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Manerebia huamanii Cerdeña & Pyrcz, n. sp. (Figs. 3, 8) Type locality. Peru, Cajamarca Department, Tres Ríos Diagnosis. This species might be confused, at first sight, only with the Colombian M. apiculata, which is of approximately the same size and shape in terms of the FW apex, and, to some extent, with the Ecuadorian M. interrupta. Both of these species apparently have individual forms with median half-moon shaped yellow patches. However, these patches in M. huamanii n. sp. are situated basally in relation to the postdiscal line, and distally in M. apiculata and M. interrupta, which makes the new species unmistakable. Considerable differences are apparent in the male genitalia, which show some similarities among M. huamanii n. sp., M. seducta, and M. ignilineata, including, among others, the stout uncus and massive basal parts of the valva. Molecular data are not available for M. huamanii n. sp. and no sister-species can be immediately identified, but its most closely related species should probably be looked for among those with the most similar genitalia. Description. MALE (Fig. 3C, D): Head: Eyes chocolate brown, naked; antennae naked, reaching 2/5 th length of costa, slender with club formed gradually, dorsally chestnut, ventrally sandy yellow, labial palpi two and a half times length of head, covered dorsally with short, ventrally with long, dense brown and sparse yellow hair-like scales. Thorax: Black, dorsally naked, legs brown, tibia covered with brown scales, femur and tarsus with sandy yellow scales. Wings: FW length: 18 mm (n=2); apex acute, distal margin slightly truncate below apex. HW oval with a slightly produced apex. FWD almost uniform medium brown, with a slightly darker basal half. FWD medium brown. FWV medium brown, a shade lighter in distal half, a faint reddish suffusion in submarginal area. HWV reddish brown, with a magenta overcast along outer margin distal to a wavy brown submarginal line, basally edged with grey; a median series of four, half-moon shaped sandy yellow patches with sharp outer edges contiguous to a faint postdiscal line. Abdomen: Dorsally covered with medium brown, ventrally with grey brown scales. Genitalia (Fig. 8B): Uncus long and arched; gnathos one third length of uncus, with a stout base and sharp apex; pedunculus small, directed downwards, saccus medium long, flattened dorso-ventrally; valva stout in basal half with a blunt dorsal process, sharply narrowing into an elongated apical one-third ending with a series of three sharp processes pointing inwards; aedeagus nearly straight, prominently widened in middle, slightly flattened dorso-ventrally, similar in length to valva. FEMALE: Unknown. Molecular data. Not available. Type material: Holotype ♂: Peru, Cajamarca, Tres Ríos, 6 o 56'41''S 78 o 52'40'' W, 3163 m, 20. viii.2010, E. Huamaní leg., MUSM; Paratype ♂: PERU, Piura, entre Las Minas y El Tambo, 2600–2900 m, 10–13.iv.1981, G. Lamas leg., MUSM. Etymology. This new species is dedicated to Erick Huamaní Villalobos, the collector of the holotype and a member of the Museo de Historia Natural Universidad Nacional de San Agustín de Arequipa (MUSA), in recognition of his invaluable contributions to different projects of the MUSA. The name is treated as a Latinized masculine noun in the genitive case. Remarks. The wing pattern characters, in particular the yellow patches situated basally in relation to the postdiscal line, and male genitalia, make this species unmistakable. The type locality lies on the western slopes of the Andes in the department of Cajamarca, half-way between two other localities where new species of Manerebia were discovered during this study, Abra de Porculla and Hacienda Udima (Fig. 16). The second known specimen, which is associated with this species, comes from a more northerly locality in Piura, some 250 km away, also situated on the western slopes of the Andes. We were, unfortunately, unable to examine its genitalia. Its size, wing shape and colour pattern match those of M. huamanii except for the lack of any HWV yellow patches. The presence or absence of HWV median patches or bands is a common, highly variable individual character in the genus Manerebia. Such variation apparently occurs, for example, in M. interrupta , M. apiculata , M. ignilineata (Dognin, 1896) and M. trimaculata (Hewitson, 1870), among other species.

Manerebia prattorum udima Pyrcz & Boyer, n. ssp. (Figs. 2, 7, 8, 11) Type locality. Peru, Cajamarca Department, 7 km S Hacienda Udima Diagnosis. The new subspecies differs from nominate M. prattorum in the paler, orange instead of reddishorange, and consistently wider HWD median band, whose outer edges are diffused and extend distally; it is also slightly larger than the nominate subspecies. Description. MALE (Figs. 2A, 11B): Head: eyes chocolate brown, naked, lustrous; labial palpi two times length of head, covered with black hair-like scales, longer ventrally, and some sparse yellow basal scales; antennae slender, 2/5 th length of costa, mostly naked, dorsally brown, ventrally chestnut, club formed gradually, ventrally sandy yellow. Thorax: black, mostly naked; legs covered with lustrous, brown and sandy yellow scales becoming dominant on tibiae and tarsi. Wings: FW length 15.5–17.5 mm, n=15, mean: 16.7 mm. FWD uniform lustrous dark brown. HWD lustrous dark brown, with long hair-like scales in basal part, crossed by an oblique, nearly straight, 3 mm wide, rich orange median band which is slightly wider in middle section and has a sharp inner and somewhat diffused distal edge, FWV dull, dark brown, slightly lighter distally, with a faint, irregular dark brown submarginal line. HWV dull dark brown, a shade lighter than forewing, with a dark brown cross-cell narrow band, a faint, sinuate dark brown submarginal line, and a pale yellow median band shaped as on dorsum. Abdomen: Dorsally and laterally covered with black, ventrally with grey brown scales. Genitalia (Fig. 7D): Uncus long, aligned with tegumen dorsum, with a blunt tip slightly curved downwards, gnathos short, one-third length of uncus, stout at base with a sharp tip; pedunculus prominent, stout, curved downwards; valva elongated with a prominent, blunt dorsal process in middle, and a narrow apical half, ending with four teeth pointing distally; saccus medium deep, narrow; aedeagus s-shaped, short, with a massive apical part. FEMALE (Figs. 2B, 11A): Sexual dimorphism is expressed in larger size of female (FW length: 18–19 mm, mean: 18.5 mm, n=4), and lighter ventral colour with prominent whitish and magenta distal suffusion. Genitalia (Fig. 8C): Anal papillae prominent, covered with long setae, with a strongly sclerotized basal plate, projecting basally and dorsally into short, sharp apophysis-like tips; membrane below papillae with a moderately sclerotized flange, postvaginal lamella moderately sclerotized