Christopher H. Dietrich

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

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.

Molecular taxonomy using single‐strand conformation polymorphism (SSCP) analysis of mitochondrial ribosomal DNA genes

Single‐strand conformation polymorphism (SSCP) analysis detects single point mutations in DNA molecules. We demonstrate that SSCP analysis of mitochondrial ribosomal DNA (rDNA) genes is a sensitive taxonomic tool because these genes often differ at numerous sites among closely related species. Using conserved primers, portions of the 12S or 16S rDNA genes were amplified using the polymerase chain reaction (PCR) in congeneric species of ticks, leaf hoppers, mosquitoes, and closely related endoparasitic wasps. SSCP was performed and products were visualized with silver staining. Species‐specific patterns were observed in all taxa. Intraspecific variation at the level of single nucleotide substitutions was detected. SSCP diagnostics are less expensive and time consuming to develop than PCR with species‐specific primers, and, unlike PCR with arbitrary primers, there is minimal concern with DNA contamination from non‐target organisms.

Phylogeny of the leafhopper subfamily Evacanthinae with a review of Neotropical species and notes on related groups (Hemiptera: Membracoidea: Cicadellidae)

Abstract.u2003 The phylogenetic analysis of ninety‐two adult morphological characters supports the treatment of Nirvaninae as a junior synonym of Evacanthinae and the redefinition of Evacanthinae to include the tribes Nirvanini, Balbillini, Evacanthini and Pagaroniini. The analysis indicates that Nirvaninae, as previously defined, is polyphyletic. A key to tribes and Neotropical genera is provided and diagnostic features for these taxa are reviewed. Jassoqualus Kramer, Neonirvana Oman, Synogonia Melichar (=Jassopronus Nielson & Godoy, syn.n.) and Tahura Melichar are retained within Nirvanini and two new Neotropical genera of this tribe are described and illustrated: Antillonirvana, gen.n., based on two new species from the Dominican Republic and one from Cuba; and Chibchanirvana, gen.n., based on two new species from Colombia. Pentoffia Kramer and Evanirvana Hill are treated as incertae sedis within Evacanthinae. Six new species of Pentoffia, a new species of Synogonia, a new species of Jassoqualus, two new species of Neonirvana and eleven new species of Tahura, all from South America, are also described and illustrated. The following taxa included previously in Nirvaninae are excluded from Evacanthinae, sensu lato: Tungurahuala Kramer to Cicadellinae; Columbonirvana Linnavuori to Typhlocybinae; Macroceratogoniini to Coelidiinae; Occinirvanini Evans to Deltocephalinae. Omaranus Distant, placed previously in Occinirvanini, is transferred to Doraturini (Deltocephalinae).

Phylogeny of the leafhopper subfamily Deltocephalinae (Hemiptera: Cicadellidae) based on molecular and morphological data with a revised family-group classification.

