Neotrop Entomol DOI 10.1007/s13744-012-0102-4 ECOLOGY, BEHAVIOR AND BIONOMICS

Ithomiini (: ) of Antioquia,

1 2 3 1 CE GIRALDO ,KRWILLMOTT ,RVILA ,SIURIBE

1Grupo de Investigación en Sistemática Molecular, Univ Nacional de Colombia, Medellín, Colombia 2McGuire Center for Lepidoptera and , Florida Museum of Natural History, Univ of Florida, Gainesville, FL, USA 3Institut de Biologia Evolutiva (CSIC-UPF), Barcelona, Spain

Keywords Abstract Biodiversity, , neotropics, Colombia is one of the most biodiverse countries on the planet. However, economic and scientific investment in completing inven- Correspondence tories of its biodiversity has been relatively poor in comparison CE Giraldo, Grupo de Investigación en with other Neotropical countries. Butterflies are the best studied Sistemática Molecular, Univ Nacional de Colombia, Calle 59A No 63 – 20 Bl 16-102, group of invertebrates, with the highest proportion of known to Medellín, Colombia; [email protected] expected . More than 3,200 species of butterflies have been recorded in Colombia, although the study of the still many unex- Edited by André VL Freitas – UNICAMP plored areas will presumably increase this number. This work pro- Received 18 September 2012 and accepted vides a list of butterflies collected in the department of 2 December 2012 Antioquia and estimates the total number of species present, based on revision of entomological collections, records in the literature * Sociedade Entomológica do Brasil 2013 and field work performed between 2003 and 2011. The list includes 99 species and 32 genera, representing 27% of all Ithomiini species. We report 50 species of Ithomiini not formerly listed from Anti- oquia, and found the highest diversity of ithomiine species to be at middle elevations (900–1,800 m). The mean value of the Chao2 estimator for number of species in Antioquia is 115 species, which is close to a predicted total of 109 based on known distributions of other Ithomiini not yet recorded from the department. Nine spe- cies are potentially of particular conservation importance because of their restricted distributions, and we present range maps for each species. We also highlight areas in Antioquia with a lack of biodiversity knowledge to be targeted in future studies. This paper contributes to mapping the distribution of the Lepidoptera of Antioquia department in particular and of Colombia in general.

Introduction in the world and contains two global biodiversity hotspots: the Tropical region, the richest and most diverse Latin America and the Caribbean possess 22% of the region on Earth, and the Tumbes-Chocó-Magdalena hot- world’s forested lands; however, 3 million of the 8.9 million spot, that includes the Chocó-Darién region (Conservation hectares lost per year (33%) throughout the world are International Foundation 2007). According to this, there is found in these regions. In Colombia, it is estimated that an urgent need to conduct biodiversity inventories in many the deforestation rate is 120,000 ha per year, more than regions, which have not been done because social conflicts 320 ha per day, a situation that severely threatens the make it difficult to access some specific areas and also for biodiversity and natural resources of the country (FAO some of the called governmental obstacles to biodiversity 2009). Colombia is one of the most biodiverse countries research (Fernandez 2011). Giraldo et al

