Identification and taxonomy.

Taxon classification

Animalia

Nymphalidae

Nakahara

Shinichi

Lamas

Gerardo

Tyler

Stephanie

Marín

Mario Alejandro

Huertas

Blanca

Willmott

Keith R.

Mielke

Olaf H. H.

Espeland

Marianne

A revision of the new genus

Amiga

Nakahara, Willmott & Espeland, gen. n., described for

Papilio

arnaca

Fabricius, 1776 (

Lepidoptera

Nymphalidae

Satyrinae

)

Zookeys3112019201982185152

10.3897/zookeys.821.31782

Amiga

arnaca

sericeella

(Bates, 1865), comb. n. et

stat. n.

Figs 2m, n, 6

Euptychia

sericeella

Bates (1865: 202). Type locality: Vera Paz, Guatemala. Lectotype (here designated) ♀: Godman-Salvin Coll. 1904.–1. B.C.A. Lep. Rhop.

Euptychia

sericeella

, Bates. // B.M. TYPE No. Rh 3181

Euptychia

sericeella

, ♀ Bates. // Forests of N. Vera Paz. F.D.G. & O.S. // Type. Sp. Figured. // ♀ // (NHMUK) [examined].

Euptychia

sericeella

: Butler 1867: 489; Kirby 1871: 53; Butler 1877: 122; Godman and Salvin 1880–1881: 89–90, pl. 8, figs 20, 21; Weymer 1911: 219; Riley and Gabriel 1924: 53; Gaede 1931: 464; D’Abrera 1988: 770–771.

Chloreuptychia

sericella

[sic]: Forster 1964: 120–121, fig. 132.

Chloreuptychia

sericeella

: R. de la Maza and J. de la Maza 1993: 182; Salinas-Gutiérrez et al. 2004: 136; Lamas 2004: 218.

Identification and taxonomy.

Bates (1865) described Euptychiasericeella based on an unspecified number of “male” specimen(s) from Vera Paz, Guatemala. Nevertheless, the only syntype specimen that we have located, in the NHMUK, is a female, a fact also noted by Godman and Salvin (1880: 89–90), who had this specimen in their possession and referred to it as “our specimen, marked as the type”. This specimen (B.M. TYPE No. Rh 3181) was also referred to as a type by Riley and Gabriel (1924: 53) and D’Abrera (1988: 770–771). Because of the complexity of euptychiine taxonomy, to provide an unambiguous reference for this name we here designate this specimen as the lectotype of Euptychiasericeella (lectotype designation). Bates explicitly stated that the taxon was closely allied to Eu[ptychia]. ebusa (= Amigaarnacaarnaca comb. n.), indicating that he clearly regarded Euptychiasericeella and Amigaarnacaarnaca as two different species. Despite the ambiguous diagnosis provided in the original description (“the fore-wing having a narrow costal border, the apex, and a broader outer border of a brown hue”), the lectotype, figured in Warren et al. (2018), exhibits several rather distinctive phenotypic differences compared to specimens from South America and Nicaragua to western Ecuador. Presumably due to these wing pattern differences, described further below, subsequent authors treated E.sericeella as a species-level taxon (e.g., Butler 1867, 1877; Weymer 1911; Gaede 1931; Forster 1964; Lamas 2004). Specimens from Zelaya department, Nicaragua (e.g., FLMNH-MGCL-263066, 263067, 263072) are phenotypically similar to specimens known from Costa Rica to western Ecuador, and A.a.sericeella thus seems to replace A.a.adela n. ssp. with little or no obvious gap or dispersal barrier separating these taxa, and conversely with no known area of sympatry. Furthermore, the presence of an iridescent bluish-lilac reflection between the postdiscal band and umbra in some specimens of the subspecies from Nicaragua, Costa Rica, and Panama might be a result of introgression with A.a.sericeella. Finally, divergence in the DNA barcode between a specimen from Honduras (LEP-16997) phenotypically similar to the lectotype of sericeella and other Central American A.arnaca from further south is no greater than between east and west Andean A.arnaca (Table 2). Based on these considerations, we downgrade E.sericeella to subspecific rank, proposing, for the first time, what we believe to be the most parsimonious hypothesis of a single species. This null hypothesis remains to be more strongly tested when new evidence, such as distributional, behavioral, or genetic data, becomes available.

