Venation typically for Cochylina (Fig. 1). Forewing (based on two slides) without costal fold; discal cell ca. 0.6 times length of wing, no M–stem, chorda obsolescent ca. 0.25 times length of wing, cross veins vestigial; all veins present except CuP; R₄ to the costa near apex, R₅ to the termen; distances between pairs of veins R₅–M_1, M₁-M₂ and M₂–M₃ on termen similar; distances between M3–CuA₁, CuA₁–CuA₂ and CuA₂–1A+2A similar; CuA₂ opposite on discal cell ca. 0.3–0.5 the distance between R1 and R₂ on the cell, approximately coincident with the point where the chorda meets Rs; anal loop ca. 0.3 times length of 1A+2A. Hindwing with Sc+R₁ somewhat parallel to Rs basally, length ca. 0.8 times length of wing; M₁ and Rs stalked in basal half; M₂, M₃ and CuA₁ obsolescent basally; M₃ and CuA₁ connate; CuA₂ well developed, CuP reduced, present only in distal portion; 1A+2A and 3A developed, anal loop ca. 0.4 times length of 1A+2A. Frenulum in males with one single bristle, three bristles in females. Male genitalia with transtilla broad and well developed; gnathos as two arms fused distally forming a short process, resulting in a plicate terminal plate; characteristic valva, elongate; cucullus with a more or less developed disc of hair like scales; sacculus with a free terminal process. Phallus with two distinctive sets of non-deciduous cornuti, one set as a ventral band of rather large aciculate cornuti basally attached; a second set in the inner part of vesica formed by microspinulate cornuti. Female genitalia with lobular lamella antevaginalis and postvaginalis; ventral spinous subpapillar sclerite on the 8-9 intersegmental membrane at the level of the ventral lobes of the anal papillae.

Figure 1. 

Venation in Brusqeulia yunkensis. Based on slides JBA20815 and JBA20836. Dashed lines indicate obsolescent veins.

Fifteen species have been described from Brazil and one from Ecuador (Razowski and Becker 2000, 2011). To that we add one species from Bolivia and another from Venezuela. Given the broad geographical and elevational range (from near sea level to ca. 2000 m), we suspect that Brusqeulia includes additional undiscovered species.

Holotype: ♂, Bolivia, Santa Cruz Department, Florida Province, Pampa Grande Municipality, locality of Hueco de la Pascana, 1575 m, 18°7.09'S; 64°3.58'W, 25 Jan 2011, J. Baixeras, A. Valdivia and G. Fernández (MNKM).

Paratypes: (15♂, 6♀). Bolivia: Santa Cruz Department, Florida Province, Mairana Municipality, locality of Yunga de Mairana, Rasete, 2000 m, 18°04'S; 63°54'W, 4 Nov 2005 (6♂, 3♀), J. Baixeras, A. Valdivia and I. García (GS USNM 124290, USNM 124291); Yunga de Mairana, ca. Bosque de Helechos, 2150 m, 18°03'S; 63°55'W , 02 Nov 2005 (1♂), J. Baixeras, A. Valdivia and I. García (GS 20724); locality of Pampa Grande, Hueco de la Pascana, 18°7.09'S; 64°3.58’W, 10 Nov 2001 (1♂), A. Valdivia and J. Baixeras; Pampa Grande, La Hoyada, 1600 m, 17°57'S; 64°06'W , 07 Nov 2005 (1♂, 1♀), J. Baixeras, A. Valdivia and I. García (GS 20727, 20728); Pampa Grande, Agua Clarita, 1554 m, 17°56.74'S; 64°7.97'W 27 Jan 2011 (5♂, 2♀), J. Baixeras, A. Valdivia and G. Fernández; Pampa Grande, El Milu, 1534 m, 17°59.36'S; 64°3.23'W, 28 Jan 2011 (1♂), J. Baixeras, A. Valdivia and G. Fernández. Paratypes deposited in MNKM, USNM, and ICBiBE.

Bolivia: Santa Cruz Department, Florida Province, Mairana Municipality, locality of Yunga de Mairana, Rasete, 2000 m, 18°04'S; 63°54'W, 4 Nov 2005 (1♂, 1♀), J. Baixeras, A. Valdivia and I. García (GS JBA20684, JBA20815, SEM stub JBA193); Pampa Grande, Agua Clarita, 1554 m, 17°56.74'S; 64°7.97'W 27 Jan 2011 (2♂), J. Baixeras, A. Valdivia and G. Fernández (GS JBA20836, JBA20844, JBA20864). Deposited in ICBiBE.

