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1 SYSTEMATICS AND BIOGEOGRAPHY OF HIGH ALTITUDE TROPICAL ANDEAN SATYRINES (LEPIDOPTERA, NYMPHALINAE: SATYRINAE) By PABLO SEBASTIAN PADRON MARTINEZ A THESIS PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE UNIVERSITY OF FLORIDA 2010
2 2010 Pablo Sebastin Padrn Mart nez
3 To my parents for their love and endless support
4 ACKNOWLEDGMENTS This study could not have been accomplished without expert guidance and support from my graduate committee. I thank Dr. Keith Willmott Dr. Marc Branham and Dr. Andrei Sourakov for their help. I want express my deep gratitude to Dr Keith W illmott for his help during my m aster s program, he has been an excellent advisor who has been able to help and advise me in the best way, always with patience and good humor. I would like to thank him for having awakened in me the interest in the study of tropical butterflies. Finally, I would like to thank him for his help with the funding for the molecular work, from a National Science Foundation grant, and for allowing me to work with the specimens that he has collect ed over several years in Ecuador. I would like to thank the Fulbright Commission and SENACYT from Ecuador for the scholarship that I received from them t hat allowed me to study in the University of Florida. I also thank Dr. Thomas E m mel, the director of The McGuire Center for Lepidoptera and Biodiversity, for allowing me to work in this institution and for providing a McGuire Research Assistantship. I thank my family, especially my parents Hernan and Mariana, for financial aid during my fieldwork in Ecuador, and primarily for giving me unconditional support during this work and throughout my life. Thanks also to my brother Santiago, who traveled with me during fieldwork, for always being willing to help and for his excellent company during those cold days and rainy days on the pramos while we collected butterflies. Finally, I would also like to express my sincere gratitude to the staff of the McGuire Center for Lepidoptera and Biodiversity, Florida Mus eum of Natural History, for the excellent work environm ent and for their willingness to help me.
5 TABLE OF CONTENTS page ACKNOWLEDGMENTS .................................................................................................. 4 LIST OF TABLES ............................................................................................................ 7 LIST OF FIGURES .......................................................................................................... 8 ABSTRACT ................................................................................................................... 10 CHAPTER 1 INTRODUCTION .................................................................................................... 12 The Subfamily Satyrinae ......................................................................................... 12 The Subt ribe Pronophilina ...................................................................................... 13 Scope of the Project ............................................................................................... 16 2 MOLECULAR PHYLOGENY OF ALTOPEDALIODES AND NEOPEDALIODES ... 17 Introduction ............................................................................................................. 17 Material and Methods ............................................................................................. 18 Specimen Acquisition, Extraction and Amplification of DNA ............................. 18 DNA Sequencing and Alignment ...................................................................... 19 Phyl ogenetic Analysis ...................................................................................... 20 Maximum Parsimony (MP) ............................................................................... 20 Bayesian Inference (BI) .................................................................................... 20 Results .................................................................................................................... 21 3 REVISION OF THE GENUS ALTOPEDALIODES .................................................. 27 Introduction to the Genus ........................................................................................ 27 Materials and Methods ............................................................................................ 27 History of classification ........................................................................................... 30 Diagno sis ................................................................................................................ 31 Generic Description ................................................................................................ 32 Biology .................................................................................................................... 33 Natural History .................................................................................................. 33 Diversity and Distribution of the Genus ............................................................ 33 Altopedaliodes species accounts ............................................................................ 34 4 DISTRIBUTION OF ALTOPEDALIODES AND NEOPEDALIODES SPECIES AND THEIR RESPONSE TO CLIMATE CHANGE IN ECUADOR ......................... 69 Introduction ............................................................................................................. 69 Methods .................................................................................................................. 70
6 Specimen Data ................................................................................................. 71 Climate Model and Calculation of Distributions ................................................ 72 Results .................................................................................................................... 74 Potential Current Distribution for Altopedaliodes Species ................................ 74 Potential Current Distribution for Neopedaliodes Species ................................ 74 Potential Distribution for Altopedaliodes Species Under an Increase in Temperature .................................................................................................. 75 Potential Distribution for Neopedaliodes Species Under an Increase of Temperature .................................................................................................. 75 Change in Range Size with Climate Change.................................................... 76 5 CONCLUSIONS ..................................................................................................... 84 Molecular Phylogeny of Altopedaliodes and Neopedaliodes .................................. 84 Distribution of Altopedaliodes and Neopedaliodes Species and their Response to Climate Change in Ecuador ............................................................................. 86 LIST OF REFERENCES ............................................................................................... 90 BIOGRAPHICAL SKETCH ............................................................................................ 98
7 LIST OF TABLES Table page 2 1 Genbank accession numbers for the gene sequences used in this study. ......... 26 3 1 Distribution of the Altopedaliodes species. ......................................................... 68
8 LIST OF FIGURES Figure page 2 1 Phylogenetic trees for 24 Pronophilina taxa based on the combined data set of three genes produced by different methods .................................................. 23 2 2 Phylogenetic tree of Altopedaliodes based on the combined data set of three genes produced by different methods .............................................................. 24 2 3 Phylogenetic tree of Neopedaliodes based on the combined data set of three genes produced by different methods .............................................................. 25 3 1 Altopedaliodes species ..................................................................................... 56 3 2 Collection sites in Ecuador. ................................................................................ 57 3 3 Altopedaliodes habitat, Parque Nacional Cajas. ................................................. 57 3 4 A. tena morphology drawings. ............................................................................ 58 3 5 Androconial patches. .......................................................................................... 59 3 6 Male genitalia of other genera of the subtribe Pronophilina ................................ 60 3 7 Female genitalia of Altopedaliodes .................................................................... 61 3 8 Female genitalia of other genera of the subtribe Pronophilina ........................... 62 3 9 A. tena morphology drawings. ............................................................................ 63 3 10 Male genitalia, lateral view, aedeagus removed and illustrated in lateral view. .. 64 3 11 Male genitalia, lateral view, aedeagus removed and illustrated in lateral view. .. 65 3 12 Male genitalia, lateral view, aedeagus removed and illustrated in lateral view. .. 66 3 13 A. nucea first instar larva. ................................................................................... 67 4 1 Maps showing current potential distribution and future potential distribution of Altopedaliodes species ...................................................................................... 77 4 2 Maps showing current potential distribution and future potential distribution of Altopedaliodes species.. ..................................................................................... 78 4 3 Maps showing current potential distribution and future potential distribution of Altopedaliodes species.. ..................................................................................... 79
9 4 4 Maps showing current potential distr ibution and future potential distribution of Neopedaliodes species.. .................................................................................... 80 4 5 Maps showing current potential distribution and future potential distribution of Neopedaliodes species ..................................................................................... 81 4 6 Maps showing current potential distribution and future potential distribution of Neopedaliodes species ..................................................................................... 82 4 7 Maps showing current potential distribut ion and future potential distribution of Neopedaliodes species ...................................................................................... 83
10 Abstract of Thesis Presented to the Graduate School of the University of Florida in Partial Fulfillment of the Requirements for the Degree of Master of Science SYSTEMATICS AND BIOG EOGRAPHY OF HIGH ALT ITUDE TROPICAL ANDEA N SATYRINES (LEPIDOPTERA, NYMPHALIDAE: SAT YRINAE) By Pablo Sebastin Padrn Mart nez August 2010 Chair: Keith Willmott Major: Entomology and Nematology The first phylogenetic hypotheses for the high Andean satyrine butterfly genera Altopedaliodes and Neopedaliodes are proposed using mitochondrial (COI and COII) and nuclear (EF 1 ) genes. These data were analyzed using Max imum Parsimony and Bayesian Inference, and the monophyly of both genera was supported and was broadly independent of analytical method or dataset used. Altopedaliodes n. sp. and A. nucea appear as sister to other taxa of A. tena, and these results, combine d with new field data that suggest sympatry between A. nucea, A. n. sp., and A. tena in Ecuador, assisted in revising the species taxonomy. A revision of Altopedaliodes is presented, all the currently recognized Altopedaliodes taxa are revised, a new speci es (A. n. sp.) and a new subspecies (A. reissi n. ssp.) are recognized, and a new status is proposed (A. nucea). The current potential distribution area was calculated for the genera Altopedaliodes (16,163.42 km2) and Neopedaliodes (46,753.88 km2) in Ecuador and the effect of future climate change was evaluated. This study showed that the likely impact of future climate change on the range sizes of these two genera of highaltitude Ecuadorian satyrines was predominantly negative. Reductions of 8.9% and 9.5% in
11 generic range size, and an average reduction of 44% and 13.6% for species range size, were predicted for Altopedaliodes and Neopedaliodes, respectively. There was also a difference between some areas in Ecuador, with southern areas of the country pres enting a greater reduction of habitat for these two genera than northern areas. These results have implications for future plans to conserve these species.