Deltocephalinae, a highly diverse and economically important subfamily of leafhoppers, contains over 6200 species and 36 tribes distributed worldwide in habitats ranging from xeric grasslands and shrublands to tropical rainforests. Recent morphological and molecular phylogenetic analyses of Cicadellidae and a morphology‐based analysis of Deltocephalinae and related subfamilies indicated that several previously recognized cicadellid subfamilies are closely related to or derived from within Deltocephalinae, but these analyses did not provide a comprehensive or well‐supported hypothesis of the phylogeny of Deltocephalinae s.l. due to either low taxon sampling or low branch support. Here, taxon sampling was increased to include members of most family‐group taxa of Deltocephalinae and molecular data (∼2800 bp 28S rDNA and ∼350 bp histone H3) were added to improve the phylogenetic estimate. Five putative outgroup taxa were included, and parsimony and Bayesian analyses of the combined molecular and morphological (119 characters) data and maximum likelihood analyses of the 28S data showed strong support for the monophyly of Deltocephalinae as defined here. Branches near the base of the tree and towards the tips were longer and better supported than many of the shorter internal branches. Similar to a previous morphological phylogenetic analysis of Deltocephalinae, all of the grass‐ and sedge‐specializing tribes were recovered in one common clade, with a few apparent reversals to nongrass feeding. Although support for this clade was low and requires further testing, the results suggest that grass/sedge specialization is a phylogenetically conservative trait within Deltocephalinae. The history of the classification of Deltocephalinae and related subfamilies is reviewed, and based on the results of the phylogenetic analyses presented here, a revised family‐group taxonomic classification is proposed. In addition to subfamilies that were recently included in Deltocephalinae, the following are considered junior synonyms of Deltocephalinae: Acostemminae syn.n., Arrugadinae syn.n., Drakensbergeninae syn.n., Mukariinae syn.n. and Stegelytrinae syn.n. The morphological characters supporting this interpretation of Deltocephalinae are provided and discussed, and a description of the subfamily is provided. A new tribe, Faltalini tribe n. (11 genera, 31 species) is described, and Magnentiini placement n. and Paraphrodini placement n. are transferred to Deltocephalinae from Nioniinae and Aphrodinae, respectively. New placements of genera include: Twiningia Ball and Eusama Oman (Athysanini: Platymetopiina), placement n.; Cerrillus Oman (Athysanini), placement n.; Scaphotettix Matsumura and Agrica Strand (Mukariini), placement n.; Loralia Evans and Phlogotettix Ribaut (Deltocephalinae, unplaced to tribe), placement n. The recognition of Scaphoideini Oman 1943 as a nomen nudum results in the following placements: Acunasus DeLong, Cantura Oman, Danbara Oman, Osbornellus Ball, Prescottia Ball, Scaphodhara Viraktamath & Mohan, Scaphoideus Uhler, Scaphoidophyes Kirkaldy, Sincholata DeLong, Sobara Oman and Soleatus DeLong (Deltocephalinae, unplaced to tribe), placement n.

Treehopper trees: phylogeny of Membracidae (Hemiptera: Cicadomorpha: Membracoidea) based on molecules and morphology

Abstract.u2003 Recent independent phylogenetic analyses of membracid relationships based on molecular and morphological data have identified monophyletic lineages within the family. However, the results of these studies have not fully resolved treehopper phylogeny, and relationships among some higher membracid lineages remain in doubt. Portions of three datasets (958 aligned nucleotides from elongation factor‐1α, 2363 aligned nucleotides from 28S ribosomal DNA, and eighty‐three morphological features of adults and nymphs) introduced in recent studies were reanalysed separately and in combination with two new molecular datasets (321 aligned nucleotides from wingless and 1829 aligned nucleotides from 18S ribosomal DNA). The results of the combined data analyses, contrary to previous analyses of morphological data alone, grouped membracids into two well‐supported lineages, one comprising Stegaspidinae and Centrotinae, the other comprising Membracinae, Darninae and Smiliinae. The analyses recovered Centrotinae, Membracinae and Darninae as monophyletic groups, but Stegaspidinae was paraphyletic with respect to Centrotinae, and Smiliinae was polyphyletic with Micrutalini placed as a sister group to the clade comprising Membracinae, Darninae and Smiliinae. These results are consistent with the following hypotheses, proposed previously based on an analysis of morphological data: (1) the posterior pronotal process was derived and lost multiple times during the evolution of Membracidae; (2) Membracidae originated in the New World and reached the Old World subsequently via dispersal; (3) maternal care evolved independently multiple times and may or may not have been preceded by the acquisition of ant mutualism.

