Manerebia granatus Willmott, Radford & 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 : 216-223

Manerebia granatus Willmott, Radford & Pyrcz , n. sp.

( Figs. 4 , 6 )

Type locality. Ecuador, Zamora-Chinchipe Department, Cordillera del Cóndor, Destacamento Paquisha Alto

Diagnosis. This species superficially resembles a number of congeners, including its sister-species M. placida n. sp., with which it may be broadly sympatric. It differs from M. placida n. sp. as follows (characters in the latter species in parentheses): the VHW is uniform reddish brown in the distal half, including distally of the submarginal ocelli and white dots (rather than just basally of these markings); the VHW ocellus in cell CuA2-CuA1 is very small, with no distinct ocellar ring (larger with distinct yellowish orange ocellar ring); the VHW ocellus in M2-M1 is not developed (developed, with black centre and ocellar ring); the VHW submarginal red-brown line is more undulate (less undulate); the VHW has virtually no trace of a pale postdiscal line (variable but distinct line present). The species is also similar to other Manerebia that lack a VHW pale postdiscal line (e.g., Manerebia lamasi Pyrcz & Willmott, 2006 n. stat., Manerebia pauperata n. stat.), but may be distinguished from them by the very reduced VHW submarginal ocelli, especially that in cell CuA2-CuA1. Two males each of M. granatus n. sp. and M. placida n. sp. were dissected and they showed consistent differences between the two species, in particular the arms of gnathos parallel in dorsal view in M. granatus n. sp. but directed inwards in M. placida n. sp., and the spines at distal tip of valvae extending further basally along the inner edge in M. granatus n. sp., but confined to the distal tip in M. placida n. sp. The male genitalia of both species may be distinguished from the otherwise similar species M. benigni Pyrcz, 2004 as described below under M. placida n. sp.

Description. MALE ( Fig 4. A, B ): 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 20–21 mm, mean: 20.4 mm, n=8; 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 Cu2-Cu1 to M1-R5, first of these in centre of a small black spot surrounded by a reddish brown ring. HWV dark blackish brown (similar to basal half of FWV) basal to an indistinct, straight, dark brown postdiscal line, distally of this line reddish brown (also extending along tornus) up to a dark reddish brown, undulate submarginal line, then dark brown, with dark reddish brown marginal line; series of seven white submarginal dots in cells 2A-CuA2 (two dots), CuA2-CuA1 to M1-Rs, first three of these in centre of small black spots. Abdomen: Covered with dense, dark brown scales dorsally and laterally, and slightly paler greyish brown scales ventrally, with long dark hair-like scales increasing in density anteriorly. Genitalia ( Fig. 6A ): 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 8 or so squat ‘teeth’ oriented subhorizontally; aedeagus curving evenly upwards, tapering anteriorly, shorter than valva, smooth, no visible cornuti. FEMALE ( Fig. 4C ): Similar to male except: slightly larger (FW length 22 mm); wings slightly paler brown, no FWD androconial patch; reddish brown scales in distal part of HWV scattered, less dense than in male; HWV with narrow, even, straight white postdiscal band; submarginal ocelli larger in cells CuA2-CuA1 of FWV and HWV, and submarginal white spot in HWV cell surrounded by a small black spot. Genitalia (not illustrated): Anal papillae prominent, covered with setae of varying length, with a strongly sclerotized, amorphous basal plate; membrane below papillae a lightly sclerotized, grooved plate terminating in a point posteriorly, bordered anteriorly by a band of fine spine-like protrusions; postvaginal lamella sclerotized and forming a broad curved plate, produced into two prominent lateral, folded flaps with smooth edges; antevaginal lamella slightly sclerotized, pocket-like and folded; antrum sclerotized; ductus bursae wide and short, opening into a large, oval corpus bursae, with two narrow bands of dorsal signa that converge slightly anteriorly and extend over half of bursa length.

Molecular data. BI ( Fig. 12 ), ML ( Fig. 13 ) trees, species-delimitation analyses (Figs. 14,15), and genetic distances (Supplementary material 2) indicate that M. granatus n. sp. is a distinct species, and on both trees it clusters with M. placida n. sp. and a new, undescribed species from Central Peru, AZ-650 Manerebia sp. , the sisterspecies to M. granatus n. sp.