12 CHAPTER 1 INTRODUCTION The Andes is one of the most diverse regions on the planet (Myers et al., 2000). This continuous chain of mountains along the western coast of South America contains innumerable habitats, which support an amazing diversity of flora and fauna. The tropical Andes contain thousands of endemic species (plants, Brako and Zarucchi, 1993; Jrgensen and Len, 1999; birds, Ridgely and Greenfield, 2001; Hilty and Brown, 1986; Schulenberg et al., 2007; amphibians, Duellman, 2004; 2005, 2007; mammals, Albuja and Patterson, 1996; insects, Hogues, 1993), including butterflies, a group of insects that is also highly diverse in the Andes (Lamas, 2004). The Subfamily Satyrinae One group of butterflies, the subfamily Satyrinae, stands out from all others as one of the most diverse butterfly subfamilies on the pl anet. Satyrinae is a subfamily of the family Nymphalidae and contains nearly half of the known diversity of brushfooted butterflies (The Lepidoptera Taxome Project, 2007). Satyrines are generally weak fliers and are particularly characteristic of forest understorey or open, grassy areas. The satyrines can be recognized by a combination of structural features. The forelegs of both sexes are poorly developed and not used for walking, the veins of the forewing usually are swollen at the base, the cells of bot h wings are closed, and very often males have patches of androconial scales on either the forewing or the hindwing, or on both (Miller, 1968; Howe 1975; DeVries, 1987). The early stages of the Satyrinae feed on monocotyledons (Howe, 1975), mainly in the fa milies Poaceae, Marantaceae, Arecaceae and Cyperaceae (Beccaloni et al., 2008), in addition to some bryophytes (DeVries, 1987). These diverse and widely
13 distributed hosts have allowed this group of butterflies to inhabit a large number of ecosystems, and i t may be one of the causes for the great diversity of these butterflies. The subfamily is also characterized by the typically dull coloration of the adults, which often have eyespots, or ocelli, on their wings; these serve as visual signals to attract mate s or function in predator avoidance (Oliver et al., 2009). The subfamily includes a number of tribes, some of which have formerly been treated as subfamilies, or even families (e.g., Miller, 1968; DAbrera, 1988; Lamas, 2004), including the Elymniini, Zet herini, Amathusiini, Brassolini, Morphini, Melanitini, Dirini, Haeterini and Satyrini (Wahlberg et al., 2009). The higher taxonomy has recently been the subject of much research, since the pioneering morphological studies of Miller (1968). Pea et al. (2006) proposed the first phylogeny for this group based on DNA sequence data. They worked with 93 taxa and 16 outgroups in order to test the monophyly of Satyrinae, and the proposed phylogeny was based on the sequences from three genes, one mitochondrial gene (COI) and two nuclear genes (EF wingless). The results of his research indicated that Satyrinae as then conceived was a polyphyletic subfamily, requiring the inclusion of several former subfamilies to maintain monophyly. More recently, Wahlberg et al. (2009), based on the phylogenetic relationships of 400 genera of Nymphalidae inferred using 10 genes and 235 morphological characters, found that the subfamily Satyrinae as conceived above was strongly supported as monophyletic. The Subtribe Pronophili na Pronophilina is a conspicuous subtribe of the Satyrini, which occurs mainly at middle to high elevations in virtually all montane areas of the Neotropical region. The subtribe is a plausibly monophyletic group of 66 genera, represented by ca. 690 species
14 (Lamas and Viloria, 2004a, 2004b; Lamas et al., 2004; Pea et al., 2006). The Pronophilina are treated as a monophyletic group with respect to other Neotropical Satyrinae based primarily on ecological features and genetic data (Pea et al., 2006). Adult size ranges from small to relatively large. The wing pattern is generally dominated by tones of brown and dark grey, but the group is also rather rich in colorful markings varying from orange to iridescent blue. Pronophilina are almost exclusively montane, whereas the other major groups of Neotropical Satyrinae, the Haeterini and the Euptychiina, are mostly lowland or premontane. Approximately 95% of all recognized species of Pronophilina occur in the tropical Andes from Venezuela to northern Argentina. T hey are by far the best represented group of butterflies in terms of species richness and abundance in cloud forest habitats (Pyrcz, 2004). The most outstanding biological feature of the pronophilines is their sensitive relationship with altitude, and most species are distributed in well defined, and sometimes very narrow, altitudinal bands (Adams, 1985, 1986; Raguso and Gloster, 1993; Pyrcz and Wojtusiak, 1999, 2002 cited in Pyrcz, 2004). Another notable feature of the pronophilines is the high percentage of endemic taxa in all surveyed mountain ranges (Pyrcz, 2004). Pronophiline butterflies are mostly associated with cloud forest habitats where poaceous plants, known or presumed to be their larval host plants, are particularly abundant (Viloria, 2004). Even though the immature stage biology of the Pronophilina remains largely unexplored, it appears that their larvae are oligophagous on montane bamboo, chiefly of the genus Chusquea (Poaceae) (Pyrcz, 2004). Several species of
15 Pronophilina apparently also use other Poaceae, such as pramo Stipa and Calamagrostis grasses, woody cane (Miller, 1986) and Guadua bamboo (Pyrcz, 2004; Beccaloni et al., 2008). Most pronophiline species, similar to other satyrines, show restricted vagility (Adams, 1986), and even thoug h this aspect of their behavior has not been studied rigorously, field observations indicate that adults keep close to their potential host plants or roosting places and move little vertically or horizontally. Some species are territorial and "hilltopping" behavior is characteristic for Junea, Steremnia and some species of Corades and Daedalma (Pyrcz, 2004) (author names for all neotropical butterfly taxa are listed in Lamas (2004) and are therefore not included in this thesis). Despite the fact that the An des have been the focus of intensive collecting during the past century, and that in recent years several researcher have spent a significant amount of time surveying this area (e.g., Lamas, Pyrcz, Viloria, Willmott and Hall), numerous new taxa are still described every year, including both species and genera. Furthermore, the recent use of molecular sequence data in several phylogenetic revisions (Pea et al., 2006, Monteiro and Pierce, 2001) has highlighted the importance of these data in better understan ding species level relationships and to provide a basis for evolutionary studies. To date, however, no genus of neotropical Satyrinae has been thoroughly studied with the use of molecular sequence data. Among the Pronophilina, a number of genera have been the subject of recent taxonomic study (e.g., Lasiophila, Pyrcz, 1999, 2006; Lymanopoda, Pyrcz et al.,1999; Pronophila, Pyrcz, 2000; Manerebia, Pyrcz et al., 2006; Pedaliodes, Pyrcz and Viloria, 1999, Pyrcz et al., 2008). However, there remain several gener a that have been historically poorly studied or
16 could benefit especially from molecular study, including Altopedaliodes and Neopedaliodes. These two genera are restricted to the highest elevation forests and grasslands, and due to the inaccessibility of these areas, and due to their rather somber aspect, they are both difficult to collect and often overlooked, and thus very poorly represented in most collections. Some species have very restricted ranges, little is known about the natural history of either g enus, and in a number of cases the taxonomy requires closer study, particularly because the simple wing patterns make association of allopatric taxa complicated. Generic monophyly is based on few and perhaps unreliable morphological characters, and studies to corroborate the generic classification using molecular sequence data would be highly desirable. Finally, both genera are of interest from ecological, evolutionary, biogeographic and conservation perspectives, but as a necessary base for such studies it is essential to have a very clear taxonomy. Scope of the Project In this project I therefore examine the systematics and biogeography of Altopedaliodes and Neopedaliodes with the following objectives: 1 Cladistic analysis. I use cladistic analyses base d on genetic characters to infer species level phylogenies for both genera, and to clarify the species classification of Altopedaliodes 2 Revision of Altopedaliodes I present a revision of the genus Altopedaliodes which summarizes known systematic and biological information about the genus. 3 Effect of climate change. Because the narrow elevational distribution and low vagility of most species in these two genera make them potentially very susceptible to changes in their environment, I also examine pos sible effects of an increase in temperature on future distribution by using ecological niche modeling techniques.
17 CHAPTER 2 MOLECULAR PHYLOGENY OF ALTOPEDALIODES AND NEOPEDALIODES Introduction Altopedaliodes and Neopedaliodes are nymphalid satyrine genera restricted to the highest elevation forests and grasslands of the Andes mountain chain (Miller, 1968; Pyrcz and Viloria, 1999; Viloria et al ., 2008). The genera are of particular evolutionary and conservation interest because a number of species have v ery restricted ranges and little is known about their natural history. From a systematic viewpoint, both genera also require close study. For example, Altopedaliodes cocytia was formerly classified in the superficially similar Punapedaliodes but current r elation ships to that genus remain unclear. At the species level, Altopedaliodes has not been the subject of any comprehensive revision, and the simple wing patterns and relatively little morphological variation make association of allopatric taxa complicated. Viloria et al (2008) revised Neopedaliodes recognizing 13 species and 13 subspecies, but the status of some of these proposed taxa needs to be tested. Altopedaliodes and Neopedaliodes taxonomy has been based principally on morphological char acters, but their potential status as synapomorphies remains to be tested by cladistic analysis. In particular, a number of characters, like wing shape and lack of sexual dimorphism (Viloria, 1998), might prove to be ecological traits that evolved in response to a shared habitat. The recent use of molecular sequence data in several phylogenetic revisions (Pea et al ., 2006, Monteiro and Pierce, 2001) has highlighted the importance of these data in better understanding species level relationships and to provide a base for
18 evolutionary studies. There are very few published species level phylogenies of any pronophiline gen us to date, and none that has used molecular sequence data. Such studies are needed not only to clarify species limits, but also to clarify the generic classification and for use in biogeographic studies. In addition to their use in taxonomy, robust phylog enetic hypotheses are necessary to provide a framework for interpreting the evolution of putatively adaptive character systems. I therefore propose to test the monophyly of the genera Altopedaliodes and Neopedaliodes and provide the first phylogenetic hypothesis of these genera, using mitochondrial and nuclear genes. By including several individuals of some widespread, taxonomically complex species, I hope also to clarify some species limits. At present, due to the availability of material for molecular study, I am only working with species from Ecuador. In the future I will attempt to include all remaining species, in addition to morphological characters. Material and Methods Specimen Acquisition, Extraction and Amplification of DNA Samples of 21 dried, unprepared specimens of Altopedaliodes and 15 of Neopedaliodes collected by Willmott in Ecuador over the past 4 years were used in this study, representing 14 species (69% of Altopedaliodes and 38% of Neopedaliodes ). DNA was extracted from the middle and hind legs using Qiagen's DNA extraction kit, following the manufacturers protocol. The remaining body and wings are kept as voucher material and deposited at the Florida Museum of Natural History (FLMNH), University of Florida, Gainesville FL, USA. Standar d PCR (Polymerase Chain Reaction) methods were used to amplify target genes, including mitochondrial cytochrome oxidase subunit I (COI), cytochrome oxidase
19 subunit II (COII) and the nuclear gene EF 1 (Hilli s et al ., 1996; Whinnett et al ., 2005), the forward and reverse primers used were: COI, Ron (f) and Hobbes (r); COII, George (f) and Eva (r); EF 1 ef44 (f) and efrcM4 (r). These genes are commonly used in phylogenetic analyses of butterflies at species and higher levels, and in a combined analysis prov ed to be very effective in resolving relationships among satyrines (Pea et al ., 2006; Monteiro and Pierce, 2001; Murray and Prowell, 2005). I also obtained sequence data for other closely related pronophiline species from GenBank, including: Parapedaliod es parepa (Hewitson, 1862), Praepedaliodes phanias (Hewitson, 1862), Punapedaliodes flavopunctata Staudinger, 1894 ; Pherepedaliodes drymaea (Hewitson, 1858) Steromapedaliodes albonotata (Godman, 1905), Redonda empetrus Thieme, 1905 and Pedaliodes phrasiclea Grose Smith, 1900. These species belong to the Pedaliodes s. l. clade according to Brower (2008). I conducted three separate analyses, thus reducing the time needed by the computer programs for each analysis. The first included Corad es enyo which belongs to the Pronophila clade, as outgroup, with the ingroup consisting of all other genera that belong to the Pedaliodes s. l. clade, and one individual of each species in my study genera. The second and third analyses included Pedaliodes phrasiclea as outgroup, and all sequenced Altopedaliodes and Neopedaliodes individuals with a separate analysis for each genus DNA Sequencing and Alignment Both strands of purified DNA fragments for each gene were sequenced for each of the species with the same primers used to amplify them. Consensus sequences generated from each reaction were edited manually using BioEdit 188.8.131.52 (Hall 1999)
20 and then aligned using the general purpose multiple sequence alignment program Clustalw2 (Thomson et al ., 1 994) in Seaview 4.2 (Galtier et al ., 1996). The final character set included for Altopedaliodes 1000 bp of EF 1, 960 bp of COII, and 930 bp of COI, with the combined dataset of three genes containing 2890 bp, and for Neopedaliodes 1000 bp of EF 1, 900 bp of COII, and 950 bp of COI, with the combined dataset of three genes containing 2850 bp. Phylogenetic Analysis Analyses based on maximum parsimony (MP) using PAUP*4.0 (Swofford, 2002), and Bayesian inference (BI), using MrBayes 3.1 (Ronquist and Huelsenbeck, 2003) were carried out for the data set of the three genes combined, as well as for individual gene data sets. Maximum Parsimony (MP) Phylogenetic trees were inferred using MP as the optimality criterion, with characters equally weighted. Tree estimati on involved heuristic searches with the treebisectionreconnection (TBR) branchswapping algorithm, and stepwise addition with up to 1000 random starting trees to reduce the problem of tree islands. Strict consensus trees were computed where there was more than one equally parsimonious tree. Bootstrap values (Felsenstein, 1985), based on a full heuristic search of 1000 pseudoreplicates using TBR branchswapping, are presented as an indicator of branch support. Bayesian Inference (BI) Bayesian Inference us ed the GTR+G+I model of sequence evolution (see above). Markov chain Monte Carlo simulations from 200,000 generations, with tree sampling every 100 generations, were performed (this was the minimum number of generations
21 and sample frequency needed to obtai n a standard deviation of split frequencies below 0.01 for these data sets). To calculate the appropriate amount of burnin to exclude I used Tracer v1.4 (Rambaut and Drummond 2007) (25%=500 generations). Bayesian topology and branch posterior probabilities were computed by majority rule consensus after deleting 500 generations as burnin. I used FigTree (Rambaut and Drummond 2007) to draw the trees from the different analyses. Results The results from MP and BI analyses of the combined data set of the thre e genes (COI, COII and EF 1) are summarized in Figure21. The results of the MP, and BI for the combined data set were consistent in showing the monophyly of Altopedaliodes + Neopedaliodes in relation to the other genera of the Pedaliodes clade (Figure 21). There was support for the monophyly of the genus Altopedaliodes in the trees produced by and MP and BI analysis (55% (MP) bootstrap support, and 99% (BI) posterior probability support ). The genus Neopedaliodes was also monophyletic in the trees produce d by both analyses (69% (MP) bootstrap support, and 64% (BI) posterior probability support, respectively). The analysis of each of the genes separately yielded similar tree topologies to those obtained with the combined data set and these results are not shown. The phylogeny of the genus Altopedaliodes inferred by the two different methods, is shown in Figure 22. The same topology was obtained with the data combined and when each gene was analyzed separately. The phylogeny is broadly congruent between the two methods used (MP and BI). There are two consistent groups, including tena and relatives, and halli + perita. Individuals from all currently recognized species
22 clustered together However, the taxon nucea forms a clade that is rather distinct fro m tena The taxa pasicles and A. n. sp. clustered within A. tena The phylogeny of the genus Neopedaliodes inferred by the two different methods, is shown in Figure 23. The same topology was obtained with the data combined and when each gene was analyzed separately. The phylogeny is broadly congruent for MP and BI. Although none of the relationships among the species was well resolved, all individuals of the currently recognized species clustered together, with the exception of N. m. citrica and N. m. inn upta whose relationships were unresolved in the BI analysis.
