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smithsonian contributions to zoology • number 649

Sympatric Diversification 

in the Upper Amazon

A Revision of the Eumaeine 

Genus Paraspiculatus 

(Lepidoptera: Lycaenidae)

Robert C. Busby, Christophe Faynel, 

Alfred Moser, and Robert K. Robbins
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smithsonian contributions to zoology • numb e r 6 4 9

Sympatric Diversification 

in the Upper Amazon

A Revision of the Eumaeine 

Genus Paraspiculatus

(Lepidoptera: Lycaenidae)

Robert C. Busby, Christophe Faynel, 

Alfred Moser, and Robert K. Robbins

WASHINGTON, D.C.

2017
ABSTRACT

Busby, Robert C., Christophe Faynel, Alfred Moser, and Robert K. Robbins. Sympatric Diversification in the Upper 

Amazon: A Revision of the Eumaeine Genus Paraspiculatus (Lepidoptera: Lycaenidae). Smithsonian Contributions to 

Zoology, number 649, viii + 65 pages, 130 figures, 4 tables, 2017.—The Neotropical lycaenid hairstreak genus Paraspiculatus is unusual because of a high frequency of sympatry in the upper Amazon Basin coupled with negligible interspecific 

variation of male genitalic structures and absence of male secondary sexual traits. Male sexual structures are postulated to 

promote species recognition by females and to contribute to reproductive isolation, for which reason a high incidence of 

sympatry in Paraspiculatus would not be expected. 

A revision of Paraspiculatus was feasible because we increased the number of study specimens more than fivefold (by 

the extensive use of rotting fish as a bait for males) and sequenced the “barcode” part of the mitochondrial gene CO1 for 

almost all species. We recognize 19 Paraspiculatus species based on male wing patterns. We partition the 19 species into ten 

species complexes, which are monophyletic in all analyses of morphological characters, CO1 sequences, and a combined 

data set. Newly described are Paraspiculatus apuya Busby & Robbins, new species; Paraspiculatus cosmo Busby, Robbins 

& Faynel, new species; Paraspiculatus transvesta Robbins & Busby, new species; Paraspiculatus grande Busby, Robbins & 

Moser, new species; Paraspiculatus honor Busby, Robbins & Hall, new species; Paraspiculatus emma Busby & Robbins, 

new species; Paraspiculatus sine Busby & Robbins, new species; Paraspiculatus azul Busby, Robbins & Faynel, new species; Paraspiculatus lilyana Busby & Robbins, new species; and Paraspiculatus noemi Busby & Robbins, new species. All 

are described from Ecuador except for P. transvesta from Guatemala. 

Male Paraspiculatus are attracted to traps baited with rotting fish in eastern Ecuador, but elsewhere this attraction is less 

frequent or absent. Using the behavior of other eumaeines as context, we briefly discuss these observations with respect to 

Paraspiculatus biology.

Ten of 19 Paraspiculatus species are sympatric in the upper Amazon Basin below 1,250 m. Five of these sympatric 

species are from a single species complex. This instance of apparent in situ diversification is responsible for much of the 

sympatric diversity in Paraspiculatus in the upper Amazon Basin. 

KEYWORDS: Adult nutrition, CO1 barcoding, Eumaeus section, in situ diversification, male secondary sexual 

structures.

Cover images (from left): Paraspiculatus orobia; P. sine; P. elis (courtesy Rob Westerduijn); and Mithras nautes (courtesy 

Will and Gill Carter).

Published by SMITHSONIAN INSTITUTION SCHOLARLY PRESS

P.O. Box 37012, MRC 957, Washington, D.C. 20013-7012

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Library of Congress Cataloging-in-Publication Data

Names: Busby, Robert C. (Robert Carroll), 1948– author. | Smithsonian Institution Scholarly Press, publisher.

Title: Sympatric diversification in the Upper Amazon : a revision of the Eumaeine Genus Paraspiculatus (Lepidoptera: 

Lycaenidae) / Robert C. Busby [and three others].

Other titles: Smithsonian contributions to zoology ; no. 649. 0081-0282

Description: Washington, D.C. : Smithsonian Institution Scholarly Press, 2017. | Series: Smithsonian contributions to 

zoology, ISSN 0081-0282 ; number 649 | Includes bibliographical references.

Identifiers: LCCN 2017017880

Subjects: LCSH: Paraspiculatus—Amazon River Watershed. | Coexistence of species—Amazon River Watershed.

Classification: LCC QL561.L8 B87 2017 | DDC 333.95/5709811—dc23 | SUDOC SI 1.27:649

LC record available at https://lccn.loc.gov/2017017880

ISSN: 1943-6696 (online); 0081-0282 (print)

ZooBank registration: 3 August 2017 (LSID urn:lsid:zoobank.org:pub:D8F7819E-8121-4D07-AE95-8936A9DA0106) 

Publication date (online): 15 August 2017

Ó The paper used in this publication meets the minimum requirements of the American National Standard for Permanence of Paper for Printed Library Materials Z39.48–1992. 
Contents

LIST OF FIGURES v

LIST OF TABLES vii

INTRODUCTION 1

MATERIALS AND METHODS 2

Abbreviations for Collections 2

Other Acronyms and Abbreviations 3

NOMENCLATURAL LIST 5

OVERVIEW OF PARASPICULATUS 6

IDENTIFICATION KEY FOR MALES 25

SPECIES-LEVEL TAXONOMY 26

Paraspiculatus elis Species Complex 26

Paraspiculatus elis (Cramer, 1779) 26

Paraspiculatus catrea Species Complex 26

Paraspiculatus catrea (Hewitson, 1874) 29

Paraspiculatus vossoroca (Bálint and Moser, 2001) 29

Paraspiculatus oroanna Species Complex 29

Paraspiculatus oroanna (Bálint, 2002) 29

Paraspiculatus apuya Species Complex 31

Paraspiculatus apuya Busby and Robbins, new species 31

Paraspiculatus hannelore Species Complex 31

Paraspiculatus hannelore (Bálint and Moser, 2001) 31

Paraspiculatus orobia Species Complex 32

Paraspiculatus cosmo Busby, Robbins, and Faynel, new species 32

Paraspiculatus orobia (Hewitson, 1867) 33

Paraspiculatus orobiana Species Complex 35

Paraspiculatus orobiana (Hewitson, 1867) 35

Paraspiculatus transvesta Species Complex 36

Paraspiculatus transvesta Robbins and Busby, new species 36

Paraspiculatus orocana Species Complex 37

Paraspiculatus grande Busby, Robbins, and Moser, new species 37

Paraspiculatus honor Busby, Robbins, and Hall, new species 37

Paraspiculatus orocana (H. H. Druce, 1912) 38
i v • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Paraspiculatus colombiensis Species Complex 38

 Paraspiculatus emma Busby and Robbins, new species 39

 Paraspiculatus sine Busby and Robbins, new species 39

 Paraspiculatus colombiensis Johnson and Constantino, 1997 40

 Paraspiculatus azul Busby, Robbins, and Faynel, new species 43

 Paraspiculatus lilyana Busby and Robbins, new species 43

 Paraspiculatus noemi Busby and Robbins, new species 44

PHYLOGENETIC ANALYSES 45

DISCUSSION 45

Morphology, Molecules, and Female Choice 45

Documenting Diversity 50

Rotting Fish, Stratification, and Nutrition 50

Distribution 51

Sympatry and Diversification 51

ACKNOWLEDGMENTS 55

APPENDIX A: CO1 DNA SEQUENCE INFORMATION 57

APPENDIX B: PARASPICULATUS GENITALIC DISSECTIONS 61

REFERENCES 63
1–4. Adults in nature: 1. ♂ Paraspiculatus orobia; 2. ♂ P. sine; 

3. ♂ P. elis; 4. ♂ Mithras nautes 2

5. Male genitalia “spiculate pad” in lateral aspect at two 

magnifications: Mithras nautes; Paraspiculatus colombiensis 7

6–13. Paraspiculatus adults, dorsal on left: 6. ♂ P. elis; 7. ♀ P. elis; 

8. ♂ P. catrea; 9. ♀ P. catrea; 10. ♂ P. vossoroca; 11. 

♀ P. vossoroca; 12. ♂ P. oroanna; 13. ♂ P. apuya 8

14–21. Paraspiculatus adults, dorsal on left: 14. ♂ P. hannelore; 

15. ♀ P. hannelore; 16. ♂ P. cosmo; 17. ♀ P. cosmo; 

18. ♂ P. orobia; 19. ♀ P. orobia; 20. ♂ P. orobia; 

21. ♂ P. orobia 9

22–30. Paraspiculatus adults, dorsal on left: 22. ♂ P. orobiana; 

23. ♀ P. orobiana; 24. ♂ P. transvesta; 25. ♀ P. transvesta; 

26. ♂ P. grande; 27. ♀ P. grande; 28. ♂ P. honor; 

29. ♀ P. honor; 30. ♂ P. orocana 10

31–40. Paraspiculatus adults, dorsal on left: 31. ♂ P. emma; 

32. ♂ P. sine; 33. ♂ P. colombiensis; 34. ♀ P. colombiensis; 

35. ♂ P. azul; 36. ♀ P. azul; 37. ♂ P. Lilyana; 38. ♂ P. noemi; 

39. ♂ P. noemi; 40. ♂ P. noemi 11

41–52. Male dorsal forewing variation in Paraspiculatus: 41. P. elis; 

42. P. oroanna; 43. P. catrea; 44. P. orobiana; 45. P. orocana; 

46. P. grande; 47. P. orobia; 48. P. emma; 49. P. sine; 

50. P. colombiensis; 51. P. azul; 52. P. lilyana 12

53–64. Male ventral forewing variation in Paraspiculatus: 53. P. elis;

54. P. oroanna; 55. P. catrea; 56. P. orobiana; 57. P. orocana; 

58. P. grande; 59. P. orobia; 60. P. emma; 61. P. sine; 

62. P. colombiensis; 63. P. azul; 64. P. lilyana 13

65–67. Male ventral forewing iridescence as a function of light from 

different angles on same specimen (light source perpendicular 

to wings, left, and at acute angle): 65. P. orobia; 66. P. sine; 

67. P. colombiensis 14

68–70. Male ventral forewing iridescence as a function of light from 

different angles on same specimen (light source perpendicular 

to wings, left, and at acute angle): 68. P. orocana; 69. P. emma; 

70. P. orobiana 15

Figures
v i • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

71–76. Male dorsal hindwing variation at the costa: 71. P. lilyana; 

72. P. azul; 73. P. noemi; 74. P. grande; 75. P. orobia; 

76. P. cosmo 16

77. Male hindwing anal lobes (arrows): P. orocana (left); 

P. colombiensis (right) 16

78–80. Male genitalia of Paraspiculatus, genital capsule in ventral 

aspect (top) and penis in lateral aspect; posterior of butterfly 

on right: 78. P. elis; 79. P. catrea; 80. P. oroanna 18

81–83. Male genitalia of Paraspiculatus, genital capsule in ventral 

aspect (top) and penis in lateral aspect; posterior of butterfly 

on right: 81. P. apuya; 82. P. orobiana; 83. P. transvesta 19

84–86. Male genitalia of Paraspiculatus, genital capsule in ventral 

aspect (top) and penis in lateral aspect; posterior of butterfly 

on right: 84. P. hannelore; 85. P. cosmo; 86. P. orobia 20

87–89. Male genitalia of Paraspiculatus, genital capsule in ventral 

aspect (top) and penis in lateral aspect; posterior of butterfly 

on right: 87. P. grande; 88. P. honor; 89. P. orocana 21

90–92. Male genitalia of Paraspiculatus, genital capsule in ventral 

aspect (top) and penis in lateral aspect; posterior of butterfly 

on right: 90. P. emma; 91. P. sine; 92. P. colombiensis 22

93–95. Male genitalia of Paraspiculatus, genital capsule in ventral 

aspect (top) and penis in lateral aspect; posterior of butterfly 

on right: 93. P. azul; 94. P. lilyana; 95. P. noemi 23

96. Variation of valvae and saccus in ventral aspect of P. azul from 

Ecuador 24

97–99. Variation in the longitudinal spiculate pad in lateral aspect 

(arrows): 97. P. noemi; 98. P. sine; 99. P. apuya 25

100–112. Female genitalia ductus copulatrix in lateral aspect, posterior 

of butterfly on right: 100. P. elis; 101. P. catrea; 

102. P. vossoroca; 103. P. hannelore; 104. P. cosmo; 

105. P. orobia; 106. P. transvesta; 107. P. grande; 

108. P. honor; 109. P. colombiensis; 110. P. azul; 

111. P. noemi; 112. P. orobiana 27

113. Neighbor-joining phenogram of 111 Paraspiculatus (17 species) 

based on CO1 barcode sequences 28

114–117. Geographic distributions: 114. P. elis, P. vossoroca; 

115. P. oroanna, P. catrea; 116. P. apuya; 117. P. orobia, 

P. hannelore 30

118–121. Geographic distributions: 118. P. cosmo; 119. P. transvesta, 

P. orobiana; 120. P. grande, P. orocana; 121. P. honor, P. emma 34

122–125. Geographic distributions: 122. P. noemi, P. lilyana; 123. P. sine; 

124. P. colombiensis; 125. P. azul 41

126. Top of a most parsimonious tree for Paraspiculatus based on 

morphological characters 46

127. Bottom of a most parsimonious tree for Paraspiculatus based on 

morphological characters 47

128. Maximum likelihood tree using the molecular data set 48

129. Most parsimonious implied-weight (default K = 3) tree based 

on the combined morphological and molecular data set 49

130. Biogeographic regions and elevation zones for each 

Paraspiculatus species 52
Tables

1. Morphological characters for Paraspiculatus 4

2. Morphological character matrix for Paraspiculatus 5

Appendix tables

A.1. CO1 sequence samples listed alphabetically by genus, species, sex, 

and country 57

B.1. Genitalic dissections of Paraspiculatus 61

INTRODUCTION

Perhaps the most unusual characteristic of the neotropical genus Paraspiculatus

Johnson and Constantino, 1997 (Lycaenidae, Theclinae, Eumaeini, Figures 1–3) is that 

many of the species are sympatric in the upper Amazon Basin with little interspecific 

variation in traditional taxonomic characters, such as genitalic structures and male secondary sexual traits. Wing patterns also show a high degree of similarity. This situation 

contrasts sharply with most eumaeine genera. For example, in genera such as Oenomaus

Hübner, diversification is accompanied by marked evolution of genitalic morphology 

among sympatric species (Faynel et al., 2012). In others, such as Arcas Swainson and 

Lathecla Robbins, sympatric species have conspicuously different male secondary structures (Robbins et al., 2012; Robbins and Busby, 2015). And in others, such as Panthiades

Hübner and Thepytus Robbins, most species can be readily distinguished by elements of 

their wing patterns (Nicolay, 1976; Robbins, 2005; Robbins et al., 2010b). 

The apparent lack of conspicuous male secondary structures in Paraspiculatus is distinctive because these structures occur in more than 95% of eumaeine species (Robbins, 

2004a). Calycopidina Duarte and Robbins, Contrafacia Johnson, Symbiopsis Nicolay, 

Dicya Johnson, and Paraspiculatus are the major eumaeine taxa in which androconia 

are lacking on male wings. However, the vast majority of species belonging to the first 

three taxa have internal abdominal male secondary sexual structures called brush organs 

(Eliot, 1973; SEMs in Robbins, 1991; Duarte and Robbins, 2010), which are lacking 

without exception in Paraspiculatus. To the extent that male secondary structures are 

postulated to promote species recognition and to contribute to reproductive isolation 

among sympatric species (e.g., Löfstedt et al., 1991; Symonds and Elgar, 2008), the apparent high frequency of sympatry in Paraspiculatus is unexpected. 

Little interspecific variation in genitalic structures, such as in Paraspiculatus, is not 

as unusual in the Eumaeini as the lack of male secondary structures. For example, Arcas

species cannot be distinguished by their genitalia (Nicolay, 1976), but no two species 

share identical male secondary structures (Robbins et al., 2012). Interspecific genitalic differences among animals are hypothesized to result from sexual selection, which 

then facilitates sympatry between sister species following secondary contact (e.g., Lande, 

1981; Hosken and Stockley, 2004; Eberhard, 2010; Simmons, 2014). Again, a low 

Sympatric Diversification in the Upper 

Amazon: A Revision of the Eumaeine Genus

Paraspiculatus (Lepidoptera: Lycaenidae)

Robert C. Busby,1 Christophe Faynel,2 Alfred Moser,3

and Robert K. Robbins4

*

1 7 Countryside Way, Andover, Massachusetts 

01810-6041, USA.

2 16 rue des Aspres, F-34160 Montaud, France.

3 Av. W. Rotermund, 1045 93030-000 São Leopoldo, RS Brazil.

4 Department of Entomology, P.O. Box 37012, 

MRC 105, Smithsonian Institution, Washington, 

D.C., 20013-7012, USA.

*Correspondence: robbinsr@si.edu

Received 4 May 2016; accepted 8 February 

2017.
2 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

incidence of sympatry in Paraspiculatus would be predicted. For 

example, in a four-species lineage of Electrostrymon Clench with 

uniform genitalia and without male secondary sexual traits on 

the wings—a situation similar to that in Paraspiculatus—sympatry was absent except for minor elevational overlap (Thompson and Robbins, 2016). 

The lack of variation in traditional taxonomic characters in 

Paraspiculatus and paucity of specimens in museum collections 

have been limiting factors in documenting diversity within the 

genus. With the exception of a few species with distinctive ventral wing patterns, such as P. elis (Cramer, 1779) and P. catrea

(Hewitson, 1874), the identification and taxonomic treatment of 

described species has been chronically incorrect, as noted in the 

species accounts below. Also, wing pattern similarity in females 

has precluded accurate association with males. 

A revision of Paraspiculatus is feasible for the first time 

because of two factors. First, we increased the study material 

fivefold. At the beginning of this study, there were fewer than 

150 specimens, including types, in major North American and 

European museums (listed below). Using traps baited with rotting fish, we added more than 450 specimens to the study series. 

Additional material was found in public museum collections in 

Latin America and in private collections. The sample size increase to 742 individuals allowed a markedly improved analysis 

of wing pattern variation. Second, we successfully sequenced the 

DNA “barcode region” of the mitochondrial gene CO1 for all 

but two Paraspiculatus species. These sequences were used primarily to associate the sexes—a nearly intractable problem for 

many species previously—and secondarily as another character 

set for distinguishing species. 

The first purpose of this paper is to propose a species-level 

classification for Paraspiculatus. Variation of wing pattern characters—albeit characters that are sometimes subtle and difficult 

to assess in worn individuals—and of CO1 DNA sequences is 

largely congruent, which provides the evidence for recognizing 

19 species, of which 10 are new. We associate the sexes, when 

the female is known, and provide a key to identify males by their 

wing pattern. We also summarize information on distribution, 

habitat, and behavior. 

The second purpose is to establish monophyletic species 

complexes within Paraspiculatus. Although phylogenetic analyses yielded a variety of trees, a number of clades were monophyletic in all results based on morphology, molecules, and both data 

sets. Recognition of these clades as species complexes provides a 

more fine-grained classification and allows us to characterize the 

composition of the set of sympatric species in the upper Amazon.

MATERIALS AND METHODS

The species-level taxonomy of Paraspiculatus is based primarily on an analysis of morphological variation among 742 

pinned specimens from the following collections (most abbreviations from Evenhuis, 2015). We also examined more than a 

hundred other specimens with redundant data. 

Abbreviations for Collections

AMNH American Museum of Natural History, New York, 

New York, USA

BMNH The Natural History Museum [formerly British Museum (Natural History)], London, UKCF Private 

collection of Christophe Faynel, Montpellier, France

CMNH Carnegie Museum of Natural History, Pittsburgh, 

Pennsylvania, USA

DZUP Museu de Entomología Pe. Jesus Santiago Moure, 

Universidade Federal do Paraná, Curitiba, Paraná, 

Brazil

LACM Los Angeles County Museum of Natural History, 

Los Angeles, California, USA

FIGURES 1–4. Adults in nature. 1. ♂ Paraspiculatus orobia feeding on a leaf coated with drops of liquid containing rotting fish, Apuya, Napo, 

Ecuador. 2. ♂ P. sine feeding on rotting fish, San Isidro, Morona Santiago, Ecuador. 3. ♂ P. elis feeding on dilute cow blood, Puerto Almendra, 

Loreto, Peru (image courtesy of Rob Westerduijn). 4. ♂ Mithras nautes on a leaf, Alta Floresta, Mato Grosso, Brazil (image courtesy of Gill 

and Will Carter). The latter two species have a similar, conspicuous transverse orange-yellow band on the hindwing. The ground color of P. 

orobia appears darker than that of P. sine because of the angle at which the image was taken, the lighting, and the worn condition of the latter 

individual.
NUMBER 649 • 3

JFLC Private collection of Jean-François Le Crom, Bogotá, 

Colombia 

JHKW Jason P. W. Hall and Keith R. Willmott collection, 

Washington, D.C., USA

MC Private collection of Alfred Moser, São Leopoldo, 

Rio Grande do Sul, Brazil

MECN Museo Ecuatoriano de Ciencias Naturales, Quito, 

Ecuador

MGCL McGuire Center for Lepidoptera and Biodiversity, 

Florida Museum of Natural History, University of 

Florida, Gainesville, Florida, USA

MNHN Muséum National d’Histoire Naturelle, Paris, France

MUSA Museo de Historia Natural de la Universidad Nacional San Agustín, Lima, Peru

MUSM Museo de Historia Natural, Universidad Nacional 

Mayor de San Marcos, Lima, Peru

RCB Private collection of Robert C. Busby, Andover, Massachusetts, USA 

SMF Forschungsinstitut und Naturmuseum Senckenberg, 

Frankfurt-am-Main, Germany 

USNM National Museum of Natural History, Smithsonian 

Institution, Washington, D.C., USA

ZMBH Museum für Naturkunde der Humboldt-Universität, 

Berlin, Germany

Other Acronyms and Abbreviations

BIN barcode index number

BOLD Barcode of Life Data System

DFW dorsal forewing

DHW dorsal hindwing

HW hindwing

ICZN International Commission on Zoological Nomenclature

SEM scanning electron microscope

VFW ventral forewing

VHW ventral hindwing

To supplement and confirm the morphological taxonomy, we 

used DNA barcode sequences belonging to the CO1 gene. Legs 

from 138 adults of 19 Paraspiculatus species in our study material 

were prepared and sequenced according to the protocol in Wilson 

(2012). Sequences longer than 200 base pairs were successfully extracted from 111 samples representing 17 Paraspiculatus species. 

The samples are listed in Appendix A with their museum vouchers 

and BOLD process numbers (Ratnasingham and Hebert, 2007). 

We visualized DNA sequence differences phenetically using the 

neighbor-joining methods on the BOLD website (Ratnasingham 

and Hebert, 2007), along with the recently introduced barcode 

index numbers (Ratnasingham and Hebert, 2013). 

On the basis of morphological variation and phenetic DNA 

sequence differences, we recognize nine Paraspiculatus species 

that were previously described plus 10 new taxa, which are described according to the rules of the ICZN (1999). We associated 

the sexes in some species, such as P. elis, by similarity of ventral 

wing pattern and distribution, and in others, such as P. orobia 

(Hewitson, 1867), by similarity of mitochondrial CO1 DNA 

sequences. 

Standard references for morphological terminology are 

Comstock (1918) for insect wing veins; Klots (1970) and Robbins 

(1991) for Lepidoptera genitalia as modified for the Eumaeini; 

Robbins et al. (2012) for male secondary sexual structures; and 

Snodgrass (1935) for all other structures. Genitalic dissections 

are listed in Appendix B. Adult size was measured from the base 

of the forewing to its apex. For new species, size is reported by 

the mean, standard deviation, and sample size. To test whether 

male P. colombiensis Johnson and Constantino, 1997 are smaller 

than male P. noemi Busby and Robbins, we did a t-test with 

unequal variances in commercially available Microsoft Excel 

software and checked the calculations online using VassarStats 

(http://vassarstats.net/index.html; accessed 10 February 2016). 

Labels on holotypes are recorded verbatim, with square 

brackets used for information not explicitly noted on the labels 

and for descriptions of the labels. Lines on holotype labels are 

separated by a forward slash (/). Otherwise, months are abbreviated by their first three letters in English. Localities and other 

data, especially within the Material Examined and Type Material 

sections, were transcribed as presented on specimen labels, with 

the result of minor inconsistencies of wording and format within 

those sections.

Nomenclaturally relevant information for Paraspiculatus

and for each species is summarized in a nomenclatural list. Citations for original descriptions can be found in Lamas (2015). 

Besides taxonomic history and morphological variation, 

species accounts include information on distribution, elevation, 

biogeography, habitat, and behavior. Maps are used to visualize distributions but, for practical reasons, unrelated species are 

sometimes placed on the same map. Biogeographic zones follow 

Brown (1982), who partitioned the forested nonmontane continental neotropics into the Transandean region, Amazon region, 

and Atlantic region. These general biogeographic regions were 

derived, in part, from vertebrate distributions (e.g., Ab’Sáber, 

1977) and have proven useful for documenting the biogeography 

of forest butterflies (e.g., Willmott, 2003; Robbins et al., 2010a). 

Although more fine-grained biogeographical regions have been 

proposed, such as those in Morrone (2014) and Chazot et al. 

(2016), they include nonforested regions, such as cerrado, where 

Paraspiculatus are not known to occur. Further, the geographic 

data for many Paraspiculatus species are insufficient to examine 

these more finely partitioned biogeographic regions. Elevation 

zones are slightly modified from Elias et al. (2009) into lowland 

species (0–1,250 m), lower montane species (500–1,700 m), and 

montane species (>1,200 m). If adults were attracted to rotting 

fish, we note whether they were attracted to traps, which were 

hung 7 to 20 m above the ground, or to leaves near the ground. 

