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Austral Entomology (2015) ••, ••–••

Thisbe silvestre sp. nov. (: ): a new myrmecophilous butterfly from the Brazilian Atlantic Forest

Lucas A Kaminski,1,2* Cristiano A Iserhard3 and André V L Freitas2

1Institut de Biologia Evolutiva, Consejo Superior de Investigaciones Científicas (CSIC) and Universidad Pompeu Fabra (UPF), Passeig Marítim de la Barceloneta, 37, Barcelona 08003, Spain. 2Departamento de Biologia , Instituto de Biologia, Universidade Estadual de Campinas, CP 6109, Campinas, SP 13083-970, . 3Instituto de Biologia, Universidade Federal de Pelotas, CP 354, Capão do Leão, RS 96160-000, Brazil. Lucas A Kaminski: http://zoobank.org/urn:lsid:zoobank.org:author:E310573F-CB6F-4C96-850E-ED7B2F93D4AE Cristiano A Iserhard: http://zoobank.org/urn:lsid:zoobank.org:author:E63CA373-990B-4EE8-AFA4-BF9E371A18EF André V L Freitas: http://zoobank.org/urn:lsid:zoobank.org:author:266CACB7-FAA8-4DC0-A44E-58002A41FF0B http://zoobank.org/urn:lsid:zoobank.org:pub:23DF558B-7CC4-471D-A19B-65EE15D856D9

Abstract This paper describes a new of Thisbe Hübner from the Atlantic Forest of south-eastern Brazil. Both adult and immature morphology are described and illustrated, and the natural history, behaviour and symbiotic interactions with ants () are documented. This new species, T. silvestre sp. nov., is known only from the type locality at Parque Nacional do Caparaó, Minas Gerais state, Brazil. Due to the isolation of the known population, specialised host plant use, restricted geographic distribution and high degree of specificity in a vulnerable habitat, T. silvestre is considered threatened. Thus, monitoring this population is urgently needed, as well as the search for other populations of this butterfly in potential localities in order to evaluate its conservation status. Key words ant-organ, conservation, mutualism, , Riodinidae, trophobiosis.

INTRODUCTION (see discussion in Hall 1999), followed by a period of sta- bility with little taxonomic novelties. This scenario has Butterfly species richness is extraordinarily high in the changed in the last 30 years, and a number of new species Neotropics, when compared with other biogeographic regions have been recently described (e.g. Brévignon 2010; Hall & (Heppner 1991; Robbins 1993). The Riodinidae is Willmott 2010; Dolibaina et al. 2013, 2014; Callaghan & remarkably diverse, with approximately 1300 described Nobre 2014; Dias et al. 2014; Siewert et al. 2014). In view species (Callaghan & Lamas 2004). Adult riodinids are of continuing new descriptions, we can assume that the notable for their morphological variation and for being usually current diversity of Riodinidae is underestimated and that a rare. In general, their populations are located in ‘ideal’ significant number of species are waiting to be unveiled in microhabitats, being temporal and spatially scarce, even in the coming years. widely distributed species (Bates 1859; Callaghan 1978; For example, the Thisbe Hübner includes seven Harvey 1987; d’Abrera 1994; Kaminski et al. 2014). Factors species that occur from through Central and South such as canopy openness and vegetation structure may influ- America (Callaghan & Lamas 2004). Although there are two ence their occurrence in a given site (Callaghan 1983; Alonso published phylogenetic proposals, there is still no consensus 2005). Furthermore, some species may present vertical strati- on the relationships of the species in relation to those formerly fication, spending much of their adult life in the tree canopy, treated in Uraneis Bates (see Hall & Harvey 2001; Penz & only occasionally descending to the understory and thus con- DeVries 2001). Recently, new species have been described in sequently being only rarely recorded (DeVries 1997; Hall & this genus (Callaghan 2001; Hall & Lamas 2001), but there are Willmott 2010; Greve et al. 2013). still some undescribed taxa in the Atlantic Forest of Brazil (D. In the second half of the 19th century there was an Dolibaina, pers. comm. 2015). increase in the rate of species descriptions, and most Expeditions (2012 and 2013) to a poorly sampled area of Riodinidae taxonomic diversity was described in this period high mountains in Minas Gerais state, southeast Brazil, uncov- ered an undescribed species of Thisbe (Riodinidae: Nymphidiini). Here, we describe this new species, providing *[email protected] information about its immature stages, natural history, behav- Early view version of record published on 1 July 2015 iour of adults and a discussion about its conservation status. © 2015 Australian Entomological Society doi:10.1111/aen.12159 2 L A Kaminski et al.

