Seven new taxa from the butterfly subtribe Euptychiina (Lepidoptera: Nymphalidae: Satyrinae) with revisional notes on Harjesia Forster, 1964 and Pseudeuptychia Forster, 1964 Author Nakahara, Shinichi Author Willmott, Keith R. Author Mielke, Olaf H. H. Author Schwartz, Johanna Author Zacca, Thamara Author Espeland, Marianne Author Lamas, Gerardo text Insecta Mundi 2018 2018-07-27 639 1 38 journal article 10.5281/zenodo.3708152 41a13689-dcc3-463c-941b-1f0ac97625aa 1942-1354 3708152 D0C05AD6-4F63-48C6-8A26-957AE9A920D2 Pseudodebis vrazi (Kheil, 1896) , new combination ( Fig. 2 , 3 , 16 ) Taygetis vrazi : Kheil 1896a: 151–152 , fig. TL: Río Atabapo [ Venezuela ] and Rio Negro [ Brazil ]. D’Abrera 1988: 754–755 , fig. [misidentified, represents Pseudodebis dubiosa Forster, 1964 ]. Table 1. GenBank accession numbers for sequences used in this study .

Voucher code	Genus	Species	Genes
			COI	EF1a	GAPDH	RPS5
MGCL-LOAN-050	Forsterinaria	pronophila	MH592919	MH592940	MH592957	MH592974
NW127-20	Forsterinaria	pronophila	DQ338797	DQ338941	GQ864975	GQ865444
CP02-57	Forsterinaria	pseudinornata	JQ392594	JQ392723	JQ392827	JQ392932
CP14-07	Forsterinaria	quantius	GQ864772	GQ864866	GQ864972	GQ865442
BC-DZ-Willmott-293	Harjesia	argentata	MH592922	MH592927	MH592944	MH592961
CP01-13	Harjesia	blanda	DQ338800	DQ338945	GQ357436	GQ357565
MGCL-LOAN-267	Harjesia	blanda	MH592913	MH592941	MH592958	MH592975
CP23-22	Harjesia	obscura	JQ392610	JQ392737	JQ392839	JQ392946
KW-140718-01	Megeuptychia	antonoe	MH592909	MH592930	MH592947	MH592964
LEP-19590	Neonympha	mitchelli	MH592912	MH592937	MH592954	MH592971
KW-15-001	Orotaygetis	surui	MH592920	MH592932	MH592949	MH592966
LEP-14787	Parataygetis	albinotata	MH592915	MH592935	MH592952	MH592969
CP04-09	Parataygetis	albinotata	JQ392615	JQ392741	JQ392843	JQ392950
CP01-06	Posttaygetis	penelea	JQ392620	JQ392745	—	JQ392955
KW-140708-01	Posttaygetis	penelea	MH592910	MH592928	MH592945	MH592962
CP22-02	Pseudodebis	celia	GU205874	GU205930	GU205988	GU206049
KW-140716-03	Pseudodebis	celia 01	MH592911	MH592929	MH592946	MH592963
PM01-23	Pseudodebis	marpessa	JQ392624	JQ392747	JQ392849	JQ392957
CP22-05	Pseudodebis	puritana	JQ392712	—	—	—
LEP-16938	Pseudodebis	valentina	MH592908	MH592936	MH592953	MH592970
CP01-94	Pseudodebis	valentina	JQ392630	—	—	—
KW-15-003	Pseudodebis	vrazi	MH592918	—	—	—
CP23-21	Sepona	punctata	JQ392607	JQ392734	JQ392838	JQ392943
MGCL-LOAN-490	Sepona	punctata	MH592917	MH592942	MH592959	MH592976
LEP-10084	Taygetina	banghaasi	MH592925	MH592933	MH592950	MH592967
KW-140719-01	Taygetina	gulnare	MH592914	MH592931	MH592948	MH592965
CP02-13	Taygetina	gulnare	JQ392613	JQ392740	JQ392842	JQ392949
MGCL-LOAN-022	Taygetina	kerea	MH592923	MH592939	MH592956	MH592973
PM02-04	Taygetina	kerea	JQ392645	JQ392763	JQ392866	JQ392976
CP22-01	Taygetina	peribaea	JQ392583	JQ392714	JQ392817	JQ392922
PM03-03	Taygetina	weymeri	JQ392708	JQ392814	JQ392918	JQ393027
MGCL-LOAN-003	Taygetina	ypthima	MH592924	MH592938	MH592955	MH592972
NW149-8	Taygetina	ypthima	GU205873	GU205929	GU205987	GU206048
BC-DZ-Willmott-015	Taygetis	chiquitana	MH592921	MH592926	MH592943	MH592960
PM04-12	Taygetis	laches	JQ392658	—	—	—
PM01-07	Taygetis	rufomarginata	JQ392679	—	—	—
LEP-10421	Taygetis	virgilia 01	MH592916	MH592934	MH592951	MH592968
PM01-03	Taygetis	larua	JQ392667	—	—	—

Taygetis Vrázi [sic]: Kheil 1896b: 230 . Taygetis valentina vrazi : Weymer 1910: 191 . Taygetis valentina var. vrazi : Gaede 1931: 434 . Taygetis rectifascia : Forster 1964: 65 , nota [as possible synonym of this species]. Harjesia vrazi : Lamas 1999: 142 , figs. 1–2; lectotype designation; Lamas 2004: 220 . Harjesia [n. sp.] Lamas MS ( Nymphalidae : Satyrinae 1369 ); Lamas 2004: 220 . Lectotype male. // LECTOTYPE ♂ Taygetis vrazi Kheil , 1896 By G. LAMAS, ‘99// atabapo [underlined] Tayg. vrázi // Mus. Nat. Pragae Inv. P5p/493/24// Genitalic vial SN-17-21 S. Nakahara // ( NMPC ) [examined] . Redescription. Male. Forewing length: 28mm ( n = 2). Head: Eyes with hair-like setae, with whitish scales at base; frons brownish, covered with whitish scales and hair-like scales; post-genal area with greyish hair-like scales and greyish scales; first segment of labial palpi adorned with white long hair-like scales and brown long hair-like scales ventrally, whitish scales