The leafhopper subfamily Typhlocybinae comprises ~6,000 described species of mostly small, delicate leafhoppers that feed preferentially on the contents of leaf parenchyma cells of their host plants. Based on described species, this is currently the sec-ond largest leafhopper subfamily (after Deltocephalinae, ZAHNISER & DIETRICH 2010), but recent sampling of tropical faunas indi-cates that the extant, but mostly undescribed, fauna of Typhlocybinae is far larger than that of any other leafhopper subfamily (DIETRICH & WALLNER 2002). The Neotropical fauna, in particular, is highly diverse but appears to remain largely un-documented (DIETRICH & DMITRIEV 2006, 2008).
Study of canopy fogging samples from lowland rainforests in eastern Peru and Ecuador (DIETRICH & WALLNER 2002) indi-cates the presence of an enormously diverse typhlocybine fauna. Composition of this fauna contrasts strikingly with that of the somewhat better known, but still poorly documented faunas of the Old World tropics. Tribes Dikraneurini, Empoascini (= Jorumini) and Alebrini, in decreasing order of species rich-ness, are well represented in the samples from Amazonia; but Erythroneurini and Typhlocybini, so abundant in north tem-perate forests and in the Old World tropics, are relatively rare.
Malaise trap samples from the eastern Andean foothills above 500 m elevation differ from those obtained from the canopy of lowland rainforest (unpublished data) in that they include larger numbers of Typhlocybini (sensu stricto). These species belong to a group of apparently closely related, endemic South American genera that includes Eualebra Baker. The group
also includes Columbonirvana Linnavuori, a genus previously placed in Nirvaninae (LINNAVUORI 1959), but transferred to Typhlocybinae by DIETRICH (2004) based on phylogenetic analy-sis of morphological characters. These two genera include a total of three previously described species (plus one new syn-onym, designated below). Recent sampling indicates that the South American fauna of Typhlocybini is much richer in both species and genera. In this paper, I review the endemic South American genera and species of Typhlocybini and discuss their possible relationships to the fauna of this tribe in other parts of the world. Considering that the 52 new species of Typhlocybini described herein were collected from only eight localities in Bolivia, Brazil, Colombia, Ecuador and Peru, fur-ther sampling in South America, especially in cloud forests of the eastern Andean foothills, will likely reveal the presence of a much more speciose assemblage belonging to this tribe.
MATERIAL AND METHODS
Morphological terminology follows that of YOUNG (1952) as modified by DIETRICH (2005) except wing veins are labeled according to the simplified system illustrated in figures 3i, j. Drawings of genitalia were made by tracing over photographs taken using a digital camera mounted on a compound micro-scope. In drawings of the male genital capsule, the entire anal complex (segments X-XI) is shown in lateral view but only the well sclerotized and pigmented parts of segment X are shown
Christopher H. Dietrich
Illinois Natural History Survey, Prairie Research Institute, University of Illinois, 1816 S. Oak St., Champaign, IL 61820, USA. E-mail: [email protected]
ABSTRACT. The fauna of Typhlocybini (sensu stricto, excluding Empoascini) endemic to South America is reviewed and comprises seven closely related genera, five described herein as new, and 55 species, 52 here described as new. The genera and species are described and keys and illustrations are provided to aid in their identification. Columbonirvana
Linnavuori comprises 17 species, 16 new. Eualebra Baker comprises 19 species, 17 new. Euzyginellagen. nov., comprises four new species. Neozyginellagen. nov., comprises six new species. Pseudhadinagen. nov., comprises one new spe-cies. Pseudozyginellagen. nov., comprises three new species. Tahurellagen. nov., comprises five new species. One new synonym is recognized: Eualebra smithii Baker, 1899 equals Dikraneura (Hyloidea) reticulata Osborn, 1928, syn. nov.
One previously described species placed in Eualebra belongs in tribe Dikraneurini; thus, a new combination is proposed:
Alconeura (Hyloidea) rubra (Van Duzee), comb. nov. Most of the specimens available for this study were from Malaise trap and canopy fogging samples obtained at very few localities in Bolivia, Brazil, Colombia, Ecuador, and Peru, sug-gesting that further sampling in South America, particularly in the Amazonian rainforest and eastern foothills of the Andes Mountains, will reveal large numbers of additional species.
in dorsal view. Material examined is deposited in the follow-ing institutions: Humboldt Institute, Villa de Leyva, Colombia (HIC); Carnegie Museum of Natural History (ICCM); Illinois Natural History Survey, Champaign (INHS); Universidad de San Marcos, Lima (USML), United States National Museum (USNM).
TAXONOMY
Typhlocybinae Kirschbaum, 1868
The tribal classification of Typhlocybinae has long been unstable, with some authors (e.g., YOUNG 1952) having recog-nized as few as four tribes and others (e.g., RUPPEL 1987) as many as ten. In his review of Western Hemisphere Typhlocybinae, YO U N G (1952) employed a relatively broad concept of Typhlocybini, including as synonyms the tribes Eupterygini Kirkaldy, 1906, Jorumini McAtee, 1926, and Empoascini Dis-tant, 1908, despite differences in the hind wing venation of these groups. Dworakowska, the principal worker on the Old World typhlocybine fauna for the past four decades, recognized tribes Typhlocybini, with synonym Eupterygini; Empoascini, with synonym Jorumini; and Zyginellini (DWORAKOWSKA 1979). The latter tribe was erected to comprise species having the hind wing submarginal vein apparently connected directly to CuA rather than being joined by a crossvein (alternatively inter-preted as the absence of the segment of vein CuA distad of its junction with the submarginal vein). Despite the unique hind wing venation, Zyginellini, as defined by DWORAKOWSKA (1979), is a rather heterogeneous assemblage. Most genera placed in this tribe strongly resemble many Typhlocybini (sensu stricto), particularly in the structure of the male genital capsule (e.g., presence of a single macroseta near the base of the plate) and aedeagus (e.g., presence of elongate paired apical processes). Moreover, some of the Neotropical species described herein vary intraspecifically (occasionally intra-individually) for the features used to distinguish Zyginellini from Typhlocybini. AHMED (1983) considered Zyginellini to be an artificial group and treated it as a synonym of Typhlocybini. Until phyloge-netic analyses can clarify the status and relationships of the various tribes proposed for groups of typhlocybine genera, it seems reasonable to adopt the classification of AHMED (1983), which recognizes five tribes in the subfamily: Alebrini, Dikraneurini (= Forcipatini), Empoascini (= Jorumini), Erythroneurini, and Typhlocybini (= Eupterygini, Zyginellini).Key to tribes of Typhlocybinae
1. Forewing appendix present ... Alebrini
1’. Forewing appendix absent ... 2
2. Hind wing with submarginal vein complete, extended along costal margin and connecting apices of all veins (except Typhlocybella: submarginal vein absent, vannal vein unbran-ched, forewing inner apical cell tapered distally) ...
... Dikraneurini
2’. Hind wing submarginal vein incomplete, either extended along costal margin but not connecting vein apices along apical margin, or connecting vein apices along apical margin but not extended along costal margin (terminated at apex of vein RP or R+M) ... 3 3. Forewing with inner apical cell parallel-sided, hind wing submarginal vein absent along apical margin, vannal vein unbranched ... Erythroneurini 3’. Forewing with inner apical cell usually distinctly tapered through most of length (often short and oblique), hind wing submarginal vein, if present, not extended along costal margin basad of RP or R+M, vannal vein branched ... 4 4. Ocelli absent or vestigial, hind wing submarginal vein absent between apices of veins MA or RP+MA and MP (Fig. 78) or, if present (Fig. 70), body strongly depressed with face nearly horizontal ... Typhlocybini 4’. Ocelli well developed (except in Paulomanus), hind wing submarginal vein present between apices of veins MA or RP+MA and MP, body not strongly depressed, face in profile oblique, not nearly horizontal ... Empoascini
Typhlocybini Kirschbaum
Typhlocybini Kirschbaum, 1868: 16.Eupteryginae Kirkaldy, 1906: 296. Zyginellini Dworakowska, 1979: 299.
Diagnosis. Species of Typhlocybini (sensu AHMED 1983) differ from other Typhlocybinae in having the following com-bination of morphological features: ocelli absent or vestigial; forewing appendix absent, inner apical cell short and oblique, not extended to apical wing margin; hind wing vannal vein branched, submarginal vein present or absent at wing apex; male pygofer with few or no macrosetae (except Eualebra).
