Descriptions of a new Brazilian Tacora species and the female of Tacora saturata, and a key to the species of the genus (Insecta, Hemiptera, Cicadellidae, Cicadellini)

Abstract Tacora johanni, a new species from Rondônia State, North Brazil, is described and illustrated. The new species can be recognized by the male genital features, especially the subgenital plates with the basal half distinctly expanded and with outer lateral margin round, the long and slender preapical pygofer process, and the styles with apical half strongly curved. Also, the genus is recorded for the first time from Venezuela, based on specimens of Tacora saturata Young, 1977, while the female of this species (here described in detail for the first time) shows two unusual features of the genitalia. A key to males of all known Tacora species and a map showing the known distribution of the genus are provided.


Introduction
The South American genus Tacora Melichar, 1926 is known from five species (Young 1977, Takiya and Mejdalani 2002, McKamey 2007: T. dilecta (Walker, 1851) (type species), T. saturata Young, 1977, T. cavichiolii Takiya & Mejdalani, 2002, T. henriquesi Takiya & Mejdalani, 2002, and T. karipuna Takiya & Mejdalani, 2002 This genus was recorded from Brazil (Acre, Amazonas, Mato Grosso, and Rondônia states), Colombia, and Peru, being restricted to the central and western portions of the Amazon region (Takiya and Mejdalani 2002). Among the Neotropical Cicadellini, Tacora species can be recognized by their distinctive color pattern: body yellow or orange with at least two conspicuous brown stripes on the forewings, one on clavus along the claval sulcus, the other transverse, transcommissural, located in the apical portion of clavus and extending over the corium towards the costal margin. Additional black, brown, red, lilac, or yellow markings are also present and vary interspecifically. According to a morphological cladistic analysis (Cavichioli 1992) of the 13 genera of the Paromenia group (established by Young 1977), Tacora is the sister group of Dasmeusa Melichar, 1926. The reader is referred to Takiya and Mejdalani (2002) for detailed comments on the taxonomy, phylogeny, and distribution of Tacora.
In the present paper, we describe a new Tacora species from Rondônia State (North Brazil). We also provide the first detailed description and illustrations of the female of T. saturata, including two unusual features of the genitalia. The genus is recorded from Venezuela (Amazonas and Bolivar states) for the first time (based on T. saturata). A new key to males of Tacora species and a map showing the known distribution of the genus, both modified from Takiya and Mejdalani (2002), are provided.

Material and methods
Techniques for preparation of male and female genital structures follow Oman (1949) and Mejdalani (1998), respectively. The dissected genital parts are stored in microvials with glycerin and attached below the specimens, as suggested by Young and Beirne (1958). The descriptive terminology adopted herein follows mainly Young (1977), except for the facial areas of the head (Hamilton 1981, Mejdalani 1998 and the female genitalia (Nielson 1965, Hill 1970. Mejdalani (1998) provided a detailed justification for the use of Hamilton's terminology for the head of the Cicadellinae and other leafhoppers. Use of the term gonoplac (= third ovipositor valvula) and the names for the processes of the dorsal and ventral sculptured areas of the first ovipositor valvula follow Mejdalani (1998).
Label data are given inside quotation marks with a reversed virgule (\) separating lines on the labels and a semicolon separating labels of a specimen. The specimens studied belong to the following institutions: American Museum of Natural History (AMNH; New York), Museu Nacional, Universidade Federal do Rio de Janeiro (MNRJ; Rio de Janeiro), and Museo del Instituto de Zoología Agrícola, Universidad Central de Venezuela (MIZA; Maracay).
The photographs of the body in dorsal view and of the general lateral views of the ovipositor valvulae were prepared with the software Automontage (Synoptics Inc., Frederick, Maryland) using a digital camera attached to a stereomicroscope. The photographs of the details of the ovipositor valvulae were taken with a digital camera at-tached to an optical microscope. Digital images of four of the five previously known Tacora species (body in dorsal view) are available in the internet site "Sharpshooter Leafhoppers of the World" (Wilson et al. 2009). These images were useful for the comparisons and identifications carried out in the present study. The orientation of the illustrations of male and female genital structures on the plates are in accordance with the monograph of Young (1977).