Figs 1–3. Habitat and adults of Thisbe silvestre sp. nov. in Serra do Caparaó, Alto Caparaó municipality, state of Minas Gerais, Brazil. (1) General view of the type locality; (2) male holotype; (3) female egg laying on Croton; note abdomen tip curved (arrow).

MATERIAL AND METHODS Cardoso’ (ZUEC), Universidade Estadual de Campinas, Campinas municipality, São Paulo state, Brazil. Study sites and rearing Morphology This species was studied in the narrow stream valley of a montane cloud forest (Figs 1–3) along the ‘Vale Verde’ trail Morphological examination and measurements were made (20°41′S, 41°84′W, 1200–1300 m a.s.l.) at the Parque using a Zeiss® Discovery V20 stereomicroscope. To study Nacional do Caparaó, Minas Gerais state, Brazil. Behavioural genitalia, abdomens were detached and soaked in a test tube of observations and collections were made in the field in Febru- hot 10% KOH solution for 5–10 min and dissections were ary 2012 and April 2013. Immature stages (eggs and larvae) stored in glycerol. Egg size is given as height and diameter. were collected directly on their host plant, Croton sect. Larval head capsule width was measured as distance between Cleodora () (Fig. 3). Larvae were reared in the most external stemmata, and maximum length for both plastic containers cleaned daily, with fresh plant material pro- larvae and pupae correspond to the distance from the head to vided every 2 or 3 days (following Kaminski et al. 2013). the posterior margin of the tenth abdominal segment in dorsal Observations and data were recorded on behaviour and devel- view. Measurements are given as a range. Colour patterns of opment times for all stages. To observe if and when the ant- immature stages were recorded using a digital camera in vivo organs were functional, we placed mature larvae (fifth and and a stereomicroscope was used for eggs and early instars. sixth instar) in a rectangular plastic arena (20 × 30 × 5 cm) Taxonomic nomenclature followed Callaghan and Lamas connected with a colony of Camponotus rufipes (Fabricius). (2004). We follow Kristensen (2003) for genitalia, Downey For details of rearing of ants in captive, see Kaminski and and Allyn (1980) for eggs, Stehr (1987) for general morphol- Rodrigues (2011). Observations and data were recorded on ogy of larvae and pupae and DeVries (1988) for ant-organs. behaviour and development times for all stages. Dry head Thisbe specimens from three public collections in Brazil capsules and pupal cases were retained in glass vials. Voucher were examined, including the Thisbe rupestre Callaghan type specimens of the immature stages were fixed in Dietrich solu- series (holotype and paratype), and additional specimens from tion and deposited in the Museu de Zoologia ‘Adão José north-east Brazil. Furthermore, we compared the new taxon © 2015 Australian Entomological Society A new myrmecophilous butterfly 3