Remarks. The endemic South American taxa treated herein fit AHMED’s (1983) concept of Typhlocybini, but form a distinctive group within this tribe. Members of this group are readily distinguished by their depressed form, produced head, and presence of two or more macrosetae on the male subgenital plate (most Old World and Nearctic Typhlocybini have 0-1).
Some of the genera included here have hind wing vena-tional patterns not known to occur among Typhlocybini in other parts of the world. One pattern (Figs 72 and 74), exem-plified by the genus Eualebra, has the submarginal vein present and connecting the apices of veins RP, MA, MP and CuA at the wing apex, but absent along the costal margin; veins RP and MA are free, connected by a crossvein.
by a crossvein (as in Eupterygini) or confluent (as in Typhlocybini, s.s.). Also, the hind wing submarginal vein may be strongly curved and colinear with CuA preapically (as in “Zyginellini”) or straight and connected to CuA by a crossvein (as in Typhlocybini, s.s.). Presence of such variation within a single species provides support for AHMED’s (1983) treatment of Eupterygini and Zyginellini as junior synonyms of Typhlocybini. The presence of a submarginal vein at the hind wing apex and relatively numerous macrosetae on the male subgenital plates (as in Alebrini and Empoascini) suggests that Eualebra and related South American genera represent a plesiomorphic lineage of Typhlocybini not represented in the Old World. Other Typhlocybini, including genera and species endemic to the Old World and North America, lack a submar-ginal vein in the hind wing and almost always have two or fewer (usually 0-1) macrosetae on the male subgenital plate.
As in some Old World Typhlocybini, the lower part of the face in most South American genera is sexually dimorphic: females have the clypellus and lorum flattened and well de-limited, but males have the lorum fused to the clypellus and both structures are markedly inflated (Fig. 59).
Based on the morphological similarities mentioned above, the South American fauna of Typhlocybini appear to be related to some Old World genera previously included in “Zyginellini” (sensu DWORAKOWSKA 1979) as well as to the “Eupteryx-complex” (sensu DWORAKOWSKA 1969) of genera that are most diverse in the Indomalayan region. These relationships need to be elucidated by phylogenetic analysis.
The endemic South American fauna of Typhlocybini comprises seven genera: Eualebra, placed by YOUNG (1952) in Typhlocybini based on the branched vannal vein and absence of a submarginal vein along the costal margin in the hind wing; Columbonirvana Linnavuori, previously placed in Nirvaninae presumably based on its depressed form and produced head, but transferred to Typhlocybinae by DIETRICH (2004); and five new genera described below, Euzyginella, Neozyginella, Pseudhadina, Pseudozyginella,and Tahurella.
Two Palearctic species of Typhlocybini known to be es-tablished in Argentina and Chile, Edwardsiana froggatti (Baker) and Ribautiana tenerrima (Herrich-Schaffer), have been well de-scribed elsewhere (CHRISTENSEN 1942, CHRISTIAN 1953) and are not treated here.
Key to endemic genera of South American Typhlocybini
1. Hind wing with submarginal vein obsolete apically (Figs 76, 78, 80, 82, 83); vein RP weak or appearing confluent with MA ... 2
1’. Hind wing with submarginal vein well developed apically (Figs 70, 72, 74, 86); vein RP well developed and separated
either forming continuous line with CuA (Fig. 82) or terminating slightly beyond CuA apex (Fig. 83) ... 3
3. Male connective Y-shaped, stem as long as or longer than arms (Fig. 264); sclerotized base of segment X, in dorsal view, U-shaped, without anterolateral projections (Fig. 262) ... Euzyginella gen. nov.
3’. Male connective U-shaped, stem not developed (Fig. 308); sclerotized base of segment X, in dorsal view, with pair of sclerotized anterolateral projections articulated to pygofer (Fig. 306) ...Pseudozyginella gen. nov.
4. Head with coronal suture not extended beyond midlength of crown, crown with paired red spots (Fig. 43) ...
...Neozyginella gen. nov.
4’. Head with coronal suture extended nearly to apex of crown; crown without spots (Fig. 49) ...Pseudhadina gen. nov.
5. Form broad and strongly depressed (Figs 18-38), face hori-zontal and flat or concave in lateral view (Figs 62, 63); male subgenital plate with numerous macrosetae in irregular longitudinal band (Fig. 8a) ...Eualebra Baker
5’. Form slender and less depressed (Figs 1-17), face oblique and usually convex in lateral view (Figs 60, 61, 68); male subgenital plate with 3-4 macrosetae in single row (Fig. 87) ... 6
6. Dorsal coloration mostly pale yellow with arcuate red and brown transverse bands; forewing apex emarginate or with distinct projection (Figs 84, 85); male subgenital plate with small stout setae at apex in addition to row of macrosetae near midlength (Fig. 317) ...Tahurella gen. nov.
6’. Dorsal coloration mostly brown with pale spots (Figs 1-17); forewing apex rounded (Fig. 69); male subgenital plate without stout apical setae (Fig. 87) ...
...Columbonirvana Linnavouri
Columbonirvana
Linnavuori,
new placement
Columbonirvana Linnavuori, 1959: 34. Type species: C. aurea Linnavuori, 1959.
Diagnosis. Small to medium-sized, depressed, slender leafhoppers. Dorsal coloration (Figs 1-17) orange to dark brown with symmetrical yellow, brown or black markings; forewing apex usually with false eye spot, costal margin with oblique dark brown false veins; ventral coloration including legs yel-low except mesepisternum brown.
anteclypeus narrow, flat, tapered distally, lorum broader than in male; frontoclypeus convex; lateral frontal sutures not ex-tended dorsad of antennal pits; antennal ledges weakly devel-oped; antennae as long as head width; ocelli absent; crown flat or slightly convex, coronal suture extended to or near apex of crown. Pronotum (Figs 1-17) with lateral margins moder-ately long, slightly divergent in dorsal view, distinctly carinate, carina even with posterior margin of eye. Front femur with AM1 enlarged and situated on ventral margin; intercalary row with few fine setae, basal seta larger than others; PV1 absent; tibia rounded dorsally, AD and PD without preapical macrosetae. Hind femur macrosetae 2+1+1; tibia row AV with 4 macrosetae near apex. Forewing (Fig. 69) with apex rounded, RA reflexed, RP confluent with MA for short distance, apical cell 2 triangular or petiolate; CuA joining M basad of its fork, inner apical cell trapezoidal; apical margin convex. Hind wing (Fig. 70) with RP and MA separate, connected by crossvein; submarginal vein extended from apex of RP to jugal lobe.
Male 2S apodemes (Figs 151-166) well developed, joined at base by transverse bridge, subparallel, extended beyond pos-terior margin of sternite III. Pygofer (Figs 87 and 88) strongly emarginate dorsally; tergite variable in length and shape among species; posterior lobe with strongly sclerotized acuminate dor-sal section variably clothed with microtrichia, unpigmented ven-tral section bearing longitidinal row or band of short fine setae; ventral appendage slender, elongate, arising near base of ventral margin and curved posterodorsad, usually areolate distally. Anal tube (Figs 87 and 88) depressed, sclerotized dorsal band short, with pair of basolateral hooks. Subgenital plate (Figs 87 and 88) constricted near base, broadest near midlength, with row of 3-4 macrosetae near lateral margin, apical macrosetae absent; fine dorsal setae divided into medial and distal groups; apical lobe compressed, parallel sided and darkly pigmented. Style (Figs 89 and 90) apodeme short; apophysis elongate, without preapical lobe, with fine setae preapically. Connective (Figs 90 and 94) U-or V-shaped, articulated to aedeagus. Aedeagus (Figs 89 and 90) with preatrium elongate, dorsal apodeme weakly developed; shaft elongate and nearly straight, with or without paired ventral pro-cesses; apical processes absent; gonopore apical.
Female (based only on type species) sternite VII with pos-terior margin angulately produced; ovipositor not extended beyond pygofer; first valvulae with strigate sculpturing dorsally near apex; second valvulae (Fig. 357) elongate, slender, curved dorsad through most of length, slightly sinuate preapically, right blade with 9-10 small, rounded, close-set teeth near apex.
Remarks. The type species of this genus was described based on a single female from Colombia (illustrated by LINNAVUORI 1959 and DIETRICH 2004). No additional specimens matching the type specimen of C. aurea Linnavuori in struc-ture or color pattern have been found, but numerous conge-neric male specimens representing 16 additional species were collected by Malaise trap in cloud forests in the Andes Moun-tains of Peru. No additional female representatives of this
ge-nus have yet been found. DI E T R I C H (2004) transferred Columbonirvana from Evacanthinae (= Nirvaninae) to Typhlocybinae but considered its tribal placement uncertain. The genus is here placed in the tribe Typhlocybini based on the concept of the tribe embodied in the above diagnosis, which is narrower than that of YOUNG (1952) but broader than that of OMANet al. (1990). It is similar to Tahurella gen. nov. but dif-fers in the smaller size, darker overall coloration, lack of trans-verse black band on the anterior margin of the head, and rounded forewing apex.