23 Figure 21. Phylogenetic trees for 24 Pronophilina taxa based on the combined data set of three genes produced by different methods: A, Maximum parsimony; B, Bayesian Inference. Support values for nodes are above the branches (bootstrap support and posterior probability).
24 Figure 22. Phylogenetic tree of Altopedaliodes based on the combined data set of three genes produced by different methods: A, Maximum parsimony; B, Bayesian Inference. Support values for nodes are above the branches (bootstrap support, and posterior probability).
25 Figure 2 3. Phylogenetic tree of Neopedaliodes based on the combined data set of three genes produced by different methods: A Maximum parsimony. B Bayesian Inference. Support values for nodes are above the branches (bootstrap support, and posterior probability).
26 Table 21. Genbank accession numbers for the gene sequences used in this study. Taxa COI EF 1 Authors Parapedaliodes parepa DQ338591 DQ339007 Pea et al., 2006 Praepedaliodes phanias DQ338592 DQ339009 Pea et al., 2006 Punapedaliodes flavopunctata DQ338861 DQ339015 Pea et al., 2006 Panyapedaliodes drymaea DQ338855 DQ339006 Pea et al., 2006 Steromapedaliodes albonotata GQ357244 GQ357310 Pea et al., 2006 Redonda empetrus GQ357243 GQ357309 Pea et al., 2006 Pedaliodes phrasiclea GQ357238 GQ357304 Pea et al., 2006 Corades enyo GQ357227 GQ357294 Pea et al., 2006
27 CHAPTER 3 REVISION OF THE GENUS ALTOPEDALIODES Introduction to the Genus Altopedaliodes belongs to the subtribe Pronophilina, which includes 66 genera, and ca. 690 species (Lamas, 2004). They are butterflies of medium size, brown in color and with simple, pale linear patterns or dots on the wings. The genus contai ns some of the highest flying butterflies of the northern Andes, from southern Venezuela to southern Peru. The species occur from 30003500m, at the forest pramo ecotone, and in open pramo to above 4000m. Like all pronophilines, Altopedaliodes typically remain low among the vegetation and fly only during warmer weather. They reach their highest diversity in eastern Ecuador, where several species can be found along a single elevational transect. The most recent classification (Lamas, 2004) recognized 11 sp ecies and six subspecies (Figure 31) in Altopedaliodes Materials and Methods I conducted fieldwork in the Ecuadorian Andes, due to the high diversity of Altopedaliodes in that country (Table 1) and to prior knowledge of sampling areas. I visited several sites in the Cordillera Occidental (the western range) and the Cordillera Oriental (the eastern range). Sample sites included (Figure 32): Azuay : Parque Nacional Cajas, Ro Blanco, Sigsig Chiginda road, Gima area; Caar : Culebrillas lake; Cotopaxi : SalcedoTena road; Chimborazo: GuamoteMacas road. All of these sites lie between 3000 to 4000m, the altitudinal range of most of the species of Altopedaliodes Sites were selected to include isolated mountain massifs and span major river valleys, which proved to be biogeographically interesting features for both genera. In most sites there was easy access by car and natural habitat was still
28 present. Finally, several sites were selected to permit specimens of many taxa to be collected for molecular analysis. Specimens were collected using an entomological net with 46 ft handle. I patrolled the forest pramo ecotone, as well as pramo grasslands (Figure 33), during sunny hour s when the butterflies are active and are easier to locate and collect. Specimens were killed with a light pinch to the thorax, and then stored in a glassine envelope with information on the place, time, weather conditions, date and other relevant collecti on information. In addition to the specimens collected in the field, I examined specimens from the McGuire Center for Lepidoptera and Biodiversity of the Florida Museum of Natural History, University of Florida, Gainesville, U.S.A. (MGCL), in addition to s pecimens collected by Willmott and Hall from Ecuador prior to 2005 (Keith Willmott and Jason Hall collection, Gainesville, Florida, USA, KWJH). From these collections I obtained distribution data and borrowed specimens for morphological and molecular study Identification of the material was accomplished by comparison wit h photographs of type speci m e n s from the Lamas collection of photographs of neotropical butterfly type specimens in Lima, Peru. Type specimens examined in this way are housed in t he Natural History Museum London, UK (BMNH), Muzeum i Instytut Zoologii Polskiej A kademii Nauk Warsaw, Poland (MIIZ), Museo de Historia Natural, Universidad de Caldas, Manizales, Colombia (MHNUNC), and Muzeum Zoologiczne Uniwersytetu Jagielloskiego, Krakw, Poland (MZUJ). Original descriptions of generic and specific names were consulted, including Forster ( 1964), Weymer ( 1890), Thieme ( 1905 ), Felder and Felder ( 1867), Krger
29 ( 1924), Hewitson ( 1869a, b ), Le Crom ( 1994 2004) and Pyrcz and Viloria ( 1999, 2007) Study of modern specimens, and photographs of the type specimens and their original descriptions were used to corroborate the identity of names. I dissected as many species belonging to these genera as available to me. I dissected males and females when b oth were available, and were able to study males of A. tena A. nucea A. pasicles A. perita A. zsolti A. reissi A. n sp A. kurti A. halli and A. kruegeri and females of A.tena, A.pasicles A. nucea A. perita sorda A.perita perita, and A. zsolti citra. To help define generic limits, I also examined specimens in the related genus Neopedaliodes, including males of N. philotera lafebreae, N. entella N. juba juba, N. margaretha margaretha N. margaretha allyni N. michaeli innupta, N. micha eli citrica N. nora nora N. nora stephani N. phoenicusa and N. parrhoebia parrhoebia. Specimens of more distantly related genera were also dissected, including males of S teromapedaliodes albonotata, Panyapedaliodes drymaea Pedaliodes phanias P. phrasi clea and Parapedaliodes parepa, and females of P. drymaea, P. parepa, P. phrasiclea and P. phanias Abdomens of males and females were soaked in hot 10% KOH solution for 10 to 20 minutes and subsequently stored in glycerol for study under binocular microsc ope. A Leica MZ 12.5 microscope was used to study genitalia and other anatomical structures. The head, legs, antennae, palpi and abdomen were removed from specimens of A. tena and N. p. lafebreae and soaked in hot 10% KOH solution for 10 minutes and subsequently stored in glycerol for study under binocular microscope. For genitalia morphological terms I follow Klots (1956). Venation was examined by using bleach to make wing scales transparent. V enation terminology follows the terms
30 proposed by Comstock and Needham (1918), cells named by the anterior vein bounding them (Figure 3 4.A). Abbreviations for the wings include: VFW (Ventral Forewing), VHW (Ventral Hindwing), DFW (Dorsal Forewing), and DH W (Dorsal Hindwing). Finally, the distribution of androconial patches on the male forewing was examined, using alcohol to help to visuali z e this ; a drop of alcohol was placed on the dorsal part of the fore wing where androconial patches occur (Figure 3 5). Morphological drawings were made using a camera lucida. S ubsequently, these preliminary drawings were scanned and the final drawings were made using Adobe Illustrator CS4 and Photoshop CS4. During my field work in Ecuador I attempted to rear Altopedaliodes species by placing mated females in plastic bags with fresh leaves of Stipa sp a plant used by Cevallos (2007) and presumed to be the hostplant for this genus. Eggs were collected and placed in individual plastic bags with fresh leaves of Stipa sp These eggs were kept in my house in Cuenca, 2500 m at a constant temperature of approximately 13 C. Altopedaliodes Forster, 1964 Type species: Pronophila tena HEWITSON, 1869: 98, by original designation. History of classification Felder & Felder (1867) described the first species now placed in Altopedaliodes as Pronophila cocytia from Colombia, a genus that then included pronophiline species that are now placed in a variety of genera. Subsequently, Hewitson (1869, 1872) described from Ecuador three species, Pronophila tena Pronophila perita and Pronophila pasicles Weymer in 1890 described Pedaliodes reissi from Colombia, and Thieme (1905) described Pedaliodes nebris from Colombia.
31 In 1964 the genus Altopedaliodes was described by Forster and originally included five species: A. tena (Hewitson, 1869), A. reissi (Weymer, 1890), A. perita (Hewitson, 1868), A. nebris ((Thieme, 1905) and A. pasicles (Hewitson, 1872). Forster defined the new genus based on the shape of the wings, and he separated the new genus from Pedaliodes primarily by the rounded wings and the elongated, narrow f ore wings. Another major difference is in the shape of the aedeagus, which is, in contrast to the species of the genus Pedaliodes slender and elongated. Pyrcz and Viloria (1999) subsequently described two further species, A. zsolti and A. kurti The complete catalog of Altopedaliodes comprises also A. kruegeri Pyrcz 1999, A. halli Pyrcz 2004, A. cocytia placed by Adams (1986) in Punapedaliodes and later transferred by Pyr cz and Viloria (1999) to Altopedaliodes, A. tamaensis Pyrcz and Viloria, 2007 and an undescribed species from Ecuador. Twelve described and one undescribed species are currently recogni z ed (Figure 31). Diagnosis According to Viloria (1998) the adults of Altopedaliodes can be recognized by their small to medium size, no marked sexual dimorphism, antennal club formed gradually, narrow forewings, wing margins rounded and smooth, rounded apex, and androconial patches present on dorsal forewing. In my study, I found a number of morphological characters that show potentially phylogenetically informative variation. In the male genitalia, the shape and length of the aedeagus, shape and size of the saccus, shape of the valva, presence of ampullar proc ess, and shape and thickness of the uncus and subuncus vary between genera (Figure 34.B, 3 6). In the female genitalia, the ostium bursa is thicker in Altopedaliodes than in other genera. The position of the ductus
32 seminalis also varies between genera, and the signa are longer in other genera in comparison with Altopedaliodes (Figure 3 7, 3 8 ). The cladistic analysis (Chapter 2) supports the monophyly of Altopedaliodes with respect to other genera from the Pedaliodes clade. However, because the Pedaliodes clade is so diverse and the relationships of its members so poorly understood, it will be very important in future to include more genera and species which could give stronger support to the results obtained here. Generic Description The wing venation of the Altopedaliodes is typical of the sub tribe Pronophilina (Viloria, 1998) (Figure 34.A). On the head, the labial palpi are covered with brown hairs and the eyes are covered with several setae (Figure 34.C). The general color of the butterflies is brown with some variation in hue. Wings of some species have white spots on both the ventral and dorsal side, and some species have yellow bands on the VHW and DHW The legs are covered with dense setae (Figure 3 9 .B, C, D) and the abdomen is covered with br own hairs and scales (Figure 39 .A). The male genitalia of Altopedaliodes are characteri z ed by a small saccus, moderately long aedeagus, straight or slightly curved dorsoventrally, only slightly laterally asymmetrical, never contorted, subrectangular valvae, dorsally slightly serrate, with rudimentary or absent ampullar process in subapical position (Pyrcz and Viloria, 2004) (Figure 34.B). I n comparison with other genera of Pedaliodes s. l. the f emale genitalia are characterised by a thicker ostium bursae, by the ductus seminalis being in a basal position under the corpus bursae, and by the shorter signa
33 Biology Natural History Immature stages of Altopedaliodes tena were collected on Stipa sp (Poaceae) by Cevallos (2007), where females were observed ovipositing, but no adults were obtained in the laboratory. Altopedaliodes species inhabit pramo areas with extensive grass and some species (e.g. A. tena A. nucea) can be found a very high altitude; during my field work I collected specimens at 4000 m. Although oxygen becomes scarce at such elevations it seems that this does not have an effect on these butterflies I observed individuals feeding on flowers during sunny days with little wind, and it is possible that these butterflies are important pollinators because they are one of the few, and the most numerous, butterfly species that inhabits these ecosystems. Diversity and Distribution of the Genus Certain species of Altopedaliodes can be relatively abundant in the pramos of Venezuela, Colombia and Ecuador, where several individuals can be found flying during sunny days over the pramo However, due to frequent cloud and rain in this type of habitat, it is often difficult to observe and c ollect individuals, and this has contributed to their scarcity in collections. For example, in the MGCL most of the species we re until recently poorly represented, and with the exception of some of the most common taxa, such as A. tena, most taxa have only a few specimens and often lack females. It is in Ecuador where this genus reaches its greatest diversity, especially in the southern part of the country, where five species of Altopedaliodes can be found in different areas.
