Genus Vol. 15 (4): 455-622 Wroc³aw, 26 XII 2004 Pronophiline butterflies of the highlands of Chachapoyas in northern Peru: faunal survey, diversity and distribution patterns (Lepidoptera, Nymphalidae, Satyrinae) TOMASZ W. PYRCZ Zoological Museum of the Jagiellonian University, Ingardena 6, 30-060 Kraków, Poland e-mail: [email protected]; [email protected] ABSTRACT. Butterflies of the tribe Pronophilini (Nymphalidae, Satyrinae) are surveyed in the highlands of Chachapoyas covering the northern part of the Andean Eastern and Central Cordilleras in Peru. 112 species are reported and discussed including their geographic distributions, altitudinal ranges and identification keys. 55 new taxa (35 species and 20 subspecies) are described: Apexacuta improvisa n. sp., A. superior n. sp., Corades tripunctata necrufa n. ssp., Corderopedaliodes corderoi exornata n. ssp., Daedalma. fraudata n. sp., D. vertex n. sp., D. boliviana peruviana n. ssp., Eretris apuleina n. sp., E. mendoza n. sp., E. truncatina n. sp., E. porphyria transmaraniona n. ssp., Junea dorinda quasinegra n. ssp., Lymanopoda araneola n. sp., L. dyari n. sp., L. inde n. sp., L. ingasayana n. sp., L. magna n. sp., L. acraeida chavezi n. ssp., L. albocincta intermedia n. ssp., Lasiophila cirta atropurpurea n. ssp., Manerebia benigni n. sp., M. diffusa n. sp., M. haywardi n. sp., M. trirufa n. sp., M. benigni tessmanni n. ssp., M. ignilineata jalca n. ssp., Mygona poeania magalyae n. ssp., Oxeoschistus iphigenia n. sp., Panyapedaliodes stellata n. sp., P. phila certa n. ssp., Pedaliodes albicilia n. sp., P. aureola n. sp., P. boettgeri n. sp., P. demathani n. sp., P. erschoffi n. sp., P. jelskii n. sp., P. maruda n. sp., P. simplissima n. sp., P. sophismata n. sp., P. stuebeli n. sp., P. sztolcmani n. sp., P. uaniuna n. sp., P. woytkowskii n. sp., P. cledonia molesta n. ssp., P. petri maasseni n. ssp., P. phrasicla rufa n. ssp., P. sztolcmani gilvaecosta n. ssp., Proboscis pomarancia n. sp., Pronophila bernardi n. sp., P. tremocrata n. sp., P. epidipnis perplexa n. ssp., Pseudomaniola mirabilis extrema n. ssp., Punapedaliodes flavopunctata minima n. ssp., Steremnia lucillae n. sp., S. agraulis agraulina n. ssp. Several new combinations, new status and synonymies are established. Species richness in the surveyed area is evaluated along potential altitudinal transects and compared to other north Andean localities. Pronophilines are sampled along an elevational transect in Molinopampa reporting 45 species belonging to 20 genera. Data on altitudinal distributions of individual species and changes in the community structure are discussed and compared with similar studies in Colombia and Venezuela. Examples of parapatric distributions along elevational gradients are pointed out. Geographic distribution patterns and faunal units within the surveyed area are identified and their affinities are discussed. 456 TOMASZ PYRCZ Key words: entomology, taxonomy, zoogeography, Lepidoptera, Satyrinae, Pronophilini, new taxa, species richness taxonomy, Andes, Peru, altitudinal distribution. INTRODUCTION THE TRIBE PRONOPHILINI MILLER (1968), in the first modern taxonomic revision of the Satyridae, characterised the tribe Pronophilini REUTER (1896) within the sub-family Satyrinae. HARVEY (1991) downranked this and several others of MILLER’s tribes to the rank of sub-tribes within the tribe Satyrini. VILORIA (Ph.D.) concurred with HARVEY but removed from the sub-tribe Pronophilina the genera Manerebia STAUDINGER, Idioneurula STRAND, Diaphanos ADAMS & BERNARD and Tamania PYRCZ and transferred them to the predominantly Holarctic Erebiina. He also suggested that all the Austral genera found in Peru, Chile and Argentina (i.e. Argyrphorus BLANCHARD, Eteona DOUBLEDAY, Neomaenas WALLENGREN etc.) considered by MILLER as Pronophilini, belong to the Australian sub-tribe Hypocystina. MILLER’s (1968) paper remains the only comprehensive taxonomic revision of the Satyrinae worldwide, despite some obvious deficiencies, as, for example, an incorrect definition of the type of venation of the Pronophilini (PYRCZ Ph.D.; VILORIA Ph.D.) and not taking into account such an important taxonomic character as male genitalia. HARVEY’s (1991) revisional classification of the Nymphalidae (sensu lato) was heavily based on larval morphology data, which was then almost completely unknown for the pronophilines and, therefore, could not be used. The conclusions of VILORIA (Ph.D.) regarding the genus Manerebia are based on few morphological characteristics, most of them quantitative, except for the lack of setose eyes in Manerebia, and are not taken into account herein. It is beyond the scope on the present paper to discuss in detail the tribal level systematics of the Satyrinae. Undoubtedly, the ultimate status