Type material: Ecuador: Holotype ♂: Zamora-Chinchipe , Destacamento Paquisha Alto, 3°54'36''S / 78°29'10''W, 2010 m, 31.viii.2010, K. Buckland leg., [PAN11; dissection, KW-20-018], FLMNH (to be deposited in INABIO) . Paratypes (7 ♂): Zamora-Chinchipe , Destacamento Paquisha Alto, 3°54'28''S / 78°29'5''W, 2100 m, 1.IX.2010, J. Radford leg., 1 ♂ [PAN52], FLMNH ; Destacamento Paquisha Alto , 3°54'28''S / 78°29'6''W, 2088 m, 31.viii.2010, K. Buckland leg., 1 ♂ [PAN13], FLMNH ; Destacamento Paquisha Alto , 3°54'36''S / 78°29'10''W, 2010 m, 3.IX.2010, K. Buckland leg., 1 ♂ [PAN69], FLMNH , 31.VIII.2010, 1 ♂ [PAN12; dissection KW-20-023], FLMNH, (Hartley, E.) , 3.ix.2010, 1 ♂ [PAN70], FLMNH , 31.viii.2010, J. Radford leg., 1 ♂ [PAN10], FLMNH ; Destacamento Paquisha Alto , 3°54'5''S / 78°28'54''W, 2299 m, 8.IX.2010, E. Hartley, 1 ♂ [PAN197], FLMNH ; PERU: 1 ♂: Amazonas , Alto Río Nieva, 2200–2500 m, vi.2002, B. Calderón leg., prep. genit. 2743, 25.06.2020 / K.Florczyk, H 397, red label saying: Manerebia benigni Pyrcz, 2004 , Paratype, det. T . Pyrcz; 1 ♂: Amazonas , R . De Mendoza , Qda. Llanohuaico, 1800–2000 m, 02.x.1998, B. Calderón leg., prep. genit. 2736, 24.06.2020 / K. Florczyk, prep. mol. CEPUJ 202006110 / K. Florczyk (lab), red label saying: Manerebia benigni Pyrcz, 2004 , Paratype, det. T . Pyrcz; 1 ♂: Amazonas , Alto Río Nieva, 2200–2500 m, VI .2002, B. Calderón leg., red label saying: Manerebia benigni tessmanni Pyrcz, 2004 , Paratype, det. T . Pyrcz .

Other specimens examined (not considered paratypes): ECUADOR: 1 ♀: Zamora-Chinchipe, km 4.3 San Andrés-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.

Etymology. The species name is the Latin word for garnet, in reference to the deep reddish brown colours of this species, and it is treated as a masculine noun in apposition.

Remarks. This species is externally most similar to M. placida n. sp., although there are a number of wing pattern differences as mentioned in the Diagnosis that are consistent in all examined specimens. Nevertheless, we initially considered that the two taxa might represent subspecies of a single species, given that no males have been collected in sympatry, and the lack of substantial differences in the dorsal androconial scales or male genitalia. However, the DNA barcode of a single female from the Jimbura-San Andrés road in southern Ecuador grouped with those of males of M. granatus n. sp. from the Cordillera del Cóndor, based on which it is tentatively considered as belonging to this species, although we exclude it from the type series. This female specimen also has reddish brown surrounding the HWV black ocellus in Cu2-Cu1, as in males of M. granatus n. sp. but in contrast to the yellowish brown ring around the ocellus in M. placida n. sp. and M. benigni . Assuming this female is conspecific with males from the Cordillera del Cóndor, M. granatus n. sp. may be relatively widespread in southern Ecuador, and perhaps even sympatric or locally elevationally parapatric with M. placida n. sp. However, so far no males are known from the area where the putative female of M. granatus n. sp. was collected. In addition, aside from the rather remarkable differences in wing pattern exhibited between the pierid Catasticta poujadei condor Radford & Willmott, 2013 , from the Cordillera del Cóndor, and the nominate subspecies in the adjacent Andes, there are few examples of butterfly species with different subspecies in these two regions. M. granatus n. sp. was also detected in northern Peru, in the highlands of Chachapoyas, where it was originally mistaken for M. benigni , and three among the known Peruvian specimens were actually included as paratypes of that species ( Pyrcz 2004). However, their genitalia match the specimens from the type locality of M. granatus n. sp. Finally, the divergence in DNA barcode between M. granatus n. sp. and M. placida n. sp. is comparable or even higher than that between other related Manerebia species. In summary, we consider that M. granatus n. sp. and M. placida n. sp. represent two distinct species.

Most known individuals of M. granatus n. sp. were collected at the type locality, a sandstone tepui in southeastern Ecuador, along a steeply climbing trail through cloud forest from 2010–2100 m. A single individual was collected on the top of the tepui in stunted elfin forest near 2300 m. A single female that may also represent this species (discussed above) was collected flying 1 m above the ground along the edge of a dirt road through cloud forest. Other congeners present at the type locality included M. benigni tessmanni Pyrcz, 2004 , 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.