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A New Tribe of Tropiduchidae (Hemiptera: Fulgoroidea) with Revision of the genus Buca and Description of Asymmetric Hind Leg Spinulation

Abstract A new tribe Bucini trib. nov. is erected for the genus Buca Walker, 1858 in Tropiduchidae (Elicinae), with a discussion of tribal features with respect to the New World fauna. Two new species of Buca are described from Ecuador (Buca asymmetrospinata sp. nov.) and French Guiana (Buca truncoptera sp. nov.). Keys to Buca species and to higher taxa of New World Tropiduchidae are provided. Male and female genitalia of Buca species are illustrated for the first time.

As a medium size family within the Fulgoromorpha, Tropiduchidae Stål, 1866 currently comprises 652 described species in 183 genera, or about 4.9% and 7.8%, respectively, of all Fulgoromorpha (Bourgoin 2016). Its global latitudinal profile is distributed between the north temperate and the south subtropical bioclimatic zones, but mainly in the warmer regions (Bourgoin 2016). Most species feed on shrubs and trees, and some are crop pests (Fennah 1982;Wilson et al. 1994;O'Brien 2002); their association with host plants is quite diverse concerning 21 plant orders, including some monocots (Bourgoin 2016).
The Neotropical genus Buca Walker, 1858 is currently considered as an incertae sedis taxon (Gnezdilov 2013a). The genus was erected for the single species Buca simplex Walker, 1858 from "Villa Nova," Brazil. To date, the genus is known only by the holotype of the type species. Originally placed under the subtribe Flatoides in the tribe Issites, the genus was first transferred into Issidae Spinola, 1839 (Issinae) by Muir (1930) but without tribal assignment. Shortly after that, Muir (1931) re-described the type specimen and illustrated it for the first time (Muir 1931, Fig. 10a-c therein). Buca was listed within the Issini by Metcalf (1958) before being finally removed from Issidae and provisionally placed in the family Tropiduchidae by Gnezdilov (2013a) (Fig. 1).
Here we revise Buca based on specimens from Ecuador, Bolivia, Brazil, and French Guiana representing 3 species, 2 of which are new and described below. The species differ by coloration and forewing shape and venation, but are little differentiated by male genitalia. We also discuss the features of the higher classification of New World Tropiduchidae as they pertain to Buca and provide a key to higher taxa.

Materials and Methods
The specimens examined were deposited in the following collections: Terminology of external morphological characters primarily follows Anufriev & Emeljanov (1988) and Gnezdilov et al. (2014), except for the female genitalia nomenclature, which follows Bourgoin (1993) and Gnezdilov (2002), and with the male terminalia nomenclature modified after Bourgoin (1988) and Bourgoin & Huang (1990). Nomenclature for wing venation follows Bourgoin et al. (2015) for the forewing and Anufriev & Emeljanov (1988) for the hind wing. Measurements and photographs at UDCC were taken using a digital imagery system consisting of a Nikon SMZ1500 mi-croscope, Nikon Digital Sight DS-U1 camera, and NIS Elements Imaging software (version 3.0), and at the National Museum of Wales (Cardiff, United Kingdom) using a Leica MZ8 with JVC video camera KY F7OB with the software Synoptics Automontage. Measurements were made directly from the specimens (for B. asymmetrospinata sp. nov.) or from calibrated photographs (for B. simplex, and B. truncoptera sp. nov.). Features ascribed to the genus are not repeated in the species level descriptions. Labels of name-bearing types are provided quoted verbatim using "/" to indicate a line break and "//" to indicate a new label and with supplemental information given in brackets. For other material examined, label data were rewritten to maintain consistency in pattern, beginning with the country, state, or province, followed by more specific locality information, collection date, and collector; the number and sex of specimens and the depository where the specimens are located are given in parentheses. Specimens at the UDCC (including loan) were provided 2D barcode labels, and label data were captured for online presentation (visualized at Discover Life at www.discoverlife.org, these data accessible from iDigBio at https://www.idigbio.org/) using "Arthropod Easy Data Capture" (Schuh et al. 2010;Schuh 2012; Arthropod Easy Capture 2013).