laterally and dorsally, second segment in length almost twice as great as eye depth and covered with whitish scales and hair-like scales laterally, dorsally adorned with whitish hair-like scales, ventrally adorned with brown long hair-like scales and long white hair-like scales 3–4× as long as segment width, third segment about one-third of second segment in length and covered with brownish scales, with band of white scales laterally; antennae approximately two-fifths of forewing length, with approximately 36 antennomeres ( n = 1), distal 9–10 antennomeres composing club, distal 5 antennomeres appear darker, club not prominent. Thorax: Dorsally and laterally covered with long light greyish hair-like scales and light greyish scales; ventrally adorned with long whitish hair-like scales and whitish scales. Legs: Foreleg whitish, with tarsus, tibia, and femur similar in length; midleg and hindleg with femur whitish ventrally, tibia and tarsus greyish dorsally, ocher ventrally, tarsus and tibia adorned with spines ventrally, pair of tibial spurs present at distal end of tibia. Abdomen: Eighth tergite appears as thin sclerotized band at base of eighth abdominal segment; eighth sternite appears as a single plate. Wing venation: Basal half of forewing subcostal vein swollen; base of cubitus inflated; forewing recurrent vein absent; hindwing humeral vein developed; origin of M 2 nearer M 1 than M 3 . Male genitalia ( Fig. 3a, b ): Tegumen subtriangular in lateral view; uncus longer than tegumen in lateral view, narrow, sparsely adorned with hair-like setae at base, slightly curved in lateral view, tapered towards terminal point; brachia reduced, appearing as somewhat “ear-like” projections at posterior margin of tegumen; combination of ventral arms of tegumen and dorsal arms of saccus almost straight, broadens in lateral view near saccus; appendices angulares developed; saccus straight, similar to uncus in length; juxta shallow “V” shaped; in lateral view, narrow apical process of valva approximately one-third of valva length, valva broadens towards terminal point, almost twice in width, distal margin appears straight in lateral view, basal two-thirds of valva appear somewhat as parallelogram in lateral view, ventral margin slightly concave, costa rectangular, distal half of valva with hair-like setae; phallus roughly straight, phallobase about one-third of phallus in length, ductus ejaculatorius visible, manica covering about half of aedeagus, vesica visible. Female. Forewing length: 29mm ( n = 1). Similar to male except as follows: Foreleg whitish, foretarsus divided into five distinct tarsomeres; ground colour of both wing surface slightly lighter. Female genitalia ( Fig. 3 c–e): Lamella antevaginalis sclerotized, appearing as “finger-like” sclerotized plate projecting posteriorly; lateral side of 8th abdominal segment sclerotized, apparently fused to lamella antevaginalis at its anterior margin; posterior one-third of ductus bursae slightly sclerotized, origin of ductus seminalis located at anterior end of this slightly sclerotized region, anterior two-thirds of ductus bursae membranous; corpus bursae roughly oval in dorsal view, together with ductus bursae extending length of entire abdomen, with two signa located in middle, virtually extending length of corpus bursae, parallel to each other. Specimens examined. ( 3 ♂ , 1 ♀ ) Brazil : Rondônia : [ Jaru ], Santa Cruz da Serra ( Vitt, L. ), 7 Aug 1985 , 1 ♂ [FLMNH-MGCL-281490], ( MGCL ). Peru : Loreto : Río Samiria , Estación Biológica Pithecia , [ 5°11′S , 74°42′W ], 180 m , ( Pacheco, V. ), 7 Nov 1979 , 1 ♀ ( Genitalic vial SN-17-97 S. Nakahara ) [MUSM- LEP-102415], (MUSM); Madre de Dios : Tambopata Research Center , La Colpa , 13°09 ʹ S , 69°37 ʹ W , 250 m , ( Aibar, P. ), 15 Nov 1999 , 1 ♂ ( Genitalic vial SN-16-63 S. Nakahara ) [MUSM-LEP- 102426 ], ( MUSM ). Venezuela : Amazonas : Atabapo , [ 4°2 ʹ 31 ʺ N , 67°42 ʹ 27 ʺ W ], 1 ♂ (Genitalic vial SN-17-21 S. Nakahara ) ( NMPC ) . Systematic placement and diagnosis. Our molecular phylogeny ( Fig. 1 ) recovered Pseudodebis as a monophyletic group with high support (bs = 100), including the type species, Taygetis valentina (Cramer, 1779) , as sister to the remainder of the sampled Pseudodebis taxa. Two