Many male eumaeines perch in “territories” at certain times of 

day to wait for receptive females to fly through and “defend” 

these areas by flying at other males that enter the territory (e.g., 
4 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Powell, 1968; Alcock and O’Neill, 1987; Cordero et al., 2000; 

Faynel, 2003, 2006a). Recorded times from our fieldwork for 

“territorial” behavior are standard time at that locality. 

We analyzed morphological characters and CO1 barcode 

sequences to identify monophyletic lineages within Paraspiculatus. Mithras nautes (Cramer, 1779) and Theorema eumenia

(Hewitson, 1865) were used as outgroups for reasons given 

below under “Taxonomic History.” 

We coded 26 morphological characters for each Paraspiculatus species, Mithras nautes, and Theorema eumenia, and 

refer to them in the species accounts by their character number 

(Table 1). Most characters refer to male wing pattern because, 

as noted, there is little variation in genitalic morphology, and 

male secondary sexual structures are lacking. The resulting matrix (Table 2) was 26 characters by 21 species (which include two 

outgroups).

We used 118 CO1 DNA barcode sequences longer than 600 

base pairs in length from our study series (downloaded from the 

BOLD database). Of these, 105 were specimens of Paraspiculatus belonging to 17 species (data were lacking for P. transvesta

Robbins and Busby and P. vossoroca Bálint and Moser, 2001); 

about 6 specimens per species. The other downloaded sequences 

belonged to the two outgroup species. We had identified each of 

the 118 adults from which DNA was extracted. We used Mesquite software to delete empty positions and corrected three base 

pairs that had been shifted to empty positions in three specimens. 

TABLE 1. Morphological characters for Paraspiculatus.

Character

number Character and states

1. Forewing shape at vein M2 (0) continuously rounded, (1) angled.

2. Male DFW basal edge dark apex extends to the distal side of discal cell (0) no, (1) yes.

3. Male DFW inner edge of black border between M3 and 2A (0) concave, (1) straight except for black scales in the middle of cell Cu2-2A 

extending basally, (2) straight without black scales in the middle of cell Cu2-2A extending basally, (3) border is a marginal dark line.

4. Male DFW blue scales between the end of discal cell and the apex (0) present, (1) absent.

5. Male DFW anterior half of discal cell entirely brown-black (0) absent, (1) present.

6. Male VFW scales along costal margin (0) same as ventral wing color, (1) light brown/tan (some separation from base color), (2) white.

7. Male VFW scales posterior of cubital vein (0) shining blue, (1) iridescent in flat plane (silvery blue), (2) iridescent only at an acute angle 

(brownish).

8. Male VFW iridescent scales, anterior of cubital vein (0) iridescent silver-blue, (1) brown, (2) iridescent shining blue.

9. Male VFW scales in limbal area just posterior of vein Cu2 (0) a shade of brown, (1) bluish gray, (2) shining blue.

10. Male VFW with scattered white scales next to the distal parts of vein 2A (0) absent, (1) present.

11. Male HW shape (angle measured from end of vein Cu1 to the anal lobe to the midpoint of inner margin) (0) angle greater than 90°, (1) 

angle 90°, (2) angle less than 90°.

12. Male HW shape (0) rounded apex, (1) less rounded, more angular at vein M1.

13. Male DHW basal side of cell Sc+R1-Rs (0) blue (same size/shape as other blue wing scales), (1) black/purple (same size/shape as blue 

wing scales), (2) gray/brown (slightly rounded tips, evenly spaced), (3) dark gray, iridescent edges (clustered in center of cell).

14. Male DHW blue scales in cell Rs-M1 (0) restricted, usually limited to basal and posterior sides, (1) extensive, fills most (75%) of cell.

15. Male DHW anal lobe shape (0) small, with rounded inner margin, (1) large, with rounded inner margin, (2) elongated like a teardrop, 

(3) with a long, “straight” inner margin.

16. Male DHW with distal greenish iridescent scales (0) absent, (1) present.

17. Male VHW with scattered blue/white scales (0) absent, (1) present.

18. Male VHW with postmedian line in cell Sc+R1-Rs displaced to the basal half of the cell (0) absent, (1) present.

19. Male VHW with transverse orange ray through the disco-cellular veins (0) absent, (1) present.

20. Male VHW postmedian line intersects vein 2A (0) closer to wing base than anal lobe, (1) much closer to anal lobe than to wing base.

21. Male VHW band of blue/green scales between the posterior part of the postmedian line and the anal angle (0) absent, (1) present.

22. Male VHW anal angle with white marginal scales at the end of 2A and at the inner margin at the edge of the anal lobe area (0) absent, 

(1) present.

23. Male genitalia with longitudinal spiculate pad (0) absent, (1) present.

24. Male genitalia with semi-hemispherical brush organs (0) present, (1) absent.

25. Density of spines on the inner surface of the corpus bursae (0) not greater in the vicinity of the origin of the ductus seminalis, (1) greater 

in the vicinity of the origin of the ductus seminalis.

26. Female ductus bursae with a narrow, lightly sclerotized medial “neck” (0) absent, (1) present.
NUMBER 649 • 5

The resulting molecular matrix was made up of 654 base pairs 

for 118 specimens belonging to 17 Paraspiculatus and two outgroup species. 

A third matrix combined the morphological and molecular 

data. We modified the molecular nexus file by adding P. transvesta and P. vossoroca, which were coded with question marks 

(?) for each molecular position. We then added the morphological data from the morphology matrix for each specimen. The 

resulting combined matrix had 680 characters (the first 654 were 

molecular sequences) for 120 specimens belonging to 19 Paraspiculatus and two outgroup species. 

The morphological matrix was analyzed with equal-weight 

maximum parsimony using traditional search (5 random seeds, 

100 replications, 1,000 trees saved per replication, collapse trees 

after search) in TNT software (Goloboff et al., 2008). To assess 

the assumption of equally weighted characters, implied weighting was performed over a range of values for the parameter K 

(3, 10, 100, 1,000). Standard bootstrap support values were also 

calculated in TNT. Strict consensus trees and mapping of character changes using unambiguous changes were done in WinClada 

software (Nixon, 2002).

The molecular matrix was analyzed with both Garli 

(Zwickl, 2006) and TNT. We ran PartitionFinder (Lanfear et al., 

2012) and determined that TrN+I, F81, and TrN+G were the 

best models for first, second, and third codon positions, respectively. The best maximum likelihood tree was derived in Garli 

using 100 search repetitions. We then analyzed the molecular 

matrix with equal-weight and implied-weight (K = 3, 10, 100, 

1,000) maximum parsimony using traditional search in TNT. 

 The combined morphology and molecular matrix was analyzed with equal-weight and implied-weight (K = 3, 10, 100, 

1,000) maximum parsimony using traditional search in TNT. 

Standard bootstrap support values were also calculated in TNT. 

NOMENCLATURAL LIST

This list summarizes relevant nomenclatural information for 

Paraspiculatus.

Paraspiculatus Johnson and Constantino, 1997

Type species: Paraspiculatus colombiensis Johnson and Constantino.

Paraspiculatus elis species complex

Paraspiculatus elis (Cramer, 1779) (Papilio), type locality: 

Suriname. [There is a possible syntype in the museum in 

Leiden.]

TABLE 2. Morphological character matrix for Paraspiculatus with Theorema and Mithras as outgroups.

Character number

Taxon 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26

T. eumenia 0 0 0 0 0 0 1 2 2 0 0 1 1 1 ? 1 0 0 0 0 1 0 0 0 0 0

M. nautes 0 0 3 0 0 0 0 2 2 0 0 0 0 0 1 0 0 0 1 0 1 1 0 0 0 0

P. elis 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 0 1 1 0 1 1 1 1 0 0

P. catrea 1 0 2 0 0 2 2 1 0 1 2 1 0 1 2 0 1 1 0 0 0 1 0/1 1 1 1

P. vossoroca 1 0 2 0 0 2 ? 1 0 1 2 0 0 0 2 0 0 1 0 0 0 ? 1 1 1 1

P. oroanna 1 0 2 0 0 2 1 1 0 1 1 0 0 0 1 0 0 1 0 0 0 1 1 1 ? ?

P. apuya 0 1 0 1 1 1 1 0 0 0 0 0 0 0 0 0 0 1 0 1 0 1 1 1 ? ?

P. hannelore 0 0 0 0 0 2 1 0 0 0 1 1 0 0 0 0 0 1 0 1 0 1 1 1 1 1

P. cosmo 0 0 0 0 0 2 1 0 0 0 1 0 2 0 0 0 0 1 0 1 0 1 1 1 1 1

P. orobia 0 0 0 0 0 2 1 2 0 0 1 0 2 0 0 0 0 1 0 1 0 1 1 1 1 1

P. orobiana 0 1 1 1 0 1 2 1 0 0 1 0 0 0 0 0 0 1 0 1 0 1 1 1 1 1

P. transvesta 0 ? ? ? ? 2 2 1 0 0 0 0 ? ? 0 0 ? 1 0 1 0 1 1 1 1 1

P. grande 0 1 1 1 0 2 1 0 0 0 1 0 3 0 3 0 0 1 0 1 0 0 1 1 1 1

P. honor 0 1 1 1 0 2 1 0 0 0 1 0 3 0 3 0 0 1 0 1 0 0 1 1 1 1

P. orocana 0 1 1 0 0 2 1 0 0 0 1 0 3 0 3 0 0 1 0 1 0 0 1 1 ? ?

P. emma 0 0 0 0 0 2 1 0 1 0 0 0 0 0 0 0 0 1 0 1 0 1 1 1 ? ?

P. sine 0 0 0 0 0 2 1 0 1 1 0 0 0 1 0 0 0 1 0 1 0 1 1 1 ? ?

P. colombiensis 0 0 0 0 0 2 1 0 1 0 0 0 0 1 0 0 0 1 0 1 0 1 1 1 1 1

P. azul 0 0 0 0 0 1 1 0 1 0 0 0 0 1 0 0 0 1 0 1 0 1 1 1 1 1

P. lilyana 0 0 0 0 0 1 1 2 1 0 0 0 0 1 0 0 0 1 0 1 0 1 1 1 ? ?

P. noemi 0 0 0 0 0 2 1 0 1 0 0 0 1 1 0 0 0 1 0 1 0 1 1 1 1 1
6 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Paraspiculatus catrea species complex

Paraspiculatus catrea (Hewitson, 1874) (Thecla), type locality: Brazil. [Syntypes in BMNH. D’Abrera (1995) called 

one of the syntype males a holotype, but under ICZN Article 74.5, this action cannot be construed as a lectotype 

designation.] 

Paraspiculatus vossoroca Bálint and Moser, 2001, type locality: Brazil, Santa Catarina, Joinville. [Holotype in DZUP.]

Paraspiculatus oroanna species complex

Paraspiculatus oroanna Bálint, 2002, type locality: Peru, 

Huánuco, Cushi. [Holotype in BMNH.]

Paraspiculatus apuya species complex

Paraspiculatus apuya Busby and Robbins, new species, type 

locality: Ecuador, Napo, 14 km Tena–Puyo Road, 1°06.7′S, 

77°46.9′W. [Holotype in USNM.]

Paraspiculatus hannelore species complex

Paraspiculatus hannelore Bálint and Moser, 2001, type locality: Brazil, Rio Grande do Sul, Dois Irmãos, Picada Veraão. 

[Holotype in DZUP.]

Paraspiculatus orobia species complex

Paraspiculatus cosmo Busby, Robbins, and Faynel, new species, type locality: Ecuador, Morona Santiago, Bosque de 

Domono, 2°11.0′S, 78°06.2′W. [Holotype in USNM.]

= cosmophila (Tessman, 1928) (Thecla), type locality: Peru, 

unavail., ICZN Art. 45.5.1.

= cosmophila (Bridges, 1988) (Thecla), type locality: Peru, unavail., ICZN Art. 45.5.1.

Paraspiculatus orobia (Hewitson, 1867) (Thecla), type locality: Brazil, Amazon. [Lectotype male unintentionally designated by Johnson and Constantino (1997) in BMNH.]

= villaanna Bálint, 2004, type locality: Colombia, Villavicencio 

(misspelled in original description). [Holotype in MNHN.]

Paraspiculatus orobiana species complex

Paraspiculatus orobiana (Hewitson, 1867) (Thecla), type locality: Brazil, Amazon, Ega (AM). [Lectotype male unintentionally designated by Johnson and Constantino (1997) in 

BMNH.]

Paraspiculatus transvesta species complex</i>

Paraspiculatus transvesta Robbins and Busby, new species, 

type locality: Guatemala, Cayuga. [Holotype in USNM.]

Paraspiculatus orocana species complex

Paraspiculatus grande Busby, Robbins, and Moser, new species, type locality: Ecuador, Esmeraldas, 12 km Lita–San 

Lorenzo Road, 0°53.1′N, 78°30.9′W. [Holotype in USNM.]

Paraspiculatus honor Busby, Robbins, and Hall, new species, 

type locality: Ecuador, Pichincha, Mindo, Río Napombillo. 

[Holotype in USNM.]

Paraspiculatus orocana (H. H. Druce, 1912) (Thecla), type 

locality: Peru, El Povenir. [Lectotype male unintentionally 

designated by Johnson and Constantino (1997) in BMNH.]

Paraspiculatus colombiensis species complex

Paraspiculatus emma Busby and Robbins, new species, type 

locality: Ecuador, Napo, 14 km Tena–Puyo Road, 1°06.7′S, 

77°46.9′W. [Holotype in USNM.]

Paraspiculatus sine Busby and Robbins, new species, type locality: Ecuador, Morona Santiago, 15 km S of Gualaquiza, 

3°27.6′S, 78°33.1′W. [Holotype in USNM.]

Paraspiculatus colombiensis Johnson and Constantino, 1997, 

type locality: Colombia, Rio Ortequaza. [Holotype in 

AMNH.]

Paraspiculatus azul Busby, Robbins, and Faynel, new species, type locality: Ecuador, Napo, 14 km Tena–Puyo Road, 

1°06.7′S, 77°46.9′W. [Holotype in USNM.]

Paraspiculatus lilyana Busby and Robbins, new species, type 

locality: Ecuador, Napo, 14 km Tena–Puyo Road, 1°06.7′S, 

77°46.9′W. [Holotype in USNM.]

Paraspiculatus noemi Busby and Robbins, new species, type 

locality: Ecuador, Pichincha, 20 km Pacto–Guayabillas 

Road, 0°11.6′N, 78°51.5′W. [Holotype in USNM.]

OVERVIEW OF PARASPICULATUS

Taxonomic History. Johnson and Constantino 

(1997) characterized Paraspiculatus—and derived the name—

from a “spiculate pad” on the posterior lateral edge of the fused 

vinculum/tegumen of the male genitalia. They applied the name 

to species that had been previously placed in the Thecla orobia

species group (Draudt, 1919–1920). Bálint and Moser (2001) 

reported that the spiculate pad was lacking in Paraspiculatus catrea and suggested as an alternative that “the degree of sclerotization and the shape of the dorsal posterior pouch beneath the 

ductus seminalis” of the female genitalia was a synapomorphy. 

Robbins (2004a,b) synonymized Paraspiculatus with the 

monotypic Mithras Hübner for two reasons. First, both genera have nonsocketed teeth on the posterior lateral edge of the 

vinculum. Second, Paraspiculatus elis possesses a ventral wing 

pattern that is very similar to that of Mithras nautes (Figures 

3, 4), as noted by Draudt (1919–1920), but has the male genitalic structures of the Thecla orobia species group. However, 

this synonymy was provisional because structure of the spiculate 

pad is different in Mithras and Paraspiculatus (Figure 5). Further, whereas Mithras nautes possesses semi-hemispherical brush 

organs that surround the male genitalia—a conspicuous trait 

shared with Eumaeus Hübner and Theorema Hewitson (Robbins, 2004a)—Paraspiculatus lacks brush organs. 

The molecular phylogenetic analysis of Quental (2008) included two species of the Thecla orobia species group, but not 

the type species of Mithras. However, the Thecla orobia species 

group did not group with Eumaeus and Theorema, although 
NUMBER 649 • 7

they were closely related in some analyses. Rather, Chalybs lineata (Hewitson) was the weakly supported sister to the Thecla 

orobia species group. However, because Chalybs lineata shares 

no morphological similarity with Paraspiculatus, the molecular 

data need to be confirmed.

Recognition of Paraspiculatus. Given the uncertainty in the phylogenetic relations of Paraspiculatus and relatives, we treat Paraspiculatus as a distinct genus characterized by 

the spiculate pad, as first proposed by Johnson and Constantino 

(1997). It is unique within the Eumaeini, so far as we are aware, 

and is a synapomorphy in the phylogenetic analyses in this paper. 

Bálint and Moser (2001) proposed a synapomorphy in the female genitalia, but we could discern no substantive basis for this 

proposal. We did code two characters of the female genitalia, but 

neither provides a synapomorphy for Paraspiculatus in phylogenetic analyses. 

Wing Pattern and Shape. Wing pattern in

Paraspiculatus is characteristically homogeneous with a few exceptions (Figures 6–40). Apart from P. transvesta Robbins and 

Busby, the dorsal wings of most males have bright blue scaling 

with a black border whereas those of most females have less blue 

scaling, oftentimes none. The ventral wings of both sexes are 

dark brown with a postmedian line composed of shining blue 

dots and dashes. On the forewing, these spots form a disjointed 

line from the costa to vein Cu2

. On the hindwing the spots are 

usually arranged in a semicircle. The anteriormost hindwing 

spot (cell Sc+R1

-Rs) is displaced basally. The blue spots are surrounded by a variable number of black scales, which enhance the 

contrast of the bright blue with the dark wing ground color. Distal of the hindwing postmedian line, there are scattered green or 

gold scales near the anal angle. A submarginal band on the hindwing is barely visible to absent because of the dark ground color. 

FIGURE 5. Male genitalia “spiculate pad” in lateral aspect (arrows) and enlarged (bottom) for Mithras nautes (left) and Paraspiculatus colombiensis (right).
8 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

However, it is present in P. hannelore Bálint and Moser, 2001 

and sometimes in other species such as P. orobiana (Hewitson, 

1867) and P. azul Busby, Robbins, and Faynel. Most individuals 

of Paraspiculatus can be recognized by these characteristic wing 

pattern traits, but individuals belonging to other genera sometimes possess some or all of these traits. 

Paraspiculatus wing shape is variable. Forewings are more 

elongate in some species, such as P. elis, than in others, such as P. 

catrea. Hindwing shape varies from round (P. apuya Busby and 

Robbins) to angular (P. vossoroca). Hindwing tails may be present in both sexes (P. elis), in females only (P. catrea), or absent 

in both sexes (P. vossoroca). In P. orobia and P. noemi, presence and length of hindwing tails vary geographically. Examples 

of other eumaeines in which tail length and number vary geographically are Arawacus jada (Hewitson) and Atlides halesus

(Cramer) (Godman and Salvin, 1887–1901; Clench, 1942). 

Variation in Male Wing Pattern. Scaling on 

the wings of male Paraspiculatus is the primary source of morphological variation, which is used for both identification and 

inference of relationships. Here we summarize important aspects 

of this variation and provide figures to illustrate these characters (Figures 41–77). As a matter of terminology, we differentiate 

bright shining blue (Figure 65 left side—scales anterior of the 

cubital vein) from iridescent blue or silver-blue (Figure 65 right 

side—scales posterior of the cubital vein). Bright shining blue is 

bright blue regardless of the angle at which it is viewed. 

FIGURES 6–13. Adults of Paraspiculatus, dorsal on left. 6. ♂ P. elis (Ecuador). 7. ♀ P. elis (Peru). 8. ♂ P. catrea (Brazil). 9. ♀ P. catrea (Brazil). 

10. ♂ P. vossoroca (Brazil). 11. ♀ P. vossoroca (Brazil). 12. ♂ P. oroanna (Ecuador). 13. ♂ P. apuya (Ecuador). Scale 1 cm (lower left).
NUMBER 649 • 9

The two wing characters we found to be most useful for 

distinguishing species groups are as follows:

(1) Scales on the basal half of the dorsal hindwing anterior of 

vein Rs are (a) blue (black in one species) and are similar 

to surrounding scales in size and shape (Figures 71–73), (b) 

dark gray with iridescent edges, concentrated in the center 

of the cell giving that area a sheen (Figure 74), or (c) brown/

gray with slightly rounded tips, evenly spaced throughout 

the cell (Figures 75, 76). 

(2) The limbal area of the ventral forewing on the posterior side 

of vein Cu2 is either dull bluish gray (Figures 60–64) or a 

shade of brown (Figures 53–59).

Other wing characters that are important for species identification include: 

(1) Scales posterior of the ventral forewing cubital vein have an 

iridescent sheen when viewed at an angle. Scales anterior of 

the ventral forewing cubital vein may be bright shining blue 

FIGURES 14–21. Adults of Paraspiculatus, dorsal on left. 14. ♂ P. hannelore (Brazil). 15. ♀ P. hannelore (Brazil). 16. ♂ P. cosmo (Ecuador). 

17. ♀ P. cosmo (Ecuador). 18. ♂ P. orobia (Brazil). 19. ♀ P. orobia (French Guiana). 20. ♂ P. orobia (Ecuador). 21. ♂ P. orobia (Peru). Scale 

1 cm (lower left).
1 0 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURES 22–30. Adults of Paraspiculatus, dorsal on left. 22. ♂ P. orobiana (Brazil). 23. ♀ P. orobiana (Peru). 24. ♂ P. transvesta (Guatemala). 

25. ♀ P. transvesta (Mexico). 26. ♂ P. grande (Ecuador). 27. ♀ P. grande (Panama). 28. ♂ P. honor (Ecuador). 29. ♀ P. honor (Ecuador). 30. ♂

P. orocana (Ecuador). Scale 1 cm (lower left).
NUMBER 649 • 1 1

FIGURES 31–40. Adults of Paraspiculatus, dorsal on left. 31. ♂ P. emma (Ecuador). 32. ♂ P. sine (Ecuador). 33. ♂ P. colombiensis (Ecuador). 

34. ♀ P. colombiensis (Ecuador). 35. ♂ P. azul (Ecuador). 36. ♀ P. azul (Peru). 37. ♂ P. lilyana (Ecuador). 38. ♂ P. noemi (Ecuador). 39. ♂ P. 

noemi (Mexico). 40. ♂ P. noemi (Costa Rica). Scale 1 cm (lower left).
1 2 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURES 41–52. Male dorsal forewing variation in Paraspiculatus. See text for character states. 41. P. elis. 42. P. oroanna. 43. P. catrea. 

44. P. orobiana. 45. P. orocana. 46. P. grande. 47. P. orobia. 48. P. emma. 49. P. sine. 50. P. colombiensis. 51. P. azul. 52. P. lilyana.
NUMBER 649 • 1 3

FIGURES 53–64. Male ventral forewing variation in Paraspiculatus. See text for character states. 53. P. elis. 54. P. oroanna. 55. P. catrea.

56. P. orobiana. 57. P. orocana. 58. P. grande. 59. P. orobia. 60. P. emma. 61. P. sine. 62. P. colombiensis. 63. P. azul. 64. P. lilyana.
1 4 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURES 65–67. Male ventral forewing iridescence as a function of light from different angles. Light 

source perpendicular to the wings (left) and at an acute angle (same specimen on right). 65. P. orobia. 66. 

P. sine. 67. P. colombiensis.
NUMBER 649 • 1 5

FIGURES 68–70. Male ventral forewing iridescence as a function of light from different angles. Light source 

perpendicular to the wings (left) and at an acute angle (same specimen on right). 68. P. orocana. 69. P. emma. 

70. P. orobiana.
1 6 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURES 71–76. Male dorsal hindwing variation at the costa. 71. P. lilyana. 72. P. azul. 73. P. noemi. 74. P. grande. 75. P. orobia. 76. P. cosmo. 

See text for explanation.

FIGURE 77. Male hindwing anal lobes (arrows). P. orocana (left). P. colombiensis (right).
NUMBER 649 • 1 7

(Figure 65) or iridescent (Figures 66–70). The iridescence 

varies from blue or silver-blue (Figures 66–69) to brownish 

bronze (Figure 70). The latter is somewhat difficult to see 

and generally requires the wing to be viewed at a very acute 

angle (45°) under bright light. 

(2) The postmedian line on the ventral forewing is composed of 

three blue/white dots near the costa with four more blue spots 

extending to cell Cu1

-Cu2 in the majority of species (Figures 

54, 55, 58–63). In others, the anterior dots are absent (Figures 53, 56, 57, 64). In some instances, the postmedian line is 

reduced to one or two posterior blue dots (Figure 57).

(3) Apical white scales on the ventral forewing may be present 

(Figures 54, 55, 57–62) or absent (Figures 53, 56, 63, 64). 

When present (despite intraspecific variation in the number 

of white scales), the white scales may form a wide (Figures 

54, 55) or narrow band (Figure 61), may be crescent shaped 

(Figure 57), or may be divided in two, with the distal patch 

at the apex (Figures 58–60, 62). 

(4) Color of blue scales on the dorsal surface is variable (Figures 

41–52). Further, the blue color in the limbal area of the dorsal 

forewing in “fresh” individuals varies from a “smooth” appearance with few interspersed black scales (Figures 42–46, 

51, 52) to a “grainy” appearance with equivalent numbers 

of black scales and blue scales (Figures 48–50). In a long series of specimens, differences among species are conspicuous. 

However, these differences “overlap” and may be difficult 

to discern in worn specimens, so color and texture tend to 

be less reliable traits for identification of a single individual.

Some conspicuous wing markings were not useful in distinguishing species in the large study samples that we accumulated. 

