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The Cissia confusa species‐group in Costa Rica and Trinidad (: )

Article in Zoological Journal of the Linnean Society · June 2008 DOI: 10.1111/j.1096-3642.1983.tb01162.x

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The Cissia confusa species-group in Costa Rica and Trinidad (Lepidoptera: Satyrinae)

M. C. SINGER, P. J. DEVRIES*

Department of

AND

P. R. EHRLICH

Department of Biological Sciences, Stanford University, Stanford, California 94305, U.S.A.

Received February 1983, accepledfor publication May 1983

The generic names Euptychia, Cissia and Argyreuptychia are reviewed in relation to the Cissia confusa species-group in Costa Rica and Trinidad, for which the generic name Czssia Doubleday is formally resurrected. Three new species of Cissia are described, C. confusa itself is resurrected from synonymy, three neotypes and three lectotypes are designated, and six new combinations established. On the basis of early instar morphology three subgroups are recognized and the general biology of the confusa species-group is summarized. A key to species in Costa Rica is provided and the ecology of the adults is reviewed.

KEY WORDS-Satyrinae - Euptychia - Argyreuptychia - Cissia - lepidopterous early stages

hostplants - new species - Costa Rica ~ Trinidad - habitats - natural history.

CONTENTS

Introduction ...... 102 Abbreviations ...... 102 Cissia Doubleday, gen. rev., ...... 102 The separation ofmyncea Cramer and confusa Staudinger...... 103 The discovery of three new Cissia species ...... 104 Subgroups of the C. confisa-group ...... 106 The species of the Cissia confusa-group ...... 106 The labe-subgroup...... 106 The confusa-subgroup ...... 111 The gomzi-subgroup ...... 114 Subgroup incertis ...... 115 A key to thr adults of the Cissia confusa-group in Costa Rira ...... 115 Discussion ...... 1 I6 Acknowledgements ...... 1 17 References...... 118

* Direct correspondence and offprint requests to: Philip J. DeVries, Department of Zoology, University of Texas, Austin, Texas 78712, USA. 101 0024-4082/83/100101+ 19 $03.00/0 0 1983 The Linnean Society of London 7 102 M. C. SINGER ET AL.

INTRODUCTION The satyrine Euptychia confusa was described by Otto Staudinger (1887) from specimens from Panama. Staudinger also illustrated the species but unfortunately, it was labelled Euptychia myncea (Cramer) on the plate (Staudinger 1886). As a result, Staudinger’s species has defied correct identification ever since. To complicate matters, there are a number of other species that resemble confusa, fly in the same habitats, and have been misidentified in museums and in the literature. We call these the ‘confusa-group’, and endeavour here to clarify the taxonomic confusion that has arisen over them. We also describe three new species from Costa Rica, their early stages and hostplant associations where known, and provide a key to the confusa-group species of Costa Rica. Some differences noted in the early stages suggest that, ultimately, the members of the confusa-group may prove not particularly closely related at all.

ABBREVIATIONS The following abbreviations of museums are used throughout: AME = Allyn Museum of Entomology, Sarasota; BMNH = British Museum (Natural History), London; CM = Carnegie Museum, Pittsburgh; MNHU = Museum fur Naturkunde der Humboldt Universitat, Berlin; MNCR = Museo Nacional de Costa Rica, San Jose; USNM = United States National Museum of Natural History, Washington, D.C.. The following abbreviations of anatomy are used: FW = forewing; HW = hindwing.

CISSIA DOUBLEDAY, GEN. REV. Euptychia Hubner, 1818: 20, sensu auctt., in part. Cissia Doubleday, 1848: 33. Type-species: Papilio clarissa Cramer, by monotypy. Argyreuptychia Forster, 1964: 123. Type-species Papilio penelope by original designation. Cissia Doubleday; Hemming, 1967: 1 16; Miller, 1968: 92-93 (Argyreupt_vchiacited as synonym). In much of the older literature concerning the confusa-group, all species are placed in the genus Euptychia Hubner. Forster (1964),however, in his account of mostly Bolivian Satyrinae, subdivided Euptychia (mainly on genitalic characters), erecting a number of new genera. Our rearing studies are consistent with some of his subdivisions. The type-species of Euptychia (mollina Hiibner) is a member of a very distiqct group of butterflies that feed on mosses and lycopsids as larvae (Singer, Ehrlich & Gilbert, 1971, and unpubl. data). All Euptychia sensu stricto reared by us ijesia Butler, westwoodi Butler, mollina and insolata Butler & Druce) have green larvae that bear projections along the dorsum. The head-forks are distinctive and the green pupae all have the same general ‘monkey face in a dunce-cap’ appearance (see Fig. 4 A,B). The larvae and pupae of the confusa- group are quite different (see below). Nonetheless, it must be recognized that Forster made several errors, two of CISSIA CONFUSA SPECIES-GROUP 103 which are pertinent here. First, one of his new genera, Argyreuptychia, has Papilio penelope as the type-species. As pointed out by Miller (1968), Forster overlooked the available name Cissia Doubleday, which has Papilio clarissa Cramer as type- species. Papilio clarissa Cramer is considered to be a synonym of Papilio penelope Fabricius (Hemming, 1967; see also below) and therefore the name Argyreuptychia becomes a synonym of Cissia. Forster’s second error was to synonymize confusa Staudinger with labe Butler. While our comparison of the lectotypes of both confusa and labe as well as the morphology and ecology of the early stages (see below) leaves no doubt that they are distinct species, there arises the problem of a generic name for confusa. According to data presented here, it appears the penelope and lube are more closely related to each other than they are to confusa. Like many of the genera erected by Forster ( 1964), Argyreuptychia is inadequately defined (and consider also that he placed hilara Butler in both Euptychia and Argyreuptychia). We therefore propose to formally resurrect the name Cissia for the confusa-group, as treated here. We do this in order that we may discuss the taxonomic problems that have arisen concerning confusa and similar species in Central America. Although not dealt with here, a number of well known Neotropical satyrines placed in various genera by Forster (1964) should probably also be placed in Cissia (e.g. libye Linnaeus and terrestris Butler). A thorough revision of all the butterflies formerly treated as Euptychia sensu lato is clearly needed. The species discussed here show an apparently unique characteristic in larval development amongst butterflies. All Cissia reared by us (with the exception of gomezi which was not reared from egg or first instar) have only four larval instars. As far as we are aware, all previously reported nymphalid butterfly species have five or more instars-including the Euptychia sensu strict0 noted above. Conceivably, this abbreviated number of instars may ultimately prove to be an excellent character for grouping together the species of the genus Cissia.

