Replace “*” with male character and “$” with female character throughout the text.
Review of the species of New World Erythroneurini (Hemiptera: Cicadellidae: Typhlocybinae).
IV. Genus Eratoneura.
Dmitry A. Dmitriev and Christopher H. Dietrich
CONTENTS
Plate 1 (Photos of Eratoneura Species)
Abstract
Introduction
Materials and Methods
Taxonomy
Genus Eratoneura Young, 1952
Key to Adult Males of Eratoneura
Eratoneura dira
Eratoneura gillettei
Eratoneura imbricariae
Eratoneura lamucata
Eratoneura maculata
Eratoneura eversi
Eratoneura noncuspidis
Eratoneura teshi
Eratoneura osborni
Eratoneura sandersoni
Eratoneura crinita
Eratoneura andersoni
Eratoneura paraesculi
Eratoneura tammina
Eratoneura texana
Eratoneura separata
Eratoneura curvata
Eratoneura restricta
Eratoneura impar
Eratoneura carmini
Eratoneura forfex
Eratoneura linea
Eratoneura rangifer
Eratoneura dimidiata
Eratoneura sanctaerosae
Eratoneura betulae
Eratoneura concisa
Eratoneura spala
Eratoneura harpola
Eratoneura minor
Eratoneura aesculi
Eratoneura bifida
Eratoneura aculeata
Eratoneura lusoria
Eratoneura pyra
Eratoneura millsi
Eratoneura longifurca
Eratoneura manus
Eratoneura staffordi
Eratoneura ardens
Eratoneura uvaldeana
Eratoneura inepta
Eratoneura firma
Eratoneura bispinosa
Eratoneura hyalina
Eratoneura harnedi
Eratoneura unica
Eratoneura mira
Eratoneura facota
Eratoneura cristata
Eratoneura mcateei
Eratoneura ungulata
Eratoneura distincta
Eratoneura spinea
Eratoneura rotunda
Eratoneura nigriventer
Eratoneura stupkaorum
Eratoneura usitata
Eratoneura incondita
Eratoneura ingrata
Eratoneura arenosa
Eratoneura uncinata
Eratoneura arpegia
Eratoneura propria
Eratoneura brevipes
Eratoneura turgida
Eratoneura delongi
Eratoneura immota
Eratoneura tantilla
Eratoneura sorota
Eratoneura solita
Eratoneura knullae
Eratoneura omani
Eratoneura mimica
Eratoneura sancta
Eratoneura inksana
Eratoneura alloplana
Eratoneura parva
Eratoneura mirifica
Eratoneura abjecta
Eratoneura hartii
Eratoneura misera
Eratoneura nevadensis
Eratoneura lata
Eratoneura levecki
Eratoneura tantula
Eratoneura beeri
Eratoneura accita
Eratoneura cera
Eratoneura rubraza
Eratoneura affinis
Eratoneura stephensoni
Eratoneura clavipes
Eratoneura retusa
Eratoneura hymac
Eratoneura externa
Eratoneura brooki
Eratoneura longa
Eratoneura sebringensis
Eratoneura parvipes
Eratoneura calamitosa
Eratoneura richardsi
Eratoneura marilandicae
Eratoneura opulenta
Eratoneura era
Eratoneura fergersoni
Eratoneura ellisi
Eratoneura acantha
Eratoneura interna
Eratoneura robusta
Eratoneura metopia
Eratoneura patris
Eratoneura econa
Eratoneura alicia
Eratoneura zioni
Eratoneura greeni
Eratoneura lenta
Eratoneura gilesi
Eratoneura maga
Eratoneura igella
Eratoneura stannardi
Eratoneura trautmanae
Eratoneura knighti
Eratoneura triangulata
Eratoneura corylorubra
Eratoneura fulleri
Eratoneura rubranotata
Eratoneura citrosa
Eratoneura clara
Eratoneura nimia
Eratoneura flexibilis
Eratoneura certa
Eratoneura direpta
Eratoneura tersa
Eratoneura luculenta
Eratoneura fausta
Eratoneura bella
Eratoneura coxi
Eratoneura torella
Eratoneura tumida
Eratoneura glicilla
Eratoneura morgani
Eratoneura arta
Eratoneura hymettana
Eratoneura lawsoni
Eratoneura continua
Eratoneura valida