Figs 4,5. Adults of Thisbe silvestre sp. nov., dorsal on the left, ventral on the right. (4) Holotype male from Serra do Caparaó, Alto Caparaó municipality, state of Minas Gerais, Brazil; (5) allotype female (ex-larva), same locality. Scale = 1 cm. with the Lamas collection of Neotropical butterfly type pho- two species by the narrower median white bands, and by the tographs at the MUSM (also available online in Warren et al. general ventral pattern, which is somewhat mottled in 2014), representing most currently relevant names and recog- T. silvestre but more ‘plain’ in T. irenea and T. molela. The nised species of Riodinidae (Callaghan & Lamas 2004). habitats of T. silvestre and T. rupestre are distinct; while The following collection acronyms are used: DZUP, T. rupestre is associated with seasonal savannah and open dry Departamento de Zoologia, Universidade Federal do Paraná, environments (Callaghan 2001; Nobre et al. 2008), T. silvestre Curitiba, Paraná, Brazil; UFP, Universidade Federal de was found in montane cloud forest (Fig. 1). Pernambuco, Recife, Pernambuco, Brazil; ZUEC, Museu de Zoologia da Universidade Estadual de Campinas, Unicamp, Description of adults Campinas, São Paulo, Brazil. Male Forewing length of holotype 19.0 mm; Paratypes: 19.0– RESULTS 21.0 mm (n = 5). Wing shape: Forewing semi-triangular with distal margin nearly straight; hind wing distal margin straight Thisbe silvestre sp. nov. to a very short rounded tail at anal angle. Dorsal surface: http://zoobank.org/urn:lsid:zoobank.org:act:AE43BB3E- Ground dark colour brown with white markings. Dorsal fore- 6CDE-4623-BB0C-9F4F2CE42B93. wing with submarginal row of small white spots between veins, large near apex, changing to almost obsolescent white marks near anal margin; a postmedian row of five white spots Diagnosis forming an arch in cells R3-R4 + R5 (spot 1), R4 + R5-M1 (2), Thisbe silvestre males are similar to those of Thisbe rupestre, M1-M2 (3), M2-M3 (4), M3-CU1 (5), spot 3 small and basally but can be easily distinguished from the latter by their larger positioned in relation to remaining; some individuals present a size, the lack of blue iridescent scales on the dorsal wing very small white spot in cell R2-R3; a postmedian white band surfaces and by the median white band on the dorsal hind from Cu1 to anal margin; discal cell brown. Hind wing with wings (Figs 2,4), which is straight in T. silvestre and convex in row of submarginal dark brown spots barely visible between T. rupestre.InT. silvestre the genitalia valvae are less robust veins outlined by fine white scales; a 2.0 mm wide median and longer than in T. rupestre, and the posterior margin of white band contiguous with forewing band; vein 2A covered uncus is rounded in T. silvestre and flat in T. rupestre. Females with conspicuous orange brown scaling in distal third. Ventral (Figs 3,5) are quite distinct from T. rupestre, whose females surface: Forewing light brown infused with pale grey scaling, lack the well-marked white band on both wing surfaces, being pattern same as dorsal surface, with some spots outlined with predominantly light brown with ochre bands (C.E.B. Nobre, light scales in half. Hind wing limbal area light brown pers. comm. 2014). Although, females of T. silvestre are with margin infused by white scales; white scales infusing into similar to those of Thisbe irenea (Stoll) and Thisbe molela basal half of wing from median white band; irregular orange (Hewitson), they can be easily distinguished from the latter brown spots outlined with light scales in basal half; fringe © 2015 Australian Entomological Society 4 L A Kaminski et al.

Figs 6–9. Male (6–8) and female (9) genitalia of Thisbe silvestre sp. nov. (6) Lateral view; (7) ventral view; (8) eighth sternite in ventral view; (9) ventral view. Scales = 0.5 mm. white and brown. Head: Brown dorsally, with long scales on male, with submarginal row of small white spots between frons, orbit and palpi white; eyes grey in life, changing to veins more marked; fringe white and brown. Ventral surface: reddish brown in museum specimens, covered with very short Ventral ground colour and pattern same as male, with submar- sparse hairs; antennae brown with white rings at shaft, a dark ginal row of small white spots between veins in forewing and brown club with an orange tip; a pair of small horn-like struc- limbal area more evident. Head and Body: Brown dorsally and tures on vertex covering antennae. Body: brown dorsally, beige white ventrally, same as male. Genitalia (Fig. 9): Sterigma and grey ventrally; legs light orange. Genitalia (Figs 6–8): heavily sclerotised; bursa copulatrix membranous; ductus Tegumen elongated, triangular in lateral view; vinculum bursae twice length of corpus bursae; signa elongated and broadly incomplete dorsally, forming a small narrow saccus symmetrical, form invaginated sclerotised spines. ventrally; uncus broad, not lobed, with posterior margin rounded. Gnathos ‘C’-shaped and as long as tegumen; valvae Description of immature stages well sclerotised, triangular in lateral view with lateral flanges, tips blunt; aedeagus long and slightly curved with cornutus; Egg (Figs 10,11): Duration 6–7 days (n = 2). Height 0.40– juxta narrower than aedeagus; eighth sternite semi- 0.44 mm; diameter 0.65–0.66 mm (n = 3). Colour light green quadrangular without lateral projections (Fig. 8). when laid, greenish white before hatching. General shape subcylindrical and flattened; exochorion with tiny projections and heptagonal cells. Micropylar area centred on top of ante- Female rior region. Very similar to male, with wings more rounded and with larger First instar (Fig. 12): Duration 4–5 days (n = 2). Head white spots. Forewing length of allotype 22.0 mm; paratypes: capsule width 0.32 mm (n = 3); maximum body length 20.0–22.0 mm (n = 2). Wing shape: Forewing and hind wing 1.9 mm. Head dark brown; body yellowish green, with pairs of same as male, with distal margins rounded. Dorsal surface: small reddish and white spots near abdominal spiracles; setae Ground colour dark brown with white markings same as in translucent, with brown base. Body dorsoventrally flattened © 2015 Australian Entomological Society A new myrmecophilous butterfly 5

