Holotype : Ecuador • 1 ♀; Santo Domingo de los Tsáchilas, Santo Domingo, Río Baba -0.30295, -79.15211, 480 m; 12 May 2023; Montalvo, J. & Rueda, M. P. leg.; Ex. Manual ZSFQ-i12112. Paratypes: Ecuador • 2 ♂; same labels as for holotype; ZSFQ-i12110, ZSFQ-i12111 • 2 ♀, 1 ♂ Santo Domingo de los Tsáchilas, Santo Domingo, Quebrada del Río Pove -0.25237, -79.156668, 570 m; 14 Aug. 2023; Rueda, M. P. & Montalvo, J. leg.; Ex. Manual; ZSFQ-i17766:17768 • 3 ♀, 1 ♂ same locality as paratypes; 20 Apr. 2024; Rueda, M. P. leg.; Ex. Manual; ZSFQ-i18855:18858.

All specimens are minuten-mounted. Dissected abdomens of the holotype and two female and two male paratypes were placed in vials with glycerol pinned beneath the specimens. Some paratypes are in poor condition, with legs or wings lost.

Ecuador • 3 5^th instars; same data as paratypes; ZSFQ-i17925, 17926.

Overall color black with whitish dorsal spot and subapical band; metopidium straight, two to four sub-equal accessory carinae. Sexually dimorphic: female with obtuse projection instead of horn, short lateral carina not surpassing humeral angles, dorsal spot 2× longer than subapical band, and longer than male in size; male with narrow horn slightly curved forwards, lateral carinae almost touching lateral margin of pronotum and dorsal spot subequal in length to subapical band.

Female holotype (ZSFQ-i12112). Measurements (mm): Length from head to wings at rest: 5.7; Total length: 6.3; Head to apex of posterior process: 4.6; Pronotal length: 4.4; Head to horn apex: 2.4; Forewing length: 4.9; Body width: 1.9; Vertex width on ocellar line: 1; Head length: 1.2; Frontoclypeus length: 0.8; Frontoclypeus width: 0.7; Prothoracic tibia length: 0.9; Metathoracic tibia length: 1.4; Metathoracic tibia width: 0.3; Prothoracic tibia width: 0.3.

Color. Overall black with whitish dorsal spot and subapical band on dorsum. Dorsal spot 2× longer than subapical band. Eyes black with dark brown margins, ocelli golden. Forewings opaque dull black, hind wings hyaline, veins black. Tarsi golden.

Surface. Head, pronotum, ventral sclerites of thorax, legs, and abdomen with dense golden pubescence; subcostal cell and veins of forewings with short, dispersed, almost indistinguishable golden pubescence. Pronotum (except metopidium) strongly punctured. Head, metopidium, forewings, and legs rough.

Head. Triangular, longer than wide (avoiding eyes); ocelli closer to eyes than each other; supra-antennal ledges arranged above clypeus; clypeus broad, longer than wide, anterior margin rounded; rostrum reaching hind coxae (Fig. 1A).

Figure 1. 

Enchenopa gennyae sp. nov. holotype female A–C habitus in frontal, dorsal view, and lateral views respectively D illustration of pronotum showing the carination. Scale bars: 1 mm (A); 2 mm (B, C).

Thorax. Pronotum somewhat compressed, in lateral view, triangular, dorsal contour arched; metopidium straight, inclined anteriorly; horn reduced to an obtuse projection, obliquely directed dorso-anteriorly, wider than long, apex broadly rounded; median carina laminated and somewhat foliaceous, especially on dorsum; lateral carina short, not extending beyond humeral angles (Fig. 1C); three parallel accessory carinae, almost as long as lateral carina extending ventroposteriorly from projection apex; posterior apex of pronotum acuminate almost reaching the apex of first apical cell (Fig. 1D); humeral angles slightly produced (Fig. 1B). Forewings with five apical cells, one discoidal cell, and one vein in the clavus, one r-m crossvein, two m-cu crossveins, and without s crossvein; apical limbus broad. Hind wings with four apical cells and one r-m crossvein. Anterior and middle tibiae foliaceous; posterior femur with apical, ventral and middle cucullate setae, metathoracic tibia compressed with spine-like cucullate setae on rows I and II, row III absent.

