Abana horvathi ( Jacobi, 1905 )

Abana horvathi ( Jacobi, 1905) View in CoL

( Figs. 27–29 View FIGURE 27 View FIGURE 28 View FIGURE 29 )

Amblydisca horvathi Jacobi, 1905: 166 View in CoL [n. sp.]. Young & Lauterer (1964): 294 [lectotype designated].

Abana horvathi, Melichar (1926): 323 View in CoL [redescription]; Schmidt (1928): 51 [listed]; Young (1968): 150 [listed, out of subgenus, illustrated: Fig. 139]; Ceotto & Mejdalani (2005): 482, 491 [phylogeny, illustrated: Figs. 23 View FIGURE 23 , 32, 38]; McKamey (2007): 263 [catalogued].

Abana (Mesobana) horvathi, Melichar (1926): 322 View in CoL [key]; Metcalf (1965): 645 [catalogued].

Amblydisca pomposula Jacobi, 1905: 167 View in CoL [n. sp.]. Young & Lauterer (1964): 295 [lectotype designated]; Young (1968): 150 [syn. of Abana horvathi View in CoL ].

Abana (Mesobana) pomposula, Melichar (1926): 322 View in CoL , 323 [key, redescription]; Metcalf (1965): 646 [catalogued].

Abana regia Melichar, 1926: 324 View in CoL [n. sp.]. Schmidt (1928): 51 [comparative note]; Young (1968): 150 [syn. of Abana horvathi View in CoL ].

Abana (Mesobana) regia, Melichar (1926): 322 View in CoL [key]; Metcalf (1965): 646 [catalogued].

Abana subfasciata Melichar, 1926: 325 View in CoL [n. sp.]. Young & Soos (1964): 467 [lectotype designated]; Young (1968): 150 [syn. of Abana horvathi View in CoL ]; Wilson & Takiya (2007): 46 [lectotype listed].

Abana (Mesobana) subfasciata, Melichar (1926): 322 View in CoL [key]; Metcalf (1965): 646 [catalogued].

Abana subfasciata var. albidipennis Melichar, 1926: 325 View in CoL [n. var.]. Young & Soos (1964): 467 [lectotype designated]. Metcalf (1965): 646 [catalogued]; Young (1968): 150 [syn. of Abana horvathi View in CoL ]; Wilson & Takiya (2007): 46 [lectotype listed].

Abana puella Schmidt, 1928: 51 View in CoL [n. sp.]. Metcalf (1965): 644 [catalogued]; Young & Nast (1963): 269 [lectotype designated]; Young (1968): 150 [syn. of Abana horvathi View in CoL ].

Diagnosis. Body coloration ( Fig. 27A, B View FIGURE 27 ) mostly ivory yellow and dark brown to black; anterior portion of crown ( Fig. 27A View FIGURE 27 ) and dorsal portion of frons ( Fig. 27B View FIGURE 27 ) yellow, crown with contrasting dark maculae posteriorly and frons with contrasting dark maculae inferiorly; pronotum ( Fig. 27A View FIGURE 27 ) reddish-brown, anterior portion with black marginal stripe complete or medially interrupted, with a pair of anterolateral yellow maculae; male forewing ( Fig. 27A, B View FIGURE 27 ) black; female forewing ( Figs. 28A, B View FIGURE 28 , 29G–R View FIGURE 29 ) with one or two ivory transverse stripes: one transcommissural at basal half (absent in some specimens or with variable width) and another not transcommissural, aligned with clavus apex. Crown anterior margin ( Figs. 27A View FIGURE 27 , 28A View FIGURE 28 , 29G, J, M, P View FIGURE 29 ) subtriangular, sometimes ending truncate; anterior portion ( Fig. 27B View FIGURE 27 ), in lateral view, inflated (lectotype, Figs. 27B View FIGURE 27 , 28B View FIGURE 28 , 29A View FIGURE 29 ), forming an obtuse angle in the transition to face, or straight, not inflated ( Fig. 29D–R View FIGURE 29 ); disk with a distinct depression; M-shaped elevation bordering posterior margin present and conspicuous. Connective ( Fig. 27E View FIGURE 27 ) arms converging anteriorly; base of arms with a U-shaped dorsal rim. Style ( Fig. 27E View FIGURE 27 ) with apodeme wide and long, almost as long as apophysis lenght; inner lobe subtriangular; apical portion as long as wide; not extending to connective apex. Aedeagal shaft ( Fig. 27F–H View FIGURE 27 ) with anterodorsal projections slender and long. Dorsal connective ( Fig. 27F–H View FIGURE 27 ) sclerotized; submedian acute process conspicuous.