Genus Buca Walker, 1858
Type species: Buca simplex Walker, 1858 Emended diagnosis. Head (including eyes) much narrower than pro-and mesothorax. Metope (~frons) heptagonal in frontal view, slightly convex, sharply widest near antennae, abruptly narrowed above compound eyes, with distinct median carina, sublateral carinae absent. Metopoclypeal suture distinct and concave (Figs. 2C and 4A). Median carina of clypeus weak or absent. Coryphe broad and projected anterior to eyes, fastigium carinate and strongly inflected, posterior margin of coryphe concave. Coryphe, pro-, and mesonotum with median carina. Antennal scape short, pedicel elongately cylindrical with sensory plaques clustered primarily on the distal dorsal surface. Ocelli present below compound eye, anterodorsad of antennae. Pronotum short, broadest medially and abruptly narrowed within eyes, paranota narrow and elongate (nearly reaching tegulae), posterior margin concave with shallow median notch. Mesonotum large, 5-7 times as long as pronotum medially. Scutellum separated from mesonotum by transverse groove. Tegulae large, curved. Macropterous, wings broad and elongate, well exceeding abdomen, held steeply tectiform (almost parallel to body) in repose. Spinulation of hind tibiae apex asymmetrical, 1 large latero-apical spine and a row of 5 or 6 small intermediate spines. First metatarsomere with 2 latero-apical spines and row of 4-6 intermediate spines between them, each intermediate spine bears a long subapical seta. Second metatarsomere asymmetric, with a single latero-apical spine that is exceeded by a large lobe projecting from its ventral surface (instead of 2nd latero-apical spine; Fig. 6).
Abdomen arched dorsally. Phallus upcurved, phallotheca almost entirely covering aedeagus, which has a pair of elongate ventral retrose hooks in its distal third. Gonostyli with long apically enlarged and medially curved capitulum, bearing a distinct lateral lobe-shaped tooth. Pygofer in lateral view relatively narrow and tall, concave on anterior margin and convex on caudal margin. Anal tube narrow and elongate in lateral view. Paraproct about half as long as anal tube.
Etymology. The origin of the genus name was not stated in Walker (1858), and the derivation of the genus name is not evident and can only be speculated upon. We treat Buca as derived as an arbitrary combination of letters of feminine gender. 2.-Forewing with RP 3-branched and MP1 3-to 5-branched (Fig. 8A); gonostylus of male genitalia with concave hind margin (Fig. 10D); male anal tube in dorsal view slightly narrowing subapically (Fig. 10C); ventral aedeagal hooks slightly enlarged proximally (Fig. 10A) (Fig. 9C); male anal tube in dorsal view not narrowing subapically (Fig. 9B); ventral aedeagal hooks not enlarged proximally (Fig. 9A) Buca simplex Walker, 1858 (Figs. 2, 7, 9) Emended diagnosis. Color dull yellowish, mid-dorsal vitta weak on mesonotum; forewings lacking dark coloration at apices. Trailing margin of forewings roundly convex, forewing venation with 2 branches of RP, ScP and RA fused, 6 branches of MP and 2 r-m crossveins. Gonostylus with convex hind margin. Ventral aedeagal hooks long and narrow. Male anal tube with convex lateral margins.
Male terminalia. Pygofer with slightly convex caudal margin. Phallus strongly curved in lateral view, dorsolateral phallothecal lobes fused and weakly sclerotized dorsally, each lobe with narrow subapical process; ventral phallothecal lobe not reaching apical phallotheca margin. Apical aedeagal processes with rounded apices, visible above phallothecal margin. Aedeagus with pair of long (2/3 aedeagal length) and narrow ventral hooks, apically acuminate. Gonostyli with long and narrow capitulum (in dorsal view), bearing a plate-shaped lateral tooth; hind margin of gonostylus convex, caudo-dorsal angle nearly right. Anal tube almost 3 times as long as wide, with convex lateral margins (in dorsal view), flattened (in lateral view). Paraproct long, almost 2.5 times as long as anal tube.