Butterflies contain more than 20,000 described species, entomological collections, literature and field work. The making this group the largest invertebrate clade for which information provided here highlights regions of Antioquia description of species is nearing completion (Kristensen et in most need of further research, in addition to increasing al 2007). A total of 3,272 species have been listed for our knowledge of species distributions throughout Colombia (Constantino & Andrade 2007), and additional Colombia. species will probably be added to the list if unexplored areas are surveyed (Gonzalez & Andrade 2008). The department of Antioquia covers an area of 62,776 km2, Material and Methods and it is divided into 125 municipalities grouped in nine ad- ministrative sub-regions. It is located in the central northwest- Data acquisition ern part of Colombia, between 05°25′–08°55′ North and 73° 53′–77°07′ West, between the Atrato and Magdalena rivers, Specimen locality data were obtained by examining major and ranges in elevation from sea level to 3,969 m (Galeano & public collections in Antioquia, through literature review, Bernal 1987). It includes part of the Caribbean coast and of the and by new field collections performed from 2003 to 2011. inter-Andean valleys, Cauca and Magdalena, covering areas in The entomological collections visited in Colombia were the the foothills (Atrato, Urabá, Magdalena, Bajo Cauca), central Museo Entomológico Francisco Luis Gallego (MEFLG- and western mountains of the Andes and the Canyon of the Medellín), Instituto de Ciencias Naturales–Universidad besides several areas of paramo (Arias 2011), Nacional de Colombia (ICN-), Colección Entomológica resulting in a highly diverse mix of climates and biogeograph- de la Universidad de Antioquia (Medellín) and Colección ical regions. Also within its boundaries are found parts of the Entomológica Universidad Católica de Oriente (). In Tumbes-Chocó-Magdalena and the Tropical Andes hotspots. addition, KRW also obtained data from a number of other There have been several papers published with lists of collections during taxonomic work on Ithomiini, including the butterflies from Antioquia (Constantino 1997, Duque 2000, following: American Museum of Natural History, New York, Henao 2006), but just one of them is focused on the USA; Collection André Victor Lucci Freitas, Campinas, ; Neotropical tribe Ithomiini (Muriel 2006). Ithomiini Booth Museum of Natural History, Brighton, UK; BMNH: The (Nymphalidae: Danainae) includes about 370 species of Natural History Museum, London, UK; Carnegie Museum of butterflies and over 1,500 geographical races occurring Natural History, Pittsburgh, USA; Florida Museum of Natural from sea level to 3,000 m, from Mexico to southern History, Gainesville, USA; Fabio Vitale Collection, Lecce, Italy; Brazil, Paraguay and across three Caribbean islands Collection Gabriel Rodríguez, Medellín, Colombia; GSM: (Lamas 2004, Willmott & Freitas 2006). Ithomiini are of Grupo de Investigación en Sistemática Molecular, special interest as entomological and ecological models Universidad Nacional de Colombia sede Medellín, Colombia; because of their diversity, abundance, great variety of Instituto de Investigación de Recursos Biológicos Alexander often precise patterns (Joron & Mallet 1998) and von Humboldt, Claustro San Agustín, – narrow host plant relationships (Drummond & Brown Colombia; Collection Jean-François Le Crom Bogotá– 1987), with host plant use apparently correlating in some Colombia; Museo de Historia Natural, Universidad Nacional cases with mimicry (Willmott & Mallet 2004). Most species Mayor de San Marcos, Lima, ; Collection Sandra Muriel, inhabit shady places in the forest (García et al 2002), Medellín–Colombia; Staatliches Museum für Naturkunde, although several species can be found at the edges of the Stuttgart, Germany; and Staatlische Museum für Tierkunde, forest, disturbed areas and in urban or suburban systems Dresden, Germany. (Brown & Freitas 2003). Ithomiines have been proposed as Localities were georeferenced using municipality web a biological indicator group at local, ecological scales pages and Google Earth®. Literature surveyed included (Brown 1991, Brown & Hutchings 1997, Fagua et al 1999, local publications such as thesis and conference abstracts, García et al 2002, Prado & Freitas 2009) and at broader, and published papers in indexed journals. Finally, informa- biogeographical scales (Beccaloni & Gaston 1995). This tion from recent collections and observations by the tribe is defined by the presence in males of an elongate authors in several localities of Antioquia since 2003 was patch of erectile, hair-like androconial scales at the anterior also included. edge of the dorsal hindwing (Willmott & Freitas 2006), and is phylogenetically related to the Tellervini (containing the Spatial analysis single Australasian Tellervo) and the largely Old World , together forming the subfamily Danainae For each locality, the political subdivision and the geomor- (Wahlberg et al 2009). phological region were identified using the following sys- This work extends the list of Ithomiini butterflies known tem described by Arias (2011). Mid-Magdalena Valley: East from the department of Antioquia, based on review of part of Antioquia department, characterized by alluvial Ithomiini butterflies of Antioquia flood plains and terraces of the and its slope of the central Colombian cordillera between 900 and tributary system below 900 m; Cordillera Central: Eastern 3,200 m in the central area of Antioquia. This region is

