Journal of the Lepidopterists' Sodell! 57(21 2003, 100-106

DESCRIPTION OF A NEW GENUS FOR "EUPTYCHIA" PECULIARIS (: ): IMMATURE STAGES AND SYSTEMATIC POSITION

ANDRE VICTOR LUCCI FREITAS Museu de Hist6ria Natural and Departamento de Zoologia, Instituto de Biologia, Universidade Estadual de Campinas, CP 6109, Campinas, Sao Pau lo 13083-970, BraziL Email: [email protected]

ABSTRACT. Based on distinct character states in life history and adult morphology, the monotypic genus, Taydebis, new genus with "Eu­ ptychia" peculiaris Butle r as the type is described, Analysis of the morphological characters and comparisons with four nearby genera suggest that the genus is closely aligned to or should he placed near Tay getis Hiibner and Pseudodebis Forster. Additional key words: life history, Poaceae, Pseudodehis, Taydehis, ,

Within the Neotropical Nymphalidae, the subf:lmily Fertile eggs were obtained from wild-captured fe­ Satyrinae is one of the most poorly understood groups, mal es that were confined in plastic bags. Larvae were with many systematic problems and undescribed reared in plastic containers cleaned daily, with fresh species, a f~lCt often noted in the literature (Forster plant material provided every two or three days (fol­ 1964, Miller 1968, DeVries 1987:257, Freitas 2002). lowing Freitas 1991). Observations and data were Adult characters have been useful for understanding recorded on behavior and development times for all relationships in some cases (Forster 1964, Miller stages. Dry head capsules and pupal castings were re­ 1968), but have been insufficient to resolve some sys­ tained in small glass vials. When there was sufficient tematic problems in the subLlmily. Since M iiller material, immatures were fixed in Kahle solution (1886) early stages have been shown as a useful source (AVLF collection). All measurements were made using of characters in systematics (Kitching 1985, a microscope fitted with a calibrated rnicrometric ocu­ Brmvn & Freitas 1994, Freitas et a1. 1997, Penz 1999) lar. Egg size is presented as length and diameter, and including for the Satyrinae (Singer et al. 1983, DeVries head capsule size is the distance between the most ex­ et a1. 1985, Freitas 2002, Freitas et al. 2002). ternal ocelli (as in Freitas 1991 ). Taxonomic nomen­ "Euptychia" peculiaris Butler 1874 is a prohlem clature follows Miller (1968) as modified by Harvey species from southeastern Brazil. This species occurs (1991), who treated the group as a subfamily, down­ at moderate elevations (800-1700 m) and is known ranking Miller's subfamilies and tribes to tribes and from only a few localities along the Mantiqueira moun­ subtribes, respectively. Nomenclature of wing veins tains and the Serra do Mar in the states of Sao Paulo follows Miller (1969), and ofbody setae follows Hin­ and Santa Catarina (Campo Alegre and Lages) (see list ton (1946). below). The record of Hayward (1973:256) from Mi­ Freitas, new genus siones, Argentina, requires further confirmation. Taydebis (Figs. 1, 2, Table 1) The present paper illustratcs and describes the criti­ cal morphological characters that distinguish this Type species: Euptychia peculiaris Butler, 1874. taxon, such as the wing venation and male genitalia. Diagnosis. Eyes hairy, reddish brown. Labial pal­ For the first time, the early stages are illustrated and pus one and a half times as long as head, brown with described in detail. A comparative discussion of sys­ light brown hairs. Antenna (8.5- 9.5 mm) up to 0.4 tematic relationships of "E. " peculiaris within the times the length of the costa; shaft dark brown dor­ Satyrinae is presented and a new genus, Taydebis, is sally, orange brown ventrally, sparse scaled dorsally; described. club not conspicuously developed, including eleven segments, vvith apical portion (last five segments) dark MATERIALS AND M ETHODS brown. Wing venation vely similar to Pseudodebis and Adults and immatures of "E. " peculiaris were stud­ Taygetis (Fig. 2). Both wings extremely rounded api­ ied at six different localities in Sao Paulo State, SE cally (Figs. 1,2). Brazil: banks of the Rio Tiete (Mogi das CnlZes, Description of adults. Male. Forewing length 20-23 mm, 700-800 m), Morro Grande Forest Reserve (Cotia, hjndwing length 16- 18 mm (n = IS). Body dark brown, abdomen 8.50-950 m) , Nlicleo Santa VirgInia (Sao Luis do ventrally light brown. U pperside ground color of wings medium brown, without marks, except felf a dark brown zigzag sub marginal Paraitinga, 900-UOO m), Campos do Jordao State line on both wings, and a light marginal line on the hindwing. Un­ Park (Campos do Jordao, 1500-1700 m), Intel-vales derside ground color lighter brown, three-tone: forewing discal area Park (Capao Bonito, 900-UOO m) and Grota Funda darker, hindwing distal halflighter. Two prominent scalloped brown lines crossing both wings 3.5 % and 60% ont from base; sub marginal Municipal Park (Atibaia, 900- 1000 m). regIOn of foreWin g with a diffuse darker brown area with four VOLUME 57, NUMBER 2 101