Eratoneura pamelae
Eratoneura protuma
Eratoneura havana
Eratoneura gemoides
Eratoneura gemina
Eratoneura malaca
Eratoneura penesica
Eratoneura parallela
Eratoneura severini
Eratoneura claroides
Eratoneura lunata
Eratoneura adunca
Eratoneura curta
Eratoneura ballista
Eratoneura rostrata
Eratoneura penerostrata
Eratoneura emquu
Eratoneura trivittata
Eratoneura anseri
Eratoneura stoveri
Eratoneura campora
Eratoneura spinifera
Eratoneura comoides
Eratoneura confirmata
Eratoneura phellos
Eratoneura bigemina
Eratoneura mensa
Eratoneura basilaris
Eratoneura micheneri
Eratoneura lucyae
Eratoneura guicei
Eratoneura lundi
Eratoneura marra
Eratoneura staminea
Eratoneura unca
Eratoneura dumosa
Eratoneura ligata
Eratoneura contracta
Eratoneura vittata
Eratoneura biramosa
Eratoneura smithi
Eratoneura prolixa
Eratoneura teres
Eratoneura amethica
Eratoneura tenuitas
Eratoneura haysensis
Eratoneura geronimoi
Eratoneura macra
Eratoneura socia
Eratoneura accola
Acknowledgements
References
Appendix I: Collections and Studied Material
Appendix II: Host Plant Index
Appendix III: Species Index
Abstract
The 197 species (including 2 new species) of the Nearctic leafhopper genus Eratoneura are reviewed. Eratoneura betulae sp.n. from New Brunswick (Canada) and E. mcateei sp.n. from the central and northeastern USA are described as new. In addition, 123 new synonyms are recognized and a lectotype is designated for Typhlocyba hartii Gillette. Key is provided for identification of males. All known species are illustrated, and data on their distributions and host plants are summarized. All species of Eratoneura are native to temperate North America, where they feed and oviposit mainly on woody deciduous trees and shrubs.
Keywords: Auchenorrhyncha, Homoptera, leafhopper, morphology, taxonomy, distribution, identification.
Introduction
This revision of the genus Eratoneura completes the series of publications on the New World Erythroneurini (Dietrich & Dmitriev, 2006a, 2007a, 2008a; Dmitriev & Dietrich, 2007a, 2009a). Eratoneura was described by Young (1952b) as a subgenus of Erythroneura Fitch to comprise the Erythroneura maculata species group designated by Robinson (1926a) and comprehensively revised by Beamer in series of publications (1931a, b, c, d, 1932a, c, d, e, f, g, h). In these papers Beamer provided illustrations of the male genitalia for all known species, incorporated these characters into a dichotomous key, and associated males with most of the species and varieties (=subspecies, see ICZN, 1999, Art. 45.6.4) that had been described by McAtee (1920a, 1924c, 1924d, 1926c) and previous authors based on females. Johnson (1935a) independently revised the species of Erythroneura sensu lato (including Eratoneura) from Ohio and provided a key for their identification. She described many new species in this and subsequent papers (Knull, 1944b, 1945b, 1949a, 1951a, 1954b, 1955a; Knull & Auten, 1937a, 1938b). After these revisions, H.H. Ross (with D.M. DeLong) and L.W. Hepner described more than 150 additional species of Eratoneura. Dietrich and Dmitriev (2006a) revised the generic classification of the New World Erythroneurini and elevated Eratoneura to the genus level.