34 Altopedaliodes species account s Altopedaliodes Forster, 1964 cocytia (C. Felder and R. Felder, 1867) kruegeri Pyrcz, 1999 kurti Pyrcz and Viloria, 1999 nebris (Thieme, 1905) perita (Hewitson, 1868) a) perita (Hewitson, 1868) b) sorda Pyrcz, 2004 reissi (Weymer, 1890) a ) reissi (Weymer, 1890) b ) flavomaculat a (Kruguer, 1924) c) salazari Le Crom, 1994 d ) n. ssp new subspecies tamaensis Pyrcz and Viloria, 2007 tena (Hewitson, 1869) nucea Pyrcz and Viloria, 1999 new status pasicles Pyrcz and Viloria, 1999 zsolti Pyrcz and Vitoria, 1999 a) zsolti Pyrcz and Viloria, 1999 b) citra Pyrcz, 2004 halli Pyrcz andViloria, 2004 n.sp new species
35 Altopedaliodes tena (Hewitson, 1869) (Figures 3 1.A, 3 5.A, 3 7 .A and 310 .D) Pronophila tena Hewitson, 1890. TL: Ecuador, Chimborazo, Puyal, 4300m. Type BMNH [photograph examined]. ? tener ; Godman and Salvin 1891, (missp.?) Altopedaliodes tena : Forster, 1964 Altopedaliodes tena : Lewis, 1974. Pedaliodes tena: Lewis, 1974. Altopedaliodes tena tena: Pyrcz and Viloria, 1999. Altopedaliodes tena tena: Lamas, 2004. Diagnosis : The male of A. tena has four to five white postdiscal spots on the DFW in the cells: R5, M1, M2, M3 and Cu1; while A. nucea has five to six smaller white dots on the VFW and four to five on the VHW, and A. pasicles has only three larger white dots which are almost fused together on the DFW in the cells R5, M1 and M2. This species is smaller than A. nucea and almost of the same size as A. pasicles. The male genitalia of A. tena is relatively smaller than in other taxa, the aedeagus is more curved, and the dorsal ampullar process in the valva is a little smaller. The male androconial patches are present in the postdiscal area between the veins C2 and M3 close to the discal cell and entering into it. Distribution and Biology: This species is widely distributed in the central and northern part of Ecuador and it seems that the Ro Caar is the southern limit of its distribution, f rom where it is replaced southwards by A. nucea. These b utterflies inhabit the forest p ramo ecotone on both slopes of the Cordillera Oriental and Occidental. I
36 collected both A. tena and A. nucea in the same place, with no evidence of intermediates, along the road to Laguna Culebrillas. Material examined: 5 males, Ecuador, Cotopaxi, Panzarumi, 3900 m, 09312007, K.R. Willmott, MGCL; 5 males, Ecuador, Napo, PapallactaOyacachi km 8, 3720m, 1102 2007, K.R. Willmott, MGCL; 2 females, Ecuador, Napo, Papallac ta Oyacachi km 8, 3720m, 1102 2007, K.R. Willmott, MGCL; 1 male, Ecuador, Morona Santiago, Cebadas Macas km 50, 3120, 0927 2007, K.R. Willmott, MGCL; 1 male, Ecuador, Napo, Quito, Baeza road east of pass 3350m 3800m, 09 271995, K.R. Willmott, KWJH; 1 ma le, Ecuador, Pichincha, Volcn Pichincha, 4100m, 0719 2009, K.R. Willmott, MGCL; 3 males, Ecuador, Carchi, Volcn Chiles, 3500m, 0624 1994, K.R. Willmott, KWJH; ; 1 female, Ecuador, Carchi, Volcn Chiles, 3500m, 06241994, K.R. Willmott, KWJH; 15 males, Ecuador, Chimborazo, GuamoteMacas road, 06262009, S. Padrn, to be deposited in MGCL; 37 males, Ecuador, Caar, Culebrillas, 3650m, 0713 2009, S. Padrn, to be deposited in MGCL; 7 males, Ecuador, Cotopaxi, SalcedoTena road, 4000m, 06222009, S. Padrn, to be deposited in MGCL; 8 males, Ecuador, Pichincha, Near Quito, 10500 ft, 03 101977, G.T. Austin, MGCL; 6 males, Ecuador, Pichincha, Pasochoa, 3300m, 1112 1938, F.M. Brown, MGCL. Altopedaliodes nucea Pyrcz and Viloria, 1999 new status (Figures 3 1 .B, 3 5.B, 3 7.C and 310 .C) Altopedaliodes tena nucea: Pyrcz and Viloria, 1999; TL: Ecuador, Azuay, Nab n; Type MZUJ [not examined]. Altopedaliodes tena nucea: Lamas, 2004.
37 Diagnosis : Altopedaliodes nucea can be differentiated from the other species by the presence of a greater number of small white submarginal dots on the wings; these are located on the DFW in cells R4, R5, M1, M2, M3 and Cu1. The male genitalia are characterized by a wider uncus and a most distinctive ampullar process in the valva, and the presence of small teeth inside this process. The DFW male androconial patches are reduced in the postdiscal area between veins Cu1 and M2. In this revision A. tena nucea is treated as species, due to morphological and molecular differences between the taxon and A. tena in addition to new field data that suggest sympatry between A. nucea and A. tena in southern Ecuador. Altopedaliodes nucea is distributed in southern Ecuador, while the other species of the tena clade A. pasicles A. tena and A. n. sp. have their distribution confined to the central and northern part of Ecuador. Distribution and Biology: This species is found in both eastern and western pramos of the Andes mountains in Azuay and Morona Santiago. I collected this species and A. tena fro m the same place but at different times; during the morning I collected principally A. tena, and at noon most of the specimens collected were A. nucea. In the Bueran area I collected several females that I kept alive in plastic bags with some fresh leaves of Stipa sp. a few hours after capture, one female laid eight eggs on leaves. Only one of the eggs hatched after 9 days, but despite being provided with fresh leaves of Stipa sp ., after another 8 days the larva died, having not fed. I kept the first instar in 70% ethanol; the larva is illustrated in Figure 3 13.
38 Material examined: 2 males, Ecuador, Azuay, GualaceoLimn road, 3215m, 104 2007, K.R. Willmott, MGCL; 1 female, Ecuador Loja, JimburaSan Andres road Km. 21, 3400m, 09221997, K.R. Willmott, KWJH; 1 female, Ecuador, Loja, JimburaSan Andres road Km. 21, 3400m, 09221997, K.R. Willmott, KWJH; 1 male, Ecuador, Morona Santiago, ChigindaGualaceo road Km. 24, 3250m, 11201997, K.R. Willmott, KWJH; 2 males, Ecuador, Morona Santiago, GimaSan Miguel Km. 16, 3320m, 10162007, K.R. Willmott, MGCL; 1 female, Ecuador, Morona Santiago, GimaSan Miguel Km. 16, 3320m, 10162007, K.R. Willmott, MGCL; 1 female, Ecuador, Morona Sant iago, Cerro Gallil, 3130m, 1015 2007, K.R. Willmott, MGCL; 2 males, Ecuador, Caar, Cuesta norte Azogues, 02091997, K.R. Willmott, KWJH; 8 males, Ecuador, Azuay, Gima, 3400m, 12222009, S. Padrn, to be deposited in MGCL; 3 females, Ecuador, Azuay, Gim a, 3400m, 12 222009, S. Padrn, to be deposited in MGCL; 23 males, Ecuador, Caar, Culebrillas, 3650m, 07132009, S. Padrn, to be deposited in MGCL; 4 females, Ecuador, Caar, Culebrillas, 3650m, 07132009, S. Padrn, to be deposited in MGCL; 10 males, Ecuador, Caar, Bueran, 3900m, 05232009, S. Padrn, to be deposited in MGCL; 5 females, Ecuador, Caar, Bueran, 3900m, 0523 2009, S. Padrn, to be deposited in MGCL; 5 males, Ecuador, Azuay, Dos Chorreras lagoon, 3800m, 05182009, S. Padrn, to be dep osited in MGCL. Other locality data: 6 males, Ecuador, Azuay, Nabn, 05031997, S. A ttal MZUJ; 1 female, Ecuador, Azuay, Nabn, 03 031997, S. Attal. MZUJ; 2 males, Ecuador, Azuay, Barabn, 0501 1994, F. Pias; 12 males, Ecuador, Azuay, Jima, 4300 m, 10 1997, I. A ldas TWP and UCP; 2 males, Ecuador, Azuay, W of Cuenca, Ro
39 Mazn, 3450 m, 0812 1986, M. J. and J. A DAMS, BMNH; 23 males, Ecuador, Azuay, Jima, 4300 m, 101997, I. Aldas, TWP (Pyrcz and Viloria, 1999). Altopedaliodes pasicles ( Hewitson, 1872) (Figures 3 1.C, 3 5.C, 3 7.B and 310 .E) Pronophila pasicles Hewitson, 1872. TL: Ecuador, Chimborazo, Allatillo. Type [photograph examined]. Altopedaliodes tena pasicles : Racheli and Racheli, 2001. Altopedaliodes pasicles : Lamas, 2004. Diagnosis : This species is easily recognized due to the presence of only three, broad, white dots on the DFW in cells R5, M1 and M2, which are almost fused between them. In addition, the overall wing color is slightly darker. The male genitalia have a wider tegumen and valva, the apical process of valva is reduced compared to related species, and the aedeagus is straight, unlike A. tena and A. nucea where it is slightly curved. The male androconial patches are in the postdiscal area between Cu2 and M3, and are broad er than in A. tena and A. nucea covering the median one fourth and entering the discal cell. Distribution and Biology: Altopedaliodes tena pasicles has a very restricted distribution; it has been collected only from a single site. Hewitson (1872), 134 years ago, described this subspecies from Allatillo (Atillo lagoons area), on the trail from the highlands to the lowland town of Macas, and subsequent ly only Pyrcz, Willmott and Petit have collected the taxon, during recent years from the same place. Although the taxon is presumably more widespread than suggested by the single locality from where it is known, it is common where it occurs and its global distribution is almost certainly
40 very restricted. The conservation status of the taxon is therefore likely to be sensitive to future environmental changes or to the degradation of habitat. Material examined: 8 males, Ecuador, Morona Santiago, Laguna Negra, Macas road, 3480 m, 10272007, K.R. Willmott, MGCL; 2 females, Ecuador, Morona Santiago, Laguna Negra Macas road, 3480 m 10 27 2007, K.R. Willmott, MGCL. Other locality data: 1 male, Ecuador, Morona Santiago, Atillo and Zuac, 3300m, 0216 ??, J.C. Peti t (Petit, 2010). Altopedaliodes kurti Pyrcz and Viloria, 1999 (Figures 3 1.D, 3 5.I and 311. D ) Altopedaliodes kurti Pyrcz and Viloria, 1999. TL: Ecuador, Loja, sector Yangana, 2850m. Holotype Altopedaliodes kurti : Lamas, 2004. Diagnosis : These butterflies are similar to A. perita but differ from that species because A. kurti does not have white dots on the DFW and DHW in the postdiscal area, the wings are of a lighter ground color and the ventral side of the wings has a uniform pattern. Altopedaliodes kurti has a white postdiscal line running from vein M2 to Cu2 in the postdiscal area on the VHW. The male genitalia is sightly similar to that of A. tena but the latter is relatively larger, especially the uncus and ampullar process, and in A. kurti the compression in the basal extremity of the aedeagus is absent. Finally, the androconial patches are present in the postdiscal area between veins Cu2 and M3 and covering the median one fourth, entering the discal cell with a small part that is adjacent to vein A2. These differ from the androconial patches present in A. tena where these are not adjacent to vein A2.