and wide, produced into two prominent lateral, folded flaps; antevaginal lamella strongly sclerotized, arched, pocketlike; antrum strongly sclerotized; ductus bursae wide and short, opening gradually into a large, oval corpus bursae, with two parallel signa in dorsal position extending over two-thirds of bursa length. Molecular data: BI (Fig. 12) and ML (Fig. 13) trees, as well as species-delimitation methods (Figs. 14, 15), and genetic distances (Suplementary material 2) support the separate specific status of M. prattorum udima n. ssp. in relation to M. inderena clara, M. inderena ssp., and M. inderena antioquiana. Nevertheless, its phylogenetic position is not clear, because M. prattorum udima n. ssp. appears as sister species to M. inderena antioquiana in the BI analysis, and according to the ML method, it is the sister group to M. inderena clara and M. inderena ssp. Both analyses have a good branch support. The nominotypical subspecies of M. prattorum has not yet been barcoded. Etymology. This subspecies is named after the village of Hacienda Udima, situated just above the type locality. The name is treated as a feminine noun in apposition. Type material: Holotype ♂: Peru, Cajamarca, 7 km S Hacienda Udima, S06°50'25 W 70°06'14 2000 m, 14– 16.v.1982, G. Lamas & E. Pérez leg., MUSM; Paratypes (35 ♂ and 7 ♀): 15 ♂ and 4 ♀: Cajamarca, La Florida, vía Hacienda Udima, 1900–1950 m, 16.vi.2018, T. Pyrcz leg., CEPUJ; 19 ♂ and 3 ♀: Cajamarca, Bosque de Udima, la Florida, S 06°50'25 W 70°06'14, 2000 m 16.vi.2018, P. Boyer leg., PBF; 9 ♂ and 3 ♀: Cajamarca, La Florida, vía Hacienda Udima, 1900–1950 m, 16.vi.2018, leg. J. Farfán, MUSA; 2 ♂ and 1 ♀: Cajamarca, La Florida, vía Hacienda Udima, 1900–1950 m, 16.vi.2018, leg. J. Cerdeña, MUSA; 2 ♂: Cajamarca, 7 km S Hacienda Udima, 2000 m, 14–16.v.1982, G. Lamas & E. Pérez leg., MUSM. Remarks. We consider M. prattorum and M. inderena as specifically different, with the latter being most likely a complex of allopatric species, despite the fact that they cluster together on both the ML and Bayesian Inference trees, because of their highly different colour patterns, with that of M. prattorum in both the nominate and the new subspecies marked by a wide HWD orange band, which is extremely conspicuous when the butterfly is on the wing, and might play an important signaling role in mating. Also, the HWV median band of M. prattorum is two times as wide as in any subspecies of M. inderena. Finally, M. prattorum flies at lower elevations than M. inderena, occurring at some 1800–2000 m, whereas most subspecies of M. inderena occur at 2200–2600 m, and the nominate is found even as high as 2800 m. M. prattorum udima n. ssp. has been reported so far exclusively from the upper valley of the Río Zaña (Fig. 11). It is possible that it is actually endemic to that area, since it is found at approximately 2000 m above sea level and there are virtually no cloud forests northwards from the type locality on the western slopes of the Andes, as far as the area of Las Minas north of the La Porculla Pass, where the type locality of M. prattorum prattorum is situated.

Manerebia punku Pyrcz & Farf��n, n. sp. (Figs. 3, 7) Type locality. Peru, Piura Department, Abra de Porculla. Diagnosis. This species is immediately recognized from its congeners by the presence of prominent HWV postdiscal yellow spots, and the absence of any median band. No closest relative can be identified at this time. Description. MALE (Fig. 3A, B): Head: eyes chocolate brown, naked, lustrous; labial palpi two times length of head, covered with black hair-like scales, longer ventrally but overall shorter than in other congeners; antennae slender, 2/5 th length of costa, naked, dorsally and ventrally brown, except for ventral side of gradually formed club, which is sandy yellow. Thorax: black; legs dull brown, tibia covered with brown, femur and tarsus with sandy yellow scales. Wings: FW length: 17.5 mm (n=2). FWD almost uniform medium brown, with a slightly darker basal half. FWD medium brown, with hair-like scales in basal half. FWV medium brown, a shade lighter in distal half, with two, faint postdiscal and submarginal regular lines, and four small postdiscal, pale yellow dots, apparently varying individually with one individual lacking these dots (Fig. 3B). HWV medium brown, a shade lighter in distal half, with dark brown postbasal, postdiscal and submarginal lines, all rather irregular, with latter fainter than others, and a series of 5 postdiscal pale yellow dots, three central ones larger than extremal ones. Abdomen: dorsally covered with black scales, laterally and ventrally with grey brown scales. Genitalia (Fig. 7C): Uncus long and arched; gnathos one-third length of uncus, with a stout base and sharp apex; pedunculus prominent, directed downwards, saccus short, bulbous; valva slender, with a gradually narrowing apical part ending with a sharp tip turned inwards, with an irregular dorsum; aedeagus stout, sinuate, short. FEMALE: Unknown. Molecular data. BI (Fig. 12), ML (Fig. 13), the species-delimitation methods (Figs. 14,15) and genetics distances (Supplementary material 2) indicate that M. punku n. sp. is a distinct species. However, based on ML analysis, M. punku n. sp. is related to the clade formed by M. germaniae, M. apiculata, M. leaena + M. inderena mirena complex + M. golondrina, M. interrupta, M. inderena similis, M. inderena fina clade + M. trimaculata, M. undulata complex + M. inderena clara, M. inderena ssp., M. inderena antioquiana, M. prattorum udima n. ssp. clade, and according to BI analysis, it is related the M. mycalesoides and M. nevadensis, but both analyses showed high branch support of the clades containing it. Type material: Holotype ♂: Peru: Piura, Abra de Porculla, 05��50���42������ S / 79��30���15������ W, 15.vi. 2018, 2247 m, J. Farf��n leg. MUSA; Paratype (4 ♂): 1 ♂: same data as the Holotype, will be deposited in MUSM; 1 ♂: Piura, Abra de Porculla, western slopes, 2200���2250 m, 15.vi.2018, T. Pyrcz leg., prep. genit. 