The Southeast Asian Lowata Dworakowska resembles some species of Columbonirvana in having a pair of narrow transverse red stripes on the anterior margin of the head but differs in having the hind wing submarginal vein absent at the wing apex.
Key to species of
Columbonirvana
1. Pronotum with large white or yellow crescent medially (Fig. 2) ... 2
1’. Pronotum uniformly brown or with pair of small white spots (Figs 1, 7) ... 8
2. Posterior margin of pronotum yellow or white (Fig. 2), aedeagus with pair of elongate processes arising near base and extended distad along shaft (Fig. 93) ... 3
2’. Posterior margin of pronotum brown (Fig. 5); aedeagus without spinelike processes (Fig. 102) ... 5
3. Aedeagus with ventral processes closely appressed to shaft in lateral view (Fig. 93) ...ameliae sp. nov.
3’. Aedeagus with ventral processes distinctly divergent from shaft in lateral view (Fig. 105) ... 4
4. Dorsal sclerotized part of pygofer apex broad and only slightly curved ventrad (Fig. 95) ...craigi sp. nov.
4’. Dorsal sclerotized part of pygofer apex slender and sharply bent ventrad (Fig. 103) ...edgari sp. nov.
5. Aedeagal shaft distinctly denticulate preapically (Fig. 129) .... 6
5’. Aedeagal shaft without denticuli (Fig. 133) ... 7
6. Aedeagus with denticuli restricted to margins of preapical flange (Fig. 150) ...vatia sp. nov.
6’. Aedeagus with denticuli scattered over preapical surface of shaft (Fig. 133) ...thomasi sp. nov.
7. Pygofer with ventral appendage gradually curved dorsad in lateral view (Fig. 127); aedeagal shaft widest at apex in la-teral view (Fig. 129) ...schulzi sp. nov.
7’. Pygofer with ventral appendage abruptly bent dorsad near midlength (Fig. 99); aedeagal shaft widest near base in la-teral view (Fig. 101) ...davidi sp. nov.
8. Anterior margin of head with pair of transverse parallel red stripes (Fig. 61) ... 9
69) ... 16
10. Aedeagus with paired ventral processes (Fig. 109) ... 11
10’. Aedeagus without processes (Fig. 117) ... 14
11. Aedeagal processes arising at or near base of shaft (Fig. 109) ... 12
11’. Aedeagal processes arising at or near midlength of shaft (Fig. 137) ... 13
12. Pygofer with dorsal process gently curved ventrad (Fig. 107); aedeagal processes straight in lateral view (Fig. 109) ...
...ernae sp. nov.
12’. Pygofer with dorsal process bent ventrad at right angle (Fig. 111); aedeagal processes distinctly curved in lateral view (Fig. 113) ...joshuai sp. nov.
13. Aedeagal processes very short, arising from ventrolateral flange, shaft apex not expanded in lateral view (Fig. 141)... ... tumida sp. nov. 13’. Aedeagal processes long, arising from tubular shaft, shaft
apex expanded in lateral view (Fig. 137) ... ...tuberculata sp. nov.
14. Aedeagal shaft in lateral view with apex narrower than base (Fig. 117) ...margaretae sp. nov.
14’. Aedeagal shaft in lateral view with apex wider than base (Fig. 121) ... 15
15. Aedeagal shaft with angulate preapical ventral projection (Fig. 121) ...matthewi sp. nov.
15’. Aedeagal shaft with ventral margin broadly rounded preapically (Fig. 145) ...urodolobrata sp. nov.
16. Forewing with round white spot near midlength of clavus and another on adjacent area of corium (Fig. 1) ...
...aidani sp. nov.
16’. Forewing with oblique white band on clavus extended onto corium (Fig. 60) ...aurea Linnavuori
Columbonirvana aidani
sp. nov.
Figs 1, 69-70, 87-90, 151
Description. Length of male 4.3 mm. Dorsum (Fig. 1) mottled orange brown with symmetrical white spots; head anterior margin with pair of thin red transverse bands between eyes; pronotum with two pairs of posterolateral white spots; forewing clavus with large round white medial spot, brachial cell with smaller white spot, veins orange bordered with dark brown distally; venter of thorax pale yellow except mesoster-num dark brown; abdomen brown ventrally except near midlength of subgenital plates. Head as wide as pronotum, crown slightly longer medially than next to eye.
length of distal lobe; dorsal sclerotized area of lobe weakly fal-cate but not produced beyond margin; ventral appendage strongly curved dorsad, apex areolate, curved posterolaterad (Fig. 87). Anal hook (Fig. 87) elongate and slender, weakly sinu-ate, extended ventrad. Subgenital plate (Fig. 87) with 3-4 macrosetae near midlength, without stout apical setae, apex abruptly curved dorsad. Style apophysis broadened preapically in ventral view (Fig. 87), apex tapered, curved ventrolaterad. Connective U-shaped (Fig. 90). Aedeagus (Figs 89 and 90) with preatrium distinctly shorter than shaft, broadened basally in ventral view; shaft slender, tubular, curved dorsad, with pair of ventral processes arising near base, closely paralleling shaft, and extended to or slightly beyond shaft apex; shaft apex slen-der, compressed.
Material examined. Holotype male, PERU: Pasco, Yanachaga-Chemillén N.P., 10°39’39.7"S 75°22’0.1"W, 2300m, 10-13 Octo-ber 2002, D. Takiya, C. Peña, R. Rakitov, Malaise trap across Rio San Alberto [USML]. Paratypes: 7 males, same data [INHS].
Etymology. This species is named in honor of my nephew, Aidan Dietrich.
Columbonirvana ameliae
sp. nov.
Figs 2, 91-94, 152
Description. Length of male 4.2 mm. Dorsum (Fig. 2) dark reddish brown with symmetrical pale yellow markings; anterior margin of head with pair of thin red transverse bands between eyes; crown yellow bordered anteriorly with brown; pronotum yellow medially with anterior and lateral margins brown and with brown transverse submarginal band posteri-orly; mesonotum yellow medially, anterolateral triangles and apex of scutellum brown; forewing clavus with small white transcommisural spot bordered posteriorly with larger crim-son area, veins orange bordered with brown in distal half; ven-ter of thorax pale yellow except mesosven-ternum orange brown. Head as wide as pronotum, crown nearly twice as long medi-ally as next to eye.
Figures 1-38. Dorsal habitus. (1-17) Columbonirvana: (1) C. aidanisp. nov.; (2) C. ameliaesp. nov.; (3) C. aurea Linnavuori; (4) C. craigi
sp. nov.; (5) C. davidisp. nov.; (6) C. edgarisp. nov.; (7) C. ernaesp. nov.; (8) C. joshuaisp. nov.; (9) C. margaretaesp. nov.; (10) C. matthewisp. nov.; (11) C. noahisp. nov.; (12) C. schulzi sp. nov.; (13) C. thomasisp. nov.; (14) C. tuberculatasp. nov.; (15) C. tumida
sp. nov.; (16) C. urodolobrata sp. nov.; (17) C. vatia sp. nov.; (18-38) Eualebra: (18) E. barbaraesp. nov.; (19) E. charlesisp. nov.; (20)
E. dorisae sp. nov.; (21) E. jessicae sp. nov.; (22) E. patriciae sp. nov.; (23) E. gingeraesp. nov., male holotype; (24) same, female specimen from Ecuador; (25) E. marilynaesp. nov.; (26) E. susanaesp. nov.; (27) E. helenaesp. nov.; (28) E. kathyaesp. nov.; (29) E. peggyae sp. nov.; (30) E. jenniferaesp. nov.; (31) E. smithii Baker, male specimen from Bolivia; (32) E. smithii Baker, female holotype of
D. (H.) reticulata Osborn from Brazil; (33) E. margaretannaesp. nov.; (34) E. albertisp. nov.; (35) E. dorothyaesp. nov.; (36) E. michaelorum
sp. nov.; (37) E. leonisp. nov.; (38) E. moralesisp. nov. Scale bars = 1 mm.