For example, black scales at the end of the forewing discal cell 

(Figures 45, 48, 52) do not provide useful information because 

of intraspecific variability. 

Finally, the brown scales that make up most of the ventral 

surface often reflect different colors when rotated in sunlight. 

For example, at about 45 degrees, the matte brown areas in the 

distal forewing of Paraspiculatus emma Busby and Robbins are 

bright bronze; at a more acute angle, they are greenish. We have 

not quantified this aspect of the ventral surface, but mention this 

trait because it could contribute to species recognition among 

females of sympatric species.

Male Genitalic Morphology. Ventrolateral 

processes of the fused vinculum and tegumen, which are typical 

of many Eumaeini, were first mentioned by McDunnough (1942: 

1), who called them “finger-like projection(s) at the base of the 

tegumen.” Nonsocketed “teeth” are found on these projections 

in a variety of presumably unrelated eumaeines in addition to 

Paraspiculatus and Mithras (McDunnough, 1942; Duarte and 

Robbins, 2010). However, the slightly projecting lateral edge of 

the fused vinculum/tegumen with nonsocketed teeth in Paraspiculatus is unique within the Eumaeini. The projection with lateral 

teeth in Mithras is considerably longer than in Paraspiculatus

and is immediately distinguishable (Figure 5). 

Other than the nonsocketed teeth on the ventrolateral edge 

of the fused vinculum/tegumen, the male genitalia of Paraspiculatus are not especially distinctive. Compared to other eumaeines, 

the gnathos are relatively stout with a conspicuous apiculus, but 

not markedly so. The vinculum strut is horizontal where the 

spiculate pad occurs, but does not distinguish Paraspiculatus

from other eumaeines. We found no structures in the male genitalia that provide evidence on the relationship of Paraspiculatus

with other genera. 

Interspecific variation of the male genitalia of Paraspiculatus is slight (Figures 78–95) and does not unambiguously distinguish any species other than P. oroanna Bálint, 2002. Shape of 

the valvae and saccus in ventral aspect is intraspecifically variable (Figure 96). The valvae of some species have the ventral 

surface at the base without setae and displaced ventrally, but the 

expression of this structure is highly variable intraspecifically. 

The penis has a terminal slender cornutus in all species. There 

is also a second minute terminal cornutus in all species except 

Paraspiculatus oroanna, P. hannelore, and P. cosmo Busby, Robbins, and Faynel. 

Expression of the distinctive nonsocketed teeth on the ventrolateral edge of the fused vinculum/tegumen also varies intraspecifically. For example, in one dissection of P. catrea the teeth 

are well expressed (Figure 79), but in two other dissections, they 

are absent (also noted in Bálint and Moser, 2001). As another 

example, in one dissection of P. noemi the teeth are well expressed (Figure 97), but in another dissection, the number of 

teeth is greatly reduced: they are almost entirely absent posteriorly. Otherwise these genitalia are indistinguishable. Other 

examples in which the location and size of the spiculate pad 

vary in P. sine Busby and Robbins and P. apuya are illustrated 

(Figures 98, 99). 

Female Genitalic Morphology. Females of 

six Paraspiculatus species are unknown (P. oroanna, P. apuya, 

P. orocana (Druce, 1912), P. emma, P. sine, and P. lilyana). The 

remaining species (Figures 100–112) share the following traits: 

(1) the signa on the corpus bursae are vestigial or absent; (2) the 

ductus seminalis attaches to the posterior corpus bursae considerably dorsal of the point where the ductus bursae attaches; and 

(3) in lateral aspect, the posterior ductus bursae is wider than the 

anterior ductus bursae.

The most distinctive female genitalia belong to Paraspiculatus. elis (Figure 100). It lacks the narrow, lightly sclerotized 

“neck” that occurs in the middle of the ductus bursae of the 

other species. It lacks a pouch on the posterior corpus bursae 

from which the ductus seminalis arises. With respect to these 

two traits, the female genitalia of P. elis are similar to those of 

Mithras nautes, as are aspects of its wing pattern, as already 

noted.

Interspecific variation of the female genitalia of Paraspiculatus, other than those of P. elis, is minimal (Figures 101–112). 

Length of the anterior and posterior parts of the ductus bursae varies quantitatively, but because of overlapping lengths, 

does not usually distinguish species. The posterior part of the 
1 8 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURES 78–80. Male genitalia of Paraspiculatus, genital capsule in ventral aspect (top) and penis in lateral aspect. Posterior 

of butterfly to the right. Scale 1 mm. 78. P. elis. 79. P. catrea. 80. P. oroanna.
NUMBER 649 • 1 9

FIGURES 81–83. Male genitalia of Paraspiculatus, genital capsule in ventral aspect (top) and penis in lateral aspect. Posterior of 

butterfly to the right. Scale 1 mm. 81. P. apuya. 82. P. orobiana. 83. P. transvesta.
2 0 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURES 84–86. Male genitalia of Paraspiculatus, genital capsule in ventral aspect (top) and penis in lateral aspect. Posterior of butterfly to the right. Scale 1 mm. 84. P. hannelore. 85. P. cosmo. 86. P. orobia.
NUMBER 649 • 2 1

FIGURES 87–89. Male genitalia of Paraspiculatus, genital capsule in ventral aspect (top) and penis in lateral aspect. 

Posterior of butterfly to the right. Scale 1 mm. 87. P. grande. 88. P. honor. 89. P. orocana.
2 2 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURES 90–92. Male genitalia of Paraspiculatus, genital capsule in ventral aspect (top) and penis in lateral aspect. Posterior of butterfly to the right. Scale 1 mm. 90. P. emma. 91. P. sine. 92. P. colombiensis.
NUMBER 649 • 2 3

FIGURES 93–95. Male genitalia of Paraspiculatus, genital capsule in ventral aspect (top) and penis in lateral aspect. Posterior of 

butterfly to the right. Scale 1 mm. 93. P. azul. 94. P. lilyana. 95. P. noemi.
2 4 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

corpus bursae varies from membranous to lightly sclerotized, 

but again, this trait does not distinguish species. The suggestion 

that morphology of the posterior corpus bursae represents a synapomorphy for Paraspiculatus (Bálint and Moser, 2001) has no 

substantive basis that we can discern.

Habitat. Lowland and montane forest. 

Caterpillar Food Plants. None recorded.

Behavior. Male Paraspiculatus in the upper Amazon 

Basin are attracted to traps baited with rotting fish and placed 

7–20 m above the ground. Occasionally, male Paraspiculatus fly 

to liquefied fish poured on leaves near the ground. Female Paraspiculatus are almost never attracted to fish. In over 25 years of 

field work, there are only six records of females being attracted 

to traps, and all occurred at a single locality. In other geographic 

regions, such as the western Andean slopes of Ecuador, collecting 

with rotting fish has been less successful. We elaborate on these 

observations in the Discussion.

Adults of Paraspiculatus are rarely encountered in the forest, so available information on adult behaviors is minimal. For 

example, male territoriality has been recorded for only P. oroanna and P. grande Busby, Robbins, and Moser. 

Identification. We recognize nine described and 

10 undescribed Paraspiculatus species based on morphology. 

The following key allows males to be identified by wing pattern. Each of these species also clusters in a CO1 DNA sequence 

neighbor-joining phenogram (Figure 113), so DNA barcodes 

provide an alternate means of identification. However, the morphological species determinations for males agreed with BINs 

FIGURE 96. Variation of valvae and saccus in ventral aspect of P. azul from Ecuador.
NUMBER 649 • 2 5

in 92 of 95 males (96.8% concordance). The three extra BINs 

referred to geographic variants of P. apuya, P. orobiana, and P. 

noemi (blue-green highlight in Figure 113). Each of the 16 females with sequenced DNA barcodes clustered with one of the 

male species (yellow highlight in Figure 113). 

Geographic Variation. We detail geographic 

variation in the morphology of P. orobia and P. noemi in 

the species accounts. As noted, there is geographic variation 

in CO1 sequences in P. orobiana, P. noemi, and P. apuya

(Figure 113). 

IDENTIFICATION KEY FOR MALES

1. With a transverse orange/yellow ray on VHW [Figure 3] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. elis

Without a transverse orange/yellow ray on VHW . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2. With an elongated anal lobe (teardrop shaped) on HW [Figures 8, 10] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

Without an elongated anal lobe (rounded) on HW . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

3. With a solid blue/white median line on VHW and scattered white markings . . . . . . . . . . . . . . . . . . . . . . . . . . . P. catrea

With a series of blue dots in the median of the VHW and no white markings . . . . . . . . . . . . . . . . . . . . . . . P. vossoroca

4. With a predominately blue dorsal surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

With a predominately brown dorsal surface (some dull blue/green scales basally placed) . . . . . . . . . . . . . . P. transvesta

5. With dark gray or gray/brown scales at the base of the DHW costa above vein Rs

 [Figures 74–76] . . . . . . . . . . . . . . . . 6

With regular blue (or black) scales at the base of the DHW costa [Figures 71–73] . . . . . . . . . . . . . . . . . . . . . . . . . . . 10

6. With tightly bunched dark gray scales in the center of cell Sc+R1

-Rs [Figure 74] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

With evenly spaced gray/brown scales; not concentrated in the center of cell Sc+R1

-Rs [Figures 75,76] . . . . . . . . . . . . 9

7. With three white/blue spots at median of VFW costal margin and four blue spots extending posteriorly to cell 

Cu1

-Cu2 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. grande

With a single blue spot (occasionally two) in the middle of the VFW and either no white spots or vestigial spots at the 

costal margin [Figure 57] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

8. With DFW black apex extending to the distal side of the discal cell . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. honor

With DFW black apex not extending to the discal cell, blue scales fill approximately 1/3 of the space between the discal 

cell and apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. orocana

9. With a narrow black border on the DFW and a broad band of shiny blue scales [Figure 59] anterior of the cubital vein 

on the VFW . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. orobia

With a black border of medium width on the DFW and no band of shiny blue scales on the VFW . . . . . . . . . . P. cosmo

10. With blue/silver-blue iridescence posterior of the cubital vein on the VFW [Figures 65–69] . . . . . . . . . . . . . . . . . . . . 11

With brownish-bronze iridescence posterior of the cubital vein [Figure 70] and with broad black outer margin on 

DFW . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. orobiana

FIGURES 97–99. Variation in the longitudinal spiculate pad in lateral aspect (arrows). 97. P. noemi (top Ecuador, bottom Mexico). 98. P. sine

(both Ecuador). 99. P. apuya (top Ecuador, bottom Rondonia, Brazil).
2 6 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

11. With dull bluish-gray scales in the limbal area of the VFW posterior to vein Cu2 [Figures 60–64] . . . . . . . . . . . . . . . 12

With brown or brownish-gray scales in the limbal area of the VFW posterior to vein Cu2

 [Figures 53–59] . . . . . . . . 17

12. With no white scales near VFW costa between median and apex [Figures 63, 64] . . . . . . . . . . . . . . . . . . . . . . . . . . . 13

With white scales near VFW costa between median and apex [Figure 60–62] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14

13. With a band of shiny blue scales above the cubital vein on VFW and with a crisply delineated black border on the 

DFW . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. lilyana

Without a band of shiny blue scales on the VFW . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. azul

14. With white scales along the costa of the VFW, which are broken into two patches—one near the median and the second 

in the apex [Figures 60, 62] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15

With a continuous, narrow band of white scales along the costa of the VFW between the median and apex [Figure 61] 

and no HW tails . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. sine

15. With mostly blue scales on the basal side of cell Sc+R1

-Rs on the DHW [Figures 71, 72] . . . . . . . . . . . . . . . . . . . . . . . 16

With mostly black (occasional purple) scales in cell Sc+R1

-Rs on the DHW [Figure 73]. . . . . . . . . . . . . . . . . . . P. noemi

16. With a narrow black outer border on the DFW and HW tails . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. colombiensis

With a black outer border of medium thickness and no HW tails . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. emma

17. With very broad black outer DFW border [Figures 12, 13] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18

With a black outer DFW border which narrows below vein Cu1

 [Figure 14] and with brown scales at the end of the discal 

cell on VHW . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. hannelore

18. With no white scales near the costa of the VFW between the median and apex and with black scales covering the anterior 

portion of the DFW discal cell . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. apuya

With a very broad band of white scales in the VFW apex between the costa and vein M3 . . . . . . . . . . . . . . . P. oroanna

SPECIES-LEVEL TAXONOMY

Paraspiculatus elis Species Complex

Paraspiculatus elis (Cramer, 1779)

FIGURES 3, 6, 7, 41, 53, 78, 100, 113, 114, 126–130

Updated Diagnosis. The lowland Amazonian 

Paraspiculatus elis and Mithras nautes have traditionally been 

treated as close relatives because of the similarity of their ventral wing patterns (Figures 3, 4), especially a single transverse 

orange-yellow band on the hindwing of both sexes (Draudt, 

1919–1920). This similarity is one reason why Paraspiculatus

was treated as a synonym of Mithras (Robbins, 2004b). The 

male genitalia of P. elis (Figure 78) and the lack of evident male 

secondary sexual traits, however, are the same as those of Paraspiculatus, which is the reason for its generic placement.

Female. (Figure 7) The female is associated by similarities with the male in ventral wing pattern, geographic distribution, and CO1 sequence.

Genitalia. (Figures 78, 100) The presence of a 

spiculate pad in a longitudinal plane in the male genitalia is the 

reason why this species is placed in Paraspiculatus. It is the only 

member of the genus that can be immediately distinguished by 

its female genitalia because it lacks a semimembranous “neck” 

in the ductus bursae. 

Distribution. (Figure 114) Widely distributed in 

the Amazon region of Brown (1982).

Habitat. Paraspiculatus elis inhabits wet forest up to 

900 m elevation.

Behavior. Males of P. elis have been recorded on 

traps and plants baited with rotting fish or dilute cow blood 

(Figure 3). 

Remarks. This species is one of the largest Paraspiculatus and has a conspicuously recognizable ventral wing pattern, but is exceedingly rare. There are fewer than 20 specimens 

in major museum collections. 

Material Examined (15♂, 3♀). ECUADOR, 

Morona Santiago, 53.1 km Santiago–Puerto Morona Rd, 

2°55.0′S, 77°42.7′W, 210 m, 17 Sep 2016, 1♂ (RCB); 3.2 km 

Mendez–Guarumales Rd, 2°41.9′S, 78°19.7′W, 550 m, 29 

Sep 2004, 1♀ (RCB); Napo, 4 km S Tena–Pano Rd, 1°00.8′S, 

77°50.1′W, 580 m, 5 Nov 1992, 1♂ (USNM); Puerto Napo, 

650 m, 20 Sep 1987, 1♂ (USNM); Río Pimpilala, 1°04.6′S, 

77°56.2′W, 600–900 m, Jun 2014, 1♂ (RCB); Sep 1996, 2♂

(JHKW), 1♀ (JHKW); 10 Sep 2006, 1♂ (RCB); Orellana, Río 

Añangu, middle Río Napo, 0°31.7′S, 76°23.7′W, 250 m, Oct–

Dec 2005, 1♂ (JHKW); 22 km Loreto–San Jose de Payamino Rd, 

0°34.9′S, 77°24.4′W, 825 m, 08 Nov 2010, 1♂ (RCB); Pastaza, 

39.4 km Puyo–Villano Rd, 1°25.6′S, 77°43.8′W, 750 m, 01 

Oct 2015, 2♂ (RCB). PERU, San Martín, Tarapoto, 1886, 1♂

(BMNH); 1♀ (BMNH); Juanjui, 1♂ (USNM); Huánuco, Tingo 

Maria, 800 m, 20 Jun 1982, 1♂ (USNM); Loreto, San Pablo, 

4°01′S, 71°06′W, 100 m, Feb 2011, 1♂ (CF).

Paraspiculatus catrea Species Complex

This lineage of two sympatric species is characterized by the 

shape of the hindwing anal angle (Table 1: Character 11) and 
NUMBER 649 • 2 7

FIGURES 100–112. Female genitalia ductus copulatrix in lateral aspect, posterior of butterfly to the right. Scale 1 mm. 100. P. elis. 

101. P. catrea. 102. P. vossoroca. 103. P. hannelore. 104. P. cosmo. 105. P. orobia. 106. P. transvesta. 107. P. grande. 108. P. honor. 

109. P. colombiensis. 110. P. azul. 111. P. noemi. 112. P. orobiana. Scale 1 mm.

NUMBER 649 • 2 9

lobe (Character 15) as well as brownish iridescence below the 

cubital vein on the ventral forewing (Character 7). 

Paraspiculatus catrea (Hewitson, 1874)

FIGURES 8, 9, 43, 55, 79, 101, 113, 115, 126–130

Updated Diagnosis and Description. This 

species has been readily identifiable for more than a century by 

its distinctive ventral wing pattern and hindwing shape (Figure 

8). The ventral characters noted by Hewitson include “a band of 

two or three linear white spots near the middle” of the forewing, 

“five or six lunular white spots” at the middle of the hindwing, 

and interspersed white scales on both wings. The white lines and 

spots are unique within Paraspiculatus (Figures 6–40). 

Bálint and Moser (2001) transferred Thecla catrea to Paraspiculatus based on female genitalic structures. The presence of a 

spiculate pad in the male genitalia (Figure 79), the morphological similarity of the female genitalia to those of other Paraspiculatus species (Figures 100–112), and the similarity of CO1 DNA 

sequences (Figure 113) place this species in Paraspiculatus. 

Female. (Figure 9) The female is associated by similarities with the male in ventral wing pattern, geographic distribution, and CO1 DNA sequences.

Genitalia. (Figures 79, 101) The spiculate pad of 

the male genitalia may be present or absent, as already noted.

Distribution. (Figure 115) Southeastern Brazil 

from the state of Rio de Janeiro to Rio Grande do Sul. 

Habitat. Paraspiculatus catrea inhabits forest from 

100 m elevation in the southern part of its range to 1,600 m 

elevation in the northern parts. It is the only lowland Paraspiculatus species that also occurs above 1,250 m elevation. 

Remarks. In all the phylogenetic analyses in this 

paper, P. catrea is placed within Paraspiculatus. 

Material Examined (8♂, 7♀). BRAZIL, Santa 

Catarina, São Bento do Sul, Rio Vermelho, 700 m, 23 Feb 1987, 

1♂ (USNM); 24 Feb 1973, 1♀ (USNM); 23–25 Apr 2009, 1♀

(MC); 900 m, 16 Feb 1993, 1♀ (MC); Praia Grande, Serra do 

Faxinal, 700 m, 20 Nov 1994, 1♀ (MC); São Bento do Sul, 

Serra do Rio Natal, 700 m, 23 Apr 2009, 1♀ (MC); Nova Teutonia 300–500 m, 26 Jan 1954, 1♂ (USNM); 30 Dec 1949, 

1♂ (USNM); Paraná, Ponta Grossa, Dec 1942, 1♀ (USNM); 

Rio Grande do Sul, Ivoti, 27 Dec 1998, 1♂ (MC); São Francisco de Paula, 900 m, 30 Mar 2002, 1♀ (MC); 3 Jun 1991, 

1♂ (MC); Morro Reuter, Jammerhal, 300 m, 24 Sep 1994, 1♂

(MC); Morro Reuter, Faz. Padre Eterno, 500 m, 24 Dec 1996, 

1♂ (MC); Guarani, 7 Jan 1973, 1♂ (USNM).

Paraspiculatus vossoroca

(Bálint and Moser, 2001)

FIGURES 10, 11, 102, 114, 126, 129, 130

Updated Diagnosis and Description. This 

species and P. catrea are immediately recognizable by their hindwing shape with an elongated anal angle (Figures 8–11). The 

ventral wing pattern of P. vossoroca differs markedly from that 

of P. catrea and is typical of the majority of Paraspiculatus species with a dark ventral surface with bright blue spots and line 

segments (Figures 10, 11). Male and female P. vossoroca lack 

tails whereas the female of P. catrea is tailed.

Female (Figure 11). The female is associated with 

the male by similarity in hindwing shape, ventral wing pattern 

(especially the white scales near the costa of the forewing), and 

geographic distribution. 

Genitalia (Figure 102). Bálint and Moser (2001) 

published a photograph of the male genitalia that does not clearly 

show a spiculate pad. We examined the male holotype genitalia, 

which are almost indistinguishable from those of P. catrea (Figure 79), and they possess a spiculate pad. The female genitalia 

are morphologically similar to those of other Paraspiculatus species (Figures 100–112). 

Distribution (Figure 114). Southeastern Brazil in 

the states of Paraná and Santa Catarina. 

Habitat. Paraspiculatus vossoroca inhabits wet Atlantic coastal forest from 200 to 900 m elevation. 

Remarks. The type locality, Vossoroca, is approximately at the midpoint on the road between Curitiba (Paraná) 

and Joinville (Santa Catarina). Despite decades of collection 

(there are literally thousands of Theclinae in museum collections 

from Joinville, Curitiba, and the area in between), only three 

individuals of P. vossoroca have been found. A fourth, very old, 

specimen has uncertain provenance. This species has not been 

collected, so far as we are aware, in almost 40 years. 

Material Examined (1♂, 2♀). BRAZIL, Santa 

Catarina, Joinville, 200 m, Jan 1969, 1♂ holotype (DZUP); 10–

200 m, 22 Feb 1977, 1♀ (USNM on loan from DZUP), Paraná, 

Tijucas do Sul, Vossoroca, 900 m, 4–13 Jan 1976, 1♀ (MC).

Paraspiculatus oroanna Species Complex

Paraspiculatus oroanna (Bálint, 2002)

FIGURES 12, 42, 54, 80, 113, 115, 126–130

Updated Diagnosis and Description. The 

male Paraspiculatus oroanna has a ventral wing pattern that is 

FIGURE 113. (Opposite page) Neighbor-joining phenogram of 111 Paraspiculatus (17 species) based on CO1 barcode sequences. Yellow 

highlights the 16 females. Pale blue-green highlights the three specimens with divergent BINs (see text).
3 0 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURES 114–117. Geographic distributions. 114. P. elis (stars), P. vossoroca (diamonds). 115. P. oroanna (flowers), P. catrea

(stars). 116. P. apuya (diamonds). 117. P. orobia (squares), P. hannelore (hearts).
NUMBER 649 • 3 1

typical of the genus, with dark brown ground color and bright 

blue spots and line segments (Figure 12). However, the extent 

of the white scales on the forewing apex is unique. These scales 

cover the distal half of the forewing from the costa to vein M3

. 

Some species, such as P. sine, have a smaller band of white scales 

extending from the costa to vein M1

. The forewing shape of P. 

oroanna is also more angular at vein M2

 than in most other Paraspiculatus species. Finally, the hindwing anal lobes in P. oroanna

are larger than those in the majority of species in the genus. 

In the original diagnosis, Bálint differentiated P. oroanna

from P. orobiana based on the “absence of the gleaming marking 

in the cell 3A+2A of the hind wing anal part on ventral side”. 

Iridescent (gleaming) scales in cell 2A-3A occur between the 

postmedian line and anal lobe in most Paraspiculatus, including 

P. oroanna and P. orobiana. The color of these markings varies from bright blue, in the case of P. oroanna and P. grande, 

to green-blue in P. orobiana and other Paraspiculatus species. 

Further, scale color can vary intraspecifically, and in worn specimens, scales are sometimes lost. 

Female. The female of P. oroanna is unknown.

Genitalia. (Figure 80) We have examined the genitalia of only one male, but the long narrow male valvae appear 

to distinguish them from all other Paraspiculatus. The penis 

lacks a second cornutus. 

Distribution. (Figure 115) Eastern slope of the 

Andes in Ecuador and Peru. 

Habitat. Paraspiculatus oroanna inhabits montane 

forest from 1,600 to 2,000 m elevation. It might be sympatric 

with P. orocana at 1,600 m, but it otherwise occurs at higher 

elevations than any other Paraspiculatus. 

Behavior. There are no records of this species being 

attracted to rotting fish. One male displayed territorial behavior 

at 11:30 hours, perching about 1 m above the ground (voucher 

in RCB). 

Material Examined (11♂). ECUADOR, Tungurahua, 1800–2200 m, Oct 2008, 2♂ (CF); Río Machay, 

1°23.9′S, 78°17.0′W, 1600–1800 m, 15 Feb 2010, 1♂ (RCB); 

28 Mar 2014, 1♂ (RCB). PERU, Huánuco, 1,500–2,000 m, Jul 

2003, 1♂ (CF); Jul 2006, 1♂ (CF); Sep 2010, 2♂ (CF); Chinchao, 

Chaupiyunca, 1,800 m, Dec 2003, 1♂ (CF); Cushi, 1900 m, 1♂

holotype (BMNH); Junín, Satipo, Nov 2009, 1♂ (CF).

Paraspiculatus apuya Species Complex

Paraspiculatus apuya Busby and Robbins, 

new species

FIGURES 13, 81, 99, 116, 126–130

Diagnosis and Description. Males of P. apuya

are distinguished from other Paraspiculatus by the dark dorsal 

forewing (Figure 13). This species has a broad black border on 

the dorsal forewing and is the only Paraspiculatus where the 

black scaling along the dorsal forewing costa covers the anterior portion of the discal cell (Character 5). The pattern on the 

ventral forewing is unusual. The anterior portion of the postmedian line is made up of three white/light-blue spots, but there 

are no additional white scales along the costa (Character 6). The 

combination of white spots and no white scales is found in only 

one other Paraspiculatus species, P. azul, and this species is easily 

distinguished from P. apuya by its bright blue dorsal surface with 

relatively narrow black borders. Mean male forewing length is 

16.6 mm (SD = 0.63, n = 8). We illustrate the wing pattern (Figure 13) and genitalia (Figure 81).

Female. Females of P. apuya are unknown.

Genitalia. (Figures 81, 99) Variation in the male 

genitalia spiculate pad are illustrated.