Fig 1 Number of Ithomiini species by municipality in Antioquia, Colombia. Giraldo et al characterized by the presence of three extensive high hills, alluvial flood plains and the terrace systems of the plateaus dissected into hills, separated by regional inclined and coast area. formations and producing a mountainous relief. The high Records were also categorized by elevation as tropical plateaus and mountainous areas are broken up by some (0–900 m), subtropical (900–1,800 m), temperate (1,800– deep lineal canyons (Medellín-Porce, Nus and Nare 2,500), cold (2,500–3,400 m) and páramo (>3,400 m). canyons). Also, a finer division of the altitudinal gradient (500 m) was used to compare diversities among altitudinal ranges Cauca Canyon employing the Jaccard similarity coefficient calculated in Past 2.17 (Hammer et al 2001). Formed by the western slope of the “Cordillera Central” Spatial analyses were performed to compare species and the eastern slope of the “Cordillera Occidental”, this composition by municipality, sub-region, geomorphology, region is a narrow and elongated basin (more than 350 km altitude and collection. Maps of species richness of long) with inclined and very inclined slopes. The canyon Ithomiini in Antioquia were obtained using ArcGis® 10.1. becomes deeper and narrower to the north. All georeferenced records were plotted using a political basemap divided into municipalities. We used a 0.2°×0.2° 2 Bajo Cauca grid (25×25 km, 625 km ) to examine patterns of species richness, with a single grid cell having approximately the Frontal foothills of the “Cordillera Central” and “Cordillera average area of the municipalities of Antioquia. Grid cells – Occidental” in the northern part of Antioquia, formed by were also used as sampling units with presence absence hills of Tertiary sedimentary rocks, and containing alluvial data to estimate the Chao 2 richness value. The Chao 2 floodplains and terrace systems of the Cauca River. estimator is a non-parametric method of incidence that does not assume any distribution of data or model and just requires absence/presence data (Moreno 2001). Chao Cordillera Occidental 2 estimator was calculated using Past 2.17 (Hammer et al 2001) Western slope of the “Cordillera Central” between 900 and 3,969 m in the west-central part of Antioquia, and includ- ing three sections: high plateau next to the Cauca Canyon around 2,000 m; the higher area on the “Triángulo Results Montañoso Tropical” including páramo areas with 3,969 m as the highest altitude; and the Murrí-Mandé A total of 1,021 distribution records within Antioquia corridor around 1,800 m next to the Atrato valley. (Online Supplementary Material Appendix 1) were obtained representing all subtribes, 32 genera and 99 species Atrato–Uraba valley (Online Supplementary Material Appendix 2), comprising 27% of Ithomiini species. The mean value of Chao 2 esti- Western and northwestern areas of Antioquia, bordered by mator (95%), calculated for the grid cells sampled, was 115 the to the north and the Antioquia Mid- species. Data were obtained for 54.4% of the municipalities Atrato area to the south. The region is characterized by of Antioquia. The average was 11 species per municipality,

Fig 2 Number of Ithomiini species by subregion in Antioquia, Colombia. Ithomiini butterflies of Antioquia

Fig 3 Number of Ithomiini species by geomorphological area in Antioquia, Colombia.