We were able to obtain partial COI sequence data (i.e., the DNA barcode) for a single specimen (GENBANK accession number MG951753), and comparison of the sequence against Genbank did not render any useful information. Interestingly, sequencing of a second sample revealed the presence of DNA related to the entomopathogenic trypanosomatid genus Crithidia Léger, 1902 (phylum Euglenozoa; GENBANK accession number MH118295).

The habitus of B. yunkensis (Fig. 2A) does not ensure discrimination from similar species of Brusqeulia (e.g., B. baeza or B. uncicera) or species of the closely related genus Limeulia Razowski & Becker, 2000. An examination of the available literature suggests that the crescent-shaped blotch of the forewing costa is present in all the species of the genus Brusqeulia and related genera. The distinctive characters at the species level are associated with the male and female genitalia. Brusqeulia yunkensis can be distinguished by the unusual configuration of the transtilla in males (Fig. 2B). The transtilla is well developed in most species of Brusqeulia and in associated genera, but a transtilla projecting posteriorly into a flat spinulous area is found only in B. crispula and presumably in B. monoloba. However, in B. yunkensis the spinulous area occupies ca. 0.2 of the total length of the transtilla, whereas in B. crispula and B. monoloba it occupies ca. 0.3 of the total length. The impression is a longer, more protruding transtilla in B. yunkensis than in the other two species. Brusqeulia crispula has a distinctive pillous disc on the cucullus that is present but only weakly developed in B. yunkensis; no disc is apparent in B. monoloba. There is wide variation in the development of the uncus in Brusqeulia species, from thin projections, as in B. bonita, B. baeza, and B. araguensis sp. n., to relatively broad finger-like projections, as in B. crispula and B. tripuncta. Brusqeulia yunkensis has a moderate development similar to that of B. teneimorpha and B. guaramiranga. A rugous spatulate projection of the gnathos has never been described in species of the group and could be a unique character. An inward curved sacculus distally projecting into a pointed process (Fig. 2C) is similar to that found in some species such as B. baeza, B. monoloba and even B. crispula, but the shape of the terminal process is diagnostic in every species of the group. Finally, the phallus of B. yunkensis seems to be a simplified organ with respect to the typical stout structure in its relatives, more elongate and without any distal ventral process. The presence of denticles on the dorsal distal part of the phallus (Fig. 2E) together with slender terminal cornuti (Fig. 2D) has been reported only in B. sebastiani. The presence of microspinulation on the inner part of the vesica is unknown in other species of the genus. So far, morphological details of the females of Brusqeulia are limited owing to the paucity of material. The only female described is B. caracagena, a species for which the male is unknown. The latter can be easily distinguished from B. yunkensis by the ductus bursae (Fig. 2F) – extremely short in B. caracagena, longer and partially sclerotised in B. yunkensis. The spinous subpapillar sclerite (Fig. 4B and D) on the intersegmental membrane, present in B. yunkensis, is absent in B. caracagena.

Figure 2. 

Morphological characters of Brusqeulia yunkensis sp. n. A habitus (Paratype, male, Bolivia, Santa Cruz, Pampagrande, 25 January 2011, ICBiBE) B male genitalia, phallus removed (GS JBA20728, z = 161 µm) C male genitalia left valva completely extended (GS JBA20684, z = 83 µm) D phallus with everted vesica (caecum removed, GS JBA20844) E phallus uneverted (GS JBA20728, same scale as D) F female genitalia (GS JBA20727). Abbreviations. ds: ductus seminalis connection to the bursa; la, lamella antevaginalis; lp, sclerite on the lamella postvaginalis; lt, lateral pocket; sf, ventral spinous field of segment 8. Scale bars: 2 mm (A); 200 µm (B, C, D, F).

Head: Vertex with long brownish scales protruding anteriorly and dorsally, fan-shaped, between antennae. Frons slightly convex covered with some whitish scales. Antennae dark brown, length ca. 0.5 as long as forewing costa, dorsally scaled, ventrally ciliated, two rows of scales per flagellomere. Palpus labialis porrect, length (all three segments combined) ca. 1.4 times diameter of compound eye, uniformly scaled; first segment short, slightly upcurved, with brown scales, second segment long, straight with mixed brown scales laterally and whitish scales dorsally, third segment short and slightly upcurved with whitish scales basally and apically and brown scales medially; opening of organ of vom Rath in apical position. Haustellum well developed. Ocelli and chaetosemata well developed.