Figs 10–21. Life history of Thisbe silvestre sp. nov. on Croton sp. (Euphorbiaceae). (10) Newly laid egg; (11) egg before hatching; (12) first instar; (13) second instar (arrow) tended by Pheidole worker; (14) third instar tended by Procryptocerus worker; (15) third instar tended by Camponotus sp.; (16) fourth instar; (17) fifth instar; (18) sixth (last) instar head in frontal view; (19) sixth (last) instar tended by C. rufipes, note the tentacle nectary organ everted (arrow); (20) in frontal view, showing tubercles and flat black spot on the prothorax; (21) pupa in lateral view. Scales = 0.5 mm (10–12), and 0.3 cm (13–21), respectively. Photos (16–21) by LL Mota.

© 2015 Australian Entomological Society 6 L A Kaminski et al.

Figs 22,23. Sequence of symbiotic interactions between Thisbe silvestre sp. nov. fifth instar larva and Camponotus rufipes ants. (22) Worker antennating the anterior part of larva body, note the everted anterior tentacle organ ATO (arrow); (23) everted nectary tentacle organ TNO (arrow) after repeated antennation by ant on the A8 segment. Scales = 0.3 cm. Photos by LL Mota.

with well-delimited segments in dorsal view; long plumose Sixth (last) instar (Figs 18,19): Duration 7–8 days (n = 4). setae laterally; remaining dorsal and subdorsal setae short. Head capsule width 2.00–2.46 mm (n = 3); maximum body Openings of anterior tentacle organs (ATOs) present on length 2.8 cm. General colour pattern and morphology same as metathoracic segment, and openings of tentacle nectary fifth instar. organs (TNOs) located on A8, but these organs are appar- Pupa (Figs 20,21): Duration 14–16 days (n = 3). Maximum ently not functional. Perforated cupola organs (PCOs) are length 1.8 cm, width at A1 0.50 cm (n = 3). Body dirty grey– located dorsally. Spiracle on the A1 segment is latero- greenish, with two conspicuous flat spots, black on prothorax ventral, whereas those on segments A2–A8 are in a dorsal and reddish brown on mesothorax and A1 abdominal segment. position. Black spiracles surrounded by white margins. Tegument Second instar (Fig. 13): Duration 5–6 days (n = 3). Head entirely corrugated, with several small tubercles, including capsule width 0.48–0.50 mm (n = 4); maximum body length basilars on basis of wings, near ocular area and associated with 3.4 mm. Head dark brown; body grey green, with small white abdominal spiracles in supra and subspiracular position. Body spots dorsally and pairs of reddish spots near spiracles; anal with several scattered stellate setae and clusters of papilliform shield with a longitudinal dark band. Body dorsoventrally setae on prothoracic and subspiracular tubercles. A silk girdle- flattened with well-delimited segments in dorsal view; crossing pupa over A1, near flat area. Fused A9 and A10 prothoracic shield covering almost completely covering head; segments constitute ventrally flattened cremaster, which has ant-organs as in first instar, but functional. short crochets in ventral position. Third instar (Figs 14,15): Duration 5–6 days (n = 4). Head capsule width 0.60–0.70 mm (n = 8); maximum body length Behaviour and natural history 4.5 mm. General colour pattern and morphology same as second instar, but with more numerous and enlarged setae. All Adults of Thisbe silvestre sp. nov. were found along a 200 m ant-organs present, including ATOs, TNOs, PCOs, dendritic stretch of a montane perennial stream (Fig. 1). In the first setae and one pair of vibratory papilla on anterior border of expedition, in February 2012, a single male was collected, and prothoracic shield. one female was observed in oviposition behaviour (Fig. 3) Fourth instar (Fig. 16): Duration 7–8 days (n = 4). Head (described below). During this expedition, dozens of plants capsule width 0.94–1.00 mm (n = 3); maximum body length were unsuccessfully inspected in search of eggs or . 