but 44 of the 68 registered municipalities had less than 10 est similarity between those altitudinal ranges too species, and 17 municipalities had only a single species. The (Table 1). highest number of species was 48 for the municipality of The entomological collection with the highest number of Amalfí (Fig 1). All of the sub-regions were represented, and species from sites in Antioquia was the MEFLG with 68 the average was 41 species per sub-region, but two of the species, increasing to 72 species including the specimens nine sub-regions had less than ten species registered under cryopreservation at the associated GSM collection. (Fig 2). The least diverse sub-region was Bajo Cauca with In contrast, several otherwise important collections in 3 species, and the most diverse was Oriente with 66 Colombia had just one record for Ithomiini butterflies col- species. The analysis by geomorphology showed notably lected in Antioquia, such as the ICN, the largest low diversity for the lowland areas of the northern and Lepidoptera collection in Colombia (Fig 5). At the scale of western parts of Antioquia, Atrato-Urabá Valley and Bajo 25×25-km grid cells, the department was divided into 135 Cauca, with nine and three species, respectively (Fig 3). In cells of which 44 (33%) have at least one species registered contrast, the richest area was Cordillera Central with 93 (Fig 6). species. The average number of species per geomorpho- logic area was 41 species. Classification of species by ele- vation showed the highest diversity for the subtropical Discussion zone with 91 species recorded, and the lowest diversity for the cold zone with just 4 species registered (Fig 4a). We report here 50 species that were not previously listed Also, the finer subdivision showed the highest diversity at for the tribe from Antioquia (Henao 2006). The number of elevation ranges of 1,000–1,499 and 1,500–1,999 m species recorded is notable because it doubles that hither- (Fig 4b). The Jaccard similarity coefficient showed the high- to reported and represents 27% of all Ithomiini species in

Fig 4 Number of Ithomiini species by altitude in Antioquia, Colombia. a Broader altitudinal classification; b Finer altitudinal classification. Giraldo et al

Table 1 Comparison of altitudi- nal diversity of Ithomiini in Range Jaccard similarity coefficient Antioquia, using the Jaccard similarity coefficient calculated 0–499 500–999 1,000–1,499 1,500–1,999 2,000–2,499 2,500–3,000 among altitudinal range of 500 m. 0–499 1 0.267606 0.314607 0.329545 0.238806 0.0454545 500–999 – 1 0.597561 0.541176 0.276316 0.0344828 1000–1499 –– 1 0.698925 0.458824 0.0740741 1500–1999 –– – 1 0.493976 0.0740741 2000–2499 –– – – 1 0.152174 2500–3000 –– – – – 1 approximately 0.4% of its distribution area (approximately 21 km2 (Elias et al 2007). Thus, additional species are 16,601,693 km2). Although 54% of Antioquia municipalities expected to be reported in Antioquia if similar studies are have been “sampled”, 25% of municipalities have just one carried out in other areas such as Urabá and Bajo Cauca. species reported. Even with one half of the municipalities The geomorphological areas analysis showed a similar sampled, the analysis by sub-region shows that there are pattern of uneven sampling. The Cordillera Central showed several zones that are significantly deficient in distribution the highest diversity, perhaps explained by the influence of data, such as Bajo Cauca and Urabá, which represent 30% faunas from the Magdalena valley, the Cauca canyon and of Antioquia. Valle de Aburra is the only sub-region that Bajo Cauca foothills, although it must be noted that in this has records in all of its municipalities. This region contains area are located the municipalities where major collecting 53 recorded species despite being the smallest sub-region, efforts have been made. The lowlands associated with representing just 2.4% of the departmental area. The high foothills and valleys appeared less diverse except for levels of collection activity here are due to the area’s Magdalena, but this pattern once again could be related proximity to Medellín, but even here collections have been to the lack of sampling in these areas, as in Bajo Cauca largely opportunistic rather than systematic. Expanding where just one locality in the municipality of Caucasia has collection efforts in other areas will likely result in higher been sampled. The Cordillera Occidental might be totals; for example, the inventory study at Porce II (Amalfi– expected to have as many species as the Cordillera Gomez Plata municipalities), mainly made by Duque Central, in part because of the influence of the Chocó (2000), lists 51 species. This number is more than a half biogeographic region. of the current list for the department, and these species The altitudinal distribution patterns are in accordance were collected in a very small area (10 km2). This locality with those from studies of ithomiines in other regions, with compares in diversity with the Napo region of eastern the highest diversity at the mid-elevation, probably related , one of the most diverse regions of the world, to overlaps of upper montane and lowland faunas at these where 58 Ithomiini species were recorded in approximately elevations (Elias et al 2009, Lisa de-Silva et al 2011). Mid-