Type material.

Holotype (♂): (Figure 13) ECUADOR: Napo/ 14 km 

Tena–Puyo Road/ 1°06.7′S, 77°46.9′W 600 m/ 23 October 2010 

(Apuya)/ I. Aldas, R. C. Busby, leg. [rectangular, white, printed], 

USNM ENT 00180817 [rectangular, white, printed barcode 

label], GENITALIA No./ 2013: 71 ♂/ C. FAYNEL [rectangular, 

green, printed], Holotype ♂/ Paraspiculatus apuya / Busby & 

Robbins [rectangular, red, printed]. Deposited in USNM.

Paratypes (3♂): ECUADOR, Napo, 14 km Tena–Puyo 

Road, 1°06.7′S, 77°46.9′W, 600 m, 06 Sep 2009, 1♂ (MECN); 

20 km Tena–Puyo Road [=14 km], 27–30 Aug 1993, 1♂

(JHKW); Morona Santiago, 54.6 km Santiago–Puerto Morona 

Rd., 2°54.7′S, 77°42.4′W, 205 m, July 2014, 1♂ (RCB).

Other Material Examined (4♂): ECUADOR, Napo, Río 

Napo, 1♂ (BMNH). PERU, Loreto, Balsapuerto, Paranapura 

River, Feb 1933, 1♂ (AMNH); Loreto, Agua Blanca, 03°56′S, 

73°28′W, 10 Jan 2004, 1♂ (MUSM). BRAZIL, Rôndonia, vic. 

Caucaulândia, 10°32′S, 62°48′W, 160–350 m, 16 Oct 1991, 1♂

(USNM). 

Etymology. This species is named for its type locality, Apuya (Napo, Ecuador). Six Paraspiculatus species have been 

found at Apuya, and a seventh (P. elis) is likely to occur there. 

The name is proposed as a nonlatinized noun in apposition. 

Distribution (Figure 116) Upper Amazon Basin in 

Ecuador, Peru, and Brazil.

Habitat. Paraspiculatus apuya inhabits wet forest 

from 160 to 600 m elevation. 

Behavior. Males are attracted to traps baited with 

rotting fish. 

Remarks. The CO1 sequence of the male P. apuya

from Rondônia, Brazil (Appendix A) clusters with those from eastern Ecuador, but is slightly divergent (Figure 113). The Brazilian 

male is worn, but we found no evident differences in wing pattern. 

Paraspiculatus hannelore Species Complex

Paraspiculatus hannelore

(Bálint and Moser, 2001)

FIGURES 14, 15, 84, 103, 113, 117, 126–130

Updated Diagnosis and Description. Paraspiculatus hannelore possesses a dark brown submarginal band 
3 2 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

and a short dark line at the distal end of the discal cell on the ventral surface of the hindwing. Although these wing pattern elements 

occur occasionally in other Paraspiculatus species, they are considerably more conspicuous in P. hannelore because the postdiscal 

area of the hindwing is a lighter brown color. Also, the ventral 

forewing subapical and postdiscal areas are a somewhat lighter 

brown, as noted in the original description. One diagnostic character used by Bálint and Moser (2001) to separate P. hannelore

from P. orobiana was a “subapical white spot” on the ventral 

hindwing of the latter, but there is no subapical “white spot” in 

our P. orobiana study series, which included the holotype.

Female. (Figure 15). The female is associated by 

similarities with the male in ventral wing pattern (Figure 14), 

geographic distribution, and CO1 DNA sequences.

Genitalia. (Figures 84, 103). The genitalic structures of P. hannelore are indistinguishable from those of P. orobia

(see Remarks), except that the minute second terminal cornutus 

in the male genitalia appears to be absent in P. hannelore. 

Distribution. (Figure 117). Southeastern Brazil in 

the states of Rio Grande do Sul and Santa Catarina. 

Habitat. Paraspiculatus hannelore inhabits forest 

along the Atlantic coast from 200 to 900 m elevation and occurs 

as far inland as Nova Teutonia (SC). 

Remarks. Paraspiculatus hannelore was synonymized with P. colombiensis (Robbins, 2004b), but phylogenetic 

analyses suggest that this synonymy was incorrect. 

Material Examined (10♂, 4♀). BRAZIL, Rio 

Grande do Sul, Dois Irmãos, Picada Verão, 350 m, 22 Mar 1992, 

1♂ holotype (DZUP); Ivoti, 200 m, 28 Dec 1994, 1♂ (MC); 

Nova Petropolis, 750 m, 16 Jan 2006, 2♂ (MC); São Francisco 

de Paula, 900 m, 1 May 1993, 1♂ (MC); Santa Catarina, Joinville, 200 m, 20 Jan 1992, 1♀ (MC); Joinville, Serrinha, 200 

m, 20 Apr 2016, 1♂ (MC); São Bento do Sul, Serra Rio Natal, 

600 m, 14 Jan 2001, 1♀ (MC); 20 Apr 2000, 1♂ (MC); 14 May 

2001, 1♀ (MC); 04 Jul 2005, 1♂ (MC); Nova Teutonia, 27°11′S, 

52°23′W, 300–500 m, 24 Feb 1954, 1♂ (USNM); Apr 1960, 1♀

(USNM); Seara, Nova Teutonia, 450–700 m, May 1977, 1♂

(USNM). One male in BMNH is labeled Mexico with no further 

data, but the known distribution of this species strongly suggests 

that the label is incorrect.

Paraspiculatus orobia Species Complex

Paraspiculatus orobia and P. cosmo have parapatric distributions and are sisters in our phylogenetic analyses. Morphologically, males in this lineage have regularly spaced gray/brown 

scales (Figures 75, 76) at the base of the male dorsal hindwing 

costa (Character 13). 

Paraspiculatus cosmo Busby, Robbins, 

and Faynel, new species

FIGURES 16, 17, 76, 85, 104, 113, 118, 126–130

Diagnosis and Description. Males of Paraspiculatus cosmo (Figure 16) and P. orobia (Figures 18, 20) differ 

by (1) the lack of a band of blue scales above the cubital vein on 

the ventral forewing (Character 8, present in P. orobia), and (2) 

a moderately broad black border on the dorsal forewing (narrow 

in P. orobia). These differences are consistent in the study series. 

Dark gray scales at the base of the male dorsal hindwing 

costa in P. cosmo also occur in P. grande, P. orocana, and P. 

honor. However, the scales in these other species are concentrated in the center of cell Sc+R1

-Rs and often have an iridescent 

sheen (Figure 74) that is less apparent in P. cosmo (Figure 76). 

Further, male hindwing tails occur in P. cosmo, but are absent in 

the other three species. 

Mean forewing length is 18.8 mm (SD = 1.35, n = 25) in 

males and 16.7 mm (SD = 0.57, n = 2) in females. We illustrate 

the wing pattern (Figures 16, 17) and genitalia (Figures 85, 104).

Genitalia. (Figures 85, 104) As in P. hannelore, the 

second terminal cornutus appears to be absent. 

Female. (Figure 17) The female is associated by geographic distribution and CO1 DNA sequences. It has blue scaling dorsally. 

The female type of the unavailable name Thecla orobiana cosmophila Tessmann also appears to be this species. This 

“type” is blue above, unlike the all-brown female of P. orobiana, 

and has conspicuous tails, unlike the females of P. grande and 

P. honor. The female of P. orocana is as yet unknown, but we 

expect it to be blue above and tailless.

Type material.

Holotype (♂): (Figure 16) ECUADOR: MoronaSantiago/ Bosque de Domono/ 2°11.0′S, 78°06.2′W 1325 m/ 

03 September 2003/ Robert C. Busby, leg. [rectangular, white, 

printed], USNM ENT 00180800 [rectangular, white, printed 

barcode label], GENITALIA No./ 2013: 79 ♂/ C. FAYNEL [rectangular, green, printed], Holotype ♂/ Paraspiculatus cosmo/ 

Busby, Robbins & Faynel [rectangular, red, printed]. Deposited 

in USNM. 

Paratypes (29♂, 2♀): ECUADOR, Morona Santiago, 

14 km W of Macas, Río Abanico, [2°15.4′S, 78°11.7′W], 1,600 

m, 08 Sep 1999, 1♂ (RCB); 09 Sep 1999, 1♂ (USNM); 15 Sep 

2006, 1♂ (RCB); 19 Sep 1999, 3♂ (USNM); 21 Sep 1998, 1♂

(RCB); 23 Sep 1998, 1♂ (RCB); 25 Sep 1998, 1♂ (RCB); 27 Sep 

1998, 1♂ (MECN); 28 Sep 1998, 1♀ (RCB); 30 Sep 1997, 1♂

(RCB); Oct 1995, 1♂ (JHKW); 1 km E of Río Abanico, [13 km 

W of Macas], 1,600 m, 11 Sep 2000, 1♂ (MECN), 1♂ (RCB); 

13 Sep 2001, 2♂ (USNM); 15 km S San Juan Bosco, 1,600 m, 25 

Sep 2001, 1♂ (RCB); 2 km N of San Isidro, 2°11.9′S, 78°09.4′W, 

1,250–1,400 m, 17 Sep 2013, 1♂ (RCB); 30 Sep 2012, 2♂

(USNM); 29 Sep 2015, 2♂ (RCB); Nueve de Octubre, 2°13.0′S, 

78°13.5′W, 1,600–1,800 m, 28 Jan 2014, 1♂ (MECN); Zamora 

Chinchipe, Zamora, 4°04.5′S, 78°58.1′W, 1,450 m, 06 Oct 

2002, 1♂ (RCB), 2♂ (USNM); Napo, 49 km Tena–Loreto Road, 

0°42.9′S, 77°44.4′W, 1,350 m, 29 Nov 2013, 1♂ (RCB); Sucumbíos, km 12 La Bonita–Rosa Florida old trail, 1,550 m, Mar 

1995 (JHKW). PERU, Huánuco, Tingo Maria, 01 Oct 2003, 1♂

(CF); Chanchamayo, La Merced, Jun 1930, 1♀ (USNM).

Other Material Examined (3♂, 1♀): PERU, Amazonas, 

Mendoza, Yanohuaico [Llanohuaico], 1,500–2,000 m, Jan 2007, 
NUMBER 649 • 3 3

1♂ (MC); Rodríguez de Mendoza, [6°25′S, 77°30′W], 1,500–

2,000 m, Dec 1998, 1♂ (MC); Río Marañon, Numparque, 1,000 

m, Oct 2006, 1♂ (MC); Pasco, between La Paz and La Salud, 

about 1,800 m, 1 ♀ (ZMHB). The last specimen is the type of the 

unavailable name, Thecla cosmophila Tessmann. The data label 

has an elevation of 2,000 m, but the original description was “in 

etwa 1800 m” [at approximately 1800 m]. 

Etymology. This name is intended to be reminiscent of the old unavailable name for this species (see Nomenclature below). It is proposed as a nonlatinized noun in apposition.

Nomenclature. The name Thecla cosmophila

Tessman, which was proposed as a female form of Thecla orobiana, is unavailable because under ICZN Articles 45.5 and 45.6 

a form relegated to one sex cannot be subspecific. Robbins and 

Lamas (2002) were incorrect in considering Thecla cosmophila

Bridges, 1988 to be an available name because it was published 

after 1985. 

Distribution. (Figure 118) Eastern slope of the 

Andes in Ecuador and Peru.

Habitat. Wet montane forest from 1,250 to 2,000 m. 

Behavior. Males are attracted to rotting fish—most 

were collected on traps, but two were on leaves. 

Remarks. Paraspiculatus cosmo and P. orobia are 

sister species (see Phylogenetic Analyses) and appear to be elevationally parapatric. In our study material, the lowest elevation for P. cosmo is 1,250 m, which is the same as the highest 

recorded elevation for P. orobia.

Paraspiculatus orobia (Hewitson, 1867)

FIGURES 1, 18–21, 47, 59, 65, 75, 86, 105, 113, 117, 126–130

Updated Diagnosis and Description. Males 

of P. cosmo and P. orobia are the only two Paraspiculatus species with regularly spaced gray/brown scales at the base of the 

male dorsal hindwing costa (Character 13, Figures 75, 76). The 

differences between these two taxa are noted in the account of 

P. cosmo. 

The “black line at the end of the [dorsal forewing] cell” 

mentioned by Hewitson (1867:103) varies intraspecifically. The 

presence of tails in P. orobia varies geographically. 

Female. (Figures 19, 21) The female is associated by 

CO1 DNA sequences. The apparent differences in wing shape in 

the two illustrated females are primarily due to poor preparation 

of the latter specimen. 

Genitalia. (Figure 86, 105) Variation among nine 

male genitalic dissections provided no evidence to suggest more 

than one species is represented by this name. The base of each 

valve is displaced ventrally, and its ventral surface lacks setae. 

Variation. Paraspiculatus orobia is the only species 

in the genus that occurs in two of Brown’s (1982) major biogeographical zones. Perhaps because of its wide distribution, it is one 

of the most geographically variable species. Males and females 

from the lower Amazon Basin, as far west as Rondonia (Brazil), 

and the Atlantic region possess tails; those from the upper Amazon Basin, from Colombia to Bolivia, lack tails (Figures 18–21). 

Those with and without tails do not differ in other morphological traits and do not differ in CO1 DNA sequences. No locality 

is known where both forms—and/or intermediate forms—occur. 

Males from the Amazon region have the characteristically welldeveloped shining blue scales anterior of the cubital vein on the 

ventral forewing (Figures 18, 59), but the expression of this blue 

patch is reduced in individuals from the Atlantic region. 

Distribution. (Figure 117) Paraspiculatus orobia 

is widely distributed in the lowland Amazon and Atlantic regions. There is one record (La Fria, Tachira, Venezuela) in the 

area of overlap between the Transandean and Amazon regions. 

Habitat. Paraspiculatus orobia inhabits wet forest 

from 100 to 1,250 m elevation. 

Behavior. Males in the upper Amazon Basin are attracted to rotting fish. Of 79 specimens collected with bait, 62 

were on traps and 17 were on leaves near the ground (Figure 1). 

Remarks. The closest relative of P. orobia is P. cosmo

and they appear to be parapatric in elevation. 

The identity of P. orobia has historically been confused. 

D’Abrera (1995:1194) illustrated six specimens as Thecla orobia, 

but they represent at least three species. Johnson and Constantino (1997) named P. colombiensis as the type of Paraspiculatus, but did not clearly differentiate it from P. orobia. Bálint and 

Moser (2001) figured a specimen of P. colombiensis as P. orobia, 

for which reason they incorrectly synonymized P. colombiensis

with P. orobia. Bálint (2004) then named P. villaanna from a tailless specimen of P. orobia. 

The “band of blue above the median nervure” on the ventral 

forewing, which was noted in Hewitson’s original description, is 

a conspicuous characteristic of this species. However, it also occurs in P. lilyana and occasionally in P. sine (two individuals).

Material Examined (207♂, 6♀). FRENCH 

GUIANA, Saül, 150 m, 7–10 Nov 2005, 1♀ (CF). COLOMBIA, Villavicencio, Ost, 400 m, 18 Dec 1924, 1♂ (MNHN). 

ECUADOR, Morona Santiago, 10 km E of Yanganza, 2°50.0′S, 

78°15.0′W, 800 m, Sep 2003, 1♂ (RCB); 10 km Santiago–Patuca 

Rd, 3°00.8′S, 78°04.5′W, 460 m, 13 Sep 2003, 1♂ (RCB); 15 

km S of Gualaquiza, 3°27.6′S, 78°33.1′W, 800 m, 15 Sep 2000, 

2♂ (RCB); 2 km N of San Isidro, 2°11.9′S, 78°09.4′W, 1250–

1450 m, 08 Jan 2012, 2♂ (CF); 29 Jan 2014, 1♂ (RCB); 12 

Sep 2016, 2♂ (RCB); 13 Sep 2016, 2♂ (RCB); 14 Sep 2015, 2♂

(RCB); 16 Sep 2013, 1♂ (RCB); 17 Sep 2013, 1♂ (RCB); 28 

Sep 2015, 1♂ (RCB); 29 Sep 2015, 6♂ (RCB); 30 Sep 2012, 1♂

(RCB); 30 Sep 2014, 1♂ (RCB); 30 Sep 2016, 6♂ (RCB); 01 Oct 

2013, 1♂ (RCB); 01 Oct 2014, 2♂ (RCB); 16 Nov 2015, 1♂

(RCB); 17 Nov 2015, 1♂ (RCB); 28 Nov 2011, 3♂ (CF); Dec 

2011, 4♂ (CF); 32.8 km Santiago–Puerto Morona Rd, 2°58.9′S, 

77°48.1′W, 670–750 m, 17 Sep 2015, 1♂ (RCB); 27 Sep 

2014, 1♂ (RCB); 28 Sep 2015, 1♂ (RCB); 38.2 km Santiago–

Puerto Morona Rd, 2°57.8′S, 77°47.2′W, 265 m, 13 Sep 2011, 

1♂ (RCB); 47.6 km Santiago–Puerto Morona Rd, 2°56.2′S, 

77°44.8′W, 245 m, 25 Sep 2014, 1♂ (RCB); 53.1 km Santiago–

Puerto Morona Rd, 2°55.0′S, 77°42.7′W, 210 m, 08 Jan 2016, 

1♂ (RCB); 27 Sep 2016, 1♂ (RCB); 54.6 km Santiago–Puerto 

Morona Rd, 2°54.7′S, 77°42.4′W, 14 Sep 2014, 1♂ (RCB); 
3 4 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURES 118–121. Geographic distributions. 118. P. cosmo (triangles). 119. P. transvesta (squares), P. orobiana (triangles). 

120. P. grande (flowers), P. orocana (stars). 121. P. honor (circle, question mark), P. emma (flowers).
NUMBER 649 • 3 5

25 Sep 2014, 1♂ (RCB); 6 km Santiago–Patuca Rd, 3°01.0′S, 

78°03.5′W, 350 m, 01 Oct 2003, 1♂ (RCB); 72 km Patuca–Santiago Rd, 3°02.1′S, 78°06.2′W, 380 m, 02 Oct 2002, 1♂ (RCB); 

Santiago (Hill North of Town), 3°02.3′S, 78°00.3′W, 350 m, 14 

Sep 2014, 3♂ (RCB); 17 Sep 2015, 1♂ (RCB); 25 Sep 2013, 1♂

(RCB); 27 Sep 2014, 2♂ (RCB); 23 Oct 2013, 1♂ (RCB); 29 Oct 

2013, 1♂ (RCB); Yakunk–Cutucú trail, 2 ̊45.11′S 78 ̊10.91′W, 

1000 m, Dec 2003, 1♂ (JHKW); Napo, 12 km Tena–Puyo Rd 

(Finca San Carlo), 1°05.3′S, 77°47.4′W, 600 m, 15 Feb 2008, 

1♂ (RCB); 21 Sep 2011, 1♂ (RCB); 23 Sep 2008, 1♂ (RCB); 

26 Sep 2007, 1♂ (RCB); Sep 1996, 1♂ (JHKW); 14 km Tena–

Puyo Rd (Apuya), 1°06.7′S, 77°46.9′W, 600 m, 10 Jan 2007, 

1♂ (RCB); 10 Jan 2013, 1♂ (RCB); 17 Jan 2011, 1♂ (RCB); 

18 Jan 2011, 1♂ (RCB); Feb 2010, 1♂ (RCB); 19 Feb 2008, 

1♂ (RCB); 27 Feb 2005, 1♂ (RCB); 28 May 2010, 1♂ (RCB); 

Aug 1993, 1♂ (JHKW); 28 Aug 2009, 1♂ (RCB); 06 Sep 2009, 

1♂ (RCB); 09 Sep 2006, 1♂ (RCB); 11 Sep 2005, 1♂ (RCB); 

12 Sep 2015, 1♂ (RCB); 13 Sep 2005, 3♂ (RCB); 15 Sep 2013, 

1♂ (RCB); 16 Sep 2012, 1♂ (RCB); 21 Sep 2011, 1♂ (RCB); 22 

Sep 2005, 1♂ (RCB); 23 Sep 2008, 1♂ (RCB); 24 Sep 2010, 1♂

(RCB); 25 Sep 1997, 1♂ (RCB); 2–3 Oct 1997, 1♂ (RCB); 04 

Oct 2013, 1♂ (RCB); 13 Oct 2011, 2♂ (RCB); 16 Oct 2010, 

1♂ (RCB); 18 Oct 2010, 2♂ (RCB); 22 Oct 2010, 2♂ (RCB); 23 

Oct 2010, 2♂ (RCB); 09 Nov 2010, 1♂ (RCB); 12 Nov 2011, 

1♂ (RCB); 28 Nov 2012, 1♂ (RCB); 29 Nov 2013, 2♂ (RCB); 

Dec 1996, 1♂ (JHKW); Dec 2010, 1♂ (RCB); 28 km Tena–Puyo 

Rd (El Capricho), 1°11.3′S, 77°49.9′W, 800 m, 10 Jan 2013, 1♂

(RCB); 24 Oct 2010, 1♂ (RCB); 25 Oct 2010, 1♂ (RCB); Chichicorrumi, 450 m, 09 Sep 1996, 1♀ (USNM); lower hill N Misahualli, Jungle Lodge Hotel, 1°01.92′S, 77°39.69′W, 500 m, 22 

Oct 2000, 1♂ (USNM); Río Pimpilala (SW of Talag), 1°04.6′S, 

77°56.2′W, 600–900 m, April 2006, 1♂ (RCB); Oct 2005, 4♂

(RCB); Nov 2008, 2♂ (RCB); Satzayacu, Tena–Puyo Rd, 700 m, 

Sep 1996, 1♂ (JHKW); Yachana Reserve, 0°50.5′S, 77°13.8′W 

, 350 m, 17 Feb 2008, 1♂ (RCB); Orellana, 1.2 km Dayuma–

Cononaco Rd, 0°40.7′S, 76°52.4′W, 325 m, 16 Jan 2012, 1♂

(RCB); 14 Nov 2010, 1♂ (RCB); 22 km Loreto–Payamino Rd, 

0°34.9′S, 77°24.4′W, 825 m 20 Nov 2011, 1♂ (RCB); mouth 

of Río Añangu, middle Río Napo, 0 ̊31.7′S 76 ̊23.7′W, 250 m, 

Oct–Dec 2005, 1♂ (JHKW); upper Río Tipitini, Coca–Tiguino 

Rd, 0°40.7′S, 76°52.4′W, Mar 1995, 1♂ (JHKW); Jul 1994, 1♂

(JHKW); Sep 1995, 1♂ (JHKW); Pastaza, 25 km Puyo–Tena Rd, 

1°20.0′S, 77°55.9′W, 1000 m, Dec 2009, 1♂ (RCB); 32 km S of 

Puyo, 1000 m, 01 Oct 1997, 1♂ (RCB); 39.4 km Puyo–Villano 

Rd, 1°25.6′S, 77°43.8′W, 750 m, 11 Sep 2016, 2♂ (RCB); 01 

Oct 2016, 1♂ (RCB); 02 Oct 2015, 1♂ (RCB); 12 Oct 2015, 

1♂ (RCB); Pitirishca, 1°57.8′S, 77°52.1′W, 800 m, 07 Sep 1999, 

1♂ (RCB); 10 Sep 2000, 1♂ (RCB); 19 Sep 2000, 1♂ (RCB); 7.1 

km E of Puyo–Macas Rd at km 32, 1°40.0′S, 77°47.7′W, 850 

m, 05 Mar 2009, 1♂ (RCB); Hills W of Santa Clara, 1°14.5′S, 

77°57.4′W, 900 m, 10 Sep 2000, 1♂ (RCB); Sucumbíos, 16km 

Lumbaqui–La Troncal Rd, 0°00.8′S, 77°15.0′W, 400–500 m, 

06 Jan 2011, 2♂ (RCB); 03 Dec 2015, 1♂ (RCB); Cerro Lumbaqui Norte, 0°01.7′N, 77°19.2′W, 900–1000 m, 30 Dec 2015, 

1♂ (RCB); Zamora Chinchipe, 5 km W of Zamora (Qbda. de 

Chorillos), 1250 m, May 2000, 1♂ (JHKW); 21 May 2000, 1♂

(RCB); Namirez Bajo, 1200 m, 18 Sep 2001, 1♂ (RCB). PERU,

Amazonas, Quebrada Chingaza, 5°22′S, 78°26′W, 500 m, 22 

Sep 1999, 2♂ (USNM); Huánuco, Tingo Maria, Aug 2001, 1♂

(CF); Aug 2003, 1♂ (CF); Sept 2009, 1♂ (CF); Aug 2011, 1♂

(CF); Tingo Maria, 670 m, Sept 2001, 1♂ (MC); Sept 2003, 1♂

(MC); 650 m, Oct 2003, 1♂ (MC); Tingo Maria, Las Pavas, 08 

Oct 1999, 1♂ (CF); Tingo Maria, Huallaga, Apr–Jun 1994, 1♂

(USNM); Loreto, Agua Blanca, 3°56′S, 73°28′W, 130 m, 06 Aug 

2007, 1♀ (MUSM); Iquitos, 1♂ (MNHN); Iquitos, 100 m, Sep 

1999, 1♂ (CF); 10 Sept 2003, 1♂ (MC); Oct 2003, 1♂ (CF); Oct 

2005, 1♂ (CF); Nov 2012, 1♂ (CF); Nov 2002, 1♂ (MC); Dec 

2006, 1♂ (MC); Iquitos, San Pablo, 04°01′S, 71°06′W, 100 m, 

Feb 2011, 2♂ (CF); Pantoja, Río Napo, Nov 2000, 1♂ (MC); 

Tamshiyacu, 100 m, Nov 2013, 1♂ (CF); Polis, Río Momon, 

100 m, Jun 2010, 1♂ (CF); Picuroyacu, 03°37′S, 73°15′W, Jun 

2010, 126 m, 1♂ (CF); May 2012, 1♂ (CF); Aug 2013, 1♂ (CF); 

Pévas, 03°19′S, 71°51′W, Jun 2010, 1♂ (CF); Aug 2010, 1♂

(CF); Río Sucusari, 22 Oct 2008, 1♀ (CF); Madre de Dios, 13 

km SW Puerto Maldonado, 300 m, 22 Oct 1983, 1♂ (USNM); 

Boca Río La Torre, 300 m, 30 Oct 1983, 1♂ (USNM); ); Parque 

Manu, Pakitza, 11°55.8′S, 71°15.3′W, 340 m, 13 Oct 1991, 1♂

(USNM); 15 Oct 1991, 1♂ (USNM); Río La Torre, Tambopata 

Reserve, 300 m, 04 Oct 1986, 1♂ (USNM); San Martin, Moyobamba, Jan–Jun, 1887, 1♀ (BMNH); Pongo de Kainaraki, 350 m, 

Dec 2004, 1♂ (MC); Cusco, Río Araza, 500–700 m, Dec 1994, 

1♂ (MC). BOLIVIA, [Santa Cruz], Buena Vista, 400 m, 01 Jul 

1914, 1♂ (CMNH); 750 m, Aug–Apr 1906–1907, 1♂ (BMNH); 

[Buena Vista], 17°46.9′S, 63°05.6′W, 1♂ (USNM); [La Paz], Mapiri, 1♂ (MNHN). PARAGUAY, [Sapucai], 02 Nov 1904, 1♂

(BMNH). BRAZIL, Amazonas, Maraã, Río Japura, Oct 1997, 

1♂ (MC); Nov 1995, 1♂ (CF); Tonantins, 100 m, Nov 2013, 

2♂ (CF); Fonte Boa, Oct 1994, 1♂ (MC); S. Paulo de Olivencia, 

11 Dec 1921, 1♂ (MNHN); Teffe, 08 Sep 1921, 1♂ (MNHN); 

Tocantins, 24 Sept 1992, 1♂ (MC); Matto Grosso, Parque F. Río 

Doce, 13 Nov 1977, 1♂ (USNM); Diamantino, Alto Río Arinos, 

350 m, 21 Sep 1995, 1♂ (MC); Nobres, Serra do Tombador, 

350 m, 21 Sep 1995, 1♀ (MC); Paraná, [Iguazu], Oct–Dec 1922, 

1♂ (BMNH); Rondônia, Ariquemes, 12 Jul 1986, 1♂ (USNM); 

62 km S Ariquemes, 10°53.0′S, 62°48.0′W, 165 m, 19–29 Sep 

1996, 2♂ (USNM); vic. Caucaulândia, 10°32.0′S, 62°48.0′W, 

160–350 m, 13 Oct 1991, 1♂ (USNM); Alto Paraiso, 9°44.0′S, 

63°16.0′W, 120 m, 08 Aug 2005, 1♂ (MC).