1613/ 24.01.2019 K. Florczyk, prep. mol. 392/ 26.06.2018 A. Zubek, CEPUJ; 1 ♂: Piura, Abra de Porculla, Chiclayo���Pucara S05��50���43��� W79��30���18���, 2200 m, 15.vi.2018, P. Boyer leg., PBF; 1 ♂: Piura, Pacaipampa, Bellavista, S04��57��� W79��33���, 1950 m, 25.vi.2003, W. Zelada leg., MUSM. Etymology. ���punku��� is a word in Quechua language (spoken currently in the Peruvian Andes) that means door, entry, access, portal, alluding to the Porculla pass that connects the two sides of the Andes at the type locality. Remarks. This species is easily distinguished from other congeners by its small size, rounded wings and, in the typical form, prominent HWV rounded yellow dots. The genitalia are also quite distinctive, being most similar to those of M. rufanalis Pyrcz & Hall, 2006 from southern Ecuador, which is otherwise externally very different. It is the only species of Manerebia known so far from the Abra de Porculla (Fig. 11C), where it occurs on the western slopes only, with the eastern slopes being extremely arid with no cloud forest vegetation. The geographic range of this species is incompletely known, and it can only be speculated that it extends both north and southwards where similar habitat is found. In the trees generated based on COI sequences this species is sister to M. apiculata, M. leaena, and M. germaniae, all externally different species occurring at higher elevations in the northern Andes., Published as part of Mahecha-J, Oscar, Florczyk, Klaudia, Willmott, Keith, Cerde��a, Jos��, Zubek, Anna, Boyer, Pierre, Farf��n, Jackie, Lachowska-Cierlik, Dorota & Pyrcz, Tomasz W., 2021, Solving the cryptic diversity of the genus Manerebia Staudinger in northern Peru description of new species and considerations on the biogeographical role of the Huancabamba Deflection (Nymphalidae: Satyrinae: Pronophilina), pp. 201-237 in Zootaxa 5072 (3) on pages 215-216, DOI: 10.11646/zootaxa.5072.3.1, http://zenodo.org/record/5744549, {"references":["Pyrcz, T. W., Willmott, K. R., Hall, J. P. W. & Viloria, A. L. (2006) A review of the genus Manerebia Staudinger (Lepidoptera: Nymphalidae: Satyrinae) in the northern Andes. Journal of Research on the Lepidoptera, 39, 37 - 79."]}

Manerebia lamasi Pyrcz & Willmott, 2006, n. stat. Manerebia satura lamasi Pyrcz & Willmott, 2006: 47. Type locality; Alfonso Ugarte, Cordillera del Cóndor, Amazonas, Peru. Holotype male: MUSM [examined]. Remarks. BI (Fig. 12), ML (Fig. 13), the species-delimitation methods, and genetics distances analysis (Supplementary material 2) indicate that the sequences of this taxon cluster in a single well-differentiated, long branch, and to represent the sister species to M. navarrae Adams & Bernard, 1979, from the Serranía de Perija in northern Colombia. Based these data we raise it to a specific rank. Manerebia lamasi n. stat. was originally described as a subspecies of Manerebia satura based on similar male genitalia and wing colour patterns. M. lamasi n. stat. is known so far exclusively from the Cordillera del Condór., Published as part of Mahecha-J, Oscar, Florczyk, Klaudia, Willmott, Keith, Cerdeña, José, Zubek, Anna, Boyer, Pierre, Farfán, Jackie, Lachowska-Cierlik, Dorota & Pyrcz, Tomasz W., 2021, Solving the cryptic diversity of the genus Manerebia Staudinger in northern Peru description of new species and considerations on the biogeographical role of the Huancabamba Deflection (Nymphalidae: Satyrinae: Pronophilina), pp. 201-237 in Zootaxa 5072 (3) on page 224, DOI: 10.11646/zootaxa.5072.3.1, http://zenodo.org/record/5744549, {"references":["Pyrcz, T. W., Willmott, K. R., Hall, J. P. W. & Viloria, A. L. (2006) A review of the genus Manerebia Staudinger (Lepidoptera: Nymphalidae: Satyrinae) in the northern Andes. Journal of Research on the Lepidoptera, 39, 37 - 79."]}

Manerebia placida Willmott & Pyrcz, n. sp. (Figs. 4, 6) Type locality. Ecuador, Zamora-Chinchipe Province, km 24 Loja-Zamora road, Quebrada San Francisco Diagnosis. This species superficially resembles a number of congeners, including M. granatus n. sp., as well as M. benigni. Diagnostic characters that distinguish it are discussed under the diagnosis of the former species. From M. benigni, M. placida n. sp. may be distinguished by not having a distinct darker postdiscal line immediately bordering the distal edge of the pale postdiscal line, and by the rings surrounding the HWV ocelli being paler yellowish brown, rather than dark reddish brown. The male genitalia of M. placida n. sp. differ from those of M. benigni as follows: its gnathos is shorter and more closely parallel to the uncus; the distal half of the valva is more elongate and narrower; the distal ���teeth��� of the valva are arranged more horizontally and within the same plane, whereas in M. benigni they point in different directions, are arranged more vertically and are less clustered together; and the aedeagus is more slender, less curved, and lacks a small median dorsal projection. Description. MALE (Fig. 4D): Head: eyes chocolate brown, naked, lustrous; labial palpi two times length of head, covered with black hair-like scales, longer ventrally; antennae slender, 2/5 th length of costa, 38 segments, mostly naked, dorsally brown, ventrally paler brown, club formed gradually of terminal 12 segments. Thorax: black, with long black hair-like scales; legs dorsally dark brown, ventrally pale yellowish brown. Wings: FW length 19���21 mm, mean: 20.4 mm, n=12; FWD uniform chocolate brown, except for patch of dense, black, elongate rectangular androconial scales in basal half to third of cells 2A-M1 and extending into adjacent posterior half of discal cell. HWD with hair-like scales in median half, uniform chocolate brown, with scattered reddish brown scaling in tornus. FWV dark brown, darker blackish brown in areas with dorsal androconial scales, with a well-marked dark reddish brown, undulating and rather irregular submarginal line, and a narrow reddish brown marginal line; series of five white submarginal dots in cells CuA2-CuA1 to M1-R5, first of these in centre of a small black spot surrounded by a pale yellowish brown ring. HWV dark blackish brown (similar to basal half of FWV) basal to a very narrow, approximately straight, white to cream postdiscal line, distally of this line reddish brown (also extending along tornus) up to series of submarginal white spots and ocelli (as described below), then dark brown, undulate dark reddish brown submarginal line and dark reddish brown marginal line; series of up to seven white submarginal dots in cells 2A-CuA2 (two dots), CuA2-CuA1 to M1-Rs, first three and last of these in centre of small black spots ringed with yellowish brown, largest spot in cell CuA2-CuA1, with dots/ocelli varying and sometimes absent in posterior half 2A-CuA2 and M1-Rs. Abdomen: Covered with dense, dark brown scales dorsally and laterally, and slightly paler grayish brown scales ventrally, with long dark hair-like scales increasing in density anteriorly. Genitalia (Fig. 6B): Uncus slightly curving and 1.5 times longer than tegumen shoulder, gnathos half-length of uncus, slightly curving upwards and pointed; pedunculus with a massive base and apex curved downwards; saccus short, bulbous; valva with a broad basal half ending in squared-off mid-dorsal process, and a narrower apical half with a distal series of 6���7 squat ���teeth��� oriented sub-horizontally; aedeagus curving evenly upwards, tapering anteriorly, shorter than valva, smooth, no visible cornuti. FEMALE: Unknown. Molecular data. BI (Fig. 12) and ML (Fig.13) trees, the species-delimitation methods (Figs. 14, 15), and genetic distances (Supplementary material 2) supports M. placida n. sp. as a valid species, and as sister species to M. granatus n. sp. These two species are also related to M. lamasi