THE SEPARATION OF MYNCEA CRAMER AND CONFUSA STAUDINGER The figure of confusa Staudinger (1886: 80) (labelled myncea Cramer) is very accurate and compared well with the lectotype and other material (Figs 1C,D; 2A). Cramer’s (1780) original figure of myncea (Fig. 1A,B) is so inaccurate that it probably caused Staudinger to base confusa on a misconception. The five eyespots on the underside of the hindwing of myncea (Staudinger apparently assumed Cramer’s figures of Satyrinae were accurate) are not almost equal in actual specimens, and there is little difference between confusa and myncea in this respect. Cissia myncea and C. confusa are similar species with non-overlapping geographical distributions; the former is common in Trinidad and South America, while the latter is restricted to Central America. They are distinguished as follows: The underside hindwing ground colour is light grey- brown in confusa and dark brown in myncea, which also has wider transverse dark bands on both wings. Typical myncea is illustrated by Barcant (1970, as palladia) and Weymer (191 1, as ocypete). Because there are also consistent differences between the larvae and pupae of myncea and confusa, we regard them as separate species rather than strongly differentiated races.

106 M. C. SINGER ETAL. canopy. After close examination of the underside patterning and the genitalia, we are confident that it represents a new species, closely allied to C. pseudoconfusa, and is described below under the name joyceae.

SUBGROUPS OF THE CISSIA CONFUSA-GROUP While species can be identified accurately from adult wing patterns and genitalia, these characters show less clear-cut patterns of variation than the early stages, which we use here to separate subgroups within the confusu-group.

The lube-subgroup This comprises labe, palladia and penelope. The larval headcapsule has a highly granulate surface and bears stout, reduced horns on the epicranium (Fig. 4C,D,E). We have details of the pupa in labe and Penelope only: it is short and stout, with no pronounced projections or knobs (Fig. 4F).

The confusa-subgroup Of the species discussed in detail in this paper, myncea, confusa and pseudoconfusa belong here. In this subgroup the larval headcapsule has a relatively smooth surface with slender head-forks directed upwards and forwards. These head- forks form a rounded arc where they join the epicranium (Fig. 4G)H)I.J). The pupae are all identical in form, with twin rows of conical projections on the abdomen (Fig. 4K,L).

The gomeri-subgroup The headcapsule is smooth, with even longer head-forks than those typical of the confusa subgroup. These forks are directed more forwards than those of confusa and form a sharper angle where they join the epicranium (Fig. 4N). The pupa is also smooth (Fig. 4M). At present, gomeri is the only species we can assign to this grouping.

Subgroup incertis At least three more species belong within the confusa-group: dvmo Schaus and agnata Schaus, together with the new species joyceae. As we have not reared these species, we are unable to place them to subgroup with any confidence.

THE SPECIES OF THE CZSSZA CONFUSA-GROUP The lube-subgroup Cissia lube (Butler) comb. nov. (Figs 2F, 3F, 4C, 4F) Euptychia lube Butler, 1870: 250, pl.l., fig.2. Lectotype 8 (no abdomen), Panama: Santa Fe, Arci (BMNH), here designated [examined]. Euptychia lube Butler; Godman & Salvin, 1880: 121, p1.8, fig.3; 1901: 652; Riley & Gabriel, 1924: 30 [nec type!]. Euptychia lube var. (or form) labe; Weymer, 191 1: 200; Gaede, 1931: 452. Argyreuptychia labe (Butler); Forster, 1964: 123. [Euptychia drymo Schaus; Emmel, 1975: fig. 126. Misidentification.] REMARKS: Euptychia labe Butler was based on three specimens in the Osbert CISSIA CONFUSA SPECIES-GROUP 107

Figure 3. Male genitalia of Cissia spp. (in all figures, small white rectangle represents 100 pm. A, Cissia CO~ZUSQ(Staudinger) with valve intact; B, C. pseudoconfusa sp. nov. with intact valvae; C, C. gomen‘ sp. nov. with nearest valve removed; D, C. joyceae sp. nov. with the nearest valve removed; E, C. palladia (Butler) with intact valvae; F, C. labe (Butler) with intact valvae.

