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smithsonian contributions to zoology • number 643

A Revision of the 

New World Species 

of Paralimna Loew 

(Diptera: Ephydridae)

Rosaly Ale-Rocha and Wayne N. Mathis

washington d.c.

2015
ABSTRACT

Ale-Rocha, Rosaly, and Wayne N. Mathis. A Revision of the New World Species of Paralimna Loew (Diptera: Ephydridae). Smithsonian Contributions to Zoology, number 643, x + 148 pages, 215 figures, 31 maps, 1 table, 2015.—The 

New World species of the genus Paralimna Loew are revised. Fifteen new species are described and four new synonyms 

are proposed (Paralimna cilifera Hendel = Paralimna sana Cresson, Paralimna sticta Hendel = Paralimna taurus Cresson, 

Paralimna bistriata Hendel = Paralimna molossus Schiner, and Paralimna multipunctata Williston = Paralimna thomae

(Wiedemann)). A key to the 34 known, New World species of the subgenera Paralimna is provided, as is a key to the 

three New World species of the subgenus Phaiosterna Cresson. Diagnoses, detailed distributional data for the species of 

the genus, notes on the biology, and illustrations (photographs and drawings) are provided to assist species identification. 

A cladistic analysis was performed to examine the monophyly of the known species groups of Paralimna. The ingroup 

includes a total of 64 exemplar species. Outgroup sampling includes the following exemplar genera: Afrolimna Cogan, 

Oedenops Becker, and Papuama Mathis and Zatwarnicki. The analyses of the 36 included characters recovered a monophyletic subgenus Phaiosterna Cresson and a paraphyletic subgenus Paralimna if the limbata species group is included. 

Except for the limbata species group, the remaining known species groups of Paralimna were not sustained.

Cover images, from left to right: Head, body, and wing of Paralimna stellata, sp. nov., from Figures 194, 1, and 207 herein, 

respectively. (Center photo by S. A. Marshall, used with permission.)

Published by SMITHSONIAN INSTITUTION SCHOLARLY PRESS

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Library of Congress Cataloging-in-Publication Data

Ale-Rocha, Rosaly, author.

A revision of the New World species of Paralimna Loew (Diptera: Ephydridae)/Rosaly Ale-Rocha and Wayne N. Mathis.

 pages cm.—(Smithsonian contributions to zoology, ISSN 0081-0282 ; number 643)

Includes bibliographical references and index.

1. Ephydridae—Classification. I. Mathis, Wayne N., author. II. Title. III. Series: Smithsonian contributions to 

zoology; no. 643.

QL537.E7A45 2015

595.77’4—dc23

2014046130

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

Publication date: 1 June 2015

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 MAPS ix

INTRODUCTION 1

Methods and Materials 2

Acknowledgments 3

SYSTEMATICS 4

Tribe Dryxini Zatwarnicki 4

 Genus Paralimna Loew 4

 Key to New World Subgenera of Paralimna 5

 Subgenus Paralimna Loew 5

 Key to New World Species of the Subgenus Paralimna 6

 1. Paralimna (Paralimna) adunca, sp. nov. 10

 2. Paralimna (Paralimna) argyrostoma Cresson 13

 3. Paralimna (Paralimna) aurantia, sp. nov. 14

 4. Paralimna (Paralimna) brunneiceps Cresson 18

 5. Paralimna (Paralimna) castanea, sp. nov. 20

 6. Paralimna (Paralimna) crinita, sp. nov. 22

 7. Paralimna (Paralimna) curta, sp. nov. 25

 8. Paralimna (Paralimna) ecuadoriensis, sp. nov. 27

 9. Paralimna (Paralimna) fellerae Mathis 28

 10. Paralimna (Paralimna) flexineuris Cresson 33

 11. Paralimna (Paralimna) fulgifrons, sp. nov. 36

 12. Paralimna (Paralimna) guttata, sp. nov. 38

 13. Paralimna (Paralimna) maculata, sp. nov. 42

 14. Paralimna (Paralimna) malleata, sp. nov. 45

 15. Paralimna (Paralimna) meridionalis Cresson 47

 16. Paralimna (Paralimna) molossus Schiner 51

 17. Paralimna (Paralimna) nigropicta Cresson 54

 18. Paralimna (Paralimna) pallida, sp. nov. 58

 19. Paralimna (Paralimna) pauca, sp. nov. 60

 20. Paralimna (Paralimna) pectinata Hendel 62
i v • smithsonian contributions to zoology

 21. Paralimna (Paralimna) piger Cresson 66

 22. Paralimna (Paralimna) pleurivittata Cresson 68

 23. Paralimna (Paralimna) plumbiceps Cresson 71

 24. Paralimna (Paralimna) puncticornis Cresson 72

 25. Paralimna (Paralimna) punctipennis (Wiedemann) 76

 26. Paralimna (Paralimna) sana Cresson 82

 27. Paralimna (Paralimna) secunda Schiner 84

 28. Paralimna (Paralimna) sera Cresson 89

 29. Paralimna (Paralimna) stellata, sp. nov. 91

 30. Paralimna (Paralimna) stigmata, sp. nov. 93

 31. Paralimna (Paralimna) taurus Cresson 96

 32. Paralimna (Paralimna) texana Cresson 98

 33. Paralimna (Paralimna) thomae (Wiedemann) 102

 34. Paralimna (Paralimna) velutina, sp. nov. 109

 Subgenus Phaiosterna Cresson 111

 Key to New World Species of the Subgenus Phaiosterna 112

 35. Paralimna (Phaiosterna) decipiens Loew 112

 36. Paralimna (Phaiosterna) longiseta Mathis 

and Zatwarnicki 117

 37. Paralimna (Phaiosterna) obscura Williston 119

 Paralimna (Coganolimna), subgen. nov. 125

PHYLOGENETIC CONSIDERATIONS 125

Characters Used in the Analysis 126

Analysis, Results, and Discussion 127

FIGURES 162–215 130

REFERENCES 145

INDEX OF SCIENTIFIC NAMES 147
Figures

1 (Frontispiece). Paralimna (Paralimna) stellata, sp. nov. (♂) x

2–5. Paralimna (Paralimna) adunca, sp. nov., male terminalia 11

6–9. Paralimna (Paralimna) argyrostoma Cresson, male terminalia 14

10–13. Paralimna (Paralimna) aurantia, sp. nov., male terminalia 16

14–17. Paralimna (Paralimna) brunneiceps Cresson, male terminalia 19

18–21. Paralimna (Paralimna) castanea, sp. nov., male terminalia 22

22–25. Paralimna (Paralimna) crinita, sp. nov., male terminalia 24

26–29. Paralimna (Paralimna) curta, sp. nov., male terminalia 26

30–33. Paralimna (Paralimna) ecuadoriensis, sp. nov., male terminalia 29

34–39. Scanning electron micrographs of Paralimna (Paralimna) 

fellerae Mathis 30

40–45. Scanning electron micrographs of Paralimna (Paralimna) 

fellerae Mathis 31

46–49. Paralimna (Paralimna) fellerae Mathis, male terminalia 32

50–53. Paralimna (Paralimna) flexineuris Cresson, male terminalia 35

54–57. Paralimna (Paralimna) fulgifrons, sp. nov., male terminalia 38

58–61. Paralimna (Paralimna) guttata, sp. nov., male terminalia 40

62–65. Paralimna (Paralimna) maculata, sp. nov., male terminalia 43

66–69. Paralimna (Paralimna) malleata, sp. nov., male terminalia 46

70–73. Paralimna (Paralimna) meridionalis Cresson, male terminalia 49

74–77. Paralimna (Paralimna) molossus Schiner, male terminalia 52

78–81. Paralimna (Paralimna) nigropicta Cresson, male terminalia 56

82–85. Paralimna (Paralimna) pallida, sp. nov., male terminalia 59

86–89. Paralimna (Paralimna) pauca, sp. nov., male terminalia 61

90–93. Paralimna (Paralimna) pectinata Hendel, male terminalia 64

94–97. Paralimna (Paralimna) piger Cresson, male terminalia 67

98–101. Paralimna (Paralimna) pleurivittata Cresson, male terminalia 69

102–105. Paralimna (Paralimna) plumbiceps Cresson, male terminalia 72

106–109. Paralimna (Paralimna) puncticornis Cresson, male terminalia 74

110–113. Paralimna (Paralimna) punctipennis (Wiedemann), male terminalia 77
v i • smithsonian contributions to zoology

114–117. Paralimna (Paralimna) sana Cresson, male terminalia 83

118–121. Paralimna (Paralimna) secunda Schiner, male terminalia 86

122–125. Paralimna (Paralimna) sera Cresson, male terminalia 90

126–129. Paralimna (Paralimna) stellata, sp. nov., male terminalia 92

130–133. Paralimna (Paralimna) stigmata, sp. nov., male terminalia 94

134–137. Paralimna (Paralimna) taurus Cresson, male terminalia 97

138–141. Paralimna (Paralimna) texana Cresson, male terminalia 100

142–145. Paralimna (Paralimna) thomae (Wiedemann), male terminalia 103

146–149. Paralimna (Paralimna) velutina, sp. nov., male terminalia 110

150–153. Paralimna (Phaiosterna) decipiens Loew, male terminalia 113

154–157. Paralimna (Phaiosterna) longiseta Mathis and Zatwarnicki, 

male terminalia 118

158–161. Paralimna (Phaiosterna) obscura Williston, male terminalia 120

162–165. Photographs of heads: Paralimna adunca, sp. nov.; Paralimna 

argyrostoma Cresson; Paralimna aurantia, sp. nov.; Paralimna 

brunneiceps Cresson 130

166–169. Photographs of heads: Paralimna castanea, sp. nov.; Paralimna 

crinita, sp. nov.; Paralimna curta, sp. nov.; Paralimna 

ecuadoriensis, sp. nov. 131

170–173. Photographs of heads: Paralimna fellerae Mathis; Paralimna 

flexineuris Cresson; Paralimna fulgifrons, sp. nov.; Paralimna 

guttata, sp. nov. 132

174–177. Photographs of heads: Paralimna guttata, sp. nov.; Paralimna 

maculata, sp. nov.; Paralimna malleata, sp. nov.; 

Paralimna meridionalis Cresson 133

178–181. Photographs of heads: Paralimna meridionalis Cresson; Paralimna 

molossus Schiner; Paralimna molossus Schiner; Paralimna thomae

(Wiedemann) 134

182–185. Photographs of heads: Paralimna nigropicta Cresson; 

Paralimna pallida, sp. nov.; Paralimna pauca, sp. nov.; Paralimna 

pectinata Hendel 135

186–189. Photographs of heads: Paralimna piger Cresson; Paralimna 

pleurivittata Cresson; Paralimna plumbiceps Cresson; Paralimna 

puncticornis Cresson 136

190–193. Photographs of heads: Paralimna punctipennis (Wiedemann); 

Paralimna sana Cresson; Paralimna secunda Schiner; 

Paralimna sera Cresson 137

194–197. Photographs of heads: Paralimna stellata, sp. nov.; Paralimna 

stigmata, sp. nov.; Paralimna texana Cresson; Paralimna 

velutina, sp. nov. 138

198–203. Photographs of wings: Paralimna castanea, sp. nov.; Paralimna 

crinita, sp. nov.; Paralimna fulgifrons, sp. nov.; Paralimna guttata, 

sp. nov.; Paralimna guttata, sp. nov.; Paralimna malleata, sp. nov. 139

204–209. Photographs of wings: Paralimna pauca, sp. nov.; 

Paralimna pleurivittata Cresson; Paralimna punctipennis

(Wiedemann); Paralimna stellata, sp. nov.; Paralimna stigmata, 

sp. nov.; Paralimna velutina, sp. nov. 140
number 643 • vii

210–213. Photographs of legs: Paralimna brunneiceps Cresson; Paralimna 

flexineuris Cresson; Paralimna molossus Schiner; Paralimna 

pectinata Hendel 141

214. Consensus cladogram, with bootstrap values, of hypothetical 

relationships among species of Paralimna 142

215. Tree chosen to represent hypothetical relationships among 

species of Paralimna 143

Distribution Maps

1. Paralimna (Paralimna) adunca, sp. nov. 12

2. Paralimna (Paralimna) argyrostoma Cresson, P. (Paralimna) 

crinita, sp. nov. 15

3. Paralimna (Paralimna) aurantia, sp. nov. 17

4. Paralimna (Paralimna) brunneiceps Cresson 20

5. Paralimna (Paralimna) castanea, sp. nov. 23

6. Paralimna (Paralimna) curta, sp. nov., P. (Paralimna) 

ecuadoriensis, sp. nov. 27

7. Paralimna (Paralimna) fellerae Mathis 33

8. Paralimna (Paralimna) flexineuris Cresson 36

9. Paralimna (Paralimna) fulgifrons, sp. nov. 39

10. Paralimna (Paralimna) guttata, sp. nov., P. (Paralimna) piger Cresson 41

11. Paralimna (Paralimna) maculata, sp. nov. 44

12. Paralimna (Paralimna) malleata, sp. nov., P. (Paralimna) 

plumbiceps Cresson 47

13. Paralimna (Paralimna) meridionalis Cresson 50

14. Paralimna (Paralimna) molossus Schiner 54

15. Paralimna (Paralimna) nigropicta Cresson 57

16. Paralimna (Paralimna) pallida, sp. nov., P. (Paralimna) stellata, sp. nov. 60

17. Paralimna (Paralimna) pauca, sp. nov., P. (Paralimna) sera Cresson 62

18. Paralimna (Paralimna) pectinata Hendel 65

19. Paralimna (Paralimna) pleurivittata Cresson 70

20. Paralimna (Paralimna) puncticornis Cresson 75

21. Paralimna (Paralimna) punctipennis (Wiedemann) 81

22. Paralimna (Paralimna) sana Cresson 84

23. Paralimna (Paralimna) secunda Schiner 89

24. Paralimna (Paralimna) stigmata, sp. nov. 95

25. Paralimna (Paralimna) taurus Cresson 98

26. Paralimna (Paralimna) texana Cresson 101

27. Paralimna (Paralimna) thomae (Wiedemann) 108

28. Paralimna (Paralimna) velutina, sp. nov. 111

29. Paralimna (Phaiosterna) decipiens Loew 116

30. Paralimna (Phaiosterna) longiseta Mathis and Zatwarnicki 119

31. Paralimna (Phaiosterna) obscura Williston 124
FRONTISPIECE (FIGURE 1). Paralimna (Paralimna) stellata, sp. nov. (Ecuador. Orellana: Tiputini, ♂). Photograph by Stephen A. Marshall, 

used with permission.
INTRODUCTION

The shore-fly genus Paralimna Loew is one of the most species rich in the subfamily 

Hydrelliinae Robineau-Desvoidy, having 98 described species worldwide (Mathis and 

Zatwarnicki 1995, 2002, and updates, including those described in this paper). Although 

the New World fauna of Paralimna is diverse, its species richness has never been adequately explored or documented, which are the primary objectives of this paper.

Few summaries of the New World fauna of Paralimna are available, and these are 

now woefully out of date. Almost one hundred years ago Cresson (1916) revised the 

genus and then, more than thirty years later (1947), he provided a synopsis of the Neotropical fauna. Although Cresson’s synopsis was then the best available, it was incomplete, including only 16 of nearly 25 described species and lacking keys, illustrations, and 

adequate descriptions. In his synopsis, and perhaps partially accounting for the incomplete synoptic treatment, Cresson (1947:47) also observed that the subgenus Paralimna

is “a group of very difficult species.” Since Cresson, the New World fauna of Paralimna

has been largely neglected, undoubtedly in part because of the difficulty in recognizing 

species, as Cresson observed.

In a preliminary study of New World fauna of Paralimna we discovered that numerous species await description. The purpose of this paper is to revise comprehensively the 

New World fauna of Paralimna, including 15 undescribed species (e.g., Figure 1) in addition to the 25 New World species that had been previously described, primarily as isolated 

descriptions. Much of this increase has resulted from a greatly improved sampling of the 

New World fauna, especially from the Neotropics. Also contributing to this expansion is 

our use of characters from structures of the male terminalia, which have revealed species 

complexes for what had been treated in some cases as a single, widespread species.

The genus Paralimna Loew, type species Paralimna appendiculata Loew, 1862a, 

was first described from specimens collected in the Nearctic Region (“Middle States” and 

“Georgia”; Loew 1862a), but the genus is now known to occur essentially pantropically 

with some species extending into north and south temperate areas (Egypt, Japan, northern Australia, and North America) (Mathis and Zatwarnicki 2002). Before the current 

study, the genus had included about 85 species, and of these 25 species had been recorded 

previously from the New World (Mathis and Zatwarnicki 1995, 2002). Paralimna is classified in the tribe Dryxini and includes most of the species diversity within this tribe. The 

diversity of species within Paralimna is accompanied by considerable structural diversity, 

A Revision of the New World Species of 

Paralimna Loew (Diptera: Ephydridae)

Rosaly Ale-Rocha, Coordenação de Biodiversidade, Instituto Nacional de Pesquisas da 

Amazônia-INPA, Caixa Postal 2223, 69080-

971, Manaus-AM, Brazil; alerocha@inpa.gov.br. 

Wayne N. Mathis, 400 East 2460 South, Price, 

Utah 84501, USA; curator emeritus of Diptera, 

Smithsonian Institution, National Museum of 

Natural History, Department of Entomology, 

Washington, D.C., USA. Correspondence: 

W. Mathis, mathisw@si.edu. 

Manuscript received 10 June 2014; accepted 18 

September 2014.
2 • smithsonian contributions to zoology

Methods and Materials

The terminology adopted in the descriptions, with the 

exceptions noted in Mathis (1986), Mathis and Zatwarnicki 

(1990a), and below, follows McAlpine (1981). Our usage of 

the term mesopleuron in the keys and descriptions is according to McAlpine (1981:26, “the entire pleuron of the mesothorax”) and is not synonymous or to be confused with the anepisternum. We have followed the terminology for most structures 

of the male terminalia that other workers in Ephydridae have 

used (see references in Mathis 1986; Mathis and Zatwarnicki 

1990a, 1990b), such as surstylus, which in Dryxini is divided 

into a presurstylus (surstylus) and postsurstylus (clasper). 

We use the term basal flagellomere for the large antennomere beyond the pedicel. The terminology for structures of 

the male terminalia is provided directly as labels on Figures 

2–5 and is not repeated for comparable illustrations of other 

species. Alternative spellings for some localities are cited in 

parenthesis, especially for locality names that were transliterated into English. The species descriptions are composite and 

not based solely on the holotype. Veins and crossveins are 

defined in McAlpine (1981). One head and two venational 

ratios that are used in the descriptions are defined below (all 

ratios are averages of three specimens: the largest, smallest, 

and one other).

1. Gena-to-eye ratio is the genal height divided by the eye height. 

Measurements are taken from the head in lateral view.

2. Costal-vein ratio is the straight line distance between the apices of radial (R) veins R2+3 and R4+5 divided by the distance 

between the apices of R1

 and R2+3.

3. Medial-vein (M-vein) ratio is the straight line distance along 

vein M between crossveins r–m and dm–cu divided by the 

distance apicad of crossvein dm–cu.

The cladistic analysis was performed using the program 

NONA (Goloboff 1999). A branch-swapping search was conducted with the characters unordered and unweighted option in 

effect. The data matrix (see Table 1) consisted of 36 morphological characters compiled for 64 species of Paralimna. Outgroups 

were selected from the genera Afrolimna Cogan, Oedenops

Becker, and Papuama Mathis and Zatwarnicki.

Distribution maps were made using Esri ArcView© 

GIS 3.2. Longitude and latitude coordinates were obtained for 

the locality where each specimen was collected and entered into 

a Microsoft Excel© spreadsheet. If unavailable directly from 

specimen labels, longitude and latitude were estimated using 

gazetteers and maps to determine the geographical coordinates. 

Localities of specimens were plotted on a world land projection, 

presented within Esri ArcView layouts, and exported as encapsulated postscript [EPS] files.

making characterization of even the genus difficult. The monophyly of the genus, for example, is supported by a single morphological character: the lateral aedeagal process (Mathis and 

Zatwarnicki 2002). Body length, which is just one indication 

of the variation, varies by a factor of four, between 1.85 and 

7.80 mm.

Paralimna is currently divided into two subgenera, the 

nominal Paralimna Loew and Phaiosterna Cresson. These 

subgenera are distinguished by the shape of the eye and height 

of the gena (eye distinctly higher than wide and short gena in 

Phaiosterna; eye nearly round, about as wide as high, and gena 

high in Paralimna). Cresson (1916) described Phaiosterna as 

a subgenus within Paralimna, and this subgenus has been 

accepted by subsequent authors. Mathis and Zatwarnicki 

(2002) revised Phaiosterna on a world basis and included six 

species of which three occur in the New World. The subgenus 

Paralimna is substantially more species rich than Phaiosterna

and currently includes nearly 80 described species (Mathis 

and Zatwarnicki 1995). Moreover, and based on morphospecies existent in collections, we knew that species diversity for 

the genus is considerably greater than described, especially 

in the New World. The forefemur of males, with series of 

distinct, often curved, setae on the anteroventral surface, is 

considered the synapomorphy of Paralimna (Paralimna). In 

the Afrotropical Region the subgenera are divided into seven 

species groups (Cogan 1968), and except for the Afrotropical 

limbata group, which is indeed monophyletic as characterized 

by the gena being secondarily short, the remaining sublineages 

within this subgenus were not characterized (Mathis and Zatwarnicki 2002).

Cresson (1915, 1916, 1929, 1933) contributed significantly 

to our knowledge of Paralimna by describing more than half 

of the New World species. Except for five species (P. decipiens

Loew, 1878; P. molossus Schiner, 1868; P. punctipennis (Wiedemann, 1830); P. secunda Schiner, 1868; and P. thomae (Wiedemann, 1830)), all other species of Paralimna from the New 

World were described mostly in the first third of the twentieth 

century (Cresson 1915, 1916, 1929, 1933; Hendel 1930; Mathis 

and Zatwarnicki 2002), and only a few species have been revised. 