n. stat. and M. navarrae. Type material: Ecuador: Holotype ♂: Zamora-Chinchipe, km 24 Loja-Zamora rd., San Francisco, casa de Arcoiris, 3��59'18''S / 79��5'42''W, 2000���2050 m, 14.x.2006, K. R. Willmott & R. Aldaz leg., [FLMNH-MGCL-111813; dissection, KW-20-021], FLMNH (to be deposited in INABIO). Paratypes (17 ♂): Zamora-Chinchipe, km 24 Loja-Zamora rd., San Francisco, casa de Arcoiris, 3��59'18''S / 79��5'42''W, 2000 m, K. R. Willmott & R. Aldaz leg., 9 Oct 2006, 1 ♂ [FLMNH-MGCL-148383], FLMNH; km 24 Loja-Zamora rd., San Francisco, casa de Arcoiris, 3��59'18''S / 79��5'42''W, 2000���2050 m, K. R. Willmott & R. Aldaz leg., 10 Oct 2006, 1 ♂ [FLMNH-MGCL-111810], 1 ♂ [FLMNH-MGCL-111814], FLMNH, 11 Oct 2006, 1 ♂ [FLMNH-MGCL-111815], 1 ♂ [FLMNH-MGCL-148384; dissection, KW-20-020], FLMNH, 14 Nov 2006, 1 ♂ [FLMNH-MGCL-111823], FLMNH, 14 Oct 2006, 1 ♂ [FLMNH-MGCL-111811], 1 ♂ [FLMNH-MGCL-111817], 1 ♂ [FLMNH-MGCL-148385], FLMNH, 15 Oct 2006, 1 ♂ [FLMNH-MGCL-111812], 1 ♂ [FLMNH-MGCL-111819], 1 ♂ [FLMNH-MGCL-111822], 1 ♂ [FLMNH-MGCL-148386], FLMNH, 31 Oct 2006, 1 ♂ [FLMNH-MGCL-111816], 1 ♂ [FLMNH-MGCL-111818], FLMNH, 6 Nov 2006, 1 ♂ [FLMNH-MGCL-111820], 1 ♂ [FLMNH-MGCL-111821], FLMNH (one male to be deposited in CEPUJ). Etymology. The species name is a feminine Latin adjective in the nominative singular, placidus, meaning calm or gentle, partly in reference to the similarity of this species to M. benigni; although that species is named for the Peruvian collector Benigno Calder��n, the root of the name is the Latin adjective benignus, meaning kind. In addition, the name alludes to the gently undulating VHW submarginal line which somewhat distinguishes this species from M. benigni, in which the line is more strongly undulate. Remarks. M. placida n. sp. is closely related to M. granatus n. sp., and we discuss under that species our decision to recognize these two taxa as distinct species. This species is known to date only from the type locality in southeastern Ecuador, where it occurs in cloud forest from 2000���2100 m. Despite continuous sampling by handnetting and trapping at that locality from 16 September to 6 December in 2006, the species was only recorded from 10 October to 14 November, during which time it was not uncommon. Males were found puddling along open as well as shady streams from 10:00���12:00, as well as flying up until 14:15 within 1 m of the ground in the forest understorey along trails near streams. Two males were collected in traps baited with rotting fish, one in the understorey and one in the canopy, both near streams. Other congeners present at the type locality included M. inderena mirena Pyrcz & Willmott, 2006, M. rufanalis, M. pauperata n. stat., and M. trimaculata., Published as part of Mahecha-J, Oscar, Florczyk, Klaudia, Willmott, Keith, Cerde��a, Jos��, Zubek, Anna, Boyer, Pierre, Farf��n, Jackie, Lachowska-Cierlik, Dorota & Pyrcz, Tomasz W., 2021, Solving the cryptic diversity of the genus Manerebia Staudinger in northern Peru description of new species and considerations on the biogeographical role of the Huancabamba Deflection (Nymphalidae: Satyrinae: Pronophilina), pp. 201-237 in Zootaxa 5072 (3) on pages 223-224, DOI: 10.11646/zootaxa.5072.3.1, http://zenodo.org/record/5744549, {"references":["Pyrcz, T. W., Willmott, K. R., Hall, J. P. W. & Viloria, A. L. (2006) A review of the genus Manerebia Staudinger (Lepidoptera: Nymphalidae: Satyrinae) in the northern Andes. Journal of Research on the Lepidoptera, 39, 37 - 79."]}

Manerebia trimaculata (Hewitson, 1870) Lymanopoda trimaculata Hewitson, 1870: 159. Type locality: Ecuador, Morona-Santiago Province, St. Rosario. Syntype male: NHML [examined]. Remarks. Various sequenced individuals identified as M. trimaculata form a widely polytomic clade among morphologically noticeably different subspecies of M. inderena and M. undulata Pyrcz & Hall, 2006. According to BI and GMYC methods (Figs. 12, 15), the M. trimaculata complex contains three putative species that are morphologically different, and indeed some are sympatric, but based on ML and bPTP analyses (Figs. 13,14), the complex was not resolved and it might represent a single species with high phenotypic variation, and the genetic distances are not high (between 1���3%). Since there are still a number of questions to be resolved, we refrain at this stage from making any taxonomic changes regarding these taxa until more data are available. This issue will be discussed alonside the status of the subspecies of M. inderena in a forthcoming paper., Published as part of Mahecha-J, Oscar, Florczyk, Klaudia, Willmott, Keith, Cerde��a, Jos��, Zubek, Anna, Boyer, Pierre, Farf��n, Jackie, Lachowska-Cierlik, Dorota & Pyrcz, Tomasz W., 2021, Solving the cryptic diversity of the genus Manerebia Staudinger in northern Peru description of new species and considerations on the biogeographical role of the Huancabamba Deflection (Nymphalidae: Satyrinae: Pronophilina), pp. 201-237 in Zootaxa 5072 (3) on page 231, DOI: 10.11646/zootaxa.5072.3.1, http://zenodo.org/record/5744549, {"references":["Pyrcz, T. W., Willmott, K. R., Hall, J. P. W. & Viloria, A. L. (2006) A review of the genus Manerebia Staudinger (Lepidoptera: Nymphalidae: Satyrinae) in the northern Andes. Journal of Research on the Lepidoptera, 39, 37 - 79."]}

Manerebia inducta Pyrcz & Willmott, n. sp. (Figs. 2, 8, 9) Type locality. Peru, Cajamarca, Sanctuario Nacional Tabaconas-Namballe, Lagunas Arrebiatadas Diagnosis. This new species is distinguished from most of its congeners by the combination of two characters: a very narrow, continuous HWV white median band and an acute FW apex. Only two species of Manerebia possess the above features, M. seducta Pyrcz & Willmott, 2006, and M. apiculata (C. Felder & R. Felder, 1867), but the latter is apparently highly polymorphic with individual forms having a broken HWV pale band or no band at all. Furthermore, in both species the HWV marginal line is displaced further from the hindwing margin, and in M. apiculata it is very undulate. Manerebia seducta can also be recognized from M. inducta n. sp. by possessing a minute single black ocellus with a white pupil in both FWV and HWV cell CuA1-CuA2, which is represented in M. inducta n. sp. by a tiny white dot on the FWV only, and absent in M. apiculata. The shape of the HWV white median band is also slightly different in M. inducta n. sp. and M. seducta, in the former being slightly indented basally in cell M3-M2 and tapering noticeably at the costa. Finally, the FWV of M. inducta is distinctly blackish in the basal two-thirds, with a somewhat clear transition to a paler ground colour in the posterior half of the wing, whereas in M. seducta the basal area is not so dark, and the ground colour changes more uniformly from dark to pale towards the distal margin. Further differences are found in male genitalia, in particular in the valves, such as the