Fig 5 Number of Ithomiini species with Antioquia records by collection or publication. Ithomiini butterflies of Antioquia elevations (subtropical-temperate climate) in Antioquia studied here, but it nevertheless contains more than represent just 25% of its surface, but they contain 97% of 1,300 specimens of ithomiines collected in Antioquia. the species here listed. Museum collections are undoubt- However, there are 12 species that are known from the edly the main source of information about Ithomiini distri- department only from specimens deposited in collections bution. The local collection MEFLG includes the highest outside of Colombia. Collections in other countries are number of species from Antioquia in any collection. located principally in Europe, such as the BMNH in Created in 1937, this is one of the younger museums London, which contains 54 species for Antioquia, and they

Fig 6 Species richness of Ithomiini in Antioquia, Colombia. Grid cells are 0.2°×0.2° (2× 25 km, 625 km2). Giraldo et al constitute a very important source of information because (Cramer), hymenaea (Prittwitz), ochretis they preserve many specimens collected before the in- (Haensch), Greta esula (Hewitson), Hyalenna sulmona crease in deforestation in Antioquia that started with the (Hewitson), exelsa (Felder & Felder), Colombian Industrial Revolution. Species that are repre- schausi (Fox), jucunda (Godman & Salvin), sented only by historical material are perhaps of particular cyrene (Latreille), Ollantaya olerioides (d'Almeida), Oleria conservation interest because of their apparent rarity in quadrata (Haensch), pacifica (Willmott & Lamas) recent years, and could be the subject of future studies in and Velamysta phengites (Fox). However, species that are local population dynamics, regional distribution and con- uncommon in Antioquia are not necessarily endemic to the servation priorities. These species include lenea department or to Colombia. Of far greater significance for