Genus Tacora Melichar, 1926
Tacora johanni sp. n. urn:lsid:zoobank.org:act:4E10B39B-9F6B-42EC-B9A7-21B25D2AA4FE http://species-id.net/wiki/Tacora_johanni Figs 1a, 2, 3 Description of the male holotype. Length 15 mm from apex of head to apex of forewings at rest. Head (Fig. 2a, b) strongly produced anteriorly; median length of crown approximately 4/5 interocular width and 1/2 transocular width; anterior margin broadly rounded in dorsal view; without carina at transition from crown to face; ocelli located behind imaginary line between anterior eye angles, each ocellus slightly closer to median line of crown than to adjacent anterior eye angle; crown with concavity between eye and ocellus, without median fovea and without sculpturing or setae; frontogenal sutures extending onto crown and attaining ocelli; antennal ledges, in dorsal view, slightly protuberant, in lateral view with anterior margin broad and rounded; frons convex, muscle impressions distinct; epistomal suture obscure; clypeus not produced, its contour continuing profile of lower portion of frons. Thorax (Fig. 2a,b) with pronotal width distinctly greater than transocular width; lateral pronotal margins convergent anteriorly; posterior margin slightly concave; dorsopleural carinae complete, rectilinear, distinctly declivous anteriorly; pronotal disk without sculpturing or pubescence; mesonotum with scutellum not swollen. Forewings ( Fig. 2c) with membrane including all of apical cells except short basal portions of second, third, and fourth; veins mostly indistinct except at wing apical portion; with four apical cells, base of fourth more proximal than base of third (not shown in Fig. 2c); without anteapical plexus of veins. Hindlegs with femoral setal formula 2:1:1; length of first tarsomere greater than combined length of two more distal tarsomeres; plantar surface with two parallel rows of small setae.
Color (Fig. 1a) of anterior dorsum orange; pronotum with transverse brown stripe on posterior margin, this stripe distinctly broader medially; mesonotum with pair of large yellow maculae laterobasally. Forewings ( Fig. 2c) with corium mostly translucent yellow, costal margin dark brown; clavus mostly yellow with two longitudinal brown stripes, both extending from basal portion to about middle area, one along claval sul-cus (broadened posteriorly) and another along inner claval margin; posterior portion of clavus with transverse, oblique brown stripe extending over corium and reaching costal margin, this stripe distinctly narrowed towards costal margin; with red area behind transverse stripe covering apex of clavus, apexes of brachial and inner discal cells and large portion of corium adjacent to inner apical cell, this red area strongly constricted posteriorly and then forming stripe (evanescent towards costal margin) across bases of apical cells; corium region between red area and costal margin mostly depigmented. Face and venter orange-yellow with brown irregular areas on legs. Genitalia with pygofer ( Fig. 3a), in lateral view, strongly produced posteriorly; posterior margin narrowly rounded; distal third of dorsal margin with strong spiniform process, apex of process located slightly before pygofer apex, process (Fig. 3b), in dorsal view, directed inward, not attaining median body line; macrosetae distributed mostly on distal third of disk, some smaller macrosetae extending anteriorly along ventral margin. Subgenital plates (Fig. 3c), in ventral view, triangular but with basal half broad, expanded, with lateral margin round; distal half extremely narrow; plates not fused to each other basally; in lateral view, plates distinctly shorter than pygofer (Fig. 3a), extending posteriorly approximately as far as 2/3 of pygofer disk length; plates with uniseriate macrosetae (very large on basal half, very small on distal half ), microsetae also present. Connective (Fig. 3d), in dorsal view, broadly T-shaped; stalk with strong median keel, the latter bifurcated basally and extending for short distance over arms. Styles (Fig. 3d), in dorsal view, extending almost as far poste- riorly as connective; strongly curved inward; without preapical lobe; apex slightly expanded, obliquely truncate, more sclerotized than remainder of apophysis; style median portion with a few setae on outer margin. Aedeagus (Fig. 3e, f ), in lateral view, narrowest in basal third, gradually broader in distal two-thirds; shaft with two pairs of longitudinal flanges, one along lateral portion, another along dorsal margin; apical portion of shaft (Fig. 3f ), in ventral view, with these flanges forming pair of projections directed medially; these projections covered by many tiny tegumentary processes; gonoduct distinctly visible in broadened portion of shaft (Fig. 3e), directed ventrally, gonopore located preapically.
Etymology. The new species is described in honor of the late Prof. Johann Becker (Museu Nacional, Universidade Federal do Rio de Janeiro) in recognition of his contri- bution to the development of Brazilian entomology. He was also one of the collectors of the holotype.
The male genitalia of the new species are similar to those of T. dilecta and T. cavichiolii. The presence of sculpturing (pegs) on the pygofer process of the male holotype of T. dilecta was mentioned and illustrated by Young (1977: 317 and Fig. 257p) as a diagnostic feature of this species. These pegs are not present in T. johanni. Other features that distinguish the new species (known only from the male) from T. dilecta include (1) the red color of the claval apex behind the transverse dark stripe (orange to yellow in T. dilecta) and (2) the rectilinear distal portion of the pygofer process in dorsal view (distinctly curved in T. dilecta). The above-mentioned features of the subgenital plates and styles of T. johanni are distinct from those of T. dilecta and T. cavichiolii. In addition, the pygofer process in T. cavichiolii is short and robust (Takiya and Mejdalani 2002: 239: Fig. 17), whereas it is long and slender in T. johanni. Young, 1977 http://species-id.net/wiki/Tacora_saturata Figs 1b, 4, 5 Tacora saturata Young, 1977: 316, figure 258 (crown, pronotum, and male genitalia).

