One new genus and three new species in the leafhopper tribe Coelidiini from the Neotropical Region (Hemiptera, Cicadellidae, Coelidiinae)

Abstract One new leafhopper genus Paracodiliagen. n. with one new species P.geniculatasp. n., and two other new species in different genera, Bolivielaexpanda and Armaturolidiasymmetricaspp. n., are described in the tribe Coelidiini (Cicadellidae: Coelidiinae) from the Neotropical region. Photographs and illustrations are provided.


Introduction
Coelidiinae is a large leafhopper subfamily found throughout the tropics. Its members are distinguished externally mainly by the very narrow vertex compared to the extremely large eyes (Figs 1,10,21) and long face with antennae situated near the lower corners of the eyes in facial view (Figs 3,12,23). In the Neotropics, there are six tribes of Coelidiinae, the largest being the Coelidiini, which includes half of the 70 known genera of the tribe (see Nielson 2011 for a key to the genera). In this paper, we describe and illustrate one new genus Paracodilia gen. n. with a new species, two new species of the genera Boliviela and Armaturolidia, and provide a brief discussion on the tribe from the region.

Materials and methods
The male genitalia of the specimens examined were macerated in 10% NaOH and observed in glycerin jelly using an Olympus SZX10 stereomicroscope. All drawings were made using an Olympus drawing tube. A Canon EOS 5D Mark II camera and lifting table controlled by CAMLIFT V2.7.0 were using to take photographs at different focal planes and then stacked by ZERENE STACKER. Photos were also taken by a ZEISS SteREO Discovery.V20 stereomicroscope equipped with a ZEISS Axiocam ICc 5 camera that also provided measurements.

Subfamily Coelidiinae Dohrn, 1859
Diagnosis. In addition to the external characters noted in the Introduction the following characters are diagnostic. Prothracic tibia with accessory row of setae between AD and AV setae rows. Forewing with three anteapical cells, usually with only the outer one closed, and appendix well developed and extended around the apex (Felix et al. 2002: 163, figs 6-9). Male genitalia with pygofer fused to valve and caudal corner produced slightly to strongly, often with processes or pair of lobes; connective Y-shaped; styles sometimes asymmetrical; dorsal connective usually present, attached dorsally to base of aedeagus, variable in length and shape (stick-like, Y-shaped, n-shaped or slightly bifurcate at apex). Female genitalia with valvulae long and slender, second valvulae with a prominent middorsal preapical tooth and additional asymmetrical teeth distally (Fan et al. 2015: 475, figs 81, 82;Viraktamath and Meshram 2017: 276, figs 29-31).

