KEYS TO THE FAMILIES OF CICADOMORPHA AND SUBFAMILIES AND TRIBES OF CICADELLIDAE (HEMIPTERA: AUCHENORRHYNCHA)

Headnote

ABSTRACT

Illustrated keys to adults reflecting the current higher classification are provided for families of Cicadomorpha (cicadas, spittlebugs, leafhoppers, and treehoppers) and for subfamilies and tribes of Cicadellidae (leafhoppers), excluding Deltocephalinae. The following families (and superfamilies) are recognized: Cicadidae and Tettigarctidae (Cicadoidea); Aphrophoridae, Cercopidae, Clastopteridae, Epipygidae, and Machaerotidae (Cercopoidea); Aetalionidae, Cicadellidae, Melizoderidae, Membracidae, Myerslopiidae (Membracoidea). The higher classification of Cicadellidae is currently undergoing revision, but a provisional key to subfamilies and tribes (except Deltocephalinae) is provided. Two new synonymies are proposed: Signoretiinae Baker, 1915 equals Phlogisinae Linnavuori, 1979, new synonym; Iassini Walker, 1870, equals Hyalojassini Evans, 1972, new synonym.

Key Words: morphology, identification, taxonomy, phylogeny, Homoptera

RESUMEN

Se provee claves ilustradados para los adultes que reflejan la clasificación jerárquica actual para las familias de Cicadomorpha (cigarras, cercopidos, chicharas y membrácidos) y para las subfamilias y tribus de Cicadellidae (chicharras), excluyendo los Deltocephalinae. Se reconoce las familias (y superfamilias) siguientes: Cicadidae y Tettigarctidae (Cicadoidea); Aphrophoridae, Cercopidae, Clastopteridae, Epipygidae y Machaerotidae (Cercopoidea); Aetalionidae, Cicadellidae, Melizoderidae, Membracidae, y Myerslopiidae (Membracoidea). La clasificación jerárquica de la familia Cicadellidae esta actualmente bajo una revision taxonomica, pero se provee una clave provisional de las subfamilias y tribus (menos los Deltocephalinae). Se propone dos sinonimos nuevos: Signoretiinae Baker, 1915 es igual que Phlogisinae Linnavuori, 1979, sinónimia nueva; y Iassini Walker, 1870, es igual que Hyalojassini Evans, 1972, sinónimia nueva.

The hemipteran (=homopteran) infraorder Cicadomorpha comprises approximately 35,000 described species of plant sap-sucking insects distributed worldwide. Species are grouped into three superfamilies that are well established based on morphological criteria: Cicadoidea (cicadas); Cercopoidea (spittlebugs, froghoppers); and Membracoidea (leafhoppers, sharpshooters, treehoppers) (Figs. 1 and 2). Cicadomorpha may be distinguished from other Hemiptera by the following combination of characters: postclypeus enlarged; antennal pedicel small, without conspicuous sensilla, flagellum aristiform; tegulae absent; forewing anal veins usually separate from base to wing margin; middle coxae small and narrowly separated. Over 3,000 species of Cicadomorpha are recorded from temperate North America, including ca. 70 exotic species, the vast majority of which are native to Eurasia. Relatively few species are economically important, but there are some major pests, such as the glassy-winged sharpshooter, Homalodisca coagulata (Say), potato leafhopper, Empoasca fabae (Harris), and beet leafhopper, Neoaliturus (=Circulifer) tenellus (Baker). Cicadomorphans injure plants either directly through feeding (Backus 1988; Backus et al. 2005) or indirectly through transmission of plant pathogens (Nielson 1968; Maramorosch & Harris 1979).

Identification of cicadomorphan species is difficult because of their tremendous diversity and the paucity of comprehensive identification keys. Recent sampling suggests that more than 90% of the extant tropical cicadomorphan species remain undescribed (Hodkinson & Casson 1991; Dietrich & Wallner 2002, unpublished data). These, as well as a large proportion of described species, have never been included in a key. Thus, when possible at all, identification of cicadomorphan species usually requires access to a large and obscure taxonomic literature and authoritatively identified reference specimens.