Descriptions of the first recognized species are comparatively 

simple and, in view of the currently known fauna, are largely 

inadequate for species’ recognition. This situation became even 

more egregious with the addition of more species descriptions. 

For the vast majority of New World congeners, description and 

illustrations of structures of the male terminalia are completely 

lacking. Following the precedent of Cogan (1968), we discovered that these structures provide additional and important characters for recognition of species. Thus, a major purpose for the 

research reported herein was to describe, illustrate, and analyze 

these structures, especially as the resulting information pertains 

to recognition of included taxa and phylogenetic placement.
number 643 • 3

were assembled by Smithsonian digital imaging specialist Karolyn Darrow, who also expertly produced these photographs. 

Their expertise and contributions are greatly appreciated. We 

thank Stephen A. Marshall for allowing use of his photograph 

of Paralimna stellata on the cover and as our frontispiece 

(Figure 1). (The remaining figures were produced by the authors: 

RAR drew the male terminalia; WNM produced the scanning 

electron micrographs.)

For reviewing a draft of this paper, we thank Tadeusz 

Zatwarnicki.

This work was funded by a postdoctoral scholarship granted 

by the Coordenação de Aperfeiçoamento de Pessoal de Nível 

Superior (CAPES), Brazil, to the first author. We also thank Flavia 

and Eduardo Lazarini Fonseca for their support to the first author 

while in Washington, D.C., for the development of this work.

We are also grateful to David Challinor (deceased), former assistant secretary for research, Smithsonian Institution, 

and Stanwyn G. Shetler, former deputy director of the National 

Museum of Natural History, for financial support to conduct 

field work and to study primary types through grants from the 

Research Opportunity Fund.

Funding for field work in Belize was provided by the Smithsonian Institution’s Caribbean Coral Reef Ecosystems (CCRE; 

Klaus Rützler, former director). This is contribution number 947 

of the CCRE project, which is partially supported by a grant from 

the Exxon Corporation. Field work in Guyana (1994, 1995, 

1997, and 2010) was supported by the Smithsonian Institution’s 

Biological Diversity of the Guianas program (publication number 

202; Vicki A. Funk, director, and Carol Kelloff, coordinator).

Funding for this research project, especially for field work 

on islands of the West Indies, in southern California (2003), in 

Texas (2004), in Idaho, Oregon, and Washington (2006), and 

in Alaska (2006) was funded largely by grants from the Biodiversity Program (Biological Surveys and Inventories), National 

Museum of Natural History, Smithsonian Institution (Lynne 

R. Parenti and George Zug, chairs). Field work in the West Indies 

was greatly expedited through the able and pleasant assistance 

of N. Dianne Mathis, Amnon Freidberg (collector nonpareil of 

Diptera), Hollis B. Williams (deceased), Kelvin Guerrero, Volker 

Hollmann, Oliver S. Flint Jr. (whose enthusiasm for collecting 

was catching), and Daniel Peréz G. Field work in the George 

Washington Memorial Parkway (2006–2008) was supported by 

a grant from the National Park Service, and we especially thank 

Brent W. Steury for his assistance, facilitation, and support.

Recent field work in Brazil (December 2009–June 2010) that 

resulted in the vast majority of Brazilian specimens studied for 

this paper was supported by a grant from Conselho Nacional de 

Desenvolvimento Científico e Tecnológico (Visiting Researcher/

Process number 401609/2009-0), which we gratefully acknowledge and thank. We thank Dianne Mathis for helping with all 

aspects of the production of this paper, especially the field work 

in Brazil. We also thank A. Bernardo Carvalho and his lab (Elisa, 

Monica, and Susana) for hosting us while we conducted field 

work along the coast of São Paulo.

Acknowledgments

Although this study was based in large part on specimens 

in the Smithsonian Institution’s National Museum of Natural 

History (USNM), Washington, D.C., numerous others were 

borrowed, particularly type specimens of the species previously 

described. To our colleagues and their institutions listed below 

who loaned specimens, we express our sincere thanks.

ABSF Archbold Biological Station, Venus, Florida 

(M. Deyrup)

AMNH American Museum of Natural History, New 

York, New York (David A. Grimaldi)

ANSP Academy of Natural Sciences of Philadelphia, 

Philadelphia, Pennsylvania (Jon K. Gelhaus and 

Jason D. Weintraub)

BMNH The Natural History Museum (formerly the British Museum [Natural History]), London, England (Brian Pitkin and John E. Chainey)

BYU Brigham Young University, Provo, Utah (Shawn 

M. Clark)

CMP Carnegie Museum of Natural History, Pittsburgh, 

Pennsylvania (Chen Young)

CNC Canadian National Collection, Ottawa, Canada 

(Jeffrey M. Cumming and James O’Hara)

CU Cornell University, Ithaca, New York (J. Liebherr)

DZUP Universidade Federal do Paraná, Curitiba, 

Paraná, Brazil (Luciane Marinoni)

FSCA Florida State Collection of Arthropods, Gainesville, Florida (Gary J. Steck)

INPA Instituto Nacional de Pesquisas da Amazônia, 

Manaus, Amazonas, Brazil (José Albertino 

Rafael)

KU University of Kansas, Lawrence, Kansas (M. Engel)

MCZ Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts (Philip D. Perkins)

MSU Montana State University, Bozeman, Montana 

(Michael A. Ivie)

MZUSP Museu de Zoologia da Universidade de São Paulo, 

São Paulo, Brazil (Carlos José Einicker Lamas)

NMW Naturhistorisches Museum, Wien, Austria (Peter 

Sehnal)

OSUC Ohio State University, Museum of Biological Diversity, Columbus, Ohio (Luciana Musetti)

SMN Staatliches Museum für Naturkunde in Stuttgart, 

Stuttgart, Germany (Hans-Peter Tschorsnig)

ZIL Zoological Institute, Lund University, Lund, Sweden (Roy Danielsson)

ZMUC Zoological Museum, Natural History Museum of 

Denmark, University of Copenhagen, Copenhagen, Denmark (Thomas Pape)

Erin Kolski of Smithsonian’s Department of Entomology 

prepared the distribution maps. The plates of heads and thoraces 
4 • smithsonian contributions to zoology

Diagnosis. This genus is distinguished from others 

of Dryxini by the following combination of characters: anterior 

proclinate fronto-orbital seta larger than posterior seta; arista 

bearing 5 or more dorsal hays; facial seta usually 1, if more, not 

well separated; notopleuron bearing 2 large setae; anepisternal 

and anterior dorsocentral (1+3) setae well developed; presutural 

supra-alar seta present; katepisternal seta present; forefemur 

lacking row of short, somewhat blunt spines along anteroventral 

surface; mid- and hindfemora normally developed, much shorter 

than length of abdomen.

Description. Small to large shore flies, body length 

1.85–7.80 mm.

Head: Frons wider than high, shallowly arched anteroventrally, not projected forward, sparsely setulose; ocelli in 

equilateral or isosceles triangle, ocellar triangle setulose; pseudopostocellar setulae well developed, usually half length of ocellar seta, divergent; paravertical seta well developed, subequal in 

length to anterior proclinate fronto-orbital seta, convergent; ocellar seta, inner and outer vertical setae, and reclinate interfrontal 

seta well developed; proclinate fronto-orbital setae 2, both shorter 

than interfrontal seta, moderately well developed, anterior seta 

about twice length of posterior seta. Basal flagellomere rounded 

apically; arista bearing 5 or more long rays along dorsal surface. 

Face with 1–3 long (subequal to length of anterior fronto-orbital 

seta), inclinate (but not cruciate) facial setae on ventral half of face 

and 3–4 smaller setulae interspersed along same vertical series as 

larger setae. Parafacial at anterior margin of eye narrow, width 

much less than length of basal flagellomere. Gena variable, from 

short (1⁄4 eye height) to high (subequal to combined length of basal 

flagellomere and pedicel); genal seta well developed.

Thorax: Mesonotum more or less uniformly setulose; 

acrostichal setulae in 6–8 irregular rows; 1 pair of large, prescutellar, acrostichal setae; dorsocentral setae 4 (1+3), well developed, 

posterior pair displaced laterally; presutural supra-alar seta well 

developed, subequal in length to anterior, dorsocentral seta; supra-alar setae 2 (1+1), presutural seta inserted more medial than 

postsutural seta; postpronotal seta well developed; notopleural 

setae 2, posterior seta shorter; anepisternum bearing 2 large setae 

along posterior margin, ventral seta greatly elongate; katepisternal seta well developed. Scutellum nearly flat, posterior margin 

truncate. Veins lacking setulae; costa long, extended to vein M; 

crossvein dm–cu regularly developed, nearly straight, longer than 

apical section of vein CuA1

. Mid- and hindfemora normally developed, shorter than length of abdomen; foretibia of male lacking 

several long, slender setae at apex on ventral surface; forebasitarsus lacking row of long, slender, pale setulae inserted along anterior surface; midtibia with 3 dorsal extensor setae (subapically, 

sub-basally, and near basal 1⁄3); mid- and hindfemora normally 

developed, much shorter than length of abdomen.

Abdomen: Male abdominal sternites 3 and 4 rectangular, longer than wide; sternite 5 narrow basally, becoming wider 

posteriorly, posterior margin shallowly concave. Male terminalia: epandrium is U-shaped in posterior view, simple, band-like, 

Field work in Valles Caldera National Preserve, New Mexico, was supported by a grant from the U.S. Department of Agriculture (Robert R. Parmenter, scientific director).

SYSTEMATICS

Tribe Dryxini Zatwarnicki

Dryxini Zatwarnicki 1992:85 [type genus: Dryxo Robineau-Desvoidy 

1830].—Mathis and Zatwarnicki 1995:115–127 [world catalog]; 

2002:1–101 [revision at generic level].

Diagnosis. Dryxini are the sister group of Notiphilini 

Bigot, and the tribe is distinguished from Notiphilini and other 

shore-fly tribes by the following combination of characters 

(Mathis and Zatwarnicki, 2002): ocellar seta stronger than the 

weak pseudopostocellar seta; reclinate fronto-orbital seta well 

developed, proclinate setae reduced or lacking; eye appearing 

bare; face wide, transversely arched, and generally projected 

anteriorly; facial setae in a serial row or two that runs more or 

less parallel to parafacial; gena high (secondarily short in some 

species of Paralimna); subcranial cavity moderately large and 

cavernous; dorsocentral setae 4 (1+3), secondarily reduced in 

some genera of Dryxini (Corythophora, Dryxo, and Omyxa); 

prescutellar acrostichal setae present, well developed; postsutural supra-alar setae strong, longer than posterior notopleural 

seta; posterior notopleural seta at same level as anterior seta; 

ventral anepisternal seta elongate, twice length of dorsal seta; 

costa elongate, extended to vein M; midtibia with prominent, 

erect, extensor setae on dorsal surface; abdominal tergites fasciate; male terminalia with surstylus divided into a presurstylus 

(clasper) and postsurstylus (surstylus); presurstylus with apex 

angulate and bifurcate; subepandrial plate reduced; pregonite 

and postgonite reduced or lacking (the structure remaining may 

represent a fused and/or reduced pre- and postgonite); hypandrium connected basally with postsurstylus, not with epandrium.

Genus Paralimna Loew

Paralimna Loew 1862a:138 [type species: Paralimna appendiculata Loew 

1862a (=Notiphila punctipennis Wiedemann 1830), monotypy].—

Osten Sacken 1878:201 [Nearctic catalog].—Becker 1896:114–115 

[review, discussion of relationships].—Williston 1908:308 [key to Nearctic genera].—Cresson 1916:101–124 [revision]; 1946:229 [key to 

Nearctic species]; 1947:37, 46 [key, review of Neotropical species].—

Curran 1934:347 [key to Nearctic genera].—Malloch 1933:11 [review].—Wirth and Stone 1956:465, 470 [generic key, review, Californian species].—Wirth 1965:748 [Nearctic catalog]; 1968:14–16 [Neotropical catalog].—Cole 1969:392, 396 [generic key, review of species 

from western United States].—Mathis and Zatwarnicki 1995:118–127 

[world catalog]; 2002:59–60 [diagnosis].
number 643 • 5

Distribution. The genus Paralimna occurs pantropically with a few species extending into temperate areas (Egypt, 

Japan, northern Australia, and North America).

Remarks. Paralimna, with 98 species (Mathis and 

Zatwarnicki 1995, 2002), includes most of the species diversity 

of Dryxini, and we know that there are numerous undescribed 

species in collections that await description, especially from 

Asia. Undoubtedly many species also await collection. Certainly 

the results of this paper document the need for ongoing field 

work with many of the new species being based on recently collected specimens.

unadorned; cercus semielliptical or ovoid; presurstylus generally L-shaped with ventral arm reduced, sometimes ventral arms 

absent; postsurstylus with apex largely bifurcate, internal lobe 

bearing 2 long and robust setae and 2 short setae inside; lateral 

aedeagal process articulated at the base of aedeagus laterally; aedeagus usually with membranous apex; hypandrium forming a 

conically shaped pouch; phallapodeme hardly sclerotized, usually 

triangular in lateral view. Female abdominal sternites 2–5 longer 

than wide, 6 rather quadrangular, 1, 7, and 8 wider than longer, 

sternite 8 with posterior margin shallowly concave; cercus small, 

ovoid; spermatheca pedunculate.

Key to New World Subgenera of Paralimna

(Adapted from Mathis and Zatwarnicki, 2002)

1. Body generally dark colored, dark brown to grayish brown; tergites without distinct dark band basally; eye distinctly 

higher than wide; gena short, usually slightly less than length of basal flagellomere (1⁄4 eye height), wholly covered by setae; 

basal flagellomere with short, inconspicuous setae dorsally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Phaiosterna Cresson

Body coloration contrastingly bicolored, silvery white to gray and dark brown; tergites distinctly banded, brown band 

basally; eye round, about as wide as high; gena high, height usually greater than length of basal flagellomere (subequal to 

combined length of basal flagellomere and pedicel), with few setae on middle and ventral margin; basal flagellomere with 

conspicuous elongate setae dorsally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Paralimna Loew

Subgenus Paralimna Loew

Paralimna Loew 1862a:138 [type species: Paralimna appendiculata Loew 

1862a (=Notiphila punctipennis Wiedemann 1830), monotypy].—

Cresson 1947:47–54 [diagnosis, review of Neotropical species].—Wirth 

1965:748 [Nearctic catalog]; 1968:14–15 [Neotropical catalog].—

Mathis and Zatwarnicki 1995:118–126 [world catalog]; 2002:61–63 

[diagnosis].

Poecilothorax Becker 1922:73 (as a genus). Type species: Poecilothorax 

angustus Becker 1922 (=Paralimna puncticollis Becker 1922), monotypy.—Cresson 1947:108 [synonymy].

Diagnosis. This subgenus is distinguished from 

other congeneric subgenera by the following combination of 

characters: small to large shore flies, body length 1.85–6.30 mm; 

thorax usually darkened dorsally and lighter laterally, frosted, 

without shining aspect.

Head: Frons usually with characteristic pattern of spots, 

brown with yellow spots at lateral margins of ocellar triangle and 

in front of anterior ocellar seta, between inner vertical and interfrontal seta, on fronto-orbits between proclinate fronto-orbital 

setae, and just above antennal bases. Ocelli arranged in equilateral or isosceles triangle; ocellar seta well developed, usually 

longer than anterior fronto-orbital seta and subequal to outer 

vertical seta, setae well separated, usually inserted anterolateral 

to anterior ocellus. Basal flagellomere usually gradually narrowed toward apex, with fringe of elongate dorsal setae; length 

about twice width; arista bearing 7–16 long hairs along dorsal 

surface. Face conspicuously convex, with carina and antennal 

groove marked, comparatively well arched; usually bearing 1, 

occasionally 2–3 long facial setae, if more than 1, setae approximate. Eye rounded, as wide as high or slightly higher than wide. 

Gena high, height slightly more than 1

⁄3 eye height, (height usually 

greater than length of basal flagellomere; subequal to combined 

length of basal flagellomere and pedicel); usually with few setae 

on ventral half.

Thorax: Mesonotum brown medially and through dorsocentral track, between yellowish to brownish gray, conspicuously vittate; pleural area predominantly gray, with or without 

spots, sometimes predominantly brown; dorsal surface of scutellum flat. Wing with coloration variable, from hyaline to distinctly patterned. Forefemur of male frequently with row of 

modified setae (curved, slightly or distinctly flattened setae) on 

anteroventral surface.

Abdomen: Frequently tergites with fasciate pattern with 

darker fascia basally. Sternite 1 shiny. Male terminalia: epandrium 

simple, band-like, unadorned; cerci subelliptical; presurstylus not 

connected to each other dorsally; postsurstylus usually widest 
6 • smithsonian contributions to zoology

to recognize different species. For example, structures of the 

male terminalia, especially the presurstylus of P. brunneiceps, 

P. fellerae, P. meridionalis, P. nigropicta, and P. punctipennis, 

are very similar. The phallapodeme should be carefully evaluated 

because it varies intraspecifically. The postsurstylus and remaining structures of the male terminalia have provided useful characters for species distinction; however, the position from which 

these structures are viewed and illustrated can mislead and result 

in misidentifications.

medially with a protuberant setulose lobe anteriorly and a variable process posteriorly in the middle; aedeagus mostly tubular, 

elongate, gently flattened laterally; lateral aedeagal process narrowed in middle, curved ventrally in lateral view.

Discussion. Structures of the male terminalia have 

been used to facilitate identification of species of Paralimna since 

Cogan (1968). Despite obvious differences in structures of the 

terminalia between species, there are cases in which some structures, mostly the presurstylus, are so similar that it is difficult 

Key to New World Species of the Subgenus Paralimna

1. Wing distinctly patterned, mostly brown with white or hyaline spots; crossvein dm–cu concave, curved to base; with short 

stump veins on longitudinal veins R4+5 and M [Figures 200–202, 207]; face high, about twice length of frons in anterior 

view [Figures 172–174, 194]; frons almost flat. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

- Wing without distinct pattern, hyaline or uniformly brown tinged; diffused infuscation mainly on transverse veins can be 

present; crossvein dm–cu not as above; short stump veins on longitudinal veins R4+5 and M rarely present; face moderately 

high, about 1.5× the length of frons; frons usually convex. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

2. Frons with a large black velvety spot on ventral portion of fronto-orbit, lateral to antenna [Figures 172, 194]. . . . . . . . 3

- Frons without a large black velvety spot [Figures 173–174]. . . . . . . . . . . . . . . . . . . Paralimna (Paralimna) guttata, sp. nov.

3. Frons with a large, shiny black ovate medial area extended from anterior ocellus to anterior margin [Figure 172]; aedeagal 

process short, shorter than half of aedeagus length, setae of presurstylus slender [Figures 54, 56]. . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paralimna (Paralimna) fulgifrons, sp. nov.

- Frons without a shiny black ovate medial area on anterior margin [Figure 194]; aedeagal process approximately half of 

aedeagal length, setae of presurstylus robust [Figures 126, 128]. . . . . . . . . . . . . Paralimna (Paralimna) stellata, sp. nov.