presence of a prominent proturberance on the ampulla of M. inducta n. sp., or its much more massive apical part, terminated by a single stout process, which is bifurcate in M. seducta. Although no molecular data are available, external and genital morphology indicate that M. seducta is the likely sister-species of M. inducta n. sp.. Description. MALE (Fig. 2C): Head: frons with a tuft of brown hair-like scales; eyes chocolate brown, naked; labial palpi two and a half times length of head, covered with medium brown hair-like scales, ventrally long, dorsally short; antennae dorsally brown with blackish clubs, ventrally chestnut. Thorax: dorsally blackish, sparsely covered with brown hair-like scales, legs orange brown, tibia and femur covered with dense medium brown scales. Wings: FW length: 17.5 mm; triangular with an acute apex, straight outer margin. HW oval with straight outer margin and smoothly rounded tornus. FWD uniform medium brown. HWD almost entirely uniform medium brown except for some orange scaling along outer margin. FWV chocolate brown with a reddish sheen in basal half, progressively turning lighter, chestnut brown from postdiscal area, and along costal margin, a series of four, faint postdiscal milky white dots from M1-M2 to CuA1-CuA2 aligned parallel to outer margin, a faint, medium brown, slightly wavy submarginal line. HWV almost uniform chestnut brown with a delicate golden sheen, with a straight, narrow transverse milky white postdiscal line with sharp outer and slightly diffuse inner margin, running from costal to anal margin, a faint, darker brown, slightly irregular submarginal line, running close to distal margin, marginal area dusted with grey. Abdomen: dorsally blackish, laterally and ventrally chestnut. Genitalia (Fig. 8A): Tegumen slender, with a slightly arched dorsum, uncus similar in length to tegumen and stout, more prominently arched with a blunt tip, gnathos half width and length of uncus, curved dorsally, sharply tipped, pedunculus short and blunt, valva stout, sharply thinning at middle with an elongated apical part sharply terminated and curved dorsally with two dorsal teeth-like processes pointing inwards, aedeagus long, thin and smooth, with a prominent ���collar��� at junction of anterior and posterior part. FEMALE (Fig. 2D): As described and illustrated in Pyrcz et al. (2006) as female of Manerebia seducta. In this species, sexual dimorphism is particularly slight. Genitalia (Fig. 9B): Anal papillae prominent, covered with long setae, with a strongly sclerotized basal plate, irregularly rounded anteriorly without apophysis; postvaginal lamella moderately sclerotized and forming a broad plate, with a stepped, tube-like protrusion just posterior of ostium bursae; antevaginal lamella forming two deep, rounded ���pockets��� with thin, broad flanges dorsally on each side that curve dorsally at their inner edge and fuse anteriorly to form a rounded, inwardly directed ���keel��� just ventral of ostium bursae; antrum very broad, tapering, strongly sclerotized ventrally; ductus seminalis origin at anterior tip of antrum; ductus bursae wide and short, opening into an oval corpus bursae, with two parallel signa in dorsal position extending over one-half of bursa length. Molecular data. Not available. Type material: Holotype ♂: Peru, Cajamarca, Santuario Nacional Tabaconas-Namballe, Lagunas Arrebiatadas, 05 o 14���05������S, 79 o 17���18������W, 3122 m, 07.x.2009, Eric Huaman�� leg., MUSA, to be deposited in MUSM. Paratypes (2 ♀): 1 ♀ (Allotype of Manerebia seducta Pyrcz & Willmott, 2006): Ecuador, Loja, Km 20 Jimbura ��� San Andr��s road, 4��42'50''S, 79��26'16''W, 3300 m, 24.ix.1997, K. Willmott leg., FLMNH; 1 ♀: Peru, Cajamarca, Santuario Nacional Tabaconas-Namballe, Lagunas Arrebiatadas, 05 o 14���05������S, 79 o 17���18������W, 3122 m, 07.x.2009, Eric Huaman�� leg., MUSA. Etymology. This species name is a feminine Latin adjective meaning ���induced��� or ���exhibited���, and is an allusion to the confusion with the externally similar M. seducta. Remarks. This new species was formerly confused with M. seducta; whose type locality is Abiseo National Park in the north-central part of the Peruvian Andes (Pyrcz et al. 2006), but the two taxa differ sufficiently externally and in genitalia to be considered as distinct species. In particular, the male of M. seducta has visible, although small, single ocelli on the FWV and HWV, while no ocelli are present in M. inducta n. sp., and the HWV submarginal line of M. seducta is much farther away from distal margin than in M. inducta n. sp. The male genitalia, although presenting several common features, differ in the morphology of the valva, in particular the grooved dorsal surface in the basal half in M. seducta, which is smooth in M. inducta n. sp., and in the apical part, which is thinner and longer in M. seducta and terminating in three prominent ���teeth��� (although only two are visible on the original figure in Pyrcz et al. 2006), whereas only a single ���tooth��� is present in M. inducta n. sp. and it is located more basally. M. inducta is found along the Peru ��� Ecuador border in uppermost forest and in shrubby gulleys in the paramo where there is Chusquea, flying with M. ignilineata. It is notable that the type locality of M. inducta n. sp., the Lagunas Arrebiatadas area in the Tabaconas-Namballe sanctuary, also contains another endemic Pronophilina species, Pedaliodes namballe Pyrcz & Cerde��a (Pyrcz et al. 2013). Finally, the discovery of M. inducta n. sp. is a reminder of the potential pitfalls of designating paratypes from distant geographical localities, even if the specimens appear similar, if not identical, in external morphology., Published as part of Mahecha-J, Oscar, Florczyk, Klaudia, Willmott, Keith, Cerde��a, Jos��, Zubek, Anna, Boyer, Pierre, Farf��n, Jackie, Lachowska-Cierlik, Dorota & Pyrcz, Tomasz W., 2021, Solving the cryptic diversity of the genus Manerebia Staudinger in northern Peru description of new species and considerations on the biogeographical role of the Huancabamba Deflection (Nymphalidae: Satyrinae: Pronophilina), pp. 201-237 in Zootaxa 5072 (3) on pages 213-214, DOI: 10.11646/zootaxa.5072.3.1, http://zenodo.org/record/5744549, {"references":["Pyrcz, T. W., Willmott, K. R., Hall, J. P. W. & Viloria, A. L. (2006) A review of the genus Manerebia Staudinger (Lepidoptera: Nymphalidae: Satyrinae) in the northern Andes. Journal of Research on the Lepidoptera, 39, 37 - 79.","Pyrcz, T. W., Cerdena, J. & Villalobos, H. (2013) Systematics, bionomics and zoogeography of high Andean pedaliodines. Part 14: Two new species of Pedaliodes Butler from the Huancabamba Deflection in southern Ecuador and northern Peru (Lepidoptera: Nymphalidae: Satyrinae). Genus, 24, 131 - 141."]}