Abdomen. Sternum III with a transverse keel extending along the sternite and slightly projected ventrally. Dorsum of tergites VII and VIII with medial tuberosities, tergites IV–VI with reduced medial tuberosities (Fig. 2A). Genitalia. Gonoplac ventrally with few setae and more sclerotized, apex rounded (Fig. 2B). First valvula blade shaped, apex ventrally rounded, dorsally acuminate extending beyond ventral margin (Fig. 2C). Second valvula broad throughout, dorsally rounded, ventrally weakly serrated with a ventral apical tooth directed upwards (Fig. 2D, E).

Figure 2. 

Abdomen and terminalia of Enchenopa gennyae sp. nov. A undissected female abdomen in lateral view B gonoplac in lateral view C first valvula in lateral view D second valvula in lateral view E close-up apex of second valvula F undissected male pygofer in lateral view G styles in dorsal view H aedeagus in lateral view. Scale bars: 1 mm (A); 0.1 mm (B–D); 0.01 mm (E); 0.2 mm (F); 0.05 mm (G, H).

Male paratype (ZSFQi-17766). Similar to female except dorsal spot as long as subapical band, pronotal horn narrow and curved forwards, lateral carina almost touching lateral margin of pronotum, metopidium with three accessory carinae at each side. Genitalia. Subgenital plate, in lateral view, 3× longer than wide, lobes diverging in first 1/4, dorsal margin concave, distally expanded (Fig. 2F). Styles 5× as long as wide, anterior projection subequal to posterior projection; shank with notch at middle of ventral margin, distally recurved and apically truncate, slightly expanded with posterior end longer and narrower than anterior end (Fig. 2G). Aedeagus U-shaped with anterior arm smaller than posterior arm and rounded; posterior arm lanceolate and abruptly narrowed at 1/3 length, anterior surface smooth, without serrations (Fig. 2H).

Measurements. Male / Female (mm): Length from head to wings: 4.4–5.1 / 5.3–5.8; Total length: 4.9–5.6 / 6.2–6.3; Head to apex of posterior process: 3.8–4.5 / 4.58–5; Pronotal length: 3.8–5.5 / 4.4–5.3; Head to horn apex: 2.2–2.7 / 2.4–2.9; Forewing length: 3.7–4.2 / 4.5–5.1; Body width: 1.6–1.7 / 1.9–2; Vertex width on ocellar line: 0.9–1 / 1.0–1.1; Head length: 1.1–1.2 / 1–1.2; Frontoclypeus length: 0.7–0.8 / 0.7–0.8; Frontolypeus width: 0.7 / 0.7–0.8; Metathoracic tibia length: 0.8–1 / 0.9–1; Prothoracic tibia length: 1.2–1.4 / 1.4–1.8; Metathoracic tibia width: 0.2–0.3 / 0.3–0.5; Prothoracic tibia width: 0.3–0.4 / 0.3.

Females longer than males, with obtuse projection instead of horn, dorsal spot 2× longer than subapical band, lateral carina not surpassing humeral angles and, in some individuals, weakly produced; two to four secondary carinae almost as long as lateral carina. Male with horn narrow and curved forwards, dorsal spot < 2× subapical band length (Fig. 3A–C), lateral carina almost attaining lateral margin of pronotum and metopidum with three accessory carinae on each side (Fig. 3D). Independent of gender, dorsal contour of pronotum is more or less arched.

Figure 3. 

Enchenopa gennyae sp. nov. paratype male and nymph A–C male habitus in frontal, dorsal view, and lateral views respectively D illustration of male pronotum showing the carination E nymph habitus in lateral view. Scale bars: 1 mm (A); 2 mm (B, C, E).

Overall color mostly white with black tarsi and scoli (Fig. 3E). One pair of abdominal scoli on each segment from III to VII; scoli length 5–6× basal width. Pronotum anteriorly with nascent horn not extended beyond head and posteriorly extended to abdominal segment III; anterior apex rounded and directed forward; posterior apex acute, dorsal margin convex in middle. Needle-like setae on chalazae distributed over whole body.