Redescription. Total length: males (n = 3) 16.7–17.5 mm, females (n = 5) 17.6–18.8 mm (without ovipositor), 18.8–19.2 mm (with ovipositor).

External structures. Anterior margin of crown ( Figs. 27A View FIGURE 27 , 28A View FIGURE 28 , 29A, D, G, J, M, P View FIGURE 29 ) subtriangular, apex sometimes truncate; anterior portion of crown, in lateral view, straight, not inflated ( Fig. 29E, H, K, N, Q View FIGURE 29 ), forming an acute angle with frons, or inflated (lectotype, Figs. 27B View FIGURE 27 , 28B View FIGURE 28 , 29B View FIGURE 29 ), forming an obtuse angle with face. Forewing ( Figs. 27A, B View FIGURE 27 , 28A, B View FIGURE 28 , 29A, G–R View FIGURE 29 ) with base of fourth apical cell aligned or slightly more distal (lectotype, Fig. 29A View FIGURE 29 ) than base of third cell. Hind leg with femoral setal formula 2:0:0 or 2:1:0. Other characters as in A. amazonica sp. nov.

Male coloration (based on lectotypes of A. horvathi and A. pomposula and a similar male at hand). Crown ( Figs. 27A View FIGURE 27 , 29A, D View FIGURE 29 ) with black stripe on posterior portion, interrupted medially, forming two triangular areas ( Fig. 27A View FIGURE 27 , 29D View FIGURE 29 ), or completely transversal between antennal ledges, not interrupted medially (lectotype of A. horvathi , Fig. 29A View FIGURE 29 ). Pronotum ( Figs. 27A View FIGURE 27 , 29A, D View FIGURE 29 ) anterior margin with a broad black stripe. Other characters as in description of A. amazonica sp. nov.

Female coloration (based on a female associated to a male similar to the lectotype of A. horvathi ). Body ( Fig. 28A, B View FIGURE 28 ) coloration mostly ivory and castaneous. Mesonotum ( Fig. 28A View FIGURE 28 ) dark brown with two black lateral maculae on the anterior portion, anterior to scutellar suture, reaching lateral margins. Forewing ( Fig. 28A, B View FIGURE 28 ) castaneous, becoming black at clavus apex and translucent yellow at wing apex; with one broad posterior ivory transverse stripe, restricted to corium, reaching costal margin, aligned to clavus apex. Other characters as in male.

Coloration in other females. Anterolateral yellow ivory maculae of pronotum ( Fig. 29M–R View FIGURE 29 ) well extended transversally, sometimes merged, forming a transversal stripe. Mesonotum ( Fig. 29J View FIGURE 29 ) completely black at anterior portion and dark brown posterior to scutellar suture. Forewing with anterior stripe very small, forming just a small central macula, not transcomissural ( Fig. 29S–T View FIGURE 29 ), or broad ( Fig. 29G–L View FIGURE 29 ), transcommissural, reaching costal margin, broadening ( Fig. 29G, H View FIGURE 29 ) or narrowing ( Fig. 29J, K View FIGURE 29 ) at clavus. Posterior stripe ( Fig. 29J View FIGURE 29 ) sometimes extending beyond brachial cell, almost reaching claval suture. Other specimens with anterior and posterior stripes very broad ( Fig. 29M–Q View FIGURE 29 ), merged at submedian posterior portion towards costal margin, and forming a large macula covering almost ( Fig. 29M, N View FIGURE 29 ) or entirely ( Fig. 29P, Q View FIGURE 29 ) the wing.

Male terminalia. Characters of the male terminalia ( Fig. 27C–H View FIGURE 27 ) as in A. amazonica sp. nov.

Female terminalia. Abdominal sternite VII ( Fig. 28C View FIGURE 28 ), in ventral view, with posterior margin between projections almost straight. Second valvula ( Fig. 28G View FIGURE 28 ), in lateral view, with 88 separate teeth on dorsal margin. Gonoplac ( Fig. 28I View FIGURE 28 ) apex rounded. Other characters as A. colombiana sp. nov.

