Remarkable sexual dimorphism, rarity and cryptic species: a revision of the ‘aegrota species group’ of the Neotropical butterfly genus Caeruleuptychia with the description of three new species (Lepidoptera, Nymphalidae, Satyrinae)

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The ‘aegrota species group’ of the Neotropical nymphalid genus Caeruleuptychia Forster, 1964, in addition to three other superficially similar, enigmatic species in the genus, are revised. A lectotype is designated for Euptychia aegrota Butler, 1867, E. aetherialis Butler, 1877 stat. rev., E. helios Weymer, 1911 and E. pilata Butler, 1867, and C. aetherialis is resurrected from its synonymy with C. aegrota. Caeruleuptychia helios caelestissima Brévignon, 2010, syn. nov., and Magneuptychia keltoumae Brévignon & Benmesbah, 2012, syn. nov. are both regarded as junior subjective synonyms of C. helios (Weymer, 1911), as a result of the discovery and first illustration of the female of this taxon. The female of C. aegrota is also described and illustrated for the first time, and three new species, C. trembathi Willmott, Nakahara, Hall & Neild, sp. nov., C. scripta Nakahara, Zacca & Huertas, sp. nov., and C. maryzenderae Lamas & Nakahara, sp. nov. are described and named. We analyze morphological and molecular data separately, in addition to combining morphological data with molecular data, to provide the first phylogenetic hypothesis for the taxa treated in this revision.

Remarkable sexual dimorphism, rarity and cryptic species: a revision of the ‘aegrota species group’ of the Neotropical butterfly genus Caeruleuptychia with the description of three new species (Lepidoptera, Nymphalidae, Satyrinae)

in Insect Systematics & Evolution

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References

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Figures

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    Schematic representation of coded characters: Adults, from left to right, C. maryzenderae sp. nov., (dorsal above, ventral below); C. scripta sp. nov., (dorsal above, ventral below); C. helios (ventral, above); C. umbrosa (ventral, below); male genitalia (lateral view, aedeagus shown below), from left to right, C. aegrota, C. scripta sp. nov., and Paryphthimoides grimon. See relevant section of the text for further information.

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    Phylogenies inferred in this study: A. Consensus tree of two parsimonious trees inferred based on morphological data, see text for details. White circle = non-unique changes (homoplasies); Black circle = unique changes (synapomorphies); number above = character; number below = character state. B. Majority-rule consensus tree inferred with MrBayes for dataset 4 (morphology and 3 genes). Posterior probabilities are shown above branches and nonparametric bootstraps values below branches. Members of the aegrota group are highlighted in bold. C. Majority-rule consensus tree inferred with MrBayes for dataset 1 (COI data). Posterior probabilities from MrBayes are shown above branches and nonparametric bootstrap values from the maximum likelihood analysis in IQ-tree are shown below branches. Members of the aegrota group are highlighted in bold.

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    A, B. Two parsimonious trees inferred based on morphological data. C. Majority rule consensus tree inferred with MrBayes for dataset 2 (morphological data). Posterior probabilities are shown above branches and nonparametric bootstraps values below branches. D. Majority rule consensus tree inferred with MrBayes for dataset 3 (COI + nuclear genes). Posterior probabilities are shown above branches and nonparametric bootstraps values below branches.