4. A quadrate to short rectangular dark brown to black velvety spot between bases of antennae; face uniformly silvery gray 

to yellowish gray, rarely an inconspicuous pale golden-brown stripe on the carina and sometimes a pair of small, pale 

golden spots between largest facial setae; wing hyaline lacking infuscate halos; head wider than high in anterior view 

[Figure 182]; male forefemur concave ventrally near apex, with anteroventral series of short, flattened, curved setae and 

posteroventral setae arranged in more than 1 row and concentrated on basal half. . . . . . . . . . . . . P. nigropicta Cresson

- Without a quadrate spot between bases of antennae, if spot present, then elongate, extended over carina; coloration of 

face variable, usually spotted; wing, head, male forefemur, and setae of male forefemur variable. . . . . . . . . . . . . . . . . . 5

5. Body covered with dense whitish-gray microtomentum; frons, scutum, and scutellum orange tinged or yellowish brown; 

face whitish gray usually with a large golden triangular or trapezoidal spot on carina extended over more than basal half 

of face. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

- Body, frons, scutum, scutellum, and face with other characters; if the body and face is covered with whitish-gray microtomentum, then the large golden spot on carina lacking . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .7

6. Large golden spot on carina present; frons with a large grayish-brown velvety spot on anterolateral margin, above and 

lateral to antenna; pedicel with a silvery white spot dorsally, basal flagellomere brown [Figure 163]; pleurae without spots; 

male forefemur not concave ventrally, bearing anteroventral row of very short, curved, flattened, and with blunt tip setae; 

posteroventral setae inconspicuous; crossvein dm–cu with infuscate halo; presurstylus not bifurcate, with horizontal process almost straight [Figure 6]. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. argyrostoma Cresson

- Large golden spot on carina present or absent; frons without a large grayish-brown velvety spot on anterolateral margin, if 

present spot on anterolateral margin is small and dark gray; pedicel predominantly gray, orange, or brown tinged dorsally, 

without a distinct white spot dorsally, basal flagellomere gray [Figure 171]; male forefemur with ventral surface concave 

on distal 1

⁄3, posteroventral setae short, arranged in more than 1 row and concentrated on basal 1⁄3 of femur [Figure 211]; 

crossvein dm–cu without infuscate halo; presurstylus with basoventral process somewhat developed, horizontal process 

shallowly curved [Figure 50]. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. flexineuris Cresson

7. Male foretibia with ventral surface irregular, wavy; male forefemur concave ventrally near apex, with anteroventral series 

of long and widely flattened setae [e.g., Figure 213]. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .8
number 643 • 7

- Male foretibia with ventral surface regular, straight, or only slightly concave; male forefemur not concave ventrally, if so, 

then the anteroventral setae of forefemur are not long nor widely flattened. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .12

8. Pleural area distinctly lighter on dorsal half (dorsal pleurites gray, only thin band on ventral margin of anepisternum and 

anepimeron brown) and darker on ventral half (ventral pleurites brown). . . . . . . . . . . . . . . . . . . . . . P. pectinata Hendel

- Mesopleuron unicolorous, ventral and dorsal pleurites mostly unicolorous, anepisternum frequently spotted. . . . . . . . .9

9. General coloration of body light, densely silvery or yellowish gray; spots of scutum pale brown; anepisternum with or 

without spots; basoventral process of presurstylus developed. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10

- General coloration of body dark gray; scutum and anepisternum with dark brown spots; basoventral process of presurstylus variable. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11

10. Male forefemur bearing a row of setae along posteroventral margin, usually interrupted for a short distance before apex; 

anepisternum usually without distinct spot; crossveins with gray infuscate halos; banded maculation pattern on tergites 

not as extensive, often lacking on tergite 5, medial stripe at most faintly evident; basoventral process of presurstylus with 

ventral margin tapered and bearing strong setae [Figure 138]. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. texana Cresson

- Male forefemur bearing a posteroventral patch of setae only near base or along entire margin; anepisternum usually 

brownish on dorsal margin; crossveins without infuscate halos; tergites with extensive brown to dark brown anterior, bilaterally symmetrical band, posterior margin of band convex on both sides and with a conspicuous medial stripe extended 

to posterior margin; basoventral process of presurstylus with ventral margin rather rounded and bearing slender setae 

[Figure 142]. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. thomae Wiedemann

11. Face dark gray with 3 brown spots, 1 on carina and 2 longitudinal parallel brown spots on ventral half, extended to margin of face; gena brown with dorsal margin silvery gray [Figure 186]; pleurae predominantly brown, anepisternum with 

ventral margin and a large, central, rounded stain brown, dorsal margin spotted by brown insertion of setae, the remaining gray; lateroventral margin of tergites brown; posteroventral surface of male forefemur with very long setae inserted on 

basal 1

⁄3 of femur; ventral surface of male foretibia not setose. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. piger Cresson

- Face gray with a golden-brown, sometimes inconspicuous, rectangular spot on carina, height equal to length of antenna; 

gena silvery gray [Figure 188]; anepisternum with a small brown spot on dorsal and ventral margin; lateral margin of 

tergites gray; setae of posteroventral surface of male forefemur long and spaced, evenly distributed along femur; male 

foretibia with conspicuous dense, rather elongate setae on posteroventral surface . . . . . . . . . . . . . P. plumbiceps Cresson

12. Gray coloration of body dark, rather bluish gray. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13

- Gray coloration of body light, silvery or yellowish gray. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .14

13. Face with a wide and short dark grayish-brown spot on carina, lateral margin brown and ventral half wholly gray; gena 

gray, lacking spots [Figure 193]; anepisternum with dorsal and ventral margins brown, central area mostly gray, sometimes with bases of setulae brown; male forefemur concave ventrally near apex; tergites predominately dark brown with 

a slender gray band on posterior margin; presurstylus with long, slender setae [Figure 122], lateral aedeagal process very 

large [Figure 124], postsurstylus slender in lateral view, lacking a developed medial posterior process [Figure 123]. . . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. sera Cresson

- Face with 3 brown spots, 1 large rectangular spot on carina, as long as length of antenna, and 2 rectangular transverse 

spots between larger facial setae; gena with a small golden-brown spot on anterior margin [Figure 175]; anepisternum 

with ventral and dorsal margins and central area mostly brown, remainder mostly gray; male forefemur not concave 

ventrally; gray band of tergites occupying about half of tergite length; presurstylus with short setae [Figure 62], lateral 

aedeagal process normally developed [Figure 64], postsurstylus widened medially in lateral view, medial posterior process 

developed [Figure 63]. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paralimna (Paralimna) maculata, sp. nov.

14. Pleural area distinctly lighter on dorsal half (dorsal pleurites gray, only thin band on ventral margin of anepisternum and 

anepimeron brown) and darker on ventral half (ventral pleurites brown). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15

- Pleural area unicolor, ventral and dorsal pleurites unicolorous, mostly gray, anepisternum frequently spotted and sometimes ventral half of anepimeron tinged. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .16

15. Wing lacking stump veins on vein M; male forefemur with anteroventral, comb-like row of flattened setae. . . . . . . . . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. fellerae Mathis

- Wing with one to several perpendicular infuscate short stump veins on second section of vein M, sometimes reduced, 

rarely absent [Figure 206]; male forefemur lacking flattened setae . . . . . . . . . . . . . . . . . . . .P. punctipennis (Wiedemann)

16. Face, anepisternum, and katepisternum predominantly brown. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17

- Face, anepisternum, and katepisternum predominantly gray, dark gray, silvery, or yellowish gray. . . . . . . . . . . . . . . . 18

17. Face uniformly dark brown, only antennal groove yellowish gray [Figure 165]; brown spots of the body faded; posteroventral surface of male forefemur bearing rows of longer, with variable length, setae [Figure 210]; lateroventral margin 
8 • smithsonian contributions to zoology

of tergites 2–5 brown; basoventral process of presurstylus developed, rounded ventrally [Figure 14]. . . . . . . . . . . . . . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. brunneiceps Cresson

- Face brown to dark brown with an inverted U shape, yellowish-gray band on middle [Figure 195]; brown spots of the 

body well distinct; forefemur of male without outstanding setae; lateroventral margin of all tergites widely brown; basoventral process of presurstylus not developed, horizontal process not bifurcate [Figure 130]. . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paralimna (Paralimna) stigmata, sp. nov.

18. Antenna black velvety with a whitish dorsal spot on pedicel; ventral fronto-orbit with a large, rounded, dark purple spot 

lateral to antennal bases; anterior margin of the frons shiny gray mediolaterally, and 1 central, small shiny gray spot 

extended until near anterior ocellus; face yellowish or silvery gray, antennal groove golden or gray, carina with a shiny 

silvery or gray spot between antennal insertion followed by a golden spot below and 2 small, rounded, diffuse golden 

stains between larger facial setae [Figure 197]; forefemur of male without outstanding setae. . . . . . . . . . . . . . . . . . . . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paralimna (Paralimna) velutina, sp. nov.

- With other character set. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .19

19. Coloration of body washed out, particularly the abdomen, whose bands do not have well-defined contours; face with 

1 short, slender pale brown spot on carina, lacking spots on distal half [Figure 183]; scutum yellowish brown; posteroventral setae of male forefemur numerous, forming more than 1 series of unaligned, elongate, and strong setae; ventral 

surface of foretibia with rather ruffled short setae; aedeagal process short and robust [Figures 84–85]; presurstylus slender 

[Figure 82]. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paralimna (Paralimna) pallida, sp. nov.

- Coloration of body distinct; face with spot on carina and usually on distal half; scutum mostly brown; posteroventral 

setae of male forefemur and ventral surface of foretibia variable; aedeagal process and presurstylus variable. . . . . . . . 20

20. Frons almost horizontal, rather protuberant on distal margin, above bases of antennae in lateral view; face long in anterior 

view, about twice height of frons. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .21

- Frons normal, convex and oblique; face shorter than twice height of frons. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22

21. Ventral surface of male forefemur distinctly concave near apex, femur bluntly narrowed on distal 1

⁄3 in lateral view; face 

with 2 golden, parallel, large spots extended along the length of face and reaching ventral margin; pedicel dark brown 

with a whitish-gray spot dorsally [Figure 167]; presurstylus bifurcate, falciform, with basoventral process long, about 2

⁄3

of length of horizontal process, horizontal process slender [Figure 22]. . . . . . . . Paralimna (Paralimna) crinita, sp. nov.

- Ventral surface of male forefemur not concave near apex, femur gradually narrowed toward apex in lateral view; coloration of face and pedicel variable; surstylus variable. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23

22. Face gray with an inconspicuous, short pale brown spot on carina; eye round, about as wide as high; pedicel lacking a 

silvery-white spot dorsally; mesopleuron and legs densely silvery gray; scutum, scutellum, and anepisternum with brown 

dot at bases of setae; wing hyaline . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. taurus Cresson

- Face yellowish gray to golden brown in the most part, silvery gray on central portion and distal margin, with 2 longitudinal, parallel golden-brown spots in middle [Figure 187]; eye rather rectangular, wider than high in lateral view; pedicel 

with a silvery-white spot dorsally; mesopleuron with a wide golden-brown stripe extended from prostigma to anepimeron, 

only ventral and dorsal margins of anepisternum paler, anepimeron wholly brown; wing slightly brown, transverse veins 

with infuscate halo [Figure 205]. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. pleurivittata Cresson

23. Face dark gray with dark brown spots, 1 widened distally on carina, and 2 elongate and parallel on distal half of face 

[Figure 168]; wing brownish; abdomen mostly brown with reduced yellow half-moon-shaped bands on lateral of tergites 

2–4, lateroventral margins of tergites brown; anepisternum brown with a distinct wide, gray, longitudinal stripe on middle; legs blackish brown, anteroventral and posteroventral setae of male forefemur very small and sparse, inconspicuous; 

aedeagal process very short, about 1⁄4 length of the aedeagus [Figure 28]. . . . . . . .Paralimna (Paralimna) curta, sp. nov.

- Coloration of face variable; wing hyaline or semi-hyaline, yellow bands of abdomen normally developed; lateroventral 

margins of tergites and anepisternum variable; legs usually brown, anteroventral series of male forefemur slightly flattened 

or only blunted apex, posteroventral series variable; aedeagal process at least half of the aedeagus length. . . . . . . . . . 24

24. Posteroventral setae of male forefemur rather disordered, organized in more than 1 series, only 1 series complete, extended 

until the apex of femur, setae numerous and short. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .25

- Posteroventral setae of male forefemur organized in 1 series extended until the apex of femur, series continuous or interrupted for a short distance before apex or more sparse on distal half, setae long or short, numerous or sparse. . . . . . . 28

25. Face gray or yellowish gray with brown spots: 1 elongate spot on carina and usually 2 spots of variable shapes on ventral half 

of face [Figure 192]; rarely ventral spots are absent; anepisternum with dorsal and ventral margins brown and with dotted 

aspect in the middle by small brown spots on insertion of setae; anepimeron usually without spots, wholly gray, and lateroventral margins of tergites usually gray; presurstylus broad, lacking basoventral process, cercus fused at the apex of dorsal 

margin of presurstylus [Figure 118]; lateral aedeagal process long and slender [Figure 120]. . . . . . . . . . . P. secunda Schiner
number 643 • 9

- Coloration of face, anepisternum, anepimeron, and lateroventral margins of tergites variable; presurstylus not as above, 

basoventral process present or absent; lateral aedeagal process variable. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26

26. Face dark gray with an elongate golden spot on carina, usually without spots on ventral half but, if present, the spots are 

very small and inconspicuous; pedicel with dorsal marginal spot whitish gray [Figure 164]; scutum with 2 dorsal stripes; 

lateral aedeagal process short and broad, shorter than half the length of aedeagus [Figures 12–13]; presurstylus not bifurcate, horizontal process slightly concave ventrally and with dorsal margin rather wavy [Figure 10]. . . . . . . . . . . . . . . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paralimna (Paralimna) aurantia, sp. nov.

- Face, antenna, and scutum variable; lateral aedeagal process long and slender, reaching half the length of aedeagus; basoventral process of presurstylus variable, dorsal margin of presurstylus not wavy . . . . . . . . . . . . . . . . . . . . . . . . . . . . .27

27. Face with coloration variable, mostly dark gray with dark brown area adjacent to or through lateral facial setae or dark to 

pale gray with a golden spot on carina and 2 triangular or rather rounded spots on ventral half of face [Figures 177, 178]; 

pleurae gray with dark gray or yellowish gray microtomentum; anepisternum with dorsal and ventral margin widely or 

narrowly brown; basoventral process of presurstylus developed, setose [Figure 70]; medial anterior lobe of postsurstylus 

small and digitiform [Figure 71]. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. meridionalis Cresson

- Face gray to dark gray with 3 brown spots, 1 rectangular on carina and 2 small rather triangular spots between larger 

facial setae [Figure 169]; pleurae dense and homogenously gray, only anepisternum with 1 brown spot on dorsal margin 

and ventral margin slightly brown tinged, stains of bases of setae brown in middle; basoventral process of presurstylus not 

developed, horizontal process slender [Figure 30]; medial anterior lobe of postsurstylus large and triangular [Figure 31]. 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paralimna (Paralimna) ecuadoriensis, sp. nov.

28. Posteroventral series of male forefemur with setae short, numerous, and proximate to each other; presurstylus with basoventral process developed or not developed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .29

- Posteroventral series of male forefemur with setae sparse, elongate, and widely separated from each other; basoventral 

process of presurstylus not developed. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32

29. Basoventral process of presurstylus not developed, presurstylus with horizontal process wide basally and gradually narrowed toward the apex [Figure 114]; face silvery gray with 1 golden-brown, elongate, drop-shaped spot on carina as apex 

of antenna, and sometimes 2 small, inconspicuous, rectangular, slender, and horizontal pale brown spots between larger 

facial setae are present [Figure 191]; male forefemur with posteroventral row of setae shortened near apex of femur, more 

concentrated on basal half. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. sana Cresson

- Basoventral process of presurstylus developed [e.g., Figures 2, 18, 66]; face silvery to yellowish gray with large spots, 1 

on carina, not drop shaped, and 2 between larger facial setae; male forefemur with posteroventral row of setae the same 

length and equally distributed along the posteroventral surface. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30

30. Ventral surface of male forefemur concave on distal half; posteroventral series short and slightly longer at base of femur, 

inserted only on basal half of femur; face silvery gray with distinct brown spots, 1 over carina, black for a short distance 

between bases of antennae and brown in the remaining, slightly longer than the antennal length and twice the width of 

the distance between bases of antennae, and 2 large triangular spots extended almost until the ventral margin of face, and 

lateral of face with a brown band [Figure 176]; abdomen mostly brown, brown bands more developed, extended dorsally, 

tergite 2 with basal brown band extended until posterior margin in middle of each side . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paralimna (Paralimna) malleata, sp. nov.

- Ventral surface of male forefemur not concave on distal half; posteroventral series extended by over the whole length of 

the femur; face gray or vaguely yellowish in the background, with variable spots; tergite 2 with basal brown band normally developed, not extended to distal margin. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31

31. Face predominantly brown due to the big brown spots, gray only on laterodistal margin and on small V-like mark in middle [Figure 166]; wing hyaline or brownish tinged with infuscate halo on crossveins r–m and dm–cu [Figure 198]; anteroventral setae of male forefemur curved, moderately long-pointed setae, not flattened; posteroventral surface with dense, 

somewhat irregular row of rather short setae along most of length of femur; anteroventral margins of tergites brown; 

basoventral process of presurstylus rather triangular [Figure 18]. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paralimna (Paralimna) castanea, sp. nov.

- Face predominantly yellowish gray [Figure 162]; wing hyaline without infuscate halo on crossveins r–m and dm–cu; anteroventral setae of male forefemur moderately elongate, only the more distal setae flattened; posteroventral setae elongate, 

conspicuous, as long as width of femur; lateroventral margins of tergites gray; basoventral process of presurstylus slender, 

hook-like [Figure 2] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paralimna (Paralimna) adunca, sp. nov.

32. Pedicel with a distinct silvery-white spot dorsally; face yellowish to golden gray with golden-brown lateral margins and 

3 large golden-brown spots, 1 on carina and 2 longitudinal, parallel, rectangular spots on ventral half [Figure 189]; 

anepisternum with 3 brown, well-delimited spots, 1 dorsal, 1 ventral, and 1 rounded spot on central portion, the latter 
1 0 • smithsonian contributions to zoology

sometimes inconspicuous; katepisternum mostly yellowish gray with a large triangular brown spot anteriorly; legs brown 

to dark brown. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. puncticornis Cresson

- Pedicel without a silvery-white spot dorsally, only distal margin of pedicel paler; face gray or yellowish gray, when present 

the spots of the ventral half are small and triangular, if the spots of the ventral half are large, the ventral half of face is

dark gray; coloration of anepisternum variable; katepisternum without large spot; legs gray. . . . . . . . . . . . . . . . . . . . 33

33. Face variable, dark gray with 1 golden, elongate spot on carina, 2 large, golden-brown spots between larger facial setae 

and lateral margins of face golden tinged, or wholly golden tinged without distinct spots or yellowish gray with goldenbrown spots, 1 on carina and 2 short inconspicuous spots on ventral half [Figures 179, 180]; ventrolateral margins of 

tergites of male usually brown; wing slightly brownish, crossvein r–m slightly infuscate; presurstylus slender and elongate, 

also sinuous [Figure 74]. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Paralimna (Paralimna) molossus Schiner

- Face dark gray with 3 golden spots, 1 elongate spot on carina and 2 parallel, elongate, rectangular spots below, in middle of face 

[Figure 184]; ventrolateral margins of tergites of male gray; wing hyaline, crossvein r–m not infuscate; presurstylus slender, with 

horizontal process abruptly narrowed in distal half [Figure 86]. . . . . . . . . . . . . . . . . . . . Paralimna (Paralimna) pauca, sp. nov.

1. Paralimna (Paralimna) adunca, sp. nov.

Figures 2–5, 162, Map 1

Diagnosis. This species is distinguished from congeners by the following combination of characters: face yellowish 

gray with 3 pale golden-brown spots—a single, long, widened 

distally spot on carina, a little longer than the length of antenna, 

and 2 small, rounded spots between larger facial setae; forefemur of male with anteroventral series of moderately long setae, 

not flattened, with rounded apex, shorter than width of femur, 

posteroventral surface with 1 series of long, sparse, and conspicuous setae, as long as width of femur; basoventral process 

of presurstylus long, falciform, about 2

⁄3 of length of horizontal 

process.

Description. Body length 4.0–5.5 mm; body generally bicolored, dark brown dorsally, lateral surfaces gray, sometimes yellowish gray.

Head: (See Figure 162) Frons brown with linear yellow 

spots at lateral margins of ocellar triangle and in front of anterior 

ocellar seta, short, not reaching the anterior margin of frons, between inner vertical and interfrontal setae, and on fronto-orbits 

between proclinate fronto-orbital setae and just above antennal 

bases; ventral fronto-orbits and dorsal parafacial with 2 small, 

inconspicuous brown spots lateral to antenna, separated by yellow spot. Antenna pale brown, covered by dense yellow microtomentum; basal flagellomere with slender, elongate, pale setae 

on dorsal margin; arista with 11 long, dorsal rays. Postcranium 

with 2 complete series of postocular setae, the second series irregular. Face yellowish gray with 3 pale golden-brown spots—a 

single, long, widened distally spot on carina, a little longer than 

the length of antenna, and 2 small, rounded spots between larger 

facial setae; parafacial and gena yellowish gray; gena with only 

few setae on middle and ventral margin; clypeus silvery gray. Eye 

round, about as wide as high, height about twice the height of 

gena. Gena high, height longer than length of basal flagellomere; 

gena-to-eye ratio 0.36–0.42.

Thorax: Mesonotum yellowish gray with dark brown 

spots, 2 stripes between dorsocentral rows, 1 short stripe on 

supra-alar area plus some small lateral spots; insertion of setae 

brown, giving a spotted aspect to scutum; postpronotum, notopleuron, and supra-alar areas yellowish gray; scutellum mostly 

yellowish gray with base brown, apex blackish brown, and 

dorsal surface spotted at insertion of setae brown. Pleurae yellowish gray, anepisternum with spotted aspect given by brown 

bases of setae. Wing hyaline; veins pale brown; costal-vein ratio 

0.39–0.43; M-vein ratio 0.87–0.90. Legs blackish brown with 

dense silvery-gray microtomentum; tarsi lighter with some yellow 

coloration, especially ventrally, last 3 segments darkened dorsally; forefemur of male with anteroventral series of moderately 

long setae, not flattened, with rounded apex, shorter than width 

of femur, posteroventral surface with 1 series of long, sparse, and 

conspicuous setae, as long as width of femur; foretibia of male 

straight.

Abdomen: Tergites distinctly bicolored, dark brown 

along anterior 2

⁄3 and medially, otherwise gray. Male terminalia: 

(Figures 2–5) Presurstylus bifurcate, bearing basoventral process 

long, falciform, about 2

⁄3 of length of horizontal process, horizontal process slender; in lateral view postsurstylus widest medially, 

medial anterior lobe developed bearing moderately elongate and 

robust setae, medial posterior process slender, digitiform; lateral 

aedeagal process about 1⁄2 length of aedeagus, rather slender, conspicuously arcuate in lateral view; aedeagus slender in dorsal view, 

progressively narrowed apically, robust in lateral view with apex 

bent backwards; phallapodeme triangular in lateral view; hypandrium moderately deeply concave.

Type Material. The holotype male of Paralimna 

adunca is labeled “MEXICO [Morelos: Puente de] Ixtla. 

400–600m 23.ii.1998 [23 Feb 1998] A. FREIDBERG/USNM 

ENT 00118261 [plastic bar code label]/HOLOTYPE ♂

Paralimna adunca Ale-Rocha & Mathis, USNM [red].” The holotype is double mounted (minuten pin in a block of plastic), is 

in excellent condition (abdomen not dissected), and is deposited 

in the USNM. Seven paratypes (3♂, 4♀; INPA, USNM) bear the 
number 643 • 1 1

97°W; 952 m), 2 May 1985, W. N. Mathis (2♂, 2♀; USNM); 

Estación biologie Los Tuxlas (~15 km N Catemaco; 18°28.8′N, 

95°07.7′W), 15–17 Sep 1987, A. L. Norrbom (2♂; INPA, 

USNM).

