Corresponding author: C. H. Dietrich (
Academic editor: Mick Webb
A new cicadellid tribe,
The leafhopper subfamily
Phylogenetic analyses of both morphological and molecular data (
In a recent taxonomic review and morphology-based phylogenetic analysis of
Morphological terminology follows
Specimens examined are deposited in the following institutions: Humboldt Institute, Villa Leyva, Colombia (HIC); Illinois Natural History Survey (INHS); North
Head with lorum extended little or no farther dorsad than clypeal suture; gena partially or entirely concealing triangular proepisternum; anteclypeus strongly convex, tapered distally; frontoclypeus without median longitudinal carina; ocelli on crown anteromesad of eyes, well separated from anterior margin; crown glabrous or punctate, without oblique lateral submarginal carinae; antennal base near anterodorsal corner of eye. Forewing (
The hind wing venation is very similar among the tribes here included in
1 | Head with margin of crown sharply carinate and encroaching onto eye laterally ( |
|
– | Head with margin of crown at most weakly carinate, not encroaching onto eye laterally ( |
2 |
2 | Body strongly depressed ( |
|
– | Body not depressed ( |
3 |
3 | Antennae longer than body; forewing crossvein r-m1 present, appendix absent ( |
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– | Antennae shorter than body; forewing crossvein r-m1 absent, appendix well developed ( |
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Medium-sized leafhoppers (~6–8 mm), body depressed, head produced, face horizontal in profile, antenna shorter than width of head, frontoclypeus with transverse carina forming distinct shelf in lateral view (
A previous cladistic analysis (
Species of
1 | Head and pronotum uniformly black dorsally; crown margin pentagonal in dorsal view | |
– | Head and pronotum pale orange, white or green with distinct reddish orange markings; crown margin parabolic in dorsal view |
Elongate, strongly depressed, leafhoppers (
1 | Anal tube without distinct apical ventrolateral spines; aedeagal shaft ( |
|
– | Anal tube with pair of distinct apical ventrolateral spines ( |
Length male 7.6–7.9. Head (male) approximately ¼ length of forewing. Forewing dark brown to black. Male anal tube with apical ventrolateral spines very weakly developed or absent. Aedeagus with gonopore-bearing shaft in lateral view broad, apex obliquely truncate, posteroapical margin concave; ventral process with pair of microtrichiate dorsolateral lobes, in ventral view abruptly expanded preapically, apex broadly bilobed, in lateral view with apex convergent toward shaft and with dorsal margin entire.
Holotype male: ECUADOR: Mt. Tungurahua, Baños, 2500m, August 20, 1937 (W. Clarke-Macintyre) [USNM]. Other material: 1 male, COLOMBIA, Cundinamarca, PNN Chingaza Charrascales,
The specimens examined from Colombia are here considered conspecific with the holotype from Ecuador, although there is slight variation among specimens in size, coloration, and the shape of the aedeagus. Given the small number of specimens available, it seems prudent to consider these minor variations to be intra-specific, despite the considerable geographic disjunction among the known populations.
Although
Length male 7.5–7.6 mm, female 7.6 mm. Head (male) approximately 1/3 as long as forewing (female with head proportionately longer compared to forewing). Forewing dark brown with tan pigment along costal area, mostly tan in female. Male anal tube with apical ventrolateral spine well developed. Aedeagus with gonopore-bearing shaft slender, apex rounded; ventral process acuminate, without dorsolateral lobes, apex in ventral view narrower than shaft.
Holotype male: COLOMBIA, Boyacá, SFF Iguaque Lagunillas,
The name refers to the acuminate ventral process of the aedeagus.
This species resembles
Medium sized, depressed leafhoppers (
The name
This genus is closely related to
The genus is described based on six species from the Andean region of South America. The species inhabit cloud forests and have been collected by sweeping grasses and other herbaceous vegetation in the understory. They are readily distinguished by differences in coloration, head proportions, and the structure of the male genitalia.
1 | Aedeagal shaft ( |
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– | Aedeagal shaft strongly asymmetrical, elongate, slender and tubular | 2 |
2 | Aedeagus ( |
|
– | Aedeagus without basal process, shaft with one or more distal processes | 3 |
3 | Aedeagus ( |
4 |
– | Aedeagus with distal process not extended laterad at right angle, either strongly recurved ventrad or more or less continuing in line with shaft | 5 |
4 | Distal process of aedeagus bifid ( |
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– | Distal process of aedeagus unbranched ( |
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5 | Aedeagus ( |
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– | Aedeagus ( |
Length male 6.6–6.9 mm, female 7.0 mm. Crown pale orange-yellow, orange-red maculae broad, overlapping ocelli, anterior margin forming acute angle; pronotum and opaque areas of forewing bright green (mottled with yellow in specimens removed from ethanol), pronotum with semicircular macula distinct. Male pygofer processes extended mesad and curved dorsad but not or only slightly crossing midline; anal tube process short, triangular. Aedeagus asymmetrical, shaft tubular, in lateral view V-shaped basally, arched and sinuate distally, terminating in two slender, bladelike processes continuing in line with shaft but asymmetrically curved, one process with angulate projection near base. Female seventh sternite with posterior margin slightly produced, rounded medially. Fifth instar nymph pale olive green dorsally with broad white median longitidinal stripe extended entire length of body; venter white.