Specimens of Enchenopa gennyae sp. nov. were found in two secondary forest remnants of the Western Foothills Forest from the urban area of ​​Santo Domingo (Fig. 9): on the banks of the Baba River (Fig. 4C) and Pove River’s ravine (Fig. 4D). Adult and nymph congregations were found on several occasions cohabiting together and perched on the underside of leaves and stems of an unidentified species of the genus Piper L. between 100 and 150 cm above the ground (Fig. 4A). Females were always more abundant than males in these congregations. Nymphs were attended by fire ants of the species Wasmannia auropunctata (Roger) (Fig. 4B).

Figure 4. 

Enchenopa gennyae sp. nov. in its natural environment and habitat A male paratype perched on a stem of its host plant B nymphs attended by Wasmannia auropunctata (Roger) C shore of Baba River D ravine of Pove River.

The species is dedicated to the mother of the first author, Genny Elizabeth Rodríguez Cueva, who helped to find the specimens of this species and has been a great support and inspiration throughout her life.

Females of Enchenopa gennyae sp. nov. have a short lateral carina that does not extend beyond the humeral angles, while males have a long lateral carina that almost reaches the lateral margin of pronotum. In the tribe Membracini, the length of the lateral carina has not been previously reported as sexually dimorphic in any species. However, in some species of Membracis, it has been noted that the lateral carina may or may not be present among individuals (Richter 1947). Likewise, Richter (1954) argued the lateral carina of some species of Enchophyllum and Enchenopa can vary in length within the same population. This species is the first known in the Enchenopa biplaga species group to exhibit sexual dimorphism in pronotum shape and lateral carina length. Like Enchenopa gennyae sp. nov., E. longimaculata Strümpel & Strümpel, 2014 has remarkable sexual dimorphism of the dorsal spot. However, in Enchenopa longimaculata the females the spot is 4× the length of the subapical band and in males 2× while in E. gennyae sp. nov. the spot is 2× as long in females but subequal in males.

Sexual dimorphism in the shape of the pronotal horn is characteristic of the Enchenopa minuta species group. However, Enchenopa gennyae sp. nov. does not belong to that group, given its coloration, the shape of the second valvula and, more importantly, the female’s lack of a pronotal horn; in the E. minuta species group, males lack the horn and females generally have a developed horn.

Enchenopa gennyae sp. nov. belongs to the E. biplaga species group due to the presence of bands on dorsum of the pronotum, the second valvula with a ventral apical tooth, and forewings with one discoidal cell. Enchenopa gennyae sp. nov. differs from E. ignidorsum (Walker, 1858) and E. vittifera (Stål, 1869) by the two white lateral bands instead of just one yellow or white one, respectively. Enchenopa gennyae sp. nov. differs from E. dubia (Fowler, 1894), E. lanceolata, E. reticornuta Strümpel & Strümpel, 2014, and E. richteri Strümpel & Strümpel, 2014 by a horn shorter than the distance between the tips of humeral angles rather than longer. Enchenopa singularis Strümpel & Strümpel, 2014 and E. longimaculata have two accessory carinae with the posterior one ~ 1/2 or almost the total length, respectively, of the lateral carina, while E. gennyae sp. nov. has from two to four somewhat subequal accessory carinae. E. gennyae sp. nov. is distinguished from E. melaleuca Walker, 1858 by the shorter horn that is not curved forward. The new species closely resembles Enchenopa biplaga Walker, 1858 due to the shape of the pronotal horn, pronotum coloration, and the disposition of the accessory carinae. However, Enchenopa gennyae sp. nov. has a straight metopidium, less produced horn, and sexual dimorphism. In contrast, in E. biplaga the metopidium is convex, the horn is large and strongly produced, and without sexual dimorphism. The females of E. gennyae sp. nov. have a short and straight horn while E. biplaga females have the horn longer than wide and curved forward. In males of E. gennyae sp. nov. the posterior arm of the aedeagus, in lateral view, is abruptly narrowed at one-third of its length, and the apical hook of the styles has the posterior tooth longer and narrower than the anterior tooth. In contrast, in the male of E. biplaga the width of the posterior arm of the aedeagus, in lateral view, slightly decreases at half of its length, and the apical hook of the styles has the posterior tooth similar in size to the anterior tooth.