Other Specimens Examined. Nearctic: UNITED 

STATES. Texas. Culberson: Guadalupe National Park, McKittrick 

Canyon (31°58.7′N, 104°45.3′W), 7 Aug 1976 (2♂; USNM); 

same label data as the holotype. Other paratypes are as follows: 

MEXICO. Chiapas: Corralito (7 km W Abasolo; 16°51′N, 

92°45.2′W), 21 May 1981, C. M. and O. S. Flint (6♀, 4♂; 

INPA, USNM); Union Juárez (9 km S; 15°02.2′N, 92°05.4′W), 

23 Apr 1983, W. N. Mathis (1♀; USNM). Veracruz-Llave:

Ocotal Chico (18°15.6′N, 94°51.5′W; 600 m), 4–5 May 1985, 

W. N. Mathis (2♂, 4♀; USNM); Fortin de las Flores (18°54′N, 

FIGURES 2–5. Structures of male terminalia of Paralimna (Paralimna) adunca, sp. nov. (Mexico. Morelos: Puente de 

Ixtla): (2) epandrium, cerci, and presurstyli, posterior aspect; (3) postsurstylus, lateral aspect; (4) aedeagus, phallapodeme, 

and hypandrium, lateral aspect; (5) same, ventral aspect. Scale bar = 0.1 mm.
1 2 • smithsonian contributions to zoology

MAP 1. Distribution map for Paralimna (Paralimna) adunca, sp. nov.

Guadalupe National Park, West Dog Canyon (31°59.8′N, 

104°49.9′W), 4 Aug 1975 (1♂; USNM).

Neotropical: GUATEMALA. Huehuetanango: Ruins at 

Zaculeu (15°20′N, 91°29.6′W), 18 Aug 1972, G. F. and S. Hevel 

(l♂; USNM).

HONDURAS. Atlantida: Lancertilla, Atlantida (15°44.1′N, 

87°27.4′W; 75–110 ft [~23–34 m]), 7 Sep 1930 (2♂; ANSP).

MEXICO. Veracruz-Llave: Lake Catemaco (18°25.3′N, 

95°06.8′W), 1 May 1969, H. J. Teskey (4♂, 1♀; CNC).

Type Locality. Nearctic. Mexico. Morelos: Puente 

de Ixtla (18°37′N, 99°19.4′W; 400–600 m).

Distribution. (Map 1) Nearctic: Mexico (Morelos), 

United States (Texas). Neotropical: Guatemala (Huehuetanango), 

Honduras (Atlantida), Mexico (Chiapas, Veracruz-Llave).

Etymology. The species epithet, adunca, is of Latin 

derivation and means bent inward or hooked, alluding to the 

shape of the basoventral process of the postsurstylus.

Remarks. Structures of the male terminalia of this 

species are very similar to those of P. crinita sp. nov., especially 

the shape of the falciform basoventral process of the presurstylus, the hypandrium, and the aedeagus; however, the lateral aedeagal process is slender in P. adunca sp. nov. while in P. crinita 

sp. nov. it is wide and elongate, and the medial posterior process is digitiform in P. adunca sp. nov. whereas in P. crinita sp. 

nov. this structure is rather truncate. These species can be distinguished further by several other characters: in P. crinita sp. nov. 

the facial coloration has 2 parallel, golden-brown, wide spots 

extended along the length of face; the frons, being almost horizontal, is slightly projected over the antennal bases; the ventral 

surface of the male forefemur is distinctly concave on the distal 1⁄3

with several series of numerous, moderately long setae; and the 

foretibia of males is densely pilose on the ventral surface. In P. 

adunca sp. nov., the face has a pale golden-brown, long, distally 

widened spot on the carina, and there are 2 small, rounded spots 

between the larger facial setae; the frons is normally convex; the 

ventral surface of the male forefemur is not concave ventrally 

and has 1 series of sparse long setae; and the foretibia of males is 

not densely pilose along the ventral surface.
number 643 • 1 3

blackish brown and yellowish gray laterally. Mesopleuron 

wholly densely silvery gray, without spots. Wing semi-hyaline; 

veins yellowish brown; crossvein dm–cu with infuscate halo; 

costal-vein ratio 0.50; M-vein ratio 0.88. Legs: Femora and tibiae brown with knees and apex of tibiae yellow, sparse silverygray microtomentum; tarsi lighter with some yellow coloration, 

especially ventrally, the more distal darkened; forefemur of male 

bearing anteroventral row of short, slightly curved, with blunt 

tip, none conspicuously flattened setae, ventral series short and 

thin; foretibia straight.

Abdomen: Tergites distinctly bicolored; tergite 1 golden 

brown in the most part and silvery gray laterally, tergites 2–5 

golden brown along anterior 2

⁄3 and medially, otherwise silvery 

gray. Male terminalia: (Figures 6–9) Presurstylus not bifurcate, 

with horizontal process almost straight, slightly widened basally; 

postsurstylus in lateral view widest medially with a medial anteriorly pointed lobe bearing long and slender setae, and a medial 

small digitiform process posteriorly; lateral aedeagal process less 

than 1⁄2 length of aedeagus, slightly curved subapically, bluntly 

rounded; aedeagus gradually tapered to membranous apex; 

phallapodeme broadly triangular in lateral view; hypandrium 

shallowly concave.

Type Material. The holotype male of Paralimna 

argyrostoma is labeled “Banana River [Río Banano] 9 XI ’09[9 

Nov1909] C[osta]R[ica. Limón:] P P Calvert/Sweeping up- per 

[sic] reservoir 100 ft. [30.5 m] alt./♂/TYPE Paralimna argyrostoma [sic] E.T.Cresson,Jr. [maroon; “Paralimna argyrostoma” 

handwritten]” (ANSP No. 6094). The holotype is double 

mounted (minuten pin in a vertical card), is in excellent condition (abdomen not dissected), and is deposited in the ANSP 

(6094).

Other Specimens Examined. COSTA RICA.

Limón: Río Banano (09°50.5′N, 82°56′W), 9 Nov 1909, P. P. 

Calvert (paratypes: 4♂; 1♀ allotype; ANSP).

Type Locality. Neotropical. Costa Rica. Limón: 

Río Banano (09°50.5′N, 82°56′W).

Distribution. (Map 2) Neotropical: Costa Rica 

(Limón), Ecuador (Cresson 1947).

Remarks. Cresson (1916) first cited this species on 

pages 106–107 with the spelling “argyrostoma.” Then on page 

120, which is the description of this species, he used the spelling “agryostoma.” Subsequent authors, including Cresson, used 

“argyrostoma” as the spelling.

The number of dorsal aristal rays varies from 10–12, although Cresson (1916) wrote that their number is from 8–10. 

Paralimna argyrostoma is very similar to P. flexineuris in general 

body coloration, but in P. argyrostoma, the male forefemur is 

not concave ventrally; there is a silvery-white spot on the pedicel; the mesopleuron lacks spots; the setae on the anteroventral 

and posteroventral surfaces of the male forefemur are very short, 

almost inconspicuous; the presurstylus is wider; and the lateral 

aedeagal process is more slender than in P. flexineuris.

2. Paralimna (Paralimna) argyrostoma

Cresson

Figures 6–9, 163, Map 2

Paralimna argyrostoma Cresson 1916:120 [Costa Rica. Limón: Río Banano 

(9°50.5′N, 82°56′W); HT ♂, ANSP (6094)]; 1918:47 [review; Costa Rica].

Paralimna (Paralimna) argyrostoma.—Cresson 1947:53 [review; Neotropical species].—Wirth 1968:14 [Neotropical catalog].

Paralimna (Paralimna) agryostoma [sic].—Mathis and Zatwarnicki 

1995:118 [world catalog].

Diagnosis. This species is distinguished from congeners by the following characters: frons, scutum, and scutellum with orange-yellow coloration; face whitish gray, with a 

single wide, golden, irregular triangular spot on the carina, ventral fronto-orbits with a large grayish-brown velvety spot that 

extends above base and lateral to antenna; pedicel with a silverywhite dorsal spot near apex; pleurae wholly silvery gray, lacking spots; forefemur of male bearing anteroventral row of very 

short, slightly curved, with blunt tip, slightly flattened setae, posteroventral setae very short; wing semi-hyaline, crossvein dm–cu 

with infuscate halo.

Description. Body length 2.65–2.80 mm; body generally bicolored, golden brown dorsally, lateral surfaces silvery 

gray.

Head: (See Figure 163) Frons predominantly orange 

yellow with anterior margin grayish brown; linear silvery spots 

at lateral margins of ocellar triangle, in front of anterior ocellar 

seta, not reaching anterior margin of frons, and on fronto-orbits above dorsal proclinate fronto-orbital seta, between proclinate fronto-orbital setae, and just above antennal bases; ventral 

fronto-orbits with a large grayish-brown spot that extends above 

base and lateral to antenna, and dorsal parafacial with a brown 

spot lateral to antenna, separated by silvery-gray spot. Antennal scape and pedicel brown with some purple shiny and sparse 

gray microtomentum, a silvery-white dorsal spot near apex of 

pedicel; basal flagellomere pale brown to brown according to 

incidence of light, with slender, elongate, pale setae on margin; 

arista with 10–12 long, dorsal rays. Postcranium with 2 irregular 

series of postocular setae. Face densely silvery to slightly yellowish gray, with a single wide, golden, irregular triangular spot on 

the carina; parafacial and gena silvery gray, but golden spots can 

occur; gena with only few setae on middle and ventral margin; 

clypeus silvery gray. Eye rounded, about as wide as high. Gena 

high, height about equal to length of basal flagellomere; gena-toeye ratio 0.38.

Thorax: Mesonotum mostly golden brown with 

silvery-gray areas, usually along setal tracks, 3 well-defined 

golden-brown stripes between dorsocentral rows, becoming 

more silvery gray laterally through postpronotum, notopleuron, and supra-alar areas; scutellum golden dorsally, with apex 
1 4 • smithsonian contributions to zoology

FIGURES 6–9. Structures of male terminalia of Paralimna (Paralimna) argyrostoma Cresson (Costa Rica. Limón): (6) 

epandrium, cerci, and presurstyli, posterior aspect; (7) postsurstylus, lateral aspect; (8) aedeagus, phallapodeme, and 

hypandrium, lateral aspect; (9) same, ventral aspect. Scale bar = 0.1 mm. (See Figures 2–5 for labels of structures.)

3. Paralimna (Paralimna) aurantia, sp. nov.

Figures 10–13, 164, Map 3

Diagnosis. This species is distinguished from 

congeners by the following combination of characters: face 

gray to dark gray with lateral margins yellowish gray, with 

3 golden spots, a single, elongate spot on carina, and 2 small, 

rounded, almost inconspicuous pale brown spots between 

larger facial setae; pedicel with silvery-gray spot dorsally; 

2 brown, slender, widely separated stripes lateral to acrostichal line; insertion of setae brown, giving a spotted aspect 

to scutum; presurstylus not bifurcate, with horizontal process 

slightly concave ventrally; in lateral view postsurstylus bearing 

an elongate, pointed digitiform process posteriorly; lateral aedeagal process short, less than 1⁄2 length of aedeagus, strongly 

swollen.

Description. Body length 2.4–3.8 mm; body generally bicolored, pale brown dorsally, lateral surfaces gray, 

sometimes dark gray.
number 643 • 1 5

Head: (See Figure 164) Frons pale brown with linear, yellow spots at lateral margins of ocellar triangle, in front of anterior 

ocellar seta, reaching the anterior margin of frons, between inner 

vertical and interfrontal seta, and on fronto-orbits above dorsal 

proclinate fronto-orbital seta, between proclinate fronto-orbital 

setae, and just above antennal bases; ventral fronto-orbits with 

a dark brown spot and dorsal parafacial with a dark gray spot 

lateral to antenna, separated by yellow spot. Antennal scape and 

pedicel blackish brown, pedicel with silvery-gray spot dorsally; 

basal flagellomere pale brown to brown according to incidence of 

light, with slender elongate pale setae on margin; arista with 11–12 

long, dorsal rays. Postcranium with 1 complete series of postocular 

setae. Face gray to dark gray with lateral margins yellowish gray, 

with 3 golden spots, a single, elongate spot on carina, as long as 

the length of antenna, and 2 small, rounded, almost inconspicuous pale brown spots between larger facial setae; parafacial silvery 

gray and gena yellowish gray; gena with only few setae on middle 

and distal margin; clypeus silvery gray. Eye round, as wide as high, 

height about twice the height of gena. Gena high, height longer 

than length of basal flagellomere; gena-to-eye ratio 0.35–0.40.

Thorax: Mesonotum yellowish gray with brown spots, 

2 brown, slender, widely separated stripes between dorsocentral rows ending before posterior dorsocentral seta, plus 1 short 

brown stripe on intra-alar area; insertion of setae brown, giving 

a spotted aspect to scutum; postpronotum, notopleuron, and 

supra-alar areas pale gray; scutellum brown basally, with apex 

blackish brown and yellowish gray laterally and in middle. Pleurae yellowish gray, sometimes darkened according to incidence of 

light, anepisternum brown tinged dorsally and ventrally, a small, 

rounded spot in middle, and bases of setae brown; anepimeron 

brown on anteroventral corner. Wing hyaline; veins brown; crossvein dm–cu slightly infuscate; costal-vein ratio 0.43–0.50; M-vein 

ratio 1.15–1.21. Legs blackish brown with sparse silvery-gray microtomentum darker dorsally; tarsi lighter with some yellow coloration, especially ventrally; forefemur of male with anteroventral 

and ventral complete series of short robust but not flattened setae; 

foretibia straight.

Abdomen: Tergites distinctly bicolored, dark brown 

along anterior 2

⁄3 and medially, medial brown portion of tergite 

2 very reduced, inconspicuous, otherwise yellowish gray dorsally 

MAP 2. Distribution map for Paralimna (Paralimna) argyrostoma Cresson and P. (Paralimna) crinita, sp. nov.
1 6 • smithsonian contributions to zoology

Type Material. The holotype male of Paralimna aurantia is labeled “BOLIVIA. La Paz: Guanay (22km SE; 15°17.8′S, 

68°15.6′W, 540m), 17 Mar 2001[,] Wayne N. Mathis/USNM 

ENT 00118268 [plastic bar code label]/HOLOTYPE ♂ Paralimna 

aurantia Ale-Rocha & Mathis, USNM [red].” The holotype is 

double mounted, is in excellent condition, and is deposited in the 

USNM. Four paratypes (3♂, 1♀; INPA, USNM) bear the same 

label data as the holotype. Other paratypes are as follows: BOLIVIA. La Paz: Guanay (1 km E; 15°29.8′ S, 67°52.7′W; 580 m), 

13 Mar 2001 (3♂, 5♀; USNM); Guanay (9 km NW; 15°29.5′S, 

67°53.8′W; 850 m), 13 Mar 2001, W. N. Mathis (1♂; USNM); 

and silvery gray laterally; lateroventral margins of tergites gray 

in males and brown in females. Male terminalia: (Figures 10–13) 

Presurstylus not bifurcate, with horizontal process long, slightly 

concave ventrally; in lateral view postsurstylus with medial anterior lobe poorly developed, bearing moderately elongate setae and 

an elongate pointed digitiform process posteriorly; lateral aedeagal 

process short, less than 1⁄2 length of aedeagus, strongly swollen, 

rounded distally, and almost straight in lateral view; aedeagus slender in dorsal view, wider and blunt, constricted apically in lateral 

view; phallapodeme broadly triangular in lateral view; hypandrium 

deeply concave, abruptly narrowed on distal half in dorsal view.

FIGURES 10–13. Structures of male terminalia of Paralimna (Paralimna) aurantia, sp. nov. (Bolivia. La Paz): 

(10) epandrium, cerci, and presurstyli, posterior aspect; (11) postsurstylus, lateral aspect; (12) aedeagus, phallapodeme, and hypandrium, lateral aspect; (13) same, ventral aspect. Scale bar = 0.1 mm. (See Figures 2–5 for 

labels of structures.)
number 643 • 1 7

Other Specimens Examined. BOLIVIA. La Paz: 

Coroico (16°11.6′S, 67°43.7′W; 1600 m), 6–7 Jan 1976, L. E. 

Peña (2♂; CNC); Guanay (9 km NW; 15°29.5′S, 67°53.8′W; 

850 m), 13 Mar 2001, W. N. Mathis (1♂; USNM).

ECUADOR. Orellana: Coca, Napo River (0°27.8′S, 76°59′W; 

250 m), 12–13 Apr 1965, L. E. Peña (2♂, 4♀; CNC). ZamoraChinchipe: Cumbaratza (03°59.5′S, 78°51.9′W; 700 m), 30–31 

Mar 1965, L. E. Peña (1♀; CNC); Río Jumboé (04°04′S, 78°55.8′W; 

1200 m), 1–2 Apr 1965, L. E. Peña (2♀; CNC); Timbara (04°01.9′S, 

78°54.1′W; 1400 m), 4 Apr 1965, L. E. Peña (1♂, ♀; CNC).

PERU. Madre de Dios: Río Manu, Erika (near Salvación; 

12°50.7′S, 71°23.3′W; 550 m), 5–6 Sep 1988, W. N. Mathis 

(1♂; USNM).

Type Locality. Neotropical. Bolivia. La Paz: Guanay (22 km SE; 15°17.8′S, 68°15.6′W).

Distribution. (Map 3) Neotropical: Bolivia (La 

Paz), Ecuador (Napo, Orellana, Zamora-Chinchipe), Peru 

(Cuzco, Huánuco, Madre de Dios).

Etymology. The species epithet, aurantia, is of 

Latin derivation and means orange, alluding to the orange coloration on the facial carina.

Guanay (8 km E; 15°30.3′S, 67°50.8′W; 510 m), 13 Mar 2001, 

W. N. Mathis (2♂, 1♀; INPA, USNM); Guanay (3 km E; 

15°30.2′S, 67°52.3′W; 500 m), 14 Mar 2001, W. N. Mathis (8♂, 

1♀; INPA, USNM); Caranavi (15°50.2′S, 67°33.4′W; 670 m), 12 

Mar 2001, A. Freidberg (1♂; USNM); Mapiri (5 km W; 15°17.8′S, 

68°15.6′W; 750 m), 16 Mar 2001 (1♂; USNM); San Pedro (3 km 

NE; 16°S, 67°35.3′W; 780 m), 12 Mar 2001, A. Freidberg (1♂; 

USNM); Tajuihui (15°40.8′S, 67°41.7′W; 590 m), 12 Mar 2001, 

S. D. Gaimari, W. N. Mathis (5♂, 2♀; USNM); N Yungas between Alcoche and Teoponte (38 km SW of Guanay; 15°33.6′S, 

67°40.1′W), 12 Apr 2001, A. L. Norrbom (2♀; USNM).

ECUADOR. Napo: Limoncocha (00°24.7′S, 76°37.5′W), 

15 Jun 1977, P. J. Spangler, D. R. Givens (1♂, 5♀; INPA, 

USNM).

PERU. Cuzco: Paucartambo, Atalaya (Río Alto Madre de 

Dios; 12°53.3′S, 71°21.6′W; 600 m), 4 Sep 1988, W. N. Mathis 

(1♂; USNM). Huánuco: Huánuco (10 km N on Rio Huallaga; 

09°51′S, 76°09.1′W), 4 Feb 1984, W. N. Mathis (5♂; INPA, 

USNM). Madre de Dios: Río Manu, Erika (near Salvación; 

12°50.7′S, 71°23.3′W; 550 m), 5–6 Sep 1988, W. N. Mathis 

(2♂; USNM).

MAP 3. Distribution map for Paralimna (Paralimna) aurantia, sp. nov.
1 8 • smithsonian contributions to zoology

a broad dorsal brown stripe, becoming more silvery gray laterally 

through postpronotum, notopleuron, and supra-alar areas; scutellum brown with apex blackish brown, silvery gray to yellowish gray laterally and on middle. Pleurae with proepisternum and 

laterotergite silvery gray, anepisternum brown with anterior and 

posterior margins silvery gray, anepimeron silvery gray on dorsal half and brown on ventral half, katepisternum brown. Wing 

semi-hyaline; veins brown, crossvein dm–cu with infuscate halo; 

costal-vein ratio 0.47–0.48; M-vein ratio 1.00–1.10. Legs blackish brown with sparse silvery-gray microtomentum; tarsi lighter 

with some yellow coloration, especially ventrally and on tarsi 1–3; 

forefemur of male (see Figure 210) bearing anteroventral row of 

conspicuous pointed, slightly curved setae, none flattened; posteroventral margin of forefemur bearing 2 rows of denser, longer, 

of different lengths setae; foretibia straight.

Abdomen: Tergites distinctly bicolored, tergite 1 brown 

in the most part and silvery gray laterally; tergites 2–5 dark brown 

along anterior 1⁄2–2

⁄3 dorsally, medially, and on lateroventral margin, otherwise silvery gray; female with lateroventral margin of the 

abdomen brown. Male terminalia: (Figures 14–17) Presurstylus 

with basoventral process somewhat developed, length about 1⁄3 of 

horizontal process length, horizontal process shallowly curved; in 

lateral view postsurstylus widest medially with medial, anterior, 

weakly produced lobe bearing long and slender setae on external surface and a medial elongate digitiform process posteriorly; 

lateral aedeagal process less than 1⁄2 length of aedeagus, slightly 

curved subapically, bluntly rounded; aedeagus gradually tapered 

to membranous apex; phallapodeme broadly triangular in lateral 

view; hypandrium somewhat deeply concave.

Type Material. The holotype male of Paralimna 

brunneiceps is labeled “Turrucares[,] 22XII[19]09 C[osta]

R[ica,] P P Calvert/Sweeping over mud/♂ /TYPE Paralimna 

BRUNNEICEPS E.T.Cresson,Jr. [maroon; “Paralimna BRUNNEICEPS” handwritten].” The holotype is double mounted (minuten pin in a vertical rectangular card), is in good condition 

(left basal flagellomere and 3 distal tarsomeres of left leg missing; 

abdomen not dissected), and is deposited in the ANSP (6093).

