Abstract
We here transfer an euptychiine taxon hitherto placed in the polyphyletic genus Magneuptychia Forster, 1964, to Caeruleuptychia Forster, 1964. Caeruleuptychia francisca (Butler, 1870), n. comb. is reclassified based on a morphology-based maximum likelihood analysis, which is consistent with ongoing analyses of molecular data. Two putative synapomorphic characters are identified for the “Caeruleuptychia umbrosa clade”, one of which appears to be an unusual characteristic of euptychiine butterflies and is tested by optimizing onto the maximum likelihood tree. We also discuss the systematic placement of three additional enigmatic Caeruleuptychia species. A lectotype is designated for Euptychia francisca, and the genitalia of this species are illustrated here for the first time.
S. Nakahara*, K. Kleckner, K.R. Willmott, McGuire Center for Lepidoptera and Biodiversity, Florida Museum of Natural History, University of Florida, Gainesville, FL 32611, USA. snakahara@ufl.edu
G. Lamas, Departamento de Entomología, Museo de Historia Natural, Universidad Nacional Mayor de San Marcos, Apartado 14-0434, Lima-14, Peru
B. Huertas, Life Sciences Department, Natural History Museum, London, UKhttp://zoobank.org/577C9E59-22A4-460D-9548-AE0DCD5D2121
Introduction
As initially highlighted by Murray & Prowell (2005), followed by Freitas (2007) and Peña et al. (2010), and confirmed by ongoing, increasingly comprehensive molecular phylogenetic studies, many genera in the diverse Neotropical satyrine butterfly subtribe Euptychiina are polyphyletic (e.g., Chloreuptychia Forster, 1964, Euptychoides Forster, 1964; Nakahara et al. 2016, 2019; Willmott et al. 2019). In part, this polyphyletic classification is a result of the convergent evolution of similar wing pattern characters, such as the lilac-blue ventral reflective coloring of Chloreuptychia and the white ventral bands of Euptychoides (sensu Lamas 2004) (Nakahara et al. 2016, 2019; Willmott et al. 2019).
Magneuptychia Forster, 1964 is another species-rich euptychiine genus described by Forster (1964), and Forster’s classification was retained by Lamas (2004) who followed Miller (1968). Along with the aforementioned two genera, and several others, Magneuptychia is also highly polyphyletic, and Costa et al. (2016) proposed that the generic name should only be applied to the type species, Papilio libye Linnaeus, 1767, in addition to two closely related taxa. However, Costa et al. (2016) did not support this proposed generic classification with phylogenetic data or provide new generic combinations for those taxa that they suggested should be removed from Magneuptychia (sensu Lamas 2004). Andrade-C. et al. (2019) also introduced four new generic names for a small number of taxa currently placed in Magneuptychia, although the relationships, monophyly, taxonomic limits and validity of those proposed genera have yet to be tested phylogenetically. Thus, we still regard Magneuptychia as one of the most diverse euptychiine genera awaiting a revisionary study in phylogenetic context to provide a robust generic classification for those species unrelated to the type species, P. libye. To contribute to ongoing revisionary work on Caeruleuptychia Forster, 1964 and Magneuptychia we here transfer “Euptychia francisca Butler, 1870” from Magneuptychia to Caeruleuptychia Forster, 1964 and thus propose a new combination based on a morphology-based phylogenetic analysis. The likely relationship of this species to other Caeruleuptychia was indicated in Espeland et al. (2019), and our phylogenetic analysis based on morphological characters performed for this study enabled us to clarify the relationships of three enigmatic taxa provisionally treated in Nakahara et al. (2018b) as well. An unusual male genitalic structure was identified, and a preliminary phylogenetic analysis was performed to trace the evolutionary history of this putative synapomorphic character for the clade.
