Lewis L. Deitz

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

Abstract.  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).

Generic revision of the New World tribe Hoplophorionini (Hemiptera: Membracidae: Membracinae)

Abstract. The tribe Hoplophorionini Goding, 1926, includes 105 species of treehoppers in 10 genera. All are apparently subsocial and lack mutualism with honeydew‐collecting hymenopterans. In many species, parental investment in offspring is unique because of a specialized kicking defence (described herein) and construction of extra‐ovipositional punctures in the host tissue through which nymphs feed. The tribe occurs from Canada to Chile, with most generic diversity in Central America but most species diversity near the equator. Three genera, Stalotypa Metcalf, Ramosella, new genus and Stirpis, new genus, are endemic to the Greater Antilles but do not constitute a monophyletic group; the first two genera appear most closely related to Turrialbia, new genus, from Costa Rica. Host plant specializations and other biological attributes are summarized for genera and species. A phylogenetic analysis of 23 hoplophorionine species produced 9 minimal‐length cladograms that were similar in many respects. Potnia Stål appears to have retained the greatest number of ancestral features. Aposematic teneral coloration of adults probably evolved once. The modification of the ancestral kicking behaviour by the first hoplophorionines appears to have released them from a constraint on pronotal form.

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.

Distribution of Karyotypes of the Cryptocercus punctulatus Species Complex (Dictyoptera: Cryptocercidae) in the Southern Appalachians: Relation to Habitat and History

Abstract The distributional pattern of the four known karyotypes (male 2n = 37, 39, 43, 45) of the Cryptocercus punctulatus Scudder species complex is reported, based on 71 sites in the Southern Appalachian Mountains with an emphasis on western North Carolina. Populations with different karyotypes are geographically structured in a mosaic, with at least one karyotype occurring in two disjunct regions. Abrupt geographic transitions between karyotypes suggest a parapatric distribution. We found no overlap in the distribution of the different karyotypes, as recently suggested. Although the boundary zones between karyotypes do not appear to coincide with physical or ecological barriers to dispersal, several transitions between karyotypes occur on or near the highest mountains in the southern Appalachians. We suggest that the different karyotypes arose by vicariance, with current boundaries formed by secondary contact when populations isolated in glacial refugia subsequently spread into high-mountain habitats. Because of their dependence on mature mesic forests, populations of the cockroach likely advance up and down mountainsides in cycles of advances and retreats dictated by climatic oscillations that raise and lower the timberline. We discuss the taxonomic status of the different karyotype groups in the C. punctulatus complex, and conclude that more exacting evidence is required to establish if species-level status is warranted. The conclusions of certain earlier studies are weak because, among other things, karyology was not examined in the sampled specimens, including those designated as types.

On Dorsal Prothoracic Appendages in Treehoppers (Hemiptera: Membracidae) and the Nature of Morphological Evidence

A spectacular hypothesis was published recently, which suggested that the “helmet” (a dorsal thoracic sclerite that obscures most of the body) of treehoppers (Insecta: Hemiptera: Membracidae) is connected to the 1st thoracic segment (T1; prothorax) via a jointed articulation and therefore was a true appendage. Furthermore, the “helmet” was interpreted to share multiple characteristics with wings, which in extant pterygote insects are present only on the 2nd (T2) and 3rd (T3) thoracic segments. In this context, the “helmet” could be considered an evolutionary novelty. Although multiple lines of morphological evidence putatively supported the “helmet”-wing homology, the relationship of the “helmet” to other thoracic sclerites and muscles remained unclear. Our observations of exemplar thoraces of 10 hemipteran families reveal multiple misinterpretations relevant to the “helmet”-wing homology hypothesis as originally conceived: 1) the “helmet” actually represents T1 (excluding the fore legs); 2) the “T1 tergum” is actually the anterior dorsal area of T2; 3) the putative articulation between the “helmet” and T1 is actually the articulation between T1 and T2. We conclude that there is no dorsal, articulated appendage on the membracid T1. Although the posterior, flattened, cuticular evagination (PFE) of the membracid T1 does share structural and genetic attributes with wings, the PFE is actually widely distributed across Hemiptera. Hence, the presence of this structure in Membracidae is not an evolutionary novelty for this clade. We discuss this new interpretation of the membracid T1 and the challenges of interpreting and representing morphological data more broadly. We acknowledge that the lack of data standards for morphology is a contributing factor to misinterpreted results and offer an example for how one can reduce ambiguity in morphology by referencing anatomical concepts in published ontologies.

New records of plant pests in New Zealand. III. Six species of Homoptera.

Four homopterous insect pests, Philaenus spumarius (Linnaeus), Acyrtho-siphon pisum (Harris), Pulvinaria hydrangeae Steinweden, and Kuwanaspis pseudoleucaspis (Kuwana) are recorded from New Zealand for the first time. The presence of Aulacaspis rosarum Borchsenius, Asiatic rose scale, and Pinnaspis aspidistrae (Signoret), fern scale, is clarified.

A review of entomophagous insects in the Cook Islands

Available information concerning entomophagous insects introduced to the Cook Islands is summarized. Host-parasite associations determined through rearing field-collected insects from Rarotonga and Aitutaki, October 1975, are given; other parasites and predators known to occur in the Cook Islands are listed.

TREEHOPPER DIVERSITY (HEMIPTERA: MEMBRACIDAE) OF LITTLE ORLEANS, ALLEGANY CO., MARYLAND

Abstract Fifty-six species (23 genera) of treehoppers (Membracidae) are recorded from the vicinity of Little Orleans, Maryland, with date and host-plant data. Eleven of the species represent new state records for Maryland, bringing the total number of species recorded for the state to 81. These data document annual collecting efforts since 1993 by the authors and others in connection with the informal treehopper conferences initiated by the late T. K. Wood. Photographs are given for 12 species representing 7 genera.