1 2 3 4 5 6 7
8 11 12 13 14 15 17
1
19 21 22 23
24 26
1
29 30 31
32 34 35 36
16
37 38
9 10
18 20
25 27 28
Figures 39-68. Dorsal habitus, lateral habitus, and face. (39-42) Euzyginella: (39) E. markisp. nov.; (40) E. quartalbertisp. nov.; (41) E. teralbertisp. nov.; (42) E. theari sp. nov. (43-48) Neozyginella: (43) N. baileyaesp. nov.; (44) N. bethaesp. nov.; (45) N. bradleyisp. nov.; (46) N. braxtonisp. nov.; (47) N. dougisp. nov.; (48) N. jenncraftae sp. nov.; (49) Pseudhadina amazonicasp. nov.; (50-52)
Pseudozyginella: (50) P. grahamisp. nov.; (51) P. davei sp. nov.; (52) P. dougbraxtoni sp. nov. (53-57) Tahurella: (53) T. josephisp. nov.; (54) T. katherinaesp. nov.; (55) T. lynnaesp. nov.; (56) T. clairaesp. nov.; (57) T. heatheraesp. nov. (58-69) anteroventral view of head: (58) Pseudhadina amazonica, male; (59) Columbonirvana noahi, male. (60-68) lateral habitus: (60) Columbonirvana aurea, holotype female; (61) C. matthewisp. nov.; (62) Eualebra margaretannaesp. nov.; (63) E. michaelorumsp. nov.; (64) Euzyginella quartalbertisp. nov.; (65) Neozyginella baileyisp. nov.; (66) Pseudhadina amazonicasp. nov.; (67) Pseudozyginella grahamisp. nov.) (68) Tahurella lynnaesp. nov. Scale bars = 1 mm.
39 40 41 42 43 44 45
46 47 48 49 50 51
58
52 53 54 55 56
59
60 61 62
63
65 64
66 67 68
straight, with pair of ventral processes arising near base, closely paralleling shaft, and extended 2/3 distance to shaft apex; shaft apex slightly compressed, rounded in lateral view, weakly emar-ginate in ventral view.
Material examined. Holotype male, PERU: Pasco, Yanachaga-Chemillén N.P., 10°39’39.7"S 75°22’0.1"W, 2300m, 10-13 Octo-ber 2002, D. Takiya, C. Peña, R. Rakitov, Malaise trap across Rio San Alberto [USML]. Paratypes: 4 males, same data [INHS].
Etymology. This species is named in honor of my niece, Amelia Dietrich.
Columbonirvana aurea
Linnavuori, 1959
Figs 3, 60, 18a-bColumbonirvana aurea Linnavuori, 1959: 34.
Remarks. This species, the type species of the genus, was described based on a single female from “Sierra San Lorenzo”, Colombia, by LINNAVUORI (1959). DIETRICH (2004) examined the holotype from the Hungarian National Museum, Budapest, and illustrated the ovipositor. No male specimens that could be considered conspecific have been found.
Columbonirvana craigi
sp. nov.
Figs 4, 95-98, 153
Description. Length of male 4.0 mm. Dorsum (Fig. 4) brown with extensive symmetrical pale markings; anterior margin of head with pair of thin red transverse bands between eyes; crown brown medially with broad white anterior and posterior margins; pronotum with white median crescent, pos-terior margin broadly white; mesonotum and scutellum white except midline anterad of scutellar suture and basolateral tri-angles brown; forewing with extensive large symmetrical white markings. Head subequal to pronotum in width, crown with median length much less than twice length next to eye.
Male 2S apodemes (Fig. 153) slender, divergent, extended beyond posterior margin of sternite IV. Pygofer dorsal emar-gination (Fig. 96) quadrate; tergite 1/3 length of distal lobe; dorsal sclerotized area of lobe relatively broad, weakly falcate with angulate dorsal preapical projection, not produced be-yond apical margin; ventral appendage gradually curved dor-sad, apex curved posteromesad (Figs 95 and 96). Anal hook (Fig. 95) well developed, triangular. Subgenital plate (Fig. 95) with 3 macrosetae near midlength and few short, stout preapical setae, apex bent posterodorsad. Style apophysis (Figs 97 and 98) parallel sided through most of length, apex blunt, curved ventrolaterad, extended slightly mesad in ventral view. Con-nective (Fig. 98) broadly V-shaped. Aedeagus (Figs 97 and 98) with preatrium distinctly shorter than shaft, nearly parallel sided in ventral view; shaft slender, tubular, nearly straight, curved slightly dorsad near apex; with pair of slender ventral processes arising near base and extended nearly to apex of shaft, only slightly divergent from shaft; shaft apex slightly com-pressed, rounded in lateral view, truncate in ventral view.
Material examined. Holotype male, PERU: Pasco, Villa Rica, 1400m, 10°43’21"S 75°15’43"W, 21 October 2002, C. H. Dietrich, vacuum, 2-28-1 [USML].
Etymology. This species is named in honor of my brother, Craig B. Dietrich.
Columbonirvana davidi
sp. nov.
Figs 5, 99-102, 154
Description. Length of male 3.4 mm. Dorsum (Fig. 5) dark brown with large symmetrical yellow markings. Anterior margin of head with two narrow transverse red bands; crown with posterior margin infused with yellow medially. Pronotum with large transverse yellow crescent. Mesonotum brown with large yellow median spot. Forewing clavus with large symmetri-cal transcommisural marking; veins mostly orange; anteapisymmetri-cal and apical cells each with medial area hyaline; brachial and discal cell each with hyaline area near apex. Head as wide as pronotum, crown twice as long medially as next to eye.
Male 2S apodemes (Fig. 154) slender, bowed laterad, ex-tended slightly beyond posterior margin of sternite IV. Pygofer emargination parallel sided (Fig. 100); tergite weakly developed and poorly sclerotized; dorsal sclerotized area of lobe gradu-ally tapered distad, apex strongly hooked ventrad but not ex-tended beyond apical margin; ventral appendage gradually curved dorsad, apex extended posterolaterad (Fig. 99). Anal hook (Fig. 99) digitiform, extended ventrad, apex hooked me-sad. Subgenital plate (Fig. 99) with 3 macrosetae near mid-length, without stout apical setae, apex gradually curved posterodorsad. Style apophysis (Figs 101 and 102) slightly broadened preapically, apex tapered, curved ventrolaterad. Connective (Fig. 102) U-shaped. Aedeagus (Figs 101 and 102) with preatrium distinctly shorter than shaft, slightly broad-ened basally in ventral view; shaft tubular, somewhat com-pressed basally, tapering toward apex in lateral view; without paired ventral processes, with indistinct ventrolateral flange extended to apex; apex tubular, subtruncate.
Material examined. Holotype male, PERU: Pasco, Yanachaga-Chemillén N.P., 10°39’39.7"S 75°22’0.1"W, 2300m, 10-13 Octo-ber 2002, D. Takiya, C. Peña, R. Rakitov, Malaise trap across Rio San Alberto [USML]. Paratypes: 4 males, same data [INHS].
Etymology. This species is named in honor of my brother, David A. Dietrich.
Columbonirvana edgari
sp. nov.
Figs 6, 103-106, 155
Figures 69-86. Wings, photographed with transmitted light to highlight venation. (69-70) Columbonirvana aidanisp. nov.) forewing and hind wing; (71-72) Eualebra jenniferaesp. nov., same; (73-74) Eualebra michaelorumsp. nov., same; (75-76) Euzyginella markisp. nov., same, absence of distal part of vein R in hind wing is aberrant; (77-78) Neozyginellajenncraftaesp. nov., same; (79-80) Pseudhadina amazonicasp. nov., same; (81-82) Pseudozyginella grahamisp. nov., same; (83) P. daveisp. nov., hind wing; (84) Tahurella lynnae sp. nov., forewing; (85-86) Tahurella heatheraesp. nov.) forewing and hind wing. (iac) Inner apical cell, (smv) submarginal vein. tended nearly to posterior margin of sternite IV. Pygofer
emar-gination quadrate (Fig. 104), tergite nearly 1/2 length of lobe; dorsal sclerotized area of lobe with digitiform distal section bent ventrad at slightly less than 90° angle, extended slightly beyond margin; ventral appendage gradually curved dorsad, apex curved slightly mesad (Figs 103 and 104). Anal hook (Fig. 103) slender, bifurcate, with slender dorsomedial branch and longer, broader ventral branch curved dorsad in lateral view. Subgenital plate (Fig. 103) with 3 macrosetae near midlength and few short, stout
shaft tubular, very slender, slightly sinuate in lateral view; with pair of acuminate ventral processes arising near midlength, curved away from, then toward shaft in ventral and lateral views; apex obliquely rounded in lateral view, blunt in ventral view.