Recent phylogenetic analyses (e.g., Dietrich & Deitz 1993; Dietrich 1999; Dietrich et al. 2001a, b; Shcherbakov 1996; Rakitov 1998; Hamilton 1999; Wallace & Deitz 2004; Cryan et al. 2004; Moulds, unpublished) have begun to elucidate the status and relationships of cicadomorphan family-group taxa, but the higher classification of Cicadomorpha remains controversial and no family-group classification has yet gained universal acceptance. Due in part to this controversy, few attempts have been made recently to develop keys for identifying the major cicadomorphan groups (families, subfamilies, and tribes). Although revised keys are available for treehopper family-group taxa (Deitz & Dietrich 1993; Dietrich et al. 2001b; Wallace & Deitz 2004), the most recent comprehensive key to leafhopper subfamilies and tribes is over 50 years old (Evans 1947). The most comprehensive keys to cicada and spittlebug family groups are nearly 100 years old (Distant 1912, 1914; Lallemand 1912). More recent keys are available for certain regional faunas (e.g., Evans 1966; Amifriev & Emeljanov 1988), but interpretations of higher taxa vary among regions and authors. The family-group classifications of cicadas (Moulds, unpublished) and cercopoids (Hamilton, Liang, unpublished) are currently being revised. Revisions to the higher classification of leafhoppers have also been made in recent years. Hamilton (1983) proposed a classification of Cicadellidae that included only ten subfamilies, but subsequent authors have not followed his system. Oman et al. (1990) recognized 40 subfamilies and 119 tribes in their provisional classification and world generic checklist. Subsequent to the 1985 cut-off date for the Oman et al. (1990) checklist, a new subfamily (Godoy & Webb 1994) and two new tribes (Theron 1986; Hamilton 1999) have been described. Also, several family-group taxa listed as valid by Oman et al. (1990) have more recently been treated as junior synonyms (e.g., Dietrich & Rakitov 2002; Dietrich & Dmitriev 2003; Dietrich 2004).

Fig. 1. Cicadoidea and Cercopoidea. A, Tettigarcta crinita (Tettigarctidae), Australia. B, Magicicada cassini (Cicadidae), Illinois. C,Paraphilaenus parallelus (Aphrophoridae), Kyrgyzstan. D, Clastoptera obtusa (Clastopteridae), Illinois. E, Hindola sp. (Machaerotidae), adult (top center) and tube constructed by nymph (lower left), Taiwan. F, Tomaspis sp. (Cercopidae), Mexico. G, spittle mass ofPhilaenus spumarius nymph. Photos by C. H. Dietrich.

Fig. 2. Membracoidea. A, Myerslopia chilensis (Myerslopiidae), Chile. B, Hylaius oregonensis (Cicadellidae), Oregon. C, Flexamia grammica (Cicadellidae), Illinois. D, Llanquihuea sp. (Melizoderidae), Chile. E, Aetalion reticulatum (Aetalionidae), Peru. F, Heteronotus quadrinodosus (Membracidae), Mexico. Photos by C. H. Dietrich.

The purpose of this paper is to provide up-todate keys to the families of Cicadomorpha and to subfamilies and tribes of Cicadellidae that reflect, to the extent possible, current consensus regarding the higher classifications of these groups.

MORPHOLOGY

The following section describes the basic morphological terminology used in keys to Cicadomorpha but is not intended as an exhaustive treatment. For a more detailed treatment of cicadomorphan morphology, see Kramer (1950). Alternative terminologies have been proposed by various authors (e.g., Blocker & Triplehorn 1985; Hamilton 1981; Mejdalani 1998). There is not yet a universally accepted system of morphological terminology for Cicadomorpha. For a detailed treatment of the morphology of cicadellid nymphs, see Dmitriev (2002).

Head. The cicadomorphan head is highly variable in form, particularly among leafhoppers (Figs. 3A, 3B, 4). For convenience, the term face is used to refer to the anterior part and the term crown refers to the dorsal part, although in different groups these areas are made up of parts of different sclerites. Some authors (e.g., Blocker & Triplehorn 1985) use crown and vertex interchangeably, but the term vertex is correctly applied only to the pair of sclerites posterolaterad of the frontal sutures and bearing the lateral ocelli. The rostrum (or beak), the modified labium, varies in length among taxa. The clypeus is almost always divided by the clypeal (or transclypeal) suture into two sclerites, the anteclypeus (clypellus) and postclypeus, both of which vary in texture and proportions among taxa. The postclypeus is usually not clearly divided from the frons dorsally, thus the term frontoclypeus is often used to refer to the combined sclerite, which is usually the largest structure of the head in anterior view. Immediately laterad of the clypeus on the lower part of the face is a pair of mandibular plates called the lora (singular-lorum), which vary in size, shape, and relative position. The lorum is situated on the maxillary plate, which, along with the more dorsally situated gena, forms the lateral margin of the head. In most leafhoppers, the gena and maxillary plate are fused, but a few groups of leafhoppers (e.g., Ulopinae), as well as most other cicadomorphans, have the maxillary plate and gena separated by a distinct suture or cleft. The antennae of Cicadomorpha consist of two short basal segments and an elongate flagellum, which varies in length and shape and may be subsegmented to various degrees. The base of the antenna may be partly covered dorsally by an outgrowth of the vertex called the antennal ledge, which also varies in shape and may extend onto the frontoclypeus. The eyes vary in size and shape, and the ocelli vary in their position relative to the eyes and to the anterior margin of the head. The lateral arms of the epicranial suture, usually termed frontal sutures, are reduced or lost in various taxa and their relative shape and position also varies. In many leafhoppers, these sutures extend to the ocelli and in such cases are often referred to as lateral frontal sutures (or laterofrontal sutures).