9.8 mm. Head brown with white marks; body grey green In the second expedition, in April 2013, males were observed similar to third instar, but without longitudinal dark band along the same sector of the river as in the previous year, and on anal shield; prothoracic shield with two reddish about 20 larvae were found on the same plants inspected in bands. General morphology same as third instar, including 2012. ant-organs. Males (n = 5) were collected at one specific sector of the Fifth instar (Figs 17,22,23): Duration 7–8 days (n = 4). river, perching on a tree about 10 m above the ground. This Head capsule width 1.40–1.82 mm (n = 6); maximum body point was particularly suitable for collecting these butterflies, length 1.48 cm. Head brown with white marks and black areas because the presence of rocks and the natural slope of the dorsally; body greyish green and covered by black and white stream helped the access to the high perching site near the tiny setae that confer a grainy appearance. General morphol- canopy. Throughout the day, several Riodinidae, ogy similar to fourth instar including ant-organs, but body and Hesperiidae species alternated occupancy of this perch setae, more numerous and enlarged, including a row of flexible site. Males of T. silvestre appeared in the perch between 13:00 dendritic setae on anterior margin of prothorax. and 15:00 h and were observed chasing any butterfly that © 2015 Australian Entomological Society A new myrmecophilous butterfly 7 crossed the stream. The only feeding behaviour observed was Allotype: Female (ex-larva, Fig. 5) from Vale Verde, Parque a male that visited flowers of Vernonia sp. Schreb. Nacional do Caparaó, Alto Caparaó, 1300 m, Minas Gerais, () in the late afternoon (Fig. 2). Brazil, also deposited in the ZUEC. Labels on the allotype (six Oviposition behaviour was observed only once between labels separated by transverse bars): / ALLOTYPUS / Vale 12:30 and 13:00 h, on Croton sp. (Euphorbiaceae). In the Verde, Base Alto Caparaó, Parque Nacional do Caparaó, Alto pre-alignment phase, the female was observed inspecting the Caparaó, MG: Brasil, 20°25′S 41°50′W, 24–25-IV-2013 – host plant, landing above mature leaves of the host plant, and 1200–1300 m, EX-LARVA – Kaminski leg / Allotypus – soon after, laying an isolated egg on the abaxial leaf surface Thisbe silvestre Kaminski, Iserhard & Freitas det. 2015 / LAK (Fig. 3). The host plant is common along the riverbanks and – 176 / DNA voucher LAK-176 / ZUEC LEP 9180 /. along the access trail. Eggs and larvae were found on plants Paratypes: All same place as holotype: 1 male, 6-II-2012 from 1.5 to 6 m in height. Larvae were found resting on the (LAK – 070) (DNA voucher NS0081), Freitas, Kaminski & abaxial leaf surface, with the head close to the extrafloral Iserhard leg., ZUEC LEP 9181; 3 males (LAK-173, LAK-174, nectaries (Figs 13–15). First to third instars larvae fed by LAK-175), 24–25-IV-2013 ZUEC LEP 9182, ZUEC LEP scraping the lower surface of the leaves. After the third instar, 9183, ZUEC LEP 9184; 2 females (LAK-182, LAK-183), larvae ate the entire leaf producing oblong leaf damage. 24–25-IV-2013, ex-larva, ZUEC LEP 9185, ZUEC LEP 9186. In the field, first instar was not tended by ants but from the second instar the tentacle nectary organs (TNOs) became func- tional, and larvae were occasionally observed being tended by DISCUSSION ants (Figs 13–15,19,22,23). A total of four ant species were recorded tending T. silvestre larvae, including one species of Several aspects of the natural history of Thisbe silvestre sp. Camponotus Mayr, (Formicinae), one Pheidole Westwood one nov., including host plant use (Croton sp., Euphorbiaceae) Procryptocerus Emery, (Myrmicinae), and one Tapinoma facultative interaction with ants, the immature and adult mor- Foerster (Dolichoderinae) – none of which were identified to phology, are very similar to those described for Thisbe irenea, species. Larvae were commonly found without tending ants the type species of the genus (see DeVries 1988, 1991). This (∼50%), and when tended, no more than one ant per larva was justifies the placement of the new species in the genus Thisbe. observed. Temporal turnover of tending ants was also We hope that the information provided on the natural history recorded: a was tended by Procryptocerus sp. 1 day and morphology of T. silvestre may help to clarify the tax- and by Camponotus sp. the following day. In the laboratory, onomy and systematics of Thisbe in the future, particularly C. rufipes workers antennated the entire larval body, but espe- because this species, and the phenotypically similar cially the posterior region near the TNOs, as well as the ante- T. rupestre, have not been included in previous phylogenetic rior region where the anterior tentacle organs (ATOs) are analyses (Hall & Harvey 2001; Penz & DeVries 2001). located (Figs 22,23). The antennation apparently stimulated Thisbe silvestre was not represented in three major scien- the eversion of the ATOs; this eversion was observed only in tific collections in Brazil. Because this is a large and con- larvae that were tended by ants (Fig. 23). The ATO eversions spicuous species, and taking into account that the Atlantic provoked clear alterations in ant behaviour, such as opening of Forest is relatively well sampled for butterflies (Santos et al. the jaws and an increase in activity and aggressiveness. 2008), it may seem surprising that this species has not been previously collected, maybe by their high perching behav- Etymology iour. On the other hand, most inventories carried out in Atlantic Forest sites are very specific and localised, thus The specific epithet is masculine Latin adjective for ‘forest some regions of this biome are unknown for butterflies. dweller’, alluding to the forest habitat of this species, which There is a need of specific inventories to fulfil these gaps of contrasts with the morphologically closely related Thisbe information. However, its apparent rarity, patchiness and rupestre that is known to inhabit rock outcrop vegetation limited geographical distribution explain the fact that (‘campos rupestres’ in Portuguese), savannahs and other open T. silvestre had remained undetected. habitats. The mountainous area of Serra do Caparaó represents the northern limit for various butterfly species that occur in the Type material highlands of south-eastern Brazil, including threatened butter- Holotype: Male (Fig. 4) from Vale Verde, Parque Nacional do flies such as quadra (Schaus) and Paulogramma Caparaó, Alto Caparaó, 1300 m, Minas Gerais, Brazil. Depos- hydarnis (Godart) (Nymphalidae) (Freitas et al. 2014; Gomes ited in the Museu de Zoologia da Universidade Estadual de et al. 2014). Although there are many common species of Campinas (ZUEC), Unicamp, Campinas, São Paulo, Brazil. Croton in the study area, we found larvae of T. silvestre only Labels on the holotype (four labels separated by transverse on a single Croton species, from the Cleodora section, sug- bars): / HOLOTYPUS / Vale Verde, Base Alto Caparaó, gesting a high degree of host specialisation. Some Croton Parque Nacional do Caparaó, Alto Caparaó, MG: Brasil, species in this section are recognised as having restricted geo- 20°25′S 41°50′W, 24–25-IV-2013 – 1200–1300 m, Kaminski graphic distributions (see Caruzo & Cordeiro 2013), suggest- & Iserhard, leg / Holotypus – Thisbe silvestre Kaminski, ing that host plant specialisation might be another factor Iserhard & Freitas det. 2015 / ZUEC LEP 9179 /. explaining the narrow geographic distribution of T. silvestre. © 2015 Australian Entomological Society 8 L A Kaminski et al.