Other Specimens Examined. COSTA RICA.

Alajuela: Alajuela (10°01′N, 84°13′W; 945 m), 15 Sep 

1909, P. P. Calvert (2♂, 1♀; ANSP); Turrúcares (09°57.6′N, 

84°19.2′W), 22 Dec 1909, P. P. Calvert (paratype: 1♀; allotype: 

1♀; ANSP); Turrúcares, Río Siquiares (09°58′N, 84°17′W), 19 

Dec 1909, P. P. Calvert (1♂, 1♀; ANSP). Guanacaste: Santa 

Cruz (14 km S; 10°10.4′N, 85°35.7′W; 160 m), La Casona, 23 

Jun 2001, W. N. Mathis (4♂, 1♀; USNM). Puntarenas: Manuel 

Antonio National Park (09°23.5′N, 84°08.7′W; coastal rainforest), 23–28 Aug 1986, L. Masner (1♂; CNC). San José: San José 

(09°56′N, 84°05′W), A. Alfaro (5♂, 3♀; ANSP); Ciudad Colón 

(09°55′N, 84°15′W; 800 m; Malaise trap), Dec 1989, P. Hanson 

(1♀; USNM); Rio Virilla (near Colon; 09°53.3′N, 84°16′W), 26 

Jun 2001, W. N. Mathis (4♂, 3♀; USNM); El Rodeo (09°54.6′N, 

84°16.2′W; 1880 m), 26 Jun 2001, W. N. Mathis (3♂, 1♀; 

USNM); Río Paraíso (09°33.8′N, 84°7.4′W; 350–400 m), 15–17 

Feb 2003, D. and W. N. Mathis (39♂, 6♀; USNM).

Remarks. Externally this species is similar to P. 

thomae but can be distinguished from it by the dark gray face, 

the spotted anepisternum, and the lack of flattened setae on the 

anteroventral surface of the male forefemur and foretibia, which 

are straight and without a ventral groove. Structures of the male 

terminalia also distinguish between these two species, mainly 

the lateral aedeagal process in P. aurantia sp. nov. is short and 

robust, the basoventral process of presurstylus is less well developed, and the hypandrium is deeper than in P. thomae.

4. Paralimna (Paralimna) brunneiceps Cresson

Figures 14–17, 165, 210, Map 4

Paralimna brunneiceps Cresson 1916:120 [Costa Rica. Alajuela: Turrúcares (9°57.6′N, 84°19.2′W); HT ♂, ANSP (6093)]; 1918:47 

[review; Costa Rica].

Paralimna (Paralimna) brunneiceps.—Cresson 1947:49 [review, Neotropical 

species].—Wirth 1968:15 [Neotropical catalog].—Mathis and Zatwarnicki 1995:119 [world catalog].

Diagnosis. This species is distinguished from congeners by the following characters: general coloration of body 

brown, faded; face uniformly dark brown with antennal groove 

gray; katepisternum and most of anepisternum brown; crossvein dm–cu with infuscate halo; femora and tibiae mostly dark 

brown; forefemur of male bearing anteroventral row of acutely 

pointed, curved or straight setae, none conspicuously flattened; 

posteroventral margin of forefemur bearing rows of longer, of 

different lengths setae; lateral margin of tergites 2–5 brown.

Description. Body length 3.2–3.9 mm; body generally bicolored, dark brown dorsally, lateral surfaces with dorsal 

half silvery gray and ventral half brown, sometimes yellowish gray.

Head: (See Figure 165) Frons brown with linear, yellow 

spots at lateral margins of ocellar triangle, in front of anterior 

ocellar seta, not reaching the anterior margin of frons, between 

inner vertical and interfrontal setae, and on fronto-orbits above 

dorsal proclinate fronto-orbital seta, between proclinate frontoorbital setae, and just above antennal bases; ventral fronto-orbits 

with a dark brown spot, the spot on dorsal parafacial inconspicuous. Antennal scape and pedicel dark brown, pedicel with pale microtomentum; basal flagellomere pale brown to brown according 

to incidence of light, with slender, elongate, pale setae on margin; 

arista with 11–12 long, dorsal rays. Postcranium with 2 complete 

series of postocular setae. Face uniformly dark brown, only antennal groove gray; parafacial gray and gena yellowish gray to pale 

brown; gena with some setae on middle and ventral margin; clypeus gray to yellowish gray. Eye rounded, about as wide as high, 

height more than the height of gena. Gena high, height subequal 

to length of basal flagellomere; gena-to-eye ratio 0.29–0.35.

Thorax: Mesonotum mostly brown with yellowish-gray 

areas, usually along setal tracks, 3 well-defined brown stripes 

along acrostichal line, sometimes the stripes join together forming 
number 643 • 1 9

margins of the tergites, which are predominantly brown, and by 

males having the forefemur with a posteroventral row of flattened 

setae. In specimens of P. meridionalis, the mesopleuron and lateral margins of the tergites are predominantly gray, and the male 

forefemur lacks posteroventral flattened setae. Paralimna brunneiceps, P. fellerae, P. meridionalis, P. nigropicta, and P. punctipennis all have very similar structures of the male terminalia, 

mainly the presurstylus, which has a well-developed basoventral 

process that is rounded along its ventral margin. However, P. 

brunneiceps, P. fellerae, and P. nigropicta also have flattened 

setae along the male forefemur. These species can also be easily 

MEXICO. Chiapas: Río Izapa (14°55.4′N, 92°10.7′W), 21 

Apr 1983, W. N. Mathis (8♂, 4♀; USNM).

Type Locality. Neotropical. Costa Rica. Alajuela: 

Turrúcares (09°57.6′N, 84°19.2′W).

Distribution. (Map 4) Neotropical: Costa Rica 

(Alajuela, Guanacaste, Puntarenas, San José), Mexico (Chiapas).

Remarks. Cresson (1916, 1947) wrote that P. brunneiceps is probably merely a form of P. meridionalis. Indeed, our 

study shows that these two species have similar male terminalia. 

Paralimna brunneiceps, however, can be distinguished from P. 

meridionalis by the coloration of the mesopleuron and lateral 

FIGURES 14–17. Structures of male terminalia of Paralimna (Paralimna) brunneiceps Cresson (Costa Rica. 

Alajuela): (14) epandrium, cerci, and presurstyli, posterior aspect; (15) postsurstylus, lateral aspect; (16) aedeagus, phallapodeme, and hypandrium, lateral aspect; (17) same, ventral aspect. Scale bar = 0.1 mm. (See 

Figures 2–5 for labels of structures.)
2 0 • smithsonian contributions to zoology

MAP 4. Distribution map for Paralimna (Paralimna) brunneiceps Cresson.

distinguished by the uniformly dark brown face with antennal 

grooves that are gray, by the infuscate halo over crossvein dm–

cu, and by the katepisternum and most of anepisternum being 

dark brown. In addition, the posteroventral margin of the forefemur bears rows of long setae in P. brunneiceps; the dorsal half 

of the mesopleuron is distinctly light colored, in contrast to being 

dark ventrally in P. fellerae; and in P. nigropicta, there is a dark 

brown to black velvety quadrangular spot between the antennal 

bases, the ventral fronto-orbits have a dark brown spot laterad 

to the antenna that form horizontal dark brown bands at the 

level of the antennal bases, and the male forefemur is distinctly 

concave on the distal half with a posteroventral row of setae 

present only on the basal half.

5. Paralimna (Paralimna) castanea, sp. nov.

Figures 18–21, 166, 198, Map 5

Diagnosis. This species is distinguished from congeners by the following combination of characters: body somewhat 

bicolored, although more extensively brown, especially frons 

and mesonotum, but also much of pleural areas, only katepisternum and anepimeron mostly gray, sometimes silvery white, 

anepisternum mostly to almost entirely brown, only central area 

sometimes gray with brown at base of setulae; face mostly to entirely brown or grayish brown, gray only on laterodistal margin 

and on small V-like mark in middle; gena with a small, rounded 

pale brown spot in the center; wing brownish to hyaline; crossveins r–m and dm–cu with a distinctly infuscate halo; forefemur 

of male bearing anteroventral, comb-like row of curved, moderately long, pointed setae, not flattened; and posteroventral surface of male forefemur bearing dense, somewhat irregular row of 

rather short setae along most of length.

Description. Body length 4.5–5.2 mm; body generally bicolored, brown dorsally, lateral surfaces gray, sometimes 

dark gray.

Head: (See Figure 166) Frons brown, reddish stained, 

mottled, with linear yellow spots at lateral margins of ocellar triangle, in front of anterior ocellar seta, short, not reaching the anterior margin of the frons, between inner vertical and interfrontal 
number 643 • 2 1

cellent condition, and is deposited in the USNM. Ten paratypes 

(10♂; INPA, USNM) bear the same label data as the holotype. 

Other paratypes are as follows: GUATEMALA. Izabal: Matías 

de Gálvez (also known as Santo Tomás de Castilla; 15°41.3′N, 

88°36.7′W), 14–15 Aug 1965, P. J. Spangler (18♂, 16♀; INPA, 

USNM).

TRINIDAD and TOBAGO. Tobago. St. John: Charlotteville (beach; 11°19.5′N, 60°32.9′W), 16 Apr–16 Jun 1993, 

1994, D. and W. N. Mathis (1♂; USNM); Charlotteville (5 km 

S; 11°18.9′N, 60°34.5′W), Hermitage River and beach, 22 Apr–

11 Jun 1993, 1994, D. and W. N. Mathis (4♂; INPA, USNM); 

Kings Bay Reservoir (11°17′N, 60°33′W), 15 Jun 1993, W. N. 

Mathis (1♀; USNM); Parlatuvier (creek; 11°17.9′N, 60°35′W), 

20 Apr 1994, W. N. Mathis (6♂; USNM); Speyside (Doctor 

River; 1 km NW; 11°18′N, 60°32′W), 12–13 Jun 1993, W. N. 

Mathis (7♂; INPA, USNM). St. Paul: Argyle Falls (11°15′N, 

60°35′W), 21 Apr 1994, W. N. Mathis (1♂; USNM); Roxborough (6 km NNW; 11°16′N, 60°35.4′W), 20 Apr 1994, W. N. 

Mathis (7♂, 1♀; USNM); Roxborough (6.5 km N; 11°17′N, 

60°35′W), 14 Jun 1993, W. N. Mathis (3♂; USNM). Trinidad. St. George: Lalaja Road (10°43′N, 61°17′W; streamlet), 

26 Jun 1993, W. N. Mathis (1♂, 1♀; INPA); Mount St. Benedict (10°39′N, 61°24′W; creek near base), 19 Jun 1993, W. N. 

Mathis (3♂, 1♀; USNM).

Other Specimens Examined. BELIZE. Stann 

Creek: Middlesex (17°02.4′N, 88°33.2′W; 125 m), 15 Mar 

1965, E. C. Welling (1♂, 2♀; CNC).

TRINIDAD and TOBAGO. Trinidad: Morne Blue 

(10°43′N, 61°17′W; 825 m), 23 Aug 1969, H. and A. Howden 

(1♂; CNC).

Type Locality. Neotropical. Trinidad and Tobago. 

Tobago, Charlotteville (2 km S; 11°19′N, 60°33′W).

Distribution. (Map 5) Neotropical: Belize (Stann 

Creek), Guatemala (Izabal), Trinidad and Tobago.

Etymology. The species epithet, castanea, is of 

Latin derivation and means the color of chestnuts, alluding to 

the chestnut-colored brown spots on the face.

Remarks. This species is generally similar to P. curta 

sp. nov. in having a brownish body and wing coloration but can be 

distinguished by the setation along the anteroventral and posteroventral surfaces of the male forefemur being moderately elongate 

and also by having conspicuous brown bands on the abdomen 

that are normally developed in P. castanea sp. nov. In specimens 

of P. curta sp. nov. the male forefemur lacks outstanding setae 

and the abdomen is mostly brown with gray-yellowish half-moonshaped spots on the lateral portions of tergites 2–4. Structures of 

the male terminalia of these two species are also quite different: in 

P. castanea sp. nov. the presurstylus is bifurcate and bears a welldeveloped basoventral process and the lateral aedeagal process is 

elongate and slender, about half the length of the aedeagus and 

almost straight in lateral view; in P. curta sp. nov. the presurstylus 

is not bifurcate and has a slender horizontal process that is almost 

straight and the lateral aedeagal process is very short.

setae, and on fronto-orbits between proclinate fronto-orbital setae 

and just above antennal bases; ventral front orbitals and dorsal 

parafacial with 2 dark brown spots lateral to antenna, separated 

by yellowish-gray spot. Antenna brown with pale brown microtomentum; pedicel paler on distal margin; basal flagellomere with 

slender, elongate, sparse pale setae on margin; arista with 12–13 

long, dorsal rays. Postcranium with 2 series of postocular setae, 

the second series irregular. Face mostly to entirely brown or grayish brown, gray only on laterodistal margin and on small V-like 

mark in middle; parafacial and gena silvery gray, gena with a 

small, rounded, pale brown spot in the center; clypeus reddish 

brown; gena with only few setae on middle and ventral margin. 

Eye rounded, about as wide as high, height about three times the 

height of gena. Gena high, height slightly greater than the length 

of basal flagellomere; gena-to-eye ratio 0.30–0.32.

Thorax: Mesonotum predominantly brown, reddish 

stained on dorsum, the same color of the frons, with 4 slender yellow stripes between dorsocentral rows, and numerous small yellow areas dorsally and laterally, postalar callus yellow; scutellum 

brown with lateral margins yellow, middle yellow spotted, and 

apex blackish brown. Pleurae and forecoxa gray; anepisternum 

with margins and a transverse spot in the center brown, sometimes the central spot faded; anepimeron brown on anteroventral 

corner. Wing (see Figure 198) brownish to hyaline with veins 

brown, crossveins r–m and dm–cu with a distinctly infuscate halo; 

costal-vein ratio 0.50–0.52; M-vein ratio 1.26–1.39. Legs brown 

with very sparse gray microtomentum, except tarsomeres lighter 

with yellowish-orange coloration ventrally and dorsally, the more 

distal segments darkened; forefemur of male bearing anteroventral, comb-like row of curved, moderately long, pointed setae, not 

flattened; and posteroventral surface of male forefemur bearing 

dense, somewhat irregular row of rather short setae along most of 

length; foretibia straight.

Abdomen: Tergites distinctly bicolored dorsally except 

tergite 1 wholly dark brown; tergite 2 dark brown on basal half 

and tergites 3–5 dark brown along anterior half and medially, 

on dorsum, otherwise yellowish gray; lateroventral margin of 

tergites brown. Male terminalia: (Figures 18–21) Presurstylus bifurcate, bearing basoventral process developed, horizontal process slender; in lateral view postsurstylus widest medially, medial 

anterior lobe poorly developed, bearing moderately elongate and 

slender setae; lateral aedeagal process long, about 1⁄2 length of aedeagus, slender almost straight in lateral view; aedeagus slender 

in dorsal view, slightly wider and bluntly constricted apically; 

phallapodeme broadly triangular in lateral view; hypandrium 

deeply concave, abruptly narrowed on distal 1⁄3 in lateral view 

and wider in dorsal view.

Type Material. The holotype male of Paralimna 

castanea is labeled “TOBAGO. St. John: Charlotteville (2 km 

S; 11°19′N, 60°33′W)[,] 10Jun1993[,] WNMathis/USNM ENT 

00118291 [plastic bar code label]/HOLOTYPE ♂ Paralimna 

castanea Ale-Rocha&Mathis, USNM [red].” The holotype is 

double mounted (minuten pin in a white plastic block), is in ex-
2 2 • smithsonian contributions to zoology

ish gray with 2 pale goldenbrown, parallel, wide spots extended 

along the length of face reaching the ventral margin, confluent 

and united on carina; pedicel dark brown with a whitish-gray 

spot dorsally; forefemur of male with anteroventral series of 

moderately long and slender, not flattened setae, ventral surface 

distinctly concave on distal 1⁄3, with several series of numerous, 

moderately long setae; foretibia of male straight, with dense and 

6. Paralimna (Paralimna) crinita, sp. nov.

Figures 22–25, 167, 199, Map 2

Diagnosis. This species is distinguished from congeners by the following combination of characters: frons almost 

horizontal, slightly projected over antennal bases; face yellowFIGURES 18–21. Structures of male terminalia of Paralimna (Paralimna) castanea, sp. nov. (Trinidad and Tobago. Tobago: St. John): (18) epandrium, cerci, and presurstyli, posterior aspect; (19) postsurstylus, lateral aspect; (20) aedeagus, 

phallapodeme, and hypandrium, lateral aspect; (21) same, ventral aspect. Scale bar = 0.1 mm. (See Figures 2–5 for labels of 

structures.)
number 643 • 2 3

MAP 5. Distribution map for Paralimna (Paralimna) castanea, sp. nov.

bristling setae on ventral surface; basoventral process of presurstylus falciform.

Description. Body length 5.2–6.0 mm; body generally bicolored, dark brown dorsally, lateral surfaces yellowish 

gray.

Head: (See Figure 167) Frons almost horizontal, slightly 

protracted over antennal bases, pale brown to dark brown with 

linear yellow spots at lateral margins of ocellar triangle and in 

front of anterior ocellar seta, extended to anterior margin of frons, 

between inner vertical and interfrontal seta, and on fronto-orbits 

above dorsal proclinate fronto-orbital seta, between proclinate 

fronto-orbital setae, and just above antennal bases; ventral frontoorbits and dorsal parafacial with 2 dark brown spots lateral to 

antenna, separated by yellow spot. Antenna inserted high on head; 

scape and pedicel dark brown; pedicel with a whitish-gray spot 

dorsally; basal flagellomere pale brown, covered by dense yellow 

microtomentum; basal flagellomere with slender, elongate, pale 

setae on margin; arista with 8 long, dorsal rays. Postcranium with 

2 complete series of postocular setae. Face yellowish gray with 

2 pale golden-brown, parallel, wide spots extended from below 

antenna and reaching the ventral margin of face, confluent and 

united on carina; parafacial and gena yellowish gray; gena with 

only few setae on middle and ventral margin; clypeus yellowish gray. Eye round, as wide as high, height more than twice the 

height of gena. Gena high, height longer than the length of basal 

flagellomere; gena-to-eye ratio 0.40–0.45.

Thorax: Mesonotum yellowish gray with 2 dark brown 

stripes between dorsocentral rows, plus some small spots laterally; insertion of setae brown, giving a spotted aspect to scutum; 

postpronotum, notopleuron, and supra-alar areas yellowish 

gray; scutellum yellowish gray with apex blackish brown, brown 

sprinkled by base of setae. Pleurae yellowish gray, anepisternum 

brown tinged on dorsal and ventral margins, brown sprinkled in 

middle by base of setae, katepisternum slightly brown tinged anteriorly. Wing (see Figure 199) hyaline; veins brown; costal-vein 

ratio 0.44–0.46; M-vein ratio 0.85–0.90. Legs blackish brown 

with dense silvery-gray microtomentum; tarsi lighter with some 

yellow coloration, especially ventrally, the more apical darkened; 

forefemur of male with anteroventral series of moderately long 

and slender, not flattened setae; ventral surface distinctly concave 
2 4 • smithsonian contributions to zoology

basoventral process long and falciform, about 2⁄3 length of horizontal process, horizontal process slender; in lateral view postsurstylus widest medially, medial anterior lobe developed, rather 

triangular, bearing elongate and robust setae, medial lobe truncate distally, becoming a wide process posteriorly; lateral aedeagal 

process long, about 2⁄3 length of aedeagus, strongly swollen and 

slightly arcuate in lateral view; aedeagus slender in dorsal view, 

on distal 1⁄3 and setulose, with several series of numerous moderately long setae; foretibia of male straight, densely pilose on 

ventral surface.

Abdomen: Tergites distinctly bicolored, tergite 1 dark 

brown with laterodistal corner yellow, tergites 2–5 dark brown 

along anterior half and medially, otherwise yellowish gray. 

Male terminalia: (Figures 22–25) Presurstylus bifurcate, with 

FIGURES 22–25. Structures of male terminalia of Paralimna (Paralimna) crinita, sp. nov. (Costa Rica. Puntarenas): 

(22) epandrium, cerci, and presurstyli, posterior aspect; (23) postsurstylus, lateral aspect; (24) aedeagus, phallapodeme, 

and hypandrium, lateral aspect; (25) same, ventral aspect. Scale bar = 0.1 mm. (See Figures 2–5 for labels of structures.)
number 643 • 2 5

grayish-brown stripes that extend to the ventral margin; parafacial 

and gena silvery gray; few setae on middle of gena; clypeus dark 

gray. Eye round, as wide as high, height about three times the 

height of gena. Gena high, height slightly longer than the length of 

basal flagellomere; ratio 0.32–0.38.

Thorax: Mesonotum dark brown with gray spots, a 

small spot on acrostichal line posteriorly and little small spots 

along of dorsocentral, supra-alar, and intra-alar tracks; postpronotal lobe gray; notopleuron brown with a little gray spot on anterior margin; scutellum brown with apex darkness, 1 small, paler 

spot on lateral margin and a preapical dorsal faint gray spot. Pleurae with dorsal and ventral margin of anepisternum, ventral half 

of anepimeron, and central portion of katepisternum brown, the 

remaining gray. Wing brownish colored; veins brown, crossvein 

dm–cu slightly infuscate; costal-vein ratio 0.42–0.50; M-vein ratio 

1.00–1.10. Legs: coxae and trochanters with dense gray microtomentum; femora black with fine brown microtomentum; tibiae 

dark brown; tarsi pale brown to orange yellow; forefemur without 

outstanding setae; foretibia straight.