Both sexes of A.arnacasericeella comb. n. et stat. n. differ from the nominotypical subspecies in the following respects: DHW feeble pearly reflection restricted to posterior one-third of hindwing and not extending into discal cell; ventral ground color somewhat paler; discal band, postdiscal band, submarginal band on the ventral forewing and ventral hindwing narrower; ventral hindwing postdiscal band not bent inwards in cell Cu₂ and gently curving towards inner margin (but see also below); ventral ocelli smaller (but see also below); bluish-lilac reflection on ventral surface appearing purplish.

Variation.

The ventral ocelli are variable in terms of size; while many specimens seem to possess ventral ocelli smaller than the nominotypical subspecies and A.arnacaindianacristoi ssp. n., some appear to have ocelli that are similar to the aforementioned two taxa in terms of size. The VHW postdiscal band is bent inwards in cell Cu₂ in some specimens, whereas it gradually curves towards the inner margin in other specimens.

Distribution

(Fig. 6). This subspecies ranges from southern Mexico to Honduras, where it appears to be uncommon.

Examined specimens

(46 ♂, 23 ♀). See Appendix for the data of these specimens.

Figure 2.

Amigaarnaca spp. specimens spanning its range (dorsal on left, ventral on right): a nominotypical subspecies from Suriname, neotype male (USNM) b nominotypical subspecies from Guyana, female (FLMNH-MGCL 263373) c nominotypical subspecies from E Ecuador, male (FLMNH-MGCL 257121) d nominotypical subspecies from Peru, female (FLMNH-MGCL 262953); e nominotypical subspecies from N Brazil, male (FLMNH-MGCL1036223) f Nominotypical subspecies from N Brazil, female (FLMNH-MGCL 207984) g Nominotypical subspecies from SE Brazil, male (FLMNH-MGCL 1036213); h nominotypical subspecies from SE Brazil, female (FLMNH-MGCL 1036218) iA.arnacaadela from Costa Rica, male (FLMNH-MGCL 207991) jA.arnacaadela from Costa Rica, female (FLMNH-MGCL 207992) kA.arnacaadela from W Ecuador, holotype male (FLMNH-MGCL 151127) lA.arnacaadela from W Ecuador, female (FLMNH-MGCL 257087) mA.arnacasericeella, male from Mexico (FLMNH-MGCL 207900) nA.arnacasericeella from Mexico, female (FLMNH-MGCL 207896) oA.arnacaindianacristoi from NW Venezuela, paratype male (FLMNH-MGCL 263107) pA.arnacaindianacristoi from N Venezuela, paratype female (FLMNH-MGCL 1036235).

Figure 6.

Distribution map of Amigaarnaca ssp.

Table 2.

Genetic distances calculated based on the Tamura-Nei model.