Thorax: Upperside with pronotum, anterior half-part of mesoscutum, and tegulae covered by dark brown scales and posterior half-part of mesoscutum and metanotum covered by white scales; smooth-scaled including tegulae, without scales tufts. Underside, including legs, whitish, male foreleg hairpencil absent. Forewing length 5.0–6.6 mm (x̄ = 6.1; n = 19) in males, 5.8–7.3 mm (x̄ = 6.7; n = 7) in females. Forewing with typical venation of Cochylina, details described for the genus. Forewing pattern not sexually dimorphic (Fig. 2A). Forewing upperside with ground colour whitish with brownish-grey marking; most marks concentrated in costal area; system of pairs of strigulae vaguely recognisable, presumably concolourous with background, only through inter strigular dark marks; some scattered marks at basal fourth of costa; marks at level of Sc fused to produce a distinctive crescent shape brownish-grey blotch projected discally in a rather conspicuous coma-like patch confluent with R2–R3; single marks between Sc and R1, R1 and R2, R2 and R3, no marks beyond R3; some scattered grey scales between marks; striae strongly fragmented; dorsal marking ill-defined; fasciae undetectable; fringe ochreous; forewing underside uniformly brownish ochreous with some pale strigulae at radial level on the costa; overlapping area whitish. Hindwing upperside and underside, including fringe, uniformly brownish-ochreous; male costal fold absent; cubital pecten not detected.

Abdomen: Dorsad greyish, paler ochreous cephalad. Segment 8 unmodified. Male genitalia (based on four preparations; Fig. 2B, C, D, E) with tegumen well developed, laterally straight; uncus developed, basally confluent with top of tegumen, progressively narrowed distally; socii membranous, conspicuous, moderately developed, hairy; gnathos as two arms distally fused distally into a short process, moderately expanded distally in a central spinous molar-like sclerite; transtilla broad, strong, with a distal moderately flat area densely covered by short strong spines; valva elongate, variable in shape (Fig. 2B and C), costa slightly concave, sclerotised; cucullus moderately lobed, membranous, slightly sclerotised, with a central area densely hairy; sacculus internally concave, well sclerotised, distally projected in a finger-like structure, with triangular ventral subdistal process; pulvinus present; vinculum well developed; juxta strongly sclerotised in a rather pentagonal plate; ampulla present; phallus with coecum penis straight, central part strongly curved down, distal part straight, presence of dorsal teeth; vesica with two clusters of non-deciduous cornuti, one proximal oriented ventrally, consisting of aciculate cornuti, basally attached, arranged in a single longitudinal band, another distal consisting of an irregular patch of microspinulate cornuti. Segment 7 in females without modified scaling (corethogyne) but with two lateral, somewhat dorsal pockets on the 7-8 intersegmental membrane. Female genitalia (based on three preparations; Fig. 2F) with sterigma broad; lamella antevaginalis as a simple but evident lobe; ostium in a short funnel like antrum; lamella postvaginalis moderately sclerotised, smooth, broad, with a distinct ventrally prominent distal lobe as a transversal plate; ductus bursae as long as corpus bursae, moderately sclerotised in proximal half, double folded (in Z) when not extended; corpus bursae subspherical, moderately covered internally by acanthae and ctenidia in variable degree of development; no signum or other specially sclerotised area; a long ductus seminalis connected ventrally to cervix, no bulla seminalis; a large globular spermatophore extracted in one of the dissections; anterior apophysis fairly short, projected internally; ventral area of segment 8 behind the sterigma heavily covered by acanthae (spinous field) continuous with distal sclerotised plate of the lamella postvaginalis; a spiny star-shaped ventral subpapillar sclerite on the 8–9 intersegmental membrane at level of ventral lobes of anal papillae (Fig. 4B and D); posterior apophysis simple, approximately same length as papillae; egg pore broad between anal papillae.

The early stages are unknown. Adults were collected in January (n = 11) and November (n = 14) at middle elevations (1554-2150 m) in Bolivia, Santa Cruz Department, Florida Province in municipalities of Mairana, El Rasete, and Pampagrande, localities of Agua Clarita, Hueco de la Pascana, and La Hoyada. The collecting sites include transition from dry to cloud forest.

