Phylogeny, taxonomy, and sympatry of Timaeta (Lycaenidae: Theclinae: Eumaeini): an Andean montane forest endemic Robert K. Robbins & Robert C. Busby Taxonomy of the Andean lycaenid hairstreak genus Timaeta Johnson et al. and its fifteen sexually dimorphic species is detailed for the first time. Species accounts summarize nomenclature, distribution, habitat, behavior, and distinguishing traits, as well as noting why each species is considered distinct under a biological or phylogenetic species concept and how males and females were associated. An identification key for males and a nomenclatural checklist are included. Nomenclatural actions include the description of Timaeta christina Robbins & Busby sp. n., Timaeta matthewi Busby & Robbins sp. n., Timaeta walteri Busby & Robbins sp. n., Timaeta roberti Busby & Robbins sp. n., Timaeta cospata Robbins & Busby sp. n., Timaeta pilosa Robbins & Busby sp. n., Timaeta gabriela Busby & Robbins sp. n., and Timaeta romero Robbins & Busby sp. n.. Timaeta neblina (Salazar & K. Johnson, 1997) is a new synonym of Timaeta eronos (H.H. Druce, 1890). A phylogenetic analysis is based on 17 coded morphological characters and yields two most parsimonious 30-step trees. Successive weighting does not change tree topology. The node chosen to delimit Timaeta is the one that had the highest jackknife support and was most stable with past usage. Generic names Trochusinus K. Johnson, Salazar & Vélez, 1997 and Jagiello Bálint & Wojtusiak, 2000 are confirmed as synonyms of Timaeta K. Johnson, Kruse & Kroenlein, 1997. Recent revisions of primarily Andean butterfly genera have had a significant proportion of previously unnamed species, and Timaeta fits this pattern with eight of 15 species previously undescribed. Most sister lineages within Timaeta are sympatric, and more than half the species occur between 1500m and 2000m in eastern Ecuador. This pattern of sympatry is similar to that of primarily lowland eumaeine genera, but contrasts with repeated reports of elevational allopatry/parapatry in Andean butterflies and birds. R.K. Robbins, Department of Entomology, PO Box 37012, NHB Stop 105, Smithsonian Institution, Washington, DC 20013-7012 USA. [email protected] R.C. Busby, 7 Countryside Way, Andover, MA 01810-6041 USA. [email protected] Introduction (Robbins 2004b). And half of the 12 genera of the The primarily Neotropical Eumaeini (Lycaenidae: Micandra Section are endemic to Andean habitats Theclinae) have diversified extensively in montane between 700 and 4,500m elevation. This paper is a habitats, especially in contrast to the primarily Neo- revision of Timaeta, one of the Andean endemics, tropical and similarly diverse Riodinidae (Robbins and is the first phylogenetic analysis of a primarily 2004a). Among the Eumaeini, about 95% of the montane lycaenid genus in the Eumaeini. 150 species in the Micandra Section are montane The Andean lepidopteran fauna is remarkably rich, Tijdschrift voor Entomologie 151: 205–233, Figs. 1– 75, Tables 1-2. [ISSN 0040–7496]. http://www.nev.nl/tve © 2008 Nederlandse Entomologische Vereniging. Published 1 December 2008. Downloaded from Brill.com09/27/2021 03:16:22PM via free access 206 Tijdschrift voor Entomologie, volume 151, 2008 but because it has been chronically undersampled, analyses of DNA sequences from 163 Eumaeini, many previously undescribed butterfly and moth including two Timaeta and one Temecla species, species are only now being discovered (Willmott et Temecla + Timaeta is monophyletic (Quental et al. al. 2001, Brehm et al. 2005, Peña & Lamas 2005). in prep.). The Andean butterfly fauna has been repeatedly The generic nomenclature of Timaeta and its pro- noted to consist of elevationally allopatric/parapatric posed synonyms, Trochusinus K. Johnson, Salazar species (Adams 1985, Pyrcz & Wojtusiak 2002, Hall & Vélez and Jagiello Bálint & Wojtusiak (Robbins 2005, Bollino & Costa 2007), and a similar pattern 2004b), has been unstable because diagnostic char- has also been reported in Andean birds (Remsen acters in the original descriptions were inaccurate. & Cardiff 1990, Bates & Zink 1994). Alternately, For example, “caecum comprising one fourth or less most species in another Andean butterfly genus were of aedeagus” was proposed as a diagnostic character sympatric with their sister lineages (Willmott et al. of male Timaeta (Johnson et al. 1997a:24), but no 2001), as also occurs in primarily lowland Neotropi- eumaeine species has been reported to have a caecum cal lycaenid genera (Robbins 1991, 2005, Robbins length greater than one fourth of the aedeagus. Lack & Duarte 2005, Robbins & Busby In Press). The of androconia was noted as a diagnostic character number of phylogenetic analyses of Andean lepidop- of male Trochusinus (Johnson et al. 