Material and methods
Specimens relevant to this study were examined in the following collections:
FLMNH: |
McGuire Center for Lepidoptera and Biodiversity (MGCL), Florida Museum of Natural History, Gainesville, USA |
HAWA: |
Haydon Warren-Gash collection, Pressac, France |
INABIO: |
Instituto Nacional de Biodiversidad, Quito, Ecuador |
MUSM: Museo de Historia Natural, Universidad Nacional Mayor de San Marcos, Lima, Peru | |
NHMUK: |
Natural History Museum, London, UK (formerly BMNH) |
These acronyms for wings are used throughout the text:
DFW: |
Dorsal forewing |
DHW: |
Dorsal hindwing |
VFW: |
Ventral forewing |
VHW: |
Ventral hindwing |
Morphological study
Morphology (genitalia, labial palpi, legs) of relevant specimens was examined by appendages being soaked in hot 10% KOH for 10–15 minutes, dissected and subsequently stored in glass tubes in glycerine. Wing venation was studied by removing scales using diluted bleach. Morphological characters were examined using a Leica MZ 16 stereomicroscope and illustrations were prepared using a camera lucida attached to this scope. The terminology for characters associated with wings (area, venation, pattern elements etc.) and genitalic structures follow Nakahara et al. (2018a). A JEOL JSM-5510LV Scanning Electron Microscope (SEM) was used to take images of relevant structures.
Phylogenetic analysis
In order to test the placement of “Euptychia francisca” in Caeruleuptychia, 34 characters were coded for phylogenetic analysis. Of these characters, five were multistate and 29 were binary (see below, subsect. Coded characters, for further information). As seen from the fact that we applied a Markov (MK) model in this study, none of these characters are considered as “ordered”. Outgroups were selected based on published information available in Espeland et al. (2019) and unpublished molecular data relevant to this group (Nakahara, unpublished data). The aforementioned 34 characters were coded for 14 ingroup taxa chosen for this study, mainly described taxa for which there is no ambiguity regarding their identity, in addition to two outgroup taxa (see Table 1 for further information). Note that the coding of female characters for C. scripta Nakahara, Zacca & Huertas, 2017 and C. pilata (Butler, 1867) was performed based on the female figured in Nakahara et al. (2018b: Fig. 2 S, T) in order to avoid missing data, although which of these two species this specimen represents has not yet been determined.
Following Lewis (2001), a phylogenetic analysis with maximum likelihood (ML) as the optimality criterion was performed for the above morphological dataset (Table 1) by specifying “MK+F+ASC” as the substitution model in IQ-TREE 2.0 (Minh et al 2020). We employed “Ascertainment bias correction (ASC)” due to the lack of constant sites. In order to initialize the candidate tree set, 98 parsimony trees were generated by random-order stepwise addition and approximate likelihood scores were calculated. Subsequently, hill-climbing Nearest-Neighbor interchange (NNI) searches on the top 20 selected trees were conducted to find the locally optimal ML trees. The tree search was completed after 112 iterations, and the tree search was finalized by finding the highest log-likelihood score for the dataset (Table 1) given the tree. Branch support was calculated using ultrafast bootstrap (UFBoot) with 1,000 replicates, in addition to assessing node support through 1,000 replications of the Shimodaira – Hasegawa-like approximate Likelihood Ratio Test (SH-aLRT) (Guindon et al. 2010; Hoang et al. 2018).We also constructed a consensus tree from 1,000 bootstrap (BS) replications as a resampling method.
Ancestral state estimation
In order to investigate the character history and infer missing character data for Caeruleuptychia maryzenderae Lamas & Nakahara, 2017 (presence/absence of hair-like scales on the valva), a Bayesian approach of estimating the ancestral state based on the threshold model was performed based on the ML tree (see Felsenstein 2012; Revell 2014). The probability of missing data for C. maryzenderae was coded as 0.5 for both character states. Ancestral state reconstruction was conducted for this particular character using the “ancThresh” function in the R studio package PHYTOOLS with Brownian motion (BM) as the liability model, generated through a run with 1,000,000 Markov chain Monte Carlo (MCMC) iterations.