McKamey (2022) reinstated Enchenopa melaleuca from the genus Enchophyllum and suggested it shares morphological similarities with some species of the E. biplaga species group. However, the holotype of this species has yet to be reviewed to confirm these affinities. Therefore, excluded E. melaleuca from the key to E. biplaga group species but compared it with E. gennyae sp. nov. in the above discussion.

In the E. biplaga species group, nymphs of E. reticornuta and E. vittifera are known (Strümpel and Strümpel 2014). They share with the nymph of E. gennyae sp. nov. the body covered with white wax-like material, black tarsi, and the presence of scoli on abdominal tergites III–VIII. However, we found the nymphs of E. gennyae sp. nov. differ from them mainly in the shape of the pronotum and scoli. The nymphs of Enchenopa vittifera have a longer horn directed forward with the posterior apex of the pronotum reaching the abdominal tergite III and have shorter scoli. While the nymphs of Enchenopa reticornuta have a straight horn with the posterior apex of the pronotum not touching the abdomen and longer scoli widened at the base.

Holotype : Ecuador • 1 ♂; Pichincha, Tandayapa Cloud Forest Station -0.009645, -78.688058, 2280 m of elevation; 3 Fbr. 2024; Rueda, M. P. leg.; Ex. Manual; ZSFQ-i18060. Paratypes: Ecuador • 1 ♂; same data as for holotype; ZSFQ-i18061 • 3 ♀, 1 ♂; Pichincha, Tandayapa Cloud Forest Station -0.009645, -78.688058, 2280 m of elevation; 9 Fbr. 2024; López-García, M. M., Montalvo, J. & Rueda, M. P. leg.; Ex. Mercury light; ZSFQ-i10862: 10865 • 1 ♀; Pichincha, Mindo, 0.04166, -78.77472, 1300 m of elevation; 11 Jun. 2022; Torres, D. leg.; Ex. Manual; ZSFQ-i8423 • 1 ♀, Imbabura, Seis de Julio de Cuellaje, 0.4509352, -78.525948, 2000 m of elevation; 13 Nov. 2021; Rubio, A. leg.; Ex. Manual; ZSFQ-i8196.

Holotype and most paratypes are minuten-mounted. The paratype female ZSFQ-i8243 was originally pinned, but later the specimen was transferred to double mounting on a minuten pin. Dissected abdomens of holotype, one male paratype, and three female specimens placed in vials with glycerol pinned with specimens.

Overall coloration black with scarlet median carina and posterior apex in females and scarlet posterior apex in males, apical 1/3 of forewing amber; pronotal horn absent, replaced by obtuse projection directed obliquely forwards, lateral carina almost touching lateral margin of pronotum; three or four weak accessory carinae, some touching lateral carinae or bifurcate.

Male holotype (ZSFQ-i10860): Measurements (mm): Length from head to wings: 5.3; Total length: 5.4; Head to apex of posterior process: 4.5; Pronotal length: 4.6; Head to horn apex: 1.5; Forewing length: 4.4; Body width: 2.2; Vertex width on ocellar line: 1.2; Head length: 1.2; Frontoclypeus length: 0.7; Frontoclypeus width: 0.1; Metathoracic tibia length: 0.9; Prothoracic tibia length: 1.8; Metathoracic tibia width: 0.3; Prothoracic tibia width: 0.2.

Color. Overall color black. Eyes brownish, ocelli golden. Posterior apex of pronotum scarlet red. Forewings almost entirely opaque black with an apical translucent amber patch restricted on the third to fifth apical cells and limbus around this area. Tarsi pale brownish.

Surface. Head, pronotum, ventral sclerites of thorax, legs, and abdomen with golden pubescence (Fig. 5B); sclerotized area of forewings with shorter pubescence. Pronotum (except by metopidium) and sclerotized area of forewings strongly punctured. Head, metopidium, legs, and abdomen rough.