		1	2	3	4	5	6	7	8	9	10	11	12	13	14	15	16	17	18	19	20	21	22	23	24	25	26	27	28	29	30	31	32	33	34	35	36
1	02-SRNP-5948 Amigaarnacaadela (Costa Rica)		0.003	0.076	0.073	0.067	0.061	0.064	0.058	0.024	0.024	0.024	0.058	0.061	0.058	0.058	0.044	0.073	0.064	0.064	0.064	0.024	0.024	0.049	0.073	0.03	0.03	0.067	0.067	0.033	0.033	0.03	0.033	0.033	0.034	0.03	0.033
2	07-SRNP-100105 Amigaarnacaadela (Costa Rica)	0.003		0.063	0.065	0.058	0.058	0.074	0.058	0.026	0.026	0.027	0.057	0.06	0.056	0.062	0.043	0.061	0.056	0.059	0.056	0.026	0.026	0.049	0.062	0.029	0.029	0.059	0.061	0.028	0.028	0.027	0.028	0.028	0.03	0.026	0.028
3	BC-DZ_Willmott-055 Amigaarnacaarnaca (Maranhao, Brazil)	0.076	0.063		0.005	0.022	0.032	0.034	0.033	0.06	0.06	0.062	0.032	0.033	0.03	0.037	0.055	0.006	0.027	0.029	0.027	0.061	0.061	0.052	0.003	0.058	0.058	0.025	0.024	0.058	0.058	0.055	0.058	0.058	0.057	0.056	0.058
4	BC-DZ_Willmott-056 Amigaarnacaarnaca (Para, Brazil)	0.073	0.065	0.005		0.024	0.032	0.034	0.031	0.06	0.06	0.06	0.03	0.032	0.03	0.035	0.053	0.006	0.027	0.024	0.024	0.061	0.061	0.049	0	0.053	0.053	0.02	0.019	0.059	0.056	0.057	0.059	0.059	0.058	0.057	0.059
5	BC-DZ_Willmott-057 Amigaarnacaarnaca (Espirito Santo, Brazil)	0.067	0.058	0.022	0.024		0.029	0.022	0.028	0.063	0.062	0.062	0.026	0.021	0.026	0.028	0.05	0.022	0.017	0.019	0.017	0.061	0.061	0.059	0.022	0.058	0.058	0.006	0.005	0.056	0.056	0.055	0.056	0.056	0.059	0.054	0.056
6	CP06-76 Amigaarnacaarnaca (Amazonas, Peru)	0.061	0.058	0.032	0.032	0.029		0.027	0.006	0.059	0.059	0.059	0.002	0.005	0.002	0.006	0.046	0.031	0.022	0.024	0.023	0.06	0.06	0.042	0.031	0.055	0.055	0.027	0.029	0.056	0.056	0.051	0.056	0.056	0.055	0.054	0.056
7	DNA99-015 Amigaarnacaarnaca (Napo, Ecuador)	0.064	0.074	0.034	0.034	0.022	0.027		0.023	0.073	0.073	0.073	0.026	0.023	0.025	0.031	0.055	0.034	0.011	0.011	0.011	0.075	0.075	0.077	0.034	0.072	0.072	0.022	0.022	0.075	0.075	0.072	0.075	0.075	0.075	0.072	0.075
8	LEP-04408 Amigaarnacaarnaca (Morona-Santiago, Ecuador)	0.058	0.058	0.033	0.031	0.028	0.006	0.023		0.06	0.06	0.056	0.002	0.005	0.002	0.01	0.043	0.031	0.023	0.023	0.021	0.06	0.06	0.039	0.031	0.054	0.054	0.026	0.028	0.056	0.056	0.053	0.056	0.056	0.057	0.055	0.056
9	LEP-09788 Amigaarnacaadela (Guayas, Ecuador)	0.024	0.026	0.06	0.06	0.063	0.059	0.073	0.06		0	0	0.059	0.061	0.057	0.064	0.041	0.06	0.061	0.061	0.061	0	0	0.036	0.06	0.014	0.014	0.061	0.063	0.015	0.015	0.012	0.015	0.015	0.014	0.013	0.015
10	LEP-09930 Amigaarnacaadela (Pichincha, Ecuador)	0.024	0.026	0.06	0.06	0.062	0.059	0.073	0.06	0		0	0.059	0.061	0.057	0.064	0.041	0.06	0.061	0.061	0.061	0	0	0.036	0.06	0.014	0.014	0.061	0.062	0.015	0.015	0.012	0.015	0.015	0.014	0.013	0.015
