SYSTEMATICS Placement of the Faltala Oman Leafhopper Group (Hemiptera: Cicadellidae: Deltocephalinae) with Descriptions of Three New Species 1 2 J. N. ZAHNISER AND M. D. WEBB Ann. Entomol. Soc. Am. 97(4): 667Ð674 (2004) ABSTRACT Three new species of the Faltala Oman leafhopper group, Kramerana junina n. sp. and K. adusta n. sp. from the Andes Mountains of Peru and Clorindaia brasileira n. sp. from the Serra da Mesa of Brazil, are described. The tribal placement of the group and the high incidence of brachyptery and reduced (or absent) ocelli in the group are discussed. A revised key to the Faltala group genera and additional characters for Kramerana are provided. KEY WORDS Deltocephalinae, Athysanini, Hecalini, Neotropical, new species OMAN (1938) DESCRIBED THE genus Faltala based on one Placement of the Faltala Group species from Argentina. Since then, four additional LinnavuoriÕs (1959) placement of the Faltala group closely related genera (Kramerana, Virganana, and in Athysanini was based on the Y-shaped connective, Aequicephalus described by DeLong and Tham- a character that cannot be considered a synapomor- bimuttu [1973], and Clorindaia described by Linna- phy of this poorly deÞned tribe, as members of several vuori[1975a]) and 12 further specieshave been de- deltocephaline tribes, indeed most, possess a similar scribed. Linnavuori and DeLong (1977) placed all Y-shaped connective. Later, Linnavuori (1975) also genera in the Faltala group of the Athysanini (Delto- noted the similarity in cephalic structure of Faltala and cephalinae). Linnavuori and DeLong (1977) gave a Clorindaia to Hecalini (Deltocephalinae) (see be- diagnosis of this group and a key to genera, and low). Recent morphological phylogenetic analyses of Blocker and Fang (1992) provided a useful synopsis of Macrostelini (Knight and Webb 1993) and Deltoceph- the included species and their distributions. In the alinae and related subfamilies (Zahniser and Dietrich, current study, two new species of Kramerana and one unpublished data) revealed several characters, espe- new species of Clorindaia are described, and newly cially of the female genitalia, that are useful in deÞning recorded details of the leg chaetotaxy and female some tribes and higher groups. In their analysis, Zah- genitalia for Kramerana are provided. The tribal place- niser and Dietrich recovered a clade of grass-special- ment of the Faltala group and the high incidence of izing leafhoppers, including the deltocephaline tribes brachyptery and reduced (or absent) ocelliinthe Hecalini, Stenometopiini (ϭStirellini), and Macros- group are discussed. In the course of identifying the telini sensu Knight and Webb in part (Balclutha species described in this work, we found that some of Kirkaldy) and the subfamilies Arrugadinae, Eupelici- the characters provided in the key to genera by Lin- nae (Eupelicini, Paradorydiini, Dorycephalini, and navuori and DeLong (1977) do not easily distinguish Listrophorini), and Drakensbergeninae, based largely the genera. A simpliÞed key is provided that reßects on the following features of the female genitalia (with previous generic concepts, adds some newly discov- some reversals, as indicated): 1) Þrst valvulae with a ered characters, and discards some ambiguous char- deÞnitely delimited apicoventral sculptured area, sub- acters. All of these genera are apparently closely re- triangular in shape (Figs. 12 and 30; reversed in Eu- lated, and as more species are identiÞed, some may be pelicini and Drakensbergeninae); 2) Þrst valvulae dor- sal sculpturing pattern maculose or scale-like, with the found to be paraphyletic. maculae or scales not overlapping and clearly sepa- rated from one another (Fig. 11) (reversed in Eupeli- cini); 3) Þrst valvulae dorsal sculpturing submarginal (Figs. 11, 12, and 30) (also present in Deltocephalinae: 1 Department of Entomology, 320 Morrill Hall, University of Chiasmusini and Doraturini); 4) second valvulae lack- Illinois at Urbana/Champaign, Urbana, IL 61801 (e-mail address: ing large dorsal teeth (Figs. 10, 28, and 33) (reversed [email protected]). in Eupelicini, Drakensbergeninae, and Dorycephalini 2 Department of Entomology, The Natural History Museum, Crom- well Road, London, SW7 5BD, United Kingdom (e-mail address: [Dorycephalus Kouchakewitch]); 5) pygofer macro- [email protected]). setae absent or reduced in number. 