41 Distribution and Biology: This species has been c ollected in the southern Ecuador (JimburaSan Andrs road and in Yangana) and appears to replace A. nucea south of the Saraguro area. Material examined: 2 males, Ecuador, Loja, km 21 JimburaSan Andrs road, 3400 m, 11221997, K. R. Willmott, KWJH. Other locality data: 1 male, Ecuador, Loja, Sector Yangana, Parque Nacional Podocarpus, above 2850 m, 10271994, Parrots in Peril Expedition, BMNH ( Pyrcz and Viloria, 1999). Altopedaliodes kruegeri Pyrcz and Viloria, 1999 (Figures 3 1.E, 3 5.J and 3 11 C ) Altopedaliodes kruegeri Pyrcz, 1999 (repl. name). TL: Colombia (Lamas, 2004), Type Pedaliodes paeonides var flavopunctata Krger, 1924 (preocc. Staudinger, 1894). Altopedaliodes flavopunctata: Adams, 1986 (misidentification). Altopedaliodes kruegeri : Lamas, 2004. Diagnosis : Th e color of the wings is a uniform brown, the VHW has five light white dots and two to three barely visible white dots in the subapical area on the VFW, which are smaller than those present in A. nebris The male genitalia differs from all other congeners because it has a relatively large and robust uncus, the ampullar dorsal process on the valva is well defined and projects apically, and the aedeagus is relatively large, though smaller than in A. nebris
42 Androconial patches are present in the postdiscal area in cells M1, M2, M3 Cu1 a small area in A2, and in the distal, posterior corner of the discal cell between the cells Cu2 and M2; there is also an androconial patch along both sides of vein A2. Pyr cz (2004) suggested that this species might not belong in the genus Pedaliodes, but be better placed in Pedaliodes This species is included within Altopedaliodes here due to morphological characteristics which are typical of the genus (shape and length of the aedeagus, shape and size of the saccus, shape of the valva, presence of ampullar process, and shape and thickness of the uncus and subuncus), and because in the cladistic analysis based on molecular sequence data it was recovered in the Altopedaliodes clade. Distribution and Biology: This species is distributed in Colombia in the Volcn Purac massif in the Central Cordillera in Hula and Cauca, and also has been found on both slopes in northern Ecuador in Carchi and Napo. Material examined: 3 males, Ecuador, Napo, PapallactaOyacachi road km 5.5, 3460m, 1102 2007, K.R. Willmott, MGCL; 2 males, Ecuador, Carchi, Volcn Chiles, 3500m, 0624 1994, K.R. Willmott, KWJH; 1 female, Ecuador, Napo, QuitoBaeza road east of pass, 3350m, 09271995, K.R. Willmo tt, KWJH. Other locality data: 1 male, Colombia, Cauca Volcano Purac massif in the Colombian Central Cordillera, T.W. Pyrcz, TWP; 1 male, Colombia, Antioqui a Pramo de Belmira, T.W. Pyrcz, TWP, ( Pyrcz and Viloria 1999). Altopedaliodes halli Pyrcz, 2004 (Figures 3 1.F, 3 5.G and 311 B )
43 Altopedaliodes halli, Pyrcz, 2004. TL: Ecuador, Loja, Cerro Toledo, 3000m; Holotype Altopedaliodes n. sp.: Lamas, 2004. Diagnosis: This species was described recently by Pyrcz ( 2004), and can be dist inguished from the most similar species A. zsolti zsolti by the dorsa color, which is slightly darker, and by the absence of white spots that are present on the VHW of A. zsolti zsolti. The latter also has pale white spots on the DFW and DHW that are absent in A. halli. The male genitalia are similar to A. perita but in A. halli the ampullar process is just visible, unlike A. perita where it is absent. The aedeagus is moderately long and wide dorsally, and the saccus is deep compared with A. perita The androconial patch covers one fourth of the wing, entering the discal cell and slightly extending along veins between Cu2 and R5, and there is also a patch along both sides of vein A2 in the cells A1 and A2. The androconial patches in this species are t he most extensive of any species in the genus. Distribution and Biology: This species is present in the southern part of Ecuador, in the province of Loja and Morona Santiago, where it has been collected only from two locations in Loja and from one location along the Gualaceo Chiginda road. Material examined: 1 female, Ecuador, Morona Santiago, ChigindaGualaceo road km 24, 3180 m 10 14200 7, K.R. Willmott, MGCL 1 male, Ecuador, Loja, Cajanuma, 2800m 10111996, K.R. Willmott, KWJH. 1 male, Ecuador, Lo ja, Cajanuma, 3000, 0812 2009, J. Radford, MGCL. Other locality data: 4 males, Ecuador, Loja, Podocarpus National Park, Cerro Toledo, 3000m, 0915 2003, I. Aldas, TWP (Pyrcz, 2004).
44 Altopedaliodes cocytia (C. and R. Felder, 1867) (Figures 3 1.G, 3 5.F, and 311. A ) Pronophila cocytia C. and R. Felder, 1867. TL : Colombia (Lamas, 2004). Type BMNH [photograph examined]. Pronophila phaesana: Hewitson, 1868. Altopedaliodes cocytia : Forster, 1964 Altopedaliodes cocytia: Lamas, 2004. Diagnosis : Individuals of this species can be separated from other members of the genus by the pattern on the VHW, where there is a postdiscal pale yellow band, which is continuous and uniform in width, extending straight from vein A2 to M2. A further distinctive characteristic is the absence of other patterns or dots on the DFW and DHW. Th e male genitalia are very different from these of other species of Altopedaliodes, especially the distinctive shape of the aedeagus and valve. The aedeagus is straight, but in the middle part the width increases several times in relation to the proximal and distal parts. The valva is very broad in relation to other species and the dorsal process is visible. The uncus is large in relation to the subuncus, and the saccus is reduced. Androconial patches are present between veins Cu2 and M3, and slightly extend into the discal cell between these two veins. Material examined: 2 males, Colombia, Cundinamarca, Bogot, Monserrat, 3000m, 0220 1944, Coll. Hovanitz, MGCL; 1 female, Colombia, Cundinamarca, Bogot, Monserrat, 3000m, 02201944, Coll. Hovanitz, MGCL; 1 male, Colombia, Tolima, Coello river basin, Coll. Hovanitz, MGCL.
45 Altopedaliodes zsolti Pyrcz and Viloria, 1999 These butterflies have the dorsal surface of both wings uniform, dark brown, with five white dots on the DFW, VFW and VHW in the cells R5, M1, M2, M3 and Cu. The HW outer margin appears slightly undulate, similar in appearance to A. perita, but the latter species is generally smaller with a lighter brown dorsal surface. On the VFW and VHW there are white distal marginal fringes. Distribution and Biology: This species occurs in the mountains of Azuay, Morona Santiago and Loja in southern Ecuador. Altopedal iodes zsolti zsolti Pyrcz and Viloria, 1999 (Figures 3 1.H, 3 5.K, and 312. B ) Altopedaliodes zsolti zolti Pyrcz and Viloria, 1999. TL: Ecuador, Loja, sector Yangana. Type Altopedaliodes zsolti : Lamas, 2004. Diagnosis: This subspecies can be differentiated from the other subspecies by the small size of the white dots on the DFW, and by the postdiscal whitish band, which extends from costa to cell M2M3 on the VHW, being less distinct than in A. zsolti citra. The male genitalia have a more visible ampullar process in the valva, the aedeagus is very long and is slightly curved and the uncus is not as long as in A. zsolti citra The male androconial patch is present in the postdiscal area and in the discal cell between Cu2 and M2, and there is also a patch along the anterior edge of vein A2. Distribution and Biology: This subspecies is present in Yangana, Podocarpus National Park in Loja (Pyrcz and Viloria, 1999) and along the Jimbura San Andrs road near the border with Peru.
46 Material examined: 1 male, Ecuador, Loja, km 21 JimburaSan Andrs road, 3400 m, 11221997, K. R. Willmott, KWJH Other locality data: 1 male, Ecuador, Loja, sector Yangana, Podocarpus National Park, above (?) 2850m, 10271994, Parrots in Peril Expedition, BMNH (Pyrcz and Viloria, 1999). Altopedaliodes zsolti citra Pyrcz and Viloria, 2004 (Figures 3 1.I, 3 5.L, 37.F and 312. A ) Altopedaliodes zsolti citra Pyrcz and Viloria, 2004. TL: Ecuador, Morona Santiago, Sigsig Chiginda road; Type Altopedaliodes zsolti n.ssp: Lamas, 2004. Diagnosis: The white postdiscal dot on the DFW in cell Cu2Cu1 is bigger than the remaining dots, being almost twice the size of the adjacent dot, whereas in A. zsolti zsolti these dots are of equal size. On the VHW, the post discal band from costa to cell M2 M3 is mo re visible, and there is a white band in the postdiscal area on the DFW between M2 and Cu1, which is more defined than in A. zsolti zsolti. The male genitalia are relatively smaller, and the uncus is longer but the subuncus smaller than in A. zsolti zsolti The ampullar process in the valva is reduced, the aedeagus is half the size of the aedeagus in the nominotypical subspecies and it is less curved. The male DFW androconial patches are similar to those in A. zsolti zsolti. Distribution and Biology: This taxon occurs in the provinces of Azuay and Morona Santiago on the eastern side of the Cordillera Oriental Material examined: 1 female, Ecuador, Morona Santiago, ChigindaGualaceo road km 23, 3150m 10 142007, K.R. Willmott, MGCL ; 2 male s, Ecuador, Morona
47 Santiago, ChigindaGualaceo road km 24, 32 50 m 11201997, K.R. Willmott, KWJH; 1 male, Ecuador, Morona Santiago, Patococha, 3600 m 1006 2007, K.R. Willmott, MGCL; 12 males, Ecuador, Morona Santiago, Gima, 3800 m 12 22 2009, S. Padrn to be d eposited in the MGCL. Other locality data: 1 male, Ecuador, Morona Santiago, Sigsig Chiginda road, Granadillas, 30003200m, 11?? 1997, I. Aldas MZUJ. 3 male s Ecuador, Morona Santiago, Sigsig Chiginda road, Granadillas, 30003200m, 12 031996, P. Boyer, TWP; 2 males: Ecuador, Azuay, GualaceoLimn road, La Virgen, 33003350 m 02 08 2002, T. Pyrcz. TWP; 3 males, Ecuador, Morona Santiago, Las Antenas road, 3500m, 12031998, P. Boyer. PB; 1 male, Ecuador, Azuay, La Virgen, 3250m, 0114 2004, M. Bollino and F. Vitale, MB ( Pyrcz and Viloria, 2004). Altopedaliodes nebris (Thieme, 1905) (Figures 3 1.J, 3 5.H, and 311. E ) Pedaliodes nebris Thieme, 1905. TL: Colombia (Lamas, 2004). Type [photograph examined]. Pedaliodes nebris var. albipunctata Apolinar, 1914; TL: Colombia (Lamas, 2004). Pedaliodes nebris var. athymi (Apolinar, 1914); TL: Colombia (Lamas, 2004). Pedaliodes nebris var. conchae (Apolinar, 1914); TL: Colombia (Lamas, 2004). Pedaliodes nebris var. abadiae (Apolinar, 1914); TL: Colombia (Lamas, 2004). Pedaliodes nebris var. pauli Apolinar, 1914; TL : Colombia (Lamas, 2004). Pedaliodes nebris var. tripunctata (Apolinar, 1914); TL: Colombia (Lamas, 2004). Pedaliodes nebris var. modesta (Apolinar, 1914); TL: Colombia (Lamas, 2004). Pedaliodes nebris var. e stanislaoi (Apolinar, 1914); TL: Colombia (Lamas, 2004).
48 Altopedaliodes nebris : Forster, 1964. Altopedaliodes nebris : Adams, 1986. Altopedaliodes nebris : Lamas, 2004. Diagnosis : This species is easily distinguished from other species by the presence of a yellow post median band on the DHW and VFW; unlike A. tamaensis in which the yellow post median band is present on the VHW, in A. nebris it is wider and reaches the anal margin. The male genitalia is characterized by the large size of the aedeagus in relation to other species; in A. nebris the aedeagus is thicker and longer than in other species, being twice the size of the valva. In addition, the tegumen is wider than in other species. The male DFW androconial patches are very reduced in the postdiscal area and in the discal cell between veins Cu2 and Cu1. Apolinar (1914) described eight forms based only on variation in the wing patterns; all of these names were synonymized by Forster (1964). Distribution and Biology: This species is distributed in eastern Colombia. Material examined: 3 males, Colombia, Cundinamarca, Bogot, Bogot pramo, 80009000 ft Hovanitz, MGCL ; 2 males, Colombia, Cundinamarca, mountains, 85009000 ft Hova nitz, MGCL ; 1 female, Colombia, mtns nr. Bogot, 8500 9000 ft, 23 XI 1943, Hovanitz, MGCL. Altopedaliodes perita (Hewitson, 1868) These butterflies are similar to A. zsolti, with the dorsal surface of both wings uniform, dark brown, with five yellow dots between the cells R5, M1, M2, M3 and Cu1 on the DFW, VFW and VHW. The species differs from A. zsolti in being smaller and
49 paler dorsally. The male genitalia are characterized by the absence of the a mpullar process in the valva, which is pres ent in some degree in the other species of Altopedaliodes Distribution and Biology: These butterflies are found in the provinces of Azuay, Morona Santiago, Loja and Zamora in the southern part of Ecuador. Altopedaliodes perita perita (Hewitson, 1868) (Figures 3 1 .K, 3 7.E and 3 10 .A) Pronophila perita Hewitson, 1868, pl. 37, fig. 25 TL: Ecuador (Lamas, 2004). Type BMNH [photograph examined]. Altopedaliodes perita: Forster, 1964. Altopedaliodes perita: Pyrcz and Viloria, 1999: 121, fig. 4 (adult), 126, fig. 27. Altopedaliodes perita perita: Lamas, 2004. Diagnosis: A. perita perita is smaller, paler and more heavily patterned than A. perita sorda. A faint irregular postdiscal yellowish band is apparent in the VHW. The male genitalia differ in the uncus and subuncus being longer and the valva being wider than in the other subspecies. Distribution and Biology: A. perita perita is distributed in the province of Morona Santiago on the east Andean slopes of Ecuador. Material examined: 2 females, Ecuador, Morona Santiago, Rio Ishpingo, 3180, 09062007, K.R. Willmott, MGCL; 1 male, Ecuador, Morona Santiago, ChigindaGualaceo road km 24, 3150, 1120 1997, K.R. Willmott, KWJH; 1 male, Ecuador, Morona Santiago, Gualaceo Limn road km 22, 3215m, 1004 2007, K.R. Willmott, MGCL.