FIG J. Adult male (top) and female (bottom) of Taydebis peculiaris from Parque Estadual de Campos do Jordiio, SP.

minute light blue centered black ocelli bordered with orange in Female. Forewing length 22- 24 mm, hindwing length 17-22 spacP' R.5-Ml, MI-M2, M2-M:3 and M3-Cul; sub marginal area mm (n = 6). Body dark hrown, ventral abdomen light brown. Gen­ of hindwing with two prominent light blue centered black ocelli with eral color and pattern very similar to but in general lighter than that orang,e margins in 'pac,"s Rs-Ml and M I- M2, minute similar ocelli of males. Wings more rounded than in Illales. in spaces M2-M3 and M3-Cul, somewhat larger in Cul-Cu2 and Variation. Variation in the dorsal wing surfaces is very low, with Cu2--] A. A dark brown zigzag sub Illarginalline and a light Illarginal IllOSt variation being recorded on the underside. The size of the line are present on botl, wings. Male genitalia (Fig. 2) with an elon­ ocelli is variable in both sexes, and in some individuals only the two gated saccus, well developcd tegulllen and long pOinted unCllS. The prolllinent ocelli of the hindwillg can be seen without magnification. gnathos appears as two long pointed processes. Valvae trapezoidal Th" wing pattern is also variable, being weakly marked in some few ending with a single well developed point. Aedeagus with one large individuals hom Campos do Jordao. Some females have the under­ cornutus. Additionallllorphological characters (legs amI labial pal­ side ground color much more yellowish, especially in the sub Illar­ pus) are shown in Fig. 2. ginal and anal areas. 102 JOURNAL OF THE LEPIDOPTERISTS' SOCIETY

R3 R4

A D ~ ___-,M3

~------1Cu1

F

FIG 2, Morphological characters of Taydebis peculiaris, A, Male wing venation- hindwing above and forewing below; B, Male midleg; C, Female foreleg; D, \'laic foreleg; E, Male labial palpas; F, Lateral view of male genitalia; G , Aedeagus in lateral view,