1997b:4), but teran genera is yet too few to determine the preva- two species that they placed in Trochusinus (eronos lence of elevational parapatry/allopatry among sister H.H. Druce and balzabamba Goodson) have con- lineages and is unexplored in the Eumaeini. spicuous androconia along the posterior edge of dor- The goals of this paper are taxonomic and biogeo- sal hindwing vein Cu2 (Draudt 1919-1920, D’Abrera graphic. The taxonomic goal is to present evidence 1995, Figs. 19, 29, 65). Jagiello (J. molinopampa for the monophyly of Timaeta and for a classification Bálint & Wojtusiak was the only included species) of its 15 sexually dimorphic species that is slightly was described “… also on the grounds of differ- modified from that in Robbins (2004b). Specifically, ences in genital morphology” (Bálint & Wojtusiak we characterize Timaeta and its included species, 2000:183), but these differences were not detailed, describe eight new species, explain why we consider and the genitalia are very similar to those of each to be distinct under a biological or phylogenetic T. timaeus (Figs. 36, 38, 51-52). species concept, note the evidence that we used to The species level taxonomy of Timaeta has also been associate males and females, and infer phylogenetic unresolved. Wing pattern sexual dimorphism makes relations among Timaeta species. The biogeographic associating males and females difficult. For example, goal is to summarize information on diversifica- Draudt (1919-1920) did not correctly associate the tion and sympatry among Timaeta species using sexes of any species known at the time, D’Abrera the inferred phylogeny and distributional data. We (1995) correctly associated the sexes of one species, specifically ask whether a pattern of sympatry or and Bálint & Wojtusiak (2000) did not correctly elevational allopatry/parapatry predominate among identify the female of T. molinopampa. Most species sister lineages in Timaeta. are rare in collections, making it difficult to assess variation. Finally, the inaccurate original generic Taxonomic history characterizations of Timaeta and its synonyms have Timaeta belongs to the Eumaeini because it has ten caused confusion, with some Timaeta species being forewing veins, “greyhound shaped” male genitalia transferred from or to genera that do not belong lacking a juxta, hairy eyes, and a male foretarsus to the Micandra Section (Robbins 2004b, detailed that is fused, stubby tipped, and used for walking below). (Eliot 1973). Timaeta was placed provisionally in the Extensive field work in Ecuadorian and Peruvian Micandra Section of the Eumaeini (Robbins 2004b) montane forest in the past decade has more than because it has female genitalia with fan-shaped signa quadrupled the number of Timaeta specimens avail- (they occur also in the Hypostrymon and Lampros- able for study, provided new data on distribution and pilus Sections), blue scales on the ventral forewing of variation, and allowed many male and female pheno- males (they occur also in the Eumaeus, Brangas, and types to be associated for the first time. This new Atlides Sections), and several male forewing vena- information makes a revision of Timaeta feasible. tion characters (Schatz & Röber 1885-1892, Clench 1971) that are not yet well-documented. Regard- less of whether this placement is correct, Temecla Materials and methods Robbins and Phothecla Robbins appear to be the The generic and species level taxonomy is based on closest relatives of Timaeta on the basis of characters an analysis of morphological variation among 171 listed in Robbins and Duarte (2004). In preliminary male and 34 female pinned specimens of Timaeta Downloaded from Brill.com09/27/2021 03:16:22PM via free access Robbins & Busby: Taxonomy of Timaeta 207 from various museum and private collections, as MIZA Museo del Instituto de Zoología Agrícola, noted below. Each species account includes notes on Maracay, Venezuela nomenclature and history, morphological variation, MUSM Museo de Historia Natural, Universidad how the sexes are associated (females of three species Nacional Mayor de San Marcos, Lima, are yet unknown or unrecognized), geographical dis- Perú tribution, elevation, and male “territorial” behavior. RCB Robert C. Busby Collection, Andover, The evidence supporting the hypothesis that each MA, USA available name represents a distinct species under a SMF Forschungsinstitut und Naturmuseum biological and/or phylogenetic species concept is dis- Senckenberg Frankfurt, Frankfurt-am- cussed. Distribution of each species is mapped. In a Main, Germany few cases, the distributions of “unrelated” Timaeta SMNS Staatliches Museum für Naturkunde, species are combined on a single map for the purpose Stuttgart, Germany minimizing the number of maps needed. Months UNCC