Although the present data are not to evaluate the conserva- Bates HW. 1859. Notes on the South American butterflies. Transactions of tion status of this species, the fact that only a single population the Entomological Society of London 2, 1–11. Brévignon C. 2010. Description de nouvelles espèces du genre Theope is known in the Atlantic Forest suggests a restricted geo- Doubleday, 1847 originaires de Guyane française. II – Le groupe de graphic distribution (one important criterion to categorise Theope thestias Hewitson, 1860 (Lepidoptera, Riodinidae, endangered species as threatened under the criteria of the , Nymphidiini). Lambillionea CX, 320–333. IUCN 2012). Accordingly, as suggested for another endan- Callaghan C & Lamas G. 2004. Riodinidae. In: Checklist: Part 4A. Hesperioidea – (ed. G Lamas), pp. 141–170. Associa- gered species in the same area (see Freitas et al. 2014), con- tion for Tropical Lepidoptera, Scientific Publishers, Gainesville, servation management actions should take the conservation Florida, USA. status of this species into account. This includes the reduction Callaghan CJ. 1978. Studies on restinga butterflies. II. Notes on the popu- of human disturbance to low levels in the area where the lation structure of Menander felsina (Riodinidae). Journal of the Lepidopterists’ Society 32, 37–48. species occurs (e.g. tourism and visitation), especially along Callaghan CJ. 1983. A study of isolating mechanisms among Neotropical the 200 m sector where adults and larvae were observed as butterflies of the subfamily Riodininae. The Journal of Research on being common in April. Other actions include reduced vegeta- the Lepidoptera 21, 159–176. tion management, scheduled cleaning of the edges and trails Callaghan CJ. 2001. New riodinids from the Central Brazilian plateau (Lepidoptera, Riodinidae). Revista Brasileira de Zoologia 18, 765– and restricted use of fire by visitors. Besides management 778. actions, it is crucial to search for additional populations of Callaghan CJ & Nobre CEB. 2014. A new species of Herrich– T. silvestre in areas of suitable habitat, such as the region of Schaeffer from Northeast Brazil (Lepidoptera, Riodinidae). Zootaxa 3780, 558–566. Serra do Brigadeiro, in Minas Gerais state. Caruzo MBR & Cordeiro I. 2013. Taxonomic revision of Croton section The present study exemplifies how much there is still to be Cleodora (Euphorbiaceae). Phytotaxa 121, 1–41. discovered in a threatened biome like the Atlantic Forest, con- d’Abrera B. 1994. Butterflies of the Neotropical region, part VI, sidered a hotspot of diversity in the world. If the current rates Riodinidae, Hill House, Ferny Creek, Victoria, Australia. DeVries PJ. 1988. The larval ant-organs of Thisbe irenea (Lepidoptera: of habitat loss are not diminished, it is likely that many species Riodinidae) and their effects upon attending ants. Zoological Journal will become extinct even before they are recognised. of the Linnean Society 94, 379–393. DeVries PJ. 1991. Mutualism between Thisbe irenea butterflies and ants, and the role of the ant ecology in the evolution of larval–ant associa- ACKNOWLEDGEMENTS tions. Biological Journal of the Linnean Society 43, 179–195. DeVries PJ. 1997. The butterflies of Costa Rica and their natural history II: Riodinidae, Princeton University, Princeton, New Jersey, USA. We thank Waldomiro de Paula Lopes, who facilitated the work Dias FMS, Dolibaina DR, Mielke OHH & Casagrande MM. 2014. Two in the Parque Nacional do Caparaó; ICMBio for providing the new species of Esthemopsis C. Felder & R. Felder, 1865 research permits (SISBIO n° 10802-5); and Ricardo Campos, (Lepidoptera: Riodinidae: ) from southeastern and Karla Yotoko and the Field Ecology Course of ‘Universidade northeastern Brazil, with taxonomic comments on Esthemopsis teras (Stichel, 1910) stat. rev. and Esthemopsis pallida Lathy, 1932 stat. Federal de Viçosa’ for the logistic support. We thank Luísa L. nov. Zootaxa 3784, 148–158. Mota who kindly took some beautiful photos; Ken Walker, Dolibaina DR, Dias FMS, Mielke OHH & Casagrande MM. 2013. Tax- Keith Willmott and Roger for critically reading the manu- onomy of the ‘Synargis axenus complex’ belonging to the ‘Synargis script; Maria B. R. Caruzo for host plant identification; regulus’ species group, with a phylogenetic reassessment of the genus Synargis Hübner, 1819 (Lepidoptera: Riodinidae: Nymphidiini). Zoo- Thamara Zacca and Diego R. Dolibaina for providing pictures logical Journal of the Linnean Society 168, 427–451. and dissections of types deposited in the DZUP; and Carlos E. Dolibaina DR, Dias FMS, Mielke OHH & Casagrande MM. 2014. A new B. Nobre for sharing unpublished data on Thisbe rupestre. species of (Lepidoptera: Riodinidae: ) from North- LAK was supported by FAPESP (10/51340-8), CNPq eastern Brazil. The Florida Entomologist 97, 1208–1212. Downey JC & Allyn AC. 1980. Eggs of Riodinidae. Journal of the Lepi- (163119/2013-9) and CAPES (3200-14-0). CAI thanks dopterists’ Society 34, 133–145. FAPESP (2011/08433-8). AVLF was supported by CNPq (fel- Freitas AVL, Kaminski LA, Iserhard CA et al. 2014. Paulogramma lowship 302585/2011-7) and the National Science Foundation hydarnis (n. comb.) (Nymphalidae: ): distribution, system- (DEB-1256742). This publication is part of the RedeLep atic position, and conservation status of a rare and endangered but- terfly. 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