Abdomen: Tergites brown with yellowish-gray halfmoon-shaped spots on lateral of tergites 2–4, on tergite 5 the 

spot is wide, covering almost completely the dorsum of segment; lateroventral margin of tergites brown. Male terminalia: (Figures 26–29) Presurstylus not bifurcate, with horizontal 

process slender and almost straight with slender setae along the 

ventral surface; in lateral view postsurstylus moderately wider 

medially with a medial anterior lobe prominent, bearing sparse, 

elongate, and slender setae, medial posterior process not developed; lateral aedeagal process very short, about 1⁄4 the length of 

aedeagus, robust, almost triangular, and slightly curved medially in lateral view, pointed apically; aedeagus rather robust and 

abruptly narrowed in the apex in posterior view; phallapodeme 

triangular in lateral view, prolonged distally; hypandrium gradually narrowed toward apex, moderately deeply concave in posterior view.

Type Material. The holotype male of Paralimna 

curta is labeled “BRASIL, AM[AZONAS], Ipixuna, Rio Liberdade, Estirão da Preta[,] 07°21′46.7′′S - 71°52′07.1′′W/11–

15.v.2011. Varredura[,] J.A.Rafael, J.T.Câmara, R.F Silva, 

A. Somavilla, R.Ale-Rocha/USNM ENT 00118262 [plastic bar 

code label]/HOLOTYPE ♂ Paralimna curta Ale-Rocha&Mathis, 

INPA [red].” The holotype is double mounted (glued to a paper 

triangle), is in excellent condition, and is deposited in INPA. 

Seven paratypes (4♂, 3♀; INPA, USNM) bear the same locality label data as the holotype but with varying dates and collectors: Ipixuna, Rio Liberdade, Estirão da Preta (07°21.8′S, 

71°52.1′W), 11–15 May 2011, Varredura, J. A. Rafael, J. T. 

Câmara, R. F. Silva, A. Somavilla, R. Ale-Rocha, R. W. H. 

Honegger, R. S. G. Hutchings. Other paratypes are as follows: 

BRAZIL. Amazonas: Rio Gregório, Comunidade Lago Grande 

(07°10.2′S, 70°49.2′W; some from light and Malaise traps), 

18–23 May 2011 (31♂, 11♀; INPA, USNM).

Type Locality. Neotropical. Brazil. Amazonas: Ipixuna, Rio Liberdade, Estirão da Preta (07°21.8′S, 71°52.1′W).

progressively narrowed apically; phallapodeme rather slender in 

lateral view; hypandrium moderately deeply concave.

Type Material. The holotype male of Paralimna 

crinita is labeled “COSTA RICA. Punta[renas].: San Pedrillo 

(8°37.2′N, 83°44.1′W), 12–15 August 2001, D.& Wayne N. 

Mathis/USNM ENT 00190644 [plastic bar code label]/HOLOTYPE ♂ Paralimna crinita Ale-Rocha&Mathis, USNM [red].” 

The holotype is directly pinned, is in excellent condition, and is 

deposited in the USNM. Eight paratypes (2♂, 6♀; INPA, USNM) 

bear the same label data as the holotype.

Type Locality. Neotropical. Costa Rica. Puntarenas: 

San Pedrillo (08°37.2′N, 83°44.1′W).

Distribution. (Map 2) Neotropical: Costa Rica 

(Puntarenas).

Etymology. The species epithet, crinita, is of Latin 

derivation and means hairy, alluding to the setose ventral surface 

of the male foretibia.

Remarks. Structures of the male terminalia of this species are very similar to those of P. adunca sp. nov., and we discuss 

distinguishing characters in the “Remarks” for the latter species.

7. Paralimna (Paralimna) curta, sp. nov.

Figures 26–29, 168, Map 6

Diagnosis. This species is distinguished from congeners by the following combination of characters: face dark gray 

laterally with margins dark brown and 3 brown spots, a single 

triangular spot on carina, slightly shorter than length of basal 

flagellomere and apex 3× width of base, and 2 parallel, slender, vertical grayish-brown stripes that extend to the ventral 

margin; wing brownish colored; abdomen mostly brown with 

gray-yellowish half-moon-shaped spots on lateral of tergites 2–4; 

presurstylus with horizontal process slender, almost straight; lateral aedeagal process very short.

Description. Body length 4.0–5.5 mm; body generally bicolored, dark brown dorsally, lateral surfaces gray, sometimes yellowish gray.

Head: (See Figure 168) Frons brown with linear yellowish spots from lateral margins of ocellar triangle to frontal anterior 

ocellar seta and extended to near anterior margin of the frons, 

1 stripe between inner vertical and interfrontal seta, on frontoorbits between proclinate fronto-orbital setae, and just above antennal bases; ventral fronto-orbits with a black spot and dorsal 

parafacial with a dark gray spot, lateral to antenna, separated by 

silvery-gray spot. Antenna brown; pedicel with dorsal rounded 

silvery-gray spot, with 1 dorsal, thicker seta and some short setae 

on the margin; basal flagellomere densely covered by pale gold 

microtomentum and slender pale setae on margin, not distinctly 

elongate; arista with 13–15 long, dorsal rays. Postcranium with 

1 complete series of postocular setae. Face dark gray, laterally 

with margins dark brown and 3 brown spots, a single triangular spot on carina, slightly shorter than length of basal flagellomere and apex 3× width of base, and 2 parallel slender vertical 
2 6 • smithsonian contributions to zoology

Distribution. (Map 6) Neotropical: Brazil (Amazonas).

Etymology. The species epithet, curta, is of Latin derivation and means short, alluding to the very short aedeagal process.

Remarks. The facial coloration of this species is 

similar to that of P. meridionalis but can be easily distinguished 

from the latter by the shape of the structures of the male terminalia and the pattern of body coloration. In P. curta sp. nov. 

the abdomen is mostly brown with gray-yellowish half-moonshaped spots on the lateral margins of tergites 2–4; the presurstylus is not bifurcate and with a slender horizontal process that 

is almost straight; the lateral aedeagal process is very short; in 

lateral view the postsurstylus has a prominent medioanterior 

lobe, and the medial posterior process is not well developed. 

In P. meridionalis the tergites have better developed brown 

bands that extend dorsally, tergite 2 has a basal brown band 

FIGURES 26–29. Structures of male terminalia of Paralimna (Paralimna) curta, sp. nov. (Brazil. Amazonas): (26) epandrium, cerci, and presurstyli, posterior aspect; (27) postsurstylus, lateral aspect; (28) aedeagus, phallapodeme, and hypandrium, lateral aspect; (29) same, ventral aspect. Scale bar = 0.1 mm. (See Figures 2–5 for labels of structures.)
number 643 • 2 7

rather triangular spots between larger facial setae; mesonotum 

mostly brown, with 3 dorsal brown stripes coalescing for most 

of the length and forming a wide brown dorsal area; forefemur 

of male with anteroventral row of very short, sparse, acutely 

pointed, curved, not flattened setae; posteroventral surface with 

complete series of somewhat elongate, robust conspicuous setae, 

more numerous, with 2–3 series, on medial portion.

Description. Body length 3.2–3.7 mm; body generally bicolored, brown dorsally, lateral surfaces gently dark 

gray.

Head: (See Figure 169) Frons brown, reddish brown 

tinged on anterior half, with linear, poorly marked pale yellow 

spots at lateral margins of ocellar triangle, in front of anterior 

ocellar seta, not reaching the anterior margin of frons, and on 

fronto-orbits between proclinate fronto-orbital setae and just 

above antennal bases; ventral fronto-orbits and dorsal parafacial with dark brown and gray spots, respectively, separated by 

yellowish-gray area. Antennal scape and pedicel dark brown, 

pedicel with distal margin paler; basal flagellomere pale brown 

to yellowish brown according to incidence of light, with slender, 

that extends to the posterior margin in the middle on each side 

(otherwise the tergites are silvery or dark gray); the presurstylus 

is bifurcate with a short, basoventral process that is somewhat 

tapered ventrally, the horizontal process is fairly arcuate, sometimes rather straight; the lateral aedeagal process is about 1⁄2 

or less the length of the aedeagus, which in lateral view is thin 

and weakly curved medially; in lateral view, the postsurstylus 

has a medioanterior, weakly produced lobe and a small, medial, 

pointed process posteriorly.

Similarities and differences between P. curta sp. nov. and P. 

castanea sp. nov. were discussed in the “Remarks” for the latter 

species.

8. Paralimna (Paralimna) ecuadoriensis, sp. nov.

Figures 30–33, 169, Map 6

Diagnosis. This species is distinguished from congeners by the following characters: face gray to dark gray with 3 

brown spots, a single rectangular spot on carina and 2 small 

MAP 6. Distribution map for Paralimna (Paralimna) curta, sp. nov. and P. (Paralimna) ecuadoriensis, sp. nov.
2 8 • smithsonian contributions to zoology

Distribution. (Map 6). Neotropical: Ecuador (El 

Oro, Guayas, Manabi).

Etymology. The species epithet, ecuadoriensis, alludes to the country of Ecuador where the type series was collected.

Remarks. We did not find external distinguishing 

characters for this species and have had to rely on structures of 

the male terminalia to make accurate identifications. The most 

conspicuous characters of the terminalia are the postsurstylus 

with a wide, rounded, medioanterior process; the medioposterior process is very prominent and triangular; the aedeagus and 

phallapodeme are elongated; and the lateral aedeagal process has 

a truncated apex (see Figures 31, 32).

9. Paralimna (Paralimna) fellerae Mathis

Figures 34–49, 170, Map 7

Paralimna (Paralimna) fellerae Mathis, 1997:54 [Belize. Stann Creek: Glovers Reef, Middle Cay; HT ♂, USNM].

Diagnosis. This species is distinguished from congeners by the following combination of characters: face silvery gray 

with 3 golden-brown spots, a single linear area on the carina and 

2 rounded spots between larger facial seta; pleural area distinctly 

lighter on dorsal half (dorsal pleurites gray, only thin band on 

ventral margin of anepisternum and anepimeron brown) and 

darker on ventral half (ventral pleurites brown); forefemur of 

male with anteroventral, comb-like row of flattened setae; crossveins r–m and dm–cu infuscate.

Description. Body length 2.20–3.30 mm; body 

generally bicolored, dark brown dorsally and ventrally, lateral 

medial surfaces mostly gray, sometimes blackish gray.

Head: (Figures 34–38, 170) Frons mostly brown to dark 

brown with linear, silvery-white spots at lateral margins of ocellar 

triangle, in front of anterior ocellar seta, and on fronto-orbits between proclinate fronto-orbital setae and, in some specimens, just 

above antennal bases; ventral fronto-orbits and dorsal parafacial 

with 2 black spots lateral to antenna, separated by silvery-white 

spot. Antenna black with some gray microtomentum medially; 

basal flagellomere with long, slender, dorsal pale setae; arista with 

11 long, dorsal rays. Postcranium with 2 complete series of postocular setae. Face, parafacial, and gena mostly silvery gray; face 

with 3 golden brown spots, a single linear area on the carina and 

2 rounded spots between larger facial seta; gena with only few 

setae on middle and ventral margin; clypeus darker gray than face. 

Eye rounded, about as wide as high, height more than twice the 

height of gena. Gena high, height subequal to length of basal flagellomere; gena-to-eye ratio 0.34–0.45.

Thorax: (Figures 39–45) Mesonotum generally brown, at 

most slightly lighter in color than frons; with some silvery-gray areas, 

usually along setal tracks, becoming grayer laterally through postpronotum and notopleuron; scutellar margins gray, apex blackish 

brown. Pleural area distinctly lighter on dorsal half (dorsal pleurites 

elongate, pale setae on margin; arista with 12–14 long, dorsal 

rays. Postcranium with 2 complete series of postocular setae. 

Face gray to dark gray with 3 brown spots, a single rectangular 

spot on carina and 2 small rather triangular spots between larger 

facial setae; parafacial and gena silvery gray; gena with only few 

setae on middle and ventral margin; clypeus silvery gray. Eye 

rounded, about as wide as high, height about three times the 

height of gena. Gena high, height subequal to length of basal 

flagellomere; gena-to-eye ratio 0.33–0.35.

Thorax: Mesonotum mostly brown, the 3 dorsal brown 

stripes coalescing for most of the length, forming a wide brown 

dorsal area with few small gray areas or slender stripes along setal 

tracks and laterally, base of setae brown; postpronotum and supraalar areas gray; scutellum mostly brown dorsally, with small pale 

gray spots dorsally, apex blackish brown and gray laterally. Pleurae gray; anepisternum with 1 brown spot on dorsal margin and 

ventral margin slightly brown tinged, stains of bases of setae brown 

in middle. Wing hyaline; veins brown; costal-vein ratio 0.50–0.51; 

M-vein ratio 0.95–0.97. Legs brown with sparse silvery-gray microtomentum; tarsi lighter with some yellow coloration, especially 

ventrally; forefemur of male with anteroventral row of very short, 

sparse, acutely pointed, curved, not flattened setae; posteroventral 

surface with complete series of somewhat elongate, robust, conspicuous setae, more numerous, with 2–3 series, on medial portion; 

male foretibia straight with rather ruffled setae on ventral surface.

Abdomen: Tergites distinctly bicolored; tergite 1 brown 

dorsally and on lateroventral margin, pale brown on anterior corner; tergites 2–5 brown along anterior half and medially, otherwise 

gray. Male terminalia: (Figures 30–33) Presurstylus not bifurcate, 

with horizontal process slender and shallowly curved, gradually 

narrowed toward distal apex; in lateral view postsurstylus widest 

medially with a medial, anteriorly wide lobe, bearing moderately 

long strong setae, medial posterior process very prominent, triangular; lateral aedeagal process about 1⁄2 length of aedeagus, slender, 

slightly curved medially in lateral view; aedeagus slender, narrowed 

on distal 1⁄3 in the membranous apex; phallapodeme triangular, 

somewhat narrow in lateral view; hypandrium moderately deep.

Type Material. The holotype male of Paralimna 

ecuadoriensis is labeled “ECUADOR: Guayas Pr[ovincia]. 

Machala (49.5 km. NNE.)[,] 40 m. elev[ation].[,] 13 Jan1978[,] 

WNMathis/USNM ENT 00118289 [plastic bar code label]/

HOLOTYPE ♂ Paralimna ecuadoriensis Ale-Rocha&Mathis, 

INPA [red].” The holotype is double mounted (minuten pin in a 

white plastic block), is in very good condition (some setae missing), and is deposited in the USNM. Nine paratypes (3♂, 6♀; 

INPA, USNM) bear the same label data as the holotype. Other 

paratypes are as follows: ECUADOR. El Oro: Pasaje (6 km 

E; 03°18.1′S, 79°47.1′W), 13 Jan 1978, W. N. Mathis (1♂; 

USNM). Manabi: Pedro Carbo (45 km NW; 01°35′S, 80°30′W), 

11 Jan 1978, W. N. Mathis (2♀; USNM); Bahia de Caraquez 

(35.6 km E; 00°35.8′S, 80°13.5′W; 20 m), 8–9 Jan 1978, W. N. 

Mathis (1♂; USNM).

Type Locality. Neotropical. Ecuador. Guayas: 

Machala (49.5 km NNE; 02°40.6′S, 79°37.7′W; 40 m).
number 643 • 2 9

FIGURES 30–33. Structures of male terminalia of Paralimna (Paralimna) ecuadoriensis, sp. nov. (Ecuador. Guayas): 

(30) epandrium, cerci, and presurstyli, posterior aspect; (31) postsurstylus, lateral aspect; (32) aedeagus, phallapodeme, 

and hypandrium, lateral aspect; (33) same, ventral aspect. Scale bar = 0.1 mm. (See Figures 2–5 for labels of structures.)

row of flattened setae, setae short, length less than width of femur; 

and posteroventral surface of forefemur bearing some setae but not 

as a distinct linear patch at basal half; foretibia straight.

Abdomen: Tergites distinctly bicolored, dark brown 

along anterior half and medially, otherwise gray, except tergite 

1 brown with lateral distal corner silvery gray. Male terminalia: 

(Figures 46–49) Presurstylus bifurcate, with horizontal process 

gray, only thin band on ventral margin of anepisternum and anepimeron blackish brown) and darker on ventral half (ventral pleurites blackish brown). Wing semi-hyaline; veins brown; crossveins 

r–m and dm–cu infuscate; costal-vein ratio 0.53–0.55; M-vein ratio 

1.00–1.22. Legs generally dark colored; coxae, femora, and tibiae 

blackish brown; tarsi lighter with some yellowish-orange coloration, especially ventrally; forefemur with anteroventral, comb-like 
3 0 • smithsonian contributions to zoology

the holotype. The holotype is double mounted (minute pin in 

a block of plastic), is in excellent condition, and is deposited 

in the USNM. Other paratypes are as follows: BELIZE. Belize: Turneffe Islands, Calabash Cays (17°17′N, 87°48′W), Mar 

1993, W. N. Mathis (2♂; USNM). Stann Creek: Wee Wee Cay 

(16°45.9′N, 88°08.6′W), Nov 1987, Mar 1988, W. N. Mathis, 

C. Feller (1♂, 1♀; USNM).

Other Specimens Examined. Nearctic. UNITED 

STATES. Florida. Marion: Ocala National Forest, Juniper 

Springs, Hwy. 40 (29°11′N, 81°42.′W), 22 Apr 1967, B. V. Peterson (1♂; CNC).

Neotropical. BELIZE. Stann Creek: Carrie Bow Cay 

(16°48.1′N, 88°04.9′W), 8–10 Apr 1977, R. J. Larson (1♀; 

long, slender, shallowly curved, extended medially, basoventral 

process short, wide, length less than 1⁄4 horizontal process; postsurstylus widest medially with medial, anteriorly produced wide lobe; 

lateral aedeagal process less than 1⁄2 length of aedeagus, thinly developed, parallel sided, slightly curved subapically, bluntly rounded; 

aedeagus gradually tapered to membranous apex; phallapodeme 

broadly triangular in lateral view; hypandrium deeply concave.

Type Material. The holotype male of Paralimna 

fellerae is labeled “BELIZE. Stann Creek District: Glover’s 

Reef, Middle Cay[,] 28 July 1989, W.N. Mathis, H.Williams/

HOLOTYPE ♂ Paralimna fellerae W. N. Mathis USNM [“♂

Paralimna fellerae” handwritten; red].” The allotype female and 

42 paratypes (30♂, 12♀; USNM) bear the same label data as 

FIGURES 34–39. Scanning electron micrographs of Paralimna (Paralimna) fellerae Mathis (Belize. Stann Creek); scale length for each figure 

follows in parentheses (scale bar for all photographs is in Figure 36): (34) head, lateral aspect (0.43 mm); (35) face, anterior aspect (0.30 mm); 

(36) antenna, lateral aspect (150 mm); (37) ommatidia and occasional interfacetal setae of compound eye, lateral aspect (30 μm); (38) enlargement of same, lateral aspect (6.0 μm); (39) mesonotum, dorsal aspect (0.50 mm).
number 643 • 3 1 FIGURES 40–45. Scanning electron micrographs of Paralimna (Paralimna) fellerae Mathis (Belize. Stann Creek; scale length in parentheses; scale bar for all photographs in Figure 43): (40) scutellum, dorsal aspect (231 μm); (41) pleural sclerites, lateral aspect (0.50 mm); (42) notopleuron, lateral aspect (167 μm); (43) left foreleg, anterior aspect (250 μm); (44) enlargement of same, anterior aspect (86 μm); (45) left foreleg, posterior aspect (0.27 mm).
3 2 • smithsonian contributions to zoology

HONDURAS. Atlantida: Corrozal (15°46′N, 85°41′W), 

30 Aug 1967, G. Lacy (1♂, 2♀; USNM). Cortés: Puerto 

Cortés/Omoa (15°49′N, 87°56.2′W), 26 Sep 1995, D. and W. N. 

Mathis (4♂, 3♀; USNM).

PANAMÁ. Cocle: Playa Santa Clara (08°22.4′N, 

80°06.3′W), 2 Jul 1967, W. W. Wirth (1♂, 4♀; USNM). Darien:

Patino Point (08°15′N, 78°17.4′W), 1–6 Sep 1952, F. S. Blanton 

(5♂, 1♀; USNM). Guna Yala: San Blas, Rio Carti Grande near 

coast (09°27.3′N, 78°57.8′W), 2 Mar 1985, O. S. Flint Jr., J. E. 

Louton (2♂; USNM).

USNM); Mullins River (17 km N Dangriga; 17°06.2′N, 

88°17.8′W), 29 Mar 1988, W. N. Mathis (1♂, 1♀; USNM).

COSTA RICA. Limón: Puerto Vargas (09°43.9′N, 

82°48.9′W; beach), 27–28 Jun 2001, W. N. Mathis (4♂, 1♀; 

USNM); Westfalia (4 km S, 09°54.5′N, 82°59′W, beach), 27 Jun 

2001, W. N. Mathis (3♂, 2♀; USNM). Puntarenas: San Pedrillo 

(08°37.2′N, 83°44.1′W), 12–15 Aug 2011, D. and W. N. Mathis 

(26♂; 2♀; USNM); Jacó (09°37.5′N, 84°38.5′W; beach), 13 Jun 

2003, D. and W. N. Mathis (1♂; USNM); Dominical (09°14.8′N, 

83°51.4′W), 12 Jun 2003, D. and W. N. Mathis (1♂; USNM).

FIGURES 46–49. Structures of male terminalia of Paralimna (Paralimna) fellerae Mathis (Belize. Stann Creek): (46) 

epandrium, cerci, presurstyli, and postsurstylus, posterior aspect; (47) same, lateral aspect; (48) phallapodeme and 

aedeagus, lateral aspect; (49) postsurstylus, lateral aspect. Scale = 0.1 mm. (See Figures 2–5 for labels of structures.)
number 643 • 3 3

MAP 7. Distribution map for Paralimna (Paralimna) fellerae Mathis.

