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Phylogeny and the Chlorocystini (Sensu Stricto) (Homoptera

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Phylogeny and the Chlorocystini (Sensu Stricto) (Homoptera Contributions to Zoology, 65 (4) 201-231 (1995) SPB Academic Publishing bv, Amsterdam The phylogeny and taxonomic status of the Chlorocystini (sensu stricto) (Homoptera, Tibicinidae) A.J. de Boer Institute for Systematics and Population Biology (Zoölogisch Museum), University of Amsterdam, P.O. Box 94766, 1009 GT Amsterdam, The Netherlands Keywords: Homoptera, Chlorocystini, Prasiini, phylogeny, taxonomy Abstract PAUP 3.1.1 (Swofford, 1993) a été utilisé pour analyser les don- dans les Prasia Muda été choisis nées; cetteanalyse genres et one comme “outgroups”. Les résultats obtenus lors de cette analyse The “Baeturia and related genera complex”, as defined earlier par ordinateur ont été légèrement modifiés a posteriori, pour (De Boer, 1990) by shared aedeagal characters, is identified as favoriser certains caractères supposés phylogénétiquementim- the tribe Chlorocystini (sensu stricto). The Prasiini (sensu stric- fluctuants. portants, au dépens de caractères puissamment Ces to) are identified as the sister group of the Chlorocystini (sensu modifications finales été réalisées à l’aide du ont programme stricto), while the genus Muda is recognized as the nearest out- d’ordinateur MacClade 3.0 (Maddison & Maddison, 1992). Une The and of the sister and group. phylogeny biogeography group matrice complète des données ainsi qu’uneliste des caractères et outgroup is briefly discussed. Baeturia kuroiwae Matsumura is des “character states” figurent dans un Appendix; on se rap- A transferred to the genus Muda. phylogeneticreconstruction of porte à des publications précédentes pour la description et pour all 147 species of the Chlorocystini (sensu stricto) is presented, l’illustration de ces caractères. based on 154 characters and 409 character states. The computer program PAUP 3.1.1 (Swofford, 1993) was used for analysing the data; the genera Prasia and Muda were used as outgroups in this analysis. The results obtained from the computer analysis Introduction were slightly modified a posteriori, favouring some presumably phylogenetically important characters over strongly fluctuating This paper presents a cladogram for the “Baeturia ones.These final modifications were carried out with the aid of and related complex", a monophyletic the computer program MacClade 3.0 (Maddison & Maddison, genera A of tibicinid cicadas from the New Guinean- 1992). complete data matrix and a list of characters and group character in for states are given an appendix; descriptions and western Pacific This here identified region. group, illustrations of these characters oneis referred to previous publi- the selected for as Chlorocystini (sensu stricto), was cations. area-cladistic study of an area comprising Maluku, New Guinea, and East-Melanesia. A well-solved Résumé cladogram is essential for such a study. The phylogenetic and biogeographic data of the genus Le “complexe de Baeturia et des apparentés”, tel and its sister genres que complex presumed group, combined précédemment défini sur la base de caractères aedéagaux par- with such data of the Cosmopsaltriaria (a similarly tagés (De Boer, 1990), est ici identifié comme tribu Chlorocystini distributed of monophyletic group cicadas) are (sensu stricto). La tribu Prasiini (sensu stricto) est considérée used in cladistic of New Guineaand des tandis an area analysis comme groupe-frère Chlorocystini(sensu stricto), que of Indo-Melanesian le genre Muda est pris en qualité de plus proche “outgroup”. adjacent areas the region (De Sont brièvement discutées la phylogénieet la biogéographie du Boer, 1995d). In that paper the area cladograms of groupe-frère et du “outgroup”. L’espèce Baeturia kuroiwae these groups are related to the palaeogeographical Matsumura est transférée au genre Muda. Est proposée une data of the region. reconstruction phylogénétiquede l’ensemble des 147 espèces de The genus under 147 Chlorocystini (sensu stricto), reconstruction basée sur 154 carac- complex study comprises into 14 Taxonomie revi- tères et 409 “character states”. Le programme d’ordinateur species, classified genera. 