Holotype male: PERU, Pasco, Yanachaga-Chemillén N.P., Refugio El Cedro, 2420 m,
The species name refers to the bifid apex of the aedeagus.
This species may be distinguished by its relatively long crown and the bifid apex of the aedeagus.
Length male 6.1 mm. Coloration as described for
Holotype male, PERU, Chanchamayo, 25 July 1960 (Young and Ramirez) [NCSU].
The species name means “crazy" and refers to the bizarre, asymmetrical aedeagus.
This species may be distinguished by its relatively short crown and by the presence of a long, unpaired ventral process arising from the base of the aedeagal shaft.
Length male 5.9–6.2 mm; female 7.1–7.5 mm. Coloration as described for
Holotype male: PERU, Chanchamayo, 25 July 1960 (Salazar and Ramirez) [NCSU]. Paratypes: 1 male, same locality, 22 July 1960 (C. Ramirez) [INHS]; 3 females, same locality except 21 and 25 July 1960 [NCSU, INHS]. Other material: 3 males, PERU, Ucayali, 8 km E Abra La Divisoria, 1250 m,
The species name refers to the long distal spine of the aedeagus.
This species may be distinguished by its moderately long crown and by the elongate, laterally directed distal spine of the aedeagus.
One specimen from Yanachaga-Chemillén National Park, Peru, has the distal spine of the aedeagus extended to the right, mirroring the condition found in other examined specimens of this species. Specimens examined from Yanachaga-Chemillén National Park have the anal tube processes considerably shorter than those in the type series from Chanchamayo, but such variation is here considered to be intraspecific. Based on the material available for study, this is the most widespread and common species of the genus.
Length male 6.5–6.8 mm. Nearly uniform pale orange dorsally, orange-red maculae slender; apical margin of crown forming acute angle, pronotal macula indistinct. Male pygofer processes robust, not crossing posteromedially; anal tube without ventrolateral processes; aedeagal shaft symmetrical, pillarlike, compressed, in lateral view extended dorsad and bent posterad at right angle, with pair of acute anterodorsal processes extended anterad, posterodorsal extension with ventral margin irregular, apex expanded with ventral spine. Female unknown.
Holotype male: COLOMBIA, Cundinamarca, PNN Chingaza Valle Del Fraylejon,
The species name refers to the mostly orange coloration of the dorsum.
This species may be distinguished by its relatively long crown, predominantly orange coloration, and broad, strongly compressed aedeagal shaft.
Length male 5.9 mm. External morphology and male terminalia similar to those of
Holotype male: PERU, Huánuco, Carpish Pass, 2600 m,
The species name refers to the strongly recurved apex of the aedeagus.
This species may be distinguished by its relatively short crown and strongly recurved aedeagal apex.
Length male 5.8–6.0, female 7.0 mm. Crown pale yellow medially, white laterally, orange/red maculae broad, overlapping ocelli; anterior margin forming approximately right angle; pronotum and opaque areas of forewing dark green (mottled with yellow in specimens removed from ethanol), pronotum with semicircular macula distinct. Male pygofer processes slender, crossing posteromedially; anal tube with pair of retrorse posterolateral spines. Aedeagus asymmetrical, shaft narrow, tubular and gradually tapered distally, in lateral view narrowly U-shaped with distal part attenuated and arcuate; apex with slender bifurcate process extended to left at right angle to shaft. Female seventh sternite posterior margin with small acute median tooth.
Holotype male: PERU, Pasco, Yanachaga-Chemillén N.P.,
The species name refers to the mostly pale green coloration of the dorsum.
This species closely resembles
In one examined male specimen, the configuration of the aedeagus is the mirror image of that of the other specimens.
I am indebted to Paul Freytag and Michael Sharkey (University of Kentucky) for access to their Malaise trap samples from Colombia, to Robert Blinn (NCSU) for the loan of additional specimens, to Carlos Peña and Pedro Lozada for help with field work in Peru, to Daniela Takiya and Dmitry Dmitriev for helpful discussions, and to Mick Webb and Chandra Viraktamath for constructive criticism of the manuscript. This work was supported in part by grants from the U.S. National Science Foundation.