Manerebia ronda Pyrcz & Boyer, n. sp. (Figs. 1, 7, 9, 10) Type locality. Peru, Cajamarca, NE Bambamarca, Laguna Salahuindo Diagnosis. This species superficially resembles a number of congeners, including M. inderena (Adams, 1986), M. leaena (Hewitson, 1861), M. undulata Pyrcz & Hall, 2006 and M. germaniae, all of which have wide HWV yellow median bands, but only some of them do not have any ventral ocelli, in particular M. inderena leaeniva Pyrcz & Willmott, 2006, M. inderena similis Pyrcz & Willmott, 2006, and M. germaniae. The last species has a series of minute HWV submarginal yellowish dots, absent in M. ronda. The most similar taxon in colour pattern is M. inderena similis (Fig. 10B), whose HWV submarginal line is smooth and parallel to the outer margin, as opposed to the more undulating line in M. ronda n. sp. In terms of male genitalia morphology, the most similar species is M. pauperata n. stat., which differs in the larger tegumen which is apparent in lateral view, and thinner subuncus (Fig. 10A, E). The nominate subspecies of M. ronda n. sp. differs from M. ronda amplia n. ssp. by the more prominent teeth on the apical part of the valva (Fig. 10A, C). Description. MALE (Fig. 1A): Head: eyes chocolate brown, naked, lustrous; labial palpi two times length of head, covered with black hair-like scales, longer ventrally, and some sparse yellow scales; antennae slender, 2/5 th length of costa, mostly naked, dorsally brown, ventrally milky white, club formed gradually. Thorax: black, naked; legs with femur covered with chestnut scales, tibia and tarsus with sandy yellow scales. Wings: FW length 18���19 mm, mean: 18.4 mm, n=7; FWD uniform chocolate brown, lustrous, with a barely noticeable submarginal darker smooth line parallel to distal margin. HWD with long hair-like scales in median half, uniform chocolate brown, with a barely noticeable undulating darker submarginal line and yellow median band slightly translucent from venter. FWV dark brown, paler along costal and particularly distal margin due to thin overcast of whitish scales, with a well-marked blackish-brown, undulating and rather irregular submarginal line, a narrow barely visible marginal line, and a faint, dark brown straight, postdiscal line. HWV dark brown, lighter along distal margin where dusted with thin whitish scales, with a zigzagging dark brown submarginal line, a barely visible, narrow marginal line, and a wide, straight, yellow (slightly more intensely coloured towards anal margin) median band, of nearly same width throughout, and a hardly visible arched black discal line angled at almost 90��. Abdomen: Covered with dense, mostly brown scales dorsally and laterally, and sparse golden brown scales ventrally. Genitalia (Fig. 7A): Uncus arched and 1.5 times longer than tegumen shoulder, gnathos half-length of uncus, strongly uplifted with a short apex; pedunculus with a massive base and apex curved downwards; saccus short, bulbous; valva with a massive basal half ending in massive mid-dorsal process terminated by several short teeth, and a narrow apical half with a dorsal crest made up of five prominent teeth; aedeagus sinuate, shorter than valva. FEMALE (Fig. 1B): Sexual dimorphism marginal, female slightly lighter and duller brown on both upper and venter. Its HWV median band is sandy yellow. Genitalia (Fig. 9A): Anal papillae prominent, covered with rather sparse setae of varying length, with a strongly sclerotized basal plate, projecting basally and dorsally into short, sharp apophysis-like tips; membrane below papillae with a moderately sclerotized flange, postvaginal lamella sclerotized produced into two prominent lateral, folded flaps with smooth edges; antevaginal lamella slightly sclerotized, pocket-like; antrum strongly sclerotized with two protruberances; ductus bursae wide and short, opening gradually into a large, oval corpus bursae, with two parallel wide, dentate signa, running close to each other in ventral position extending over half of bursa length. Molecular data. BI (Fig. 12) and ML (Fig. 13) trees, as well as species-delimitation methods (Figs. 14, 15) and genetic distances (Sumplementary material 2) do not support the separate specific status of M. ronda n. sp., which is placed within the Manerebia pauperata n. stat. clade with zero or close to zero genetic distance. Type material: Holotype ♂: Peru, Cajamarca, NE Bambamarca, Laguna Salahuindo, S 06��36���897/ W78��26���159, 2700���2750 m, 11.vi.2018, T. Pyrcz leg., CEPUJ (to be deposited in MUSM); Paratypes (17 ♂ and 2 ♀:): 7 ♂: same data as the holotype, CEPUJ; 3 ♂: Cajamarca, ouest de Laguna Salahuindo, nord Bambamarca, S 06��36'897 W78��26'159, 2700 m, 11.vi.2018, P. Boyer leg., PBF; 7 ♂ and 2 ♀: Cajamarca, Bambamarca, La Ramada, 06��36���53������S / 78��26���09���W, 11.vi.2018. 2715 m, leg. J. Cerde��a and J. Farf��n, MUSA. Etymology. This species is named for a local political institution, a peasant patrol or meeting of local authorities in rural Peru, which was particularly active in the Bambamarca area during the insurgency of the Sendero Luminoso in the 1980s. It is treated as a feminine noun in apposition. Remarks. This species is externally most similar to M. inderena mirena but its male genitalia and COI barcode data (Figs. 12, 13, 14, 15) indicate its close affinities to M. pauperata n. stat. (Fig. 10E), which occurs at lower elevations and is externally markedly different, characterized by much large size, no HWV median yellow band, and large submarginal ocelli. Therefore, the two are considered as specifically distinct. Another species that has similar genitalia and somewhat similar colour patterns is M. germaniae, which occurs throughout Ecuador and Colombia, which can be recognized by the shorter distance between the base of the valva and the tip of the dorsal process, and by the shorter teeth on the apical part of the valva, as well as by the presence of HWV submarginal yellow dots, which are totally lacking in M. ronda n. sp. So far, the nominate subspecies of M. ronda n. sp. has been collected only in the north-central part of the department of Cajamarca, NW of Bambamarca, within the watersheds of western tributaries of the R��o Mara����n (Fig. 16)., Published as part of Mahecha-J, Oscar, Florczyk, Klaudia, Willmott, Keith, Cerde��a, Jos��, Zubek, Anna, Boyer, Pierre, Farf��n, Jackie, Lachowska-Cierlik, Dorota & Pyrcz, Tomasz W., 2021, Solving the cryptic diversity of the genus Manerebia Staudinger in northern Peru description of new species and considerations on the biogeographical role of the Huancabamba Deflection (Nymphalidae: Satyrinae: Pronophilina), pp. 201-237 in Zootaxa 5072 (3) on pages 205-206, DOI: 10.11646/zootaxa.5072.3.1, http://zenodo.org/record/5744549, {"references":["Pyrcz, T. W., Willmott, K. R., Hall, J. P. W. & Viloria, A. L. (2006) A review of the genus Manerebia Staudinger (Lepidoptera: Nymphalidae: Satyrinae) in the northern Andes. Journal of Research on the Lepidoptera, 39, 37 - 79."]}

