Hyphinoini, Haupt, 1929

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 : 531-532

Membracidae :

Darninae :

Hyphinoini

Amended diagnosis of the tribe Hyphinoini . ADULTS. Forewings entirely exposed or slightly concealed by pronotum; membrane densely covered in erect setae ( Eualthe , Hyphinoe and Hanstruempelia ), partially glabrous on posterior half ( Tomogonia ) or apical third ( Bubalopa ), row of erect setae alongside each vein; forewing vein R initially divided in R 1+2+3 and R 4+5, one s crossvein, one r-m crossvein and two m-cu crossveins, s crossvein distad of r-m. Femora lacking cucullate setae, metathoracic coxa and trochanter unarmed; metathoracic tibia bearing three complete rows of cucullate setae, row III single basally, double or triple on apical third, plantar surface of first metathoracic tarsomere with cucullate setae (absent in Bubalopa and Hanstruempelia bitumina Sakakibara, 2004 ). Abdominal segments III–VIII (each segment or all) often showing middorsal spots, fenestrae, scars or tubercles (except absent in Hanstruempelia ).

4 TH /5 TH INSTAR NYMPH. Tuberculate chalazae widely distributed, denser on scoli; scoli with or without setal portion. Vertex with 8–10 scoli, arranged into bilaterally symmetrical pairs. Prothorax with pair of premetopidial scoli, and two conical and robust postmetopidial scoli; meso- and metathorax with pair of scoli each. Abdominal terga III–IX (each tergum or all) bearing pair of scoli, ventrolateral lamellae absent. Abdominal segment IX dorsal length almost equal to combined lengths of segments IV–VIII.

Remarks. Prior to this work, the defining features of Hyphinoini were the densely pubescent forewing membrane and the presence of cucullate setae on the ventral surface of the first metathoracic tarsomere ( Deitz 1975). These were also inferred as synapomorphies supporting the monophyly of the tribe, although the sampling of that study ( Dietrich et al. 2001) only included representatives of Hyphinoe . In Bubalopa , the forewing membrane is partially glabrous, and cucullate setae are absent on the plantar surface of the first metathoracic tarsomere. We believe that the erect forewing setae still represent a key trait in diagnosing Hyphinoini because forewings are noticeably pubescent in all tribal constituents. In instances where the membrane is partially glabrous, veins exhibit a line of erect setae along their entire length. This amended diagnosis, which is intended to complement the revision of Deitz (1975), provides additional detail into the variation of wing pubescence in different hyphinoine groups, as well as cases in which key diagnostic traits are absent.

Other important exceptions include Tomogonia vittatipennis ( Fairmaire, 1846) and T. camposiana Goding, 1920 , whose forewings lack setae in the posterior half, except along the veins. Cucullate setae on the plantar surface of the first metathoracic tarsomere are absent in Hanstruempelia bitumina , although 1–5 cucullate setae may be present at the distal margin ( Sakakibara 2004). Members of the genus Hanstruempelia also lack paired fenestrae, spots or tubercles on abdominal terga III–VIII, which are observed in other species of Hyphinoini .

We inferred the placement of Bubalopa within Hyphinoini based on the wing setae and venation, as well as unique features of immature stages. Nymphal traits are newly reported for Bubalopa and Hyphinoe . The arrangement of scoli and chalazae in the head, thorax, and abdomen show an unambiguous topological correspondence between these two groups. Immature features show great potential to inform the classification of darnine subfamilies and tribes, especially at the early stages, when the size and appearance of scoli make comparisons more straightforward. Scoli decrease in size throughout development in hyphinoine nymphs, with a different pattern of reduction in each genus. For instance, most scoli decrease equally and in proportion to body size in H. obliqua ( Fig. 2F ). Contrastingly, in B. furcata , the mesothoracic and abdominal scoli on segments VI–VIII decrease significantly in size compared to other scoli in later nymphal stages ( Fig. 6F ). Nymphal traits which are informative for tribal-level diagnosis include a similar arrangement of body scoli ( Figs. 1A , 3A ): 4–5 bilaterally symmetrical pairs on the head, one pair of premetopidial scoli, one pair of postmetopidial scoli, one pair on mesonotum and metanotum, and one pair on abdominal segments. Moreover, the differentiation of scoli in late instar nymphs can prove useful for recognizing genera. Hyphinoini nymphs can be distinguished from other similar immatures in unrelated tribes (e.g., Ceresini , which also have a laterally compressed body) in having smaller, stalked or tuberculate chalazae on scoli, a greater number of scoli on the head, and smaller scoli on the metanotum as compared to other thoracic and abdominal segments in late instar nymphs.

Our decision to retain Bubalopa in Hyphinoini also takes into consideration the fact that these species do not fit into the current concept of any other darnine tribe. Representatives of Bubalopa do not exhibit the femoral cucullate setae diagnostic of Darnini and Procyrtini; their coxa and trochanter are unarmed, lacking the apposed and spinelike processes observed in Cymbomorphini and Procyrtini; and their metathoracic tibia bears three rows of cucullate setae, unlike members of Hemikypthini.A phylogenetic revision of Darninae is warranted to evaluate potential tribal synapomorphies and to determine when multiple acquisitions or secondary losses may have occurred, especially concerning forewing characters and leg chaetotaxy.

Presented here is a provisional key to aid the identification of Hyphinoini genera. A comprehensive revision of Hyphinoe Stål is warranted to reassess the placement of species which strongly diverge in their pronotal morphology. The distinction between Hyphinoe and Hanstruempelia Sakakibara also needs to be further clarified. We based our concept of genera on their corresponding type species, and we believe a reappraisal of Hyphinoini genera will result in the description of new taxa.

Figures

FIGURE 1. Live specimens of Hyphinoe obliqua (Walker, 1858). Nymphs: A) and B) 1/3rd instar, C) 4/5th instar, D) Last instar exuvia.

FIGURE 2. Hyphinoe obliqua (Walker, 1858). Adult: A) Frontal view, B) lateral view, C) dorsal view. Last instar exuvia: D) dorsal view, E) frontal view, F) lateral view, G) close-up frontal view, H) frontolateral view. as: abdominal scoli; hs: head scoli; mss: mesothoracic scoli; mts: metathoracic scoli; pms: premetopidium scoli; pos: postmetopidium scoli.

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 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.