0013-8746/04/0667Ð0674$04.00/0 ᭧ 2004 Entomological Society of America 668 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 97, no. 4 Figs. 1–5. K. junina n. sp. (1) Male, dorsal habitus. (2) Female, same. (3) Female, lateral habitus. (4) Nymph, dorsal habitus. (5) Male, face. All scale bars ϭ 1.0 mm. All the Faltala group taxa have the characters listed lini and, in combination with an elongate connective, above and, together with some hecalines, have the the following were also removed: Tenucephalus De- head produced anteriorly; the anterior margin of head Long, Onura Oman, Acrolithus Freytag and Ma, and foliacious, carinate, or transversely striate; the tergum Egenus Oman. All of these taxa were transferred by with several longitudinal stripes; and strong sexual Hamilton to Deltocephalini sensu Hamilton (1975). Of dimorphism in head length and brachyptery (see be- these genera, Hecullus and Tenucephalus have been low). However, although the inclusion of the Faltala examined in the current study (Hecullus sp.: Figs. 33 group in Hecalini would Þt in with this wider concept and 34) and found to have the same characters of the of the group (see also Linnavuori 1959, Freytag and female genitalia noted above for the clade found by Ma 1988, Nielson 1996), it would not conform to the Zahniser and Dietrich (unpublished data). most recent and more restrictive deÞnition of Ham- In addition to the morphological characters cited ilton (2000). This deÞnition was based on the ocelli above, the limited ecological data for the Faltala group being closer to the eyes than to the facial sutures. also support placement in the grass-feeding clade de- Although this feature is consistently observed in Heca- scribed above. All records of hosts or habitat for spe- lini sensu Hamilton, it also occurs in other Deltoceph- cies of the Faltala group indicate that they feed on alinae, e.g., Balclutha Kirkaldy, Cerrillus Oman, Drak- grasses. DeLong and Thambimuttu (1973) reported ensbergena Linnavuori, and Scaphytopius Ball. that they “occur upon grasses and are found in mead- Contrary to Linnavuori (1975b), Hamilton (2000) also ows where sheep or cattle are grazing.” Of the species considered Hecalini to be closely related to Eupelici- described in this study, the two from Peru were swept nae (Dorycephalini) based on the following synapo- or vacuumed from grasses, and the Brazilian species morphies: male apodemes of second sternite as long as was collected in yellow pan traps in the Serra da Mesa tergal apodemes, yet narrow; “anal tube” withdrawn region, characterized by cerrado/cerradinho vegeta- into the pygofers; male pygofers with broad ventral tion, which is typiÞed by grasses, herbs, bushes, and lobes that meet or cross at lower edges; ovipositor small trees. curiously shaped, being straight or concave ventrally In conclusion, based on the above evidence, the and hump-backed near midlength (Readio 1922, Plate Faltala group taxa, together with Hecullus and Tenu- 29, Figs. 4Ð8). cephalus, are at present a nondeÞned assemblage that Based on the absence of the above ocellicharacter, could be placed as a subgroup in Hecalini sensu lato, Hamilton (2000) removed Hecullus Oman from Heca- having many of the characters noted above for this July 2004 ZAHNISER AND WEBB:THE Faltala OMAN LEAFHOPPER GROUP 669 Figs. 6–13. K. junina n. sp. (6) Connective and styli, ventral view. (7) Valve and plate, ventral view. (8) Aedeagus, lateral view. (9) Female seventh sternite. (10) Second valvulae. (11) Detail of Þrst valvula sculpturing pattern. (12) Apex of Þrst valvula. (13) First valvula, lateral view. group, but lacking the distinct hump-backed ovipos- considered adult characters in hemimetabolous insects, itor, ocelli closer to the eyes than the facial sutures, and that their absence in the adult is a kind of neoteny. and other characters found in Hecalini sensu stricto This might be determined genetically or in combination (Hamilton 2000). Alternatively, these taxa could form with environmental factors (e.g., temperature) affecting a separate lineage within the grass-feeding clade noted hormonal levels in the insect. Members of the Faltala above. However, for the present, these taxa are re- group corroborate the correlation of brachyptery with tained in Athysanini pending further studies. reduced or absent ocelli, because its members charac- teristically have reduced ocelli. Brachyptery Key to the Faltala Group Genera With respect to brachyptery in the grass-specializ- ing clade described above, subbrachypterous or sub- 1. Ocellinear to eyes, separated by Ϸ3ϫ their own macropterous species are prevalent in some groups diameter from eyes .............Faltala (e.g., Hecalini), but fully brachypterus species (e.g., Ocellidistantfrom eyes, 1⁄3 to 1⁄2 distance from some Dorycephalini) are less common. However, in eye to crown apex ................. 2 the Faltala group (all genera) and in Acrolithus and 2. Frontoclypeus glabrous or with indistinct mi- Hecullus (females only), fully brachypterous species crosculpturing ..............Clorindaia have been reported. An interesting character related Frontoclypeus distinctly shagreened .....3 to brachyptery is the