Material examined. Holotype male, PERU: Pasco, Villa Rica, 1400m, 10°43’21"S 75°15’43"W, 21 October 2002, C.H. Dietrich, vacuum, 2-28-1[USML].
Etymology. This species is named in honor of my father, Edgar H. Dietrich.
69 71 73
70 72 74
75
77
79
76 78 80
81
82
83 85
84
Columbonirvana ernae
sp. nov.
Figs 7, 107-110, 156
Description. Length of male 3.8 mm. Dorsum (Fig. 7) brown, without distinct pale markings; head anterior margin with pair of thin red transverse bands between eyes; forewing veins yellow distally, apical and anteapical cells hyaline in middle, discal and brachial cell each with hyaline spot near
apex. Head subequal to pronotum in width, crown less than twice as long medially as next to eyes.
Male 2S apodemes (Fig. 156) long, slightly divergent, par-allel sided, extended nearly to posterior margin of sternite V. Pygofer emargination with sides divergent posteriorly (Fig. 108), base truncate; tergite ca. 1/3 length of distal lobe; dorsal sclero-tized area of lobe moderately broad, gradually tapered distad, apex hooked ventrad and extended slightly beyond margin; ventral Figure 87-106. Columbonirvana, male genitalia. (87-90) C. aidanisp. nov.: (87) genital capsule) lateral view; (88) pygofer and basal sclerites of anal tube) dorsal view; (89) genitalia) lateral view; (90) genitalia, ventral view, only right style shown. (91-94) C. ameliaesp. nov., same. (95-98) C. craigisp. nov., same. (99-102) C. davidisp. nov., same; (103-106) C. edgarisp. nov., same.
87
91
95
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103 104
100 96 92
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105
few smaller stout setae distally, apex gradually curved dorsad. Style (Figs 109 and 110) somewhat expanded beyond midlength, apex hooked ventrolaterad. Connective (Fig. 5d) V-shaped, stem poorly developed. Aedeagus (Figs 109 and 110) with preatrium nearly as long as shaft; shaft very slender, sinuate in lateral view, with pair of long slender straight processes arising near base and extended distad slightly more than half distance to shaft apex, divergent from shaft in lateral view, parallel in ventral view; shaft apex obliquely truncate in lateral view.
Material examined. Holotype male, PERU: Pasco, Villa Rica, 1400m, 10°43’21"S 75°15’43"W, 21 October 2002, C. H. Dietrich, vacuum, 2-28-1 [USML]. Paratype: 1 male, PERU: Pasco, Yanachaga-Chemillén N.P., 10°39’39.7"S 75°22’0.1"W, 2300m, 10-13 October 2002, D. Takiya, C. Peña, R. Rakitov, Malaise trap across Rio San Alberto [INHS].
Etymology. This species is named in honor of my niece, Erna Dietrich.
Columbonirvana joshuai
sp. nov.
Figs 8, 111-114, 157
Description. Length of male 3.8 mm. Dorsum (Fig. 8) brown, without distinct pale markings; head anterior margin with pair of thin red transverse bands between eyes; forewing veins yellow distally, apical and anteapical cells hyaline in middle, discal and brachial cell each with hyaline spot near apex. Head subequal to pronotum in width, crown approxi-mately twice as long medially as next to eyes.
Male 2S apodemes (Fig. 157) parallel sided, parallel to each other, extended nearly to posterior margin of sternite IV. Pygofer emargination with sides divergent posteriorly (Fig. 112), base truncate; tergite ca. 1/2 length of distal lobe; dorsal scle-rotized area of lobe moderately broad, gradually tapered dis-tad, apex abruptly hooked ventrad and extended slightly beyond margin; ventral appendage acuminate, evenly curved dorsad, apex curved slightly mesad (Figs 111 and 112). Anal hook (Fig. 111) relatively long, tapered, extended ventrad, apex hooked mesad in dorsal view. Subgenital plate (Fig. 111) with 3 macrosetae near midlength, apex gradually curved dorsad. Style (Figs 113 and 114) somewhat expanded beyond midlength, apex hooked ventrolaterad. Connective (Fig. 114) V-shaped, stem absent. Aedeagus (Figs 113 and 114) with preatrium nearly as long as shaft; shaft very slender, sinuate in lateral view, with pair of long tapered evenly curved processes arising near base and extended distad half distance to shaft apex, divergent from shaft in lateral view, parallel in ventral view; shaft apex obliquely truncate in lateral view.
Material examined. Holotype male, PERU: Pasco, Villa Rica, 1400m, 10°43’21"S 75°15’43"W, 21 October 2002, C. H. Dietrich, vacuum, 2-28-1 [USML].
Figs 9, 115-118, 158
Description. Length of male 3.9 mm. Dorsum (Fig. 9) mostly dark brown; head anterior margin with pair of thin red transverse bands between eyes; forewing with oblique pale or-ange mark along posterior part of brochosome field and smaller orange areas more distad along costal margin, veins orange, bordered with brown, central areas of distal cells hyaline; ven-ter of head and thorax stramineous except mesosven-ternum dark brown, abdomen brown ventrally except submedial areas of sternite III, posterior margins of pregenital sternites and basal 2/3 of subgenital plate stramineous. Head narrower than pronotum, crown less than twice as long medially as next to eyes.
Male 2S apodemes (Fig. 158) weakly divergent, extended to posterior margin of sternite IV. Pygofer emargination paral-lel sided, base irregularly rounded (Fig. 116); tergite ca. 1/4 length of distal lobe; dorsal sclerotized area of lobe slender, parallel sided through most of length, apex tapered and de-curved, extended slightly beyond apical margin; ventral ap-pendage gradually curved dorsad, apex directed posteromesad (Figs 115 and 116). Anal hook (Fig. 115) with short antero-ventral process, longer irregularly sinuate antero-ventral extension hooked mesad. Subgenital plate (Fig. 115) with 3-4 macrosetae near midlength. Style (Figs 117 and 118) apophysis with slight preapical gibbosity, apex tapered blunt, weakly hooked ventrolaterad. Connective (Fig. 118) U-shaped with arms strongly bent mesad distally. Aedeagus (Figs 117 and 118) with preatrium much shorter than shaft, narrowest at base in ven-tral view; shaft weakly compressed, nearly straight, somewhat broadened medially, with indistinct angulate dorsolateral flange preapically without paired ventral processes, apex expanded in lateral view, compressed and subtruncate.
Material examined. Holotype male, PERU: Pasco, Yanacha-ga-Chemillén N.P., 10°39’39.7"S 75°22’0.1"W, 2300m, 10-13 October 2002, D. Takiya, C. Peña, R. Rakitov, Malaise trap across Rio San Alberto [USML].
Etymology. This species is named in honor of my grand-mother, Margaret R. Baka.
Columbonirvana matthewi
sp. nov.
Figs 10, 61, 119-122, 159
larger transcommisural band near midlength, veins mostly orange, cells hyaline medially, costal margin with orange pig-ment in distal third. Head subequal to pronotum in width, crown only slightly longer medially than next to eye.
Male 2S apodemes (Fig. 159) weakly divergent, extended to near midlength of sternite IV. Pygofer emargination with lateral margins concave, widest posteriorly (Fig. 120); tergite 1/5 length of distal lobe; dorsal sclerotized area of lobe slen-der, tapered distally, apex curved ventrad and extended slightly beyond margin; ventral appendage with weakly sclerotized jointlike area near midlength, distal portion strongly curved dorsad (Figs 119 and 120). Anal hook (Fig. 119) somewhat elon-gate, divided preapically into digitiform posterolateral and flat-tened anteromedial process. Subgenital plate (Fig. 119) with 3-4 macrosetae near midlength, apex gently curved dorsad. Style (Figs 121 and 122) apophysis parallel sided, apex tapered, blunt, weakly hooked ventrolaterad. Connective (Fig. 122) V-shaped with tiny digitiform medial lobe. Aedeagus (Figs 121 and 122) with preatrium much shorter than shaft, only slightly expanded basally in ventral view; shaft tubular, slender, with indistinct ventrolateral flange ending 1/4 distance from apex, without paired ventral processes, apex compressed with broad triangu-lar ventral keel.
Material examined. Holotype male, PERU: Pasco, Yanachaga-Chemillén N.P., 10°39’39.7"S 75°22’0.1"W, 2300m, 10-13 Octo-ber 2002, D. Takiya, C. Peña, R. Rakitov, Malaise trap across Rio San Alberto [USML]. Paratypes: 3 males, same data [INHS].