Salvin collection, two having been collected by Arck in Panama (Veraguas, at Calobre and Santa Fe), the third in the Polochic Valley, Guatemala, by Hague. In the BMNH we find no trace of the last named specimen. A female labelled “Veraguas, Panama, Arce” was later identified by Godman & Salvin (1901:652) as confusa. A male, lacking the abdomen, is labelled “Santa Fe, Panama, Arc&”. In addition, there is a second female labelled “Chiriqui, Panama. Arc&/E. labe Butl. type/Type, sp. figured/BM Type no. Rh. 3146, Euptychia labe 9 Butl.”. The last specimen is that quoted by Riley & Gabriel (1924: 30) as the ‘type’. Thus, of the three syntypes, one cannot be found, one lacks an abdomen, and the third no longer corresponds to the taxon in question, due to the subsequent restriction of Godman & Salvin (1901). Finally, the Figure 4. Larvae and pupae of Euptychia and Cissia spp. A, Larva ofE. westwoodi Butler from Finca la Selva, Costa Rica (note dorsal and lateral papillae and shape of head capsule); B, pupa of E. westwoodz from Finca la Selva (note ‘monkey in a dunce-cap’ form of pupa and prominent ridge at interface of abdomen and venter); C, larva of C. labe (Butler) from Finca la Selva, Costa Rica (note rounded, granulate head capsule with very reduced head-forks); D, larva of C. penelope (Cramer) from Trinidad (this is the type species of Cissiat note similarity to C. labe; E, larva of c. palladia (Butler) from Trinidad (note similarity of head capsule to C. labe); F, pupa of C. hbe from Finca la Selva (note more rounded shape and very reduced abdominal protuberances compared with C. confusa and allies); G, last instar larvae of C. confusa (Staudinger) (left) and C. pseudoconfusn sp. nov. (right) from Parque Corcovado, Costa Rica (note differences in coloration and dorsal patterning); H, last instar larva of C. confusa immediately after moulting (the dark chocolate colour will appear within 24 h and obscure the dorsal pattern); I, mature larva of C. myncea (Cramer) from Trinidad (note differences in patterning from C. confusa); J, detail of head capsule of C. confusa showing head-fork configuration; K, pupa of C. confusa (see text for details); L, pupa of C. pseudoconfusa (see text for details; M, pupa of C. gomexi sp. nov. (note differences from pupae of the confusa and hbe subgroups; N, mature larva of c. gorneri (note major differences in head capsule from congeners). ClSSlA CONFCJSA SPECIES-GROUP 109 specimen accepted as ‘type’ since 1924 does not appear to be part of the original type series! We accordingly have no option other than to select the damaged Santa Fe male as Lectotype, the confusa female becomes paralectotype, and the type status of the Riley & Gabriel specimen is rejected. All three specimens have been labelled accordingly. Cissia lube is distributed from Mexico to Panama, occurring in Costa Rica from sea level to about 800 m in association with disturbed areas in rainforest habitats. The adults (Fig. 2F) avoid deep shade and are usually found along riparian edges, old second-growth forest, or in large lightgaps in rainforest. We have reared C. lube from egg to adult on several unidentified bambusoid grasses. EGG:Similar to confusa (see below). LARVA:Ground colour of the mature larva is warm, mid-brown, with a slight blush of orange (cf. second instar of C. pseudoconfusa). A prominent dark brown dorsal midline runs from the head to in between the short caudal tails on the last segment. Four very‘dark dorsolateral spots are present on some larvae. Headcapsule highly granulate with reduced head-forks forming a pair of epicranial bumps (Fig. 4C). PUPA: Ground colour very dark purple-brown to almost black; wingpads and abdominal segments slightly marked with striations of light brown; abdominal protuberances very small and rounded. In general form (Fig. 4F) it is more rounded and stockier than species of the confusa-subgroup. HOSTPLANT: Paspalurn spp., Ichnanthus sp. (Poaceae: Bambusoidea) and probably other grasses. Females typically oviposit where the grasses are moderately tall (15 cm or more) and form dense clumps. A female alights, then closes her wings and drops inside the clump, laying single eggs on dead leaves and twigs, finally emerging some distance from the point of entry.