Description of early stages. The follOwing descriptions are Third instar. Head black with green front and two short black based on immatures reared from Santa Virginia and Morro Grande, diverging scoJi on the vertex, Head capsule width 1.20- 1.42 mm The typical features of the immatures are very similar at these two (mean = 1.32 mm, SD = 0,053, n = 18); scoli 0.40-0,56 mm (mean = sites, Howp,ver, the number of instars was vmiable with four instars in 0.46 mm, SD = 0,050, n = 18), Body dark green with many longitu­ Morro Grande (Table 2) and five in Santa Virginia (see discussion), dinal yellow stripes; caudal filaments short. Maximum length 16 Females laid individual eggs when confined in plastic bags, suggest­ mm, Duration: 6-8 days, ing the ovipOSition of isolated eggs as the usual situation in nature, Fourth instar, Head green with a pair of short scoli with red Egg. Sphelical, light green, without visible ridges or marks under tips, Head capsule width 1.70-1.90 mm (mean = 1.78 mm, SD = the optic microscope. Height and diameter 0,92 mm (n = 30)~ Dn­ 0,055, n = 17); scali 0,58-0,74 mIll (mean = 0,6,3 mm, SD = 0,044, n ration: 5-7 days, = 17), Body emerald green, with many longitudinal thin yeJlow and First instar (Figs~ 3a, b, 4), Head capsule black, with enlarged light green stripes; candal filaments short, Maximum length 22 mm, chalazae, bearing a pair of short scoli on vertex, each with two long Duration: 9-10 days, narrow setae ending into a fine point. Third stemma larger than the Fifth (last) Instar (Fig, 3d-f) , Head the same as in previous in­ other stemmata, Head capsule width 0.66- 0,72 mm (mean = 0,69 star, Head capsule width 2.45-2,75 mm (mean = 2,54 mm, SD = mm, SD = 0,017, n = 30); scoli 0,08-0,12 mm (mean = O,ll mm, SD 0,090, n = JO ); scoli 0,75-0,88 mm (mean = 0,82 mm , SD = 0,0.39, n = 0,013, n = 30), Body beige, becoming light grecn after feeding, = 10), Body color same as fourth instar, Maximum length 33 mm, smooth, with many weak white longitudinal stripes and a pair of Duration; 9-10 days, short caudal filam ents, Setae XO, D, SD and L thickened with Pupa ( Fig~ 3g- h), Entirely green, elongated, smooth, with short clubbed tips; body chaetotaxy illustrated in Fig, 4, Maximum length ocular caps and slightly projecting alar caps bordered with a thin yel­ 5 mm, Duration: H-9 days, low line, Total length 12-14 mm, Duration 10 days (n = 7), Second instar (Fig, 3c), Head black with two diverging scoJi on veltex. Head capsule width 0,90-1.02 mm (mean = 0,97 mm , SD = Etymology. The name is a reduced combination of 0,039, n = 26); scoli 0,28- 0.40 mm (mean = 0,:34 nUTI, SD = 0,031, n = 26), Body slender, light green \\~th many longitudinal white stripes; Taygetis and Pseudodebis, pOSSibly the two most caudal Hlam ents short, Maximum length 10 mm, Duration: 7---8 days. closely related genera, c; r c ~ tTl :-1ClI Z c ~ IJ:) tTl ::v K)

TABLE l. Comparisons of Taydebis with related genera.

Forewing Hindwing Shape of Aedeagus: Shape Shape Saccus: ratio Species Eyes apex margin aedeagus length/width of uncus of gnathos Tegumen length/width Taydebis peculiaris 1 hairy rounded not wavy straight 12 straight, slender elongated, pointed pronounced 13 Pseudodebis griseola' few sparse rounded slightly wavy strongly curved 23 curved, broad elongated, pointed pronounced 11 hairs Psudodebis euptychidia3 hairy rounded slightly wavy straight 18 curved, slender short, rounded pronounced 6 Pseudodebis valentina' hailY rounded slightly wavy straight 12 curved, slender short, broad slightly pronounced 5 Taygetis kerea 5 hairy acute wavy straight 13 straight, broad elongated, pointed pronounced 7 3. 6 hairy tnmcate wavy straight 25 straight, slender elongated, pOinted slightly pronounced 8 Taygetis ypthima7 very short acute wavy straight 12 straight, slender elongated, pointed pronounced 4 s'parse hai rs in lateral portion ' hairy acute wavy straight 10 straight, slender elongated, pOinted not pronounced 6 Taygetis celia' hailY truncate wavy straight 10 straight, slender short, rounded not pronounced 4 Taygetis sylvia' hairy acute wavy straight 6 straight. broad elongated, pointed not pronounced 4 Taygetis echo 8 few sparse rounded wavy straight 14 straight, slender elongated, pointed not pronounced 5 hairs Harjesia blanda 4 hairy rounded wavy straight 8 curved, slender elongated, pointed not pronounced 4 Posttaygetis penelea" 9 hairy rounded wavy cu rved at the 15 straight, broad short, rounded not pronounced 4 distal end Source of material (all localities in Brazil): I- this paper; 2-Jaru, RO; 3-Linhares, ES; 4-Marechal Thaumaturgo, AC; 5-Jatai, SP; 6---Campinas, SP; 7-Campos do Jordao, SP; 8-A1ta Floresta, MT; 9- Morro do Diaho, SF.

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FIr. 3. Early stages of Taydebis peculiaris. a , b, First in star (lateral, dorsal); c, Second instar; d, e, f, Fifth (last) instar (lateral and two dor­ sal views); g, h, Pupa (lateral, ventral ). All specimens from Santa Virginia, SF.