The specific epithet refers to the Quechuan word yun-ka, which translates as warm valley, a band of forest on the slopes of the Andes Mountains. This zone is of enormous interest from a conservation perspective.

Holotype: ♂, Venezuela, Aragua State, locality of Rancho Grande, 10°7'N; 67°20.63'W, 10–21 Feb 1969, D. Duckworth and E. Dietz (GS USNM 69274).

Paratypes: (4♀). Venezuela, Aragua State, locality of Rancho Grande, 1100 m, 10°7'N; 67°20.63'W, 24-31 Oct 1966 (1♀) (SEM stub JBA202); 22-31 Jul 1967 (3♀), R.W. Poole (GS USNM 85011).

The habitus of B. araguensis (Fig. 3A) has more extensive dark brown scaling in the wing pattern compared to B. yunkensis, resulting in a more diffuse and ill-defined pattern. A similar pattern is found in B. teneimorpha and B. caracagena. Species more closely related to B. yunkensis (e.g., B. baeza and B. uncicera) have a more defined, contrasting pattern. The forewing costal crescent-shaped blotch allows clear discrimination between the two species (well defined in B. yunkensis and diffuse in B. araguensis), but in the context of the genus, these differences could be assumed to represent variation. More diagnostic characters are associated with the male and female genitalia. Brusqeulia araguensis can be distinguished by the extremely narrow uncus, the narrowest in the genus, even compare to closely related species such as B. bonita and B. baeza. The transtilla and gnathos are well developed in B. araguensis, similar to most species in Brusqeulia, and it is not diagnostic. Teeth or lobes are developed in the distal part of the sacculus coincident with the ventral part of the cucullus in most, if not all, species of the genus. Among congeners, B. araguensis, B. costispina, and B. tripuncta all have several teeth, but their development in B. araguensis is moderate compared to the other two species. Finally, the phallus in B. araguensis is simpler than in most species of the genus. So far, morphological features of the females of Brusqeulia are limited by the paucity of material. The only females available are B. caracagena, B. yunkensis, and B. araguensis (the last two described in this paper). Both share a broad sterigma, but B. araguensis and B. yunkensis are definitely more closely related to each other than either is to B. caracagena, even though differences between them are conspicuous. Both B. caracagena and B. yunkensis lack the spiny cushion-like asymmetrical areas on the lamella antevaginalis found in B. araguensis. B. caracagena can be easily distinguished from B. araguensis and B. yunkensis by the ductus bursae, short in B. caracagena, long and convoluted in B. araguensis and B. yunkensis. The position of the ductus seminalis is clearly different in B. yunkensis (from cervix) and B. araguensis (from mid-corpus bursae); no information about the ductus seminalis in B. caracagena is available. The subpapillar spiny sclerite of the 8–9 intersegmental membrane is pointed in B. yunkensis and truncate in B. araguensis.

Head: Vertex with long whitish scales protruding anteriorly and dorsally, fan-shaped, between antennae. Frons slightly concave covered with a whitish scales. Antenna dark brown, length ca 0.4 as long as forewing costa, dorsally scaled, ventrally ciliated, two rows of scales per flagellomere. Labial palpus porrect, length (all three segments combined) ca. 1.3 times diameter of compound eye, uniformly scaled; first segment short, slightly upcurved with ochreous scales, second segment long, straight with ochreous scales, third segment short, slightly upcurved with a mixed of dominant ochreous scales and a few whitish scales only basally; opening of organ of vom Rath in apical position. Haustellum well developed. Ocelli and chaetosemata well developed.

Thorax: Dorsum whitish ochreous with a dorso-apical dark brownish band. Smooth scaled including tegulae, with no tufts. Legs whitish, unmodified, male foreleg hairpencil absent. Forewing length 5.7 mm (n = 1) in males, 5.7–6.2 mm (x̄ = 5.9; n = 4) in females. Forewing pattern (Fig. 3A) not sexually dimorphic. Forewing upperside general background colour whitish with scattered greyish-brown marks; marking ill defined; pairs of strigulae ill defined, concolourous with general background, vaguely detectable, with variable degree of suffusion; basal and subbasal fasciae poorly developed, median fascia as an irregular costal blotch projected tornally, with a small group of dark scales at the level of cubital cell; some coma-like marks on the costa as postmedian and preterminal fasciae; fringe concolourous with general background; forewing underside uniformly brownish ochreous with some pale strigulae on the costa; overlapping area whitish. Hindwing upperside and underside, including fringe, uniformly brownish-ochreous; male costal fold absent; cubital pecten not detected.

