Manerebia placida Willmott & Pyrcz, 2021

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), Zootaxa 5072 (3), pp. 201-237 : 223-224

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 Calderón, 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 .

Figures

FIGURE 4. Adults (left—dorsum, right—venter) A. M. granatus n. sp. ♂ Holotype, Ecuador, Paquisha Alto B. M. granatus n. sp. ♂ Paratype, Peru, Alto Río Nieva C. M. granatus n. sp.♀, Ecuador, San Andrés D. M. placida n. sp. ♂ Holotype, Ecuador, Zamora - Arcoiris

FIGURE 6. Male genitalia (top—lateral view, middle—plan view, bottom—aedeagus extracted, lateral view) A. M. granatus n. sp. Peru, Rodríguez de Mendoza, prep. genit. KF-2736 B. M. placida n. sp. Ecuador, Zamora, prep. genit. KW-20-021 C. M. benigni tessmanni, Peru, Alto Río Nieva, prep. genit. KF-H_241 D. M. benigni tessmanni, Ecuador, Río Troya, prep. genit. KF-6282

FIGURE 12. Bayesian Inference tree (BI) reconstructed from a fragment of COI mitochondrial gene. Posterior probability values indicated on nodes.

FIGURE 13. Maximum likelihood tree (ML) reconstructed from a fragment of COI mitochondrial gene. Ultrafast bootstrap support values indicated on nodes.

FIGURE 14. The Bayesian implementation of the Poisson Tree Processes (bPTP) for delimiting species based on a fragment of COI mitochondrial gene.

FIGURE 15. The generalized mixed Yule-coalescent (GMYC) for delimiting species based on a fragment of COI mitochondrial gene.