Fig. 3. Leafhopper morphology. A, dorsal habitus. B, head, anterior view (face). C-D, apex of male abdomen, Deltocephalinae, lateral and ventral views, respectively. E, Typhlocybinae, male genital capsule, lateral view. F, apex of female abdomen, lateral view (vl = valvula, vlf = valvifer, s = sternite, t = tergite). Drawings A-B modified from Oman (1949), C-E modified from Anufriev & Emeljanov (1988), F from Kramer (1950).

Fig. 4. Leafhopper head morphology. A-F, head, anterior view: A, Evacanthus (Evacanthini); B, Krisna (Krisnini); C, Tartessus (Tartessinae); D, Platyproctus (Adelungiini); E, Bathysmatophorus (Errhomenini); F, Thymbrus (Thymbrini). G-H, head and prothorax, lateral view: G, Cicadella (Cicadellini); H, Matsumurella (Athysanini). I-K, head, pronotum, mesonotum, and scutellum, dorsal view: I, Populicerus (Idiocerinae); J, Pediopsoides (Macropsinae); Oniella (Evacanthini). Drawings A-F original; G-K from Anufriev & Emeljanov (1988).

Thorax (Fig. 5). The pronotum of Cicadomorpha varies in shape, ornamentation, texture and proportions, particularly in treehoppers (Fig. 2F), which usually have a well developed posterior process that partially or completely conceals the scutellum and may also overlap the wings. The mesonotum and scutellum, divided by the scutellar suture (and often incorrectly referred to collectively as the "scutellum"), vary in proportions and the degree to which they are concealed by the pronotum. Important pleural sclerites include the proepisternum, the mesepisternum, and mesepimeron, which vary in proportions and, in the latter, may bear tubercles or other processes. The mesepisternum is usually divided into an anterodorsal anepisternum and a posteroventral katepisternum, but these two sclerites are fused in treehoppers. The forewing, or tegmen (pl., tegmina), and hind wing vary in proportions, texture, and venation. Important features of the wings include the branching pattern of veins and the shape and texture of the cells (areas between veins).

Fig. 5. Morphology of cicadomorphan thorax (from Kramer 1950): A-C, Lepyronia (Aphrophoridae), dorsal, lateral, and ventral view (in A, pronotum removed to show mesonotum). D-F, same, Paraulacizes (Cicadellidae). G, Stictocephala (Membracidae), lateral view of meso- and metathorax. Abbreviations: acs = antecostal suture; acx = antecoxal area; aes = anepisternum; anp = anterior notai wing process; aw = prealar bridge; ax = axillary sclerite; ba = basalare; bst = basisternite; ex = coxa; cxp = pleural coxal process; epm = epimeron; es = episternum; fp = furcal pit; fst = furcasternite; kes = katepisternum; mr = meron of coxa; pbs = prebasisternite; pc = precosta; pis = pleural suture; pnp = posterior notai wing process; pnt = postnotum; pocx = postcoxal bridge; prnt = pronotum; psc = prescutum; pw = postalar bridge; pwg = pleural wing groove; sp = spiracle; st = sternite; tn = trochantin; tr = trochanter; twg = tergal wing groove

The legs vary in shape, proportions, spination, and chaetotaxy (setal arrangement). Different systems have been proposed for naming the rows and groups of setae on the legs of leafhoppers (Rakitov 1998) and treehoppers (Deitz 1975). In Rakitov's system, adopted here, longitudinal rows of setae are labeled according to their position assuming that the leg is extended perpendicular to the mid-saggital plane of the body; i.e., anterodorsal (AD), posterodorsal (PD), anteroventral (AV) and posteroventral (PV) (Figs. 7G, 7J). The front femur of leafhoppers often bears additional anteromedial (AM) and intercalary (IC) setal rows (Fig. 7K). When appropriate, individual setae are numbered sequentially, beginning with the most distal (e.g., AVl referrs to the distal seta of the anteroventral row). Setal formulae are often used to describe the arrangement of enlarged setae, or macrosetae, particularly on the front tibia and hind femur. In treehoppers, the number and arrangement of rows of cucullate setae (setae with dark, hoodlike basal sockets) on the hind tibia are important for distinguishing tribes.