Figure 5. 

Enchenopa chocoandina sp. nov. holotype male A–C habitus in frontal, dorsal view, and lateral views respectively D approach to base of horn in lateral view, the white arrows indicate the accessory carinae. Scale bars: 1 mm (A); 2 mm (B, C).

Head. Triangular blunt, longer than wide (excluding eyes); distance between ocelli subequal to ocelli-eye distance; supra-antennal ledges arranged above clypeus; clypeus broad, longer than wide, anterior margin rounded; rostrum reaching posterior coxae (Fig. 5A).

Thorax. Pronotum, in lateral view, triangular; metopidium straight directed forwards; pronotal horn absent instead an obtuse projection with rounded apex; humeral angles slightly produced (Fig. 5B); median carina sharp; lateral carina parallel to median carina, running from apex of anterior projection to middle of the lateral margin of pronotum, almost touching the margin (Fig. 5C); three accessory carinae short (1/10 length of lateral carina), weak, irregular, sub-perpendicular to primary lateral carina, the last two diverging from the lateral carina (Fig. 5D), left side with the anterior two accessory carinae convergent; posterior apex of pronotum acuminate, just surpassing first apical cell. Forewing with one vein on clavus, two discoidal cells, two m-cu cross veins, one s cross veins which enclose second discoidal cell, and five apical cells; apical limbus broad. Anterior and middle tibiae foliaceous; posterior femur with apical ventral and middle cucullate setae, posterior tibia with spine-like cucullate setae on rows I and II, row III absent.

Abdomen. Sternum III with a transversal keel extended along the sternite, strongly projected downwards and medially invaginated. Tergites III to VI with a pair of medial tuberosities, tuberosities of tergite VI strongly developed (Fig. 6D). Subgenital plate, in lateral view, 3× longer than wide, lobes diverging since the base, dorsal margin concave (Fig. 6A). Aedeagus with posterior arm lanceolate, 2× longer than anterior arm and strongly inclined forwards; anterior face of posterior arm armed at apical1/3 with small dorsal apical denticles, gonopore subapically (Fig. 6C). Styles distally recurved, apically truncate with anterior part longer than posterior part, spine tuft on dorsal margin just anterior to apex (Fig. 6B).

Figure 6. 

Abdomen and terminalia of Enchenopa chocoandina sp. nov. A undissected male pygofer in lateral view B styles in dorsal view C aedeagus in lateral view D undissected female abdomen in lateral view E gonoplac in lateral view F first valvula in lateral view G second valvula in lateral view H close-up apex of second valvula. Scale bars: 0.1 (A, H); 0.05 (B, C); 1 mm (D); 0.2 mm (E–G).

Female paratype (ZSFQ-i8423): Similar to male except for the pronotal projection more angulated and produced, central carina reddish behind humeral angles and forewings with amber patch extended from the second to fifth apical cells and limbus around this area. Genitalia. Gonoplac ventrally setose and more sclerotized than dorsally (Fig. 6E). First valvula blade shaped, basal 2/3 broad, apex acuminate (Fig. 6F). Second valvulae blade shaped with two dorsal tubercles in apical 1/2 (Fig. 6G, H).

Nymph unknown.

Measurements. Female / male (mm): Length from head to wings: 5.7–5.9 / 5.3–5.5; Total length: 5.9–6.4 / 5.4–5.7; Head to apex of posterior process: 5–5.6 / 4.3–4.5; Pronotal length: 4.9–5.5 / 4.5–4.6; Head to horn apex: 1.6–1.9 / 1.5–1.8; Forewing length: 5–5.4 / 4.4–4.8; Body width: 2.3–2.7 / 2.1–2.2; Vertex width on ocellar line: 1.2–1.4 / 1.1–1.2; Head length: 1.3–1.5 / 1.0–1.2; Frontoclypeus length: 0.7–0.9 / 0.7–0.8; Frontoclypeus width: 0.8–1.0 / 0.7–0.9; Metathoracic tibia length: 1–1.2 / 0.9–1.1; Prothoracic tibial length: 1.5–1.9 / 1.8–1.9; Metathoracic tibia width: 0.3–0.4 / 0.3–0.3; Prothoracic tibia width: 0.2–0.4 / 0.2–0.3.