11	LEP-09931 Amigaarnacaadela (Esmeraldas, Ecuador)	0.024	0.027	0.062	0.06	0.062	0.059	0.073	0.056	0	0		0.058	0.061	0.057	0.064	0.041	0.06	0.059	0.059	0.059	0	0	0.036	0.06	0.013	0.013	0.06	0.062	0.015	0.015	0.012	0.015	0.015	0.014	0.013	0.015
12	LEP-10696 Amigaarnacaarnaca (Zamora-Chinchipe, Ecuador)	0.058	0.057	0.032	0.03	0.026	0.002	0.026	0.002	0.059	0.059	0.058		0.003	0	0.005	0.045	0.03	0.021	0.021	0.021	0.059	0.059	0.039	0.03	0.053	0.053	0.025	0.026	0.055	0.055	0.05	0.055	0.055	0.054	0.054	0.055
13	LEP-10697 Amigaarnacaarnaca (Pastaza, Ecuador)	0.061	0.06	0.033	0.032	0.021	0.005	0.023	0.005	0.061	0.061	0.061	0.003		0.003	0.005	0.046	0.032	0.018	0.018	0.018	0.062	0.062	0.046	0.032	0.055	0.055	0.02	0.021	0.058	0.058	0.053	0.058	0.058	0.057	0.056	0.058
14	LEP-10703 Amigaarnacaarnaca (Zamora-Chinchipe, Ecuador)	0.058	0.056	0.03	0.03	0.026	0.002	0.025	0.002	0.057	0.057	0.057	0	0.003		0.005	0.044	0.03	0.023	0.023	0.023	0.058	0.058	0.039	0.03	0.053	0.053	0.025	0.026	0.054	0.054	0.049	0.054	0.054	0.052	0.052	0.054
15	LEP-15082 Amigaarnacaarnaca (Morona-Santiago, Ecuador)	0.058	0.062	0.037	0.035	0.028	0.006	0.031	0.01	0.064	0.064	0.064	0.005	0.005	0.005		0.051	0.035	0.023	0.023	0.023	0.064	0.064	0.042	0.035	0.058	0.058	0.026	0.028	0.06	0.06	0.052	0.06	0.06	0.055	0.058	0.06
16	LEP-16997 Amigaarnacasericeella (Copan, Honduras)	0.044	0.043	0.055	0.053	0.05	0.046	0.055	0.043	0.041	0.041	0.041	0.045	0.046	0.044	0.051		0.053	0.044	0.044	0.044	0.041	0.041	0.028	0.053	0.041	0.041	0.046	0.048	0.044	0.044	0.044	0.044	0.044	0.047	0.042	0.044
17	LEP-34357 Amigaarnacaarnaca (St-Laurent du Maroni, French Guiana)	0.073	0.061	0.006	0.006	0.022	0.031	0.034	0.031	0.06	0.06	0.06	0.03	0.032	0.03	0.035	0.053		0.024	0.027	0.024	0.059	0.059	0.052	0.003	0.056	0.056	0.024	0.022	0.056	0.056	0.053	0.056	0.056	0.055	0.054	0.056
18	LEP-37404 Amigaarnacaarnaca (San Martin, Peru)	0.064	0.056	0.027	0.027	0.017	0.022	0.011	0.023	0.061	0.061	0.059	0.021	0.018	0.023	0.023	0.044	0.024		0.003	0	0.059	0.059	0.052	0.024	0.054	0.054	0.016	0.017	0.054	0.055	0.053	0.054	0.054	0.055	0.052	0.054
19	LEP-37411 Amigaarnacaarnaca (San Martin, Peru)	0.064	0.059	0.029	0.024	0.019	0.024	0.011	0.023	0.061	0.061	0.059	0.021	0.018	0.023	0.023	0.044	0.027	0.003		0	0.061	0.061	0.052	0.024	0.051	0.051	0.012	0.014	0.058	0.055	0.057	0.058	0.058	0.058	0.056	0.058