Paraspiculatus orobiana Species Complex

Paraspiculatus orobiana (Hewitson, 1867)

FIGURES 22, 23, 44, 56, 70, 82, 112, 113, 119, 126–130

Updated Diagnosis and Description. Males 

of Paraspiculatus orobiana (Figure 22) are most easily distinguished by (1) the absence of white scales along the distal portion of the ventral forewing costal margin (Character 6) and (2) 
3 6 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

the absence of three white/light-blue spots at the anterior portion 

of the ventral forewing postmedian line (an uncoded character 

due to intraspecific variability, but three spots are present in most 

species). The area along the costa is often a lighter color distal 

of the postmedian line than basal of it, but this is due to lighter 

brown or tan scales, not white scales. We found slight variation 

in the second character noted above as one of eight male specimens had two of the three spots at the anterior portion of the 

postmedian line. The other seven had none. 

According to Hewitson (1867:103), the dorsal surface of 

the forewing has “a black line at the end of the cell.” However, 

we could clearly distinguish this line in only two of the eight 

males in our study series because of variation in the width of the 

distal border. 

Tessman (1928) described Thecla orobiana female form cosmophila, but, as already explained, this female belongs to the 

species that we name P. cosmo.

Female. (Figure 23) The female of P. orobiana was 

associated by similarity with the male in CO1 DNA sequences. 

As with the male, the ventral forewing of the female lacks white 

scales on the distal side of the costa and lacks three white/lightblue spots at the anterior portion of the postmedian line. 

Hewitson (1867:103) described the female of P. orobiana

“like the male except for some white spots on the costal margin 

at the commencement of the band of blue on the underside of the 

anterior wing”. Hewitson associated the female because Bates 

collected it at the type locality of P. orobiana (Ega, Amazonas, 

Brazil), but the wing pattern description is inconsistent with the 

female wing pattern associated by DNA sequences. Both P. orobia and P. colombiensis occur in the Brazilian state of Amazonas 

(Figures 117, 124) and possess white spots at the costa of the 

ventral forewing.

Genitalia. (Figure 82, 112) The male and female 

genitalia are typical of Paraspiculatus, but have no distinctive 

traits. 

Distribution. (Figure 119) Paraspiculatus orobiana is widely distributed in the lowland Amazon region. 

Habitat. Paraspiculatus orobiana inhabits wet forest from 100 to 350 m elevation.

Behavior. Two males from Ecuador were attracted 

to traps baited with rotting fish. 

Remarks. The CO1 barcode sequence of the male P. 

orobiana from Amazonas, Brazil (Appendix A) is slightly divergent from the sequences of Ecuadorian and Peruvian specimens, 

but clusters with them (Figure 113). We found no evident wing 

pattern differences among these specimens. 

Material Examined (8♂,1♀). COLOMBIA, 

Amazonas, Río Caquetá, La Pedrera, 1°18′S, 69°42′W, 120 m, 

26 Apr 1992, 1♂ (MUSM). ECUADOR, Morona Santiago, Santiago (Hill North of Town), 3°02.3′S, 78°00.3′W, 350 m, 27 Sep 

2014, 1♂ (RCB); 27 Sep 2015, 1♂ (RCB). PERU, Amazonas, 

Cavallo Cocho, May–Jul 1884, 1♂ (BMNH); Loreto, Iquitos, 

4°16′S, 73°25′W, 100 m, Oct 2005, 1♂ (CF); Picuroyacu, 3°37′S, 

73°15′W, 10 Nov 2008, 1♀ (USNM). BRAZIL, Amazonas, Ega, 

1♂ holotype (BMNH); Maraã, Río Japurá, May 1995, 1♂ (MC); 

Manicore, 16 Aug 1976, 1♂ (USNM).

Paraspiculatus transvesta Species Complex

Paraspiculatus transvesta

Robbins and Busby, new species

FIGURES 24, 25, 83, 106, 119, 126, 129, 130

Diagnosis and Description. Males of Paraspiculatus transvesta are distinguished by the dull dorsal pattern, 

which lacks blue except for some greenish-blue scales near the 

base of the wings. The ventral forewing has brownish iridescent 

scaling posterior of the cubital vein that is visible when the wing 

is held at an acute angle under a bright light (Character 7), a trait 

that it shares with P. orobiana. The ventral wing pattern is typical 

for the genus and does not have unique characters. Mean forewing length is 17.4 mm (SD = 0.0, n = 1) in the male and 17.3 mm 

(SD = 0.64, n = 2) in the females. We illustrate the wing pattern 

of each sex (Figures 24, 25) and their genitalia (Figures 83, 106).

Female. (Figure 25) The female is associated by similarities with the male in dorsal wing pattern and in geographic 

distribution. The pattern of basal green-blue scales on the dorsal 

wings of both the female and male is unique in the genus. The 

ventral pattern is the same as in the male. We were unable to 

obtain a CO1 sequence from either sex due to the age of the 

specimens. 

Type material.

Holotype (♂): (Figure 24) GUAT[EMALA]/ Cayuga 

[rectangular, white, printed], aug [rectangular, white, printed], 

Schaus and/ Barnes/ coll [rectangular, white, printed], USNM 

ENT 00179436 [rectangular, white, printed barcode label], 

GENITALIA No./ 1982: 210 ♂/ R. K. ROBBINS [rectangular, 

green, printed], Holotype ♂/ Paraspiculatus transvesta/ Robbins 

& Busby [rectangular, red, printed]. Deposited in USNM.

Paratype (1♀): MEXICO, [Veracruz], Sontecomapan, 

[18°30′N, 95°02′W], May 1955, 1♀ (MGCL).

Other Material Examined (1♀): HONDURAS, San 

Pedro Sula, 1895, 1♀ (BMNH).

Etymology. We name this species P. transvesta because the male looks like a female. It is proposed as a nonlatinized noun in apposition. 

Genitalia. (Figures 83, 106) The male and female 

genitalia are typical of Paraspiculatus. 

Distribution. (Figure 119) Mexico to Honduras.

Habitat. This species appears to be a denizen of lowland forest. Elevation within 10 km of the type locality is 50–500 

m, within 10 km of Sontecomapan is 10–800 m, and within 10 

km of San Pedro Sula is 50–1,100 m. The wide range of elevations in Figure 130 for P. transvesta is an artifact of imprecise 

labeling of specimens. 

Remarks. Many characters could not be scored for 

P. transvesta because we could not homologize the male’s femalelike wing pattern with males of other species. 
NUMBER 649 • 3 7

Paraspiculatus orocana Species Complex

Paraspiculatus orocana, P. grande, and P. honor form a 

monophyletic lineage of allopatric/parapatric species. Males are 

distinguished by three characters. First, the dark gray scales on 

the dorsal hindwing cell Sc+R1

-Rs are concentrated in the middle 

of the cell and have iridescent edges that give off a sheen under a 

bright light (Character 13, Figure 74). The gray/brown scales in 

male P. orobia and P. cosmo are less iridescent and more evenly 

spaced so that the center of the cell is not distinct (Figures 75, 76). 

Second, the P. orocana group males have a relatively straight inner 

margin of the anal lobe that contrasts in shape with the smaller, 

more rounded lobes of other Paraspiculatus species (Character 

15, Figure 77). Finally, the ventral anal lobes in the P. orocana

group have long, dark fringe scales that lack white scales at the 

end of vein 2A or anteriorly on the inner margin (Character 22) 

as in most other Paraspiculatus species. Other identifying male 

characters shared by the P. orocana group as well as other species 

in the genus are: (1) A broad black border and apex on the dorsal 

surface of the forewing (Figures 45, 46), with the border having a straight inner edge (Character 3), (2) a less rounded, more 

angular hindwing (Character 11), (3) an area of iridescent, silverblue scales located both anterior and posterior of the cubital vein 

on the ventral forewing (Characters 7, 8, Figure 68), and (4) the 

absence of hindwing tails. The P. orocana species group is monophyletic in a phylogenetic analysis of CO1 sequences.

Paraspiculatus grande Busby, Robbins, 

and Moser, new species

FIGURES 26, 27, 46, 58, 74, 87, 107, 113, 120, 126–130

Diagnosis and Description. Males of Paraspiculatus grande differ from other species in this complex by a 

postmedian line consisting of three light blue spots at the costal 

margin with four posterior blue spots (Figure 58). In P. orocana

and P. honor, the anterior spots are absent or vestigial, and the 

number of posterior blue markings is limited to one or two spots 

(Figure 57). The white scales near the costa of the ventral forewing in P. grande form two separate patches, one on the distal side of the three median spots and the other at the apex. In 

contrast, the white scales in both P. orocana and P. honor form 

more of a single crescent-shaped cluster between the median and 

apex. Mean forewing length is 21.3 mm (SD = 0.64, n = 21) in 

males and 20.4 mm (SD = 0.0, n = 1) in the female. We illustrate 

the wing pattern of each sex (Figures 26, 27) and their genitalia 

(Figures 87, 107).

Female. (Figure 27) The female is associated by 

CO1 DNA sequences. The only known female is from Panama 

whereas all males are from western Ecuador. Panama and western Ecuador are part of the Transandean biogeographic zone and 

have a high incidence of faunal overlap (Brown, 1982).

Type material.

Holotype (♂): (Figure 26) ECUADOR: Esmeraldas/ 12 

km Lita–San Lorenzo Road/ 0°53.1′N, 78°30.9′W 850 m/ 26 

March 2011 Río Chuchuví/ I. Aldas, R. C. Busby, leg. [rectangular, white, printed], USNM ENT 00180789 [rectangular, white, 

printed barcode label], GENITALIA No./ 2013: 86 ♂/ C. FAYNEL [rectangular, green, printed], Holotype ♂ / Paraspiculatus 

grande / Busby, Robbins, and Moser [rectangular, red, printed]. 

Deposited in USNM.

Paratypes (22♂, 1♀): PANAMA, Bocas del Toro, nr. 

Chiriquí Grande, 1100 m, 19 Jun 1982, 1♀ (USNM). ECUADOR, Esmeraldas, 12 km Lita–San Lorenzo Road, 0°53.1′N, 

78°30.9′W, 850 m, March 2001, Río Chuchuví, 1♂ (USNM); 

May 2011, 1♂ (USNM), 1♂ (RCB); 15 May 2014, 1♂ (RCB); 

19 Jun 2011, 1♂ (RCB); 22 Jun 2011, 1♂ (RCB); Jul 1999 1♂

(JHKW); Jul 2005, 1♂ (RCB); Jul 2006, 1♂ (USNM); 14 Jul 2011, 

2♂ (USNM); Aug 2006, 1♂ (MECN); Aug 2008, 1♂ (USNM); 

Oct 2008, 1♂ (MECN); 25 Oct 2014, 1♂ (RCB); 28 Oct 2014, 

1♂ (USNM); El Encanto, km 17 Lita–San Lorenzo Road, 850 

m, Jul 2003, 1♂ (JHKW); Alto Tambo, [0°54.7′N, 78°32.8′W], 

[725 m], Aug 2000, 2♂ (RCB); Lita, [0°52.5′N, 78°28.0′W], 

600–1000 m, Apr 2002, 1♂ (MC); Carchi, El Baboso [Carmelo], 

0°53.1′N, 78°26.5′W, 950 m, 17 Jun 2011, 1♂ (RCB).

Other Material Examined (2♂): ECUADOR, Pichincha, Pacto–Guayabillas Road, 01 May 2003, 1♂ (MC); Pacto, 

1100 m, 20 Mar 2001, 1♂ (CF). These specimens (the southernmost data points) were excluded from the type series because of 

uncertainty regarding the exact locations.

Etymology. Paraspiculatus grande is among the 

largest species in the genus. The name is proposed as a nonlatinized noun in apposition. 

Genitalia. (Figures 87, 107) The male and female 

genitalia are typical of Paraspiculatus. The ventral base of the 

valvae is similar to that of P. orobia, but this may be a superficial 

resemblance. 

Distribution. (Figure 120) Panama to the western 

slope of the Andes in Ecuador. 

Habitat. Wet lower montane forest from 725 to 

1,100 m elevation.

Behavior. The vast majority of specimens in our 

study group were collected with a net. Several males exhibited 

territorial behavior, perching 3 to 4 m above the ground along 

a ridgetop in the early afternoon. Specific records for Río Chuchuví (Ecuador, Esmeraldas Province) are 12:30, 13:10, and 

13:35 hours (vouchers in RCB and USNM). Only one male of P. 

grande was collected on a trap baited with fish.

Remarks. Paraspiculatus grande occurs at localities 

below 1,100 m; P. honor occurs at elevations above 1,200 m. 

Also, P. grande is allopatric with P. orocana (Figure 120). 

Paraspiculatus honor Busby, Robbins, 

and Hall, new species

FIGURES 28, 29, 88, 108, 113, 121, 126–130

Diagnosis and Description. Although males of 

P. grande have four blue spots at the posterior end of the ventral forewing postmedian line, P. honor and P. orocana typically 
3 8 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

possess only one or two of these blue spots. The postmedian-line 

white spots at the ventral forewing costa in P. honor are indistinct 

because they blend in with the median crescent-shaped patch of 

white scales that extends distally toward the apex. This pattern 

contrasts with the sharply defined spots in P. grande and the lack 

of spots in P. orocana (Figure 57). The black dorsal forewing 

apex of P. honor extends basally to the distal end of the discal cell 

(Character 2), in which it is similar to P. grande, but differs from 

P. orocana. In the latter, blue scales separate the discal cell from 

the black apex (Figure 45). CO1 DNA sequences of P. honor also 

distinguish this taxon from P. grande and P. orocana. Mean forewing length is 18.4 mm (SD = 1.55, n = 3) in males and 18.8 mm 

(SD = 0.0, n = 1) in the female. We illustrate the wing pattern of 

each sex (Figures 28, 29) and the genitalia (Figures 88, 108). 

Type material.

Holotype (♂): (Figure 28) ECUADOR: Pichincha/ 

Mindo, Río Napombillo/ 1200 m, 26 Aug 1999/ K. R. Willmott 

[rectangular, white, printed], USNM ENT 00171673 [rectangular, white, printed barcode label], Holotype ♂/ Paraspiculatus 

honor/ Busby, Robbins & Hall [rectangular, red, printed]. Deposited in USNM.

Paratypes (2♂, 1♀): ECUADOR, Pichincha, 5 km 

Nanegal–García Moreno Rd, 0°09.4′N, 78°39.4′W, 02 Jun 2015, 

(1,375–1,700 m), 1♂ (RCB); 16 Jun 2012, 1♀ (RCB); Charchi, 

Chical, 0°57.7′N, 78°12.2′W, 1391 m, 19 Aug 2016, 1♂ (RCB).

Other Material Examined (2♂): COLOMBIA, [no further locality data], 2♂ (AMNH). 

Female. (Figure 29) The female of P. honor was recognized by its CO1 DNA sequence. It was found in the same 

location as the male paratype, a montane habitat 30 km distant 

from where the male holotype was collected.

Etymology. This beautiful, rare butterfly is named 

for Honor Leslie-Melville, at the request of Keith Willmott and 

Julia Robinson Willmott, in celebration of the friendship and 

support of the Leslie-Melville family over many years. Keith 

Willmott and Jason Hall were first to discover this species in 

Ecuador. The name is a noun in apposition. 

Genitalia. (Figures 88, 108) The male and female genitalia are typical of Paraspiculatus. The distal cornutus is minute. 

Distribution. (Figure 121) Northwestern Ecuador 

and Colombia. We assume the two specimens labeled “Colombia 

S.A., collected by Felipe Ovalle” are from western Colombia as 

most specimens collected by him occur in western Colombia. 

Habitat. Paraspiculatus honor inhabits wet montane forest from 1,200–1,700 m elevation. 

Behavior. The holotype was attracted to a trap 

baited with rotting fish (JHKW, personal communication). Another male was attracted to a leaf baited with rotting fish. 

Paraspiculatus orocana (H. H. Druce, 1912)

FIGURES 30, 45, 57, 68, 77, 89, 113, 120, 126–130

Updated Diagnosis and Description. Males 

of P. orocana (Figure 30) are separated from P. honor and 

P. grande by the presence of blue scales on the dorsal forewing 

between the distal end of the discal cell and broad black border 

(Character 2). Two characters on the ventral forewing that distinguish P. orocana from the other species in Paraspiculatus (except for P. honor) are (1) a crescent-shaped patch of white scales 

along the distal side of the costa and (2) a single postmedian blue 

spot (occasionally two) instead of a line composed of between 

four and seven spots and line segments. 

The original description by Druce (1912) distinguishes 

P. orocana from P. orobia based on “black [dorsal] veins and 

tail” in the latter. Our study material shows that the dorsal black 

veins occur in many Paraspiculatus species including P. orocana

and that the amount of black varies intraspecifically and with 

age of the specimen. Druce also mentions the scarcity of “metallic green scales towards the anal angle” in P. orocana. As in the 

case of black dorsal veins, this character is highly variable and of 

little diagnostic value.

Female. Unknown. Females of P. honor and P. grande 

share the same dorsal wing pattern (dark gray with basal blue), 

suggesting that females of P. orocana will be similar. 

Genitalia. (Figure 89) The male genitalia are typical of Paraspiculatus.

Distribution. (Figure 120). Paraspiculatus orocana is widely distributed in the eastern Andes. 

Habitat. Paraspiculatus orocana inhabits forest 

from 1,250 to 1,600 m elevation in Ecuador, but the holotype 

from Peru was collected at 900 m. For this reason, we tentatively 

classify it as a lower montane species.

Behavior. In Ecuador, the males in our study material were attracted to rotting fish in traps (12 individuals) and on 

leaves (one individual). 

Material Examined (31♂). ECUADOR, Morona Santiago, 14 km W of Macas (Río Abanico), [2°15.4′S, 

78°11.7′W], 1600 m, 12 Sep 1999, 1♂ (RCB); 2 km N of San Isidro, 

2°11.9′S, 78°09.4′W, 1250–1450 m, 07 Jan 2013, 1♂ (RCB); 14 

Aug 2011, 1♂ (CF), 12 Sep 2016, 1♂ (RCB); 13 Sep 2016, 2♂

(RCB); 14 Sep 2015, 1♂ (RCB); 19 Sep 2012, 1♂ (RCB); 29 Sep 

2015, 1♂ (RCB); 29 Sep 2016 1999, 1♂ (RCB); 30 Sep 2016, 

1♂ (RCB); 28 Nov 2011 1999, 2♂ (CF); Bosque de Domono, 

2°11.0′S, 78°06.2′W, 1325 m, 16 Sep 2003, 1♂ (RCB); Napo, 49 

km Tena–Loreto Rd., 0°42.9′S, 77°44.4′W, 1350 m, Nov 2012, 

1♂ (RCB); Pastaza, N of Mera, 1350 m, Jul 2015, 1♂ (JHKW);

Tungurahua, El Tigre, Topo, 14 Aug 1998, 1♂ (CF); Zamora 

Chinchipe, 5 km W of Zamora (Qbda. de Chorillos), 1250 m, Apr 

1995, 1♂ (JHKW); May 2000, 3♂ (JHKW); 17 May 2000, 3♂

(RCB); 19 Sep 2001, 1♂ (RCB); Zamora, 4°04.5′S, 78°58.1′W, 06 

Oct 2002, 1♂ (RCB); 08 Oct 2002, 1♂ (RCB). PERU, San Martin, 

Afluente, 7°18′S, 77°12′W, 1,350 m, 01 Aug 2007, 1♂ (CF); El 

Porvenir, 900 m, Oct 1909, 1♂ holotype (BMNH); Jorge Chaves, 

1400 m, Nov 2003, 1♂ (MC). BOLIVIA, [La Paz], Nor Yungas, 

Caranavi, 1000–1500 m, Jan 2003, 1♂ (MC).

Paraspiculatus colombiensis Species Complex

Paraspiculatus colombiensis, P. azul, P. sine, P. noemi, 

P. lilyana, and P. emma form a monophyletic lineage we call the 
NUMBER 649 • 3 9

P. colombiensis species complex. In males, the scales on the distal 

part of the ventral forewing posterior to vein Cu2

 are dull bluish 

gray (Character 9, Figures 60–64), in contrast to other members 

of the genus, where these scales are brown or brownish gray 

(Figures 53–59). Other characteristic male traits (not necessarily unique within Paraspiculatus) are (1) blue and black wing 

scales (Figures 71–73) in the basal half of the dorsal hindwing 

anterior to vein Rs (Character 13) that do not differ from scales 

on other parts of the wing, (2) an area of iridescent, blue or 

silver-blue scales (Figures 66, 67) posterior to the cubital vein 

of the ventral hindwing (Character 7), (3) a rounded hindwing 

(Character 11), and (4) a black forewing border (Figures 48–52) 

with a concave inner edge (Character 3). Phylogenetic analysis 

of DNA sequences also shows this lineage to be monophyletic. 

The females of P. colombiensis and P. azul are brown dorsally without blue scales as in P. orobia and P. orobiana. The 

other females in this complex are unknown. 

Paraspiculatus emma Busby and Robbins, 

new species

FIGURES 31, 48, 60, 69, 90, 113, 121, 126–130

Diagnosis and Description. Males of Paraspiculatus emma are distinguished from others in the P. colombiensis species group by (1) a wider black border on the dorsal 

forewing, and (2) less blue at the hindwing apex in cell Rs-M1

(Character 14). The silver-blue iridescent scales found on the 

posterior side of the cubital vein (Figure 69) are not as prominent on the anterior side as in the other species of this group. The 

white scales along the costa of the ventral forewing in P. emma

are lumped into two patches, a trait shared with P. colombiensis

and P. noemi. The hindwing in P. emma has no tails, a character 

also found in P. sine and P. lilyana. Mean forewing length is 16.3 

mm (SD = 0.92, n = 38) in males. We illustrate the wing pattern 

(Figure 31) and male genitalia (Figure 90).

Female. Females of P. emma are unknown. 

Type material.

Holotype (♂): (Figure 31) ECUADOR: Napo/ 14 km 

Tena–Puyo Road/ 1°06.7′S, 77°46.9′W 600 m/ 21 September 2005 (Apuya)/ Robert C. Busby, leg. [rectangular, white, 

printed], 11:00 hrs/ 5 m [rectangular, white, handwritten, blue 

ink], USNM ENT 00180812 [rectangular, white, printed barcode label], GENITALIA No./ 2013: 69 ♂/ C. FAYNEL [rectangular, green, printed], Holotype ♂/ Paraspiculatus emma/ Busby 

& Robbins [rectangular, red, printed]. Deposited in USNM.

