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Journal of the Lepidopterists' Society 46(1), 1992, 44-53

PARADULCEDO, A NEW GENUS OF SATYRINAE (NYMPHALIDAE) FROM WESTERN COLOMBIA

Luis M. Constantino

Apartado Aereo 1676, Cali, Colombia

ABSTRACT. Paradulcedo, new genus, is proposed for the satyrid butterfly originally described as Callitaera mimica Rosenberg & Talbot. This little known species, endemic to western Colombia, has long been considered a member of the genus Cithaerias, from which it is quite distinct. Descriptions of the wing venation and male genitalia are given. Paradulcedo is compared to members of the genera Dulcedo, Pseudohaetera, Haetera, Cithaerias, and Pierella.

Additional key words: Dulcedo, Pseudohaetera, Haetera, Cithaerias, Pierella.

The tribe Haeterini contains five genera: Cithaerias Hiibner, Dulcedo d'Almeida, Pseudohaetera Brown, Haetera Fabricius, and Pierella Her-rich-Schaeffer. All five are confined to the Neotropical Region (Miller 1968, Masters 1970, Smart 1976, D'Abrera 1989) and all occur in Colombia. The butterflies of this tribe are, for the most part, readily distinguished from all other groups of the Satyrinae by having largely transparent wings with one or two ocelli and patches of color on the hindwing margin. The only exception to this description is the genus Pierella which has brown coloration on the upper wings and brilliant patches of color on the hindwing distal area with a variable number of ocelli. The genus Dulcedo is monotypic containing the single species D. polita (Hewitson, 1869) (Fig. 11), which ranges from Nicaragua to Western Colombia (DeVries 1987). The genus Pseudohaetera also contains a single species, P. hypaesia (Hewitson, 1868), distributed from Colombia to Bolivia (Weymer 1924, Brown 1942, Smart 1976) where it is confined to the Andean region between 1200-2500 m. This species is the only known high-altitude haeterine butterfly found commonly in cloud-forest relicts above 2000 m (Fig. 12). In Colombia, P. hypaesia occurs on the east slope of the western cordillera and on the central and eastern cordillera (J. A. Salazar pers. comm.).

In the last ten years, butterfly collectors working on the west side (Pacific slope) of the western cordillera have found a species that closely resembles P. hypaesia both in the color markings on the hindwing and in flight behavior. Rosenberg and Talbot (1914) described this species as Callitaera mimica from specimens collected at La Selva, located at 1400 m on the upper San Juan River, Choco, Colombia. This rare and poorly known species, endemic to western Colombia, has been retained in the genus Cithaerias virtually since its original description, and was listed in Cithaerias by D'Abrera (1989). Takahashi (1981), in a list of Haeterinae collected by two Japanese expeditions to Colombia, did not

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Cul

Figs. 1-4. Paradulcedo mimica (Rosenberg & Talbot). 1, 6 adult, upper side. 2, 6 genitalia, lateral view with aedeagus removed (aedeagus, lateral view, beneath). 3, 6 forewing venation. 4, 6 hindwing venation. Scale lines = 1 mm for Fig. 2 and 5 mm for Figs. 3-4.

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mention this species. In the most recent treatment of Neotropical Sa-tyridae (D'Abrera 1989), the male type of C. mimica Rosenberg & Talbot is illustrated. Keith S. Brown Jr. (pers. comm.) suggested that C. mimica could belong to the genus Dulcedo based on superficial observations.

As members of neither Cithaerias nor Dulcedo are believed to exist above 1500 m in the Andean mountains and because they have distinctive flight behaviors, I began a detailed study to determine the proper generic affinities of mimica. Comparisions of the genitalia, wing venation, and adult behavior of C. mimica with those of species representing its close relatives, Dulcedo, Cithaerias, Haetera, Pseudohae-tera, and Pierella, suggests that C. mimica does not belong to either Cithaerias or Dulcedo. The results of these studies led to the conclusion that C. mimica should be placed in a new genus closely related to Dulcedo and Pseudohaetera. I therefore propose the following new genus.

Paradulcedo Constantino, new genus

(Figs. 1-4, 5, 10, 22)

Type species: Callitaera mimica Rosenberg & Talbot, 1914:677.