Distribution. Bolivia (Cochabamba [new record], La Paz, and Santa Cruz [new record] departments), Ecuador (Morona-Santiago [new record] and Pastaza provinces), and Peru (Cusco, Huánuco [new record], Junín, Madre de Dios [new record], Pasco and San Martín departments). Before this work, A. horvathi was known from Bolivia in Coroico, Nor Yungas province, and Mapiri, Larecaja province, both in La Paz department ( Melichar 1926, Ceotto & Mejdalani 2005); from Colombia without specific locality data ( Young 1968); from Ecuador in Canelos and Rio Villano, Pastaza Province ( Schmidt 1928); and from Peru in Marcapata, Cusco region ( Jacobi 1905; Melichar 1926); Chanchamayo province, Junín Region ( Schmidt 1928); “Juanfué” [= Juanjuí, misspelled] and “Cumbasé” [= Cumbaza river] in San Martín region ( Melichar 1926); and Pasco and Junín regions ( Ceotto & Mejdalani 2005). Here, we extend the distribution of this species in Bolivia to Cercado and Ichilo provinces (Cochabamba and Santa Cruz departments respectively); in Ecuador to Orellana and Morona Santiago provinces; and in Peru to Huánuco and Madre de Dios departments ( Fig. 30 View FIGURE 30 ). The record of this species by Young (1968) from Colombia is probably based on the similar species A. colombiana sp. nov. described here, as no specimens of A. horvathi s.s. were recorded from that country.

