Bubalopa furcata ( Fairmaire, 1846 )

Flórez-V, Camilo & Evangelista, Olivia, 2021, A revision of the treehopper genus Bubalopa Stål illuminates the systematics of Hyphinoini (Hemiptera: Auchenorrhyncha: Membracidae), Zootaxa 5052 (4), pp. 529-551 : 537-545

Bubalopa furcata ( Fairmaire, 1846)

( Figs. 3–8 )

Hemiptycha furcata Fairmaire, 1846: 314 .

Bubalopa furcata Stål, 1869c: 255 ; Funkhouser, 1927: 117; Metcalf and Wade, 1965: 701; McKamey, 1998: 476.

Diagnosis: Overall color ochraceous yellow, punctation reddish black, antero-dorsal surface of suprahumeral horns dark brown with slight red tinge, tibia black, spotted yellow medially, anterior and posterior margins orange. Suprahumeral horn orientation: 35–50º (frontal view), 25–40º (dorsal view); dorsal outline slightly arched, descending in almost straight line from above humeral angles to apex of posterior process. Males: parameres Tshaped, subgenital plate abruptly narrow apically.

Description: ADULTS. Color. Head and pronotum ochraceous yellow with reddish dark brown punctation. Eyes grey, ocelli yellow, surrounding cuticle forming a brown ring. Antero-dorsal surface of suprahumeral horns dark brown with slight red tinge. Forewings slightly to entirely opaque, amber to reddish dark brown with black punctation. Thoracic sternites, coxa, trochanter, femora and tarsomeres ochraceous yellow; dorsal surface of tibiae black spotted yellow, anterior and posterior margins orange, ventral surface ochraceous yellow. Head: Triangular, wider than long; suprantennal ledges discreetly sinuous, apex of frontoclypeus acute. Thorax: Suprahumeral horns 2–2.5x as long as wide basally, orientation: 35–50º (frontal), 25–40º (dorsal); dorsal carina of suprahumeral horns less sharp than others; dorsal contour weakly arched, highest above humeral angles, descending in almost straight line to apex of posterior process; semicircular impression at each side of median carina, approximately at middorsum. Abdomen: Terga III–V with a pair of oval dorsal pits (scars) ( Figs. 7A, 7D , 8A, 8F ). Male: Apodeme reaching posterior margin of sternite III. Lateral plate fused with pygofer. Subgenital plates wider at basal half, with slight dorsal hump, abruptly narrow in lateral view. Parameres broad, T-shaped, bifurcate distally into fingerlike lobes, weakly curved and round apically. Aedeagus U-shaped, several rows of minute teeth along entire length of anterior surface of posterior arm. Female: Gonoplac blade-shaped, covered in setae, distinctly more sclerotized along posteroventral margin. First valvulae blade-shaped, broad throughout; dorsal margin, from mid-length to apex, finely sculpted in short diagonal lines, gradually turning into well-marked striations, extended transversally across triangular and acute apex. Second valvulae blade-shaped, slightly broader at apex, two prominent teeth on apical third, quadrangular teeth on dorsal contour apically, located after apical prominent tooth, apical margin dorsally serrated, ramus extended up to apical 1/5.

