Lycoderides santainensis, Flórez-V & Lapèze & Urban, 2025

Flórez-V, Camilo, Lapèze, Jérémie & Urban, Julie M., 2025, Taxonomic and ecological notes on Lycoderides Sakakibara (Hemiptera: Membracidae), including two new species from the highlands of the Colombian Andes, Zootaxa 5665 (2), pp. 151-186 : 164-169

Lycoderides santainensis Flórez-V sp. nov.

urn:lsid:zoobank.org:act:330647A0-E0C5-4FB6-83B7-9B00DA2FB450

( Fig. 3A–B , 4E–F , 5E–F , 6C–D , 10D, 10H , 11D, 11H , 12K–O , 14J–M , 15 )

Diagnosis: Sexually dimorphic. Warts distinct in color from surrounding body area. Forewing veins with wart-like protuberances. Male. General color reddish dark brown to dark brown. Metopidium vertical; pronotum elevated above head 1× head length, descending almost from above humeral angles; suprahumeral horns reduced to white wart-like protuberances. Forewing veins with warts; hyaline area from mid-length to apex, densely speckled around veins at third apical; veins lighter on non-speckled areas; sclerotized and punctuated until behind half-length. Female. General color light grizzled brown to brown. Metopidium vertical, with one pronotal anterior process obliquely directed, bearing two triangular, carinate suprahumeral horns. Forewing opaque hyaline from apex of basal cell M to apex; brownish on fourth and fifth apical cells and apical area of second and third apical cells; sparsely speckled with brown; densely speckled with dark brown spots around R 2+3, M 1+2, M 3+4, and m-cu.

Description. Holotype male ( Fig. 3A , 4E , 5E , 6C ). Color: General color reddish dark brown to dark brown. Head light brown, with one dark brown spot on coronal suture above frontoclypeus and one pair of dark brown spots oblique to ocelli; supra-antennal ledges lighter. Metopidium, supraocular callosities, and postocular lobes light brown; dorsal and posterior surface dark brown with warts; suprahumeral horns and lowest area of dorsal margin white. Forewing sclerotized, punctuated, reddish dark brown until behind mid-length; membrane from mid-length to apex hyaline, densely speckled around veins at third apical; veins lighter in non-speckled areas. Hindwings hyaline. Femora and tibiae light brown, tibiae with two darker bands; coxae dark brown. Abdominal segments III–VI orange, segments VII–IX dark brown.

Sculpture: Head, pronotum, sclerotized area of forewing, forewing veins, dorsal surface of tibiae and tarsomeres with light brown pubescence; surface densely and coarsely punctuated (opaque appearance); scattered wart-like protuberances around pronotal median carina and forewing veins.

Head: Hexagonal, twice as wide as long; eyes hemispheric; ocelli above transocular line, closer to posterior margin and to eyes than to each other; supra-antennal ledges arched; frontoclypeus slightly emarginate above supra-antennal ledges; in lateral view, head slightly projected posteriorly; beak extending to metacoxae.

Thorax: Ventral base of mesepimeron with finger-shaped process. Pronotum in frontal view ( Fig. 3A ), more or less triangular, elevated above head almost 1.5x head length; humeral angles triangular; anterior process with two suprahumeral horns reduced to white wart-like protuberances; in lateral view ( Fig. 4E ), metopidium vertical, truncate at level of suprahumeral horns; summit behind suprahumeral horns, descending more or less straight until above forewing vein A3, then forming small crest before reaching posterior margin of posterior process, terminating before claval suture of forewing; scutellum completely covered; in dorsal view ( Fig. 5E ), dorso-medial carinae irregular, zig-zag-line. Tibia with flattened dorsal surface, metatibia with row II of cucullate setae.

Abdomen ( Fig. 12K–O ): Male genitalia. Lateral plate totally fused to pygofer. Stlyes attached to subgenital plate on basal 1/3 of subgenital plate. Subgenital plate bilobed, divided almost from base, attached with VIII abdominal sternite. Styles in lateral view, mesally expanded, apex hook-shaped; tooth obliquely directed lateral- and ventrally. Aedeagus with anterior arm reduced, posterior arm in lateral view, basally expanded, then slightly narrow until apex; anterior and antero-lateral surface of apical 1/3 of posterior arm with denticles; in posterior view, subcylindrical.

Female ( Fig. 3B , 4F , 5F , 6D ). Color: Variegated light grizzled brown to brown. Post-clypeus dark brown. Pronotal posterior process tip light brown; warts dark brown. Forewing sclerotized, punctuated, and light grizzled brown until behind half-length; membrane from apex of basal cell M to apex opaque hyaline, sparsely speckled with brown; fourth and fifth apical cells and apical area of second and third apical cells brownish; veins R2+3, M1+2, M3+4, m-cu densely speckled with dark brown spots. Legs light grizzled brown; tibiae with two darker bands; femora and tarsomeres brown; coxae dark brown.