Fig 7 Distribution maps of priority Ithomiini species for study in and around Antioquia. Ithomiini butterflies of Antioquia conservation efforts in Antioquia are those species that are is 99, an additional 10 species are expected to occur in endemic, or almost endemic, to Colombia that also occur in Antioquia based on their known distributions in Colombia, Antioquia: iolaia (Hewitson), nepos including: Callithomia hydra (Felder & Felder; known from (Weymer), Greta n. sp., Ithomia hymettia (Staudinger), to , but not yet recorded in Colombia), Ithomia lagusa (Hewitson), benigna Episcada sylpha (Haensch; known from Magdalena and (Weymer), medellina (Haensch), Pteronymia César departments), confusa (Butler) (known laura (Staudinger) and Oleria thiemei (Oberthür). Because from Panama, and Amazon region), of their high endemism and low frequency in collections, Napeogenes cranto (Felder & Felder; known from Nariño, these species should be priorities for research into their Boyacá, Cundinamarca, Magdalena, Tolima and Valle del distribution, biology and local population dynamics, and we Cauca departments), Oleria n. sp. (known from Nariño and provide distribution maps in Fig 7. Valle del Cauca departments), Pteronymia alissa In contrast, there are several species that may be con- (Hewitson; known from Cundinamarca and Nariño depart- sidered common because they have been collected in more ments), Pteronymia alida (Hewitson; known from Valle del than 25% of the municipalities with specimen records. Cauca and probably the Cordillera Oriental), Pteronymia However, species records by municipality provide a rela- fumida (Schaus; known from Valle del Cauca and tively unreliable measure of distribution because munici- Risaralda departments), Pteronymia obscuratus (Fabricius; palities differ in area. Using an equal-area analysis (25× known from ) and Velamysta 25 km grid) (Fig 6), we identified several “common” species pupilla (Hewitson; known from Quindío department). including: clearista (Doubleday), With these additional species, the estimated number of (Hewitson), dero (Hübner), Dircenna olyras ithomiine species for Antioquia department reaches 109, (Felder & Felder), humboldt (Latreille), Episcada close to the mean value of Chao 2 estimator (115 species; polita (Weymer), Greta andromica (Hewitson), Ithomia see Fig 9). Spatial patterns of species richness in Ithomiini iphianassa (Doubleday), polymnia (Linnaeus), in Antioquia show a similar pattern to those found for the (Hewitson), Mechanitis lysimnia vascular plants in this department, where collection efforts (Fabricius), Napeogenes stella (Hewitson), Oleria fumata are highly correlated with proximity to the capital city (Haensch), Oleria amalda (Hewitson), timna (Medellín) (Idarraga & Callejas 2001). Since areas near to (Hewitson), Pteronymia aletta (Hewitson), Pteronymia lat- the capital are among the most deforested in Antioquia, illa (Hewitson), lavinia (Hewitson), Hypoleria oca- areas of the department with the best preserved forest lea (Doubleday), euclea (Godart) and Hypothyris areas, likely to have the most intact ithomiine faunas lycaste (Fabricius) (Fig 8). The grid cell analysis showed (Brown 1991, Beccaloni & Gaston 1995, García et al 2002; once more the undersampling in much of Antioquia, with Brown & Freitas 2003), have yet to be intensively sampled 67% of grid cells without records. Grid cells were also used (Fig 10). In addition, these little-studied areas are located in as sampled units of presence/absence data to estimate the Tumbes-Chocó-Magdalena hotspot, particularly in the richness. The lower value of Chao 2 estimator (95%) sug- western (next to the Atrato river and Urabá region) and gests a maximum of 104 species for the cells sampled northeastern regions (Bajo Cauca region). Most of the spe- (Fig 9). Even though the number of species here registered cies listed here appear to have been collected in forest relicts

Fig 8 Proportion of all grid cells occupied by common species of Ithomiini in Antioquia, Colombia. Giraldo et al

Fig 9 Species accumulation curve and Chao 2 estimator of Ithomiini richness, using grid cells as units of analysis with specimen record presence/ absence data. within the most deforested region of Antioquia, highlighting Firstly, areas to the north and northeast, including munici- the importance of these relict forests for biodiversity conser- palities of the Bajo Cauca and Nordeste sub-regions, espe- vation and demonstrating that protection and management cially municipalities with well-preserved forest such as Anorí, measures directed to these regions are worthwhile. In addi- , Nechí, Tarazá, Zaragoza and Zegovia, included in the tion, it is likely that surveys of unexplored well-preserved Tumbes-Chocó-Magdalena hotspot and currently a target forested areas will also result in new discoveries. area for mining. Secondly, unstudied areas next to the Finally, several priority areas for future research on but- Atrato river in the municipalities of Vigía del Fuerte and terfly biodiversity in Antioquia department are suggested. Murindó, also part of the Tumbes-Chocó-Magdalena