202 A.J. de Boer - Phytogeny and taxonomicstatus of Chlorocystini s. s. sions, with descriptions of the species, have been are presumably not monophyletic, most of the in for most of these in tribe are close- published separate papers generaplaced any single probably Aedeastria 1990 Boer, related to each and the classifica- genera: De Boer, (De 1990; ly other, present 1993a), Guineapsaltria De Boer, 1993 (De Boer, tion into tribes can certainly form the basis for 1993b), Gymnotympana Stài, 1861 (De Boer, modern phylogenetic systematic work. The current 1995a), Mirabilopsaltria De Boer, 1995 (De Boer, subdivision of cicadas in six families (Cicadidae, 1995b), Papuapsaltria De Boer, 1995 (De Boer, Platypedidae, Plautillidae, Tettigadidae, Tettigarc- 1995c), Scottotympana De Boer, 1991 (De Boer, tidae, and Tibicinidae) is even more disputed than 1991), and Thaumastopsaltria Kirkaldy, 1900 (De the classification in tribes, although the two largest Boer, 1992a). The large genus Baeturia Stài, 1866 families, the Cicadidae and the Tibicinidae, and the was subdivided into seven monophyletic species smallfamily Tettigarctidae are generally considered groups that were discussed in separate papers: the to be sound groups (see Duffels, 1993). bloetei group (De Boer, 1989), conviva group (De Several tibicinid genera from New Guinea and Boer, 1986), exhausta group (De Boer, 1994a), gut- Eastern Australia were recognized to form one tulinervis group (De Boer, 1994b), loriaegroup (De monophyletic group, based on apomorphous ae- Boer, 1994c), nasuta group(De Boer, 1982; 1994d), deagal characters (De Boer, 1990). This group, of viridis Boer, the Australian which the in four and group (De 1992b); genera were previously placed genera of the complex (Chlorocysta West wood, different tribes (viz., the Chlorocystini, Gym- 1851, Cystopsaltria Goding & Froggatt, 1904, Cys- notympanini, Hemidictyini, and Taphurini), was tosoma Westwood, 1842, Glaucopsaltria Goding & not given the official status of tribe or subtribe, since its Froggatt, 1904, Owra Ashton, 1912, and Venustria sister-group relationships were not known. will Goding & Froggatt, 1904) be treated in a single Pending the discovery of its sister group, the group in discussion of the the of “Baeturia and paper (De Boer, prep.). For a was given preliminary name characters used in the present phylogenetic analysis related genera complex." one is referred to these taxonomie revisions. In order to polarise the characters, two out- have been selected. These dis- groups outgroups are A quest for the sister group of the “Baeturia and cussed preceding the phylogenetic analysis proper. related genera complex” Some short remarks on their phylogeny and bio- geography are made and some of the consequences The species of the “Baeturia and related genera of the presumed relationships between ingroup and complex" possess, apart from their apomorphous for the in- which outgroups traditional classifications are aedeagal shape, one character separates them dicated. from the vast majority of other tibicinid cicadas: they lack a medial uncus lobe. The presence of a distinct and posteriorly directed medial uncus lobe, The existing classification of cicadas often larger than either the lateral uncus lobes or the claspers, is found throughout the cicadas, both The higher classification of cicadas, as presented in in the Cicadidae and in the Tibicinidae. This the catalogues of Metcalf (1963a, b) and Duffels & character is very unstable within the Cicadidae Van der Laan is (1985) mainly based on the work of since presence and absence often occur among between related it be Distant, performed roughly 1888 and closely taxa, but can possibly regarded 1920. Many of the "divisions" proposed by Distant plesiomorphous in the Tibicinidaewhere a total ab- survive, virtually unchanged, as tribes in the sence occurs less frequently. The mere size of this modern classification. This classification thus pre- lobe varies strongly within both families, which tribes in dates phylogenetic systematics and the thus makes it questionable whetherthe lobes as found formed rather are phenetic than phylogenetic these families all have a monophyletic origin. groups. Nevertheless, though most of these tribes The “Baeturia and related genera complex" Contributions to Zoology, 65 (4) - 1995 203 of cicadas. The first five shares the absence of a medial uncus lobe with the tered in other groups Southeast Asian Muda 1897 and characters all be for the genera Distant, might apomorphies Kumanga Distant, 1905, while this lobeis either ab- “Baeturia and related genera complex" and the sent or strongly reduced in size in the oriental Prasiini together (4-8 in Fig. la), but then each of must have reversed several times in Prasiini (consisting of the genera Arfaka Distant, the characters swollen 1905, Jacatra Distant, 1905, Lembeja Distant, these two groups. The angularly post- is also found in Muda. One character in 1892, and Prasia Stài, 1863, see De Jong, 1985). clypeus cruciform eleva- Tibicinid species without a distinct medial uncus particular, viz., the very narrow foundin several far- tion the close lobe can also be geographically on mesonotum, strongly suggests a Prasiini and the ther removed genera, for example Fractuosella relationship between the oriental Boulard, 1979, AbromaStài, 1866 (Boulard, 1979), “Baeturia and related genera complex". That and Dinarobia Mamet, 1957(Orian, 1954) from the remarkable feature is shared by the
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