Cissia palladia (Butler) comb. nov. (Figs 2E, 3E, 4E.) Euptychia palladia Butler, 1866: 461, pl. 39, fig. 21. Lectotype Q, Brazil: Tapajos, H. W. Bates (BMNH), here designated [examined]. Euptychia palladia Butler; Weymer, 191 1: 200, [nec pl. 47b, as palladia]; Riley & Gabriel, 1924: 42, Gaede, 1931: 459. Argyreuptychza palladia (Butler); Forster, 1964: 123. REMARKS: Euptychia palladia was described by Butler from an unstated number of specimens. A single female now in the BMNH corresponds with the description; amongst others, it bears the labels “Tapajos, Amazons, H.W. Bates/ 9 Tapajos/Type of Species/BM Type no. Rh. 3145, Euptychia palladia 9 Butl.”. This specimen is here designated as lectotype, and has been labelled accordingly. Note: the species illustrated by Weymer (191 1: pl 47b) as “palladia” is not this species, but probably represents C. confusa. Cissia palladia (Fig. 2E) ranges from Nicaragua to the Amazon Basin and the Island of Trinidad. In Costa Rica it occurs very locally in association with deciduous forest on the Pacific slope, along riparian edges. In Trinidad it occurs at Andrew’s Trace (800 m elevation) and around the research station at Simla, near Arima, where it reaches peak abundance in the dry season. MALE GENITALIA (Fig. 3E). EGG:Round, pale yellow, with fine hexagonal markings on the surface. LARVA (Fig. 4E): All instars have the ground colour medium brown and bear an overall pattern of alternating light and dark rectangles that become more I10 M. C. SINGER ETAL. apparent in the later instars. This pattern differs from those seen in all other taxa of the confusa-group. The headcapsule surface is very rough and granulate; head-forks as in labe but somewhat larger. PUPA: Unfortunately, we have no record or notes on pupal morphology. HOSTPLANT: This species was reared from eggs obtained in the laboratory; the larvae were fed on numerous grasses.

Czssia penelope (Fabricius) (Fig. 4D). Papilio penelope Fabricius, 1775: 493. Neotype 3, [Surinam] ex Sehested CY 70nder Lund (Zoologisk Museum, Copenhagen), here designated. Papilio clarissa Cramer, 1780: 10, pl. 293, figs D,E. Neotype 9,Surinam: ex van Lennep (BMNH), here designated [examined; objective synonym of Pupilio penelope]. Cissia clarissa (Cramer); Doubleday, 1848: 33. EUPTYCHIACLARISSA (Cramer) (=penelope Fabricius, 1793); Butler. 1868: 16. Euptychia penelope (Fabricius) ( = clarissa Cramer, 1780); Kirby, 187 1 : 48; Weymer, 191 1: 200, pl. 476; Gaede, 1931: 459. Argyreupt_ychiapenelope (Fabricius); Forster, 1964: 123. Cissia penelope (Fabricius);Miller, 1968: 92/3. REMARKS: Papilio penelope was described by Fabricius from material in the Drury collection. Most of Drury’s material has been lost subsequently, except that portion that passed through Macleay to the Macleay Museum, Sydney, and CSIRO Canberra. According to R. I. Vane-Wright (pers. comm.) it seems most unlikely, from the Macleay records which he examined in Sydney during 1980, that the type material ofpenelope was amongst the Drury butterflies which reached Australia. We therefore presume that the original Fabrician material is lost or unrecognizable. A single male Cissia labelled ‘Penelope ex Am: Ma: Schmid’, ex Sehested & Tmder Lund Collection, exists in the Zoologisk Museum, Copenhagen. We have examined photographs (taken in 1983 by R. I. Vane-Wright) of this contemporary Fabrician specimen (almost certainly identified by Fabricius himselfi, and here designate it as neotype of Papilio penelope Fabricius. Papilio clarissa was described by Cramer from Surinam material then in the Caspar Stoll collection. Stoll’s material seems largely, if not entirely, lost or destroyed. The majority of extant specimens associated with Cramer and Stoll are those, mainly ex van Lennep, which have reached the BMNH via the Felders and Walter Rothschild (Vane-Wright, 1975). A female specimen now in the BMNH, which corresponds well with both the published Cramer figures and the original paintings (BMNH Cramer MS, volume 3, pl. 268, figs C,C), is labelled “Lennep Surin./No. 140 Clarissa Cr. IV. 293. D.E. [a ‘Cramer label’- Vane-Wright, 1975:56] /Felder Colln./Rothschild Bequest BM 1939-1 ”, is here designated as neotype of Papilio clarissa, and has been labelled accordingly. Conceivably, this specimen is part of the original Cramer and Stoll material, but this is not open to proof. The neotypes ofpenelope and clarissa both represent the same species. Weymer ( 19 1 1) and Gaede ( 1931 ) include Euptychia pitheus Moschler, 1882 (type-locality: Brazil, Paramaribo) as a variety or form ofpenelope; we have not examined the type-material of this nominal taxon. Museum specimens indicate that C. penelope is widely distributed in tropical CISSIA CONFUSA SPECIES-GROUP 111 South America. M.C.S. observed this species in Trinidad, where it is characteristic of highly disturbed sites, avoiding deep shade. EGG: Yellow, round and very small compared to the other species. Interestingly, the eggs of this species have no glue to stick them to the hostplant. LARVA (Fig. 4D): Polymorphic in all instars, ranging in ground colour from cream to deep orange-brown; some have undulating dark lines and dots dorsolaterally, while others lack this pattern. Headcapsule surface very rough and granulate; head-forks much reduced and similar to those of labe. PUPA: Ground colour very dark brown; abdominal protuberances smaller than in confusa and each bearing a small white dot. No other pattern. HOSTPLANT: Polyphagous on many species of grass in Trinidad. Females land on a hostplant, tap the leaf surface with their small forelegs and release an egg which falls to the ground.