Fig 10 Comparison of patterns of a Ithomiini richness and b extent of forested areas in 2006 in Antioquia (Modified of Instituto Geográfico Agustín Codazi 2007a, b). Ithomiini butterflies of Antioquia hotspot. Unfortunately, these areas mentioned above have environmental correlates and conservation. J Conserv – reduced accessibility because of social conflict, and biodiver- 6:217 231 Brown KS JR, Hutchings RW (1997) Disturbance, fragmentation and sity studies in such regions are not easily accomplished. the dynamics of diversity in Amazonian forest butterflies. In: It is clear that a considerable amount of sampling is still Laurance WF, Bierregaard RO Jr (eds) Tropical forest remnants. needed to obtain a reliable inventory of Ithomiini species Ecology, management, and conservation of fragmented communi- – for Antioquia and, eventually, for Colombia. The major ties. The University of Chicago Press, Chicago, pp 91 110, 632 Conservation International Foundation (2007) Biodiversity Hotspots. issues preventing an increase in our knowledge of Web site: http://www.biodiversityhotspots.org/xp/hotspots/ Colombian biodiversity are not the lack of training or moti- Pages/default.aspx. Accessed 23 Jan 2012 vated researchers, but the social conflict in many areas that Constantino LM (1997) Conocimiento de los ciclos de vida y plantas limits accessibility, the lack of financial support and the hospederas de lepidópteros diurnos de Colombia como estrategia para el manejo, uso y conservación de poblaciones silvestres, pp. recent numerous governmental barriers to studying biodi- 259–278. In: Seminario Aconteceres Entomológicos, para compren- versity in Colombia. Antioquia is one of the best studied der los insectos: estudiarlos. Universidad Nacional de Colombia, departments for insect diversity in Colombia, together with Medellín, p 282 Cundinamarca and Valle del Cauca departments, so an even Constantino LM, Andrade G (2007) Mariposas diurnas y nocturnas, p.107-143. In Amat G, Andrade G, Amat E (eds) Libro Rojo de los worse panorama is expected for the rest of the country. It is Invertebrados Terrestres de Colombia. Bogotá, Instituto de Ciencias becoming clear that much of the biodiversity of the country Naturales – Universidad Nacional de Colombia, Conservación is still unknown while deforestation and mining exploitation Internacional Colombia, Instituto Alexander von Humbolt, Ministerio continue and habitats are reduced and fragmented. It is thus de Ambiente, Vivienda y Desarrollo Territorial.Colombia, 216p Drummond BA III Jr, Brown KS (1987) Ithomiinae (Lepid.: urgent to increase efforts to document and protect biodiver- Nymphalidae): summary of know larval foodplants. Ann Mo Bot sity in Colombia and, to this end, government strategies Gard 74:341–358 should be directed to support scientific investigation. Duque P (2000) Diversidad de Mariposas diurnas en diferentes frag- mentos de bosque en la zona de influencia del proyecto hidroeléctrico Porce II. Tesis de Maestría Universidad Nacional de Acknowledgments We thank Efraín Henao, Gonzalo Andrade, Jean Colombia sede Medellín Francois LeCrom, Marta Wolff, Oscar Ortega and John Quiroz for letting Elias M, Hill RI, Willmott KR, Dasmahapatra KK, Brower AVZ, Mallet J, us review local collection. We also thank Luis Alberto Arias for his Jiggins CD (2007) Limited performance of DNA barcoding in a support in the geomorphological description of Antioquia and Alba Lucía diverse community of tropical butterflies. Proc R Soc London Ser Marín for her help in maps design. Support for this research was B 274:2881–2889 provided by the Sciences School of the Universidad Nacional de Colom- Elias M, Joron M, Willmott KR, Silva-Brandão KL, Kaiser V, Arias CF, bia sede Medellín and the Fundación BBVA (MARIPOSA Proyect: BIO- Gomez LM, Uribe S, Brower AV, Jiggins CD (2009) Out of the CON08_021). KRW acknowledges support from the Leverhulme Trust, Andes: patterns of diversification in clearwing butterflies. Mol and thanks to the many museum curators and individuals who facilitat- Ecol 18:1716–1729 ed access and research in ithomiine collections. Gerardo Lamas, Jim Fagua G, Aarillo AR, Andrade MG (1999) Las mariposas (LEPIDOPTERA: Mallet and Fraser Simpson provided particularly important data and RHOPALOCERA) como indicadores del grado de intervención en la support for databasing efforts at the BMNH. cuenca del Río Pato (Caquetá, Colombia), p. 285–315, In Amat G, Andrade-C G, Fernandez F (eds) "Insectos de Colombia. Vol. 2". 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