Cithaerias mimica (Rosenberg & Talbot); D'Abrera, 1989:740.

Cithaerias gilmouri Okano; Okano, 1986:1.

Description. Male and female with same wing color pattern; female larger. Eyes: Naked, black-copper in life. Palpi: Slender, appressed to head, not extending beyond frontal vestiture; third segment small, one-seventh length of second. Antenna: Slender, eight-tenths length of forewing discal cell, comprising 47-48 segments, the terminal 7 slightly compressed without forming a club. Forewing (Figs. 1,3): Completely transparent with two strong lines running across; wing shape rounded (elongated in Cithaerias); M, and R3+5 connate at the point of origin from the discal cell (D) (in Cithaerias Mx departs from R3+5 and not from D; in Dulcedo Mx and R3+5 arise independently from the discal cell). Hindwing (Figs. 1, 4): Transparent with black markings (actually a strong, wide submarginal line) similar to Pseudohaetera; viewed ventrally, the markings are brown in Pseudohaetera, black in Paradulcedo; two ocelli on the hindwing margin (Cithaerias and Dulcedo have one); M3 curved, originating from discal cell (M3 straight in Cithaerias); Cuj and M3 originating separately from discal cell (arising from same point in Pseudohaetera and Haetera); discal cell acute distad (rounded in Dulcedo). Length of forewing (from base to apex): male, 23.5 to 26.0 mm (n = 11); female, 28.0 to 32.0 mm (n = 6).

Male genitalia (Fig. 2): Uncus curved and elongate without lateral projections (in Dulcedo there are lateral projections or horns); gnathos small (large and prominent in Cithaerias); aedeagus pointed apically (rounded in Dulcedo, Fig. 5; Table 1).

Relationships. Morphologically, Paradulcedo can be separated from Cithaerias by the characters noted in the description above. Although the wing venation of Paradulcedo shows affinities with Dulcedo, differences in genitalia indicate that they are not congeneric. On the other hand, the genitalia of Paradulcedo reveal affinities with Pseudohaetera, but the wing venation is quite different.

Type specimens. Lectotype, 6 and 9 COLOMBIA: Choco Dept, La Selva (Pacific slope), upper San Juan River, 1400 m (BMNH). Paralectotypes: All from Colombia, 1 <$, 1 9, Risaralda Dept., Pueblo Rico, 1580 m (BMNH); 1 6, Risaralda Dept., Siato, 1585 m (BMNH). All type material is in the Natural History Museum, London (BMNH).

Additional specimens. COLOMBIA: CHOCO: San Jose del Palmar, 1000 m, 2 S (J. H. Velez) (Museo de Historia Natural, Manizales), 2 9 (J. A. Salazar leg.), 1 6 (J. F. Lecrom

Table 1. Morphological characters to differentiate Paradulcedo from Cithaerias, Dulcedo, Hae

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Figs. 5-9. Male genitalia comparisons of five haeterine genera from Colombia. 5, Paradulcedo mimica (Rosenberg & Talbot) lateral view of genitalia with aedeagus removed (aedeagus, lateral view, beneath) Alto Calima, Valle. 6, Dulcedo polita (Hewitson), Alto Anchicaya, Valle. 7, Pseudohaetera hypaesia (Hewitson), Pefias Blancas, Farallones de Cali, 2000 m. 8, Haetera piera (Linnaeus), Rio Amacayacu, Amazonas. 9, Cithaerias aurorina (Weymer), Puerto Narino, Amazonas. Scale lines = 0.5 mm.

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leg.); El Tabor, 1 6 (J. F. Lecrom leg.); RISARALDA: Santa Cecilia, 900 m, 1 2 (J. H. Velez) (MHNM); San Antonio del Chami, 1600-2000 m, 2 2 (J. A. Salazar leg.); VALLE: Rio San Juan, km 56, 1200 m, 2 S, 6. viii. 1985 (L. M. Constantino leg.); Calima III, 1400 m, 1 6, 16.v. 1984 (E. Constantino leg.); Alto Rio Pepitas, 1600 m, 1 6, 15.vi. 1986 (M. Linares leg.); cerro Los Chancos, 1600 m, 2 6, 1 2, 10.vii.1983 (J. I. Martinez leg.); Rio Bravo, Calima, 1300 m, 1 8, 12.iv.1985 (L. M. Constantino leg.).