4 TH /5 TH INSTAR NYMPH. Color: Mostly green in live specimens, brown in dry preserved specimens. Surface: vertex, pronotum and abdomen covered in minute, stout and subcylindrical setae, sparser on metathorax and mesonotum, abdominal segment I+II glabrous; elongate and slender setae covering ventral surface of body and legs, more densely distributed on ventral side of head. Tuberculate chalazae present on post-ocular lobes, suprahumeral buds, entire prothorax (except for pre-metopidium), anteroventral margin at base of forewing pads, mesonotal and abdominal scoli, ventrolateral margins of abdominal terga ( Fig. 6E , ach), and anterodorsal and posterodorsal margin of tibia; chalazae and their setal portion varying in size, larger and prominent along the anterior surface of thoracic and abdominal scoli. Head: subtriangular; vertex bearing 10 scoli, arranged into 5 bilaterally symmetric pairs: 2 slender scoli on upper margin, equidistant from eyes and coronal suture, curved upwards ( Fig. 6D , hs1); 2 stout scoli, each at external anterior margin, juxtaposed to eyes ( Fig. 6D , hs3); 2 slender scoli (longest in vertex), each near external anterior margin, adjacent to latter pair, distinctly curved downwards ( Fig. 6D , hs2); 2 smaller scoli, each adjacent to eyes, right below the latter pair ( Fig. 6D , hs4); 2 small and stout scoli at inferior margin, right below eyes, slightly curved posteriorly ( Fig. 6D , hs5). Thorax: prothorax well developed, tall and laterally flattened antero-dorsal projection, bearing 2 robust, conical scoli, curved forward apically (=postmetopidium scoli sensu McKamey et al. 2015) ( Fig. 6F , pos); pair of minute scoli on metopidium (=premetopidium scoli sensu McKamey et al. 2015) ( Fig. 6F , prs); suprahumeral buds flap-like, folded posteriorly onto sides of prothorax ( Fig. 6C , shb); posterior process triangular, reaching anterior margin of metanotum; meso- and metanotum with 1 pair of small scoli each ( Fig. 6F , mss, mts); ventral margin of forewing pads slightly emarginate anteriorly, sheltering metathoracic leg in repose, venation barely perceptible (tinged bright green in live specimens), pads reaching fourth abdominal segment; hindwing pads subtriangular, as long as forewing pads, anal margin mostly exposed. Abdomen: Abdominal terga III–V with pair of greatly enlarged dorsal scoli, each pair slightly arched in frontal and dorsal view in U-shaped outline ( Fig. 6C, 6F, 6I , asIII-asV), terga VI–VIII with pair of small spinelike scoli ( Fig. 6F , sps). Segment IX tubular, elongate (more than half the length of rest of abdomen), bearing one pair of apical spine-like scoli directed posteriorly ( Fig. 6F , asIX) and slightly diagonally in V-shape.

Measurements (in mm): Female (n=3, including lectotype)/male (n=1): Body length: 13.48/11.62; forewing length: 11.94/10.34; pronotal length: 11.91/9.99; pronotum height: 3.41/2.64; pronotal width: 4.50/3.46; head width: 4.08/3.28; vertex width: 2.81/2.19; vertex length: 2.17/1.72.

Lectotype designation for Hemiptycha furcata Fairmaire, 1846: 314 . In the original description of Hemiptycha furcata, Fairmaire (1846) specified Bogota as the type-locality, and the Signoret Collection as the repository of the examined material. When erecting Bubalopa and redescribing Bubalopa furcata, Stål (1869) also listed Bogota as the type-locality and mentioned that the specimen he examined had a mutilated head. The single specimen currently housed at the Naturhistorisches Museum Wien closely matches the description of Fairmaire and Stål both in terms of morphological features and its state of preservation. However, while the determination label of this specimen correctly states ‘ furcata ’, the locality label reads ‘ Bolivien’, contradicting the records from both of these authors.

The Signoret collection was sold to the Naturhistorisches Museum Wien, Vienna, Austria (NMW) in 1890 after the death of Victor Signoret in 1889. After its acquisition, Signoret’s specimens were re-labelled by Anton Handlirsch ( Rabitsch 2000). Each specimen received at least two labels, which were highly characteristic in terms of appearance and style. One label features a handwritten locality (usually one word, e.g., ‘ Bolivien ’) and a printed collection source (i.e., ‘Coll. Signoret’). The second label includes the species identification, containing a handwritten epithet (i.e. ‘ furcata ’), and the printed name of the identifier/author (i.e., ‘det. Signoret’) ( Figure 4C ). Fairmaire and Stål’s manuscripts, published between 1846 and 1869, indicate that they examined the type before it was transferred to NMW. The authors, therefore, correctly reported the locality from Signoret’s original labels before it was incorrectly transcribed by Handlirsch. Transcription errors, although rare, have been observed in other hemipteran specimens that Handlirsch curated (e.g., Reduviidae : Phymatinae; Rabitsch 2000). Despite this label inconsistency, there is strong indication that this exemplar is the syntype of Hemiptycha furcata , from Bogota, not Bolivia, which was examined first-hand by Stål. The term ‘Bogota’ was often used in the 19 th century to refer to multiple sites in Colombia, and therefore will not be included in the distribution of this species.