All species of Eratoneura are native to temperate North America, where they feed and oviposit mainly on woody deciduous trees and shrubs. Ross (1957c, 1958c) did an extensive study of coexisting Eratoneura species on sycamore in southern Illinois. Adults overwinter in leaf litter, and in the spring, they emerge and feed on the new leaves of early-emerging deciduous plants before migrating to their “definitive” summer host plants where they mate and lay eggs (Ross & DeLong 1953a). Most species oviposit and undergo nymphal development on a single, or a group of closely related, plant species. In the southern USA, they complete two or more generations per year, but in the north there may be but a single generation. In the fall, adults may again feed on a variety host plants prior to seeking out winter shelter in leaf litter. Most species of Eratoneura have no known economic importance, but a few are recorded as pests of apple (Beamer, 1930a, 1932b; Hamilton, 1985a).
Material and Methods
Few previous workers have specifically targeted Eratoneura in their collecting, and specimens from trap catches and other general collecting that find their way into curated collections are often in very poor condition. Because of this, and due to time and budget constraints, this study focused only on collections known to contain large numbers of well-curated specimens of Eratoneura, as well as those housing primary types: Illinois Natural History Survey (INHS), Ohio State University (OSU), University of Kansas Natural History Museum (KSEM), Mississippi State University, Mississippi Entomological Museum (MEM), Canadian National Collection of Insects, Arachnids and Nematodes (CNC), Smithsonian National Museum of Natural History (USNM), California Academy of Sciences (CAS), and Colorado State University (CSUC). The numbers of studied specimens from each collection are summarized in Appendix 1. Future collecting will undoubtedly show that the distributions of most species are much broader than indicated on the maps accompanying individual species treatments. Although these maps show regional biases reflecting the locations and holdings of the studied collections, they are based on vouchered collection records and, thus, accurately reflect current knowledge of species distributions. On the maps the type locality is marked with a star.
Identification of species was mainly based on type material. In some cases, when the type was not located, or the holotype is a female (e.g., most of McAtee’s species), we followed Beamer’s interpretation, based on study of dissected male specimens that he labeled “allotype” to indicate that they had been compared to the female primary types of previous workers. Although Beamer’s “allotypes” have no official standing in nomenclature, these dissected male specimens facilitate unambiguous interpretation of Beamer’s concept of the species.
Morphological terminology follows Dietrich and Dmitriev (2006a, see also Fig. 1). Although individual species have a characteristic color pattern, details and intensity may be highly variable both inter- and intraspecifically. Overwintering individuals tend to be more brightly colored than adults of the summer generation of the same species. This has resulted in many species being described multiple times based on different color forms. In the descriptions below, the pattern of fully colored individuals is described, although completely or almost completely discolored forms are known for most species. Thus, identification keys are based mainly on male genitalia, with external characters used only for supplemental purposes.
This work recognizes numerous synonyms treated as valid species by previous workers. Most junior synonyms represent color variants, or forms differing slightly in the shape of the aedeagus, or the shape and length of the pygofer appendages. A few species were described based on aberrant forms with distorted genitalia, possibly caused by parasitism. In some cases where genitalia were mounted on slides, species were described based on contaminant particles embedded in the balsam.
Each species is illustrated by one or more habitus photos taken using a Microptics digital imaging system. Original drawings were prepared only in cases where those available from other sources were deemed inaccurate. Thus, numerous figures are reproduced from other sources, as noted in the figure captions. Inconsistencies (e.g., in line thickness) among line drawings reflect differences in the drawing styles of previous authors. In all cases, figures reproduced from previous publications are either in the public domain or are reproduced with permission.
Line drawings of the male genitalia accompanying each species treatment are labeled as follows:
a – pygofer or pygofer appendage, lateral view;
b – pygofer or pygofer appendage, dorsal view;
c – style or style apex, broad aspect;
d – aedeagus, lateral view;
e – aedeagus, posteroventral or ventral view.
