Manerebia benigni tessmanni Pyrcz, 2004

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 : 230

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, Río Troya , 2100 m, 15.viii.2017 , P. Boyer leg., PBF; 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; 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 Andrés-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 Río 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 Río 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.

Figures

FIGURE 5. Adults (left—dorsum, right—venter) A. M. benigni tessmanni ♂ Ecuador, Río Troya B. M. benigni tessmanni ♀, Ecuador, Jimbura —San Andrés C. M. pauperata n. stat. ♂ Ecuador, Zamora —Chinchipe D. M. lamasi n. stat. ♂, Ecuador, Morona—Santiago

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.