Abdomen. The basal segments of the abdomen and adjacent parts of the metathorax are modified, particularly in males, for sound production. In cicadas, the structure of these sound producing organs, or tymbals, varies among species. The organs of sound detection in cicadas, or tympana, are located ventrally at the base of the abdomen and may be concealed by opercula, which are flaplike outgrowths of the metathorax. Similar structures may occur in other cicadomorphans, but they are not as well developed. In some groups of leafhoppers, the shapes of the internal apodemes of male abdominal segments I-III, associated with the production of species-specific courtship signals transmitted through the substrate, are diagnostic at the species level.

Fig. 6. Leafhopper wings. A-G, forewing: A, Hortensia (Cicadellini); B, Errhomus (Errhomenini); C, Deltocephalus (Deltocephalini); D, Hamana (Scarini); E, Idiocerus (Idiocerinae); F, Typhlocyba (Typhlocybini); G, Jikradia (Coelidiinae). H-O, hind wing: H, Agallia (Agalliini); I, Macropsis (Macropsinae); J, Penestragania (Iassini); K, Typhlocyba (Typhlocybini); L, Protalebrella (Alebrini); M, Hymetta (Erythroneurini); N, Empoasca (Empoascini); O, Joruma (Jorumini). Drawings from Oman (1949).

The male genital capsule (Figs. 3C-E) consists of the tergite of segment IX, or pygofer (pygophore), and the sternite of segment IX, or valve, which may or may not be separated from each other by a suture. The pygofer varies in shape and chaetotaxy, and may bear various lobes or processes. In Membracoidea and Cercopoidea, there is a pair of posteroventral lobes called subgenital plates (or, simply, plates) connected to the posterior margin of the valve; these vary in shape, chaetotaxy, and degree of fusion to each other and to the valve. The sclerotized parts of the genitalia of Membracoidea and Cercopoidea consist of a pair of lateral styles, a median connective, and an aedeagus or penis, all of which vary in shape and proportion and may be highly modified with various processes and accessory structures. In cicadas, the styles and connective are vestigial. Segment X, which forms the major part of the anal tube, also varies in shape and proportion and may bear spines or processes.

The female ovipositor (Fig. 3F) consists of two pairs of blade-like structures, the first and second valvulae, and an outer sheath, the third valvulae (gonoplacs). The second valvulae vary in shape, proportion and armature (dorsal teeth or serrations). The first valvulae, which enclose the second, vary in shape, proportion, and texture. When not in use, the ovipositor is partially enclosed by the enlarged ninth abdominal tergite, or pygofer. The seventh abdominal sternite usually overlaps the bases of the ovipositor and pygofer ventrally and varies in shape among species in some groups.

Fig. 7. Leafhopper leg morphology. A-E, apex of hind femur, dorsal view, showing variation in macrosetal formula: A, 2+2+1; B, 2+2+1; C, 2+1+1; D, 2+1; E, 2+1. F-J, right hind tibia, posterior view, except G and J, cross-section: F-G, Bathysmatophorus (Errhomenini); H, Bothrogonia (Cicadellini); I-J, Diplocolenus (Paralimnini). K-N, left front femur, anterior view: K, Doratura (Doraturini); L, Thagria (Thagriini); M, Alebra (Alebrini); N, Xestocephalus (Xestocephalini). O-P, hind tarsus, ventral view: O, Kybos (Empoascini); P, Balclutha (Balcluthini). Drawings A-N from Rakitov (1998), copyright Russian Entomological Journal, used with permission; O-P from Anufriev & Emeljanov (1988).

IDENTIFICATION

The following key will separate adults of the superfamilies and families of Cicadomorpha. The family classification of Cicadoidea follows that of Moulds (1990); see Duffels & van der Laan (1985) for an alternative classification. That of Cercopoidea follows Metcalf (19601962); but see Hamilton (2001) for an alternative classification. That of Membracoidea follows Deitz & Dietrich (1993) and Hamilton (1999).