The confusa-subgroup Cissia myncea (Cramer) comb. nov. (Figs lA,B, 41). Papilio myncea Cramer, [1780]: 10, pl. 293, fig. C. Neotype 8,Surinam: Fruhstorfer (BMNH), here designated [examined]. Xeonympha clerica Herrich-Schaffer, 1865: 69. [Unnecessary replacement name for, and junior objective synonym of Papilio myncea Cramer.] Eupptychia myncea (Cramer); Butler, 1868: 15; Weymer, 1911: 199; Gaede, 1931: 456. [Euptychia ogpete (Fabricius); Weymer, 191 1 : pl. 476. Misidentification.] [Euptychia palladia Butler; Barcant, 1970: pl. 13, fig. 9. Misidentification.] REMARKS: Papilo myncea was described by Cramer from Surinam material in the Stoll collection. No material directly attributable to this source has reached the BMNH via the Felder Collection-although two old specimens (one without data, the other labelled “Bahia”) ex Felder Collection do correspond well with Cramer’s published figures (allowing for inaccuracies noted above and even better with his original paintings (BMNH MS). Because it is essential to have a suitable type specimen for all the taxa in this confusing set of siblings, we have selected a specimen from the BMNH collection of myncea, which also corresponds quite closely with the old Felder material and the Cramer figure, to act as neotype. This male specimen bears the labels “Surinam, v.-ix., Fruhstorfer/Fruhstorfer Coll. BM 1937-285” and has been appropriately labelled as the neotype of Papilio myncea Cramer. Note: the specimen illustrated by Staudinger (1886: pl. 80) as “myncea” is the lectotype of confusa. In Trinidad, C. myncea (Fig. lA,B) occurs from sea level to at least 800 m elevation, in heavily-shaded sites such as overgrown cacao plantations or along trails through the forest. The butterfly does not enter Central America. EGG: Round, smooth and yellow, as in confusa. The female oviposits on the hostplant. LARVA (Fig. 41): First instar larvae are yellow without pattern; second instar yellow-green without pattern; third instar olive green to brown with a dark brown diamond pattern on the dorsum; fourth instar slightly lighter brown than confusa. The diamond pattern is prominent in the early fourth instar but later becomes faint. Headcapsule smooth, with the prominent brown head-forks tinged whitish, and shorter than those of confusa. I12 M. C. SINGER ETAL PUPA:Shaped and patterned as in confusa but lighter brown, without purplish hue; an irregular rectangular mark on the wingpads stands out very clearly. HOSTPLANT: In Trinidad several species of grasses, including Paspalum and two Panicum spp. (one is panicum pilosum Swartz).

Cissia confusa (Staudinger)comb. nov., sp. rev. (Figs 1 C, D, 2A, 3A, 4G, H, J, K) [Euptychia labe Butler, 1870: 250; Godman & Salvin, 1880: 121. In part.] [Euptychia myncea (Cramer); Staudinger, 1886: pl. 80. Misidentification.] Euptychia confusa Staudinger, 1887:225 Lectotype & Panama: Chiriqui, Ribbe (MNHU, Berlin), here designated. [examined]. Euptychia confusa Staudinger; Godman & Salvin, 1901 : 652/3. Euptychia lube var. confusa Staudinger; Weymer, 191 1:200, pl. 47b; Gaede, 193 1 :452. [?Euptychia palladia Butler; Weymer, 191 1:pl. 476. Misidentification.] [Argyreuptychia lube (Butler); Forster, 1964: 123. Misidentification: confusa given as synonym of lube.] REMARKS: Although Staudinger’s original description of Euptychia confusa mentions a specimen sent to him by Hewitson from Chiriqui, it is not clear how many other specimens he then had before him. A male which apparently belonged to his original series (and later dissected by Forster) was received on loan by P.J.D. (at BMNH). This specimen bears various labels, including “Chiriqui Ribbe/abgebildet [apparently it is the specimen figured by Staudinger, 1886, as myncea] /Origin. [mauve Staudinger type-label] /ex. collect. Staudinger/Praparat. Nr. 148 Zoolog. Staatssammlung Munchen”, and is here designated lectotype of E. confusa; it has been labelled accordingly. Because of the confusion over the identity of this species, we include a brief redescription of the adults. MALE: Eyes moderately hairy. Antennae densely scaled, light brown, with lateral tufts of white scales between the segments, distal 1/3 pale rufous brown on ventral surface. Forewing, underside ground colour greyish with two prominent reddish brown transverse bands (one post-basal, the other post- median) (Fig. 2A), with an ochraceous post-median patch. Hindwing, underside with five oddly-shaped ocelli which bear two metallic pupils and of which two (cells M!, Cu,?) are larger and darker than the others; area basal and distal to the ocelli greyish, with the submarginal grey area narrow; at the anal angle, a reddish brown submarginal line that touches the posterior ocellus. Genitalia (Fig. 3A). Length of FW: 21.8-24.22 mm (mean 22.8, n = 20). FEMALE:The female differs slightly from the male by having a rounder FW apex and a redder cast to the upperside. FW length: 22.5-24.2 mm. In Costa Rica, C. confusa is encountered uncommonly as solitary individuals in rainforest, from sea level to about 700 m elevation. This butterfly is intolerant of dry or open forest habitats. Its geographical range is confined to Mexico and Central America.