Etymology. This species is named in honor of my brother, Matthew M. Dietrich.
Columbonirvana noahi
sp. nov.
Figs 11, 59, 123-126, 160
Description. Length of male 4.0-4.1 mm. Dorsum (Fig. 11) brown marked with white and orange; head anterior mar-gin white; pronotum with anterior marmar-gin orange; mesonotum and scutellum orange; forewing base brown infused with or-ange, costal area with two large white oblique markings; veins orange, cells mostly dark brown distally; venter mostly tan, mesosternum dark brown, abdomen tan ventrally, medial and anterior parts of sternites and apex of subgenital plate brown. Head subequal to pronotum in width, crown less than twice as long medially as next to eyes.
Male 2S apodemes (Fig. 160) straight, slightly divergent, extended slightly beyond posterior margin of sternite IV. Pygofer emargination parallel sided, base broadly V-shaped (Fig. 124); tergite short, ca. 1/6 length of distal lobe; dorsal sclero-tized area of lobe broad, gradually tapered distad, apex strongly hooked ventrad but not extended beyond apical margin; ven-tral appendage broadened medially and abruptly narrowed and somewhat twisted preapically, curved dorsad, apex bent posterad (Figs 123 and 124). Anal hook (Fig. 123) bifid with two processes digitiform and subequal in length, extended ventromesad. Subgenital plate (Fig. 123) with 3-4 macrosetae
near midlength, without stout apical setae, apex gradually curved dorsad. Style (Figs 125 and 126) apex somewhat ex-panded near apex, apex hooked ventrolaterad. Connective (Fig. 126) broadly V-shaped. Aedeagus (Figs 125 and 126) with preatrium nearly as long as shaft; shaft compressed, relatively broad in lateral view, without paired ventral processes, with weak ventrolateral flange; apex tubular and subtruncate.
Material examined. Holotype male, PE R U: Pasco, Yanachaga-Chemillén N.P., 10°39’39.7"S 75°22’0.1"W, 2300m, 10-13 October 2002, D. Takiya, C. Peña, R. Rakitov, Malaise trap across Rio San Alberto [USML]. Paratype: 1 male, same data [INHS].
Etymology. This species is named in honor of my nephew, Noah Dietrich.
Columbonirvana schulzi
sp. nov.
Figs 12, 127-130, 161
Description. Length of male 4.6 mm. Dorsum (Fig. 12) dark brown with large symmetrical yellow markings. Anterior margin of head with two narrow transverse red bands; crown with posterior margin infused with yellow medially. Pronotum with large transverse yellow crescent. Mesonotum yellow ex-cept dark brown anterolateral triangles; scutellum with ante-rior half yellow. Forewing clavus with large diamond-shaped transcommisural marking; veins mostly orange; anteapical and apical cells each with medial area hyaline; brachial and discal cell each with hyaline area near apex. Head as wide as pronotum, crown twice as long medially as next to eye.
Male 2S apodemes (Fig. 161) broad, tapered, not ex-tended beyond posterior margin of sternite III. Pygofer emar-gination nearly parallel sided (Fig. 128); tergite weakly developed and poorly sclerotized; dorsal sclerotized area of lobe abruptly tapered distad, apex hooked ventrad at 90° angle but not extended beyond apical margin; ventral appendage areolate distally, gradually curved dorsad, apex extended posterolaterad well beyond pygofer apex (Fig. 127). Anal hook (Fig. 127) short, weakly bidentate, extended anteromesad. Subgenital plate (Fig. 127) with 4 macrosetae near midlength, without stout apical setae, apex gradually curved posterodor-sad. Style (Figs 129 and 130) apophysis abruptly narrowed preapically, apex tapered, curved ventrolaterad. Connective (Fig. 130) Y-shaped; stem broad, slightly shorter than arms. Aedeagus (Figs 129 and 130) with preatrium shorter than shaft, parallel sided basally in ventral view; shaft strongly com-pressed, in lateral view nearly straight, parallel sided through most of length, apex broadened and subtruncate; without pro-cesses or flanges.
Material examined. Holotype male, PE R U: Pasco, Yanachaga-Chemillén N.P., 10°39’39.7"S 75°22’0.1"W, 2300m, 10-13 October 2002, D. Takiya, C. Peña, R. Rakitov, Malaise trap across Rio San Alberto [USML].
Figure 107-126. Columbonirvana, male genitalia. (107-110) C. ernaesp. nov.: (107) genital capsule) lateral view; (108) pygofer and basal sclerites of anal tube, dorsal view; (109) genitalia, lateral view; (110) genitalia, ventral view, only right style shown; (111-114) C. joshuaisp. nov., same; (115-118) C. margaretaesp. nov., same; (119-122) C. matthewisp. nov., same; (123-126) C. noahisp. nov., same.
Columbonirvana thomasi
sp. nov.
Figs 13, 131-134, 162
Description. Length of male 3.4-3.9 mm. Dorsum (Fig. 13) dark brown with large symmetrical yellow markings. Ante-rior margin of head with two narrow transverse red bands; crown with posterior margin infused with yellow medially. Pronotum with large transverse yellow crescent. Mesonotum and scutellum without pale markings. Forewing clavus with large pentagonal transcommisural marking; veins mostly or-ange; anteapical and apical cells each with medial area
hya-line; brachial and discal cell each with hyaline area near apex. Head as wide as pronotum, crown twice as long medially as next to eye.
Male 2S apodemes (Fig. 162) robust, parallel-sided, ex-tended beyond posterior margin of sternite IV. Pygofer emar-gination nearly parallel sided (Fig. 132); tergite weakly developed and poorly sclerotized; dorsal sclerotized area of lobe broadend preapically, abruptly bent ventrad, apex tapered, not extended beyond apical margin; ventral appendage areolate distally, abruptly bent dorsad near midlength, apex not extended be-yond pygofer apex (Fig. 131). Anal hook (Fig. 131) short,
sinu-107 108 110
112 111
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123
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126 125 122 121
118 117
114 113
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ate in lateral view, hooked mesad. Subgenital plate (Fig. 131) with 3 macrosetae near midlength, without stout apical setae, apex gradually curved posterodorsad. Style (Figs 133 and 134) apophysis narrowed preapically, apex tapered, curved ventrolaterad. Connective (Fig. 134) V-shaped. Aedeagus (Figs 133 and 134) with preatrium shorter than shaft, slightly broad-ened basally in ventral view; shaft strongly compressed, in lat-eral view nearly straight, gradually tapered through most of length, apex broadened and obliquely truncate; with pair of preapical ventrolateral flanges and numerous preapical denticuli. Material examined. Holotype male, PERU: Pasco, Yanachaga-Chemillén N.P., 10°39’39.7"S 75°22’0.1"W, 2300m, 10-13 Octo-ber 2002, D. Takiya, C. Peña, R. Rakitov, Malaise trap across Rio San Alberto [USML]. Paratypes: 4 males, same data [INHS].
Etymology. This species is named in honor of my brother-in-law, Thomas Braxton.
Columbonirvana tuberculata
sp. nov.
Figs 14, 135-138, 163
Description. Length of male 4.3 mm. Dorsum (Fig. 14) mostly brown, venter pale yellow except medial part of frontoclypeus, gena below eye, and mesepisternum brown. Head anterior margin with pair of transverse red bands, crown, pronotum, mesonotum and scutellum brown infused with stramineous; forewing with indistinct stramineous band ex-tended from base of clavus across mesonotum and another larger transcommisural band near midlength, veins mostly orange, cells hyaline medially, costal margin with orange pig-ment in distal third. Head subequal to pronotum in width, crown only slightly longer medially than next to eye.
Male 2S apodemes (Fig. 163) robust, parallel sided, ex-tended to posterior margin of sternite IV. Pygofer emargination with lateral margins slightly divergent posteriorly (Fig. 136); tergite weakly sclerotized; dorsal sclerotized area of lobe with dorsal tooth adjacent to anal hook, apex curved ventrad and extended slightly beyond margin; ventral appendage areolate distally, abruptly bent dorsad near midlength and extended posterodorsad slightly beyond dorsal pygofer margin (Figs 135 and 136). Anal hook (Fig. 135) short, weakly bidentate, curved anteromesad. Subgenital plate (Fig. 135) with 3-4 macrosetae near midlength, apex gently curved dorsad. Style apophysis (Figs 137 and 138) parallel sided, apex tapered, blunt, weakly hooked ventrolaterad. Connective (Fig. 138) V-shaped, stem not devel-oped. Aedeagus (Figs 137 and 138) with preatrium much shorter than shaft, slightly expanded basally in ventral view; shaft tu-bular, slender, somewhat compressed distally, apex slightly broadened and obliquely truncate in lateral view; with pair of slender ventral processes arising near midlength, extended dis-tad and slightly divergent from shaft, not reaching shaft apex. Material examined. Holotype male, PERU: Pasco, Yanacha-ga-Chemillén N.P., 10°39’39.7"S 75°22’0.1"W, 2300m, 10-13 October 2002, D. Takiya, C. Peña, R. Rakitov, Malaise trap across Rio San Alberto [USML].