50 Altopedaliodes perita sorda Pyrcz, 2004 (Figures 3 7.D and 31 0 .B) Altopedaliodes perita sorda Pyrcz, 2004. TL: Ecuador, Zamora Chinchipe, El Paso, LojaZamora road, 2700m Type MZUJ [not examined]. Altopedaliodes perita n.s sp : Lamas, 2004. Diagnosis: This subspecies is typically larger than A. perita perita the wing color is darker and it lacks the five submarginal yellow dots that are present between the cells R, M1, M2, M3 and Cu1 on the DFW in the other subspecies. In the male genitalia, the valvae have a blunt tip and are slightly shorter than in the nominotypical subspecies, and the subuncus is t hin and reduced. Distribution and Biology: Altopedaliodes perita sorda is t he southern subspecies occurring in Podocarpus National Park and the Cordillera de Lagunillas along the border of the provinces of Loja and Zamora Chinchipe. Material examined: 1 female, Ecuador, Loja, Cajanuma, 3000, 0922 2007, K.R. Willmott, MGCL. Oth er locality data: 1 female, Ecuador, Loja, C ordillera Lagunillas, 3200, 09??2003, P. Boyer, PB; 8 males, Ecuador, Zamora Chinchipe, El Paso, 2800, 11221996, I. Aldas, TWP; 1 female, Ecuador, Zamora Chinchipe, El Paso, 2800, 11221996, P. Boyer, PB (Py rcz, 2004). Altopedaliodes tamaensis Viloria and Pyrcz, 2007 (Figures 3 1.L, 3 5.E and 3 11 F ) Altopedaliodes tamaensis Viloria and Pyrcz, 2007 TL: Colombia, Departamento Norte de Santander, Pramo del Tam. Holotype
51 Altopedaliodes n. sp. Pyrcz and Viloria, MS no. 539; Lamas et al. (2004: 207). Diagnosis: According to Pyrcz and Viloria (2007) this species is similar to A. nebris but can be distinguished by the absence of the white submarginal dots on the DFW and in the DHW, and also by the yellow post median band on the VHW being more diffused and extending along the anal margin. The male genitalia are characterized by a long aedeagus and by a reduced ampullar process in the valva, unlike in A. nebris where the ampullar process in the valva is more visible and the aedeagus i s thicker Distribution and Biology: This species is known from Colombia in the pramos of the eastern Cor dillera in the Tam region. Material examined: None. Other locality data: 1 male, Colombia, Santander, P ramo del Tam, 31003200m, 0216 1992, A. Viloria, MALUZ; 1 male, Venezuela, TchiraApure, Pramo del Tam, 3100 3350m, 02 181992, A. Viloria, MALUZ; 2 males, Venezuela, TchiraApure, Pramo del Tam, 3300m, 08201996, A. Viloria, MALUZ ( Viloria and Pyrcz, 2007) Altopedaliodes reissi (Weymer, 1890) Diagnosis: Individuals belonging to this species can easily be differentiated from other species in the genus by the presence of a large white spot on the DFW in the cells R1, R2 and R3 Another characteristic that can help distinguish this species is the presence o f a white dot on the VHW in cell Cu1, this is parallel to the fourth dot, but closer to the discal cell. Altopedaliodes reissi reissi (Weymer, 1890)
52 (Figure 3 1.M) Pedaliodes reissi Weymer, 1890. TL: Colombia (Lamas, 2004). Type uncertain] [ photograph examined]. Altopedaliodes reissi : Forster, 1964. Altopedaliodes reissi : Adams, 1986 Altopedaliodes reissi reissi : Lamas, 2004. Diagnosis: This subspecies can be distinguished from the other subspecies by the square white costal spot on the DFW between the cells R1, R2 and R3, which is smaller than in A. reissi salazari and A. reissi flavomaculata. Distribution and Biology: This subespecies is distributed in Colombia and has been collected by Adams (1986) in the Tolima area. Stbel also found A. reissi in the mountains of Popay n ( Adam s, 1986). Material examined: 1 male, Colombia, Pramos [photograph of specimen identified by G. Lamas, collection uncertain]. Altopedaliodes reissi salazari Le Crom, 1994 (Figures 3 1.N and 3 12 D ) Altopedaliodes reissi salazari, Le Crom, 1994. TL: Colombia, Caldas, P ramo de Letras, 3200m. Type MHNUNC] [photograph examined]. Altopedaliodes reissi salazari : Lamas, 2004. Diagnosis: According to Le Crom (1994), this subspecies can be distinguished from the other subspecies by the square white costal spot on the DFW, which is larger than in other subspecies, and by the male genitalia, where the tegumen is rounder and the tip of the vinculum is thinner than in other subspecies
53 Distribution and Biology: This subspecies is found in Colombia in Caldas, where it flies in the P ramo de Letras. Material examined: None. Other locality data: 1 male, Colombia, Caldas, P ramo de Letras, 3200m 02011991, MHNUNC; 1 female, Colombia, Caldas, P ramo de Letras, 3200m 08231993, MHNUNC (Le Crom, 1994). Altopedaliodes reissi flavomaculata (Krger, 1924) (Figure 3 1.O) Pedaliodes reissi var. flavomaculata Krger, 1924. TL: Colombia (Lamas, 2004). Lectotype Altopedaliodes reissi flavomaculata: Pyrcz, 1999. Altopedaliodes reissi flavomaculata: Lamas, 2004. Diagnosis: The characteristics that can be used to distinguish this subspecies, based on the photograph of the type specimen, include a difference in color in the pale markings of the DFW: in A. reissi reissi these are white, whereas in A. reissi flavomaculata they are yellow. T his feature is the only one that can be distinguished in the specimen photographed. Distribution and Biology: Nothing has been reported on the biology of this taxon, except that the label on the type specimen indicates that it occurs at 3000m. Material examined: None. Other locality data: 1 male, Neiva, 30.x.1917 (Pyrcz, 1999). Altopedaliodes reissi n. ssp. (Figures 3 1.P, 3 5.M and 312. C )
54 Diagnosis : This new subspecies is distinguished from the other subspecies by the small white costal spot on the DFW, which is very small in comparison with A. reissi reissi A. reissi flavomaculata and A. reissi salazari. The pattern on the DHW is different to the other subspecies because four white dots are visible between the cells R5, M1, M2, M3 and Cu1 in the postdiscal area. The male genitalia differ from A. reissi salazari in the size of the ampullar process of the valve; in A. reissi salazari this is very small, unlike the new species in which this is well developed. Distribution and Biology: This new subspecies is found in Ecuador on the PapallactaOyacachi road, a location that is very distant from other known localities for the species. Mol ecular sequence data were not available from other subspecies to further clarify the status of this new taxon. Material examined: 9 males, Ecuador, Napo, PapallactaOyacachi road km. 8, 3720m, 1102 2007, K.R. Willmott, MGCL. Altopedaliodes new species (F igures 3 1.Q, 3 5.D and 310.F) Diagnosis : This new species can be distinguished from the other species by the simple pattern on the DFW; there is a faint white band that runs from the costal border in the subapical area to reach vein R4. On the VFW this band is also visible and reaches vein R5 The male genitalia of this species possess the diagnostic charc teristics of Altopedaliodes including a small saccus, a moderately long aedeagus that is straight or slightly curved dorsoventrally, only slightly laterally asymmetrical, and sub rectangular valvae which are dorsally slightly serrate (Pyrcz and Viloria, 2004). Molecular sequence data also suppor t the inclusion of the new taxon in the genus
55 Altopedaliodes, with close relatives including A. tena A. nucea and A. pasicles This taxon is treated as a species due to the simple pattern of the wings, which lack the white dots that are present on the dorsal and ventral surfaces of the wings in A. tena a species that has been collected near the location of the new species and is appar ently sympatric or parapatric. Th e new taxon l acks the three fused white dots that are present on the dorsal and ventral s urfaces of the wings in A. pasicles and also the male genitalia differ from that species in the shape of the valva. Distribution and Biology: This taxon has been collected from a single site with precise, reliable locality data (Plano de Navarro on the SalcedoTena road). Although there are other specimens in MGCL that were collected in surroundings of Baos these data are vague. The distribution of this species is allopatric with the other species Altopedaliodes since the type locality is located in t he eastern part of the Cordillera Oriental, whereas specimens of A. tena have be en collected along the same road but in the west side of the Cordillera Oriental Material examined: 1 male, Ecuador, Napo, SalcedoTena road, 3290m, 09 302007, K.R Willmott, MGCL. 2 males, Ecuador, Tungurahua, Baos surroundings, R. Lafebre, MGCL.
56 Figure 31. Altopedaliodes species: A) A. tena; B) A. nucea; C) A. pasicles; D) A. kurti; E) A. kruegeri; F) A. halli; G) A. cocytia; H) A. zsolti zsolti; I) A. zsolti citra; J) A. nebris; K) A. perita perita; L) A. tamaensis (from Viloria and Pyrcz, 2007); M) A. reissi reissi; N) A. reissi salazari; O) A. reissi flavomaculata; P) A. reissi n. ssp; Q) A. n. sp.
57 Figure 32. Collection sites in Ecuador: A) Parque Nacional Cajas; B) Ro Blanco; C) Sigsig Chiginda road; D) Gima area; E) Culebrillas lake; F) Bueran; G) SalcedoTena road; H) GuamoteMacas road. Figure 33. Altopedaliodes habitat, Parque Nacional Cajas.
58 Figure 34. A. tena morphology drawings: A) Wing veins: A: anal, C: costal, Cu: cubital, Dc: discal cell, Hu: humeral, M: medial, R: radial, Sc: subcostal; B) Mal e genitalia(folowing Pyrcz and Viloria, 2004) : Ae:aedeagus, ap: apical process of valva, dp: dorsal(ampullar) process of the valva, Ss: saccus, Su: subuncus, T: tegumen, U: uncus, V: valva, Vm: vinculum; C) Head; D) Female genitalia.
59 Figure 35. Androconial patches: A) A. tena ; B) A. nucea ; C) A. pasicles ; D) A. n. sp, E) A. tamaensis ; F) A. cocytia ; G) A. halli; H) A. nebris ; I) A. kurti ; J) A. kruegeri ; K) A. zsolti zsolti; L) A. zsolti citra ; M) A. reissi n. ssp.
60 Figure 36. Male genitalia of other genera of the subtribe Pronophilina: A) N. philotera lafebreae; B) P. parepa ; C) S. albonotata; D) P. drymaea; E) P. phanias ; F, P. phrasiclea.
61 Figure 37 Female genitalia of Altopedaliodes : A) A.tena ; B ) A.pasicles ; C) A. nuc ea ; D) A. perita sorda ; E) A.perita perita ; F) A. zsolti citra
62 Figure 38. Female genitalia of other genera of the subtribe Pronophilina: A) P. drymaea; B) P. parepa ; C) P. phrasiclea; D) P. phanias
63 Figure 39 A. tena morph ology drawings: A) abdomen; B) f oreleg; C) midleg; D) hindleg.
64 Figure 310. Male genitalia, lateral view, aedeagus removed and illustrated in lateral view. A) A. perita perita; B) A perita sorda; C) A. nucea; D ) A. tena; E) A. pasicles ; F) A n.sp.