Tribe Coelidiini Dohrn, 1859
Diagnosis. Coelidiini can be differentiated from other tribes by the following combination of characters. Crown moderately long, midlength produced distally beyond anterior margin of eyes, distal length 1/4 to 1/3 of entire median length. Eyes large, occupying approx. 2/3 of entire dorsal area of head (more than 2/3 in Tinobregmini and less than 2/3 in Youngolidiini). Clypeus elongate, maximum width more than 1/2 but less than 3/4 of length, without median longitudinal carina or present but incomplete in a few species (complete in Teruliini). Pygofer triangle with 0-3 pairs of caudal processes. Dorsal connective usually present, stick-like, sometimes bifurcated at base or apex. Subgenital plate compressed, not segmented (depressed and segmented in Thagriini and Tharrini). Segment X slightly shorter to longer than pygofer dorsal length (very short in Thagriini).
Etymology. The generic name refers to the similarity between the new genus and Codilia Nielson, 1982. Diagnosis. General color deep brown. Crown light brown with distal 1/4 orange, pronotum and mesonotum pitchy without spots; eyes piceous; forewing with distinct yellow mark on clavus, veins without spots (Figs 1, 2). Face fuscous (Figure 3).
Anterior margin of head obtusely angled. Crown midlength produced distally beyond eyes, approx. 1⁄ 3 of entire length; area between eyes as wide as eye width, surface rugulose, elevated above eyes; lateral margins slightly carinate, not surpassing anterior margin of eyes, slightly convergent basally. Medial length of pronotum distinctly longer than crown and quite shorter than mesonotum ( Figure 1). Forewings typical of subfamily ( Figure 2). Clypeus long and narrow, lateral margins slightly convex; clypellus broad, maximum width approx. equal to midlength, base inflated, lateral margins constricted medially ( Figure 3).
Male genitalia. Valve triangulate in ventral view. Pygofer narrow in lateral aspect, caudodorsal margin produced and tapered in lateral view, without triangular plates at base (Figure 4), caudoventral processes short, covered with short spines, adjacent to base of caudodorsal processes ( Figure 5). Segment X elongate, lateral margins parallel in ventral view ( Figure 4). Aedeagus moderately long and narrow, somewhat tubular, bent at midlength in lateral aspect, apex asymmetrically bifurcate; gonopore apical on ventral surface between bifurcations (Figs 6, 7); dorsal connective absent. Connective large, Y-shaped with stem broad, shorter than length of arms ( Figure 6). Style very long, much longer than aedeagus, sinuate in lateral view, tapered to acute apex (Figs 6,8). Subgenital plate narrow, lateral margin abruptly tapered distally with dense setae (Figure 9).
Remarks. This new genus is similar to genus Codilia Nielson, 1982 externally, but has very different male genitalia. The aedeagus is distinctly curved at midlength in lateral view and apex asymmetrically bifurcate, but in Codilia the aedeagus is slightly curved with a large retrorse spine at the apex. Also in the new genus the subgenital plate is highly setose laterally in its apical half but sparsely setose in Codilia. Diagnosis. Pygofer with caudoventral processes short, covered with short spines, adjacent to base of caudodorsal processes; aedeagus somewhat tubular, bent at midlength in lateral aspect, apex asymmetrically bifurcate, gonopore apical on ventral surface between bifurcations, dorsal connective absent; style much longer than aedeagus, tapered to acute apex; subgenital plate with dense setae.
Etymology. The species epithet is derived from the Latin word geniculus, referring to the aedeagus being geniculate or bent medially in lateral view. Diagnosis. Pygofer with caudodorsal margin produced as a curved, tapered process, caudoventral processes subbasally, apex slightly enlarged in lateral aspect, extended beyond caudodorsal processes; aedeagus large and broad, lateral edge rolled ventrally, tapered to compressed apical 1/4 in lateral view, apex curved dorsally, gonopore subapical on lateral surface, dorsal connective bifurcate at base, enclosing aedeagus subbasally; style reaching to apex of aedeagus, needle-like with apical third enlarged, corrugated, a small process at subapex; subgenital plate with dense hair-like setae at apical half.
External morphology. Medium species, body slender. General color dark brown. Crown, pronotum and mesonotum deep brown with trivial light-brown spots, some gathered in patches; eyes piceous; forewing brown with pellucid markings, veins dark brown with pale spots (Figs 10, 11). Face fuscous with dull yellow tiny spots (Figure 12).
Anterior margin of head obtusely angular. Crown midlength produced distally approx. 1⁄ 3 of entire length; area between eyes approx. as wide as eye width, surface without wrinkles, elevated above level of eyes; lateral margins parallel, slightly carinate, surpassing anterior margin of eyes distally. Medial length of pronotum longer than crown but shorter than mesonotum ( Figure 10). Clypeus long, narrow, lateral margins convex basally; clypellus broad, maximum width approx. equal to midlength, base inflated, lateral margins constricted medially, tapered apically ( Figure 12).