Sculpture: Surface densely and coarsely punctuated (opaque appearance). Light brown pubescence, sparsely distributed on head, pronotum, sclerotized area of forewing, forewing veins, dorsal surface of tibiae, tarsomeres. Waxy pubescence present on gena, post-ocular lobes, thoracic pleurites and sternites, coxae (leaving variable smooth spots), trochanter, ventral and dorsal surface of femora (leaving triangular smooth bands on anterior and posterior surface), and abdominal sternites. Wart-like protuberances scattered around dorso-median carina and forewings.

Thorax: Mesepimeron lacking finger shape process on ventral base (different to males). Pronotum in frontal view ( Fig. 3B ), more or less triangular; with pronotal anterior process, elevated 1× head length above metopidium, bearing two triangular, carinate suprahumeral horns, each carina surrounded by warts, dorso-median carina from anterior margin of metopidium; in lateral view ( Fig. 4F ), pronotal horn directed obliquely forward and upward, 0.5× length of remainder pronotum length; dorsal margin descending almost straight until tip of pronotal horn to posterior third, then forming a small crest before reach posterior margin of posterior process; scutellum completely covered; in dorsal view ( Fig. 5F ), suprahumeral horns triangular. Left forewing with two additional r-m cross-veins, and on right forewing with one complete and one partial additional m-cu crossvein.

Abdomen ( Fig. 14J–M ): Female genitalia. Terga with sclerotized areas and pits ventrally. Sternites, pleurites, ventral margin of pygofer and gonoplac with thick and rolled setae. Genitalia: Second valvulae blade-shaped, basal half narrower than apical half; apical half separated, dorsal margin of apical half with sub-quadrangular teeth (TE); ramus extended to apical portion; pores over ramus and below dorsal margin of apical half and ventral margin of 1/3 apical area.

Variations: There is no difference in color between live and pinned specimens.

Late instar nymph ( Fig. 10D, 10H , 11D, 11H ): Dorsoventrally compressed. Color: brownish orange. Sculpture: Dorsal tegument sparsely covered with chalazal setae and plant indument, moss, and lichen. Head: small subtriangular processes above ventro-anterior margin; in frontal view; in ventral and dorsal view, ventro-anterior margin forming a continuous and widely arched margin with ventrolateral lobes. In lateral view, projected obliquely forward and downward; in dorsal view, more or less five times wider than long, rectangular. Thorax: Pronotum with metopidium obliquely directed dorso-posteriorly; dorso-median margin forming a sinuous plateau with two rows of short chalazae; two humps (suprahumeral horns) at each side of anterior plateau. Abdomen: In dorsal view, tergal segments IV–VIII with large subtriangular lateral lamellae, directed postero-laterally, smaller on tergum IV; terga III–VIII with three pairs of longitudinal rows of short tuberculate chalazae: one pair near the midline, one above the lateral lamellae, and one between the previously mentioned pairs. Live specimens: Densely covered by plant indument, moss, and lichen. Color varies from green to brownish orange, seemingly influenced by location on host plant.

Measurements: Holotype male/ paratype female (mm): Body length: 7.47/8.71; pronotum length: 4.57/5.15; pronotum height: 1.73/2.55; forewing length: 6.28/7.42; width between humeral angles: 2.50/2.94; head width: 2.29/2.61; vertex length: 0.85/0.86; vertex width: 1.49/1.76.

Biology: The nymphs of this species are highly camouflaged against their host plant, Quercus humboldtii ( Fagaceae ) ( Fig. 15A, C ). They are covered with the plant’s indument and moss and are commonly found in leaf axils, often beneath the stipules ( Fig. 15C ). Nymphs do not form aggregations , but they are often found in close proximity, with up to three individuals near each other. They were always associated with Crematogaster sp. (Myrmicinae) ( Fig. 15A ). This ant species is usually present in small numbers (1–3 ants per nymph). Frequently, when the nymphs or the nearby branch are disturbed, the ants raise their abdomens and move them as if vibrating, eventually secreting a white substance. However, some ants flee when the nymphs are disturbed. On one occasion, three ants were observed feeding on a late-instar nymph ( Fig. 15B ).

Observations were made during four field trips to the ‘Páramo de Santa Inés’ (3250 masl), the first in February of 2015, the second in August of 2015, the third in February 2016 and the last in December 2016. The first time, nymphs were found on a seedling of Q. humboldtii , tended by Crematogaster sp. In this trip, nymphs or adults were not found on other nearby individuals of Q. humboldtii . During the second field trip, nymphs were found with ants and a male in the same individual of Quercus ( Fig. 15D ). Also, on the upper part of this plant, an aggregation of nymphs and a female of Metcalfiella sp. ( Membracinae ) were observed ( Fig. 15G ). Ants were not observed tending these treehoppers, but ants were feeding on honeydew drops that fell on a nearby leaf. On another individual Quercus plant, one male was found.