species formerly placed in Taygetomorpha Miller, 2004 ( P. celia (Cramer, 1779) and P. puritana (Weeks, 1902)) are found to be part of the monophyletic Pseudodebis . This is in accordance with Matos-Maraví et al. (2013) , where Taygetomorpha was synonymized under Pseudodebis . Pseudodebis vrazi n. comb. is placed as sister to P. puritana (Weeks, 1902) with a relatively high support (bs = 91). Its placement in Pseudodebis is also supported based on morphology, namely the reduction of the brachia. All other described species of Pseudodebis appear to have reduced (or almost absent) brachia, although the degree of reduction varies across species ( Forster 1964 ; SN pers. obs.). Pseudodebis vrazi n. comb. is readily distinguished from P. puritana by the following characters (in addition to others): 1) relatively small size (forewing length of P. puritana : 38–40 mm ( n = 3)); 2) VHW postdiscal band appears as a pair of very close, parallel lines in P. vrazi n. comb. (appears as a regular, single band in P. puritana ); 3) ocellus in VHW cell Cu 1 present with a pupil in a black area, ringed with orange in P. vrazi n. comb. (present as a creamy-white smudge in P. puritana ); 4) apical process of valva broadens at terminal point (in lateral view) in P. vrazi n. comb. (rather consistent in width in P. puritana ); 5) lamella antevaginalis appears as a “finger-like” sclerotized plate projecting posteriorly in P. vrazi n. comb. (appears as a broad, rather rectangular plate in P. puritana ). Pseudodebis vrazi n. comb. is also a lowland species, whereas P. puritana occurs in cloud forest habitats mostly above 1300 m . Forster (1964) considered Taygetis rectifascia Weymer, 1907 to be a possible synonym of Taygetis vrazi , although these two species are clearly not conspecific. Distribution. This species is known to date from four localities scattered across the Amazon basin in Venezuela ( Amazonas ), Brazil ( Rondônia ) and Peru ( Loreto and Madre de Dios ). Remarks. Until the present study, P. vrazi n. comb. was a rather enigmatic taxon known only from two specimens, the male lectotype from the Río Atabapo, Venezuela , housed in NMPC, and a female specimen from the Río Samiria, Peru , in the MUSM ( Lamas 1999 ). Kheil (1896a) described Taygetis vrazi based on the previously mentioned Venezuelan male lectotype and a paralectotype from Rio Negro, Brazil , although we were unable to locate this Brazilian specimen supposedly housed in MNHU ( Lamas 1999 ). Despite the fact that subsequent authors (e.g. Weymer 1910 ; Forster 1964 ) provided differing taxonomic opinions about this species, it remained in Taygetis for more than 100 years. Lamas (1999) reviewed its history of classification, designated a lectotype for T. vrazi , and reported the aforementioned Peruvian female for the first time. Moreover, Lamas (1999) transferred the species from Taygetis to the genus Harjesia Forster, 1964 , stating that the “male syntype held at NMPC shows conclusively that T. vrazi is a member of the genus Harjesia ”, although no further justification for this taxonomic change was provided. Subsequently, Lamas (2004) retained this systematic placement, and the species remained in Harjesia until now (e.g. Warren et al. 2017 ). Recently, during the course of examining the euptychiine collection at MUSM, SN discovered that one worn male specimen from La Colpa, Peru , identified as “ Harjesia sp. n. 4” (listed as one of the undescribed Harjesia species in Lamas (2004)) actually represents P. vrazi n. comb. based on comparison of the male genitalia of this specimen and the male lectotype . This male specimen from La Colpa, Peru , was sequenced (KW_15_003_ Pseudodebis _vrazi_01; see Fig. 1 ) to help clarify its systematic status, and we consequently regard this species as a member of Pseudodebis Forster, 1964 . Morphology also reinforces removal of the species from Harjesia since the type species of that genus, H. blanda ( Möschler, 1877 ) , does have a rather developed brachia unlike P. vrazi n. comb. and other Pseudodebis . In addition, a male specimen from Rondônia , Brazil , in MGCL was discovered as the only known specimen of P. vrazi n. comb. from that country, increasing the number of known specimens for this species to four.