Distribution: COLOMBIA: Antioquia: Medellín (Corregimiento de Santa Elena, 2500 masl); Puerto Berrío (Vereda Cristalina, 400 masl) ( Fig. 11 ).

Examined Material: Lectoype female (here designated): “ Bolivien \ Coll. Signoret.”, “ furcata \ det. Signoret.” [original type locality: ‘ Bogota’, incorrectly transcribed by A. Handlirsch] ( NMW). COLOMBIA: Antioquia: Medellín: “ COLOMBIA. Antioquia, Medellín, Corregimiento Santa Elena, Reserva SMA San Pedro , 6.194048ºN, 75.499603ºW, 2600 msnm, Manual, en Senna pistaciifolia (Fabaceae) , Jun. 2019, leg. Semillero de Entomología 2019-I (C. Flórez-V), CBUCES-F 9036 (1 male, 1 female, CBUCES); “ COLOMBIA. Antioquia, Medellín, Corregimiento Santa Elena, Reserva SMA San Pedro , 6.194048°N, 75.499603°W, 2600 msnm, Manual, en Senna pistaciifolia (Fabaceae) , Jun. 2020, leg. C. Flórez-V, A. Ospina, CBUCES-F 8002 ”, “ CBUCES-F 8003 ”, “ CBUCES-F 8023 ” (1 male, 2 females, CBUCES); “ COLOMBIA. Antioquia, Medellín, Corregimiento Santa Elena, Reserva SMA San Pedro , 6.194048°N, 75.499603°W, 2600 msnm, Manual, en Senna pistaciifolia (Fabaceae) , Sep. 5/2020, leg. C. Flórez-V, A. Ospina, CBUCES-F 8004 ” (1 nymph, CBUCES). Puerto Berrío: “ COant. Puerto Berrío Vda. / Cristalina R.U.N.A. 400–500m / Manual 22-sep-2005 / A. Acosta / CEUA 66707 ” (1 female, CEUA) .

Biology: Adults and immatures were observed feeding on branches of Senna pistaciifolia (Kunth) H.S. Irwin & Barneby (Fabaceae) .Adults and nymphs are highly cryptic. The color, shape, and ultrastructure of scoli and chalazae help nymphs blend into the apical indumenta of the host plant. Nymphs closely resemble host plant stipules, where they usually feed. Adults were consistently observed on ligneous plant tissue feeding on subapical branches. Both nymphs and adults are solitary, not associated with ants or other hymenopterans, and were sparsely distributed in the surveyed transects. For example, only five adults and five nymphs were collected after thorough inspection of 50 host plants in that area, which suggests low population density. Other treehopper species were found to co-occur in the same host plant individual: Bordoniana nigricosta (Goding, 1926) ( Smiliinae : Acutalini), an unidentified species of Membracinae , and several aggregations of Aetalion cf. nigromarginatum (Aetalionidae) including multiple adults and nymphs.

Remarks: Representatives of B. furcata are easily distinguished from other Bubalopa by their overall color, ochre yellow and reddish brown punctation, acute frontoclypeus, slender suprahumeral horns, and a nearly diagonal dorsal outline. The supernumerary crossveins on the forewings vary intraspecifically and can be as numerous as in B. obscuricornis but still fewer than in B. iguaque . In males, suprahumeral horns are more horizontal (35º–40º, frontal view; n=2) than in females. A larger sampling of females indicates that there is considerable variation in horn orientation (35º–50º, frontal view; n=5), although four of the five examined females showed an angle from 40º–50º. While those differences are evident in frontal view, horns appear similar in lateral and dorsal view in both sexes for all exemplars. As observed in other species in the genus, B. furcata bears prominent oval pits on the dorsolateral area of abdominal terga III–V, which are thought to be scars resulting from nymphal scoli ( Lencioni-Neto and Sakakibara 2013). The small semicircular impressions on the pronotal posterior process in adults, which appear to be topologically correlated with scoli in the nymphs, are also visible in A. robustus . These traits are not observed in other species of Bubalopa (see remarks after the redescription of the genus).