EGG: Shiny yellow, round. At Finca La Selva, Costa Rica, the egg is laid singly off the hostplant on associated dead vegetation. At Parque Corcovado, Costa Rica, about 20% are laid on the hostplant. LARVA (Fig. 4G,HJ): First instar larvae have a yellow body; second instars CZSSIA COflFUSA SPECIES-GROUP 113 ochraceous; remaining instars have the body brown; mature larva is chocolate brown with a light brown diamond pattern along the dorsum; a series of small white dots dorsolaterally; area behind headcapsule constricted; headcapsule surface relatively smooth; head-forks prominent and uniformly dark brown. During the day, first to third instar larvae rest on the underside of the hostplant leaves. Last instar larvae rest at the base of the hostplant, on the stem, with the head held to the ground. PUPA (Fig. 4K): Dark purplish brown with dark patterns on the wingpads and dorsum; more strongly marked on the wingpads than C. pseudoconfusa, less so on the head area; abdomen with pyramidal projections on the dorsum. HOSTPLANT: At Finca La Selva, Costa Rica, usually on Euterpe macrospadix (Palmae) seedlings, with a few on the seedlings of We@ageorgei (Palmae), and a single record on Panicum sp. (Poaceae). Occasionally found throughout the country on seedlings of Calathea spp. (Marantaceae).At Parque Corcovado, only found on Panicum sp. (Poaceae), but larvae can be transferred in the laboratory to all the hostplants listed above.

Cissia pseudoconfusa sp. nov. (Figs 2B, 3B, 4G,L) [Euptychia confusa Staudinger; Weymer, 191 1 :pl. 47b. Misidentification.] MALE: Head similar to C. confusa but differs by having antennae scaled black on the basal 2/3 with white tufts of scales laterally, terminal 7 segments black with pale rufous tip. Underside similar to C. confusa (see Fig. 2B) but differs distinctly by having a double submarginal line in the HW tornus that does not touch the posterior ocellus; the area basal and distal to the submarginal ocelli is cream-yellow. Genitalia (Fig. 3B). Length of FW: 19.0-21.1 mm, n= 10. FEMALE: There are very slight differences between the sexes. The female has the forewing apex more rounded, and the ochraceous patch on the underside is sometimes brighter. VARIATION: Forewing upperside occasionally has a distinct subapical ocellus ringed in ochre; postbasal and postmedial lines of the HW underside may converge along the anal margin without narrowing; postmedial area on HW underside may in some specimens be almost white. HOLOTYPE: Male, Costa Rica, Cartago, Turrialba, C.A.T.I.E., 600 m, 4 Sept. 1978, P.J. DeVries, (BMNH). PARATYPES: Ecuador: 9 La Chima, Rio de las Juntas, nr Bahahoyo, Prov. 10s Rios, June-July 1893, M. de Mathan, Oberthur Coll. (BMNH). Panama: 28, Bugaba, 800-1500 ft, Champion, Godman & Salvin Coll. (BMNH); 1 8 Canal Zone, Farfan 12 Sept. 1971, Coll. Jae (BMNH). Costa Rica: 1 d, Limon Prov. Rio Hondo and Highway, 60 m, 15 Sept. 1976. P.3. DeVries (MNCR); 1 8,Osa Peninsula, Puerto Jimenez, 8 Nov. 1977, P.J. DeVries (AME); 2 8,Prov. Heredia, 3 km. SW Puerto Viejo, 75 m., Finca la Selva, 29 May 1971, P. Opler (USNM); 1 9, Costa Rica [no further data], C. Underwood, Rothschild Coll. (BMNH). Mexico: Teapa, Tabasco, April, H.H. Smith, Godman & Salvin Coll. (BMNH). REMARKS: Cissia pseudoconfusa has been found from Mexico to Panama, with one record from Ecuador. In Central America, it occurs from sea level to about 700 m in association with a variety of habitats, ranging from rainforest to 111 M. C. SINGER ET AL. disturbed tropical deciduous forest. Encountered as solitary individuals, but apparently more common than C. confusa (evidence of museum collections and field observations). EGG: Similar to confusa. Laid singly on the hostplant where the hostplant is sparse and growing in deep shade. LARVA (Fig. 4G): Differs from confusa as follows: first instar yellow) second instar yellow with several pale dorsolateral spots and a dorsal stripe from behind the head to about one-third way along the back; third instar and mature larva progressively more strongly marked, having body creamy brown, never dark like confusa; headcapsule relatively smooth) head-forks prominent but having dorsal patterns lighter in colour than rest of the head; head-forks shorter than confusa. Feeding behaviour as in confusa. PUPA (Fig. 4L): Shape as in confusa, but coloured yellow to brown rather than purplish. HOSTPLANT: Only known to feed on Punicum sp. (Poaceae). At La Selva and Corcovado it is a specialist on this one species, but the butterfly also occurs in a number of sites where this grass is not present.