Manerebia ronda amplia Pyrcz & Boyer, n. ssp. (Figs. 1, 7, 10) Type locality. Peru, Lambayeque Department, Ka��aris Diagnosis. This subspecies of M. ronda n. sp. is externally most similar to M. undulata milaena (Fig. 10D), from the western slopes of the Andes in the extreme south of Ecuador. Both are characterized by a milk-white and rather wide HWV median band gradually broadening from the costa to the anal margin, no trace of ocelli and rather smaller size compared to most other similar congeners such as M. germaniae Pyrcz & Hall, 2006, or M. inderena (Adams, 1986). M. undulata milaena has a magenta suffusion on both the FWV and HWV, not apparent in M. ronda amplia n. ssp. In addition, the submarginal dark brown line of the HWV of M. ronda amplia n. ssp. is more undulating than in M. undulata milaena, and the two differ markedly in male genitalia as described below. The subspecific status of M. ronda amplia n. ssp. and M. ronda ronda is strongly supported by COI data (Figs. 12, 13, 14, 15), and male genitalia (Fig. 7). Description. MALE (Fig. 1C): Head: eyes chocolate brown, naked, lustrous; labial palpi two times length of head, covered with black hair-like scales, longer ventrally, and with some sparse yellow scales; antennae slender, 2/5 th length of costa, dorsally brown, ventrally somewhat orange, club formed gradually. Thorax: black, dorsally mostly naked, with just some sparse hair-like scales along sides; femora sparsely covered with dark-brown scales, tibiae and tarsi densely with milky-white scales. Wings: FW length 17.5 mm, n=2. FW apex subacute, outer margin straight. HW oval with smooth outer margin. FWD uniform medium brown. HWD uniform medium brown with white median band slightly translucent from venter. FWV blackish brown gradually turning medium brown towards distal margin, with an undulating dark brown submarginal line. HWV chestnut, pale brown along distal margin, with a nearly straight milky white to pale yellow medium band, gradually broadening from costal to anal margin, and a delicately wavy submarginal dark brown line. Abdomen: dorsally and laterally covered with black, ventrally with grey-brown scales. Genitalia (Fig. 7B): Uncus arched and 1.5 times as long as tegumen shoulder, gnathos halflength of uncus, strongly uplifted with a short apex; pedunculus with a massive base and apex curved downwards; saccus short, bulbous; valva with a massive basal half ended with a massive mid-dorsal process terminated by several short teeth, and a narrow apical half with a dorsal crest made up of several (5���6) prominent teeth; aedeagus sinuate, shorter than valva. FEMALE: Unknown. Molecular data. BI (Fig. 12), ML (Fig. 13) tree as well as bPTP (Fig. 14) and GMYC (Fig. 15) do not support the separate specific status of M. ronda amplia n. ssp., which is placed alongside M. ronda n. sp. within the M. pauperata n. stat., clade with zero or close to zero genetic distance (Suplementary material 2). Type material: Holotype ♂: Peru, Lambayeque, Ka��aris, Bosque de Ka��aris, 06��03���43������S / 79��15���09������W, 14.vi. 2018, 2687 m, leg. J. Cerde��a, MUSA; Paratypes (6 ♂) 1 ♂: Lambayeque, Pucara v��a Ka��aris, 2700���2800 m, 14.vi.2018, T. Pyrcz leg., prep. genit. 1872/ 18.01.2019 K. Florczyk, prep. mol. 20190303 A. Zubek, CEPUJ; 1 ♂: Lambayeque, route de Ka��aris, S06��03'47 W79��14'21, 2400���2500 m, 14.vi.2018, P. Boyer leg., PBF; 1 ♂: Cajamarca, 10���15 km SE Hda. Udima, Qda. El Palmo, 2400���2600 m, 15.v.1982, G. Lamas & E. P��rez leg., MUSM; 1 ♂: Lambayeque, Geopampa, geographical coordinates (wrong), 2824���3135 m, 24.vii.2017, J. Grados, MUSM; 2 ♂: Peru, Lambayeque, 3,1km SSE Ka��aris, 06 o 04���28������S / 79 o 15���02������, 2824 m, 24.vii.2017, G. Espinoza & E. Gambos, MUSM. Etymology. The subspecies name is a feminine Latin adjective meaning ���wide���, in reference to the HWV median pale yellow band. Remarks. This subspecies is known so far only from a handful of specimens collected in the upper Chamaya river valley in an area where several other new species of Pronophilina have been discovered (within the genera Pronophila Doubleday, [1849], Lasiophila C. Felder & R. Felder, 1859, Pedaliodes Butler, 1867). When first collected, it was identified as M. undulata milaena, which is known from southernmost Ecuador, because of its extremely similar appearance, in particular the shape of the HWV median band. However, its male genitalia are very distinct from those of M. undulata milaena, bearing a close resemblance to those of M. pauperata n. stat., with an arched uncus, the basal projection of the valva and multiple crown-like teeth on apex. COI barcode data confirm their close affinity, as M. ronda amplia n. ssp., falls within the M. pauperata n. stat. clade. Externally, however, M. pauperata n. stat., is markedly different, being considerably larger, with prominent ventral ocelli and no yellow median band (Fig. 10E). So far, M. ronda amplia n. ssp. is known from two localities, in the western Peruvian Andes in the departments of Lambayeque and Cajamarca (Fig. 16), separated by some 100 km., Published as part of Mahecha-J, Oscar, Florczyk, Klaudia, Willmott, Keith, Cerde��a, Jos��, Zubek, Anna, Boyer, Pierre, Farf��n, Jackie, Lachowska-Cierlik, Dorota & Pyrcz, Tomasz W., 2021, Solving the cryptic diversity of the genus Manerebia Staudinger in northern Peru description of new species and considerations on the biogeographical role of the Huancabamba Deflection (Nymphalidae: Satyrinae: Pronophilina), pp. 201-237 in Zootaxa 5072 (3) on pages 206-207, DOI: 10.11646/zootaxa.5072.3.1, http://zenodo.org/record/5744549, {"references":["Pyrcz, T. W., Willmott, K. R., Hall, J. P. W. & Viloria, A. L. (2006) A review of the genus Manerebia Staudinger (Lepidoptera: Nymphalidae: Satyrinae) in the northern Andes. Journal of Research on the Lepidoptera, 39, 37 - 79."]}

Manerebia pauperata Pyrcz & Willmott, 2006, n. stat. Manerebia satura pauperata Pyrcz & Willmott, 2006: 47. Type locality: Km 40, road Loja – Zamora, Zamora-Chinchipe, Ecuador. Holotype male: CEPUJ [examined]. Remarks. In both BI (Fig.12) and ML (Fig. 13) trees M. pauperata n. stat. and M. satura situate in distant branches of the tree and cluster in highly resolved clades with other congeners. Both M. lamasi n. stat., and M. pauperata n. stat. have also been collected in sympatry in the Cordillera del Cóndor in southern Ecuador. Additionally, the two taxa also consistently differ in wing colour patterns (Fig. 5C, D) and male genitalia. We therefore formally recognize M. lamasi n. stat. as a distinct species. Despite an overall similar colour pattern in comparison with M. satura from central and southern Peru, with the exception of the absence of the HWV yellow median band, M. pauperata n. stat. shows some genitalic differences. Moreover, the species-delimitation methods (Figs. 14, 15) and genetic distances (Supplementary material 2) clearly support treating the two taxa as separate species.