Remarks. Walker (1858) specifies the type locality as "Villa Nova" from the [Henry Walter] Bates collection. This locality is on the Amazon River and appears to be located at 3.3580°S, 64.7052°W in Estado do Amazonas. This species was previously re-described by Muir (1931).
Male terminalia. Pygofer in lateral view relatively narrow, broadest ventrally (underside concave from ventral view), with convex cau-dal margin. Phallus recurved in lateral view, dorsolateral phallothecal lobes fused dorsally, each with a narrow subapical process; ventral phallothecal lobe narrow, rounded apically, not reaching apical phallotheca margin. Apical aedeagal processes truncate. Aedeagus with pair of long (2/3 aedeagal length) ventral hooks, apically acuminate, sometimes with marginal teeth. Gonostyli with long, apically enlarged capitulum (in dorsal view), bearing a plate-shaped lateral tooth; hind margin of gonostylus angularly concave, caudo-dorsal angle right. Anal tube narrow and elongate (in lateral view), nearly parallel-sided, distal portion usually slightly expanded (in dorsal view). Paraproct long, almost half as long as anal tube.
Remarks. Specimens from the Ecuador canopy fogging study (e.g., Erwin & Geraci 2009) are currently housed at the USNM (representatives retained at UDCC and ZIN), but it is expected that representatives will subsequently be repatriated to the Quito Catholic Zoology Museum at the Pontifical Catholic University of Ecuador (PUCE) by the USNM.
Buca truncoptera sp. nov. (Figs. 5,8B) Diagnosis. Color green yellowish with very dark dorsal vitta across coryphe, pronotum, and mesonotum; apical cells of wing with dark brown patches. Trailing margins of forewings truncately rounded with ScP+RA separated before the margin in ScP and 2 branches of RA, 9 branches of MP and 4 r-m crossveins.
Description. Color. General coloration light green yellowish. Metope with black spot medially below its upper margin. Third segment of rostrum orange apically. Coryphe, pro-, and mesonotum with wide dark median stripe, black with orange margins; stripe continuing on claval margins of forewings. First anal vein of forewings orange. Apical cells of forewing with dark brown patches. Hind wings with postcubitus and anal veins black. Abdominal tergites V-VII with red median stripe. Leg spines with black apices. Structure. Body length (with wings;  holotype) 8.5 mm, body length (without wings), 5.4 mm. Metope sharply enlarged near level of antennae in frontal view, with distinct median carina running from fastigium to level of antennae, obsolete ventrad of antennae. Lateral margins of metope sharp, laterally projected, hiding scape in frontal view. Clypeus with carina absent. Coryphe broad (Fig. 5B), anterior margin of coryphe convex, posterior margin concave. Pronotum short with median carina, disc small, paranota narrow, acutely angulate, nearly reaching tegulae. Mesonotum very large, approximately 6 times as long as pronotum medially (ratio of mesonotum length excluding scutellum to pronotum length = 6.08), with weak carinae. Forewings long (6.9 mm), extending well past end of abdomen, and wide (3.8 mm), truncately rounded (Figs. 5A and 8B). Forewing venation: forewing with crossveins absent in costal area; basal cell large; ScP+RA (branch ScP+RA forked from RP in proximal third of wing) branched into ScP, RA1, and RA2 distally; RP 3-branched with rp1-rp2 crossvein; 4 r-m crossveins; MP 9-branched (MP1 with 5 branches and MP4 with 2), 1 m-cu crossvein, CuA only very distally forked for a minute C5, 3 cup-pcu crossveins near apex of clavus. Hind tibiae with 7 spines apically.
Remarks. This species is described from a single female specimen with the right side of the head damaged.
Etymology. The specific epithet is derived from the Latin word "truncus" (maimed, cut off) combined with the Greek term "pteron" with the feminine termination "-a"; a reference to the relatively truncate forewings.