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    Adults of Caeruleuptychia species discussed herein: A, B. C. aegrota Male, Lectotype, Brazil, Para (NHMUK, BMNH(E) 1267012); C, D. C. aegrota Female, Ecuador, Rio Chalayacu (MGCL, FLMNH-MGCL-209991); E, F. C. aetherialis Male, Lectotype, Ecuador, Sarayacu (NHMUK, BMNH(E) 1267014); G, H. C. aetherialis Male: Ecuador, Sucumbíos, km 10.5 Lumbaquí-Baeza rd. (MGCL, FLMNH-MGCL-209989); I, J. C. trembathi sp. nov., Male, Holotype, Ecuador, 2 km N San Isidro (MGCL, FLMNH-MGCL-195361); K, L. C. trembathi sp. nov., Female, Ecuador, Río Abanico (DATA, No. 20141); M, N. C. helios Male, Peru, Tambopata Research Center (MUSM-LEP-100121); O, P. C. helios Female, Peru, Estación Biológica Los Amigos (MUSM-LEP-103073); Q, R. C. pilata Male, Peru, Pucallpa (MGCL, FLMNH-MGCL-278163); S, T. C. pilata or C. scripta sp. nov., Female, Brazil, Rondônia (MGCL, FLMNH-MGCL-278166); U, V. C. scripta sp. nov., Male, Holotype, Peru, 30 km SW Puerto Maldonado (USNM, LEP-18637); W, X. C. maryzenderae sp. nov., Male, Paratype, Peru, Pakitza (MUSM, MUSM-LEP 100329).

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    Caeruleuptychia female abdomen and genitalia. A–C, C. aegrota Ecuador, Río Chalayacu (KW-15-144). A. Abdomen, lateral aspect. B. Abdomen, dorsal aspect at posterior tip. C. Genitalia, dorsal aspect. D–F, C. trembathi Ecuador, Río Abanico (KW-15-137). D. Abdomen, lateral aspect. E. Abdomen, dorsal aspect at posterior tip. F. Genitalia, dorsal aspect.

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    Locality records for Caeruleuptychia species. ‘?’ indicates a locality point based only on a female specimen putatively identified as that species.

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    Caeruleuptychia male genitalia: la, lateral aspect; al, aedeagus lateral aspect; ad, aedeagus dorsal aspect; va, ventral aspect; da, dorsal aspect. A. C. aegrota, Peru, Madre de Dios (SN-16-61) (aedeagus from SN-16-64). B. C. aetherialis, Peru, Cuzco (SN-16-81). C. C. trembathi sp. nov., HT, Ecuador, San Isidro (KW-15-127). D. C. helios, Peru, Pakitza (SN-16-80). E. C. pilata, Brazil, Amazonas (TZ 35880). F. C. scripta sp. nov., PT, Brazil, Vila Rondônia (Billota 1997; DZ-5267). G. C. maryzenderae sp. nov., PT, Peru, Pakitza (SN-16-40).

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    Caeruleuptychia female abdomen and genitalia. A–C, C. helios, Peru, Estación Biológica Los Amigos (SN-14-17). A. Abdomen, lateral aspect. B. Abdomen, dorsal aspect at posterior tip. C. Genitalia, dorsal aspect. D–H, C. pilata/scripta Brazil, Fazenda Rancho Grande (KW-15-156). D. Abdomen, lateral aspect. E. Abdomen, dorsal aspect at posterior tip. F. Abdomen, ventral aspect at posterior tip. G. Genitalia, dorsal aspect. H. Corpus bursae, perpendicular to bands of cornuti.

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    Caeruleuptychia morphology. A. C. aegrota, Peru, Tambopata, male wing venation (FLMNH-MGCL-210072). B. C. aegrota, Ecuador, Río Chalayacu, lateral view head and labial palpi (FLMNH-MGCL-209993). C. C. pilata, Brazil, Boa Vista, male right forewing androconial scales (FLMNH-MGCL-278164). D. C. scripta, male left forewing androconial scales. E. C. maryzenderae, male right forewing androconial scales. F. C. pilata/scripta first instar head capsule, frontal view, from female dissection KW-15-156. G. As F, lateral view.

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    Locality records for Caeruleuptychia species.

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    Caeruleuptychia helios male attracted to bait of fermented fish and urine. Villa Carmen Biological Station, Pilcopata, Cuzco, Peru, April 2016 (photo: A. Neild).

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    Trail 2 at Villa Carmen Biological Station, Pilcopata, Cuzco, Peru, showing secondary regrowth habitat where numerous species of Caeruleuptychia have been recorded.

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    View of the Río Piñipiñi valley, Villa Carmen Biological Station, Pilcopata, Cuzco, Peru. The secondary forest along the banks of the river supports several species of Caeruleuptychia.

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