Material examined. Type material: Lectotype of Amblydisca horvathi , ♂ (based on photographs of habitus): “Peru S\ Marcapata\ Garlepp c.”, “ Coll. A. Jacobi \ 1913-9”, “ A. Jacobi \ Typus”, “ LECTOTYPE \ Amblydisca \ horvathi \ Jacobi\ Young+Lauterer”, “ a. horvathi \ Jac.”, “Staatl. Museum für\ Tierkunde, Dresden” ( SMTD, Fig. 8A–C View FIGURE 8 ) . Lectotype of Amblydisca pomposula , ♂ (based on photographs of habitus): “Peru S\ Marcapata\ Garlepp c.”, “ Coll. A. Jacobi \ 1913-9”, “ LECTOTYPE \ Amblydisca \ pomposula \ Jacobi\ Young+Lauterer”, “Staatl. Museum für\ Tierkunde, Dresden” ( SMTD, Fig. 8D–F View FIGURE 8 ) . Lectotype of Abana puella , ♀ (based on photographs of habitus): “Ecuador\ Canelos”, “ Abana \ puella Schmidt \ Edm. Schmidt\ determ. 1928”, “ puella \ Schmidt\ 1928”, “Typus”, “ Lecto- \typus” ( MZPW, Fig. 29G–I View FIGURE 29 ) . Lectotype of Abana regia , ♀ (here designated): “Sud-America\ Cumbase”, “regia Mel \ det. Melichar ”, “ Coll. Breddin ”, “ Syn- \ typus”, “Collectio\ Dr. L. Melichar \ Moravské museum Brno”, “Invent. c. \ 3025 /Ent. \ Mor. Museum, Brno” ( MMBC, Fig. 29J–L View FIGURE 29 ) . Paralectotype of Abana sonora , ♀ (based on photographs of habitus): “Peru\ Marcapata”, “ Abana \ sonora Mel ”, “ sonora M. \ DET. L. MELICHAR”, “ TYPUS ” [paralectotype, see Wilson & Takiya 2007: 45], “hung” ( HNHM, Fig. 29S–U View FIGURE 29 ) . Lectotype of Abana subfasciata , ♀ (based on photographs of habitus): “Bolivia”, “ LECTOTYPE \ Abana \ subfasciata \ Mel. \ Young & Soós ‘63”, “ Abana \ subfasciata Mel ”, “ subfasciata M. \ DET. L. MELICHAR”, “ TYPUS ” ( HNHM, Fig. 29M–O View FIGURE 29 ) . Lectotype of Abana subfasciata var. albidipennis , ♀ (based on photographs of habitus): “Bolivia”, “ LECTOTYPE \ Abana subfas- \ ciata var. albid- \ ipennis Mel. \ Young & Soós ‘63”, “ v. albidipennis \ M.\ DET. L. MELICHAR”, “ TYPUS ” ( HNHM, Fig. 29O–R View FIGURE 29 ) . Other specimens: BOLIVIA: 1 ♂, colored as a ♀, with a detached abdomen and genitalia on a same pin ( MMBC: coll. L. Melichar). Cochabamba [new record] : 1 ♀, Cristal Mayu , 1949-x-03, L.E. Pena ( NCSU) ; 1 ♀, Osunto, 1921-22, G. Mc Creigh ( USNM) . Santa Cruz [new record]: 1 ♂, Ichilo province, Santa Cruz, Campamento Macuñucu GP-PNA - Río Saguayo-Rio Yapage , 1997-ix-06, J.C. Salvatierra ( MHNNK, based on photographs of habitus) . ECUADOR: 1 ♀, Eastern Ecuador, F.W. Gading ( USNM) . Orellana [new record]: 1 ♀, Reserva Étnica Waorani, Transect Ent, 1 km S. Onkone Gare Camp , lot#95, 00º39‘10“S, 76º26‘00“W, 220 m a.s.l., 1995-ii-08, Terre firme forest, fogging, T. L. Erwin, DNA voucher: Takiya, Rakitov & Dietrich PR 106 ( INHS) GoogleMaps ; 1 ♀, same data, except: Tiputini Biodiversity Station, nr. Yasuni National Park , lot#204, 00º37‘55“S, 76º08‘39“W, 200–250 m a.s.l., 1999-ii-07 ( USNM) GoogleMaps . Morona-Santiago [new record]: 1 ♂, Macas, Oriente Ecuador , 1921-vii, J.B. Rorer ( USNM) ; 2 ♀♀, same data . 1 ♂, Ecuador ( USNM) . PERU: 1 ♀, N. E. Peru , 1935-xi-24 ( NCSU) ; 1 ♂, Peru ( DZRJ) ; Junín: 1 ♀, Chanchamayo ( NCSU) . Cusco: 1 ♂, Marcapata ( MMBC: coll. L. Melichar) ; 4 ♂♂, Cusco, 3 rd km E Quincemil , 13º13‘03“S, 70º43‘40“W, 633 m a.s.l., 2012-viii-20 / 2012-ix-01, sweep, A.P.M Santos & D. M. Takiya ( MUSM) GoogleMaps ; 1 ♂, same data, except: 8 km W Quincemil, Pte. Manire , 13º17‘56“S, 70º47‘49“W, 773 m a.s.l., 2012-viii-31, DNA voucher Entomologia DZRJ ENT6385 ( DZRJ) GoogleMaps ; 1 ♀, same data, except: Central Hidroeléctrica de Quincemil , 13º17‘03“S, 70º46‘53“W, 757 m a.s.l., 2012-viii-26–28, Varredura, DNA voucher Entomologia DZRJ ENT6602 ( DZRJ) GoogleMaps ; 1 ♂, same data, except: 19 rd km W Quincemil, Río Araza tributary, 13º20‘10“S, 70º50‘57“W, 874 m a.s.l., 2012-viii-23–31, Malaise, R. R. Cavichioli, J.A. Rafael, A.P.M. Santos & D. M. Takiya ( DZRJ) GoogleMaps ; 1 ♂, Cusco ( DZRJ) ; 1 ♂, Cusco , Hacienda María, 900 m .a.s.l., 1952-xii, F. L. Woytknowski ( NCSU) ; 1 ♂, Yahuarmayo , 2010-ii-12, C.H. T. Townsend ( USNM) . Huánuco [new record]: 1 ♂, Tingo Maria , 1962-vii-09, W. T. Van Velzen ( USNM) ; 1 ♂, same data, except: 1944-ix-17, E.J. Hambleton ( USNM) ; 1 ♂, same data, except: 13.–23.ii.2013, V. Hula & J. Niedobová ( MMBC) ; 1 ♂, same data, except: 9 km S Tingo Maria, Pte. Cuevas , 9°21‘27“ S, 75°59‘32“ W, 620 m a.s.l., 2002-x-04, G. Solis, DNA voucher: Takiya, Rakitov & Dietrich PR 123 ( INHS) GoogleMaps . Madre de Dios [new record]: 1 ♀, Tambopata Res. Zone Tambopata Research Cntr on Rio Tambopata , 13º08.305‘S, 69º36.502“W, 622 ft, 2004-x-03–07, C. R. Bartlett ( DZRJ) . San Martín: 1 ♂, Cumbasé [= Cumbaza river], coll. Breddin ( MMBC: coll. L. Melichar) ; 1 ♂, Juanfué [= Juanjuí], coll. Breddin ( MMBC: coll. L. Melichar) .