65 Figure 311. Mal e genitalia, lateral view, aedeagus removed and illustrated in lateral view. A ) A. cocytia ; B ) A. halli; C ) A kruegeri; D ) A. kurti; E ) A. nebris ; F ) A. tamaensis (re drawn from Viloria and Pyrcz, 2007).
66 Figure 312. Mal e genitalia, lateral view, aedeagus removed and illustrated in lateral view. A ) A. zsolti citra ; B ), A. zsolti zsolti; C ) A reissi n. ssp.; D ) A.reissi salazari (re drawn from Le Crom, 1994).
67 Figure 31 3. A. nucea first instar larva.
68 Table 31. Distribution of the Altopedaliodes species. Genus Species Subspecies Country Altopedaliodes cocytia Colombia nebris Colombia tamaensis Colombia kruegeri Colombia reissi reissi Colombia reissi flavomaculata Colombia reissi salazari Colombia reissi n.ssp Ecuador tena tena Ecuador nucea Ecuador pasicles Ecuador perita perita Ecuador perita sorda Ecuador zsolti zsolti Ecuador zsolti citra Ecuador halli Ecuador kurti Ecuador n.sp Ecuador
69 CHAPTER 4 DISTRIBUTION OF ALTOPEDALIODES AND NEOPEDALIODES SPECIES AND THEIR RESPONSE TO CL IMATE CHANGE IN ECUA DOR Introduction Prediction and understanding of species distributions are central to diverse applications in ecology, evolution and conservation (Holt and Keitt, 2005; Young, 2007, Rdder et al ., 2010). In the past century, global average temperature has increased by approximately 0.6 C ( Intergovernmental Panel on Climate Change, 2001), and future increases could have an important effect on the distribution of organisms o n the planet (Bachelet et al ., 2001; Lister and Stuart, 2008; Kannan and James, 2009). To predict some of the likely impacts of future climate change, we can compare current and future potential distributions of species. Impacts will dep end on the dispersal ability of individual taxa, relative to the speed and magnitude of shifts ( Lewis, 2006) and the availability of new suitable habitats after the shift. A number of studies have shown that butterflies are very susceptible to climate change and may respond by migrating to more suitable environments ( Hill et al ., 1999) or become extinct. Butterflies with a narrow elevational distribution tend to advance to higher altitudes in response to an increase in temperature. Parmesan and Yohe (2003) found evidence that climate change is already affecting living systems and predicted that in the future the average rangeshift will be 6.1 km per decade towards the poles and the same quantity in meters in altitude; these shifts might obviously have implications for the conservation of species. Recently, Thomas et al. (2004) assessed the extinction risks of several species in different regions of the planet under three different climatewarming scenarios. They found that an average of 1537% of species will be committed to extinction under these
70 scenarios. Although these studies have evaluated a large number of species and ecosystems, the vast majority of them have been focused on the effect of climate change on temperate species in highly disturbed habitats, rather than tropical species in less disturbed habitats. The study presented here is the first to evaluate the effect of climate change on butterflies in tropical high Andean ecosystems. The altitudinal distribution of Pronophilina satyrine butterfly species seems to be highly dependent on the presence of pristine forest, and it has been observed that patterns of vertical distribution are strongly altered by disturbance of the original vegetation (Adams, 1985 cited in Viloria, 2 004). I chose two s tudy genera, Altopedaliodes and Neopedaliodes that inhabit the very narrow ecotone between the highest elevation forests and pramo grassland, from 25004000m above sea level. In both genera, a number of species are known from very restricted ranges, and some from only a single site. These characteristics suggest that Altopedaliodes and Neopedaliodes are likely to be significantly affected by future climate change. The purpose of this study is to predict the future distributions of select Altopedaliodes and Neopedaliodes species, explore for general patterns and discuss the conservation implications of the results. Methods The response of each species geographic range to climate and vegetation change can be predicted from point locality data. There are many different methods to predict species current and future distribution (e.g. Elith et al ., 2006), but two of the most commonly used are MAXENT (Phillips et al ., 2004) and BIOCLIM ( Busby, 1991; Nix, 1986) I decided to use the BIOCLIM method within the free GIS computer program DIVA GIS (Hijmans et al ., 2004) to calculate the areas of each species' range according
71 to models based on their current distribution, and to estimate the habitable area after assumed climate change. The BIOCLIM method has been used in different organisms to predict distribution (Campbell et al. 1999; Elith et al ., 2006; Echarri et al ., 2009), including butterflies in Australia (Beaumont and Hughes, 2002) and USA (Sierra Nevada) ( Scheingross, 2007) BIOCLIM is a prediction system that uses bioclimatic parameters derived from mean monthly climate estimates, to approximate energy and water balances at a given location (Nix, 1986). The present version can produce up to 35 bioclimatic parameters based on the climate variables maximum temperature, minimum temperature, rainfall, solar radiation and pan evaporation. BIOCLIM was chosen because it has been demonstrated to be easy to use, accurate in its predictions, and, in contrast with other models, to produce only one outcome for a parti cular dataset. This allowed me to identify the effect of the different variables on the area predicted. Although BIOCLIM can interpolate up to 35 climatic parameters to predict potential distribution, the progressive addition of climatic parameters results in progressively smaller potential distributions, which may lead to misrepresentations of the potential distribution of species. Therefore, the inclusion or exclusion of parameters is an important consideration in implement ing this model, as I discuss further below. Specimen Data I selected nine species of Altopedaliodes ( A. nucea, A. tena A. halli, A. kurti A. n.sp, A. kruegeri A.reissi A. perita and A. zsolti ) and six species of Neopedaliodes ( N.entella N. philotera N. phoenicusa, N. parrhoebia N. michaeli and N. juba ) for which I was able to obtain precise and reliable distribution data.
72 The data came mainly from specimens collected by Willmott in Ecuador (unpublished data) and from my own fieldwork in that country. Data from the MGCL and publications were also used. Modern localities were georeferenced using Google Earth, and historical localities using a gazetteer of entomological stations in Ecuador (Brown, 1941). These data were imported into DIVA GIS to create a layer with the collection points. Climate Model and Calculation of D istributions I used climate data from the WorldClim database (version 1.4, Hijmans et al ., 2005) including current climate (~19502000) version 1.3, October 2004, with a resolution 2.5 minutes, and future climate under the increase of greenhouse gases (2x current CO2 conditions, CCM3 model projected to 2100 AD) from Govindasamy et al. (2003), with a resolution of 2.5 minutes. Elevation data for Ecuador at 2.5 minute resolution and admins trative boundaries for South America were also obtained from Hijmans et al. ( 2004) and land cover data were obtained from the Global 2000 Land Cover (GLC) dataset ( European Commission, 2003). To predict the actual and future distribution I selected two cl imatic variables in BIOCLIM: annual precipitation and annual temperature. I decided to work with only two variables, because the number of climatic variables used to predict the distribution of species is negatively correlated with the number of parameters incorporated in the model, with progressive addition of variables resulting in progressively narrower potential distributions (Beaumont et al ., 2005). I chose t hese two variables because they have been de monstrated to be important in the determining the distribution of butterflies (Thomas et al ., 2004).
73 I ran the analysis using the BIOCLIM option, which classifies grid cells into one of two categories. Those grids where all climate parameters lie within the 0 100 percentile of the species climate envelope are classified as suitable. Grids where at least one climate parameter within a grid cell is outside of the species climate envelope are classified as unsuitable (Nix, 1986). Some areas predicted to b e climatically suitable were excluded as likely to be unsuitable due to other factors, including environmental factors such as type of vegetation, presence of geographical barriers and state of conservation of habitat, or ecological factors, such as species replacement. I attempted not to exclude areas where the species may occur but hasn't been recorded yet because of poor sampling. T o exclude unsuitable areas I created a mask to include only the areas that are suitable for the species. The mask was based on the GLC landcover categories, considering the following vegetation coverages suitable for Altopedaliodes and Neopedaliodes : Shrub Cover, closedopen, deciduous; Herbaceous Cover, closedopen; Sparse Herbaceous or sparse Shrub Cover. I also considered th e influence of some geographic barriers such as river basins and valleys when creating masks I then reclassified the predicted distribution layer to reflect presence or absence only. This new classification has values of 0 for categories where the species is believed not to occur, and 1 for those where it is believed to occur. This new layer was overlaid on to the land cover dataset to obtain a final predicted distribution. To calculate the current and future distribution area and the difference between th em, I obtained a histogram of the frequency of grid cells in each of the vegetation cover categories, using the number of 2.5minute grid cells in a species distribution as
74 an approximation of area. A 2.5 minute grid cell at the equator has an area of appr oximately 21.44 km2 (4.63 km x 4.63 km). Then I compared the change in the number of grid cells (area) in the potential current distribution with the potential future distribution. This analysis was done for each genus, as well as for each species individu ally. I decided to do the former so as to increase the number of records by combining the sampling points of all species of each genus. Results Potential Current D istribution for Altopedaliodes S pecies The area of the potential current distribution of the 9 Altopedaliodes species in Ecuador was 754 grid cells, representing 16,163.42 km2 ( Figure 41). The potential current distribution area for each Altopedaliodes species in Ecuador was: A. tena 453 grid cells with an area of 9710.9 km2, A. nucea 254 grid cells with an area of 5445.4 km2, A. halli 13 grid cells with an area of 278.7 km2, A. zsolti 19 grid cells with an area of 407.3 km2, A. perita 7 grid cells with an area of 150.1 km2, A. n sp 1 grid cell with an area of 21.4 km2, A. kruegeri 5 grid cells with an area of 107.2 km2, A. kurti 1 grid cell with an area of 21.4 km2, and for A. reissi 1 grid cell with an area of 21.4 km2 (Fig ures 42, 4 3 and 44 ). Potential Current D istribution for Neopedaliodes S pecies The area of the potential current distribution of the 6 Neopedaliodes species in Ecuador was 2181 grid cell, representing 46,753.88 km2 (Fig ure 45). The potential current distribution area for each Neopedaliodes species in Ecuador wa s: N. phoenicusa 218 grid cells with an area of 4673.2km2, N. entella 147 grid cells with an area of 3151 km2, N. juba 693 grid cells with an area of 14,855.8 km2, N. philotera 409 grid cells with an area of 8767.7 km2, N. michaeli 214 grid cells with an
75 a rea of 4587.5 km2, and N. parrhoebia 500 grid cells with an area of 10,718.5 km2 (Fig ures 4 6 and 4 7). Potential D istribution for Altopedaliodes S pecies Under an Increase in T emperature Under a scenario of an increase in temperature, the potential distri bution for the genus Altopedaliodes in Ecuador was 687 grid cells, 14,727.15 km2 (Fig ure 4 1 ). This means a reduction in total distribution area of 1436.27 km2 (8.9% of the current potential distribution) (Fig ure 41) The potential future distribution area of Altopedaliodes species in Ecuador was: A. tena 406 grid cells with an area of 8703.4 km2, A. nucea 242 grid cells with an area of 5187.7 km2, A. halli 8 grid cells with an area of 171.5 km2, A. zsolti 11 grid cells with an area of 235.8 km2, A. perita 12 grid cells with an area of 257.2 km2, A. nsp 1 grid cell with an area of 21.4 km2, A. kruegeri 3 grid cells with an area of 64.3 km2, A. kurti 1 grid cell with an area of 21.4 km2, and for A. reissi 3 grid cells wit h an area of 64.3 km2 (Fig ures 4 2, 4 3 and 4 4). Potential D istribution for Neopedaliodes S pecies Under an Increase of T emperature Under a scenario of an increase in temperature, the potential distribution for the genus Neopedaliodes in Ecuador was 1974 grid cells, 42,316.4 km2 (Fig ure 45). This means a reduction in total distribution area of 4437.4 km2 (9.5% of the current potential distribution). The potential future distribution of Neopedaliodes species in Ecuador was: N. phoenicusa 236 grid cel ls with an area of 5059.1 km2, N. entella 102 grid cells with an area of 2186.6 km2, N. juba 573 grid cells with an area of 12.283.3 km2, N. philotera 382 grid cells with an area of 8188.9 km2, N. michaeli 189 grid cells with an area of
76 4051.6km2,and N. pa rrhoebia 492 grid cells with an area of 10547 km2 (Fig ures 46 and 4 7). Change in Range Size with Climate C hange Under a scenario of increasing global temperature, the potential future distribution area for both genera is reduced, by 8.9% for Altopedaliodes and by 9.5% for Neopedaliodes Most species of Altopedaliodes ( A. halli, A. kruegeri A. tena A. nucea and A. zsolti ) were predicted to suffer contractions in their distributional range under climate change. The average reduction in distri bution area was 44%. In the genus Neopedaliodes most species ( N. entella N. juba, N. philotera N. michaeli and N. parrhoebia) were also predicted to suffer contractions in their distributions under climate change, with an average reduction of 13.6 % Se veral species ( A. kurti and A. n sp ) where not predicted to have any change in their distribution in future. For a minority of Altopedaliodes species ( A. reissi and A. perita), the change in the size of their distribution was positive, with an average increase in their distribution of 54.4%. Within Neopedaliodes, only N. phoenicusa showed a positive change in distribution area, with an increase of 8.3%.
