta Nel, 1996, syn.n. tum expanded posteriorly, venation generalized, male Shaanxiarcta nom.n. is proposed for Involuta Zhang, genitalia with harpagones, tarsal empodia present, and 1993, non Cox, 1931. The hindwing from the Middle thoracic ganglia separated [Evans, 1941; Moulds, 1990, Eocene of Eckfeld, Germany is tentatively placed in the 2005]. Unlike Cicadidae, hairy cicadas (of both sexes) extant tribe Tettigarctini. Eleven fossil genera are ex- produce only substrate-transmitted vibrational signals cluded from the family. [Claridge et al., 1999] and lack auditory tympana. They ÐÅÇÞÌÅ. Äàíà êðàòêàÿ õàðàêòåðèñòèêà Tettig- prefer cool to cold climates and are confined mainly to arctidae, ñàìîãî ïðèìèòèâíîãî ñåìåéñòâà Cicadoidea, subalpine forests; adults are active at dusk and after dark èçâåñòíîãî íà÷èíàÿ ñ òåðìèíàëüíîãî òðèàñà (îêîëî (hiding under loose bark during the day), common even in 200 ìëí. ëåò íàçàä) è ïðåäêîâîãî äëÿ ïåâ÷èõ öèêàä snowy season, and tolerate subzero temperatures [Moulds, (Cicadidae). Ïðèâåäåíû êëþ÷è äëÿ ñåìíàäöàòè îïè- 1990]. Tettigarcta was separated into a tribe by Distant ñàííûõ âûìåðøèõ ðîäîâ òåòòèãàðêòèä è åäèíñòâåí- [1905], raised to subfamily rank by Tillyard [1926] and to íîãî ñîâðåìåííîãî; äàíû çàìå÷àíèÿ ïî íåêîòîðûì a full family by Becker-Migdisova [1947: 448]. âûìåðøèì òàêñîíàì. Ïî æèëêîâàíèþ ïåðåäíèõ êðû- Tettigarctidae were first recognized in the fossil record ëüåâ ñåìåéñòâî ðàçäåëåíî íà äâà ïîäñåìåéñòâà, by Zeuner [1944] (Cenozoic of Europe) and Becker-Migd- Cicadoprosbolinae ñ òðèáàìè Cicadoprosbolini stat.n., isova [1949] (Mesozoic of Asia); earlier authors described Architettigini trib.n. è Turutanoviini trib.n., è Tettig- them as Diptera, Megaloptera, or Cicadidae. Becker-Migd- arctinae ñ òðèáàìè Protabanini stat.n., Meunierini è isova [1947] first described Cicadoprosbole in Prosbol- Tettigarctini. Îïèñàíû Sunotettigarcta kudryashevae idae, but later transferred it to Tettigarctidae along with sp.n. (âåðõíÿÿ þðà Êàðàòàó) è Magrebarcta gen.n. äëÿ three other Mesozoic genera [Becker-Migdisova, 1949]. Liassotettigarcta africana Nel, Zarbout, Barale et Evans [1956] separated Cicadoprosbole at the family Philippe. Liassocicada Bode, 1953 = Liassotettigarcta level, and some other authors followed him, placing most Printed in 2009. or all Mesozoic genera into Cicadoprosbolidae [e.g. Hamil- 344 D.E. Shcherbakov Table. Some tegminal characters of tettigarctid tribes and genera compared to the type genera of both subfamilies and to Cicadidae. Òàáëèöà. Íåêîòîðûå ïðèçíàêè ïåðåäíåãî êðûëà â òðèáàõ è ðîäàõ Tettigarctidae â ñðàâíåíèè ñ òèïîâûìè ðîäàìè îáîèõ ïîäñåìåéñòâ è Cicadidae. C as in Cicadoprosbole; T as in Tettigarcta; i intermediate state; a, b, d aberrant; ? unknown; a and T homoplastic to a and T of extant cicadoids; Cc last appearance of C condition in transformation series. First four taxa show more characters in Cicadoprosbole state, last two more characters in Tettigarcta condition, and Protabanini are about halfway between the nominate genera of two subfamilies. C êàê ó Cicadoprosbole; T êàê ó Tettigarcta; i ïðîìåæóòî÷íîå ñîñòîÿíèå; a, b, d óêëîíÿþùååñÿ; ? íåèçâåñòíî; a è T ïàðàëëåëèçì ïî îòíîøåíèþ ê a è T ñîâðåìåííûõ öèêàäîèäîâ; Cc ïîñëåäíåå ïîÿâëåíèå ñîñòîÿíèÿ C â òðàíñôîðìàöèîííîì ðÿäó. Ó ÷åòûð¸õ ïåðâûõ òàêñîíîâ ïðåîáëàäàþò ïðèçíàêè â ñîñòîÿíèè êàê ó Cicadoprosbole, ó äâóõ ïîñëåäíèõ â ñîñòîÿíèè êàê ó Tettigarcta, à Protabanini íàõîäÿòñÿ ïðèìåðíî íà ïîëïóòè ìåæäó òèïîâûìè ðîäàìè äâóõ ïîäñåìåéñòâ. ton, 1990, 1996]. However, it is better to treat these groups Architettix) [Hamilton, 1990] provided with jumping as subfamilies [Boulard & Nel, 1990], because diverse armature (long laterotibial spines) more developed than Mesozoic forms fill