Superfamily Membracoidea (Homoptera: Auchenorrhyncha). I. Introduction and revised classification with new family-group taxa

Abstract. A key and descriptions are given for the four families here recognized for the superfamily Membracoidea Rafinesque, 1815: Cicadellidae Latreille, 1825 (sensu Oman et al., 1990), Melizoderidae, fam.n., Aetalionidae Spinola, 1850 redefined here), and Membracidae Rafinesque, 1815 (redefined here). The following placements are corroborated: Cicadelloidea Latreille, 1825, is a junior synonym of Membracoidea Rafinesque, 1815; Biturritiidae Metcalf, 1951, is a junior synonym of Aetalionidae Spinola, 1850; Nicomiidae Haupt, 1929, is a junior synonym of Membracidae Rafinesque, 1815. The new family Melizoderidae includes two genera: Melizoderes Spinola and Llanquihuea Linnavuori & DeLong. Within Aetalionidae, the subfamily Biturritiinae is redefined to include only five genera. Within the family Membracidae, the subfamily Endoiastinae, subfam.n., is described; the subfamilies Centronodinae Deitz, 1975, status n. (with tribe Centronodini Deitz, 1975, including the genus ParacentronodusSakakibara, 1971, new Placement), and Centrodontinae Deitz, 1975, status n. (with tribe Centrodontini Deitz, 1975), are recognized for the first time; and the following groups are redefined: subfamilies Stegaspidinae Haupt, 1929, Nicomiinae Haupt, 1929, and Membracinae Rafinesque, 1815; and tribe Stegaspidini Haupt, 1929 (Stegaspidinae). The new subfamily Endoiastinae includes three genera: Endoiastus Fowler, Scytodepsa Stål, and Stictodepsa Stål. Endoiastus productus Osborn, 1922, is placed as a junior synonym of Stictodepsa neotropicalis Kirkaldy, 1909, syn.n. Lectotypes are designated and illustrated for Tettigonia muscaria Fabricius, 1803 (now Lophyraspis muscaria) and Cicada fuscata Fabricius, 1803 (now Stictodepsa neotropicalis Kirkaldy, 1909).

Morphology-based phylogeny of the treehopper family Membracidae (Hemiptera: Cicadomorpha: Membracoidea)

A parsimony‐based phylogenetic analysis of eighty‐three morphological characters of adults and immatures of seventy representatives of the tribes and subfamilies of Membracidae and two outgroup taxa was conducted to evaluate the status and relationships of these taxa. Centrotinae apparently gave rise to Nessorhinini and Oxyrhachini (both formerly treated as subfamilies, now syn.n. and syn.reinst., respectively, of Centrotinae). In contrast to previous analyses, a clade comprising Nicomiinae, Centronodinae, Centrodontinae, and the unplaced genera Holdgatiella Evans, Euwalkeria Goding and Antillotolania Ramos was recovered, but relationships within this clade were not well resolved. Nodonica bispinigera, gen.n. and sp.n., is described and placed in Centrodontini based on its sister‐group relationship to a clade comprising previously described genera of this tribe. Membracinae and Heteronotinae were consistently monophyletic. Neither Darninae nor Smiliinae, as previously defined, was monophyletic on the maximally parsimonious cladograms, but constraining both as monophyletic groups required only one additional step. The monophyly of Stegaspidinae, including Deiroderes Ramos (unplaced in Membracidae), was supported on some but not all equally parsimonious cladograms. More detailed analyses of individual subfamilies, as well as morphological data on the undescribed immatures of several membracid tribes and genera, will be needed to elucidate relationships among tribes and genera. A key to the subfamilies and tribes is provided.

Superfamily Membracoidea (Homoptera: Auchenorrhyncha). II. Cladistic analysis and conclusions

Abstract. Homologies among traditional morphological characters in the Membracoidea (sensu lato) are reassessed and the phylogenetic relationships among higher membracoid taxa are explored, incorporating new morphological evidence from nymphs and adults. Weighted and unweighted parsimony analyses of a matrix of sixty–three characters and thirty‐nine OTUs representing the families Aetalionidae, Cicadellidae, Melizoderidae and Membracidae, and an outgroup (superfamily Cercopoidea) yielded various topologies that were largely congruent but presented alternative hypotheses of relationships among the Membracidae. These analyses indicate that the superfamily consists of the following clades: Cicadellidae + (Melizoderidae + (Aetalionidae + Membracidae)). The family Membracidae, traditionally characterized by the presence of a posterior pronotal process, apparently gave rise to Nicomia Stål and other genera that lack this process.