Etymology. The species name refers to the small dorsal preapical tooth (tubercle) on the male pygofer lobe.
Columbonirvana tumida
sp. nov.
Figs 15, 139-142, 164
Description. Length of male 4.2 mm. Dorsum (Fig. 15) mostly brown, venter pale yellow except mesepisternum brown. Head anterior margin with pair of transverse red bands, crown, pronotum, mesonotum and scutellum brown infused with stramineous; forewing with indistinct stramineous band ex-tended from base of clavus across mesonotum and another larger transcommisural band near midlength, veins mostly orange, cells hyaline medially, costal margin with orange pig-ment in distal third. Head subequal to pronotum in width, crown only slightly longer medially than next to eye.
Male 2S apodemes (Fig. 164) relatively slender, parallel-sided, extended slightly beyond sternite IV. Pygofer emargin-ation with lateral margins slightly divergent posteriorly (Fig. 140); tergite weakly sclerotized; dorsal sclerotized area of lobe constricted preapically, apex curved ventrad and extended slightly beyond margin; ventral appendage areolate distally, abruptly bent dorsad near midlength and extended postero-dorsad slightly beyond dorsal pygofer margin (Figs 139 and 140). Anal hook (Figs 139 and 140) short, sinuate in lateral view, hooked mesad. Subgenital plate (Fig. 139) with 3-4 macrosetae near midlength, apex gently curved dorsad. Style apophysis (Figs 141 and 142) parallel sided, apex tapered, blunt, weakly hooked ventrolaterad. Connective (Fig. 142) V-shaped. Aedeagus (Figs 141 and 142) with preatrium short, slightly expanded basally in ventral view; shaft slender, somewhat compressed distally, apex slightly broadened and obliquely rounded in lateral view; with pair of ventrolateral flanges broadening distad in ventral view and each giving rise to spine extended distad and slightly diver-gent from shaft, not reaching shaft apex.
Material examined. Holotype male, PERU: Pasco, Yanacha-ga-Chemillén N.P., 10°39’39.7"S 75°22’0.1"W, 2300m, 10-13 October 2002, D. Takiya, C. Peña, R. Rakitov, Malaise trap across Rio San Alberto [USML].
Etymology. The species name refers to the tumid (swol-len) medial section of the aedeagal shaft.
Columbonirvana urodolobrata
sp. nov.
Figs 16, 143-146, 165
Figures 127-150. Columbonirvana, male genitalia. (127-130) C. schulzi sp. nov.: (127) genital capsule) lateral view; (128) pygofer and basal sclerites of anal tube, dorsal view; (129) genitalia, lateral view; (130) genitalia, ventral view, only right style shown; (131-134) C. thomasisp. nov., same; (135-138) C. tuberculatasp. nov., same; (139-142) C. tumidasp. nov., same; (143-146) C. urodolobrata sp. nov., same; (147-150) C. vatiasp. nov., same.
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Male 2S apodemes (Fig. 165) robust, short, tapered, ex-tended slightly beyond sternite III. Pygofer emargination with lateral margins slightly divergent posteriorly(Fig. 144); tergite sclerotized in basal fifth; dorsal sclerotized area of lobe very slender, apex abruptly curved ventrad and extended slightly beyond margin; ventral appendage areolate distally, abruptly bent dorsad near midlength, bowed mesad and extended posterodorsad slightly beyond dorsal pygofer margin (Figs 143 and 144). Anal hook (Fig. 143) short, sinuate in lateral view, abruptly constricted preapically, hooked mesad. Subgenital plate (Fig. 143) with 3 macrosetae near midlength, apex gently curved dorsad. Style apophysis (Figs 145 and 146) parallel sided, apex tapered, blunt, weakly hooked ventrolaterad. Connective (Fig. 146) V-shaped. Aedeagus (Figs 145 and 146) with preatrium shorter than shaft, tapered basally in ventral view; shaft strongly compressed, nearly straight, slightly tapered in lateral view; apex strongly expanded with acute posterodorsal angle and rounded posteroventral lobe; without processes.
Material examined. Holotype male, PERU: Pasco, Yanacha-ga-Chemillén N.P., 10°39’39.7"S 75°22’0.1"W, 2300m, 10-13 October 2002, D. Takiya, C. Peña, R. Rakitov, Malaise trap across Rio San Alberto [USML].
Etymology. The species name combines uro, meaning “tail”, with dolobrata, meaning “hatchet”, referring to the hatchetlike apex of the aedeagus.
Columbonirvana vatia
sp. nov.
Figs 17, 147-150, 166
Description. Length of male 3.4-3.9 mm. Dorsum (Fig. 17) dark brown with symmetrical white markings; anterior margin of head with pair of thin red transverse bands between eyes; crown white bordered anteriorly with brown; pronotum white medially with anterior and lateral margins brown and with brown transverse submarginal band posteriorly; mesonotum white medially, anterolateral triangles and apex of scutellum brown; forewing clavus with large white transcommisural spot, veins orange bordered with brown in distal half; venter of thorax pale yellow except mesosternum orange brown. Head as wide as pronotum, crown slightly longer medially than next to eye.
Male 2S apodemes (Fig. 166) robust, subparallel, extended to posterior margin of sternite IV. Pygofer dorsal emargination quadrate, lateral margins constricted medially (Fig. 148); terg-ite 1/3 length of distal lobe; dorsal sclerotized area of lobe fal-cate, produced slightly beyond apical margin; ventral appendage gradually curved dorsad, apex curved dorsolaterad (Figs 147 and 148). Anal hook (Fig. 147) long, tapered, sinu-ate, apex hooked mesad. Subgenital plate (Fig. 147) with 3 macrosetae near midlength and few short, stout preapical se-tae, apex bent dorsad. Style apophysis (Figs 149 and 150) slightly sinuate, apex tapered, curved ventrolaterad then hooked mesad. Connective (Fig. 150) U-shaped. Aedeagus (Figs 149 and 150) with preatrium much shorter than shaft, not
expanded basally in ventral view; shaft slender, tubular, nearly straight, with pair of rounded, finely serrate ventrolateral flanges arising near apex; shaft apex compressed, slightly ex-panded, obliquely rounded in lateral view.
Material examined. Holotype male, PERU: Pasco, Yanachaga-Chemillén N.P., 10°39’39.7"S 75°22’0.1"W, 2300m, 10-13 Octo-ber 2002, D. Takiya, C. Peña, R. Rakitov, Malaise trap across Rio San Alberto [USML]. Paratypes: 2 males, same data [INHS].
Etymology. The species name is derived from the Latin word vatius, meaning “bent”, and refers to the sharply bent pygofer appendages.
Eualebra
Baker
Eualebra Baker 1899: 402.Type species: E. smithii Baker 1899.
Diagnosis. Small to medium sized, strongly depressed, somewhat ovoid leafhoppers (Figs 18-38) broadest across midlength of resting forewings in dorsal view. Color pale yel-low to dark brown with symmetrical white, yelyel-low, orange, brown, and/or black markings dorsally.