Paratypes (51♂): ECUADOR, Napo, 14 km Tena–Puyo 

Road, 1°06.7′S, 77°46.9′W 600 m, Apuya, 05 Aug 2012, 1♂

(USNM); 28 Aug 2009, 2♂ (RCB); 06 Sep 2009, 1♂ (RCB); 9 

Sep 2006, 1♂ (USNM); 9 Sep 2010, 1♂ (USNM); 10 Sep 2010, 

1♂ (RCB); 11 Sep 2016, 1♂ (RCB); 13 Sep 2005, 1♂ (USNM); 

20 Sep 2011, 1♂ (RCB); 21 Sep 2005, 1♂ (USNM); 21 Sep 

2011, 2♂ (USNM), 1♂ (RCB); 23 Sep 2008, 5♂ (RCB), 2♂

(USNM); 23 Sep 2010, 1♂ (RCB); 24 Sep 2011, 1♂ (RCB); Oct 

2005, 1♂ (USNM); 2–3 Oct 1997, 1♂ (RCB); 11 Oct 2011, 1♂

(USNM); 12 Oct 2011, 2♂ (RCB); 18 Oct 2010, 1♂ (RCB); 22 

Oct 2010, 1♂ (USNM); Nov 2011, 1♂ (CF); 12 Nov 2011, 1♂

(MECN); 03 Jan 2012, 1♂ (RCB); 08 Jan 2011, 1♂ (MECN), 

1♂ (RCB); 13 Jan 2010, 1♂ (MECN); 17 Jan 2011, 2♂ (RCB); 

18 Jan 2011, 1♂ (USNM); 07 Mar 2009, 1♂ (USNM); 11 Mar 

2005, 1♂ (RCB); 28 km Tena–Puyo Road, 1°11.3′S, 77°49.9′W, 

800 m, El Capricho, 05 Aug 2012, 1♂ (RCB); 11 Sep 2006, 

1♂ (USNM); 04 Jan 2007, 1♂ (USNM); Río Pimpilala (SW of 

Talag), 1°04.6′S, 77°56.2′W, 600–900 m, Jul 2004, 1♂ (USNM); 

23 Sep 2010, 1♂ (MECN); Dec 2005, 1♂ (RCB); 07 Jan 2006, 

1♂ (RCB); 12 km Tena–Puyo Road (Finca San Carlo), 1°05.3′S, 

77°47.4′W, 600 m, 02 Oct 2013, 1♂ (RCB), 23 Sep 2008, 1♂

(USNM); Morona Santiago, Bomboiza nr. Gualaquiza, 850 m, 

Jul 1993, 1♂ (JHKW).

Other Material Examined (2♂): ECUADOR, Morona 

Santiago, Nueva Tarqui, 15 Sep 2000, 1♂ (RCB). PERU, [Loreto], Balsapuerto, [5°50′S, 76°36′W], 1♂ (MNHN). These 

specimens were not made paratypes because their elevations are 

uncertain. The elevation of Nueva Tarqui is 900 m with nearby 

ridges over 1,150 m. The elevation of Balsapuerto, Peru is 200 m 

(Lamas, 1976). However, at the time that the Peruvian male of 

P. emma was collected, there was a well-frequented trail between 

Moyobamba and Balsapuerto that went through the mountains 

southwest of Balsapuerto. It is unclear if this specimen was collected in Balsapuerto, which is at lower elevation than the paratypes listed above, or at higher elevation on this trail. 

Etymology. We name this beautiful species 

P. emma in honor of Emma Rose Hughes, granddaughter of 

Robert C. Busby. The name is a noun in apposition. 

Genitalia. (Figure 90) The genitalia are typical of 

Paraspiculatus, but lack evident distinctive traits.

Distribution. (Figure 121) Eastern slope of the 

Andes in Ecuador and Peru. 

Habitat. Wet lower montane forest. The majority of 

specimens were sampled in a relatively small area in Napo Province at an elevation range of 600–900 m. 

Behavior. Males are attracted to traps baited with 

rotting fish. 

Remarks. Paraspiculatus emma is sympatric with 

four of the other five species in the P. colombiensis species complex (excluding P. noemi) in eastern Ecuador at 800 m elevation. 

Paraspiculatus sine Busby and Robbins, 

new species

FIGURES 2, 32, 49, 61, 66, 91, 98, 113, 123, 126–130

Diagnosis and Description. Males of Paraspiculatus sine are distinguished from the other species in the P. colombiensis group by the unique pattern of white scales along the 

costa of the ventral forewing (Character 6). These scales form a 

narrow band that extends distally from the anterior part of the 

postmedian line to the apex (Figure 61). White scales almost always cover part of the dark submarginal band in the apex. In contrast, P. colombiensis (Figure 62), P. noemi, and P. emma (Figure 

60) have two separate patches of white scales, one median and one 

at the apex, and P. azul (Figure 63) and P. lilyana (Figure 64) lack 
4 0 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

white scales along the ventral forewing costa. Another trait of P. 

sine is the round hindwing with no tail (shared with P. lilyana and 

P. emma). The dorsal color in the limbal part of the forewing in P. 

sine has a “grainy” appearance caused by the intermixing of many 

black scales (Figure 49), which contrasts with the much smoother 

look of P. azul and P. lilyana (Figures 51, 52). Mean male forewing length is 16.9 mm (SD = 1.11, n = 18). We illustrate the wing 

pattern (Figure 32) and male genitalia (Figure 91).

Female. Females of P. sine are unknown.

Type material.

Holotype (♂): (Figure 32) ECUADOR. Morona Santiago/ 15 km S of Gualaquiza/ 3°27.6′S, 78°33.1′W 850 m/ 

16 September 2000/ Robert C. Busby, leg. [rectangular, white, 

printed], USNM ENT 00180851 [rectangular, white, printed 

barcode label], GENITALIA No./ 2013. 58 ♂/ C. FAYNEL [rectangular, green, printed], Holotype ♂/ Paraspiculatus sine / Busby 

and Robbins [rectangular, red, printed]. Deposited in USNM.

Paratypes (23♂): ECUADOR, Morona Santiago, 15 

km S of Gualaquiza [Bomboiza], 3°27.6′S, 78°33.1′W, 850 

m, Jul 1993, 1♂ (JHKW); 16 Sep 2000, 1♂ (USNM); 29 Sep 

2000, 1♂ (USNM); 05 Oct 2002, 1♂ (RCB); 2 km N of San 

Isidro, 2°11.9′S, 78°09.4′W, 1,250–1,400 m, 29 Jan 2014, 1♂

(RCB); 13 Sep 2014, 1♂ (MECN), 1♂ (RCB); 16 Sep 2013, 1♂

(RCB); 17 Sep 2013, 1♂ (USNM); 30 Sep 2013, 1♂ (USNM), 

1♂ (MECN); 30 Sep 2014, 1♂ (RCB); Napo, 25 km S of Tena 

[nr. Pimpilala], 1°04.3′S, 77°56.5′W, 875 m, 08 Oct 2003, 1♂

(RCB); Río Pimpilala, 1°04.6′S, 77°56.2′W, [900 m], Nov 2008, 

1♂ (USNM); Aug 2016, 2♂ (RCB); 28 km Tena–Puyo Road 

(El Capricho), 1°11.3′S, 77°49.9′W, 800 m, 05 Jan 2007, 1♂

(USNM); Zamora Chinchipe, Zamora, 4°04.5′S, 78°58.1′W, 

1,450 m, 08 Oct 2002, 1♂ (RCB); 10 km Los Encuentros–El 

Pangui Rd., 3°42.2′S, 78°36.0′ , 1,000–1,200 m, 04 Oct 2007, 

1♂ (RCB); Pastaza, 25 km N of Puyo, 1,000 m, 9–11 Oct 1988, 

1♂ (RCB); 37 km Puyo–Arajuno Road, 1°22.6′S, 77°42.6′W, 

1,100 m, 11 Jan 2012, 1♂ (USNM); Sucumbios, Cerro Lumbaquí 

Norte, 0°01.7′N, 77°19.2′W, 900–1,000 m, 01 Jan 2007, 1♂

(RCB). PERU, Cusco, Carretera a Manu km 87, Chontachaca, 

13°01.8′S, 71°29.7′W, 972 m, 24–25 Oct 2014, 1♂ (CF). 

Etymology. The name P. sine is derived from a 

nickname that we gave this species when we first began to work 

on Paraspiculatus. The name is proposed as a nonlatinized noun 

in apposition. 

Variation. Two of 24 males examined have a band 

of shining blue scales on the anterior side of the cubital vein on 

the ventral forewing, similar to those in P. orobia and P. lilyana.

Genitalia. (Figures 91, 98) The genitalia are typical 

of Paraspiculatus and lack distinctive characters. Variation in the 

male genitalia spiculate pad is illustrated (Figure 98).

Distribution. (Figure 123) Eastern slope of the 

Andes in Ecuador.

Habitat. Paraspiculatus sine inhabits wet lower 

montane forest from 800 to 1,450 m elevation.

Behavior. Males are attracted both to traps and 

plants (Figure 2) baited with rotting fish.

Remarks. Paraspiculatus sine appears to be the 

highest-elevation species in the P. colombiensis complex. However, its range overlaps that of P. emma, P. lilyana, P. azul, and 

P. colombiensis from 800 to 900 m elevation in eastern Ecuador.

Paraspiculatus colombiensis

Johnson and Constantino, 1997

FIGURES 5, 33, 34, 50, 62, 67, 77, 92, 109, 113, 124, 126–130

Updated Diagnosis and Description. Males 

of Paraspiculatus colombiensis are best diagnosed using the following combination of characters: (1) blue wing scales in the 

basal half of the dorsal hindwing anterior to vein Rs (Character 13, shared with five species in the P. colombiensis species 

group; the exception is P. noemi whose scales are mostly black), 

(2) white scales along the costa in the distal part of the ventral 

forewing (Character 6, excludes P. azul and P. lilyana), (3) the 

white scales on the ventral forewing divided into two parts—one 

median and one near the apex (character not coded, excludes P. 

sine), and (4) relatively narrow dorsal forewing black margin 

near the tornus (character not coded, excludes P. emma whose 

forewing margin is much broader). Also of note, P. colombiensis

has tails (shared with P. noemi and P. lilyana) and a somewhat 

darker dorsal blue color than P. azul, P. lilyana, or P. sine. Unfortunately, characters such as the presence or absence of tails and 

color differences become less reliable in old specimens. 

The original diagnosis by Johnson and Constantino is confusing because it mainly focuses on differences between P. colombiensis and the “traditional orobia-Group” (P. orobia, 

P. orocana, and P. orobiana). These species have different wing 

patterns, and the diagnostic comparisons do not apply in every 

case. The only specific comparison of P. colombiensis to P. orobia is based on ventral ground color, which is highly variable and 

of no diagnostic value. 

Female. (Figure 34) The female was associated with 

the male using CO1 DNA sequences. 

Genitalia. (Figures 92, 109) The genitalia of P. colombiensis are typical of Paraspiculatus, but lack evident distinctive traits.

Distribution. (Figure 124) Paraspiculatus colombiensis is widely distributed throughout the Amazon region. 

Habitat. Paraspiculatus colombiensis inhabits wet 

forest from approximately 100 to 1,300 m elevation. 

Behavior. Males in the upper Amazon Basin are attracted to traps baited with rotting fish. Recently, six females 

of P. colombiensis were collected on traps at Santiago, Morona 

Santiago, Ecuador. These are the first records of female Paraspiculatus coming to bait and is especially interesting because 

Santiago has been sampled for years without encountering 

females. 

Remarks. Bálint and Moser (2001) synonymized 

P. colombiensis with P. orobia based on a misidentified individual of the latter. Robbins (2004b) corrected this error, listing 

both as distinct species. 
NUMBER 649 • 4 1

FIGURES 122–125. Geographic distributions. 122. P. noemi (diamonds), P. lilyana (stars). 123. P. sine (squares). 124. P. colombiensis (circles). 125. P. azul (bullets).
4 2 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Material Examined (210♂, 10♀). FRENCH 

GUIANA, Saül, 12 Jun 2005, 1♂ (CF). COLOMBIA, [Putumayo], Umbría, 1♂ (MNHN); Mocoa, Oct 1927, 1♂ (MNHN); 

[Caqueta], Río Ortequaza, 400 m, 15 Sep 1947, 1♂ holotype (MNHN). ECUADOR, Morona Santiago, 1.8 km Santiago–Puerto Morona Rd, 3°02.4′S, 77°59.7′W, 350–500 m, 

13 Jan 2011, 1♂ (RCB); 19 Sep 2006, 2♂ (RCB); Aug 2015, 

1♂ (JHKW); 10 km E of Yanganza, 2°50.0′S, 78°15.0′W, 800 

m, Sep 2003, 3♂ (RCB); 15 km S of Gualaquiza, 3°27.6′S, 

78°33.1′W, 800 m, May 1994, 1♂ (JHKW); 15 Sep 2000, 2♂

(RCB), 1♀ (RCB); 16 Sep 2001, 2♂ (RCB), 27 Sep 2000, 1♂

(RCB); 29 Sep 2000, 3♂ (RCB); 05 Oct 2002, 1♂ (RCB); 24.5 

km Santiago–Puerto Morona Rd, 2°58.1′S, 77°50.5′W, 600 

m, 13 Sep 2011, 1♂ (RCB); 17 Sep 2010, 1♂ (RCB); 22 Sep 

2012, 2♂ (RCB); 26 Sep 2013, 1♂ (RCB); 26 km Santiago–

Puerto Morona Rd, 2°58.4′S, 77°49.9′W, 550 m, 17 Sep 2005, 

2♂ (RCB); 32.8 km Santiago–Puerto Morona Rd, 2°58.9′S, 

77°48.1′W, 670–750 m, 26 Sep 2014, 1♀ (RCB); 26 Sep 2016, 

2♂ (RCB); 27 Sep 2014, 1♂ (RCB); 28 Sep 2014, 1♂ (RCB); 

29 Sep 2014, 1♂ (RCB); 47.6 km Santiago–Puerto Morona 

Rd, 2°56.2′S, 77°44.8′W, 245 m, 24 Sep 2013, 4♂ (RCB); 24 

Sep 2014, 1♂ (RCB); 25 Sep 2014, 1♂ (RCB); 53.1 km Santiago–Puerto Morona Rd, 2°55.0′S, 77°42.7′W, 210 m, 17 Sep 

2016, 1♂ (RCB); 27 Sep 2016, 1♂ (RCB); 54.6 km Santiago–

Puerto Morona Rd, 2°54.7′S, 77°42.4′W, 205 m, 14 Sep 2014, 

1♂ (RCB); 25 Sep 2014, 1♂ (RCB); Santiago (Hill North of 

Town), 3°02.3′S, 78°00.3′W, 350 m, 06 Jan 13, 1♂ (RCB); 06 

Jan 13, 1♂ (RCB); 07 Jan 16, 1♂ (RCB); 08 Jan 15, 2♂ (RCB); 

11 Jul 14, 1♂ (RCB); 31 Aug 2009, 1♂ (RCB); 01 Sep 2009, 

1♂ (RCB); 14 Sep 2014, 1♂ (RCB); 14 Sep 2015, 1♂ (RCB); 

15 Sep 2011, 2♂ (RCB); 16 Sep 2011, 2♂ (RCB); 16 Sep 2014, 

2♂ (RCB); 16 Sep 2016, 1♂ (RCB); 17 Sep 2011, 3♂ (RCB); 

17 Sep 2015, 1♂ (RCB); 18 Sep 2011, 2♂ (RCB); 18 Sep 2015, 

1♂ (RCB); 19 Sep 2015, 1♂ (RCB); 20 Sep 2010, 1♂ (RCB); 

21 Sep 2012, 2♂ (RCB); 22 Sep 2016, 2♂ (RCB); 23 Sep 2012, 

1♂ (RCB); 23 Sep 2014, 2♂ (RCB), 1♀ (RCB); 23 Sep 2015, 

1♂ (RCB); 24 Sep 2015, 1♂ (RCB); 25 Sep 2012, 1♂ (RCB); 

25 Sep 2013, 7♂ (RCB); 25 Sep 2015, 1♀ (RCB); 26 Sep 2014, 

1♂ (RCB), 1♀ (RCB); 26 Sep 2016, 2♂ (RCB); 27 Sep 2013, 

4♂ (RCB); 27 Sep 2014, 1♀ (RCB); 28 Sep 2012, 1♂ (RCB); 

28 Sep 2014, 1♀ (RCB); 29 Sep 2012, 1♂ (RCB); 29 Sep 2014, 

1♂ (RCB); 15 Oct 2011, 4♂ (RCB); 23 Oct 2013, 4♂ (RCB); 

29 Oct 2013, 1♂ (RCB); 3 km Bella Union–Patuca Rd, 650 m, 

20 Sep 2012, 2♂ (RCB); Nueva Tarqui, [1000 m], 15 Sep 2000, 

1♂ (RCB); Yaupi, 300 m, Sep 2010, 1♂ (JHKW); Napo, 12 km 

Tena–Puyo Rd (Finca San Carlo), 1°05.3′S, 77°47.4′W, 600 m, 

18 Jan 2011, 1♂ (RCB); Sep 1996, 1♂ (JHKW); 12 Sep 2010, 

1♂ (RCB); 23 Sep 2008, 1♂ (RCB); 01 Oct 2012, 2♂ (RCB); 

14 km Tena–Puyo Rd (Apuya), 1°06.7′S, 77°46.9′W, 600 m, 03 

Jan 2006, 1♂ (RCB); 03 Jan 2012, 1♂ (RCB); 03 Jan 2016, 1♂

(RCB); 11 Jan 2013, 1♀ (RCB); 14 Jan 2012, 1♂ (RCB); 17 Jan 

2011, 3♂ (RCB); Apr 2006, 1♂ (RCB); May 2009, 1♂ (RCB); 

06 Sep 2009, 2♂ (RCB); 06 Sep 2011, 2♂ (RCB); 10 Sep 2016, 

2♂ (RCB); 11 Sep 2005, 1♂ (RCB); 12 Sep 2005, 1♂ (RCB); 

13 Sep 2005, 1♂ (RCB); 15 Sep 2013, 1♂ (RCB); 21 Sep 2005, 

1♂ (RCB); 23 Sep 2008, 3♂ (RCB); 24 Sep 2011, 1♂ (RCB); 

Oct 2010, 2♂ (RCB); 02 Oct 2013, 1♂ (RCB); 03 Oct 2013, 

2♂ (RCB); 04 Oct 2013, 2♂ (RCB); 12 Oct 2011, 1♂ (RCB); 

12–14 Oct 1996, 1♂ (RCB); 13 Oct 2011, 1♂ (RCB); 17 Oct 

2010, 1♂ (RCB); 18 Oct 2010, 1♂ (RCB); 12 Nov 2011, 1♂

(RCB); Dec 1996, 2♂ (JHKW); 28 km Tena–Puyo Rd (El Capricho), 1°11.3′S, 77°49.9′W, 800 m, 12 Sep 2006, 1♂ (RCB); 

6.7 km SW of Puerto Napo (Yutzupino), 1°04.3′S, 77°49.8′W, 

600 m, 22 Sep 2011, 2♂ (RCB); 02 Oct 2012, 1♂ (RCB); NW 

of Misahuallí, 1°01.5′S, 77°39.4′W, 610 m, 18 Oct 2000, 1♀

(USNM); Río Pimpilala (SW of Talag), 1°04.6′S, 77°56.2′W, 

900 m, 04 Jan 2006, 1♂ (RCB); 600 m, Sep 1996, 1♂ (JHKW); 

20 km Puerto Napo–La Punta Rd, 1°03.0′S, 77°40.8′W, 500 

m, 23 Sep 2005, 1♂ (RCB); Orellana, 22 km Loreto–Payamino 

Rd, 0°34.9′S, 77°24.4′W, 825 m, 15 Jan 2012, 1♂ (RCB); nr. 

Coca (Río Napo), 0°28.0′S, 76°59.0′W, 300 m, 22 Oct 2005, 

1♂ (JHKW); upper Río Tiputini (Coca–Tiguino Rd), 300 m, 

Sep 1996, 1♂ (JHKW); Pastaza, 39.4 km Puyo–Villano Rd, 

1°25.6′S, 77°43.8′W, 750 m, 03 Jan 2016, 1♂ (RCB); 11 Sep 

2016, 1♂ (RCB); 02 Oct 2015, 2♂ (RCB); 02 Oct 2016, 2♂

(RCB); Pitirishca, 1°57.8′S, 77°52.1′W, 800 m, 07 Sep 1999, 1♂

(RCB); 20 Sep 1998, 2♂ (RCB); 24 Sep 1999, 2♂ (RCB); Sucumbios, 16 km Lumbaqui–La Troncal Rd, 0°00.8′S, 77°15.0′W, 

400–500 m, 25 Aug 2009, 1♂ (RCB). PERU, Huánuco, Panguana, 230 m, Jun 2013, 1♂ (MC); Junín, Chanchamayo (Río 

Ulcumayo), 10°59.0′S, 75°27.0′W, 1250–1370 m, 3♂ (USNM); 

Pampa Hermosa, 11°02.0′S, 75°24.0′W, 1300 m, 09 Oct 2003, 

1♂ (MUSM); 10°59.0′S, 75°25.4′W, 1230 m, 30 Sep 2008, 

1♂ (MUSM); Loreto, Contaya, 16 Oct 2008, 1♂ (CF); 24 Oct 

2008, 1♂ (CF); Iquitos, 100 m, Jul 2002, 1♂ (MC); Sept 2006, 

1♂ (MC); El Milagro, 21 km Iquitos–Nuata Rd, Oct 2010, 1♂

(CF); km 28 Iquitos–Nauta Rd, 03°59.0′S, 73°26.0′W, 180 m, 30 

Oct 2003, 1♂ (USNM); Nueva Esperanza, Feb 2012, 1♂ (CF); 

San Pablo, 04°01′S, 71°06′W, 100 m, Feb 2011, 1♂ (CF); Pévas, 

03°59.0′S, 73°26.0′W, 180 m, 01 Feb 2011, 1♂ (CF); Picuroyacu, 03°37.0′S, 71°15.0′W, 126 m, 01 Feb 2011, 1♂ (CF); 10 

Nov 2008, 1♂ (CF); Madre de Dios, 30 km SW Puerto Maldonado, 300 m, 27 Oct 1983, 1♂ (USNM); 50 km WSW Puerto 

Maldonado, 12°45.0′S, 69°35.0′W, 250 m, Sep–Nov 1992, 1♂

(USNM); Boca Río La Torre, 300 m, 26 Oct 1983, 1♀ (USNM); 

29 Oct 1983, 1♂ (USNM); Parque Manu, Pakitza, 11°55.8′S, 

71°15.3′W, 340 m, 05 Oct 1991, 1♂ (USNM); 10 Oct 1991, 1♂

(USNM); 13 Oct 1991, 1♂ (USNM); 14 Oct 1991, 1♂ (USNM); 

400 m, 1♂ (USNM); Pasco, La Salud, [1200 m], 28 May 1942, 

1♂ (LACM); San Martin, Rioja, Dec 2004, 1♂ (MC); Ucayali, 

Pucallpa, 150 m, Aug 2005, 1♂ (MC). BOLIVIA, [Santa Cruz], 

Buena Vista, 450 m, 22 Feb 1927, 1♂ (MNHN). BRAZIL, Acre, 

Bujari, Sena Madureira, 200 m, 12 Sept 2003, 1♂ (MC); Amazonas, Manicore, 18 Dec 1924, 1♂ (MNHN); Tonantins, Jul–

Sep 1880, 1♂ (BMNH); Uypiranga, Jul–Sep 1880, 1♂ (BMNH); 

Rôndonia, Alto Paraiso, Rio Candeias, 6 Aug 1998, 1♂ (MC).
NUMBER 649 • 4 3

Paraspiculatus azul Busby, Robbins, 

and Faynel, new species

FIGURES 35, 36, 51, 63, 72, 93, 96, 110, 113, 125, 126–130

Diagnosis and Description. Males of Paraspiculatus azul are distinguished from other members of the P. 

colombiensis group by (1) the lack of white scales in the distal 

part of the ventral forewing (Character 6), shared with P. lilyana (Figures 63, 64), (2) the presence of blue scales (in most 

specimens) anterior of vein Rs in the apex of the dorsal hindwing 

((Figure 72), character not coded, shared with some specimens of 

P. noemi), and (3) a lighter dorsal blue color with a very smooth, 

non-grainy appearance ((Figure 51), character not coded, shared 

with P. lilyana). The “smooth look” is a result of a uniform pattern of blue wing scales, with few intermixed black scales. This 

phenotype contrasts with P. noemi and P. sine (Figure 49) which 

have significant intermixing of black scales, especially in the distal area of the forewings. Another character that is almost always present in P. azul is a tiny patch of shining blue scales on 

the ventral forewing at the junction of veins Cu1

and Cu2 (Figure 

63). However, the diagnostic value of this trait is less important because similar blue scales are occasionally found in P. sine, 

P. colombiensis, and P. noemi. Paraspiculatus azul is separated 

from its nearest relative, P lilyana, by the lack of a broad band 

of shining blue scales anterior of the cubital vein on the ventral 

forewing and the presence of tails on the hindwing. Paraspiculatus azul is most easily confused with P. colombiensis, but is 

differentiated by its lighter blue dorsal color, lack of white scales 

along the ventral forewing costa, and more pronounced submarginal band on the ventral forewing. Mean forewing length is 

16.9 mm (SD = 0.68, n = 15) in males and 15.7 mm (SD = 1.41, 

n = 2) in the females. We illustrate the wing pattern of each sex 

(Figures 35, 36) and their genitalia (Figures 93, 110).

Female. (Figure 36) The brown female of P. azul was 

recognized by its CO1 DNA sequence. As in the male, the female 

ventral forewing lacks white scales along the costa. 

Type material.