During the third field trip, nymphs were once again observed on the same Quercus individual, as well as on four additional plants. On one of these plants, a female was spotted, well-camouflaged and mimicking the color of the oak stem ( Fig. 15E–F ). Thus, nymphs, males, and females exhibit mimicry with the plant’s indument, though in different ways. Additionally, on the same plants, two different species of Metcalfiella and a female of Aetalion nigromarginatum ( Aetalionidae ) were observed sitting on eggs ( Fig. 15H ). During the last field trip, only nymphs were found on the same Quercus individuals.

Examined material: Holotype male in CBUCES. COLOMBIA: Antioquia: Belmira : “ COLOMBIA. Antioquia, Belmira, Páramo de Santa Inés, por el lado de la Laguna en Sabanas, 6.615899°N, 75.648855°W, 2800-3250 msnm, manual, en Quercus humboldtii, Ago. 16/2015, leg. J. Cardona & C. Flórez-V, CBUCES-F 8030” ( CBUCES).

Paratypes in CBUCES : “ COLOMBIA. Antioquia, Belmira, Páramo de Santa Inés por el lado de la Laguna en Sabanas , 6.615899°N, 75.648855°W, 2800–3250 msnm, manual, en Quercus humboldtii, Feb. 14/2016, leg. C. Flórez-V, CBUCES-F 8031” (1 female in CBUCES); “ COLOMBIA. Antioquia, Belmira, Páramo de Santa Inés , Sector Sabanas, cerca de la Laguna, 6.615899°N, 75.648855°W, 2800–3250 msnm, manual, en Quercus humboldtii, Feb. 4–6/2015, leg. N. Arcila, C. Flórez-V, I. Olivares \ CBUCES-F 8024” (1 nymph in CBUCES); “ COLOMBIA. Antioquia, Belmira, Páramo de Santa Inés , Sector Sabanas, cerca de la Laguna, 6.615899°N, 75.648855°W, 3200–3250 msnm, manual, en Quercus humboldtii, Ago. 16/2015, leg. J. Cardona, V. Correa, C. Flórez-V, H. Londoño, CBUCES-F 447” (1 male in CBUCES); “ COLOMBIA. Antioquia, Belmira, cerca del Páramo de Santa Inés , robledal subiendo a Sector Sabanas, cerca de la Laguna, 6.615899°N, 75.648855°W, 2900 msnm, manual, en Quercus humboldtii, Sep. 2–7/2017, leg. V. Correa, C. Flórez-V, A. Ospina, D. Taborda, CBUCES-F 1708” (1 female in CBUCES) .

Holotype minuten mounted, in excellent state of preservation. Paratypes adults minuten mounted. One male and one female paratype with dissected abdomens placed in vials with glycerin pinned with specimens. Two nymphs and 1 male in 75% ethanol.

Distribution: COLOMBIA: Antioquia: Belmira (Páramo de Santa Inés: 6.621161°N, 75.645648°W; 2800– 3250 masl) ( Fig. 1 ).

Etymology: The name refers to the type locality in ‘Páramo de Santa Inés’, located in the northern part of the Cordillera Central in Antioquia, Colombia. This remarkable and ecologically significant páramo is one of the main water sources for Medellín and 11 municipalities in central Antioquia.

Remarks: Males of L. santainensis sp. nov. and L. abditus are similar. However, males of L. santainensis sp. nov. can be distinguished by their larger size, darker and more reddish coloration, vertical metopidium (as opposed to the forward-reclined metopidium in L. abditus ), suprahumeral horns reduced to small warts, forewing veins with warts, and relatively longer forewings in proportion to pronotal length. Additionally, the tibiae are more foliaceous. Males of L. santainensis also resemble those of L. serraticornis and L. aburraensis , but they can be distinguished by their larger size, pronotal shape, smaller suprahumeral horns, forewing sclerotization, coloration, and wart distribution, as well as the male genitalia, where the aedeagus, which has a more prominent hump at the base of the posterior arm ( Fig. 12N ), as well as by its styles, which are broader along their extension in both lateral and dorsal views ( Fig. 12L–M ). Males of these four species share a distinct pronotal shape compared to other Lycoderides , lacking a well-developed anterior process and having reduced suprahumeral horns.

The female of L. santainensis differs from other Lycoderides species and can be distinguished by its pronotum, which has an anterior horn elevated to a height equal to the head length above the metopidium, the presence of warts throughout the pronotal horn and the dorso-medial carinae, and an elevated posterior region of the posterior pronotal process. Additionally, the thick and rolled setae on the sternites, pleurites, pygofer, and gonoplac have not been observed in any of the other species examined in this study ( Fig. 14M ). The female of this species is morphologically similar to those of L. abditus and L. serraticornis , sharing features such as a pronotum with an anterior horn, a scutellum concealed by the pronotum, and partially hyaline and speckled forewings.