The gomezi-subgroup Cissia gomezi sp. nov (Figs 2C, 3C, 4M, N) MALE: Underside with general pattern and colour as in confusa but differs by having the two anterior ocelli on the HW with only a single pupil each; the middle two ocelli are indistinct yellowish smudges. The HW tornus has a double marginal line, of which the distal most line is thickest. The FW underside has a submarginal brown band that runs from the apical ocellus almost to the inner margin (Fig. 2C). Genitalia (Fig. 3C). FW length: 23.0-23.5 mm. FEMALE: FW length 22.5. The single available female differs slightly from males by having more strongly contrasting underside coloration and by the HW ocelli having somewhat wider ochre rings around them. How consistent these differences are is not known. HOLOTYPE: Male, ‘Costa Rica’, Rothschild Bequest, B.M. 1939-1 (BMNH). PARATYPES: Costa Rica: 2 3, same data as holotype; 1 9)Parque Nacional Corcovado, Osa Peninsula, Sirena, ex. larva 24 Sept. 1980, M.C. Singer & P.J. De Vries. All type-material in BMNH. We name this species, apparently only known from Costa Rica, for Luis Diego Gomez Pignaterio, in recognition of his extensive work on the flora and fauna of that country. EGG: Unknown. LARVA (Fig. 4N): The (presumed) third and fourth instars are bright orange) with darker diagonal and dorsolateral stripes. Head-forks longer and more forward projecting than any other species of the confusa-group. PUPA (Fig. 4 M): Ground colour pale brown, with fine striations of darker brown; a thick lateroventral brown line on abdomen, which is without protuberances. In general, shape more elongate (especially the abdomen) than any of the other taxa treated here. HOSTPLANT: We found a single larva feeding on an unidentified grass. CISSIA CONFUSA SPECIES-GROUP 115

Subgroup incertis We here transfer to Cissia two very rare Costa Rican species, both described by William Schaus in 1913, which to date have not been recorded from any other countries. Additionally, we include C. joyceae, sp. nov. Cissia drymo (Schaus) comb. nov. Euptychia dryrno Schaus, 1913:341, pl. 50, fig. 6. 2 6 syntypes, Costa Rica, Guapiles, W. Schaus (BMNH and USNM) [examined]. Euptychia drymo Schaus; Seitz, 1924:1029, pl. 193e; Gaede, 1931:445. REMARKS:Euptychia drymo was described from an unstated number of males, evidently collected by Schaus himself, from Guapiles, Costa Rica. According to the description, at least one specimen was “in British Museum”. We have traced single males in the BMNH and USNM: these we regard as syntypes. Note: the species illustrated by Emmel (1975: fig. 125) as “drymo” is not this species, but lube.

Cissia agnata (Schaus) comb. nov. Euptychia agnata Schaus, 1913:342, p1.50. fig.4. 6 syntype(s), Costa Rica: Guapiles, W. Schaus (type-depository uncertain) [not examined] Euptychia agnata Schaus; Seitz, 1924: 1029, pl. 1934 Gaede, 1931:437. REMARKS:Euptychia agnata was described by Schaus from an unstated number of males collected at Guapiles, Costa Rica. No type-material of agnata is in the BMNH; P.J.D. did not find any material of this species in the USNM in 1978- but conceivably the type(s) were then on loan. We have identified this species from the good original figure; the only old museum specimen we have seen is in the BMNH (Costa Rica: Estrella, C.H. Lankester). P.J.D. has collected C. agnata at Finca la Selva, Heredia Province and Carrillo, San Jose Province (Costa Rica).

Cissiajoyceae sp. nov. (Figs 2D, 3D) MALE:Differs from pseudoconfusa by the following: forewing underside with a richer ochraceous wash on the discal patch; grey marginal line running from R, to anal cell is wider and darker grey, and more strongly angled basally between the veins. Hindwing underside ground colour bone white, except for the area between the submarginal ocelli and distal margin which is wider and more yellow than in pseudoconfusa. The reddish brown marginal line in the tornus single as in confusa, but does not touch the posterior ocellus (Fig. 2D). Genitalia (Fig. 3D). FW length 20.3 mm. FEMALE: Unknown. HOLOTYPE: Male, Costa Rica: Heredia Province, Finca la Selva, canopy trap No. 1, 20 Nov. 1979, P.3. DeVries. Unique specimen deposited in BMNH. We name this butterfly in memory of Joyce, who loved nature and whose mother helped support our work.