Male genitalia. Pygofer short, caudodorsal margin produced as a curved, tapered process (Figure 14), basally with a triangular plate on each side, caudoventral processes subbasally, very long, apex slightly enlarged in lateral aspect, extended beyond caudodorsal processes; segment X broad and long with lateral margins parallel in ventral view ( Figure 13). Aedeagus large and broad, lateral edge rolled ventrally, tapered to compressed apical 1/4 in lateral view, apex curved dorsally; gonopore subapical on lateral surface; dorsal connective bifurcate at base, enclosing aedeagus subbasally (Figs 15, 16). Connective very small, Y-shaped, anterior stem very short (Figure 19). Style long, reaching to apex of aedeagus, needle-like with apical third enlarged, corrugated, a small process at subapex (Figs 17, 18). Subgenital plate narrow, outer lateral margin convex at apical third, tapered to apex, dense hair-like setae at apical half ( Figure 20).
Etymology. The specific epithet is descriptive of the expanded aedeagus in dorsal and lateral views.
Remarks. The new species keys to Collasuyusana Nielson in Nielson (2011), but its aedeagal shaft is atypical of this genus. We therefore provisionally assign the species to Boliviela based on the inflated aedeagal shaft, despite the dorsal connective being atypical of this genus. According to Prof. Nielson, retired faculty of Brigham Young University (pers. com.) the aedeagus holds historically a higher diagnostic value than the dorsal connective. Diagnosis. Pygofer with caudodorsal margin moderately produced, apex rounded, with two long and narrow processes subbasally, surpassing caudodorsal processes, apex acute; aedeagus somewhat tubular, shaft with serrated flange from subbase to subapex laterally on right, apex symmetrical, bifurcated, gonopore dorsal at apex between bifurcations; style approx. equal to length of aedeagus, very narrow with apical third enlarged, surface rugulose, apex lanceolate; Subgenital plate with very long hair-like setae densely.
External morphology. Medium sized, slender species. General color brown. Crown light brown with dark brown marks; pronotum and mesonotum brown with pale spots; eyes gray; forewing with pale spots along veins ( Figure 21); face brown with dense light brown spots on clypeus and clypellus ( Figure 23).
Anterior margin of head angular. Crown in middle produced distally approx. 1⁄3 of entire length; area between eyes apparently narrower than width of eyes, surface without wrinkles, elevated above level of eyes; lateral margin slightly carinate, surpassing anterior margin of eyes distally, slightly convergent basally. Median length of pronotum about equal to length of crown and mesonotum ( Figure 21). Clypeus elongate, lateral margins convex; clypellus narrow, basal width less than 2/3 midlength, base flattened and narrow, apex slightly expanded (Figure 23).
Etymology. The species epithet is named for the aedeagus apex being symmetrically bifurcate.
Remarks. The new species keys to genus Armaturolidia Nielson in Nielson (2011), but the general color of brown with pale marks is different from the original diagnosis of that genus (unicolorous or with black band). This species is close to A. denticulata Nielson, 1986 in some male genitalia characters but can be separated by the lateral margin of crown extending beyond eyes, caudodorsal processes of pygofer not bilobed, style equal to aedeagus in length and apex enlarged, and aedeagus apex symmetrically bifurcate.

Discussion
The Neotropical Coelidiini fauna is very rich in general, and highly variable morphologically, e.g., in some genera such as Coelidia Germar the crown is elevated above the level of the eyes and its anterior margin is obtusely angulate while in some other genera such as Spinolidia Nielson the crown is approximately even with the eyes, slightly depressed medially with anterior margin broadly rounded and more similar to the Afrotropical and Oriental species. Two of genera treated here (Boliviela and Armaturolidia) belong to the Boliviela genus group which also comprises Collasuyusana Nielson, Ventrolidia Nielson, Carinoscapula Nielson, Paralidia Nielson, Dicodia Nielson, Gracilidia Nielson and Tinocripus Nielson. This group is diagnosed by a narrower angulate head with elevated crown. Also, most species in the group have the male pygofer with separate triangular plates basally and long ventral processes subbasally (Figs 13, 24).
The former, previously neglected, character is also found in some Youngolidiini such as Youngolidia (see Nielson 1992: figs 25, 34, 37) and Rikana (see Nielson 1992: fig 1), suggesting a close relationship between the two tribes. A similar plate is also present in the cicadellid subfamily Hylicinae, e.g., Kalasha Distant and Hemisudra Schimdt, but the relationship between these characters in these taxa is unexplored.