Holotype (♂): (Figure 35) ECUADOR: Napo/ 14 km 

Tena–Puyo Road/ 1°06.7′S, 77°46.9′W 600 m/ 12 November 

2011 (Apuya)/ I. Aldas, R. C. Busby, leg. [rectangular, white, 

printed], USNM ENT 00180848 [rectangular, white, printed 

barcode label], Trap [rectangular, white, handwritten, black ink], 

GENITALIA No./ 2013: 54 ♂/ C. FAYNEL [rectangular, green, 

printed], Holotype ♂/ Paraspiculatus azul / Busby, Robbins, and 

Faynel [rectangular, red, printed]. Deposited in USNM.

Paratypes (11♂, 1♀): ECUADOR, Napo, 14 km Tena–

Puyo Road, 1°06.7′S, 77°46.9′W, 600 m, (Apuya), 07 Mar 2009, 

1♂ (USNM); 13 Sep 2005, 1♂ (RCB); 24 Sep 2005, 1♂ (USNM); 

22 Oct 2010, 1♂ (RCB); 1 km NE of Misahuallí, 1°01.5′S, 

77°39.4′W, 575 m, 09 Oct 2003, 1♂ (USNM); 12 km Tena–Puyo 

Road (Finca San Carlo), 1°05.3′S, 77°47.4′W, 600 m, Aug 2005, 

1♂ (RCB); Pimpilala, Río Jatunyacu, [1°05.3′S, 77°47.4′W], 

600 m, 14–15 Sep 1996, 1♂ (JHKW); Tena–Puyo Rd. [28 km], 

[1°05.3′S, 77°47.4′W], 800 m, 26 Oct 1996, 1♂ (JHKW); Orellana, 1.2 km Dayuma–Cononaco Rd., 0°40.7′S, 76°52.4′W, 325 

m, 06 Nov 2011, 1♂ (MECN); 16 Nov 2011, 1♂ (RCB). PERU, 

M. de Dios, Parque Manu Pakitza, 11°55.5′S, 71°15.2′W, 340 m, 

20 Oct 1991, 1♀ (USNM); Loreto, Picuroyacu, 3°37′S, 73°15′W, 

126 m, 01 Feb 2011, 1♂ (CF). 

Other Material Examined (8♂): ECUADOR, Napo, 14 

km Tena–Puyo Road, 1°06.7′S, 77°46.9′W, [600 m], (Apuya), Nov 

2011, 2♂ (CF); Morona Santiago, 53.1 km Santiago–Puerto Morona Rd., 2°55.0′S, 77°42.7′W, 210 m, 26 Sep 2015, 1♂ (RCB); 27 

Sep 2016, 1♂ (RCB); Pastaza, 39.4 km Puyo–Villano Rd., 1°25.6′S, 

77°43.8′W, 750 m, 02 Oct 2015, 1♂ (RCB); 02 Oct 2016, 2♂

(RCB). PERU, Loreto, Iquitos, 100 m, Nov 2012, 1♂ (CF). 

Etymology. Paraspiculatus azul was originally recognized by its bright blue dorsal coloration, which distinguished 

it from the darker blue hues found in other species, such as P. 

colombiensis. This name is proposed as a nonlatinized noun in 

apposition. 

Genitalia. (Figures 93, 96, 110) The genitalia of P. 

azul are typical of Paraspiculatus, but lack evident distinctive 

traits. Variation in shape of the valvae and saccus are illustrated. 

Distribution. (Figure 125) Eastern slope of the 

Andes in Ecuador and Peru.

Habitat. Wet lowland and montane forest from 126 

to 800 m elevation. 

Behavior. Males are attracted to traps baited with 

rotting fish. 

Remarks. Paraspiculatus azul and P. lilyana are 

closely related (see Phylogenetic Analyses) and sympatric in eastern Ecuador between 600 and 800 m. 

Paraspiculatus lilyana

Busby and Robbins, new species

FIGURES 37, 52, 64, 71, 94, 113, 122, 126–130

Diagnosis and Description. Males of Paraspiculatus lilyana are distinguished from others in the P. colombiensis group by (1) a broad band of shining blue scales on the 

ventral forewing anterior of the cubital vein (Character 8, Figure 

64), (2) a light dorsal blue color with a crisp black border (Figure 

52), which is shared with P. azul (this character was not coded 

because it was difficult to quantify in species with more diffuse 

borders), (3) a ventral forewing postmedian line that often lacks 

white spots near the costa (Figure 64), which is shared with no 

other species in the P. colombiensis group, and (4) light brown 

or tan scales (not white) in the distal part of the ventral forewing 

along the costa (Character 6), which is shared with P. azul. Mean 

male forewing length is 16.7 mm (SD = 0.50, n = 10). We illustrate the wing pattern (Figure 37) and male genitalia (Figure 94).

Female. The female is unknown.

Type material.

Holotype (♂): (Figure 37) ECUADOR: Napo/ 14 km 

Tena–Puyo Road/ 1°06.7′S, 77°46.9′W 600 m/ 20 September 
4 4 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

2011 (Apuya)/ Robert C. Busby, leg. [rectangular, white, printed], 

USNM ENT 00180831 [rectangular, white, printed barcode 

label], 1550 hrs/ Trap [rectangular, white, handwritten, black 

ink], GENITALIA No./ 2013: 66 ♂/ C. FAYNEL [rectangular, 

green, printed], Holotype ♂/ Paraspiculatus lilyana / Busby and 

Robbins [rectangular, red, printed]. Deposited in USNM.

Paratypes (10♂): ECUADOR, Napo, 14 km Tena–Puyo 

Road, 1°06.7′S, 77°46.9′W, 600 m, 28 Aug 2009, 1♂ (RCB); 10 

Sep 2010 1♂ (MECN); 12 Sep 2005, 1♂ (USNM), 1♂ (RCB); 

04 Oct 2013, 1♂ (RCB); 13 Oct 2011, 1♂ (RCB); 12 km Tena–

Puyo Road, 1°05.3′S, 77°47.4′W, 600 m, (Finca San Carlo), 14 

Sep 2004, 1♂ (USNM); Chichicorrumi nr. Jatun Sacha, 450 m, 

Sep 1993, 1♂ (JHKW); Pastaza, Pitirishca, 800 m, 1°57.8′S, 

77°52.1′W, 10 Sep 2000, 1♂ (RCB); 19 Sep 2000, 1♂ (USNM). 

Etymology. We name this beautiful species P. lilyana in honor of Lily Elizabeth Hughes, granddaughter of Robert 

C. Busby. The name is a noun in apposition. 

Genitalia. (Figure 94) The genitalia of P. lilyana are 

typical of Paraspiculatus, but lack evident distinctive traits.

Distribution. (Figure 122) The eastern slope of 

the Andes in Ecuador.

Habitat. Paraspiculatus lilyana inhabits wet montane forest from 450–800 m elevation. 

Behavior. Males are attracted to traps baited with 

rotting fish. 

Remarks. Paraspiculatus lilyana and P. azul are 

closely related and sympatric in eastern Ecuador at 450–600 

m elevation. Paraspiculatus lilyana and P. orobia share a broad 

band of shining blue scales on the ventral forewing anterior to 

the cubital vein (Character 8), but this represents convergence 

according to the phylogenetic analyses.

Paraspiculatus noemi

Busby and Robbins, new species

FIGURES 38–40, 73, 95, 97, 111, 113, 122, 126–130

Diagnosis and Description. Males of Paraspiculatus noemi are distinguished from the other species in the 

P. colombiensis group by the predominance of dark wing scales 

at the base of the dorsal hindwing costa (Character 13, Figure 

73). This pattern contrasts with the other species in the group 

in which a majority of the scales are normally bright blue (Figures 71, 72). The scales in P. noemi are the same size and shape 

as the “regular” blue wing scales, but are predominately black, 

occasionally purple. The overall appearance of this area is dark 

and is easily distinguished from the other species in the P. colombiensis group. These dark scales differ from both the dark gray, 

iridescent scales in the P. orocana species group and the gray/

brown scales in P. orobia and P. cosmo. Another diagnostic factor which helps to separate P. noemi from the other species in the 

P. colombiensis species group is the bold white triangle which is 

formed by a large patch of white scales (Figure 38) at the median 

of the ventral forewing along with the three white/light-blue spots 

at the anterior of the postmedian line. This basic pattern is also 

shared by P. colombiensis and P. emma, but tends to be more 

conspicuous and consistent in P. noemi, especially in specimens 

from western Ecuador and Colombia. There is more intraspecific 

variation in the amount of white scales in both P. colombiensis

and P. emma, although occasional specimens have a well-formed 

white triangle similar to those in P. noemi. Finally, the tails in 

Ecuadorian P. noemi are short, a character that is unique within 

the P. colombiensis species group. Unfortunately, this trait is of 

limited practical value because tail length is geographically variable in P. noemi and because the tails of worn specimens (of other 

“tailed species”) may be broken or absent. Mean forewing length 

is 19.0 mm (SD = 0.74, n = 14) in males and 18.1 mm (SD = 0.0, 

n = 1) in the female from Costa Rica. We illustrate the wing pattern (Figures 38–40) and genitalia (Figures 95, 111) of P. noemi.

Female. (Figure 40) The female is associated by geographic distribution.

Type material.

Holotype (♂): (Figure 38) ECUADOR: Pichincha/ 20 

km Pacto–Guayabillas Road/ 0°11.6′N, 78°51.5′W 900 m/ 14 

July 2011/ I. Aldas, R. C. Busby, leg. [rectangular, white, printed], 

USNM ENT 00180855 [rectangular, white, printed barcode 

label], GENITALIA No./ 2013: 60 ♂/ C. FAYNEL [rectangular, 

green, printed], Holotype ♂/ Paraspiculatus noemi/ Busby and 

Robbins [rectangular, red, printed]. Deposited in USNM.

Paratypes (13♂): ECUADOR, Pichincha, 21 km 

Pacto–Guayabillas Road, 0°11.2′N, 78°51.8′W, 875 m, 28 May 

2008, 2♂ (RCB); Esmeraldas, 12 km Lita–San Lorenzo Road, 

0°53.1′N, 78°30.9′W, 850 m, Río Chuchuví, Mar 2003, 2♂

(RCB); 25 Mar 2011, 1♂ (USNM); 27 Mar 2011, 1♂ (USNM); 

May 2003, 1♂ (MECN); Jun 2004, 1♂ (RCB); Jul 2002, 2♂

(JHKW); Aug 2008, 1♂ (MECN); Calderón, 01.06 N, 78.42 W, 

[100 m], 18 Nov 2014, 1♂ (CF); Carchi, Río Sabalera, [0°11.2′N, 

78°51.8′W], 500 m, Aug 2014, 1♂ (RCB).

Other Material Examined (4♂, 1♀): MEXICO, Oaxaca, Sierra Juarez (Gulf Slope), 2500 ft, April 1992, 1♂ (USNM). 

COSTA RICA, Heredia, La Selva, 10°26′N, 84°01′W, 50–100 m, 

11 Jul 1993, 1♀ (USNM). PANAMA, Veraguas, 1♂ (BMNH). 

COLOMBIA, Cauca, Juntas, [2°27′N, 76°36′W], fin [Aug or 

later] 1897–Jan 1898, 1♂ (BMNH). VENEZUELA, Merida, 

Briceno, 1♂ (BMNH). 

Etymology. We name P. noemi in honor of Noemi 

Patiño Artega, wife of Ismael Aldas Villafuerte, who made significant contributions to the study of Paraspiculatus through his 

field work at the type locality and elsewhere in Ecuador. The 

name is a noun in apposition. 

Variation. In contrast to specimens from western 

Ecuador, the three P. noemi from Mexico and Central America 

have fewer white scales along the ventral costa of the forewing. 

Two of these three specimens have longer tails than those from 

Ecuador. The Mexican male has more blue at the apex of the 

dorsal hindwing anterior to vein Rs than the Ecuadorian males. 

Although CO1 DNA sequences from the Mexican male cluster 

with those from western Ecuador, they are also slightly divergent 

(Figure 113). 
NUMBER 649 • 4 5

Genitalia. (Figures 95, 97, 111) The genitalia of P. 

noemi are typical of Paraspiculatus, but lack evident distinctive 

traits. Variation in the spiculate pad is illustrated.

Distribution. (Figure 122) Transandean. Mexico 

throughout Central America to the western slope of the Andes in 

northwestern Ecuador and to northern Venezuela. 

Habitat. Wet montane forest from 50–800 m 

elevation.

Behavior. Males are occasionally attracted to traps 

with rotting fish. Two individuals from Ecuador were attracted 

to leaves baited with fish (vouchers in RCB).

Remarks. The Mexican and Central American part 

of the range of P. noemi is representative of the remarkable rarity 

of most Paraspiculatus in museum collections. In the approximately 135 years since Godman and Salvin (1887–1901) first 

reported it in Panama (originally identified as Thecla orobia), 

only one male and one female P. noemi have subsequently been 

collected in Mexico and Central America. 

As documented, P. noemi varies geographically. That the 

Mexican and Central American specimens represent a species distinct from P. noemi—rather than a geographic variant—is a viable 

alternate hypothesis, but we lack sufficient supporting evidence.

Paraspiculatus noemi and P. colombiensis are not sister 

species on the consensus cladogram shown below in Phylogenic 

Analyses. Male wing pattern differences between them are consistent, and male P. colombiensis (mean wing size 17.4 mm) are 

statistically smaller than male P. noemi (t test, unequal variances, 

t = 5.13, df = 43, p < 0.0001). Despite these differences, phylogenetic resolution in the P. colombiensis group is not robust, 

and we would not be surprised if further molecular data showed 

that these two species are geographically allopatric phylogenetic 

sisters.

PHYLOGENETIC ANALYSES

Equal-weight parsimony for the morphological matrix 

analysis yielded four trees of 57 steps. Implied-weight parsimony 

analysis (K = 1000, 100, 10, 3) resulted in one tree, which is also 

one of the equal-weight trees (Figures 126, 127). Since it is the 

shortest tree in all analyses, we consider it to be the best estimate 

of phylogeny based on morphology. Bootstrap support values 

are given in Figure 126. Paraspiculatus monophyly was supported by five synapomorphies, but the only one unique to Paraspiculatus within the Eumaeini is the spiculate pad (Character 

23). It is appropriate that the genus was named for this feature. 

Analyses of the molecular data set yielded a variety of trees. 

We derived the best maximum likelihood tree in Garli (Figure 

128). There were 20 trees of 271 steps in the equal-weight parsimony analysis, two trees (271 equal-weight steps) with the 

implicit-weight parameter K = 1000 or 100, two trees with K = 

10 (both 272 steps), and one tree with K = 3 (276 steps). 

Analysis of the combined morphology and molecular matrix resulted in 40 trees of 334 steps in equal-weight parsimony, 

one 334-step tree in implied-weight parsimony (K = 1000, 100, 

and 10), and one 335-step tree in implied-weight parsimony (K 

= 3) (Figure 129).

Despite the number of trees resulting from the analyses 

of the molecular and morphology data sets, some results were 

consistent. Each of the 17 Paraspiculatus species for which we 

had molecular data (an average of six specimens per species) 

was monophyletic in the resulting trees. The six-species P. colombiensis complex (P. emma, P. sine, P. colombiensis, P. azul, 

P. lilyana, and P. noemi) was monophyletic in all analyses. The 

three-species P. orocana species complex (P. grande, P. honor, 

and P. orocana) was monophyletic in all analyses. The twospecies P. catrea species complex (P. catrea and P. vossoroca) 

was monophyletic in all analyses of morphology (molecular 

data are lacking for P. vossoroca). Finally, the P. orobia species complex (P. orobia and P. cosmo) was monophyletic in all 

analyses. These multispecies lineages are reflected in the classification of species complexes.

DISCUSSION

Morphology, Molecules, and Female Choice

Eumaeini systematics has historically been based on three 

major morphological character sets: (1) wing pattern and shape, 

(2) genitalic structures, and (3) male secondary sexual traits (e.g., 

Ziegler, 1960; Clench, 1961, 1964; Nicolay, 1971a,b; Johnson, 

1989; Robbins, 1991). These structures were used because they 

are frequently variable interspecifically, relatively conspicuous, 

and easily scored. Other characters, such as those of the head 

and legs, infrequently provided useful taxonomic information 

(Eliot, 1973; Robbins, 1986, 1991; Robbins and Venables, 

1991). Characters of the immatures were rarely used because 

of the paucity of preserved immatures and because scoring larval characters and assessing their variation was labor intensive 

(Ballmer and Pratt, 1988). This pattern of morphological character usage in the Eumaeini remains largely unaltered (e.g., Duarte 

and Robbins, 2009, 2010; Faynel et al., 2012; Robbins et al., 

2012; Robbins and Busby, 2015).

The morphological, ecological, and behavioral traits that 

adult females use to recognize and reject courting non-conspecific 

males are the bases of reproductive isolation among sympatric 

species. Direct evidence that lycaenid females can use each of the 

three major morphological characters sets listed in the previous 

paragraph to choose between conspecific and non-conspecific 

males is limited. For wing pattern and shape, female lycaenids can distinguish conspecific males by wing pattern details 

(Fordyce et al., 2002). For genitalic structures, sexual selection 

is the primary explanation for evolving differences (e.g., WestEberhard, 1983; Eberhard, 1985, 2010; Hosken and Stockley, 

2004; Simmons, 2014), but direct evidence for sexual selection 

among lycaenids is lacking. For male secondary sexual structures, pheromones dispersed by lycaenid wing androconia are 
4 6 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 126. Top of a most parsimonious tree for Paraspiculatus based on morphological characters (Table 2), showing where characters 

evolved (open circles are homoplastic changes). This was the only most parsimonious equal-weight and implied-weight tree with a variety 

of values for the parameter K from 3 to 1,000. Bootstrap values noted.
NUMBER 649 • 4 7

used during courtship (Lundgren and Bergström, 1975; Ômura 

et al., 2013). 

The indirect evidence that female Eumaeini routinely use at 

least one of the three major morphological character sets to determine male conspecificity is accumulating. When wing patterns 

are interspecifically uniform, such as within most of the genus 

Oenomaus, each species has conspicuously different male genitalia (Faynel et al., 2012). Further, in those few cases within Oenomaus where wing pattern has changed markedly, change in the 

male genitalia is slight (C. Faynel et al., unpublished data). When 

FIGURE 127. Bottom of a most parsimonious tree for Paraspiculatus based on morphological 

characters (Table 2), showing where characters evolved (open circles are homoplastic changes). 

This was the only most parsimonious equal-weight and implied-weight tree with a variety of 

values for the parameter K from 3 to 1,000. Bootstrap values noted.
4 8 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 128. Maximum likelihood tree using the molecular data set. Numbers following the taxa are the number 

of specimens of that species. All species were monophyletic. Bootstrap values noted.
NUMBER 649 • 4 9

FIGURE 129. Most parsimonious implied-weight (default K = 3) tree based on the combined morphological and molecular data set. The taxa in red are sympatric in the upper Amazon Basin below 1,250 m 

elevation. Bootstrap values noted.
5 0 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

male genitalic structures are interspecifically uniform in the Eumaeini, such as within the genus Arcas (Nicolay, 1971a), each 

species has a unique set of male secondary sexual traits as well 

as wing pattern/shape differences (Robbins et al., 2012). Finally, 

when sexual structures are uniform, as in the Electrostrymon 

guzanta species complex (Thompson and Robbins, 2016), males 

of each species can be distinguished by their wing pattern.

Analysis of DNA sequences provides an independent test 

of the traditional morphological character sets in the Eumaeini 

for biased selection and misinterpretation, especially as it relates 

to species delimitation. Mitochondrial CO1 barcode sequences 

have been widely adopted for species delimitation (Janzen et al., 

2009). Discordance between species delimited by morphology 

and by CO1 sequences in the Lepidoptera is generally in the 10–

15% range (Wiemers and Fiedler, 2007; Zahiri et al., 2014). The 

recent introduction of the barcode index number (BIN) (Ratnasingham and Hebert, 2013) makes the identification and measurement of discordance easier. 

We determined species limits in Paraspiculatus based on 

male wing pattern and shape. We then determined species limits 

for each of the 17 Paraspiculatus species (Figure 113) for which 

we had data based on the clustering of CO1 barcode sequences. 

The morphological species determinations agreed with BINs in 

92 of 95 males (3.2% discordance). Intraspecific CO1 sequences 

for P. orobiana, P. apuya, and P. noemi varied more than was 

recognized by the refined single linkage analysis that is used to 

determine BINs (Ratnasingham and Hebert, 2013). Geographical variation in DNA sequences is a likely explanation for this 

discordance. For each of the three species, the second BIN was 

assigned to a male from a different geographic area. 

Paraspiculatus adds another piece of indirect evidence that 

some female eumaeines use wing pattern to recognize conspecific 

males. Paraspiculatus primary male sexual structures vary little 

interspecifically. Secondary male sexual structures are lacking. 

Presumably, females do not use sexual structures to recognize 

conspecific males. Alternatively, variation in male wing pattern 

is concordant with species delimitation by CO1 barcode sequences, so the hypothesis that females recognize males by wing 

pattern is plausible. 

Documenting Diversity

Robbins (2004a) noted that more than 25% of neotropical eumaeine species represented in museum collections were 

undescribed. In subsequent taxonomic revisions of Timaeta

Johnson, Kruse, and Kroenlein, Thepytus Robbins, Lathecla

Robbins, Porthecla Robbins, and Oenomaus, more than 50% 

of the recognized species were newly described (Robbins and 

Busby, 2008, 2015; Robbins et al., 2010b; Faynel, 2006b, 2008; 

Faynel and Moser, 2008; Faynel et al., 2011, 2012). Paraspiculatus continues this trend, with 10 of 19 species treated newly 

described in this paper. Although butterflies are considered to be 

the best documented large group of insects (Robbins and Opler, 

1997), there is still a significant proportion of undocumented 

diversity.

We noted in the introduction that a revision of Paraspiculatus was possible only because new sampling techniques using fish 

bait resulted in a fivefold increase in the number of specimens 

available for study. It would be hard to overemphasize the need 

for expanded and more intensive collecting of eumaeines, especially in South America. 

Rotting Fish, Stratification, and Nutrition

Rotting fish in a trap 10 m above the ground is not a “natural” food source for hairstreak butterflies. This collecting method 

has been used primarily to increase sample sizes of otherwise rare 

species and not as a way to study biology. Despite these limitations, the attraction to rotting fish is related to various biological 

aspects of Paraspiculatus and other eumaeines. The significance 

of adult male butterfly food sources has been discussed in the Lycaenidae and Riodinidae (Beck et al., 1999; Hall and Willmott, 

2000), but needs to be expanded to account for variation in the 

attractiveness of rotting fish as a eumaeine bait depending upon 

geographic area, taxon, and sex. The purpose of this section is 

to summarize what we know about the attraction of adult eumaeines, especially Paraspiculatus, to rotting fish and to discuss 

how these observations may relate to Paraspiculatus biology.

The attractiveness of rotting fish bait to eumaeines is geographically variable. On the eastern slope of the Andes, especially 

in Ecuador, it has been effective at 200 to 1,800 m elevation. On 

the western slopes of the Ecuadorian Andes, it has been much less 

effective. In the mountains of northern Venezuela and in French 

Guiana, it did not attract eumaeines. We do not know whether 

these results are due to geographic variation in naturally occurring sources of adult nutrition or to other geographically variable factors, such as availability of larval food. Regardless, this 

geographic variability in the attractiveness of rotting fish to eumaeines is one reason for the limited number of records of Paraspiculatus in areas such as the northern parts of South America. 

Trapping adult eumaeines with fish-baited traps is different 

from trapping other adult butterflies. Nymphalidae attracted to 

traps baited with fish usually land on the bait and fly up into the 

trap. Adult eumaeines apparently fly downward toward the fish 

(Robbins, 2001), and land on the top and sides of the trap, not 

on the bait dish. If the netting has rotting fish on it, the landed 

butterfly extends its proboscis, presumably feeding. If the trap is 

disturbed, it flies away. To sample these adult lycaenids, the trap 

has to be lowered carefully to the ground, where the butterflies 

are caught on the trap netting. 

The efficacy of rotting fish bait for eumaeines depends upon 

taxon and sex. In Paraspiculatus, Ocaria Clench, and Penaincisalia Johnson, the ratio of baited males to females is almost 1:0 

(the few female exceptions in Paraspiculatus have been noted). 

In Ignata Johnson, Strephonota Johnson, Austin, Le Crom, and 

Salazar, Siderus Kaye, and Erora Scudder, the ratio of baited individuals is reversed, close to 0:1 (again with a few exceptions, 

this time male). In the majority of Eumaeini, however, both sexes 

are attracted to rotting fish, although the sex ratio is variable 

from species to species. In contrast, the majority of riodinids 
NUMBER 649 • 5 1

attracted to fish bait in the same area were males (Hall and Willmott, 2000). Finally, in some eumaeine genera, such as Timaeta

and Micandra Staudinger, neither sex is attracted to rotting fish. 

Adult butterflies and other animals may spend most of their 

time in a restricted stratum of the forest (cf. DeVries, 1988 and 

included references). For slow-flying butterflies, flight height 

above the ground can be recorded and has been shown to be a 

function of wing pattern and/or size (Papageorgis, 1975; Medina 

et al., 1996). For faster-flying butterflies, two methods have been 

used. A tower in tropical forest in Africa allowed observation 

and trapping of butterflies at different forest strata (Jackson, 

1961). For Lycaenidae, this was apparently the first time that it 

was demonstrated that some species are frequent and active in 

the higher forest strata, but not at ground level. In other words, 

Lycaenidae may be rare to butterfly collectors on the ground, but 

are not necessarily rare in the forested habitats. Second, traps 

placed at the ground and in the upper forest strata may attract 

different sets of species. Those caught primarily in ground-level 

traps are presumed to be primarily active at ground level, not 

in the upper strata, and vice versa (DeVries, 1988; DeVries and 

Walla, 2001; DeVries et al., 2012; Alexander and DeVries, 2012; 

Fordyce and DeVries, 2016). Exceptions to this presumption 

were summarized in DeVries and Walla (2001). 