Coded characters
- 1. Bluish scales on dorsal wing surface: absent (0); present (1)
- 2. (following char. no.1) If bluish scales on dorsal surface present: absent anterior of DHW Rs (0); covering area anterior of DHW Rs (1)
- 3. DFW androconial patch, as compacted scales in cell 2A: absent (0); present (1)
- 4. DFW androconial patch, as hair-pencil in cell Cu2, absent (0); present (1)
- 5. DFW subapical black spot in cell M1: absent (0); present (1)
- 6. VFW ocellus in cell M1: reduced (0); present (1)
- 7. VFW ocellus in cell M1, if present, pupil: absent (0); present (1)
- 8. Orangish umbra on VFW: absent (0); present (1)
- 9. DHW androconial patch at tornus: absent (0); present (1)
- 10. (following char. no. 9) If DHW androconial patch present, color: black (0); orange (1)
- 11. VHW ocellar ring in cells M1 and Cu1: skewed (0); evenly broad (1)
- 12. Ocellus in VHW cell M2 and M3: appearing as incomplete ocellus (e.g., lacking black central area, pupil) compared to ocellus in cells M1 and Cu1 (0); similar in appearance to ocellus in cells M1 and Cu1 (1)
- 13. Ocellus in VHW cell Cu2: absent (0); present (1)
- 14. DFW and DHW violet tinge: absent (0); present (1)
- 15. FW venation: origin of R2 at base of discocellular vein (0); closer to R1 compared to origin of discocellular vein (1); closer to R3 compared to origin of discocellular vein (2)
- 16. Male eighth sternite: single plate (0); divided into two plates (1); V-shaped (upwards) (2); V-shaped upside down (3)
- 17. Prominently protruding bulge from posterior edge of tegumen above uncus: absent (0); present (1)
- 18. Hair-like scales on tegumen: absent (0); present (1)
- 19. Uncus: straight (0); arched downwards (1)
- 20. Brachia: absent/reduced (0); developed (1)
- 21. Juxta: plate-like in ventral view (0); stripe-like in ventral view(1)
- 22. Valva shape in lateral view, roughly: boat-shaped (0); bean-shaped (1)
- 23. Valva, hair-like scales: absent (0); present (1)
- 24. Valva, ventral margin of apical process: rather straight (0); curved inwards (1); curved outwards (2)
- 25. Valva, dorsal hump of apical process: absent (0); present (1)
- 26. Phallus, sclerotized region of vesica: absent (0); present as weakly sclerotized cornuti (1); spiny cornuti (2)
- 27. Female, eighth tergite: membranous (0); sclerotized (1)
- 28. Female 7th and 8th abdominal intersegmental membrane pleated and fully expandable (0); partially expandable (1)
- 29. Rectangular weakly sclerotized region on folded portion of intersegmental membrane: absent (0); present (1)
- 30. Papillae analis, posterior apophysis: absent (0); present (1)
- 31. (following char. no. 30) Posterior apophysis, if present: as rounded point (0); rather elongated projection (1)
- 32. Lamella antevaginalis: membranous (0); sclerotized (1)
- 33. Ductus seminalis: origin within basal one-fourth of ductus bursae (0); located within distal three-fourths of ductus bursae (1)
- 34. Signa located: ventrally (0); dorsally (1); laterally (2)
Results
The Robinson–Foulds distance between our ML tree and the consensus tree was 6, and thus the ML tree and consensus tree were topologically almost identical (Fig. 1). The branch support of the consensus tree was identical to the ML tree’s UFBoot support at all nodes; therefore, only the ML tree is presented herein. “Euptychia francisca” was recovered as a member of a moderately well supported “Caeruleuptychia umbrosa clade” (Fig. 1; SH-aLRT/UFBoot=94.4/56). Three enigmatic species, C. maryzenderae, C. pilata and C. scripta, were also recovered as members of this clade, with C. pilata and C. scripta as a sister species pair. The consensus tree recovered Caeruleuptychia as a moderately supported clade (BS=78), including the senior subjective synonym of the type species (“E. caerulea Butler, 1869”), namely “Euptychia urania Butler, 1867” (Nakahara, unpublished data). The type species of Magneuptychia, “Papilio libye”, was shown to be distantly related to Caeruleuptychia in Espeland et al. (2019) (which analysis included C. urania and C. umbrosa (Butler, 1870)), and we thus consider this proposed taxonomic change, C. francisca, n. comb., as justified. Ancestral state estimation strongly indicated that the presence of setae on the male genitalic valva is a synapomorphy for a small group of Caeruleuptychia, containing, among other species, C. francisca, n. comb. (Fig. 2). Furthermore, the probability of this character state being “present” in C. maryzenderae was estimated to be high (Fig. 2).