Natural History

Paradulcedo mimica is restricted to undisturbed premontane and montane forest (commonly cloud-forest relicts) on the west side (Pacific slope) of the western cordillera between 900 and 1600 m, with a mean annual precipitation of 5000-6000 mm. Its geographic distribution ranges from Choco to Cauca (Fig. 22). The range may include Narino and northwestern Ecuador, but there are no specimens reported from these two regions. The flight behavior is very similar to that of P. hypaesia, characterized by being slow and erratic, whereas that of Dulcedo polita (which occurs locally on the Pacific Coast in association with pluvial forest from sea level to 1000 m) is very fast and straight. Flight activity of P. mimica is during day light hours, from 1000 to 1500 h for males and from 1300 to 1600 h for females. The flight period is restricted mainly to the dry season between June and August, but in some years P. mimica can be found during the first rainy season between March and May. Despite many hours of field observation, neither courtship nor oviposition activities were observed, so the larval food plant and immatures remain unknown.

Discussion

My field observations of P. mimica over the last five years indicate that it is restricted to a narrow transitional habitat on the Pacific drainage between montane cloud forest and the foothills of the western cordillera. This habitat, which I term the "belt of endemism," shows a high degree of endemism and biotic peculiarity for butterflies, plants (Gentry 1982), and birds (Haffer 1967), in contrast to other areas of Colombia. This perhaps is an indication that P. mimica is an endemic Colombian species. The known distribution of P. mimica (Fig. 22) agrees with the "Choco Quaternary Refugia" proposed for neotropical lepidoptera by Brown (1975). Very likely the Choco region was a "forest refugium" during pleistocene glaciations (Brown 1982), explaining the occurrence of a high number of endemic butterflies there and in immediately adjacent areas. The Choco region is located on the Pacific slopes of the western cordillera of Colombia from north of Quibdo to near Lago Calima, across Upper Atrato and San Juan River systems, and its influence is seen as far south as central-western Ecuador (Brown 1975).

Although P. mimica was so named because of the black markings of

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Figs. 10-21. Wing venation of some haeterine species from Colombia. Note the shape variation of the hind wing discal cell and the vein M3 for each genus. 10, Paradulcedo mimica (Rosenberg & Talbot) 6, Alto Calima, Valle. 11, Dulcedo polita (Hewitson) 9, Rio Tatabro, Bajo Anchicaya, Valle. 12, Pseudohaetera hypaesia (Hewitson) 9, Penas Blancas, Farallones, Valle. 13, Haetera macleannania Bates 6, Rio Tatabro, Valle. 14, Haetera macleannania Bates 9, Rio Raposo, Valle. 15, Haetera piera piera (Linnaeus) 9, Leticia, Amazonas. 16, Haetrera piera ecuadora Brown 6, Puerto Umbria, Putumayo. 17, Haetera piera ecuadora Brown 9, Orito, Putumayo. 18, Cithaerias menander (Drury) 6, Rio Sabaletas, Valle. 19, Cithaerias aurorina (Weymer), 6, Puerto Narino, Amazonas. 20, Cithaerias pyritosa ssp. (Clifton ms.) 6, Villavicencio, Meta. 21, Cithaerias pyritosa ssp. 9, same locality.

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Fig. 22. Known distribution of Paradulcedo mimica in Colombia. 1, Risaralda, San Antonio del Chami, 1600-2000 m. This locality is the farthest north known for P. mimica. The habitat where the collections were made is on the upper Mistrato River in montane cloud forest. 2, Risaralda, Santa Cecilia. On the west side (Pacific slope) of the western cordillera in premontane cloud forest. 3, Risaralda, Pueblo Rico, 1584 m, the Paratype locality, on the road to Santa Cecilia. 4, Choco, San Jose del Palmar, 1000 m on the Pacific slope in cloud forest. 5, Valle, Rio Bravo (upper Calima) in premontane and montane cloud forest. 6, Valle, Cerro los Chancos, near Calima Lake, 1600 m, in montane cloud forest. 7, Valle, Rio San Juan, km 56 near Queremal, 1200 m, in montane cloud forest.