Abdomen: Dorsally greyish, pale ochreous cephalad. Segment 8 unmodified in males. Male genitalia (based on one preparation; Fig. 3C) with tegumen well developed, laterally straight; uncus slender, straight, basally confluent with top of tegumen to drastically slimmed distally; socii membranous, hairy, obvious, moderately developed; gnathos as two arms distally fused and projected in a short process distally spatulate; transtilla broad, naked; appreciable pulvinus, valva elongate, costa concave, moderately sclerotised, cucullus subrectangular, membranous ventrally, costal area slightly sclerotised, central area densely hairy, sacculus basally convex, distally concave, well sclerotised, transition area of sacculus to cucullus with several tooth like distal process, one of them larger and basal clearly associated to the sacculus, the distal one assignable to the cucullus, a variable number of smaller teeth in between; vinculum broad but rather weakly developed; juxta strongly sclerotised horseshoe shaped; phallus (Fig. 3D) (fragmented in three pieces in the slide) presumably straight with simple caecum, central part broken; no teeth detected on the external surface; vesica simple with two clusters of cornuti, one distal (vesica not evaginated) consisting of non-deciduous (not detected in female corpus bursae) cornuti arranged in a single longitudinal band, another proximal consisting in an irregular patch of microspinulate cornuti. Segment 7 in females without modified scaling (corethogyne) but with two inconspicuous laterodorsal pockets on the 7–8 intersegmental membrane. Female genitalia (based on two preparations; Fig. 3B) with sterigma broad, complex, slightly asymmetrical, ostium simple, slightly on the right; sterigma broad extended laterally in pockets ventrally covered by acanthae continuous laterally with two asymmetrical membranous cushion-shaped areas densely covered by acanthae (Fig. 4A); lamella antevaginalis with a moderately sclerotised convex plate; lamella postvaginalis moderately sclerotised, broad, with a distinct ventrally prominent but smooth dome like plate; ductus bursae rugose, sinuous, posterior half more sclerotised, internally covered by ctenidia continuous with internal vestiture of corpus bursae; corpus bursae subglobular, densely internally covered by ctenidia; no signum or any other sclerotised area detected; ductus seminalis from central area of corpus bursae; no bulla seminalis detected; no spermatophore found; anterior apophysis short projected internally; behind the sterigma the ventral area of segment 8 as a densely spiny lobe; 8–9 intersegmental membrane densely covered by acanthae; densely spiny crescent shape ventral sclerite on the 8–9 intersegmental membrane at the level of the ventral lobes of the anal papillae; posterior apophysis simple, approximately as long as anal papillae; presence of evident broad egg pore between anal papillae.

Figure 3. 

Morphological characters or Brusqeulia araguensis. A habitus (Paratype, female, Venezuela, Rancho Grande, 22–31 August 1967, USNM) B female genitalia (GS USNM85011) C male genitalia (GS USNM69274) D phallus (fragments photographically assemblage, may not correspond to the real order or orientation) (GS USNM6274). Abbreviations. ds: ductus seminalis connection to the bursa; la, lamella antevaginalis; lp, sclerite on the lamella postvaginalis; lt, lateral pocket; sf, ventral spinous field of segment 8; sp, subpapillar sclerite. Scale bars: 3 mm (A); 200 µm (B, C, D).

The early stages are unknown. Adults have been collected in February (n = 1), July (n = 2), August (n = 1), and October (n = 1) at middle elevation (1100 m) in Aragua State, Venezuela.

The specific epithet refers to the state of Aragua in Venezuela.

Figure 4. 

Female terminalia of Brusqeulia yunkensis and B. araguensis, ventral view, under scanning electron microscopy. A B. yunkensis B B. araguensis C subpapillar sclerite of B. yunkensis D same in B. araguensis. Abbreviations; la, lamella antevaginalis; lp, lamella postvaginalis; sf, ventral spinous field of segment 8; sp, subpapillar sclerite. Scale bars 100 µm.