Genus Caeruleuptychia Forster, 1964
Type species: Euptychia caerulea Butler, 1869 (p. 6, pl. III, figs. 1–2) – by original designation.
Caeruleuptychia francisca (Butler, 1870), new combination
Euptychia francisca (1870: p. 49, pl. 18, fig. 3):
Type material: Lectotype (designated here) ♂: // ‘Ecuador Hewitson Coll. 70–69. Euptychia Francisca, Butl. 1’ // ‘SYN-TYPE’ // ‘BMNH (E) 787670’ // ‘NHMUK 014172457’ // (NHMUK) [examined]; Paralectotypes ♂: // ‘B. M. TYPE No. Rh3182 Euptychia francisca; ♀ Butl.’ // ‘Ecuador Hewitson Coll. 70–69. Euptychia Francisca, Butl. 2’ // ‘E. Francisca Butl. type.’ // ‘Type H. T.’ // ‘SYN-TYPE’ // ‘BMNH (E) 1266950’ // ‘NHMUK 014172458’ // (NHMUK) [examined].
Euptychia francisca: Butler 1877: 117; Kirby 1871: 643; 1879: 134; Weymer 1911: 196; Gaede 1931: 446; D’Abrera 1988: 769, fig.
Magneuptychia francisca: Lamas, 2004: 220; Costa et al. 2016: 204; Espeland et al. 2019: 118, fig. 1.
Identification and taxonomy
Caeruleuptychia francisca, n. comb. is distinguished from other species in the genus by the ochre ground color of the VFW and VHW. The presence of hair-like setae on the valva and the violet tinge mainly on the DHW (also slightly visible along the DFW’s inner margin) of males can also be used to distinguish this species from other Caeruleuptychia species, and indeed from all euptychiine species. Caeruleuptychia francisca, n. comb. can perhaps be confused with species in the “C. umbrosa clade” when the ground color is somewhat faded, but its rather large adult size (forewing length 21–22 mm (n = 3), whereas the forewing length of many species in this clade is smaller than 20 mm) compared to many other species in the clade is useful in distinguishing this taxon from congeners. Despite the absence of androconial scales and/or hair pencils on the DFW of male C. francisca, n. comb. being atypical among species in the “C. umbrosa clade”, several undescribed species in this clade are known in which males lack androconial scales and/or hair pencils on the DFW. These undescribed species can be distinguished from C. francisca, n. comb. by the characters mentioned above, in addition to several other phenotypic characters (Nakahara, unpublished data).