Paraspiculatus are rarely encountered in the forest understory in the eastern Andes, as determined by the paucity of netcollected individuals in museums. Less than 2% of the specimens 

in our study series from eastern Ecuador were collected with a 

net. Although we used rotting fish poured on leaves near ground 

level as well as in traps hung 7 to 20 m above the ground, the latter method was more than twice as productive in attracting male 

Paraspiculatus. Controlled experiments with paired traps near 

ground level and in the forest upper strata might be used to test 

the hypothesis that adult Paraspiculatus are rare in collections 

because they spend most of their time in the upper forest strata.

DeVries (1988:99) wrote “Excluding those species where 

males visit wet sand or plant material for non-nutritional resources . . , any tropical forest community of butterflies can be 

divided into two adult feeding guilds: those species that obtain 

the bulk of their nutritional requirements from flower nectar (all 

Papilionidae, Pieridae, Lycaenidae, Riodinidae and some Nymphalidae), and those species that feed upon the juices of rotting 

fruits, fermenting sap, or animal waste (several subfamilies of 

the Nymphalidae . . .).” This guild partition needs modification, 

at least for Riodinidae (Hall and Willmott, 2000) and Lycaenidae in eastern Ecuador. Whereas many neotropical forest eumaeine lycaenids nectar at flowers seasonally, especially when 

certain species of Cordia (Boraginaceae) are flowering (Opler et 

al., 1975; Faynel, 2003, 2005, 2006a, 2010), it is premature to 

conclude that flower nectar makes up the bulk of adult eumaeine 

nutritional requirements. In our experience, adult eumaeines in 

wet forest feeding on flowers is a rarely observed occurrence. 

Adults of a Colombian eumaeine species feed on exudates from 

Hemiptera and on plant fluids oozing from wounds made by 

sucking insects (Heredia and Robbins, 2016). Nonfloral adult 

feeding behavior is widely observed in miletine lycaenids 

(Lohman and Samarita, 2009 and included citations) and has 

recently been noted in North American eumaeines (Wagner and 

Gagliardi, 2015; Gagliardi and Wagner, 2016). Finally, many lycaenid adults are attracted to and eat liquefied decaying animals, 

such as rotting fish (and cow’s blood in one instance). The limited previous observations of this phenomenon may be due to 

the geographical and seasonal variation in its occurrence or due 

to the observation that lycaenid adults do not fly into traps, but 

land on the tops and sides. As noted, unless the trap is lowered 

with great care, such adults fly away and would not be recorded 

as being attracted to fish as bait (fish-bait). 

Rotting fish attracts a significant proportion of the lycaenid 

fauna in eastern Ecuador. It may attract members of either sex, 

unlike previous work on non-nectar attractants (Beck et al., 1999; 

Hall and Willmott, 2000). Perhaps the most interesting and unusual observation is the intergeneric variation in sex ratio of adult 

eumaeines attracted to fish-bait. Adult male butterflies devote up 

to 10% of their adult weight to spermatophores transferred to females during mating (Rutowski et al., 1983; Caballero-Mendieta 

and Cordero, 2013). It might be reasonable to hypothesize that 

males in genera, such as Paraspiculatus, transfer the nutrition ingested to females in spermatophores. Alternately, in genera where 

females predominate at fish-bait, we might expect less nutritional 

transfer in spermatophores. Intergeneric variation in sex ratio of 

adult eumaeines attracted to fish-bait suggests that eumaeine reproductive biology is worth investigating. 

Distribution

Based on current data, most Paraspiculatus species have 

restricted geographic distributions and occur over a relatively 

narrow range of elevations (Figure 130). Only P. orobia occurs 

in more than one of the major biogeographic regions of Brown 

(1982). Only three species (P. orobia, P. colombiensis, P. catrea) 

occur over an elevation range of more than 1,000 m (Figure 

130). The wide elevational range of P. transvesta is an artifact of 

imprecise data, as noted in its species account. However, the use 

of traps baited with rotting fish in areas other than Ecuador may 

significantly expand current geographic distributions. 

The most widespread Paraspiculatus species are those that 

occur in the lowlands, as seems generally true for Eumaeini 

(Robbins et al., 2010b; Robbins and Busby, 2015). Paraspiculatus elis, P. orobia, P. noemi, and P. colombiensis occur throughout their respective biogeographic region (Figures 114, 117, 122, 

124) and inhabit lowlands. However, the distributions of these 

species are restricted when compared with species in some eumaeine genera, such as Rekoa Kaye in which four of the seven 

species range from Mexico to southern Brazil (Robbins, 1991). 

Sympatry and Diversification

In the Introduction, we noted the high incidence of sympatry among Paraspiculatus species in the upper Amazon under 

1,250 m elevation (10 of 19 species). The set of sympatric Paraspiculatus species could result from in situ diversification in the 
5 2 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

upper Amazon, from the diversification of clades, each with one 

Amazonian representative, or a combination of these processes. 

Although a comprehensive explanation for the high frequency 

of sympatry in Paraspiculatus is beyond the scope of this paper, 

revising the genus is a first step. 

According to our results, the 10 Paraspiculatus species that 

occur in the upper Amazon under 1,250 m elevation represent 

all three possibilities. Three species (P. elis, P. apuya, and P. orobiana) belong to single species complexes because they have 

no evident very close relative within the genus. The P. orobia

FIGURE 130. Biogeographic regions and elevation zones for each Paraspiculatus species. Paraspiculatus orobia also occurs in the northern part 

of the Atlantic Region. Green bars represent lowland species, violet are lower montane species, and red are montane species. Paraspiculatus 

catrea is an elevational generalist over its entire range, but this may be an artifact because it occurs at higher elevations in the northern part of 

its range and at lower ones in the southern part.
NUMBER 649 • 5 3

and P. orocana species groups fit the traditional “superspecies” 

concept of Mayr (1963), in which no two component species 

are sympatric. Each of these superspecies has one species in the 

upper Amazon Basin under 1,250 m elevation (P. orobia and 

P. orocana). Finally, five of the six species in the P. colombiensis

species complex (P. colombiensis, P. lilyana, P. azul, P. emma, and 

P. sine) are sympatric at 800 m elevation in eastern Ecuador. This 

species complex is the main reason for the original observation 

of a high incidence of sympatry in the upper Amazon Basin. In 

situ diversification is usually examined on ecological or physical 

islands (Simon et al., 2009; Gómez-Díaz et al., 2012; Pante et 

al., 2012; Blaimer et al., 2015), but perhaps it is also relevant to 

biogeographic areas. A comprehensive explanation for the high 

frequency of sympatry in Paraspiculatus would undoubtedly 

focus on this lineage.

For allowing us to examine specimens in collections under their care, we are grateful 

to Mirna Casagrande (DZUP), David Grimaldi (AMNH), Blanca Huertas (BMNH), 

Gerardo Lamas (MUSM), Olaf Mielke (DZUP), Jacques Pierre (MNHN), and 

Wolfgang Nassig (SMF). For giving us access to their private collection and providing 

specimens for DNA sequencing, we thank Jason Hall and Keith Willmott. For providing 

photographs, specimens, or other data, we are grateful to Pierre Boyer, Michael Büche, 

Mauro Costa, Will and Gill Carter, Jose Cerdeña, Jean-Yves Gallard, Kim Garwood, 

Jean-François Le Crom, Jean-Claude Petit, Thibault Rosant, and Rob Westerduijn. For 

illustrative and technical help at Smithsonian, we are indebted to Karie Darrow, Brian 

Harris, and Vichai Malikul. We thank Gerardo Lamas for information on the history of 

collecting in Balsapuerto, Peru. We thank two anonymous reviewers, who made numerous suggestions and corrections, for their time-consuming contributions. For help dealing 

with the BOLD database and sequencing at the Canadian Centre for DNA Barcoding, we 

thank Jeremy DeWaard, Vlad Dinca, Evgeny V. Zakharov, and their many coworkers. 

For facilitating specimen loans among the authors, we thank Helena Romanowski and 

Vanessa Schaeffer at the federal university (UFRGS) in Porto Alegre, Brazil. For supporting the project “Diversidad y Biologica de Lepidopteros en el Ecuador,” we thank 

the Museo Ecuatoriano de Ciencias Naturales, especially Santiago Villamarín-Cortez, 

Curador de Invertebrados, who has provided valuable assistance in obtaining permits for 

our fieldwork in Ecuador. Faynel expresses his appreciation to the Smithsonian Institution Short-Term Visitor Program, which facilitated this project. To all, we are grateful. 

Acknowledgments

TABLE A.1. The CO1 sequence samples listed alphabetically by genus, species, sex, 

and country of origin, with collection voucher numbers for the U.S. National Museum (USNM) and the private collection of Christophe Faynel (CF); and online BOLD 

database–assigned process identification numbers.

Species, sex, country Voucher number BOLD ID

Mithras nautes, f, French Guiana CF-LYC-162 NLYCA162-12

Mithras nautes, m, Ecuador USNM ENT 00180865 EUM070-13 

Mithras nautes, m, Peru CF-LYC-161 NLYCA161-12

Paraspiculatus apuya, m, Brazil USNM ENT 00180888 EUM081-13 

Paraspiculatus apuya, m, Ecuador USNM ENT 00180817 EUM027-13

Paraspiculatus apuya, m, Ecuador USNM ENT 00180818 EUM028-13 

Paraspiculatus apuya, m, Ecuador USNM ENT 00181672 EUM116-14

Paraspiculatus apuya, m, Ecuador USNM ENT 00181767 EUM301-15

Paraspiculatus azul, f, Peru USNM ENT 00180894 EUM087-13 

Paraspiculatus azul, m, Ecuador USNM ENT 00180848 EUM052-13 

Paraspiculatus azul, m, Ecuador USNM ENT 00180871 EUM056-13

Paraspiculatus azul, m, Ecuador USNM ENT 00180878 EUM102-14 

Paraspiculatus azul, m, Ecuador USNM ENT 00181669 EUM113-14

Paraspiculatus azul, m, Ecuador USNM ENT 00181671 EUM115-14

Paraspiculatus azul, m, Ecuador USNM ENT 00181700 EUM118-14

Paraspiculatus azul, m, Ecuador USNM ENT 00181701 EUM119-14

Paraspiculatus azul, m, Ecuador USNM ENT 00181702 EUM120-14

Paraspiculatus azul, m, Ecuador USNM ENT 00181703 EUM121-14

Paraspiculatus azul, m, Ecuador USNM ENT 00181704 EUM122-14 

Paraspiculatus azul, m, Ecuador USNM ENT 00181706 EUM124-14

Paraspiculatus azul, m, Peru CF-LYC-158 NLYCA158-12

Paraspiculatus catrea, f, Brazil USNM ENT 00181667 EUM111-14

Paraspiculatus catrea, f, Brazil USNM ENT 00181668 EUM112-14 

Paraspiculatus catrea, m, Brazil USNM ENT 00179443 EUM089-13

Appendix A: CO1 DNA 

Sequence Information
5 8 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE A.1. (Continued)

Species, sex, country Voucher number BOLD ID

Paraspiculatus colombiensis, f, Ecuador USNM ENT 00179432 EUM074-13

Paraspiculatus colombiensis, f, Ecuador USNM ENT 00180840 EUM045-13

Paraspiculatus colombiensis, f, Ecuador USNM ENT 00180877 EUM036-13

Paraspiculatus colombiensis, f, Ecuador USNM ENT 00181768 EUM302-15

Paraspiculatus colombiensis, f, Ecuador USNM ENT 00181769 EUM303-15 

Paraspiculatus colombiensis, m, Ecuador USNM ENT 00180841 EUM046-13

Paraspiculatus colombiensis, m, Ecuador USNM ENT 00180842 EUM047-13

Paraspiculatus colombiensis, m, Ecuador USNM ENT 00180843 EUM048-13

Paraspiculatus colombiensis, m, Ecuador USNM ENT 00180844 EUM050-13

Paraspiculatus colombiensis, m, Ecuador USNM ENT 00180847 EUM051-13

Paraspiculatus colombiensis, m, Ecuador USNM ENT 00180872 EUM049-13

Paraspiculatus colombiensis, m, Ecuador USNM ENT 00181670 EUM114-14 

Paraspiculatus colombiensis, m, Peru CF-LYC-157 NLYCA157-12

Paraspiculatus colombiensis, m, Peru CF-LYC-159 NLYCA159-12

Paraspiculatus cosmo, f, Ecuador USNM ENT 00180806 EUM018-13

Paraspiculatus cosmo, m, Ecuador USNM ENT 00180798 EUM012-13

Paraspiculatus cosmo, m, Ecuador USNM ENT 00180799 EUM013-13

Paraspiculatus cosmo, m, Ecuador USNM ENT 00180800 EUM014-13 

Paraspiculatus cosmo, m, Ecuador USNM ENT 00180804 EUM016-13

Paraspiculatus cosmo, m, Ecuador USNM ENT 00180805 EUM017-13

Paraspiculatus cosmo, m, Peru CF-LYC-156 NLYCA156-12

Paraspiculatus elis, m, Ecuador USNM ENT 00180863 EUM069-13

Paraspiculatus elis, m, Peru CF-LYC-163 NLYCA163-12

Paraspiculatus emma, m, Ecuador USNM ENT 00180807 EUM019-13

Paraspiculatus emma, m, Ecuador USNM ENT 00180808 EUM020-13

Paraspiculatus emma, m, Ecuador USNM ENT 00180809 EUM021-13

Paraspiculatus emma, m, Ecuador USNM ENT 00180811 EUM022-13

Paraspiculatus emma, m, Ecuador USNM ENT 00180812 EUM023-13

Paraspiculatus emma, m, Ecuador USNM ENT 00180813 EUM024-13

Paraspiculatus emma, m, Ecuador USNM ENT 00180815 EUM025-13 

Paraspiculatus grande, f, Panama USNM ENT 00179438 EUM078-13

Paraspiculatus grande, m, Ecuador CF-LYC-425 NLYCA425-13

Paraspiculatus grande, m, Ecuador USNM ENT 00180788 EUM002-13

Paraspiculatus grande, m, Ecuador USNM ENT 00180789 EUM003-13

Paraspiculatus grande, m, Ecuador USNM ENT 00180790 EUM004-13

Paraspiculatus grande, m, Ecuador USNM ENT 00180791 EUM005-13

Paraspiculatus grande, m, Ecuador USNM ENT 00180792 EUM006-13

Paraspiculatus grande, m, Ecuador USNM ENT 00180793 EUM007-13

Paraspiculatus hannelore, f, Brazil USNM ENT 00181663 EUM107-14

Paraspiculatus hannelore, m, Brazil USNM ENT 00181664 EUM108-14

Paraspiculatus hannelore, m, Brazil USNM ENT 00181665 EUM109-14

Paraspiculatus hannelore, m, Brazil USNM ENT 00181666 EUM110-14

Paraspiculatus honor, f, Ecuador USNM ENT 00180862 EUM068-13

Paraspiculatus honor, m, Ecuador USNM ENT 00181673 EUM117-14 
NUMBER 649 • 5 9

TABLE A.1. (Continued)

Species, sex, country Voucher number BOLD ID

Paraspiculatus lilyana, m, Ecuador USNM ENT 00180831 EUM037-13

Paraspiculatus lilyana, m, Ecuador USNM ENT 00180832 EUM038-13

Paraspiculatus lilyana, m, Ecuador USNM ENT 00180833 EUM039-13

Paraspiculatus lilyana, m, Ecuador USNM ENT 00180834 EUM040-13

Paraspiculatus lilyana, m, Ecuador USNM ENT 00180835 EUM042-13

Paraspiculatus lilyana, m, Ecuador USNM ENT 00180838 EUM043-13

Paraspiculatus lilyana, m, Ecuador USNM ENT 00180839 EUM044-13

Paraspiculatus noemi, m, Ecuador USNM ENT 00180855 EUM063-13

Paraspiculatus noemi, m, Ecuador USNM ENT 00180857 EUM064-13

Paraspiculatus noemi, m, Ecuador USNM ENT 00180858 EUM065-13

Paraspiculatus noemi, m, Ecuador USNM ENT 00180859 EUM066-13

Paraspiculatus noemi, m, Ecuador USNM ENT 00180860 EUM067-13

Paraspiculatus noemi, m, Mexico USNM ENT 00180898 EUM093-13 

Paraspiculatus oroanna, m, Ecuador CF-LYC-151 NLYCA151-12

Paraspiculatus oroanna, m, Ecuador USNM ENT 00180787 EUM001-13 

Paraspiculatus oroanna, m, Peru CF-LYC-152 NLYCA152-12

Paraspiculatus orobia, f, French Guiana CF-LYC-150 NLYCA150-12

Paraspiculatus orobia, f, Peru USNM ENT 00181972 EUM380-15

Paraspiculatus orobia, f, Peru USNM ENT 00180889 EUM082-13

Paraspiculatus orobia, m, Brazil USNM ENT 00179431 EUM073-13

Paraspiculatus orobia, m, Ecuador USNM ENT 00180819 EUM029-13

Paraspiculatus orobia, m, Ecuador USNM ENT 00180821 EUM030-13

Paraspiculatus orobia, m, Ecuador USNM ENT 00180823 EUM031-13

Paraspiculatus orobia, m, Ecuador USNM ENT 00180825 EUM032-13

Paraspiculatus orobia, m, Ecuador USNM ENT 00180826 EUM033-13

Paraspiculatus orobia, m, Ecuador USNM ENT 00181997 EUM188-14

Paraspiculatus orobia, m, Ecuador USNM ENT 00181998 EUM189-14

Paraspiculatus orobia, m, Ecuador USNM ENT 00181999 EUM190-14

Paraspiculatus orobia, m, Peru CF-LYC-160 NLYCA160-12

Paraspiculatus orobiana, f, Peru USNM ENT 00180901 EUM095-13

Paraspiculatus orobiana, m, Brazil USNM ENT 00179441 EUM096-14

Paraspiculatus orobiana, m, Ecuador USNM ENT 00181753 EUM287-15

Paraspiculatus orobiana, m, Peru CF-LYC-155 NLYCA155-12

Paraspiculatus orocana, m, Ecuador CF-LYC-154 NLYCA154-12

Paraspiculatus orocana, m, Ecuador USNM ENT 00180794 EUM008-13

Paraspiculatus orocana, m, Ecuador USNM ENT 00180795 EUM009-13

Paraspiculatus orocana, m, Ecuador USNM ENT 00180796 EUM010-13

Paraspiculatus orocana, m, Ecuador USNM ENT 00180797 EUM011-13

Paraspiculatus orocana, m, Peru CF-LYC-153 NLYCA153-12

Paraspiculatus sine, m, Ecuador USNM ENT 00180830 EUM035-13

Paraspiculatus sine, m, Ecuador USNM ENT 00180849 EUM057-13 

Paraspiculatus sine, m, Ecuador USNM ENT 00180851 EUM059-13

Paraspiculatus sine, m, Ecuador USNM ENT 00180852 EUM060-13

Paraspiculatus sine, m, Ecuador USNM ENT 00180853 EUM061-13

Paraspiculatus sine, m, Ecuador USNM ENT 00180854 EUM062-13
6 0 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE A.1. (Continued)

Species, sex, country Voucher number BOLD ID

Theorema eumenia, m, Costa Rica 15-SRNP-30546 MHMYK7751-15

Theorema eumenia, m, Costa Rica 09-SRNP-30866 MHMYC1167-09

Theorema eumenia, m, Costa Rica 12-SRNP-31404 BLPEF1814-13

Theorema eumenia, m, Costa Rica 09-SRNP-30865 MHMYC1168-09

Theorema eumenia, f, Costa Rica 09-SRNP-30789 MHMYC1175-09

Theorema eumenia, f, Costa Rica 09-SRNP-30787 MHMYC1176-09

Theorema eumenia, f, Costa Rica 09-SRNP-32018 MHMYH037-10

Theorema eumenia, f, Costa Rica 09-SRNP-3212 MHMYH038-10

Theorema eumenia, f, Costa Rica 11-SRNP-30697 MHMYM1434-11

Theorema eumenia, f, Costa Rica 11-SRNP-30696 MHMYM1435-11
TABLE B.1. Genitalic dissections of Paraspiculatus with dissection number, country 

(and province if known) where the specimen was collected, and museum in which it is 

deposited (with barcode identifier if available). An asterisk (*) indicates a genitalic photograph was used for comparison of stuctures; a hashtag (#) indicates the specimen was 

drawn and is illustrated herein. Museum abbreviations are defined in text under “Materials and Methods” (ENT is part of the USNM barcode number).

Dissection Museum, with

Sex, species number Locality barcode if available

♂ P. apuya CF71*# Napo, Ecuador USNM ENT 00180817

♂ P. apuya CF72* Napo, Ecuador USNM ENT 00180818

♂ P. apuya CF73* Rondonia, Brazil USNM ENT 00180888

♂ P. azul CF54*# Napo, Ecuador USNM ENT 00180848

♂ P. azul CF55* Napo, Ecuador USNM ENT 00180871

♀ P. azul CF56*# Madre de Dios, Peru USNM ENT 00180894

♂ P. catrea 1982:74*# Santa Catarina, Brazil USNM ENT 00181696

♀ P. catrea WDF2411 Rio de Janeiro, Brazil USNM 

♀ P. catrea 1982:75# Paraná, Brazil USNM 

♂ P. catrea 2013:68 Rio Grande do Sul, Brazil USNM 

♂ P. catrea 2013:69 Santa Catarina, Brazil USNM ENT 00179443

♂ P. colombiensis CF50*# Morona Santiago, Ecuador USNM ENT 00180842

♂ P. colombiensis CF51* Morona Santiago, Ecuador USNM ENT 00180843

♂ P. colombiensis CF52* Pastaza, Ecuador USNM ENT 00180847

♀ P. colombiensis CF53* Pastaza, Ecuador USNM ENT 00179432

♀ P. colombiensis CF49*# Morona Santiago, Ecuador USNM ENT 00180840

♂ P. colombiensis CF 504 Colombia MNHN

♂ P. cosmo CF78* Morona Santiago, Ecuador USNM ENT 00180798

♂ P. cosmo CF79*# Morona Santiago, Ecuador USNM ENT 00180800

♂ P. cosmo CF80* Morona Santiago, Ecuador USNM ENT 00180804

♀ P. cosmo CF81*# Morona Santiago, Ecuador USNM ENT 00180806

♀ P. cosmo 1982:312 Peru USNM ENT 00179440

Appendix B. Paraspiculatus

Genitalic Dissections
6 2 • SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE B.1. (Continued)

Dissection Museum, with

Sex, species number Locality barcode if available

♂ P. elis 1982:208# Peru USNM ENT 00181697

♂ P. elis 1983:135 Peru USNM 

♀ P. elis 1983:136# Peru USNM ENT 00179430

♂ P. emma CF68* Napo, Ecuador USNM ENT 00180807

♂ P. emma CF69*# Napo, Ecuador USNM ENT 00180812

♂ P. emma CF70* Napo, Ecuador USNM ENT 00180813

♂ P. emma CF509 Peru USNM ENT 00180813

♂ P. grande CF85* Esmeraldas, Ecuador USNM ENT 00180788

♂ P. grande CF86*# Esmeraldas, Ecuador USNM ENT 00180789

♀ P. grande CF87*# Panama USNM ENT 00179438

♂ P. hannelore CF63 Santa Catarina, Brazil USNM ENT 00180895

♂ P. hannelore CF64*# Santa Catarina, Brazil USNM ENT 00180896

♀ P. hannelore CF65*# Santa Catarina, Brazil USNM ENT 00180897

♂ P. honor 1982:311 Colombia USNM ENT 00179442

♂ P. honor 2013:71# Ecuador RCB

♀ P. honor CF84*# Pichincha, Ecuador USNM ENT 00180862

♂ P. lilyana CF66* Napo, Ecuador USNM ENT 00180831

♂ P. lilyana CF67*# Napo, Ecuador USNM ENT 00180832

♂ P. noemi CF60*# Pichincha, Ecuador USNM ENT 00180855

♂ P. noemi CF61* Esmeraldas, Ecuador USNM ENT 00180858

♂ P. noemi CF62* Mexico USNM ENT 00180898

♀ P. noemi 2013:74# Costa Rica USNM ENT 00180899

♂ P. oroanna 2013:40# Ecuador USNM ENT 00180787 

♂ P. orobia CF74*# Napo, Ecuador USNM ENT 00180819

♂ P. orobia CF75* Napo, Ecuador USNM ENT 00180821

♂ P. orobia CF53145 Bolivia CMNH

♂ P. orobia CF209 Bolivia USNM

♂ P. orobia CF76* Rondonia, Brazil USNM ENT 00179431

♂ P. orobia CF501 Peru MNHN

♂ P. orobia CF502 Bolivia MNHN

♂ P. orobia CF503 Amazonas, Brazil MNHN

♂ P. orobia CF507 Colombia MNHN 

♀ P. orobia CF77*# Peru USNM ENT 00180889

♂ P. orobiana 1982:313*# Amazonas, Brazil USNM ENT 00179441

♀ P. orobiana CF2013:28# Peru USNM ENT 00181901

♂ P. orocana CF82*# Zamora Chinchipe, Ecuador USNM ENT 00180794

♂ P. orocana CF83* Morona Santiago, Ecuador USNM ENT 00180795 

♂ P. sine CF57* Napo, Ecuador USNM ENT 00180849

♂ P. sine CF58*# Morona Santiago, Ecuador USNM ENT 00180851

♂ P. sine CF59* Napo, Ecuador USNM ENT 00180830

♂ P. transvesta 1982:210*# Guatemala USNM ENT 00179436

♀ P. transvesta 2013:70# Mexico MGCL

♀ P. vossoroca CF88*# Santa Catarina, Brazil USNM ENT 00179437
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