7 7 Figure 4 1. Maps showing current potential distribution and future potential distribution of Altopedaliodes species. A) Current potential distribution for the genus Altopedaliodes in Ecuador. B) Future predicted distribution under a climate change scenario.
78 Figure 4 2. Maps showing current potential distribution and future potential distribution of Altopedaliodes species. A) Current potential distribution of A. halli in Ecuador. B) Future potential distribution of A. halli in Ecuador. C) Current potential distribut ion of A. kruegeri in Ecuador. D) Future potential distribution of A. kruegeri in Ecuador. E) Current potential distribution of A. kurti in Ecuador. F) Future potential distribution of A. kurti in Ecuador.
79 Figure 4 3. Maps showing current potential dist ribution and future potential distribution of Altopedaliodes species. A) Current potential distribution of A. tena in Ecuador. B) Future potential distribution of A. tena in Ecuador. C) Current potential distribution of A. nucea in Ecuador. D) Future potential distribution of A. nucea in Ecuador. E) Current potential distribution of A. reissi in Ecuador. F) Future potential distribution of A. reissi in Ecuador.
80 Figure 44 Maps showing current potential distribution and future potential distribution of Neopedaliodes species. A) Current potential distribution of A.zsolti in Ecuador. B) Future potential distribution of A. zsolti in Ecuador. C) Current potential distribution of A. perita in Ecuador. D) Future potential distribution of A. perita in Ecuador. E) Current potential distribution of A. n.sp in Ecuador. F) Future potential distribution of A. n.sp in Ecuador.
81 Figure 4 5. Maps showing current potential distribution and future potential distribution of Neopedaliodes species. A) Current potential distribution for the genus Neopedaliodes in Ecuador. B) Future predicted distribution under a climate change scenario.
82 Figure 4 6 Maps showing current potential distribution and future potential distribution of Neopedaliodes species. A) Current potential distribution of N. entella in Ecuador. B) Future potential distribution of N. entella in Ecuador. C) Current potential distribution of N. philotera in Ecuador. D) Future potential distribution of N. philotera in Ecuador. E) Current pot ential distribution of N. juba in Ecuador. F) Future potential distribution of N. juba in Ecuador.
83 Figure 4 7. Maps showing current potential distribution and future potential distribution of Neopedaliodes species. A) Current potential distribution of N. michaeli in Ecuador. B) Future potential distribution of N. michaeli in Ecuador. C) Current potential distribution of N. parrhoebia in Ecuador. D) Future potential distribution of N. parrhoebia in Ecuador. E) Current potential distribution of N. phoenicusa in Ecuador. F) Future potential distribution of N. phoenicusa in Ecuador.
84 CHAPTER 5 CONCLUSIONS Molecular P hylogeny of Altopedaliodes and Neopedaliodes This study is the first phylogenetic analysis of the genera Altopedaliodes and Neopedaliodes and shows that the monophyly of both genera are well supported (Figure 2 1). This result was unaffected by either the optimality criterion used or the dataset that was analyzed, and corroborates previous studies that have recognized these genera as distinct taxa (Lamas, 2004; Forster, 1964 and Pea et al ., 2006, Viloria et al ., 2008). Furthermore, these results are supported by potential morphological synapomorphies, especially in the male and female genitalia, in comparison with the outgroup genera included in this analysis (in C hapter 3). One morphological character that might support a sister relationship between Altopedaliodes and Neopedaliodes as in the molecular results, is the ductus bursae, which is unsclerotized in Neopedaliodes and Altopedaliodes but sclerotized in examined outgroup species (in C hapter 3) ( Parapedaliodes parepa, Praepedaliodes phanias Punapedaliodes flavopunctata, Pherepedaliodes drymaea, Steromapedaliodes albonotata Redonda empetrus and Pedaliodes phrasiclea). The inferred phylogeny of Altopedaliodes was broadly independent of analytical method or dataset. An interesting result is that the taxon A. pasicles formerly considered a subs pecies of A. tena is recovered as sister to one individual of A. tena The latter specimen of A. tena was collected in the same place (km. 50 Cebadas Macas road) as the specimens of A. pasicles Although A. tena and A. pasicles seem to be morphologically distinct and sympatric, this close relationship between sympatric individuals might suggest recent hybridization, or that A. pasicles diverged from the
85 population of A. tena with which it is now sympatric. These results point to the necessity of a broader geographic sampling and genes to resolve this issue. It is also interesting that a taxon formerly considered a subspecies, A.tena nucea, appears sister to all other A. tena taxa. Combined with new field data that suggest sympatry between A. nucea and A. tena in southern Ecuador, it is seems most reasonable to treat A. nucea as a distinct species. Altopedaliodes nucea is distributed in southern Ecuador, while the other species of the tena clade; A. pasicles A. tena and A. n. sp. have their distribution confined to the central and northern part of Ecuador. The inferred phylogeny of the genus Neopedaliodes was also broadly independent of analytical method or dataset, and congruent with the current taxonomy. However, due to the few taxa evaluated in this analysis it is difficult to draw firm conclusions as to whether the current classification is reliable. In this analysis, N. p. lafebreae was considered a subspecies of N. philotera based on the morphology of the specimens studied, but since no specimens of N. philotera philotera were available for the molecular study, it is therefore very important in the future work to attempt to include additional taxa. The use of three genes in the combined analysis and separate analyses proved to be informative in the r econstruction of the phylogeny of these genera, although it is clear that it is necessary to use more genes and better taxonomic and geographic sampling in future analyses. It is also necessary to include all the recognized species in these genera, as this can help give more support to these preliminary results. More data and taxa will be needed to clarify the taxonomic status of certain problematic taxa, some of which have been suggested to be included in other genera. Finally, as part of my
86 future researc h, I intend to include morphological characters in a further phylogenetic analysis. The combinations of morphological and molecular characters have been proved to be very effective in resolving relationships among butterflies ( Wahlberg et al ., 2005). Distr ibution of Altopedaliodes and Neopedaliodes Species and their Response to Climate Change in Ecuador The current potential distribution area for the genera Altopedaliodes ( 16,163.42 km2) and Neopedaliodes ( 46,753.88 km2) represent 6.3% and 18.25%, respectively, of Ecuadorian mainland national territory. These areas are relatively small, considering that most of the future projected areas have some degree of intervention by man (cropland, deforested areas) which are likely not suitable for establishm ent of these species. The predicted range sizes for Altopedaliodes and Neopedaliodes species in Ecuador differ greatly among species, ranging from a single grid cell ( 21.4 km2, A. kurti A. nsp A. reissi ) to 693 grid cells ( 14,855.8 km2, N. juba ). Predi cted range area is highly influenced by the difference in the number of collection points across species, with greater area predicted for those species that have a greater number of sampling points. For example, the projected area for A. nucea with 9 collection points was 254 grid cells and for A. zsolti with 4 collection points were 19 grid cells. However, these results are influenced by factors other than sampling artifacts. The restricted global distribution that some species have, with sever al being known only from Ecuador, or even from only one or two localities, results in a small projected distribution area (e.g. A. pasicles and A. n.sp). Finally, differences in range size are also caused becauses certain species are restricted to specific habitats ( A. pasicles ), unlike other species ( A. tena and A. nucea )
87 that may use a broad variety of habitats, which allows them to have a wider distribution. A. pasicles has been collected only in one place in the GuamoteMacas road, unlike A. tena that has been collected in multiple sites on both Andean slopes in the northern part of Ecuador. Finally, in some cases geographic barriers such as rivers (e.g. Paute river basin, Caar river basin, Pastaza river basin) and mountain ranges were assumed to prevent the future displacement of species to areas that could be suitable. For example, A. nucea was predicted to occur west of the Cordillera Occidental in Azuay, but because of the large barrier of these high mountains, it is unlikely that this species in the future will cross the Cordillera Occidental to inhabit these new areas. This study showed that the likely impact of future climate change on the range sizes of two genera of highaltitude Ecuadorian satyrines was predominantly negative. This was true for each genus as well as the majority of species. Results from analysis of the entire genus differ from separate analyses of each species. Although only a reduction of 10% was predicted for the genus, an average reduction of 44% for individual species was predicted This difference can be explained by the fact that for each species I took into account more precise factors like geographic barriers lack of habitat, presence of replacement species etc which could not be taken into account for the analysis at the genus level. The results were not used to predict which species would go extinct, as other studies have done (Thomas et al ., 2004). Other studies have used a species area relationship to predict how area reduction will affect species richness, but we lack a
88 sufficient understanding of species area relationships in the Andes at present to allow this to be done. There was a notable difference the predicted range changes between Altopedaliodes and Neopedaliodes in the generic analysis with the genus Neopedaliodes predicted to have the bigger reduction in its range. However, it seems that the genus Altopedaliodes will be the most affected given that Altopedaliodes generally fly at higher elevations which makes them more at risk for future climate chang e, because these species may thus not be able to migrate to higher altitudes in response to an increase in temperature. There was also a difference in predicted impact between different areas in Ecuador. The southern areas of the country suffered the greatest reduction in habitat for these two genera, caused by the fact that in this area the Andes m ountains are of relatively low altitude, which reduces the future predicted area for these two genera. For the species that showed no difference in distributio n between now and future predictions, this was likely an artifact of very few collection points (see above). In the case of the few species that showed an increase in range size, these results should be taken with caution, since a large majority of the fut ure projected range might lack suitable habitat or be inaccessible because of geographic barriers Viloria et al. (2004) have emphasized the fact that adult pronophiline butterflies are very sedentary and heavily dependent on certain microhabit ats and that high elevation species have not only very low relative vagility, but in some cases females show varying degrees of flight loss (Adams and Bernard, 1981; Viloria et al ., 2003 cited in Viloria, 2004).
89 These results have implications for future pla ns to protect these species. For example, similar studies have been used to help guide conservation decisions (e.g. Rutherford et al ., 1999; Dockerty et al ., 2003; Hossell et al ., 2003), and to identify species most at risk of extinction (Busby, 1988; Thom as et al ., 2004). The predicted distributions can also be used as a tool to define priority areas for conservation. In the case of the butterfly species studied here, it is important to propose certain measures of protection for both existing habitats and those that have been projected in the future to be potentially inhabited by these butterflies. One of the priority measures would be to prohibit the burning of pramo and forest, which is a technique used by the rural inhabitants of the Andes to prepare the soil where potatoes will be planted; it is also a popular belief that burning pramo and forests attracts rain in times of drought. In future, this analysis should be expanded with more records for each species, to help to reduce sampling bias. In additi on, different climate scenarios corresponding to different CO2 accumulation predictions should be evaluated to test how robust are the conclusions to uncertainties in future predictions (Thomas et al ., 2004).
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98 BIOGRAPHICAL SKETCH Pablo Sebastin Padr n M. was born in 1983, in Caar Ecuador; he has always been attracted and fascinated by the complex shapes and beautiful colours in which nature is presented, and s ince childhood he has always enjoyed contact with nature. In 2001 he decided to study b iology at the Universidad del Azuay, Cuenca, Ecuador. D uring his career he has had the opportunity to work in several projects about insects and their relationship with the environment and he has spent several months collecting butterflies and insects in the jungles of Ecuador. In 2005 he obtained a scholarship to travel to Panama f or three months to work as a field assistant in the Smithsonian Tropical Research Institute. In 2008 he discovered and described a new subspecies Hyposcada illinissa tundayme, Padrn 2008. In the same year he was admitted into the Master of Science program in the University of Florida, for which he obtained a Fulbright Scholarship. His research focus es on Lepidotera taxonomy.