A KEY TO THE ADULTS OF THE CISSIA CONFUSA-GROUP IN COSTA RICA The following key is prepared for the identification of all the species in the confusa-group in Costa Rica. While using the key, the reader should bear in mind that worn specimens tend to fade and rufous lines may then appear 116 M. C. SINGER ETAL brown. The characters of the hindwing tornus are generally most helpful for separation. Cissia dryma is keyed out twice because worn specimens of this rare species may show fading on the upperside. Forewing upperside with a discal rufous patch drymo (Schaus) Forewing upperside without discal rufous patch ...... 2 Forewing underside without submarginal ochraceous patch ...... 3 Forewing underside with submarginal ochraceous patch ...... 4 Hindwing underside having the largest subtornal ocellus with a single blue pupil and ringed thickly in ochre ...... agnata (Schaus) Hindwing underside having the largest subtornal ocellus with double pupils and finely ringed in ochre ...... gomezi sp. nov. Forewing underside ground colour almost entirely ochraceous ...... drymo (Schaus) Forewing underside ground colour grey but with an ochraecous patch .....5 Hindwing underside with a small ocellus posterior to the postmedial line along inner margin ...... palladia (Butler) Hindwing underside without small ocellus at inner margin ...... 6 Hindwing underside with a prominent rufous square in the tornus...... lube (Butler) Hindwing underside without prominent square in the tornus...... 7 Hindwing underside with a single rufous marginal line in the tornus .....8 Hindwing underside with a double rufous marginal line in the tornus .....pseudoconfwsa sp. nov Area between hindwing margin and submarginal ocelli cream-yellow and as broad as the second largest ocellus; marginal line in tornus does not touch posterior ocellus . . joyceae sp. nov. Area between hindwing margin and submarginal ocelli grey; marginal line in tornus touches posterior ocellus ....confusa (Staudinger)

DISCUSSION Butterflies in the genus Cissia are representative of the numerous tropical satyrine butterflies that primarily feed, as larvae, on grasses. Many of them are CISSIA CONFUSA SPECIES-GROUP 117 amenable for studies in both the field and in laboratory culture, and could thus be a useful group for investigation of the relationships between herbivorous and man’s most important group of crop plants. As this work demonstrates, much remains to be done on the alpha and beta taxonomy of the group before any such potential can be realized. It is clear from a study of the entire butterfly fauna of Costa Rica (DeVries, in prep.) that a revision of the Neotropical Satyrinae, especially ‘Eu/@hz’ sensu lato and allied genera is needed. The utility of early stages in ascertaining systematic relationships of butterflies has been recognized by numerous authors (e.g. Denis & Schiffermiiller, 1775; Muller, 1886; Edwards, 1868- 1897; Scudder, 1889; Moss, 1920, 1933; Kitching, 1983; DeVries et a!., in prep.). The present paper highlights the use of early stages for the confusa-group and, as far as we are aware, is the first taxonomic investigation of Neotropical satyrines using such data. Far greater effort in butterfly systematics should be committed to careful comparative studies of eggs, larvae and pupae. It is hoped that future revisionary studies will heed the advice of Denis & Schiffermuller (1775) (‘one eye on the larva, one eye on the adult’), and be critical of genera erected solely upon the basis of male genitalia, as has been recently commented upon for the North American butterflies (Ehrlich & Murphy, 1981). There is very little ecological work published on Cissia and allies. The oviposition behaviour has been investigated (Mackay & Singer, 1982; Singer & Mandracchia, 1982), and the hostplant specificity of the satyrine community in Trinidad has also received some attention (Singer & Ehrlich, 1983). However, it is unclear if the herbivory patterns revealed are widespread in Neotropical satyrine communities, since the basic population biology and life histories of most remain entirely unknown. Of the taxa treated here under the confusa-group, some are rare in museum collections and very seldom seen in the field. For example, C. drymo is known from only two specimens. Both were collected by Schaus at Guapiles, Costa Rica, and the species has not been found subsequently despite intensive field work and the examination of numerous collections. The reasons for its rarity are unknown, but there are two distinct ecological possibilities. Cissia drymo may have been restricted to the former rain-forest habitats at Guapiles, as certain other Costa Rican butterfly species appear to have been. Unfortunately, Guapiles now has only secondary vegetation and banana plantations, hence the question is academic. A second, and more hopeful possibility, is that drymo may be a forest canopy species, as found for a number of rare Costa Rican butterflies in a study of the rain-forest canopy and understory (DeVries, unpubl. data). The Costa Rican Atlantic slope habitat termed the Carrillo Belt (DeVries, 1980; DeVries & Chacon, 1981) should contain some of the species which are known from a few specimens-C. ugnata is very rare in collections and appears to be restricted to this area.

ACKNOWLEDGEMENTS We thank I. A. Chacon, R. Hesterberg, P. A. Opler, G. Otis and G. B. Small for help collecting material from the field. D. J. Harvey photographed and printed the genitalic figures. The manuscript was greatly improved by information and suggestions from P. R. Ackery, D. J. Harvey, L. E. Gilbert, 118 M. C. SINGER ElAL. L. Patterson, G. Robinson and R. I. Vane-Wright. Dr H. J. Hannemann kindly sent material on loan from the MNHU, Berlin. The photograph of the original Cramer plate of Pupilio myncea is reproduced through the kindness of the BMNH. We also thank the staff of Servicios de Parques Nacionales and the Museo Nacional for field assistance to DeVries. This paper was supported in part by a Fulbright Scholarship to DeVries while at the BMNH.

REFERENCES

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