the gap between Cicadoprosbole in recent cicadoids. and Tettigarcta (Table). The hair covering in diverse, well preserved tettig- The oldest records of Tettigarctidae s.l. are from the arctids was apparently never as long and abundant as in earliest Jurassic and terminal Triassic (ca. 200 Myr) of modern hairy cicadas (Tettigarcta spp.). A common Eurasia. Jurassic insects of Gondwana are virtually name of Mesozoic hairy cicadas could rather be ap- unknown, and fossil tettigarctids in the Southern Hemi- plied to another, unrelated family of large, cicada- and sphere were found only in the Lower Cretaceous of even moth-like Auchenorrhyncha, Palaeontinidae, be- Brazil [Hamilton, 1990; Menon, 2005] and Tunisia cause at least some palaeontinids (as seen in specimens [Nel et al., 1998]. There are no tettigarctids among ten from the Upper Jurassic of Karatau) were extremely auchenorrhynchan specimens from the well-known Cre- hairy, with very long beak and unarmed legs. They taceous locality Koonwarra, Australia. The family per- presumably were crepuscular and well camouflaged on sisted in Europe at least until the Middle Eocene (ca. 45 tree trunks, like living Tettigarcta, and the niche parti- Myr) [Wappler, 2003], whereas the earliest record of tion between Palaeontinidae and Tettigarctidae in the Cicadidae is from the Paleocene (ca. 65 Myr) of North Mesozoic might be analogous to that between Tettig- America [Cooper, 1941]. Eskov [1987: 98] briefly sum- arcta and Cicadidae now. marized the former distribution of Tettigarctidae in the Tettigarcta was contrasted to Cicadidae as the most Northern Hemisphere. Nel [1996] listed extinct genera primitive living genus of the superfamily, and Mesozoic which belong to (or are excluded from) the family. tettigarctid genera permit to trace the ancestry of Cica- Tettigarctids were structurally diverse and not rare doidea even more clearly. All three extant superfamilies in the Jurassic and Cretaceous, like are singing cicadas of Cicadomorpha Clypeata (Cercopoidea, Cicadoidea, in the Cenozoic and nowadays. They varied in size from Membracoidea) descended from the fourth, Mesozoic small forms, with transparent wings less than 15 mm Hylicelloidea [Shcherbakov, 1996]. Cercopoids and long (Turutanovia, Architettix) to large cicadas (larger cicadoids united by several sound synapomorphies than modern Tettigarcta) with dark-patterned wings up [Emeljanov, 1987] enter the record almost synchro- to 60 mm long (Figs 14). Cretaceous Elkinda shows nously and possibly represent closely related lineages; tegminal veins profusely branched, like in some cica- membracoids had separated from hylicelloids some- dids (e.g. Polyneura). Tettigarctids had the beaks usual- what later. Phylogenetic position of tettigarctids was ly very long, and hind legs at least sometimes (e.g. also discussed by Hamilton [1996]. Review of fossil and extant Tettigarctidae 345 3 All veins profusely branched beyond nodal line; about 30 apical cells. CuA with more than 2 branches; apical cell behind posterior CuA branch much wider than next apical cells. M and CuA forming short stalk beyond basal cell ................................................... *Elkinda Shcherbakov Less than 15 apical cells. CuA with 2 branches; apical cell behind CuA2 narrower than next apical cell .............. 4 4 Transverse ir nearer to nodal line than to wing tip. Basal CuA1 section converging to M3+4. Apical cell behind CuA2 short and narrow. RA bent at nodal line. Both M and CuA slightly sigmoidal, cross-joined beyond basal cell ....................... *Cicadoprosbole Becker-Migdisova Reclined ir nearer to wing tip than to nodal line. Basal CuA1 section subparallel
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