Head (Figs 18-38) narrower than pronotum, depressed, produced, anterior margin parabolic in dorsal view; crown slightly convex, declivous, glabrous, coronal suture restricted to posterior third; face horizontal (Figs 62-63); rostrum not extended beyond front trochanters; lower part of face sexually dimorphic, male anteclypeus inflated and expanded laterad, parallel-sided and weakly convex in female; lorum very nar-row; frontoclypeus flat or concave medially; lateral frontal su-tures not extended dorsad of antennal pits; antennal ledges curved, nearly vertical; antennae slightly longer than head width; ocelli vestigial, on crown margin approximately mid-way between eye and midline. Pronotum (Figs 18-38) with lat-eral margins long, divergent in dorsal view, distinctly carinate, carina even with posterior corner of eye. Front femur with AM1 enlarged and situated on ventral margin; intercalary row with few fine setae, basal seta larger than others; PV1 well devel-oped; tibia somewhat flattened dorsally, AD and PD without preapical macrosetae. Hind femur macrosetae 2+1+1; tibia row AV with 3-4 macrosetae near apex. Forewing (Figs 71 and 73) with RA slightly to strongly reflexed, RP and MA separate and connected by crossvein or confluent for short distance preapically. Hind wing (Figs 72 and 74) with anterior branch of R absent, veins RP and MA separate, joined by crossvein; MP and CuA separate, joined by oblique m-cu crossvein; sub-marginal vein extended from apex of RP to jugal lobe.
append-age usually well developed, arising from base of anal tube (seg-ment X), articulated to posterodorsal margin of pygofer later-ally and to dorsal connective medilater-ally, usulater-ally with at least one separate projection extended mesad and/or ventrad. Scle-rotized dorsal connective present (Fig. 214), usually U-shaped (rarely platelike) and articulated between dorsal apodeme of aedeagus and paired basal processes of anal tube. Valve very short, rectangular, fused to pygofer. Subgenital plate (Fig. 167) depressed, ovoid in ventral view, distal half tapered with 5 or more macrosetae arranged in lateral band, fine setae short and sparse on dorsal surface, apex rounded or subtruncate. Style (Figs 169 and 170) apodeme short, broad; apophysis elongate, slender, apex simple, tapered and slightly hooked, without con-spicuous setae. Connective (Fig. 170) U-shaped with pair of posterodorsal lobes articulated to aedeagus. Aedeagus (Figs 169 and 170) with preatrium well developed; dorsal apodeme short, lobelike; shaft short, tubular, with or without processes; go-nopore apical.
Female with posterior margin of sternite VII (Fig. 361) angulately produced medially; ovipositor not extended beyond pygofer; first valvulae Figs 359, 364, and 369) with dorsal
sculp-turing strigate; second valvulae (Figs 362, 370, and 374) vari-able interspecifically in shape and dentition.
Remarks. DIETRICH (2004) noted that Eualebra and Columbonirvana share a unique hind wing venational pattern different from that of other Typhlocybinae and therefore treated both genera as unplaced to tribe within Typhlocybinae. Based on the revised tribal diagnosis given above, both of these genera are now included in Typhlocybini (also see Remarks under Columbonirvana). Species of Eualebra resemble species of the Old World genus Eurhadina Haupt in their ovoid shape in dorsal view and strongly depressed body form. However Eurhadina differs in having the structure of the lower part of the face similar in males and females, the forewing narrow dis-tally, the male subgenital plates usually with a single subbasal macroseta, and the pygofer lacking an elongate ventral pro-cess. Also, like all other known Old World Typhlocybini, Eurhadina lacks a submarginal vein at the apex of the hind wing. Species of Eualebra occur in both lowland Neotropical rainforest and premontane cloud forests.
Eualebra previously included five species. YOUNG (1952) excluded E. notata Baker from Guatemala, noting that the hind Figures 151-166. Columbonirvana, base of male abdomen, ventral view. (151) C. aidanisp. nov.; (152) C. ameliaesp. nov.; (153) C. craigisp. nov.; (154) C. davidisp. nov.; (155) C. edgarisp. nov.; (156) C. ernaesp. nov.; (157) C. joshuaisp. nov.; (158) C. margaretae
sp. nov.; (159) C. matthewisp. nov.; (160) C. noahisp. nov.; (161) C. schulzi sp. nov.; (162) C. thomasisp. nov.; (163) C. tuberculata
sp. nov., right apodeme broken; (164) C.tumidasp. nov.; (165) C.urodolobrata sp. nov.; (166) C.vatia sp. nov.
154
162 158
165 164
161 157 153 152
151
155
156
160 159
wing venation of the female holotype is consistent with that of tribe Dikraneurini, but did not suggest an appropriate ge-neric placement for the species so it remains incertae sedis. Ex-amination of the holotype of Eualebra rubra Van Duzee from Jamaica, deposited in the California Academy of Sciences col-lection, indicates that it belongs in the subgenus Alconeura (Hyloidea) of tribe Dikraneurini. Therefore, A. (H.) rubra (Van Duzee) is proposed as a new combination. As a result, only three taxa previously included, E. smithii Baker, E. reticulata Osborn, and E. rufoornata (Stål), all described from Brazil, are retained in Eualebra. Seventeen new species, described below, are also included. Based on the species known at present, the genus appears to be restricted to South America.
Eualebra, as presently defined, is somewhat heteroge-neous morphologically, comprising species that range in size from 2.7 to 4.7 mm and exhibit a variety of color patterns, shapes, and male genitalia configurations. Given the still poor state of knowledge of the genus, and particularly considering that females are known for only a few species, I have opted not to recognize additional genera or subgenera but, rather, place the known species into five informal species groups that may represent distinct lineages. Recognition of some of these groups as separate genera may be warranted once the fauna becomes better known. In particular, the ovipositors of the few female specimens available for study exhibit considerable variation and may provide genus-level characters.
Key to species of
Eualebra
1. Body length >4.5 mm, head in profile with anterior margin thick (Fig. 63), hind tibia row AV with three macrosetae (michaelorum species group) ... 2
1’. Body length < 4.5 mm, head in profile with anterior margin thin (Fig. 62), hind tibia row AV usually with four macrosetae ... 4
2. Dorsal coloration mostly pale yellow with few brown or black markings (Fig. 34) ... 3
2’. Dorsal coloration mostly brown with smaller pale markings (Fig. 36) ...michaelorum sp. nov.
3. Pronotum with narrow median posterior macula; forewing clavus unmarked (Fig. 35) ...dorothyae sp. nov.
3’. Pronotum with broad transverse macula on posterior margin; forewing clavus with distinct transverse brown macula at midlength (Fig. 34) ...alberti sp. nov.
4. Dorsum color pale stramineous marked with numerous minute brown spots (Fig. 31), forewing with irregular cream-colored spots, crown unmarked or with small symmetrically arranged pale orange markings (smithii species group) .. 5
4’. Dorsal color pattern not consisting of small brown spots on a stramineous background, forewing with cream-colored spots absent or large and symmetrically arranged, crown with orange markings, if present, large ... 6
5. Forewing clavus with small round white spots (Fig. 33); aedeagus with paired lateral flanges extended full length of shaft (Fig. 10t); female sternite VII with posterior margin broadly but shallowly produced (Fig. 18p) ...
...magaretannae sp. nov.
5’. Forewing clavus with irregular white vermiform areas (Fig. 31); aedeagus without paired lateral flanges (Fig. 10p); female sternite VII with short median lobe .smithii Baker
6. Crown and pronotum with pair of orange or brown longitu-dinal stripes extended onto pronotum (Fig. 20); length >4.0 mm (dorisae species group) ... 7
6’. Crown and pronotum without pair of longitudinal stripes or, if stripes present, length <3.0mm ... 9
7. Dorsum predominantly bright yellow (Fig. 22); male tergite IX with narrow, parallel-sided posterior emargination (Fig. 184); aedeagus without processes (Fig. 185) ...
...patriciae sp. nov.
7’. Dorsum predominantly brown; male tergite IX with broad, obtusely angulate emargination (Fig. 8j); aedeagus with pair of processes (Fig. 178) ... 8
8. Crown width between eyes much less than 2X median length (Fig. 21); aedeagus with paired processes arising near apex (Fig. 182) ...jessicae sp. nov.
8’. Crown width between eyes ~2X median length (Fig. 20); aedeagus with paired processes arising near base (Fig. 178) ...dorisae sp. nov.
9. Pronotum with narrow white median cruciform or trident-shaped mark surrounded by brown (Fig. 18); length <3 mm (barbarae species group) ... 10
9’. Pronotum with white marking, if present, broad and not cruciform or trident-shaped; length >3.0 mm ... 11
10. Male pygofer with elongate dorsal spine and short, bifurcate ventral process (Fig. 167); aedeagus strongly curved in la-teral view (Fig. 169) ...barbarae sp. nov.
10’. Male pygofer with dorsal spine absent, ventral process long and acuminate (Fig. 8e); aedeagus nearly straight in lateral view (Fig. 173) ...charlesi sp. nov.
11. Crown brown with pale marking, if present, restricted to narrow area near apex (Fig. 38) ... 12
11’. Crown with large medial lobed or wedge-shaped pale area (Fig. 23, 29) (gingerae species group, part) ... 14
12. Pronotum uniformly brown, without white spots (Fig. 30); forewing clavus with pair of triangular yellow transcommi-sural markings (gingerae species group, part) ...
...jenniferae sp. nov.