Some structures of the male genitalia of B. furcata are highly characteristic, including the subgenital plate, with a large dorsolateral indentation, and the robust and T-shaped paramere. These two features are not observed in any other Membracidae . The internal lateral margin of abdominal pleurites, and the lateral and anterior margin of abdominal sternites are somewhat invaginated into the abdomen. The spiracles are located in the invaginations of the pleurite. The second valvulae of B. furcata ( Figs. 8I–J ) differs from the figure presented in Deitz (1975) for the same species (Fig. 24O, pp. 85). We believe that Deitz (1975) illustrated a specimen of Eualthe , which had been previously misidentified by Funkhouser as Bubalopa .

This is the first species of Bubalopa for which nymphal morphology is described. The relevance of these characters to tribe and genus classification are discussed elsewhere in this paper. Nymphs were collected on the same date and same host plants as adults.

The specimen from Puerto Berrío was incorrectly referred to in Flórez-V et al. (2015) as a representative of Sundarion . The authors commented that this exemplar did not exhibit femoral cucullate setae as in Sundarion but listed other characters as deep pronotal punctations or supernumerary forewing crossveins, which are characteristic of Bubalopa .

Figures

FIGURE 3. Live specimens of Bubalopa furcata. Nymphs: A) habitus of an early instar. B–D). Fourth/fifth instar nymph: B) habitus in lateral view, C) 4/5th instar nymph resting near the stipule of its host plant (Senna pistaciifolia), D) habitus in lateral view. Adult: E) frontal habitus, F) lateral habitus.

FIGURE 4. Lectotype of Hemiptycha furcata Fairmaire, 1846. A) Frontal view. B) Lateral view. C) Labels. Note: the provenance was incorrectly transcribed by Anton Handlirsch. The original locality is ‘Bogota’ (Fairmaire 1846, Stål 1869). See remarks in the text. Female specimen of Bubalopa furcata. D) Wings. E) Frontal view. F) Dorsal view. G) Lateral view. H) Frontolateral view.

FIGURE 5. Bubalopa furcata (Fairmaire, 1846). Male: (A–E) A) Frontal view. B) Dorsal view, pronotal scars indicated by a rectangle. D) Lateral view. E) Frontolateral view. Female: (E–H). E) Frontal view. F) Dorsal view. G) Lateral view. H) Frontolateral view.

FIGURE 6. Bubalopa furcata (Fairmaire, 1846). A) male abdomen, oval pits indicated by a rectangle. B) Female, ventral view. Fourth-fifth instar nymph: C) Frontal view. D) Head, anteroventral view. E) Head, ventral view. F) Lateral view, left side. F) Lateral view, right side. G) Frontolateral view. H) Dorsal view. ach: lateral abdominal chalazae; as: abdominal scoli; hs: head scoli; mss: mesothoracic scoli; mts: metathoracic scoli; pms: premetopidium scoli; pos: postmetopidium scoli; shb: suprahumeral horns bud; sps: small spinelike scoli.

FIGURE 7. Male of Bubalopa furcata (Fairmaire, 1846). Abdomen:A) Dorsolateral view, oval pits indicated by a rectangle, B) ventral view, C) close-up of abdominal spiracles, D) close-up of scars on abdominal terga III and IV, E) lateral view. F) Pygofer in lateral view. G) Subgenital plate in ventral and dorsal view.Aedeagus: H) Lateral view, I) caudal view, J) anterolateral view. K) Subgenital plate in lateral view. Parameres: L) Lateral view, M) dorsal view, N) ventral view.

FIGURE 8. Female of Bubalopa furcata (Fairmaire, 1846). Abdomen: A) Dorsal view, B) ventral view, C) lateral view, oval pits indicated by a rectangle, D) Pygofer. E) Close-up of spiracle. F) Close-up of scars on terga III–V. First valvulae: G) lateral view, H) close-up apex. Second valvulae: I) lateral view, J) close-up apex.

FIGURE 11. Distribution map of Bubalopa furcata, B. iguaque sp. nov., and B. obscuricornis.