Type Locality. Neotropical. Belize. Stann Creek: 

Glover’s Reef, Middle Cay (16°44.5′N, 87°48.7′W).

Distribution. (Map 7) Nearctic: United States 

(Florida). Neotropical: Belize (Belize, Stann Creek), Costa Rica 

(Limón, Puntarenas), Honduras (Atlantida, Cortés), Panamá 

(Cocle, Darien, Guna Yala).

Remarks. All specimens of this species came from 

Wee Wee Cay and were collected from a large muddy area with 

a covering mat of blue-green algae. Structures of the male terminalia of P. fellerae are very similar to those of P. brunneiceps, 

P. meridionalis, P. nigropicta, and P. punctipennis. We discuss 

distinguishing characters of these species in the “Remarks” for 

P. brunneiceps. The external colorational pattern of P. fellerae

is also very similar to that of P. punctipennis and can easily be 

confused with the latter. Both species have a mesopleuron that is 

distinctly light colored dorsally in contrast to a darkened ventral 

half. The wing of P. fellerae, however, lacks short perpendicular 

stump veins along the second section of vein M (the wing of P. 

punctipennis has a stump vein or two), and the ventral surface 

of the male forefemur has flattened setae, unlike P. punctipennis.

10. Paralimna (Paralimna) flexineuris Cresson

Figures 50–53, 171, 211, Map 8

Paralimna flexineuris Cresson 1916:111 [Peru. Lima: Chosica (11°56.2′S, 

76°42.1′W); HT ♂, USNM (19684)]; 1947:49–50 [review, Neotropical species].—Wirth 1968:15 [Neotropical catalog].—Mathis and Zatwarnicki 1995:120 [world catalog].

Diagnosis. This species is distinguished from congeners by the following combination of characters: face usually 

wholly whitish gray, without spots, but sometimes with two parallel, distally widened, very close spots forming a large, golden, 

rather trapezoidal spot on carina, extended until near ventral 

margin of face; frons and dorsum of thorax orange tinged, frons 

with conspicuous dark gray rounded spot on ventral frontoorbital; scutum with 2 golden-brown stripes between dorsocentral rows; pleurae whitish gray, only a small, inconspicuous, 

lightly brown to orange-brown stripe on dorsal margin of anepisternum; forefemur with ventral surface convex, anteroventral 
3 4 • smithsonian contributions to zoology

phallapodeme wider on basal 2⁄3 and slender on the remainder; 

hypandrium deeply concave; cerci long and narrowed, almost 3× 

longer than wide.

Type Material. The holotype male of Paralimna 

flexineuris is labeled “TD 4197 [written in pencil]/Chosica[,] 

Peru [handwritten]/ CHTTowsend coll/HoloTYPE Paralimna 

flexinervis [sic,] ETCressonJr [pink; “Paralimna flexinervis” 

handwritten]/Type No 19684 U S N M [red; “19684” handwritten].” The holotype is pinned directly, is in fair condition 

(apex of left wing torn, abdomen missing), and is deposited in 

the USNM (19684). Two paratypes bear the same data as the 

holotype (1♀; ANSP), (1♀ allotype; USNM).

Other Specimens Examined. CHILE. Arica and 

Parinacota: Arica (18°28.5′S, 70°18.9′W), Oct 1932, F. Jaffuel 

(1♂; USNM).

ECUADOR. El Oro: Pasaje (6 km E; 03°18.1′S, 79°47.1′W), 

13 Jan 1978, W. N. Mathis (11♂, 4♀; USNM); Rio Grande (15 

km S Zaruma; 03°42.8′S, 79°36.8′W; 850 m), 9 Apr 1965, L. E. 

Peña (2♂, 4♀; CNC). Guayas: Boliche (14.5 km S; 02°07.7′S, 

79°35.5′W), 14 Jan 1978, W. N. Mathis (4♂; USNM); El Triunfo 

(60 km N Guayaquil; 01°55.3′S, 80°0.7′W), 11 Feb 1973, M. A. 

Deyrup (1♀; USNM); Guayaquil (airport; 02°09.5′S, 79°53′W), 

27 Feb 1973, M. A. and E. Deyrup (8♂; USNM); Machala (49.5 

km NNE; 02°40′S, 79°13.4′W; 40 m), 13 Jan 1978, W. N. Mathis 

(4♂, 12♀; USNM). Manabi: Chone (00°41.7′S, 80°05.3′W), 9 

Jan 1978, W. N. Mathis (4♂; USNM); Pedro Carbo (01°48.7′S, 

80°13.9′W), 11 Jan 1978, W. N. Mathis (4♂, 7♀; USNM); Bahia 

de Caraquez (35.6 km E; 00°35.8′S, 80°13.5′W), 8–9 Jan 1978, 

W. N. Mathis (5♂, 3♀; USNM).

MEXICO. Morelos: Cuernavaca (12 km E; 18°58.1′N, 

99°07.7′W; 1310 m), 14 Aug 1954, J. G. Chillcott (1♂, 3♀; 

CNC). Sonora: Ciudad Obregón (27°29.6′N, 109°56.3′W), 17 

May 1961, W. W. Gibson (1♀; CNC). Tamaulipas: Morelos 

Antiguo (22°33.1′N, 99°05.1′W; 305 m), 21 Jul 1954, J. G. 

Chillcott (1♀; CNC).

PERU. Lima: Callao (12°02′S, 77°08′W), 17 Nov 1950, 

E. S. Ross, A. E. Michelbacher (1♂; USNM); Cocachacra (18 km 

E Chosica; 11°55.3′S, 76°32′W), 11 Feb 1984, W. N. Mathis 

(1♂; USNM); Lima (12°01.6′S, 77°04.6′W), 2 Feb 1984, W. N. 

Mathis (1♂; USNM).

VENEZUELA. Lara: Barquisimeto (S Medina; 10°03.8′N, 

69°19.3′W; on vegetation, sugarcane field), 7 Jun 1965, I. Gaud, 

L. F. Martorell (1♂; USNM); Carora (10°10.7′N, 70°04.4′W), 5 

Apr 1981, L. Hollenberg, A. S. Menke (1♂; USNM).

Type Locality. Neotropical. Peru. Lima: Chosica 

(11°56.2′S, 76°42.1′W).

Distribution. (Map 8) Nearctic: Mexico (Morelos, 

Sonora, Tamaulipas). Neotropical: Chile (Arica and Parinacota), 

Ecuador (El Oro, Guayas, Manabi), Peru (Lima), Venezuela 

(Lara).

Remarks. Cresson (1916) described this species 

from a specimen that was missing its abdomen and included 

only females in the paratype series. We have examined the holotype and two paratypes, one being the allotype female. The 

series of flattened setae short, posteroventral surface with rather 

numerous setae concentrated in basal 1⁄3, and foretibia with ventral surface slightly wavy.

Description. Body length 3.6–4.2 mm; body generally 

bicolored, pale orange brown dorsally, lateral surfaces light gray.

Head: (See Figure 171) Frons brown, orange stained, 

grayish on anterior margin, with linear gray spots at lateral margins of ocellar triangle, in front of anterior ocellar seta and weakly 

extended anteriorly but not reaching the anterior margin of the 

frons, and on fronto-orbits between proclinate fronto-orbital setae 

and just above antennal bases; ventral front orbitals and dorsal 

parafacial with 2 dark gray spots lateral to antenna, separated 

by silvery-gray spot. Antenna brown with gray microtomentum; 

pedicel with brown microtomentum on dorsum, with 1 dorsal, 

thicker seta and some short setae on the margin; basal flagellomere with slender, sparse, pale setae on margin; arista with 14 

long, dorsal rays. Postcranium with 2 complete series of postocular setae. Face, parafacial, gena, and clypeus densely silvery 

gray, face usually without spots but sometimes with two parallel, 

distally widened, very close spots forming a large, golden, rather 

trapezoidal spot on carina, extended until near ventral margin of 

face; gena with only few setae on middle and ventral margin. Eye 

rounded, slightly higher than wide, height about twice the height 

of gena. Gena high, height subequal the length of basal flagellomere; gena-to-eye ratio 0.40–0.44.

Thorax: Mesonotum whitish gray, orange stained on 

dorsum, the same color of the frons, with 2 golden brown stripes 

between dorsocentral rows, visible according to incidence of light, 

and numerous small silvery-gray areas dorsally, lateral of scutum silvery gray, except the notopleuron orange tinged; scutellum with lateral margins gray and apex blackish brown. Pleurae 

and forecoxa light gray, dorsal margin of anepisternum lightly orange brown. Wing semi-hyaline, slightly orange tinged; veins orange tinged; costal-vein ratio 0.46–0.48; M-vein ratio 0.90–0.97. 

Legs brown with sparse gray microtomentum, except tarsomeres 

lighter with yellowish-orange coloration ventrally and dorsally; 

forefemur of male (see Figure 211) with ventral surface convex, 

anteroventral series of flattened setae short, of same length, anteroventral surface with rather numerous setae concentrated in 

basal 1⁄3; foretibia with ventral surface slightly wavy.

Abdomen: (Lost in holotype.) Tergites distinctly bicolored, dark brown along anterior half and medially on dorsum, 

otherwise whitish gray; anterior brown portion of tergite 2 reduced to 3 small spots, 1 in middle and 2 dorsolateral. Male 

terminalia: (Figures 50–53) Presurstylus bearing long, slender setae, with basoventral process somewhat developed with 

rounded ventral margin, length about 1⁄3 of horizontal process 

length; horizontal process shallowly curved; in lateral view postsurstylus widest with medial anteriorly and posteriorly produced 

lobes, anterior lobe bearing long and robust setae on ventral 

margin; lateral aedeagal process less than 1⁄2 length of aedeagus, 

somewhat robust in lateral view, slightly curved in middle 

and rounded apically; aedeagus slightly constricted in the apical 1⁄3 and gently widened to membranous apex; in lateral view 
number 643 • 3 5

This species is similar to P. thomae, especially the male forefemur, but can be distinguished from it by the orange coloration 

of the frons and scutum and the gray face. The characters of the 

male terminalia also contribute to distinguishing between these 

two species. In P. flexineuris, the presurstylus has a basoventral 

process with a rounded ventral margin, the anterior lobe of the 

postsurstylus bears a long, strong seta on the ventral margin, 

and the lateral aedeagal process is somewhat robust and slightly 

paratypes, however, are not conspecific with the holotype. 

Other specimens have been erroneously identified as this species, probably based on study of the paratypes (e.g., Cresson, 

1947). Despite lacking its abdomen, the other diagnostic characters of the holotype are adequate to distinguish this species 

from congeners and to identify other conspecific specimens, 

thus allowing us to illustrate and analyze structures of the male 

terminalia.

FIGURES 50–53. Structures of male terminalia of Paralimna (Paralimna) flexineuris Cresson (Venezuela. Lara): (50) 

epandrium, cerci, and presurstyli, posterior aspect; (51) postsurstylus, lateral aspect; (52) aedeagus, phallapodeme, and 

hypandrium, lateral aspect; (53) same, ventral aspect. Scale bar = 0.1 mm. (See Figures 2–5 for labels of structures.)
3 6 • smithsonian contributions to zoology

MAP 8. Distribution map for Paralimna (Paralimna) flexineuris Cresson.

curved in the middle with rounded apices. In P. thomae, the 

presurstylus has a basoventral process with a truncate ventral 

margin, the anterior lobe of the postsurstylus bears short and 

very thin setae along the ventral margin, and the lateral aedeagal 

process is somewhat slender, almost straight, and has a pointed 

apex.

Cresson (1916) wrote about the similarity of this species 

and P. texana and also noted differences found in the setation of 

the male foreleg. Along the ventral surface the setae are located 

on the basal half in P. flexineuris; in P. texana, the series extends 

to the femoral apex. Both species have numerous, conspicuously 

flattened setulae along the anteroventral surface of the forefemur. Additionally, the face of P. flexineuris lacks a spot between 

the antennae, whereas in P. texana there is a golden-brown, 

elongate, and slender spot between the antennae. The abdominal pattern may also help in distinguishing these two species. 

In P. texana the basal brown tergal bands are reduced and very 

slender and are lacking on tergite 5. Additionally in P. texana,

the presurstylus has a basoventral process with a pointed ventral 

margin that bears strong setae (with rounded ventral margin and 

thin setae in P. flexineuris), the posterior lobe of the postsurstylus is short and pointed (wide and rounded in P. flexineuris), 

and the lateral aedeagal process is very thin and almost straight 

(somewhat robust and slightly curved in middle with rounded 

apex in P. flexineuris).

11. Paralimna (Paralimna) fulgifrons, sp. nov.

Figures 54–57, 172, 200, Map 9

Diagnosis. This species is distinguished from congeners by the following combination of characters: brown 

wing with hyaline halos; face brownish bronze, in profile carina strongly protuberant; scape and pedicel velvety black, 

basal flagellomere orange colored; anterior half of frons velvety black lateral of antenna and with a ovate, shiny black, 

slightly protuberant area between bases of antennae and anterior ocellus; crossvein dm–cu concave, curved to base; lateral aedeagal process short and slender, about 1⁄3 length of 

aedeagus.
number 643 • 3 7

Rocha&Mathis, USNM [red].” The holotype is double mounted 

(minuten pin in a red block of plastic), is in excellent condition, 

and is deposited in the USNM. Twenty-four paratypes (21♂, 

3♀; INPA, USNM) bear the same label data as the holotype. 

Other paratypes are as follows: BRAZIL. Amazonas: Barcelos, 

Bacuquara (00°09.1′S, 63°10.6′W; Malaise trap; igarapé), Jul 

2007, A. S. Filho, T. Krolow (1♂, 3♀; INPA); Barcelos, Serrinha 

(00°25.1′S, 63°23.1′W; Malaise trap; forest), Jul–Aug 2007 

(1♂, 2♀; INPA); Barcelos, Igarapé Ererê/Coruja (00°06.3′N, 

63°51′W; Malaise trap), 18–25 Jun 2008, F. F. Xavier-Filho (1♀; 

INPA); Rio Demeni, Pirico (00°19.5′S, 62°47.3′W; Armadilha 

suspense lamina d’água), Aug 2008, A. Silva, R. Machado (14♀, 

1♂; INPA, USNM).

PERU. Loreto: Iquitos (12 km W; 03°48.4′S, 73°20.5′W), 

16 Feb 1984, W. N. Mathis (5♂; USNM); Iquitos (14 km W; 

03°52.1′S, 73°28.2′W), 16 Feb 1984, W. N. Mathis (2♂; USNM).

VENEZUELA. Territorio Federal Amazonas: Basecamp, 

Cerro de la Neblina (00°54.4′N, 66°10′W; 140 m; Malaise trap 

over small stream at east side of basecamp), 20–24 Mar 1984, 

O. S. Flint Jr., J. E. Louton (1♂; USNM).

Other Specimens Examined. BRAZIL. Pará:

Belém (01°26.7′S, 48°35.2′W), Sep 1952, Th. Dobzhansky (2♂, 

2♀; USNM).

Type Locality. Neotropical. Peru. Loreto: Río 

Momon (~25 km NW Iquitos; 03°41.2′S, 73°15.9′W).

Distribution. (Map 9) Neotropical: Brazil (Pará, 

Amazonas), Peru (Loreto), Venezuela (Territorio Federal Amazonas).

Etymology. The species epithet, fulgifrons, is of 

Latin derivation and means flash (fulgeo) and frons (frons), alluding to the shiny mark on the frons.

Remarks. This species is part of a monophyletic 

group that includes P. guttata sp. nov. and P. stellata sp. nov. 

The group shares a concave, basally curved crossvein dm–cu, 

eyes that are higher than wide, a brown wing with hyaline halos, 

and a slightly protuberant facial carina among others characters. Structures of the male terminalia are very similar with a 

simple, horizontal presurstylar process that lacks a developed 

basoventral process and a shortened lateral aedeagal process. 

Paralimna fulgifrons sp. nov., however, can be easily distinguished from the other species of the group by the ovate, shiny 

black, slightly protuberant area between the antennal bases and 

the anterior ocellus, and the basal flagellomere is tinged with 

orange. Specimens from Venezuela have a complete series of 

flattened setulae on the male forefemur, covering the entire anteroventral surface, and the scutellum is darkened. With respect 

to male terminalia, the most conspicuous difference between 

this species and the other mentioned species can be seen in the 

length of the lateral aedeagal process, which is shorter, in the 

setae of the horizontal process of the presurstylus, which are 

slender and short (long and robust in P. stellata sp. nov.), and 

in the anteromedial lobe of the postsurstylus, which is large 

(small in P. guttata sp. nov.).

Description. Body length 3.5–4.7 mm; body predominantly dark brown with small pale gray spots dorsally and 

on mesopleuron and yellowish-gray spots on lateral surfaces of 

abdomen.

Head: (See Figure 172) Frons bicolored, posterior half 

brown with pale gold spots, a triangular spot lateral to posterior 

ocellus, a triangular spot in front of anterior ocellar setae, and 

a stripe between inner vertical seta and lateral margin of frons 

below ventral proclinate fronto-orbital seta; anterior half of frons 

velvety black lateral of antenna and with a ovate, shiny black, 

slightly protuberant area between bases of antennae and anterior 

ocellus. Antennal scape and pedicel velvety black; pedicel with 1 

dorsal, thicker seta and some short setae on the margin; basal flagellomere orange with slender, elongate, pale setae on margin; arista 

with 14–15 long, dorsal rays. Postcranium with 1 complete series 

of postocular setae. Face, parafacial, and gena reddish brown, pale 

orange spot between bases of antennae and silvery gray on antennal 

groove; parafacial with a pale gold spot just below the level of antenna; gena with dorsal margin pale golden, few setae on middle of 

gena; clypeus concolorous with face. Eye rounded, about as wide as 

high, height more than three times the height of gena. Gena height 

equal to length of basal flagellomere; gena-to-eye ratio 0.26–0.32.

Thorax: Mesonotum brown with small pale yellow spots 

on dorsocentral line and other lateral, irregular pale yellow spots; 

notopleuron with 3 small pale spots; scutellum brown with apex 

blackish brown. Pleurae predominantly brown, margins of katepisternum with gray microtomentum, laterotergite gray, some 

little gray spots on lateral and dorsal margins of anepisternum and 

dorsal margin of anepimeron. Wing (see Figure 200) brown with 

hyaline rounded halos; veins brown, crossvein dm–cu concave, 

curved to base; costal-vein ratio 0.37–0.57; M-vein ratio 0.80–

0.82. Coxae and trochanters with silvery microtomentum, femora 

and tibiae with brown microtomentum; tarsomeres 1 and 2 with 

yellowish-orange coloration, the remaining brown. Forefemur of 

male with short, flattened, sparse setae on distal half of anteroventral surface; foretibia straight.

Abdomen: Tergites mostly brown with triangular 

yellowish-gray spots on sides of tergites 2–5; lateroventral margins 

of tergites brown. Male terminalia: (Figures 54–57) Presurstylus 

not bifurcate, with horizontal process slightly widened basally and 

rather constricted in middle; in lateral view postsurstylus widest 

medially with a medial anterior large lobe bearing quite long and 

slender setae, posterior process rounded, poorly developed; lateral aedeagal process short, about 1⁄3 length of aedeagus, slender 

and slightly curved medially in lateral view; aedeagus somewhat 

slender, not tapered to membranous apex, quite truncate distally 

in dorsal view; phallapodeme broadly triangular in lateral view, 

prolonged distally; hypandrium widened basally, shallowly concave in lateral view.

Type Material. The holotype male of Paralimna 

fulgifrons is labeled “PERU. Loreto: Rio Momon (~25 km NW 

Iquitos)[,] 17 Feb 1984[,] W.N.Mathis/USNM ENT 00118267 

[plastic bar code label]/HOLOTYPE ♂ Paralimna fulgifrons Ale-
3 8 • smithsonian contributions to zoology

spot; postcranium with 1 series of postocular setae; mesonotum 

predominantly brown with small yellow stripes along the setal 

tracks; pleurae predominantly brown; wing pale brown tinged 

with brown spots, 2 above discal cell and 1 bigger spot on distal 

half of R1

; veins infuscate, mainly crossveins r–m, dm–cu, and 

vein CuA1

, frequently small infuscate stumps below vein R4+5 and 

above vein M; crossvein dm–cu concave, curved to base; forefemur of male with anteroventral series of short, arcuate, and 

slightly flattened setae.

12. Paralimna (Paralimna) guttata, sp. nov.

Figures 58–61, 173–174, 201–202, Map 10

Diagnosis. This species is distinguished from congeners by the following combination of characters: face brown 

with paler small area in middle or predominantly brown with 

laterodistal region paler; gena generally dark brown, yellowish 

in the margin of eyes, sometimes with a small, rounded yellow 

FIGURES 54–57. Structures of male terminalia of Paralimna (Paralimna) fulgifrons, sp. nov. (Peru. Loreto): (54) epandrium, 

cerci, and presurstyli, posterior aspect; (55) postsurstylus, lateral aspect; (56) aedeagus, phallapodeme, and hypandrium, lateral 

aspect; (57) same, ventral aspect. Scale bar = 0.1 mm. (See Figures 2–5 for labels of structures.)
number 643 • 3 9

on middle and distal margin; clypeus golden brown with distal 

margin paler. Eye rounded, about as wide as high, height three 

times the height of the gena. Gena high, height subequal to length 

of basal flagellomere; gena-to-eye ratio 0.27–0.33.