Nomenclatural, distributional, morphological and host-plant data summarized below were extracted from a relational specimen-level database of Erythroneurini (Dmitriev & Dietrich, 2003 onwards), developed using the 3I software package (Dmitriev, 2006a, Dmitriev & Dietrich, 2008a). The on-line database provides more detailed information for each species, including a complete list of specimens examined, photos of type specimens, and interactive key to species.
In the species treatments below, only summer host plants are listed, although most species have also been collected from plants other than their oviposition hosts.
Specimens of newly described taxa (E. betulae sp.n. and E. mcateei sp.n.) are deposited in the insect collections of the Illinois Natural History Survey (Champaign), Canadian National Collection of Insects, Arachnids and Nematodes (Ontario), and Mississippi State University (State College).
Taxonomy
Subfamily Typhlocybinae
Tribe Erythroneurini Young, 1952
Genus Eratoneura Young, 1952
Erythroneura maculata species group Robinson, 1926a:109
Erythroneura (Eratoneura) Young, 1952b:84 (Type: Erythroneura dira Beamer, 1931)
Eratoneura Dietrich & Dmitriev, 2006a:133
Diagnosis: Ground color usually pale, often with red or orange maculae distributed on dorsum and black or dark brown spots near costal margin and in inner apical cell; crown without pair of brown or black spots. Male pygofer without ventral appendage or sclerotized ridge; dorsal appendage immovably fused to margin, simple or bifurcated, with additional small projection at base; basolateral setae well developed. Style apex usually with three angulate projections.
Description: Length 2.3–3.7 mm, slender. Head narrower than pronotum. Crown fore margin produced, angulate or rounded. Ocelli absent or vestigial. Face depressed in profile, less than 45° from horizontal. Pronotum smooth, without conspicuous pits. Forewing outer apical cell about 2X as long as wide or longer; second apical cell basally truncate (ir crossvein present); third apical cell parallel sided, straight; CuP vein longer than segment of CuA between Cu and MP; inner apical cell with oblique base, basal segment of CuA and CuP veins forming continuous line; Pcu vein not visible. Hind wing apex truncate; submarginal vein not extended to wing apex; ScR vein present; MP and CuA veins touching at one point, fused for short distance or separated by m-cu crossvein. Front femur anteroventral row with basal seta distinctly larger than others. 2S abdominal apodemes variable, small, narrow and short to large, broad, and extended beyond posterior margin of sternite III (3S). Pygofer not extended to apex of subgenital plate; lobe rounded or angulate; dorsal emargination extended to base of segment; dorsal membrane without fine setae; oblique dorsolateral internal ridge not developed; basolateral setae in distinct group, small; distal and dorsal setae undifferentiated; internal surface of lobe with setae; sparse long fine setae present; microtrichia well developed. Pygofer dorsal appendage immovably fused to margin, without basal suture; appendage simple or bifurcated, with additional small projection at base; ventral appendage or sclerotized ridge absent. Sternite IX with median longitudinal internal ridge. Subgenital plates free; lateral margin with angulate subbasal projection; section basad of medial constriction shorter than distal section; with 4 basal macrochaetae uniseriate along margin; distinct marginal subbasal rigid setae forming continuous row; distal macrochaetae absent. Style free; preapical lobe prominent; apex smooth with 3 points or rarely truncated. Aedeagus articulated to connective; with dorsal apodeme broadly expanded in lateral view; apodeme parallel sided in ventral view, connection to pygofer membranous. Aedeagus shaft symmetrical, usually denticulate distally, sometimes with basal or distal processes. Connective U- or V-shaped, without median anterior lobe, with long arms. Anal tube without processes. Coloration variable, but all known species with crown lacking pair of fuscous preapical spots. Usual color pattern (see E. dira Beamer, Plate 1a, 2a, for example) consisting of pale or dull yellow dorsum with red or orange maculae distributed on crown thorax and forewings and black or dark brown spots near costal margin and near base of inner apical cell.
Distribution: Temperate North America.
Host plants: Deciduous trees, shrubs.