“Euptychia francisca” was described by Arthur Gardiner Butler in his “Lepidoptera Exotica, or descriptions and illustrations of exotic Lepidoptera” published in August 1870 (Butler 1870). Butler did not explicitly state the sex or number of specimens he examined, but he did state that the specimens came from Ecuador, were collected by Buckley, and were housed in Hewitson’s collection. However, his rather brief description of “E. francisca” does include references to some diagnostic characters for this taxon mentioned above, such as “Alae subtus ochraceae”, (wings below ochre) and with a slight tinge of violet on the dorsal wings (“alae supra fuscae violaceo-tinctae”). The mention of these phenotypic characters combined with the illustration of this species on pl. 18, fig. 3 allows us to confidently conclude that Butler examined and described a specimen of what we regard here as C. francisca. The syntype (in the NHMUK) figured in Warren et al. (2017) is identified as a female, perhaps based on the rather weak violet tinge on the dorsal wing. We located another male syntype in better condition (listed above and figured in Fig. 3), and thus we here designate this syntype as a lectotype of “Euptychia francisca” in order to settle the nomenclature (lectotype designation). Note that apparently the abdomen of the paralectotype is glued on and it is not clear whether it belongs to the same specimen or even the same species, but wing pattern characters leave no doubt as to its identity.
Variation
The VHW postdiscal and submarginal bands are apparently fused in cell 2A in some specimens (e.g., FLMNH-MGCL-269500), whereas they appear not to merge in other specimens (e.g, FLMNH-MGCL-269502).
Specimens examined (31♂, 2♀)
Ecuador: Orellana: Loreto-Tena rd., Pasourcu, [0°44’4” S, 77°30’44” W], 900–1000 m, (Hall, J. P. W., Willmott, K. R., J. C. R., J. I. R), 23, 26.vii.2016, 1 ♂ [FLMNH-MGCL-209992] (FLMNH); Napo: km 14 Tena–Puyo rd., Apuya, [1°6’18” S, 77°46’42” W], 600 m, (Warren-Gash, H.), 22.viii.2010, 2 ♂, (HAWA); Tena–Puyo rd., El Capricho, [1°11’14” S, 77°49’53” W], 800 m, (Willmott, K. R.), 2.x.1997, 2 (FLMNH); Pastaza: Puyo–Macas rd., Pitirishca, [1°48’18” S, 77°49’15” W], 1000 m, (Willmott, K. R.), 26.vii.1998, 1 ♂ [dissection, SN-19-33] (FLMNH); Puyo, [1°28’ S, 77°59’ W], 1000 m, (Brown, F. M. & H. H.), 6.xii.1938, 1 ♂ [FLMNH-MGCL-269502], 1 ♂ [FLMNH-MGCL-269503], 1 ♂ [FLMNH-MGCL-269504; dissection, SN-17-164], 1 ♀ [FLMNH-MGCL-269505] (FLMNH); Río Bobonaza, Sarayacu, [1°44’ S, 77°29’ W], 200 m, (Buckley, C.), 1 ♂ [BMNH-E-787671] (NHMUK); Río Bobonaza, Sarayacu, [1°44’ S, 77°29’ W], 400 m, (Buckley, C.), 1 ♂ [BMNH(E)-1498419] (NHMUK); Morona-Santiago: c. 6 km along Bolivarense track, [1°40’25” S, 77°46’27” W], 850 m, (Warren-Gash, H.), ix.2011, 1 ♂, (HAWA); km 32.8 Santiago–Puerto Morona rd., [2°58’54” S, 77°48’6” W], 