Habits. This species is frequently found in grass DISCUSSION fields and swampy areas at medium to high altitudes, The supposed systematic position of Taydebis pecu­ independent of the conservation status of the area. liaris was based on both adult and immature morphol­ Oviposition behavior was not observed, and the host ogy. The male genitalia are very similar to two other plant in the fi eld is unknown. In the laboratOl)" larvae genera as discussed above, and different from the readily accepted the Carpetgrass Axonopus compres­ other series of genera of euptychiines. The last ins tar is sus (Sw. ) P. Beauv. ("gram a misioneira") , a common similar to that of Pseudodebis marpessa and some grass with soft leaves used in shaded lawns in Brazil. Taygetis (Murray 2001), but also has remarkable simi­ Systematic position. A genus near Taygetis and larities with Godartiana (unpublished results). The Pseudodebis (Table 1); distinguished from Pseudode­ first instar has long narrow setae on the head capsule, bis by the elongated pointed gnathos (short, rounded most similar to those of Posttaygetis penelea (D. Mur­ in Pseudodebis), the longer saccus and a straight un­ ray in prep. ), and different from Taygetis that has CllS in lateral view (curved in Pseudodebis) . Distin­ wide, Battened fan-shaped setae. The pupa also has a guished from most Taygetis by the longer saccus, few characters very similar to some Taygetis species, in­ rounded foreWing apex, and the presence of a well­ cluding the slightly projecting alar caps bordered with developed tegumen (weakly developed in most a yellow line (Young 1984, AVLF unpublished data). Taygetis ). Characters from immatures also support The general appearance of the immatures of T pe­ the affinities of Taydebis with Pseudodebis and culiaris is not divergent from those of most known Taygetis (see below). euptychiines, including first instar larva with two VOLUME 57, N UMBER 2 105

TABLE 2. Data Ii'om a larval lot of Taydebis peculiaris with only four instal's (Morro Grande, Cotia, SP).

Duration (days) Head capsule width (mm) Length of scoli (mm) Maximum length (mm) n 1st instal' 7-K 0.68 0.08-0.10 6 4 2nd instar 6-7 102 0.36-0.38 12 4 3rd illstar 6-7 16 0.44 17 4 4th instal' 10 2.28-2.22 0.62-0.66 30 :3 short head horns, an elongated striped mature larva Campos do Jordao), Fundac;ao Florestal (Intelvales), and Sabesp and a smooth pupa (Young 1984, DeVries et al. 1985, (Morro Grande).