20	LEP-37416 Amigaarnacaarnaca (San Martin, Peru)	0.064	0.056	0.027	0.024	0.017	0.023	0.011	0.021	0.061	0.061	0.059	0.021	0.018	0.023	0.023	0.044	0.024	0	0		0.06	0.06	0.052	0.024	0.052	0.052	0.013	0.014	0.055	0.055	0.054	0.055	0.055	0.055	0.053	0.055
21	LEP-37525 Amigaarnacaadela (Esmeraldas, Ecuador)	0.024	0.026	0.061	0.061	0.061	0.06	0.075	0.06	0	0	0	0.059	0.062	0.058	0.064	0.041	0.059	0.059	0.061	0.06		0	0.036	0.06	0.014	0.014	0.061	0.063	0.014	0.014	0.012	0.014	0.014	0.014	0.013	0.014
22	LEP-55465 Amigaarnacaadela (Carchi, Ecuador)	0.024	0.026	0.061	0.061	0.061	0.06	0.075	0.06	0	0	0	0.059	0.062	0.058	0.064	0.041	0.059	0.059	0.061	0.06	0		0.036	0.06	0.014	0.014	0.061	0.063	0.014	0.014	0.012	0.014	0.014	0.014	0.013	0.014
23	MGCL-LOAN-028 Amigaarnacaadela (Antioquia, Colombia)	0.049	0.049	0.052	0.049	0.059	0.042	0.077	0.039	0.036	0.036	0.036	0.039	0.046	0.039	0.042	0.028	0.052	0.052	0.052	0.052	0.036	0.036		0.049	0.03	0.03	0.056	0.059	0.03	0.03	0.03	0.03	0.03	0.03	0.03	0.03
24	MGCL-LOAN-090 Amigaarnacaarnaca (Para, Brazil)	0.073	0.062	0.003	0	0.022	0.031	0.034	0.031	0.06	0.06	0.06	0.03	0.032	0.03	0.035	0.053	0.003	0.024	0.024	0.024	0.06	0.06	0.049		0.053	0.053	0.021	0.019	0.056	0.056	0.054	0.056	0.056	0.055	0.055	0.056
25	MGCL-LOAN-139 Amigaarnacaadela (Choco, Colombia)	0.03	0.029	0.058	0.053	0.058	0.055	0.072	0.054	0.014	0.014	0.013	0.053	0.055	0.053	0.058	0.041	0.056	0.054	0.051	0.052	0.014	0.014	0.03	0.053		0	0.051	0.053	0.005	0.002	0.003	0.005	0.005	0.005	0.003	0.005
26	MGCL-LOAN-144 Amigaarnacaadela (Choco, Colombia)	0.03	0.029	0.058	0.053	0.058	0.055	0.072	0.054	0.014	0.014	0.013	0.053	0.055	0.053	0.058	0.041	0.056	0.054	0.051	0.052	0.014	0.014	0.03	0.053	0		0.051	0.053	0.005	0.002	0.003	0.005	0.005	0.005	0.003	0.005
27	MGCL-LOAN-162 Amigaarnacaarnaca (Minas Gerais, Brazil)	0.067	0.059	0.025	0.02	0.006	0.027	0.022	0.026	0.061	0.061	0.06	0.025	0.02	0.025	0.026	0.046	0.024	0.016	0.012	0.013	0.061	0.061	0.056	0.021	0.051	0.051		0.002	0.058	0.055	0.057	0.058	0.058	0.06	0.056	0.058
28	MGCL-LOAN-217 Amigaarnacaarnaca (Bahia, Brazil)	0.067	0.061	0.024	0.019	0.005	0.029	0.022	0.028	0.063	0.062	0.062	0.026	0.021	0.026	0.028	0.048	0.022	0.017	0.014	0.014	0.063	0.063	0.059	0.019	0.053	0.053	0.002		0.059	0.056	0.058	0.059	0.059	0.062	0.058	0.059
29	YB-BCI23691 Amigaarnacaadela (Panama)	0.033	0.028	0.058	0.059	0.056	0.056	0.075	0.056	0.015	0.015	0.015	0.055	0.058	0.054	0.06	0.044	0.056	0.054	0.058	0.055	0.014	0.014	0.03	0.056	0.005	0.005	0.058	0.059		0	0.002	0	0	0	0.002	0
30	YB-BCI35406 Amigaarnacaadela (Panama)	0.033	0.028	0.058	0.056	0.056	0.056	0.075	0.056	0.015	0.015	0.015	0.055	0.058	0.054	0.06	0.044	0.056	0.055	0.055	0.055	0.014	0.014	0.03	0.056	0.002	0.002	0.055	0.056	0		0.002	0	0	0	0.002	0