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the hindwing which resemble P. hypaesia, there is no evidence of a mimetic association between the two species. The two are parapatric, P. mimica occurring on the west slope and P. hypaesia on the east slope of the western cordillera. The genera Dulcedo, Paradulcedo, and Pseu-dohaetera are monotypic, with the latter two restricted to cloud-forest relicts in the Andean mountains. Lowland rain-forests are the province of the other neotropical Haeterini, Cithaerias, Haetera, and Pierella, with the greatest diversity in the Amazon Basin.

The restriction of haeterine adults to the deep shade of the forest understory (commonly undisturbed rain forests) renders them highly vulnerable to habitat disturbance. Most adults tend to stay within a confined area in the forest at all times. Adults cannot tolerate direct sunlight, and once forests are cleared the species disappear from their natural habitat. In secondary forests, haeterine species are seen rarely or are absent. As a result, P. mimica is seriously threatened as most of its natural habitats have been taken over for agriculture and pastures.

Acknowledgments

I thank J. A. Salazar for originally pointing out to me that this species corresponded to a new genus and for helpful comments and continual encouragement on this paper; J. H. Velez, Museo de Historia Natural de Manizales, Colombia, for providing facilities and access to the collection under his care; and B. A. Drummond, P. J. DeVries, and an anonymous reviewer for useful comments on the manuscript. Thanks also to J. F. Lecrom and E. Schmidt-Mumm for their assistance in checking data and specimens in their collections; P. R. Ackery, The Natural History Museum, London, for providing the reference for the type-species and for his comments on the manuscript; K. S. Brown Jr. (Universidade Estadual de Campinas, Brazil) for making suggestions on the taxonomic status of this distinctive species; my brother Emilio and sister Sandra Constantino for collecting and providing specimens; and German Ramirez for the photographs of the adults.

Literature Cited

Brown, F. M. 1942. Notes on Ecuadorian butterflies. J. New York Entomol. Soc. 50:

304-333. Brown, K. S. 1975. Geographical patterns of evolution in neotropical Lepidoptera.

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phalinae). J. Entomol. 44:201-242. ---------- 1982. Paleoecology and regional patterns of evolution in Neotropical forest

butterflies, pp. 255-308. In Prance, G. T. (ed.), Biological diversification in the tropics.

Columbia Univ. Press, New York. 714 pp. D'Abrera, B. 1989. Butterflies of the Neotropical Region, Part 5, Nymphalidae (Cone.)

and Satyridae. Hill House, Victoria, Australia. 202 pp. D'Almeida, R. F. 1951. Ligeras observacoes sobre o genero Cithaerias Hiibner, 1819,

(Lep. Satyridae). Arquiv. Zool. Estado Sao Paulo 7:493-505. DeVries, P. J. 1987. The butterflies of Costa Rica and their natural history. Princeton

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112-136. In Prance, G. T. (ed.), Biological diversification in the tropics. Columbia

Univ. Press, New York. 714 pp. Haffer J. 1967. Speciation in Colombian forest birds west of the Andes. Am. Mus.

Novit. 2294:1-57.

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Masters, J. H. 1970. Bionomic notes on Haeterini and Biini in Venezuela (Satyridae).

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with the list of the Neotropical transparent winged Satyridae. Tokurana 12:1-10. Rosenberg, W. F. H. & G. Talbot. 1914. New South American butterflies. Trans.

Entomol. Soc. Lond. 1913:671-682. Smart, P. 1976. Encyclopedia of the butterfly world. Hamlyn, London. 275 pp. Takahashi, M. 1981. A list of the butterflies of the Haeterinae and Biinae (Lep.:

Satyridae) collected by two Japanese expeditions in Colombia and Peru, South America. Tyo to Ga. 32:108-116. Weymer, G. 1924. Familie Satyridae, pp. 173-283. In Seitz, A. (ed.), Die Gross-Schmet-

terlinge der Erde. Vol. 5. A. Kernen, Stuttgart.

Received for publication 14 September 1991; revised and accepted 2 February 1992.