650–750 m, (Busby, R. C., Aldas, I.), 15.9.ix.2014, 1 ♂ [FLMNH-MGCL-195341] (FLMNH); nr. Gualaquiza, Bomboiza, [3°25’36” S, 78°31’0” W], 850 m, (Hall, J. P. W., Willmott, K. R.), 26–29.vii.1993, 1 ♂ [FLMNH-MGCL-269501] (FLMNH); Río Waiwaim, Taisha, [2°22’36” S, 77°30’0” W], 600 m, (Hall, J. P. W., Willmott, K. R.), 1–4.vi.1994, 1 ♂ [FLMNH-MGCL-269500] (FLMNH); Zamora-Chinchipe: c. 3 km W Guayguayme Alto, ridge above San Luís, [3°55’14” S, 78°54’49” W], 1470 m, (Willmott, K. R., J. I. R., J. C. R.), 23.vi.2013, 1 ♂ [FLMNH-MGCL-157422], 1 ♂ [FLMNH-MGCL-157423], 1 ♂ [FLMNH-MGCL-157424], 1 ♀ [FLMNH-MGCL-157425; dissection, SN-19-110] (FLMNH), (INABIO); c. 4 km N Guayguayme Alto, [3°53’50” S, 78°53’25” W], 1470 m, (Busby, R. C.), 20.ix.2014, 1 ♂ [FLMNH-MGCL-195342] (FLMNH); km 11.5 Los Encuentros–Zarza, La Libertad, [3°47’54” S, 78°36’26” W], 1250 m, (Willmott, K. R., Hall, J. P. W.), 6,8.viii.2009, 1 ♂ [FLMNH-MGCL-145690], 1 ♂ [FLMNH-MGCL-145692], 1 ♂ [FLMNH-MGCL-145693] (FLMNH); lower Río Numpatakaime, c. 3 km S Shaime, [4°21’0” S, 78°39’28” W], 900 m, (Willmott, K. R., Hall, J. P. W.), 31.vii.2009, 1 ♂ [FLMNH-MGCL-145691] (FLMNH); ridge SE La Primavera, [3°43’19” S, 78°36’16” W], 1175 m, (Hall, J. P. W., Willmott, K. R., J. I. R.), 23.vii.2018, (INABIO); Zamora, ridge to west, [4°4’30” S, 78°58’7” W], 1400–1450 m, (Willmott, K. R.), 20.v.2000, 1 ♂ [dissection, SN-17-175] (FLMNH), 1 ♂ (FLMNH); Zamora–Yantzaza rd., Namírez Bajo, [3°58’6” S, 78°49’48” W], 1050 m, (Willmott, K. R.), 21.v.2000, 1 ♂, (FLMNH); not located: ‘Ecuador’, 1 ♂ [BMNH(E)-1498420], 1 ♂ [BMNH-E-787670] (NHMUK), 1 ♂ [‘E. Francisca Butl. Type’ // ‘Ecuador Hewitson Coll. 79-69. Euptychia francisca, Butl. 2‘ // ‘Type H.T.‘ // ‘B.M. TYPE No. Rh3182 Euptychia francisca ♀ Butl.‘] (NHMUK). Peru: Amazonas: Cordillera del Cóndor, Quebrada Kegkem, [3°38’ S, 78°18’ W], 700 m, (Grados, J.), 20.xi.2003, 1 ♂ [MUSM-LEP-102947] (MUSM); Falso Paquisha, [3°58’ S, 78°25’ W], 800 m, (Lamas, G.), 31.x.1987, 1 ♂ [MUSM-LEP-102946] (MUSM).
Other records. Ecuador: Pastaza: Zamora-Chinchipe: nr. Zamora, Chachacoma, [4°4’30” S, 78°57’36” W], 1350 m, (Willmott, K. R., Aldaz, R.), 1.xi.2006, (sight record) (Willmott & Hall, unpublished data).
Distribution and natural history
Caeruleuptychia francisca is known from Napo in eastern Ecuador as far south as Amazonas in extreme northern Peru on the Ecuadorian border (Fig. 22). It has been reliably recorded from 600–1470 m in rainforest and cloud forest habitats in the foothills of the eastern Andes. Most known sites for the species consist of primary or selectively logged forest on ridgetops, where males were observed, often in small groups, perching in light gaps or relatively open forest from 1–4 m above the ground from 12:30 to 15:00 h. A pair was found in copula at 12:30 h on a forest ridgetop in the vicinity of other perching males.