Manerebia benigni tessmanni Pyrcz, 2004 (Figs. 5, 6) Manerebia benigni tessmanni Pyrcz, 2004: 495. Type locality: Abra Pardo Miguel, Amazonas, Peru. Holotype male: CEPUJ [examined]. Other specimens examined: ECUADOR: 1 ♂: Zamora-Chinchipe, Jimbura���Zumba, R��o Troya, 2100 m, 15.viii.2017, P. Boyer leg., PBF; 1 ♀: Zamora-Chinchipe, km 4.3 San Andr��s-Jimbura rd., [4��47'59''S, 79��18'18 ''W], 2020 m, 13.x.2010, K. R. Willmott leg., [FLMNH-MGCL-145842; dissection KW-20-022], FLMNH; 1♂: Morona-Santiago: km. 9.5 Chiguinda-Gualaquiza rd., river, [3��14'38''S, 78��40'15''W], 1650 m, (Willmott, K. R.), 11 Oct 2007, [FLMNH-MGCL-118316], (FLMNH), 12 Oct 2007, 1 ♂ [FLMNH-MGCL-118307], 1 ♂ [FLMNH-MGCL-118308], 1 ♂ [FLMNH-MGCL-118309], 1 ♂ [FLMNH-MGCL-118310], 1 ♂ [FLMNH-MGCL-118312], 1 ♂ [FLMNH-MGCL-118313], 1 ♂ [FLMNH-MGCL-118314], (FLMNH); San Martin, Chiguinda, [3��13'41''S, 78�� 41'59''W], 2030 m, (Willmott, K. R.), 11 Oct 2007, 1 ♂ [FLMNH-MGCL-118311], 1 ♂ [FLMNH-MGCL-118315], (FLMNH); Zamora-Chinchipe: Destacamento Paquisha Alto, [3��54'28''S, 78��29'5''W], 2100 m, (Radford, J.), 3 Sep 2010, 1 ♂ [PAN72; dissection, KW-20-017], (FLMNH); km 4.3 San Andr��s-Jimbura rd., [4��47'59''S, 79��18'18''W], 2020 m, (Willmott, K. R.), 13 Oct 2010, 1 ♀ [FLMNH-MGCL-145840], (FLMNH). Remarks. M. benigni tessmanni was described from the northern part of the highlands of Chachapoyas, from the Abra Pardo Miguel area specifically, and during the course of subsequent field work it has also been recorded in several localities in the southern Ecuadorian provinces of Morona-Santiago and Zamora-Chinchipe. In the ML and bPTP trees M. benigni sequences clustered in a highly resolved clade (Figs.13, 14), even if the only sampled specimen of M. benigni tessmanni from the type locality clusters in an external position in an internal clade of nominate M. benigni. This, in our opinion shows that the more northerly populations of M. benigni tessmanni are better differentiated on the molecular level from the nominate, which is logical from a geographical point of view, although they do not differ in colour patterns from the topotypical specimens of this subspecies. We illustrate two specimens from R��o Troya and Jimbura, male and female respectively (Fig. 5A, B), associated with this taxon based on male genitalia (Fig. 6C, D), and the fact that the two have matching HWV colour patterns, that are different from both M. granatus n. sp. and M. placida n. sp. Such a distribution pattern is not unfrequent among pronophiline butterflies at the subspecific level, as exemplified by Eretris porphyria transmaraniona Pyrcz 2004, which is also found in the two areas on the opposite sides of the R��o Chamaya valley (Pyrcz, 2004). However, according to BI and GMYC analyses (Figs. 12, 15), the sample DL- 457 M. benigni benigni appears as sister species to M. satura and the other samples of M. benigni, and it represents a separate species, as indicated by high genetic divergence (>3.7%) (Supplementary material 2). More samples from the Peruvian department of Amazonas are, however, needed in order to have a better insight on the relationships of the taxa within this clade., Published as part of Mahecha-J, Oscar, Florczyk, Klaudia, Willmott, Keith, Cerde��a, Jos��, Zubek, Anna, Boyer, Pierre, Farf��n, Jackie, Lachowska-Cierlik, Dorota & Pyrcz, Tomasz W., 2021, Solving the cryptic diversity of the genus Manerebia Staudinger in northern Peru description of new species and considerations on the biogeographical role of the Huancabamba Deflection (Nymphalidae: Satyrinae: Pronophilina), pp. 201-237 in Zootaxa 5072 (3) on page 230, DOI: 10.11646/zootaxa.5072.3.1, http://zenodo.org/record/5744549, {"references":["Pyrcz, T. W. (2004) Pronophiline butterflies of the highlands of Chachapoyas in northern Peru: faunal survey, diversity and distribution patterns (Lepidoptera, Nymphalidae, Satyrinae). Genus, 15, 455 - 622."]}

The species of the Otiorhynchus clavipes (Bonsdorff, 1785) group (Coleoptera: Curculionidae) treated here, are characterized by their unusually high phenotypical variation, which often caused taxonomic problems and controversies. Molecular markers COI and EF1-$\alpha$, karyological analysis, as well as morphological, biogeographical and ecological data are used to study weevils collected in the Alps, Carpathians, Sudetes and different areas of Germany. In the investigated populations of the flightless species O. fagi Gyllenhal, 1834 and O. clavipes, we detected an interspecific genetic distance of 11.3-15.8% (COI) and 3.1-3.7% (EF1-$\alpha$) depending on geographical distance. The phylogenetic trees indicate that both species are monophyletic and that they were correctly delimited from each other. Both species have also separate geographical ranges in Central Europe. Male specimens differ in the morphology of the aedeagus and the last abdominal sternite. Our study supports the legitimacy of species delimitation of O. fagi and O. clavipes as separate species, which can be treated as stable taxonomic hypotheses. The determination of the species status required the re-examination of spe cies ranges and allowed together with data on biology and altitudinal preferences a better biogeographical and ecological characterization of the species.

Bird schistosomes are commonly established as the causative agent of swimmer’s itch - a hypersensitive skin reaction to the penetration of their infective larvae. The aim of the present study was to investigate the prevalence of the genus Bilharziella in comparison to other bird schistosome species from Lake Drawsko - one of the largest recreational lakes in Poland, struggling with the huge problem of swimmer’s itch. In total, 317 specimens of pulmonate snails were collected and examined. The overall digenean infection was 35.33%. The highest bird schistosome prevalence was observed for Bilharziella sp. (4.63%) in Planorbarius corneus, followed by Trichobilharzia szidati (3.23%) in Lymnaea stagnalis and Trichobilharzia sp. (1.3%) in Stagnicola palustris. The location of Bilharziella sp. on the presented phylogeny showed that it is with high probability a different species than known so far B. polonica. Our finding complements the confirmed occurrence of bird schistosomes in European water bodies. Overall, presented research reveals the special importance of P. corneus as a source of the bird schistosome cercariae. This study suggests that the health threat connected with the blood flukes need to be further investigated by constant monitoring of their occurrence in intermediate hosts.

A survey of helminths associated with terrestrial slugs focusing on the invasive Arion vulgaris and the native A. ater was conducted on populations from France, Germany, Netherlands, Norway and Poland. In total, 648 terrestrial slugs were collected from 18 sample sites, and identified by means of morphological examination, dissection of genitalia and molecular analysis using mitochondrial DNA. In addition to A. vulgaris and A. ater, also A. vulgaris/A. rufus hybrids and A. ater/A. rufus hybrids were collected. Helminth species were identified based on morphological features and sequencing of the 18S and ITS rDNA regions. The parasites included four nematode species: Alloionema appendiculatum, Angiostoma sp., Phasmarhabditis hermaphrodita, Entomelas sp., two trematode species: Brachylaima mesostoma, Eurytrema sp., and one cestode (tapeworm) species: Skrjabinia sp. Alloionema appendiculatum was the most common helminth in the investigated slug populations. Furthermore, we found higher prevalence of trematodes in the invasive A. vulgaris compared with the native A. ater, while differences in the prevalence for nematodes were not as clear. publishedVersion

Class EUTARDIGRADA Richters, 1926 Order PARACHELA Schuster, Nelson, Grigarick and Christensen, 1980 Superfamily HYPSIBIOIDEA Pilato, 1969 (in Marley et al. 2011) Family HYPSIBIIDAE Pilato, 1969 Subfamily ITAQUASCONINAE Rudescu, 1964 Genus Mesocrista Pilato, 1987, Published as part of G��siorek, Piotr, Stec, Daniel, Morek, Witold, Zawierucha, Krzysztof, Kaczmarek, ��ukasz, Lachowska-Cierlik, Dorota & Michalczyk, ��ukasz, 2016, An integrative revision of Mesocrista Pilato, 1987 (Tardigrada: Eutardigrada: Hypsibiidae), pp. 2803-2828 in Journal of Natural History (J. Nat. Hist.) (J. Nat. Hist.) 50 on page 2809, DOI: 10.1080/00222933.2016.1234654, http://zenodo.org/record/5189370, {"references":["Richters F. 1926. Tardigrada. Handbuch Der Zoologie. III: 58 - 61.","Pilato G. 1969. Evoluzione e nuova sistemazione degli Eutardigrada. Boll Zool. 36: 327 - 345.","Marley NJ, McInnes SJ, Sands CJ. 2011. Phylum Tardigrada: A re-evaluation of the Parachela. Zootaxa. 2819: 51 - 64.","Rudescu L. 1964. Tardigrada. Fauna Republicii Populare Romine IV: 7 - 308."]}