Females are longer and have more produced pronotal projections than males (Fig. 7A, C), three or four accessory carinae (Fig. 7D), lateral carina behind humeral angles reddish rather than just the posterior apex of pronotum, and the amber membrane is most extended (Fig. 7B). Independent of gender, some accessory carinae are bifurcate or converged.

Figure 7. 

Enchenopa chocoandina sp. nov. paratype female A–C habitus in frontal, dorsal view, and lateral views respectively D approach to base of horn in lateral view, the white arrows indicate the accessory carinae. Scale bars: 1 mm (A); 2 mm (B, C).

This species is distributed in the Montane forests of northwest Ecuadorian Andes (Fig. 9), between 1300 to 2300 m elevation. It inhabits the borders of secondary forests (Fig. 8B) and is a solitary species. The species has been recorded perched on the leaves or stems of different species of Asteraceae and Araceae, but more oftenly on Munnozia pinnatipartita (Hieron.) H.Rob. & Brettell (Asteraceae) (Fig. 8A), an endemic Ecuadorian species (Barriga et al. 2011). The species has been observed active during the day and attracted to mercury light traps at night between 8 pm and 2 am (Fig. 8C).

Figure 8. 

Enchenopa chocoandina sp. nov. in its natural environment and habitat A paratype female from Mindo by David Torres B border of a secondary forest at Tandayapa Cloud Forest Station C female paratype attracted by a light trap.

Figure 9. 

Distribution of the new species in Ecuador. Red stars = records of Enchenopa gennyae sp. nov.; blue stars = records of Enchenopa chocoandina sp. nov.

The species, a noun in apposition, is named after the Andean Choco Biosphere Reserve declared by UNESCO as the seventh biosphere reserve of Ecuador, where this species lives. It honors the people who defend this territory from the metal mining that threatens the ecosystems and biodiversity of this important area.

Enchenopa chocoandina sp. nov. belongs to E. andina species group based on the pronotum with a horn or projection shorter than the distance between the humeral angles, the forewing with transparent patches, and the blade-shaped second valvulae. Moreover, some species of this group, such as E. pilosa and E. eurycephala, have dense pubescence, shared with E. chocoandina sp. nov.

Enchenopa chocoandina sp. nov. differs from Enchenopa loranthacina (Sakakibara & Marques, 2010) by the obtuse projection obliquely directed forwards rather than a horizontally inclined horn and from E. pilosa Strümpel & Strümpel, 2014 and E. eurycephala Strümpel & Strümpel, 2014 by the head longer than wide instead of as long as wide or wider than long respectively. Enchenopa chocoandina sp. nov. has a reddish median pronotal carina and amber forewing patches which separate it from E. monoceros (Germar, 1821) and E. luizae (Lencioni-Neto & Sakakibara, 2015) which have the median carina concolorous and forewing patches hyaline. The new species resembles E. andina (Schmidt, 1924) by the black overall coloration with median carina and posterior apex reddish, and the amber forewing apex; however, E. chocoandina sp. nov. does not have a horn instead an obtuse projection, longer and denser pubescence, and three or four weak and irregular accessory carinae instead of two or three. Moreover, the new species is considerably shorter than E. andina and differs in the male and female genitalia.

Enchenopa chocoandina sp. nov. lacks a distinctive horn; instead, it has an obtuse projection with a rounded apex, resembling species of the E. beebi species group or the males of E. minuta species group (Strümpel and Strümpel 2014). However, E. chocoandina sp. nov. properly does not fit within the E. beebi species group due to the absence of large punctation on the upper portion of the pronotum and dorsum, a short translucent apical patch and yellow tarsi. Neither does it belong to the E. minuta species group due to the absence of sexual dimorphism in pronotal horn shape; in E. chocoandina both females and males lack a horn.

We suggest E. andina and E. chocoandina sp. nov. could be related species by the scarlet median carina and posterior apex only shared in both species. Both inhabit mountain forests of the north Andes of Ecuador however are geographically separated by the Interandean Valley.