DeVries 1987). The body setae with clubbed tips in LITERATURE CTTED the first instar are also present in many Satyrinae BHOWN, K. S. JR. & A. V L. FHEITAS. 1994. Juvenile stages ofIthomi­ (Murray 2001, and AVLF unpublished data from illae: ovelview and systematics. Trop. Lep. 5:9-20. IlIore than 60 neotropical species); their function is DEVRIES, P. J. 1987. The of Costa Rica and their natural still unknown. hist01Y. Papilionidae, l'icridae, Nymphalidae. Prince ton Uni­ versity Press, Princeton, New Jersey. Even if Taydehis is distinguishable from most DEVR1ES, P. J., 1. J. KITCH1NG & R. 1. VANE-WRIGIlT. 1985. The sys­ species of the nearby genera, the present scenario tematic position of' Antirrhea ancl Caerois, with comments on shows that the boundaries among these and other gen­ the higher classification of the Nymphalidae (). Syst. Entomol. 10:11-32. era in this series are still not established. A more care­ FORSTER , \V 1964. Beitrage zur Kenntnis del' Insektenhmna Bo­ ful comparison (Table 1) suggests that many different liviens XIX. Lepidoptera III. Satyridae. Verb±fentlichungcn del' taxonomic entities may be included under the genera zoologischcn Staatssammillng Miinchen 8:51- 188, pis. 27-35. Pseudodebis and Taygetis , including "Pseudodebis" FREITAS , A. V L. 1991. Varia<;iio morfo16gica, eiclo de vida e sis­ tematiea de Tegosa claudina (Eschscholtz) (Lepidoptera, griseola and "Taygetis celia", which should be placed Nymphalidae, Mclitaeinae) no Estado de Sao Paulo, Brasil. in two new different genera. Rev. bras. Entomol. 35:301-306. Thus, the correct position of Taydebis within the --. 2002. Immature stages of Eteona tisiphone (Nymphalidae: Satyrinae) . .I. Lepid. Soc. 56:286-288. Euptychini may well need filrther investigation, and FREITAS, A. V L., K. S. BROWN JR. & L. D. OTEHO. 1997. Juve nile additional cladistic studies (morphological anclJor stages of Cybdelis, a key genus uniting the diverse branches of the Eurytelinae. Trup. Lep. 8(1):29-34. molecular) could help to further clarify this placement. FREITAS, A. V. L., D. MURRAY & K. S. BROWN TR. 2002. Immatures, natural history and the systematic pos'ition of ACKNOWLEDCMENTS (Nymphalidae). J. Lcpid. Soc. 56:117-122. This study was conducted as part of a Post-Doctoral project on HAHVEY, D. J. 199] . Highe r classification of the Nymphalidae (Ap­ Satyrinae ecolot,'Y and systematics (BIOTA-FAPESP program, grants pendix B), pp. 2.55-273. Tn Nijhollt, H. F. (ed.), The development 98/0.5101-8 and 00/01484-1). I would like to thank Drs. Keith S. and evolution of butte rAy wing pattems. Smithsonian Press. Brown Jr. , Marcio Uehara-Prado, Gerardo Lamas, Lee D. Miller, HAYWARD, K. J. 1973. Catalogo de los ropal6ceros argentinos. Jacqueline Y. Miller, Debra Murray and Carla Penz for their help in Opera LilJoalla 23: l-318, 1 map. diverse phases of the mannsclipt. Drs. Lee D. Mille r and Jacqueline HI~noN, II. E. 1946. On the homology and nomenclature of the Y. Miller greatly improved the final version of the manuscltipt. T grate­ setae of lepidopterous larvae, with some notes on the phy­ fully acknowledge the invaluable assistance of the follOWing agcncics logeny of the Lepidoptera. Trans. R. Entomol. Soc. Lonclon to complete these studies: Instituto Florestal (Santa Virginia and 97: 1-37.

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S02 S01 S02 S:/' soy S01/ soy SOy S01" \t01 " 0 L1 _L1 " L1 /' I •, / •, • /\ L2 L1 L2 L1 /\ L2 L1 L2 L2~L1 L2 L1 ,SV1 ,SV1 / SV1 I SV1 SV1 ... /\ SV1 " - SV2 SV1 /SV2 ISV2 SV2, "" SV2

T1 T2 T3 A1 A2 A3-6 A7 A8 A9 A10

FIG 4. Chaetotaxy of the first instar lalVa of Taydehis peculiaris. 106 JOliHNAL OF THE LEPlDOPTEHISTS' SOCIETY

KlTCH INC, I. J. 1985. Early stages and the classification of the milk­ PEN!., C. 1999. Higher level phylogeny for the passion-vine butter­ weed butterflies (Lepidoptera: Danainae). Zool. J. Linn. Soc. flies (I\yrnphalidae, Heliconiinae) based on early stage and 85:1-97. adult morphology. Zoo!. J. Linn. Soc. 127:277-344. MILLER, L. D. 1968. The higher classification, phylogeny and zoo­ SINGER, M. c., P. r. DEVRIES & P. R. EHRLrCH. 1983. The Cissia geography of the Satyridae (Lepidoptera). Mem. Am. Entomo!' conji.lsa species-group in Costa Rica and Trinidad (Lepidoptera: Soc. 24:iii + 174 pp. Satyrinae). Zool. J. Linn. Soc. 79: 101-119. - --. 1969. Nomenclature of wing veins and cells. J. Res. Lepid. YOCNG, A. M. 1984. Natural history notes for Taygetis andromeda 8(2) ::37-48. (Cramer) (Satyridae) in Eastern Costa Rica. J. Lepid. Soc. MULLE R, W. 188fi. Sudamerikanisc\w Nymphalidenraupen: Ver­ .38:102-113. such eines naturlichen Systems de l' I\ymphaliden. Zoologische Jahrbucher (,lena) 1 (:3/4) :4J7- fi78, pIs 12-1.5. Receiued Jor publication 28 August 2002; revised find accepted 30 M URRAY. O. 2001. Innnatnre stages and biology of Taygctis Hub­ Novemher 2002. ner (Lepidoptera: Nymphalidae). Proc. Entomol. Soc. \Vasb­ ington 103(4):932-94.5.