Auchenorrhyncha: (Cicadas, Spittlebugs, Leafhoppers, Treehoppers, and Planthoppers)

Publisher Summary This chapter discusses hemipteran suborder Auchenorrhyncha its life history, behavior, and ecology, which is the group of sapsucking insects comprising the modern superfamilies, Cercopoidea, Cicadoidea, Membracoidea, and Fulgoroidea. Together, these groups include more than 40,000 described species. Morphologically, Auchenorrhyncha differ from other Hemiptera in having the antennal flagellum hair-like, the rostrum arising from the posteroventral surface of the head, a complex sound-producing tymbal apparatus, and the wing-coupling apparatus consisting of a long, downturned fold on the forewing and a short, upturned lobe on the hind wing. Adult male and female Auchenorrhyncha locate each other by means of species-specific acoustic courtship signals. These signals are produced by specialized organs at the base of the abdomen called tymbals, present in both sexes. A few cicadas and planthoppers are also able to use the stridulatory surfaces of their wings to produce sound. The loud, sometimes deafening, calls of many male cicadas are well known. In noncicadoids, the courtship calls are usually inaudible, being transmitted through the substrate, and distinct tympana are absent. The calls of some leafhoppers and planthoppers, audible only with special amplifying equipment, are among the most complex and beautiful of any produced by insects.

Systematics and phylogeny of the Neotropical treehopper subfamily Nicomiinae (Hemiptera, Membracidae)

Morphological characters of adults of the treehopper subfamily Nicomiinae Haupt, 1929 (Hemiptera, Membracidae) including seven genera (Eudonica gen. nov.; Euwalkeria Goding, 1926; Holdgatiella Evans, 1962; Nicomia Stal, 1858; Nodonica Dietrich, McKamey& Deitz, 2001; Stalomia gen. nov.; and Tolania Stal, 1858) and 22 species (16 new) are described and illustrated. Keys are provided for genera and for species of Euwalkeria, Holdgatiella, and Nicomia. Nomenclatural changes, based on study of the primary type material of 15 species, include three new combinations, one new synonymy, and reinstatement of one junior synonym. Eudonica has one species, Eudonica nanella sp. nov.; Euwalkeria has five species, including four new species: E. colorata sp. nov., E. distincta sp. nov., E. perdita sp. nov., E. rubrica sp. nov.; Holdgatiella has two species, one of which is described as new: Holdgatiella chiloensis sp. nov.; Nicomia has twelve species, nine of which are described as new: N. buccina sp. nov., N. harenosa sp. nov., N. inscripta sp. nov., N. jucunda sp. nov., N. monticola sp. nov., N. nigrifasciata sp. nov., N. notidana sp. nov., N. pulchella sp. nov., N. serrata sp. nov.; Nodonica has one species, Nodonica bispinigera Dietrich, McKamey & Deitz; and Stalomia has one species, Stalomia veruta sp. nov. Tolania contains eleven previously described species and nearly 60 new species, which will be treated in a later publication. Three new combinations are proposed: one species described in Nicomia is placed in the tribe Abelini (Centrotinae) as Abelus retrospinosus (Lethierry) comb. nov.; one species previously placed in Nicomia is transferred to the genus Tolania as T. obliqua (Walker, 1858), comb. nov.; one species described in Holdgatiella is placed in the genus Tolania as T. stria (Cryan & Deitz, 2002), comb. nov. One new synonymy is proposed: Hoplophera [sic] cicadoides Walker, 1862, syn. nov., a junior synonym of Nicomia interrupta Stal, 1858. Nicomia subfasciata Stal, 1858, is reinstated. The results of a phylogenetic analysis based on morphology are presented, illustrating the hypothesized relationships among species and genera of Nicomiinae. The analysis supports the broader concept of Nicomiinae proposed here. The monophyly of this group is supported by eleven characters, and all of the included genera are also monophyletic.