31	YB-BCI35436 Amigaarnacaadela (Panama)	0.03	0.027	0.055	0.057	0.055	0.051	0.072	0.053	0.012	0.012	0.012	0.05	0.053	0.049	0.052	0.044	0.053	0.053	0.057	0.054	0.012	0.012	0.03	0.054	0.003	0.003	0.057	0.058	0.002	0.002		0.002	0.002	0.002	0	0.002
32	YB-BCI46591 Amigaarnacaadela (Panama)	0.033	0.028	0.058	0.059	0.056	0.056	0.075	0.056	0.015	0.015	0.015	0.055	0.058	0.054	0.06	0.044	0.056	0.054	0.058	0.055	0.014	0.014	0.03	0.056	0.005	0.005	0.058	0.059	0	0	0.002		0	0	0.002	0
33	YB-BCI46628 Amigaarnacaadela (Panama)	0.033	0.028	0.058	0.059	0.056	0.056	0.075	0.056	0.015	0.015	0.015	0.055	0.058	0.054	0.06	0.044	0.056	0.054	0.058	0.055	0.014	0.014	0.03	0.056	0.005	0.005	0.058	0.059	0	0	0.002	0		0	0.002	0
34	YB-BCI49395 Amigaarnacaadela (Panama)	0.034	0.03	0.057	0.058	0.059	0.055	0.075	0.057	0.014	0.014	0.014	0.054	0.057	0.052	0.055	0.047	0.055	0.055	0.058	0.055	0.014	0.014	0.03	0.055	0.005	0.005	0.06	0.062	0	0	0.002	0	0		0.002	0
35	YB-BCI6807 Amigaarnacaadela (Panama)	0.03	0.026	0.056	0.057	0.054	0.054	0.072	0.055	0.013	0.013	0.013	0.054	0.056	0.052	0.058	0.042	0.054	0.052	0.056	0.053	0.013	0.013	0.03	0.055	0.003	0.003	0.056	0.058	0.002	0.002	0	0.002	0.002	0.002		0.002
36	YB-BCI766 Amigaarnacaadela (Panama)	0.033	0.028	0.058	0.059	0.056	0.056	0.075	0.056	0.015	0.015	0.015	0.055	0.058	0.054	0.06	0.044	0.056	0.054	0.058	0.055	0.014	0.014	0.03	0.056	0.005	0.005	0.058	0.059	0	0	0.002	0	0	0	0.002

Butler

(1867) A monograph of the genus

Euptychia

, a numerous race of butterflies belonging to the family

Satyridae

; with descriptions of sixty species new to science, and notes to their affinities, etc.Proceedings of the Zoological Society of London1866: 458–504.

Kirby

(1871) A Synonymic Catalogue of Diurnal

Lepidoptera

.John Van Voorst, London, 690 pp.

Butler

(1877) On new species of the genus

Euptychia

, with a tabular view of those hitherto recorded.Journal of the Linnean Society of London (Zoology)13(67): 116–128. https://doi.org/10.1111/j.1096-3642.1877.tb02375.x

Godman

Salvin

(1880–1881) Biologia Centrali-Americana.

Insecta

Lepidoptera

Rhopalocera

. Dulau & Co., Bernard Quaritch (London) 1(1): 1–32, pls 1, 2 [September 1879], 1(2) [November 1879]: 33–56; 1(3)[February 1880]: 57–72; 1(6)[August 1880]: 73–88; 1(9)[February 1881]: 89–96.

Weymer

(1911) 4. Familie:

Satyridae

. In: Seitz

(Ed.) Die Gross-Schmetterlinge der Erde.A.Kernen (Stuttgart)5: 173–192.

Riley

Gabriel

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