Tacora saturata
Description of the female. Length 13 mm from apex of head to apex of forewings at rest [male with about same length]; median length of crown approximately 9/10 interocular width and 1/2 transocular width. Head and thorax (Fig. 4a) much as in the above-described male holotype of T. johanni sp. n. Forewings extending well beyond apex of ovipositor.
Color (Fig. 1b) of anterior dorsum orange; pronotum with transverse, submarginal brown stripe on posterior portion, this stripe broader medially; mesonotum with pair of large brown maculae laterobasally. Forewings (Fig. 4b) with corium mostly translucent orange, costal margin dark brown; in rest position with transcommissural, brown W-shaped figure in basal half of clavi with anterior angles enclosing large, bright yellow maculae that extend to wing bases; large, bright yellow rounded transcommissural macula located just behind W-shaped figure, followed by transverse, oblique brown stripe extending over corium and reaching costal margin; apical portion of clavus red, with or without pair of small orange spots adjacent to claval sulcus just behind trans-verse brown stripe; distal half of corium with red stripe in inner anteapical cell (along outer margin of inner apical cell) and then descending across bases of apical cells.
Genitalia with abdominal sternite VII (Fig. 4c) broad, lateral margins convergent posteriorly, posterior margin broadly convex. Internal sternite VIII without distinct, well-defined sclerotized areas. Pygofer (Fig. 4d), in lateral view, well produced posteriorly, strongly narrowed towards apex; posterior margin very narrow, subacute; macrosetae located mostly on distal third, a few extending anteriorly along ventral margin. Valvifers I (Fig. 5a), in lateral view, of quadrate form, except for distinct lobe on posteroventral portion. Valvulae I, in ventral view, distinctly expanded basally; in lateral view (Fig. 5a), valvulae with broad basal lobe, blade distinctly expanded in distal half in comparison with basal half; dorsal sculptured area (mostly scale-like processes, except for linear processes basally, Fig. 5b, c) extending from basal portion of blade to apex, ventral sculptured area (scale-like processes, Fig. 5d) restricted to apical portion of blade; basal portion of blade with group of distinct setae located below ramus; apex of blade forming distinct dentiform projection (Fig. 5a); ventral interlocking device distinct, elongate, restricted to basal half of blade, located along ventral blade margin but with distal portion directed dorsally. Valvulae II (Fig. 5e), in lateral view, with dorsal margin regularly convex beyond basal curvature; without preapical prominence; apex obtuse; ventral outline of apical portion slightly concave; about 63 teeth (Fig. 5e, f, g,  h), mostly subtriangular or subrectangular, distributed on dorsal portion of blade, with clear space between them; denticles (Fig. 5g) on posterior portions of teeth and on inferior half of apical portion; blade with numerous curved ducts (Fig. 5f, g) extending to teeth or terminating below the latter, ducts also extending towards apex (most teeth receive a single duct, others two or none). Gonoplacs, in lateral view, extending posteriorly slightly beyond pygofer apex; basal half with ventral margin convex and dorsal margin concave, the latter abruptly expanded towards distal half; ventral margin of distal half concave; apex of blade rounded; apical portion with few small setae and tiny tegumentary processes, the latter extending anteriorly along ventral margin.
Remarks. Young (1977) provided a detailed description of the male of T. saturata. Our identification is based on his description. The male genitalia and color pattern (male and female) of our Venezuelan specimens agree with the features and illustrations given by him. He recorded T. saturata from Colombia and Brazil. This species is the only one in the genus in which the transverse pronotal stripe is submarginal (Figs  1b, 4a).

Concluding remarks
Six species of Tacora are now known. The most relevant features for the recognition of males from each of these species are given in our key, which provides an adequate comparison among the previously described species and T. johanni sp. n. These features are from the aedeagus and its flanges, pygofer and its processes, subgenital plates and styles, as well as from the color pattern. Females of Tacora are of difficult identification when they can not be associated to males from the same collecting series, due perhaps to our still incomplete comparative knowledge of the female genitalia morphology and color pattern intraspecific variation. In addition to our description of T. saturata, the female genitalia of T. dilecta, T. henriquesi, and T. cavichiolii were described in detail (the first by Young 1977 and the second and third by Takiya and Mejdalani 2002). Our study of the T. saturata female revealed peculiar, previously undescribed genital features of potential taxonomic interest that deserve further comparative investigation, i.e. the lobe on the posteroventral portion of valvifer I and the dentiform projection on the apex of valvula I (Fig. 5a). Unfortunately, these parts of the female genitalia were not described for the other species.
The previously known distribution of Tacora (Brazil, Colombia, and Peru) is now extended to Venezuela (Fig. 6). Takiya and Mejdalani (2002) suggested that a vicariant event isolated the Tacora ancestor in Western Amazonia in an area that included the Napo and Inambari endemism centers, and possibly also the Imerí center (where no records of the genus were known at that time). Interestingly, the present results appear to support the proposal of Takiya and Mejdalani (2002), as we found that T. saturata is distributed in the Imerí center (Fig. 6).