Remarks. The concept of A. horvathi s.s. was restricted here to males with body coloration ( Fig. 27A, B View FIGURE 27 ) mostly ivory yellow and dark brown to black, the anterior part of the crown ( Fig. 27A View FIGURE 27 ) and the dorsal part of the frons ( Fig. 27B View FIGURE 27 ) yellow, the crown with contrasting dark maculae posteriorly and the frons with contrasting dark maculae inferiorly, the pronotum ( Fig. 27A View FIGURE 27 ) reddish brown, with an anterior part with a black marginal stripe complete or medially interrupted, with a pair of anterolateral yellow maculae, and the forewing ( Fig. 27A, B View FIGURE 27 ) black; and females with similar coloration, except for the forewing ( Figs. 28A, B View FIGURE 28 , 29G–R View FIGURE 29 ), which is castaneous and has one or two ivory transverse stripes: a transcommissural one on the basal half (of variable width, sometimes very small and non-transcommisural or absent) and another non-transcommissural one aligned with the clavus apex; sometimes both stripes can fuse and be extended, covering the wing area almost entirely to completely. Abana horvathi s.s. also differs from other species of the A. horvathi s.l. complex defined here by the anterior part of the crown, which is inflated in lateral view in the lectotype of A. horvathi and most specimens found close to the type locality of A. horvathi ( Figs. 27B View FIGURE 27 , 28B View FIGURE 28 , 29B, T View FIGURE 29 ), a character only shared with A. inornata sp. nov., a species that does not belong to the A. horvathi complex. We believe that the inflated anterior part of the crown may indeed be an exclusive characteristic of A. horvathi s.s. within this complex and that specimens showing the straight state of the crown potentially represents other species but this needs to be further investigated, as specimens showing both states have been found in Peru, for example.

After examining the external morphology of the type material of A. horvathi and its synonyms, it is clear that females of A. horvathi s.s. as defined here, exhibit a high degree of polymorphism. However, the results of the species delimitation analyses of A. horvathi s.l. indicate that the coloration pattern of females may still contain important taxonomic information, whereby different female morphotypes may represent distinct species ( Figs. 26A–D View FIGURE 26 , 28A, B View FIGURE 28 ). Therefore, it is likely that A. horvathi s.s. as defined here is still a group of several species, with the different morphotypes of females ( Figs. 29G–R View FIGURE 29 ) representing different species. However, this question could not be addressed in depth in the present work, as it was not possible to obtain genetic sequences for the majority of these different female morphotypes.

Young & Lauterer (1966) published a list of types of Cicadellinae species described by L. Melichar and deposited in the MMBC, in which they designated lectotypes for most species. For some reason, they omitted Abana regia Melichar from their list. Abana regia was described by Melichar (1926) based on an unspecified number of females from “ Peru: Cumbase (Coll. m.)”. In Melichar‘s collection in MMBC there is only one female that corresponds to this information ( Fig. 29J–L View FIGURE 29 ). According to Article 74 of the ICZN (1999), we designate this specimen as the lectotype for A. regia in order to stabilize the nomenclature.

Abana sonora Melichar was described by Melichar (1926) based on an unspecified number of females from “ Peru: Marcapata (Mus. Budapest); Bolivien (Coll. m.)”. Young & Lauterer (1966) designated the single female from Bolivia, which was present in Melichar’s collection at the MMBC, as a lectotype ( Fig. 3A–C View FIGURE 3 ). Consequently the female from Peru: Marcapata deposited in the HNHM ( Fig. 30S–U View FIGURE 30 ) was considered a paralectotype ( Wilson & Takiya 2007). While the paralectotype from Peru: Marcapata in the HMHM belongs to A. horvathi s.s., the lectotype of A. sonora from Bolivia in MMBC does not. Instead, the lectotype superficially resembles the genus Zaruma Melichar , but is clearly not conspecific with any of the described species of the latter. Therefore, A. sonora Melichar , stat. rev. is here removed from synonymy with A. horvathi proposed by Young (1968) and is treated as a valid species and a member of the Cicadellini incertae sedis in the absence of males.