Discussion
The spinal arrangement on the apex of the hind tibiae of Buca shows a peculiar asymmetry, and the 2nd metatarsomere presents a reduction of the internal latero-apical spine and replacement of the 2nd external one with an enlargement of ventral surface. This arrangement may be unique within the Tropiduchidae, but not within the Fulgoroidea. Reductions of 1 latero-apical spine on 1st and 2nd metatarsomeres are documented for the genera Gelastissus Kirkaldy, 1906 andNubianus Gnezdilov &Bourgoin, 2009 from the Caliscelidae (Gnezdilov 2008; Gnezdilov & Bourgoin 2009). Impair reduction or even complete loss of the 2nd metatarsomere lateral spines is also reported in some tettigometrid species  but not constantly enough to make it a specific character for these species as in Buca.
Buca has a rounded ovipositor with gonoplacs missing marginal teeth which is consistent with Elicinae sensu Gnezdilov (2013b), but the structure of male genitalia and wing venation discussed below are not consistent with either of the described tribes (Elicini and Parathisciini), and so a new tribe is described to include the genus.
The general structure of male genitalia in Buca (phallus with welldeveloped phallotheca, aedeagus bearing ventral hooks, and gonostylus with plate-shaped lateral tooth) is similar to that of the Issidae. However, the genus Buca Walker and accordingly the tribe Bucini trib. nov. do not belong to the family Issidae as this genus has hind wings with simple venation including separated postcubitus (Pcu) and 1st anal vein (A1) (Fig. 7B), whereas the Issidae have much more complex venation with many transverse veins and for many species a characteristic partial fusion of postcubitus and 1st anal vein in their mid-length (Gnezdilov 2012, Figs. 2 and 3 therein). Also the general pattern of forewing venation with characteristic steps of intermediate veins between branches of median in distal half of the wing and shape and carination of metope (Buca lacks the sublateral carinae of the metope) do not allow us to place this taxon into the Issidae as it is defined by Gnezdilov (2013c) and Gnezdilov et al. (2014).
According to the pattern of hind wing shape and venation, Buca species are close to Colpopterinae of Nogodinidae and Parathisciini of Tropiduchidae (Gnezdilov 2013b). In general aspect, Buca resembles Parathisciini (Elicinae; see Gnezdilov, 2013b, Figs. 8-11 therein); however, the Parathisciini are restricted to Africa and have reticulate forewing venation (leaf-like wings). The Nogodinidae also have reticulate forewing venation (near the wing apex in Colpopterinae).
Buca does fit within the general concept of the Elicinae of the Tropiduchidae (sensu Gnezdilov 2013b). Within the New World fauna, all the Elicinae are placed in the Elicini (which include the Gaetuliina of Fennah 1978); however, the Elicini are often brachypterous, have reticulate forewing venation (including crossveins in the costal area), with wings just exceeding the abdomen and phallus with short phallotheca. Thus at the moment, we place this very peculiar genus as a separate tribe in the subfamily Elicinae Melichar (according to the rounded ovipositor) of the family Tropiduchidae, which is in need of phylogenetic revision.
Apparently these taxa of so-called "issidoid group of families" sensu Gnezdilov (2013c) are a complex of related groups that evolved recently, and perhaps rapidly, and represent different lineages of morphological evolution of male and female genitalic structures (convergence is possible as well). Phylogenetic analyses are needed to help solve the problem of the evolution of these structures and to support the classificatory solutions proposed here. However, recent investigations of the evolution of the higher Fulgoroidea using molecular data (Gnezdilov 2013c;Gnezdilov et al. 2015) show the difficulties in building a phylogeny of this complex of families.
Of the 76 specimens of Buca asymmetrospinata sp. nov. examined, 74 were from Terry Erwin's canopy fogging studies in Orellana Province, Ecuador (e.g., Lucky et al. 2002;Erwin & Geraci 2009). Few other specimens of Buca were available for study, suggesting that this is principally a canopy-dwelling species, not likely to be taken by most common collecting methods. Still few data on forest canopy planthop-per fauna are published. Thus, this is first confirmed canopy record of Elicinae. For the Neotropics, recently 1 issid species, Oronoqua ibisca Gnezdilov, Bonfils, Aberlenc & Basset, 2010 was described from mid-canopy of Panama and 3 more issid species were recorded from canopy of China and Brunei (Gnezdilov et al. 2010;Meng et al. 2013;Gnezdilov 2015). Apparently, we are just at the beginning of discovering canopy planthopper biodiversity.