Discussion
Despite our decision to transfer “Euptychia francisca” to Caeruleuptychia being based in this paper solely on morphological data, this placement is also partly supported by published (Espeland et al. 2019) and the senior author’s unpublished molecular data. C. francisca, n. comb. is recovered as a member of the strongly supported “Caeruleuptychia umbrosa clade” in a preliminary maximum likelihood analysis based on four genes (Nakahara, unpublished data). The molecular matrix is still in development and will be published in forthcoming revisions of the “Caeruleuptychia caerulea clade” and “C. umbrosa clade”, so we focus attention here on the morphological characters, which will be important in placing several rare species which lack sequence data. In fact, the placement of C. maryzenderae, C. pilata and C. scripta in the “C. umbrosa clade” is also supported by molecular data, but only based so far on DNA “barcode” sequences, which were not available while preparing Nakahara et al. (2018b). Owing partly to the molecular evidence for these taxa being members of the “C. umbrosa clade”, two putative morphological synapomorphies for the clade were subsequently identified and are described here. These synapomorphies are the presence of hair-like scales on the valva (Fig. 6), and the appearance of the posterior apophysis of the papilla analis as a rounded point (Fig. 12). Unfortunately, the hair-like scales on the valva were overlooked during our previous study and illustration of the male genitalia of C. pilata, C. scripta and C. maryzenderae in Nakahara et al. (2018b). We here confirm the presence of this character in C. pilata and C. scripta, although it still remains to be confirmed for C. maryzenderae. Nevertheless, the ancestral state reconstruction conducted for this study indicated the character state to be “present” for C. maryzenderae with a high probability (Fig. 2). Furthermore, the presence of hair-like setae on the valva appears to be synapomorphic for the remainder of the “C. umbrosa clade” based on our Bayesian analysis. However, the distribution of this character state must be tested on a more comprehensive, well-resolved phylogeny of the group, and the results here should be regarded as preliminary. The presence of such hair-like scales on the valva has not been observed in any other euptychiine species, although the setae can apparently easily be lost, even by the gentle rubbing with a paint brush typically used to clean genitalia prior to preparing the illustration, explaining our omission in Nakahara et al. (2018b). Examination of the setae using a Scanning Electron Microscope (SEM) shows that these setae are structurally similar to the hair-like scales of the DFW hair-pencil possessed by many species in the “C. umbrosa clade” in possessing horizontally aligned grooves (termed “longitudinal ridges” sensu Downey & Allyn 1975; Figs 18–21). In contrast to the showy and colorful species in two other major clades in the genus Caeruleuptychia (“C. caerulea clade” and “C. aegrota clade”), the species in the “C. umbrosa clade” are usually drab and dull, typical of euptychiine species. As an alternative to the iridescent bluish scales that could serve in other Caeruleuptychia taxa in mate recognition, members of the “C. umbrosa clade” perhaps use these hair-like setae on the valvae in species recognition and to ensure reproductive isolation. Clearly, a well-resolved taxonomy and phylogeny is needed for such a group where many interesting evolutionary and biological questions remain to be addressed, and we hope that this study will contribute towards our steadily improving understanding of euptychiine diversity and relationships.
Acknowledgements
We are grateful to Sebastián Padrón (Cuenca, Ecuador) and Andrew Neild (London, UK) for help with translating relevant portions of original descriptions, Thamara Zacca (Campinas, Brazil) for checking characters at University of Campinas, David Plotkin (Gainesville, FL, USA) and Ryan St Laurent (Gainesville, FL, USA) for support and fruitful discussion. KRW thanks Jason Hall, Julia and Jamie Robinson Willmott, Ismael Aldas, Raul Aldaz, and Robert Busby, for companionship in the field in Ecuador and for providing specimens and/or data. SN acknowledges the National Science Foundation (Grant No. DEB–1256742) and the University of Florida’s Entomology and Nematology Department for support. Field work in Ecuador was done under the permit “Diversity and Biology of Lepidoptera of Ecuador” (MAE-DNB-CM-2016-0045) with the support of INABIO and the Ministerio del Ambiente, and supported by grants from the National Geographic Society, National Science Foundation (DEB-0103746, DEB-0639861, DEB-1256742), the Darwin Initiative and the Leverhulme Trust.
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