KEY TO FAMILIES OF CICADOMORPHA

The interpretation of leafhopper subfamilies and tribes in the following key is based on that of Oman et al. (1990), but with several important exceptions that reflect ongoing revisionary work on the higher classification of Cicadellidae. Following Hamilton (1983), the typhlocybine tribe Helionini is treated as a synonym of Empoascini, and Petalocephalini is treated as a synonym of Ledrini. Following Hamilton (1983, 1999), Agalliinae, Evansiolinae, and Adelungiinae are considered synonyms of Megophthalminae; these taxa, sensu Oman et al. (1990), are here recognized as tribes within Megophthalminae (sensu lato). Neopsinae is retained at subfamily rank, following Linnavuori (1978) rather than as a tribe of Macropsinae.

The interpretation of Neocoelidiinae follows Dietrich (2003). Following Dietrich (2004), Nirvaninae is treated as a synonym of Evacanthinae, which includes tribes Evacanthini, Nirvanini, Pagaroniini, and Balbillini. Also following Dietrich (2004), Macroceratogoniini is included as a tribe of Coelidiinae rather than of Evacanthinae. Following Young (1968, 1986) and Linnavuori & DeLong (1977), Errhomeninae (= Bathysmatophorini) and Mileewinae are treated as subfamilies distinct from Cicadellinae. Thus, the present interpretation of Cicadellinae follows Young (1968, 1977) in including only the tribes Cicadellini and Proconiini.

Scarini (=Gyponini) is treated as a tribe of Iassinae following Linnavuori & Quartau (1975), rather than as a distinct subfamily. Following Hamilton (1975), Xestocephalinae is treated as a synonym of Aphrodinae. Following Dietrich & Rakitov (2002) and Dietrich & Dmitriev (2003), the following subfamilies (sensu Oman et al. 1990) are considered synonyms of Deltocephalinae: Eupelicinae, Koebeliinae, Paraboloponinae, Penthiniinae, and Selenocephalinae. The name Penthimiinae Kirschbaum, 1868, has priority over Deltocephalinae Dallas, 1870, but the latter name is provisionally retained in view of its long-term usage and because the classification of Deltocephalinae is undergoing revision.

KEY TO SUBFAMILIES AND TRIBES OF CICADELLIDAE

The monobasic subfamily Phlogisinae Linnavuori, 1979, is treated as a junior synonym of Signoretiinae Baker, 1915 (NEW SYNONYMY), and the monobasic tribe Hyalojassini Evans, 1972, is treated as a junior synonym of Iassini Walker, 1870 (NEW SYNONYMY), based on shared characters given in the key.

Two family-group taxa described after the publication of the Oman et al. (1990) checklist are added: Tinterominae, a subfamily described by Godoy & Webb (1994); and Sagmatiini, a tribe of Euacanthellinae described by Hamilton (1999). The endemic South African tribe Equeefini Theron, 1986 (Coelidiinae), is excluded because sufficient material was not available for study. The subfamilies Acostemminae, Arrugadinae, Drakensbergeninae, Mukariinae, and Stegelytrinae, and the tribe Paraphrodini (Aphrodinae) are also excluded. These taxa key to Deltocephalinae and are the subject of an ongoing phylogenetic study and revision of the "deltocephaline-like" leafhoppers (Zahniser & Dietrich, unpublished).

KEY TO SUBFAMILIES AND TRIBES OF CICADELLIDAE

KEY TO SUBFAMILIES AND TRIBES OF CICADELLIDAE

KEY TO SUBFAMILIES AND TRIBES OF CICADELLIDAE

KEY TO SUBFAMILIES AND TRIBES OF CICADELLIDAE

KEY TO SUBFAMILIES AND TRIBES OF CICADELLIDAE

References

ACKNOWLEDGMENTS

I am grateful to D. Dmitriev, R. Rakitov, D. Takiya, J. Zahniser, and the participants in the Gainesville "Homoptera" workshop for helpful suggestions that greatly improved the keys; to the Russian Entomological Journal for permission to use Figs. 7A-N; and to G. A. Anufriev and A. F. Emeljanov for permission to use Figs. 3CE, 4J-K, and 7O-P. Figures reproduced from Kramer (1950) and Oman (1949) are in the public domain. This work was supported in part by grants DEB 9978026 and DEB 0089671 from the National Science Foundation.

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