This represents the first record of a Stegaspidinae species occurring above 3000 masl, as well as the first record of the genus Lycoderides on Fagaceae , specifically on the genus Quercus . Both adults and nymphs camouflage themselves in different ways with their host plant. The dorsoventrally flattened shape of the nymphs resembles that of other groups of Membracidae in which nymphs are camouflaged when lying on the twigs of their host plant, such as certain species of Nicomiinae (e.g., Nicomia , Flórez-V pers. obs.) and other Stegaspidinae (e.g., Microcentrini, Bocydium , Stylocentrus , Problematode ; Flórez-V & Evangelista 2017, Flórez-V 2019). On the other hand, males and females mimic the plant bark in different ways: females resemble the stem bark in more sclerotized areas, while males resemble the axillary buds.

The accumulation of moss or plant indument in nymphs may be facilitated by the presence of chalazae and lateral abdominal lamellae, as well as by the flattened body shape and possibly by specific behaviors that allow individuals to actively collect these materials, since this external covering would be lost with each molt.

Figures

FIGURE 1. Distribution map in Colombia of Lycoderides aburraensis sp. nov., L. amazonicus, L. brevilobus, L. brulei, L. fernandezi / Lycoderes argutus, L. luteus, L. marginalis, L. phasianus, L. santainensis sp. nov. and L. serraticornis.

FIGURE 3. Frontal habitus of the new species of Lycoderides related with L. serraticornis. Lycoderides santainensis sp. nov. A) male holotype and B) female paratype. Lycoderides aburraensis sp. nov. C) male holotype. Scale: 1 mm.

FIGURE 4. Lateral habitus of the species of Lycoderides related with L. serraticornis. Lycoderides abditus A) male, B) female. Lycoderides serraticornis C) male and D) female. Lycoderides santainensis sp. nov. E) male holotype and F) female paratype. Lycoderides aburraensis sp. nov. G) male holotype. Scale: 1 mm.

FIGURE 5. Dorsal habitus of the species of Lycoderide related with L. serraticornis. Lycoderides abditus A) male, B) female. Lycoderides serraticornis C) male and D) female. Lycoderides santainensis sp. nov. E) male holotype and F) female paratype. Lycoderides aburraensis sp. nov. G) male holotype. Scale: 1 mm.

FIGURE 6. Frontolateral habitus of Lycoderides species related to L. serraticornis. Lycoderides serraticornis A) male and B) female. Lycoderides santainensis sp. nov. C) male holotype and D) female paratype. Lycoderides aburraensis sp. nov. E) male holotype.

FIGURE 10. Nymphs of some Lycoderides species. Frontal habitus of: L. luteus A) morph present on Miconia sp. (Melastomataceae) and B) morph species present on Andesanthus lepidota (Melastomataceae); C) L. phasianus, D) L. santainensis sp. nov. Lateral habitus: L. luteus E) morph present on Miconia sp. (Melastomataceae) and F) morph species present on A. lepidota (Melastomataceae); G) L. phasianus, H) L. santainensis sp. nov.

FIGURE 11. Nymphs of some Lycoderides species. Ventral habitus of: L. luteus A) morph present on Miconia sp. (Melastomataceae) and B) morph species present on Andesanthus lepidota (Melastomataceae); C) L. phasianus, D) L, santainensis sp. nov. Dorsal habitus: L. luteus E) morph present on Miconia sp. (Melastomataceae) and F) morph species present on A. lepidota (Melastomataceae); G) L. phasianus, H) L. santainensis sp. nov.

FIGURE 12. Male pygofer, subgenital plates and styles (dorsal and lateral view), and aedeagus (lateral and caudal view) of some Lycoderides species. A–E) Lycoderides serraticornis, F–J) L. aburraensis sp. nov., K–O) L. santainensis sp. nov.

FIGURE 14. Gonoplac, first and second valvulae of Lycoderides species.A–C) Lycoderides serraticornis, D–F) L. luteus, G–I) L. phasianus, J–M) L. santainensis sp. nov.

FIGURE 15. Biology of Lycoderides santainensis sp. nov. A) Nymph tended by Crematogaster sp. B) Nymph predated by Crematogaster sp. C) Nymph of a late instar camouflaged with the host plant indumentum. D) Male. E) and F) Females camouflaged with branch coloration. G) Female of Metcalfiella sp. guarding their nymphs, on the same plant individual with nymphs of L. santainensis sp. nov. H) Female of Aetalion nigromarginatum on the same plant individual with L. santainensis sp. nov.