Key to adult males of Eratoneura1
1. Aedeagus with pair of slender distal processes (Fig. 2e, 3e). 2
1'. Aedeagus without slender distal processes (Fig. 1e, 9e). 8
2(1). Aedeagus with pair of processes arising at base or near midlength of shaft (Fig. 2d, e). 3
2'. Aedeagus without processes arising at base or near midlength of shaft (Fig. 5d, e). 4
3(2). Aedeagal shaft curved dorsad, processes arising near midlength of shaft (Fig. 2d, e). Dorsal pygofer appendage extended to pygofer apex (Fig. 2a, b). 2. E. gillettei (Beamer)
3'. Aedeagal shaft curved ventrad, processes arising at base of shaft (Fig. 3d, e). Dorsal pygofer appendage not extended to pygofer apex (Fig. 3a, b). 3. E. imbricariae (Ross & DeLong)
4(2). Tip of aedeagal shaft with short dorsal projection; distal processes extended ventrolaterad (Fig. 4d, 5d). 5
4'. Tip of aedeagal shaft without dorsal projection; distal processes extended dorsolaterad (Fig. 6d). 6
5(4). Distal processes of aedeagus about as long as shaft (Fig. 4e). 4. E. lamucata (Ross & DeLong)
5'. Distal processes of aedeagus about half as long as shaft (Fig. 5e). 5. E. maculata (Gillette)
6(4). Third point of style apex longer than half distance between other two points (Fig. 6c). Distal processes of aedeagus arising subapically, shorter then half length of shaft (Fig. 6e).
6. E. eversi (Ross & DeLong)
6'. Third point of style apex shorter than half distance between other two points (Fig. 7c, 8c). Distal processes of aedeagus arising apically, almost as long as shaft (Fig. 7e). 7
7(6). Third point of style apex short toothlike, subequal in size to second point (Fig. 7c). Aedeagal shaft with prominent lateral lobes, smooth (Fig. 7d, e). Pygofer appendage compressed (Fig. 7a, b).
7. E. noncuspidis (Beamer)
7'. Third point of style apex almost as long as half distance between other two points (Fig. 8c). Aedeagal shaft without lateral lobes, with teeth (Fig. 8d, e). Pygofer appendage not compressed (Fig. 8a, b). 8. E. teshi (Hepner)
8(1). Aedeagus with pair of processes arising at base of shaft (Fig. 9d, e, 10d, e). 9
8'. Aedeagus without processes arising at base of shaft (Fig. 17d, e). 15
9(8). Pygofer appendage not extended to pygofer apex, bulbous subapically (Fig. 9a, b). Forewing basal half uniformly red (Plate 2i). 9. E. osborni (DeLong)
9'. Pygofer appendage extended to or beyond pygofer apex, not bulbous subapically (Fig. 10a, b). Forewing basal half yellow with reddish maculae (Plate 2a). 10
10(9). Third point of style apex longer than distance between other two points (Fig. 10c). Pygofer appendage curved upward, widest at midlength (Fig. 10a). Basal processes of aedeagus small toothlike (Fig. 10d, e). 10. E. sandersoni (Ross)
10'. Third point of style apex shorter than distance between other two points (Fig. 11c). Pygofer appendage straight or curved downward, widest at base (Fig. 11a). Basal processes of aedeagus longer (Fig. 11d, e). 11
11(10). Pygofer appendage with ring of long spines near midlength (Fig. 11a, b). Aedeagal shaft strongly compressed, with dorsal carina or distal lobe; basal processes appressed to shaft (Fig. 11d, e). 12
11'. Pygofer appendage without ring of long spines (Fig. 14a, b). Aedeagal shaft without dorsal carina or distal lobe; basal processes divergent from shaft (Fig. 14d, e). 13
12(11). Aedeagal shaft broad in lateral view, only about twice as long as wide; basal processes extended along dorsal margin of shaft (Fig. 11d). 11. |