and the generic composition of the Pronophilini require a comprehensive revision, which can be carried out in the future when more comparative data, especially on the morphology of larval stages become available. For the purposes of this paper, the Pronophilini are treated as a cohesive group identified from other Neotropical Satyrinae based primarily on ecological features. They are exclusively tropical montane, whereas other tribes of Neotropical Satyrinae, the Haeterini and the Euptychiini, are mostly lowland or premontane (except for the euptychiine genus Forsterinaria HAYWARD). Approximately 95% of all recognised species of Pronophilini occur in the tropical Andes from Ven- ezuela to northern Argentina (LAMAS, VILORIA & PYRCZ 2004). They are by far the best-represented group of butterflies in terms of species richness and abundance in cloud forest habitats (ADAMS 1985, 1986; PYRCZ & WOJTUSIAK 1999, 2002). The systematics and the zoogeography of North Andean pronophilines (Ecuador, Colombia and Venezuela) are relatively well known thanks to the contributions of PRONOPHILINE BUTTERFLIES 457 BROWN (1941, 1943, 1944), ADAMS & BERNARD (1977, 1979, 1981), ADAMS (1985, 1986), VILORIA (1994), PYRCZ (1995, 1999, 2000), PYRCZ & VILORIA (1999), PYRCZ & WOJTUSIAK (1999, 2002), PYRCZ et al. (1999), VILORIA & PYRCZ (2002) and VILORIA et al. (2003). Bolivian and Argentine faunas were monographed by HAYWARD (1958, 1963) and FORSTER (1964). Their papers are however far from being exhaustive and have identification errors. There are no published surveys of the pronophiline fauna of Peru at all. The only recent references on this tribe are descriptive papers dealing with a few central (PYRCZ 2003) and southern Peruvian species (LAMAS 1997, 1999) and incomplete local surveys of Lepidoptera in general (LAMAS et al. 1999, 2003). The most outstanding characteristic of the pronophilines is their altitudinal distribution pattern. Most species are distributed in well-defined and sometimes very narrow bands of altitude (ADAMS 1985, 1986; RAGUSO & GLOSTER 1993; PYRCZ & WOJTUSIAK 1999, 2002). This aspect of their ecology will be discussed in the second part of this paper. Another notable feature of the pronophilines is the high percentage of endemic taxa in all surveyed ranges. Endemism ratio is the highest in the peripheral ranges of the northern Andes, especially in the Sierra Nevada de Santa Marta, Cordillera de Mérida and Sierra de Perijá (ADAMS 1985). Pronophiline butterflies, similarly to other satyrines, are mostly sedentary (ADAMS 1986; DEVRIES 1987) and are often stenobionts associated with perhumid habitats within the cloud forest environment (ADAMS 1986; PYRCZ & WOJTUSIAK 2002). Most species show restricted vagility and even though this aspect of their behaviour has not been studied rigorously, field observations indicate that adults keep close to their potential host plants or roosting places and move little verti- cally or horizontally. Some species are territorial and hilltopping behaviour is characteristic for Junea, Steremnia and some species of Corades and Daedalma. Even though the biology of the Pronophilini remains largely unexplored, it appears that their larvae are oligophagous on montane bamboo, chiefly of the genus Chusquea (Poaceae) (SCHULTZE 1929; ADAMS & BERNARD 1981; DEVRIES 1987, HEREDIA & VILORIA 2004) and Swallenchloa (PYRCZ, WILLMOTT & HALL 1999). Several species of Pronophilini seemingly use also other Poaceae, such as paramo Stipa and Calamagrostis grasses (BROWN 1941; VILORIA & PYRCZ 1999), woody cane (MILLER 1986) and Guadua bamboo (PYRCZ 2000). PELZ (1997) reared Pedaliodes parepa (HEWITSON) in Germany on a substitute food plant, a grass Poa annua (Poaceae). GREENEY & PYRCZ (in prep.) reared on Chusquea various Ecuadorian pronophilines including Daedalma, Corades, Pronophila, Eretris and Pedaliodes. Montane bamboos are more abundant in clearings and ecotones and are typical plants for the early stages of natural succession. Human activity may, in some circumstances, favour Chusquea and therefore acts in favour of maintaining Pronophilini populations or even locally increasing their densities (ANDRADE 1994; PYRCZ & WOJTUSIAK 1999). A reliable way of obtaining sufficient material for statistical analysis is a crucial factor in ecological and zoogeographical studies (BROWN 1982). Fortu- nately, adults of all species of Pronophilini are readily attracted to decomposing 458 TOMASZ PYRCZ organic matter, particularly to faeces, carrion and rotten fruits. Therefore, the use of baits provides a very good method of sampling (PYRCZ & WOJTUSIAK 1999, 2002). THE HIGHLANDS OF CHACHAPOYAS The highlands of Chachapoyas in northern Peru do not form a clear-cut geographic entity; therefore, the area of study is