Thorax: Mesonotum predominantly brown with small 

yellow stripes along the setal tracks, postpronotal lobe yellow; scutellum brown to yellowish gray with apex blackish brown, lateral 

margins and a small, rounded dorsal spot yellowish gray. Pleurae dark brown with some small areas yellow: anepisternum and 

katepisternum with dorsal margin yellow, anepimeron yellow on 

dorsal half, laterotergite silvery gray. Wing (see Figures 201–202) 

pale brown to brown tinged with hyaline rounded spots; veins 

strongly infuscate, mainly crossveins r–m, dm–cu, and vein CuA1

, 

frequently small infuscate stumps below vein R4+5 and above vein 

M; crossvein dm–cu concave, curved to base; costal-vein ratio 

0.62–0.69; M-vein ratio 1.00–1.10. Legs brown except tarsi yellow, especially ventrally; forefemur of male with only slender setae 

ventrally, flattened setae absent; foretibia straight.

Abdomen: Tergites distinctly bicolored, golden 

brown or brown bronze with rather triangular band of silvery-gray to yellowish-gray microtomentum on laterodistal 

Description. Body length 3.2–4.5 mm; predominantly 

dark golden brown with numerous small gray or yellowish-gray 

areas dorsally and laterally on thorax and yellowish-gray areas on 

lateral surfaces of abdomen.

Head: (See Figures 173–174) Frons brown to dark brown 

with linear silvery-white or yellow spots at lateral margins of ocellar triangle in front of anterior ocellar seta and slightly convergent 

distally until near anterior margin of the frons, between inner vertical and interfrontal setae, and on fronto-orbits between proclinate fronto-orbital setae and just above antennal bases; ventral 

fronto-orbits and dorsal parafacial with 2 brown spots lateral to 

antenna, separated by yellowish-gray spot. Antennal scape and 

pedicel brown to dark brown; pedicel brown with distal margin 

paler; basal flagellomere pale brown to yellow according of incidence of light, with dorsal slender elongate hairs; arista with 

11–15 long dorsal rays. Postcranium with 1 series of postocular 

setae. Face brown with small paler area in middle and antennal 

groove yellowish, sometimes brown mainly in middle and lateral 

margins paler; parafacial silvery gray and gena wholly brown or 

with a small yellowish-gray spot in middle, a silvery-gray microtomentum around margin of the eyes; gena with only few setae 

MAP 9. Distribution map for Paralimna (Paralimna) fulgifrons, sp. nov.
4 0 • smithsonian contributions to zoology

broadly triangular in lateral view; hypandrium wide and moderately concave in posterior view.

Type Material. The holotype male of Paralimna 

guttata is labeled “BRASIL: AM[AZONAS.] C. UNIV [Campus Universitário] 7-VI-1982[,] J.A. RAFAEL/AR: MALAISE/

USNM ENT 00118271 [plastic bar code label]/HOLOTYPE ♂

Paralimna guttata Ale-Rocha & Mathis, INPA [red].” The holotype is double mounted (glued to a paper triangle), is in excellent 

condition (some Lepidoptera scales), and is deposited in INPA. 

Twenty-four paratypes (8♂, 16♀; INPA, USNM) bear the same 

locality data as the holotype. Other paratypes are as follows: 

BRAZIL. Amazonas: Comissão Executiva do Plano da Lavoura 

margin, tergite 2 with distal half yellowish gray and tergites 

3–5 brown bronze shiny along anterior 2⁄3 and medially; ventrolateral margin of tergites brown to bronze shiny. Male 

terminalia: (Figures 58–61) Presurstylus not bifurcate, with 

horizontal process slender and shallowly curved; in lateral 

view postsurstylus widest medially with a medial, anteriorly 

wide lobe bearing moderately long, slender, sparse setae, 

medial posterior process developed, triangular, or poorly developed; lateral aedeagal process shortened, about 1⁄2 length 

of aedeagus, slightly curved medially in lateral view, bluntly 

rounded; aedeagus gradually tapered to membranous apex, 

moderately robust or slender in posterior view; phallapodeme 

FIGURES 58–61. Structures of male terminalia of Paralimna (Paralimna) guttata, sp. nov. (Brazil. Amazonas): (58) epandrium, cerci, and presurstyli, posterior aspect; (59) postsurstylus, lateral aspect; (60) aedeagus, phallapodeme, and hypandrium, lateral aspect; (61) same, ventral aspect. Scale bar = 0.1 mm. (See Figures 2–5 for labels of structures.)
number 643 • 4 1

GUYANA. Paramakatoi (04°42′N, 59°42.8′W), 24–25 Aug 

1997, W. N. Mathis (11♂, 2♀; INPA, USNM).

Other Specimens Examined. BRAZIL. Amazonas: Reserva Ducke (26 km N of Manaus; 02°55.8′S, 

59°58.5′W), 31 Aug 1982 (1♀; INPA); Rio Nhamundá 

(01°35.2′S, 57°37.5′W; 25 m; Malaise trap), 17–20 May 2008, 

J. A. Rafael and team (2♂, 2♀; INPA).

GUYANA. Mazaruni (06°25.2′N, 58°37.7′W; high forest), 

13 Aug 1937, Richards and Smarts, (3♀; BMNH). New River 

(03°21.2′N, 57°35′W; 228 m), 7–8 May 1938, C. A. Hudson 

(1♀; BMNH). Essequibo River (05°22.2′N, 58°43.4′W; source), 

J. Ogilvie (1♀; BMNH).

VENEZUELA. Bolívar: Gran Sabana, Akuriman (08°16′N, 

62°45.3′W), Nov–Dec 1940, P. Anduze (2♂, 3♀; USNM). Merida: Tovar (08°19.7′N, 71°45.6′W), 7 Aug 1943, P. Anduze 

(1♂; USNM).

Type Locality. Neotropical. Brazil. Amazonas: 

Manaus, Universidade Federal do Amazonas (Campus Universitário; 03°06′S, 59°58.5′W).

Distribution. (Map 10) Neotropical: Brazil (Amazonas, Pará, Roraima), Guyana, Venezuela (Bolívar, Merida).

Cacaueira (km 31, Est. AM 010 BR; 02°52.7′S, 59°57.1′W), 3 

May 1976, M. Dantas (1♀; INPA); Reserva Ducke (26 km NE 

Manaus; 02°55.8′S, 59°58.5′W), 24 Aug–16 Sep 1982, 1986, 

U. Barbosa, L. Aquino (3♂, 15♀; INPA); Reserva Ducke (26 

km NE Manaus; 02°55.8′S, 59°58.5′W; 40 m), 5 May 2010, 

D. and W. N. Mathis (9♂; USNM); Reserva Biológica da 

Campina (02°30′S, 60°00′W; km 60 da Rodovia BR-174, 

Manaus/Boa Vista), 17–20 Feb 1976, L. P. Albuquerque (1♀; 

INPA); Borba, Rio Abacaxis Paxiúba (04°28.8′S, 58°34.4′W; 

Malaise trap), 2–4 Jun 2008, J. A. Rafael and team (2♂, 3♀; 

INPA, USNM); Borba, Rio Abacaxis Paxiúba (05°15.1′S, 

58°41.9′W; Malaise trap), 26–28 May 2008, J. A. Rafael and 

team (1♀; INPA); Rio Abacaxis, Flona Pau Rosa (05°15.1′S, 

58°41.9′W), 27–29 May 2008, J. A. Rafael and team (1♂, 2♀; 

INPA).Pará: Óbidos, Fazenda Pajurá (01°37.4′S, 55°23.2′W; 

Malaise trap), 5–11 Sep 2001, J. A. Rafael, J. F. Vidal (1♂; 

INPA); Rio Nhamundá (01°35.2′S, 57°37.5′W; 25 m; Malaise trap), 17–20 May 2008, J. A. Rafael and team (6♂, 9♀; 

INPA). Roraima: Rio Uraricoera, Ilha de Maracá (03°21.7′S, 

61°26′W), 18–28 Aug 1987, J. A. Rafael, L. S. Aquino, J. F. 

Vidal, E. Binda (1♀; INPA).

MAP 10. Distribution map for Paralimna (Paralimna) guttata, sp. nov., and P. (Paralimna) piger Cresson.
4 2 • smithsonian contributions to zoology

brown and a small, rounded brown spot in middle. Wing hyaline; 

veins brown; costal-vein ratio 0.44–0.48; M-vein ratio 0.96–1.04. 

Legs brown with sparse silvery-gray microtomentum; foretarsus 

brown, mid and hind tarsi lighter with some yellow coloration, 

especially ventrally; forefemur of male with anteroventral series of 

moderately long setae, shorter and slightly flattened on distal half 

of the femur, ventral surface without outstanding setae, posteroventral setae very slender and short, inconspicuous; foretibia of 

male slightly arcuate.

Abdomen: Tergites distinctly bicolored except tergite 

1 wholly dark brown, tergites 2–5 dark brown along anterior 1⁄2–2⁄3

and medially, otherwise yellowish gray; lateroventral margins of 

tergites brown. Male terminalia: (Figures 62–65) Presurstylus not 

bifurcate, with horizontal process slender and shallowly curved; 

in lateral view postsurstylus widest medially with a medial, anteriorly wide lobe bearing moderately long strong setae, medial 

posterior process very prominent; lateral aedeagal process about 

1⁄2 length of aedeagus, slightly curved medially in lateral view; 

aedeagus gradually tapered before apex and widened in the membranous apex; phallapodeme broadly triangular in lateral view, 

prolonged distally; hypandrium wide and shallowly concave in 

posterior view.

Type Material. The holotype male of Paralimna 

maculata is labeled “DOMINICAN R[E]P[UBLIC]. Monseñ[or]. 

Nouel: nr. Jima, 670m, 19°01.6′N, 70°28.9′W[,] 29Mar1999, 

W. Mathis/USNM ENT 00090309 [plastic bar code label]/HOLOTYPE ♂ Paralimna maculata Ale-Rocha &Mathis, USNM 

[red].” The holotype is double mounted (minuten pin in a white 

block of plastic), is in excellent condition, and is deposited in the 

USNM. Eighteen paratypes (13♂, 5♀; INPA, USNM) bear the 

same label data as the holotype. Other paratypes are as follows: 

CUBA. Havana: Baracoa (23°03.7′N, 82°21.2′W), Sep 1901 

(1♂; USNM).

DOMINICAN REPUBLIC. Barahona: Baoruco (8.3 km 

S; 18°01.9′N, 71°08.4′W), 15 May 1995, W. N. Mathis (9♂; 

USNM); Baoruco (beach and river; 18°04.6′N, 71°05.5′W), 19 

May 1998, D. and W. N. Mathis (8♂; USNM); Baoruco (Sierra de Neiba, Los Guineos; Rio Colorado; 18°35′N, 71°11′W), 

11–12 Aug 1990, J. Rawlins, S. Thompson (8♂, 8♀; CMP); 

Cortico (La Mina; 18°06.7′N, 71°13.4′W; 1300 m), 23 Mar 

1999, W. N. Mathis (1♂; USNM); Paraíso (5 km N; 18°01.5′N, 

71°11.6′W; 150 m), 21 Mar 1999, W. N. Mathis (1♂; USNM); 

Paraíso (6 km NW; Rio Nizao; 18°02′N, 71°12′W; 170 m), 

25–26 Jul 1990, J. Rawlins, S. Thompson, C. Young (16♂, 

34♀; CMP); Paraíso (9.2 km NW; 18°03′N, 71°12′W; Rio 

Nizao and Rio Coltico; 230 m), 9–10 Aug 1990, J. Rawlins, 

S. Thompson (1♀; CMP); San Rafael (18°01.9′N, 71°08.4′W), 

22 Mar 1999, W. N. Mathis (1♂; USNM). El Seibo: El Seibo 

(5 km E; 18°44.73′N, 68°59.2′W; 120 m), 12 May 1995, 

W. N. Mathis (5♂; USNM); Pedro Sáchez (18°51.4′N, 69°6.5′W), 

26 May 1998, D. and W. N. Mathis (3♂, 1♀; USNM). Hato 

Mayor: El Valle (9.5 km W; 18°59′N, 69°29′W; 170 m), 6 Jul 

1992, R. Davidson, J. Rawlins, S. Thompson, C. Young (1♀; 

CMP). Independencia: Angel Feliz (5 km NNW; Sierra de Neiba, 

Etymology. The species epithet, guttata, is of Latin 

derivation and means drop or spot, alluding to the hyaline halos 

of the wing.

Remarks. This species forms a monophyletic group 

with P. fulgifrons sp. nov. and P. stellata sp. nov. In the “Remarks” for P. fulgifrons sp. nov., distinguishing characters are 

noted. Additionally, P. guttata sp. nov. can be distinguished 

from the other two species by having a brown to pale brown face 

with gray areas on the dorsal half, gray antennal grooves laterad 

of the facial carina, a brown pedicel with yellowish ventral and 

distal margins, and a frons that lacks a large black velvety spot 

on the ventral portion of the fronto-orbits.

13. Paralimna (Paralimna) maculata, sp. nov.

Figures 62–65, 175, Map 11

Diagnosis. This species is distinguished from congeners by the following combination of characters: face and gena 

dark bluish gray, face with 3 brown spots, a single large rectangular spot on carina, as long as the length of antenna, and 2 also 

rectangular transverse spots between larger facial setae, lateral 

margin brown, gena with a small golden-brown spot on anterior margin; anepisternum with ventral and dorsal margins and 

central area mostly brown, remainder mostly gray; forefemur of 

male with anteroventral series of moderately long setae, shorter 

and slightly flattened on distal half of the femur.

Description. Body length 2.6–3.2 mm; body generally 

bicolored, dark brown dorsally, lateral surfaces dark bluish gray.

Head: (See Figure 175) Frons almost to entirely uniformly chestnut brown with linear yellow spots at lateral margins 

of ocellar triangle and in front of anterior ocellar seta, short, not 

reaching the anterior margin of frons, between inner vertical and 

interfrontal seta, and on fronto-orbits between proclinate frontoorbital setae and just above antennal bases; ventral fronto-orbits 

and dorsal parafacial with a small dark brown spot lateral to antenna. Antenna brown, covered by yellow microtomentum; basal 

flagellomere with slender, elongate, pale setae on margin; arista 

with 9–11 long, dorsal rays. Postcranium with 2 series of postocular setae, the second series irregular. Face dark bluish gray to 

gray with 3 brown spots, 1 large rectangular spot on carina, as 

long as the length of antenna, and 2 also rectangular transverse 

spots between larger facial setae, and lateral margin brown, toward parafacial; parafacial and gena dark gray, gena with a small 

golden-brown spot on anterior margin; gena with only few setae 

on middle and ventral margin; clypeus brown. Eye rounded, about 

as wide as high, height about three times the height of the gena. 

Gena high, height subequal to length of basal flagellomere; genato-eye ratio 0.30–0.33.

Thorax: Mesonotum predominantly dark brown with 

small yellow spots along setal tracks and laterally; postpronotum silvery gray; scutellum brown dorsally, paler on distal 1⁄3, 

with apex blackish brown and yellowish gray otherwise. Pleurae 

gray to dark gray, anepisternum with dorsal and ventral margins 
number 643 • 4 3

FIGURES 62–65. Structures of male terminalia of Paralimna (Paralimna) maculata, sp. nov. (Dominican Republic. Barahona): (62) epandrium, cerci, and presurstyli, posterior aspect; (63) postsurstylus, lateral aspect; 

(64) aedeagus, phallapodeme, and hypandrium, lateral aspect; (65) same, ventral aspect. Scale bar = 0.1 mm. 

(See Figures 2–5 for labels of structures.)

71°17′W; 1500 m), 27 Dec 1955, J. Maldonado-Caprilles (8♂, 

5♀; USNM); Constanza (9 km SW; 18°52′N, 70°43′W; road to 

Pinar Bonito; 1433 m), 29 Jul 1991, D. A. Grimaldi, J. Stark 

(130♂, 153♀; AMNH); Constanza (11.5 km S; 18°51.7′N, 

70°41′W; 1410 m), 27 Mar 1999, W. N. Mathis (1♀; USNM); 

El Rio (9.5 km E; 19°0.9′N, 70°33.5′W; 980 m), 6 May 1995, 

W. N. Mathis (1♂; USNM); Jarabacoa (1–2 km S; 19°06.9′N, 

70°37′W; 520 m), 8–21 May 1995, 1998, D. and W. N. Mathis 

(43♂, 6♀; INPA, USNM); Jarabacoa (5 km S; 19°05.8′N, 

just S crest; 18°41′N, 71°47′W; 1780 m), 13–15 Oct 1991, R. 

Davidson, J. Rawlins, S. Thompson, C. Young (1♂, 2♀; CMP); 

Angel Feliz (5.5 km NNW; Sierra de Neiba, near crest; 18°41′N, 

71°47′W; 1750 m), 21–22 Jul 1992, R. Davidson, J. Rawlins, 

S. Thompson, C. Young (1♂; CMP); La Descubierta (32 km 

NW; Sierra de Neiba; 18°38.6′N, 71°46.4′W; 1850 m), 1–5 Dec 

1991, S. Peck (1♀; CMP); Los Bolos (18°37.8′N, 71°39.2′W; 

1370 m), 24 Mar 1999, W. N. Mathis (4♂; USNM). La Vega:

Paso de la Vaca, Monseñor-Nouel-Constanza Road (19°04′N, 
4 4 • smithsonian contributions to zoology

MAP 11. Distribution map for Paralimna (Paralimna) maculata, sp. nov.

70°36.5′W; 640 m), 8–20 May 1995, W. N. Mathis (24♂, 4♀; 

USNM); La Cienega de Manabao (19°03.9′N, 70°51.8′W; 1050 

m), 28 Mar 1999, W. N. Mathis (20♂; USNM); Rio Camu (3.5 

km NW La Vega; 19°13.8′N, 70°35.2′W; 100 m), 18 May 1998, 

D. and W. N. Mathis (2♂; USNM); Salto Baiguate (near Jarabacoa; 19°05.5′N, 70°36.9′W; 570 m), 9 May 1995, W. N. Mathis 

(13♂, 3♀; USNM); Salto Baiguate (near Jarabacoa; 19°05.7′N, 

70°37′W, 570m), 16 May 1998, D. and W. N. Mathis (8♂, 1♀; 

USNM); Salto Guasara (near Jarabacoa; 19°04.4′N, 70°42.1′W; 

680 m), 9 May 1995, W. N. Mathis (11♂, 1♀; USNM); Salto 

de Jimenoa (19°06′N, 70°35.9′W; 575 m), 20 May 1995, W. N. 

Mathis (1♂, 1♀; USNM). Monseñor Nouel: Bonao (8.7 km 

W; 19°01.8′N, 70°29.4′W), 10 May 1995, W. N. Mathis (1♂; 

USNM); dam near Rodeo (18°53.1′N, 70°33.5′W), 22 May 

1998, D. and W. N. Mathis (1♂, 1♀; USNM); Rodeo (1 km NE; 

18°52.9′N, 70°32.7′W), 22 May 1998, D. and W. N. Mathis 

(3♂; USNM). Pedernales: Alcoa Road (km 30; N Cabo Rojo; 

18°07.3′N, 71°35.8′W; 1080 m), 20 Mar 1999, W. N. Mathis 

(1♀; USNM); Rio Mulito (13 km N Pedernales; 18°09′N, 

71°46′W; 230 m), 17 Jul 1992, R. Davidson, J. Rawlins, S. 

Thompson, C. Young (2♂, 8♀; CMP). Peravia: Rio Ocoa (San 

José Ocoa; 18°31.7′N, 70°30.4′W), 21 May 1998, D. and W. N. 

Mathis (1♂; USNM); San José de Ocoa (18 km N; 830 m), 24 

Jul, 1986, D. A. Grimaldi (18♂, 19♀; AMNH); San José Ocoa 

(10 km NE; 18°35′N, 70°25.6′W), 21 May 1998, D. and W. N. 

Mathis (18♂, 1♀; USNM). Puerto Plata: Sonador (19°35.9′N, 

70°36.2′W; 440 m), 18 May 1995, W. N. Mathis (3♂; USNM).

Type Locality. Neotropical. Dominican Republic. 

Monseñor Nouel: near Jima (670m, 19°01.6′N, 70°28.8′W).

Distribution. (Map 11) Neotropical: West Indies 

(Cuba, Dominican Republic).

Etymology. The species epithet, maculata, is of 

Latin derivation and means spot or stain, alluding to the brown 

spot on the gena.

Remarks. This species is similar to P. sera in having 

a darkened body that is somewhat bluish with gray microtomentum. It can be distinguished from P. sera, however, by the 

pattern of facial spots, including a brown genal spot, and by 

the simple male forefemur. The male terminalia offers definitive 

characters to distinguish these species. The horizontal process of 

the presurstylus in P. maculata sp. nov. is slender and has short 

setae, the postsurstylus is widened medially with an anterior and 
number 643 • 4 5

laterally from level of first dorsocentral seta to last dorsocentral 

seta, plus some small spots laterally; postpronotum yellowish 

gray; scutellum brown dorsally, yellow spotted, with apex blackish brown and yellowish gray laterally. Pleurae densely gray or 

silvery gray, with dorsal margin of anepisternum and insertion 

of numerous setae brown tinged. Wing (see Figure 203) subhyaline; veins pale brown; costal-vein ratio 0.36–0.41; M-vein ratio 

1.07–1.15. Legs blackish brown with dense silvery-gray microtomentum; tarsi lighter with some yellow coloration, especially 

ventrally, the apical segments darkened; forefemur of male concave ventrally on distal half, with a posteroventral row of short 

setae at basal half and apex of femur, absent on concave region, 

slightly longer at base of femur; anteroventral row of curved and 

moderately short and slightly flattened setae; foretibia of male 

straight.

Abdomen: Tergites distinctly bicolored, tergite 1 dark 

brown with lateral corner yellowish gray; tergite 2–5 dark brown 

along anterior 2⁄3 and medially, otherwise yellowish gray. Male terminalia: (Figures 66–69) Presurstylus bifurcate, with basoventral 

process large, as long as 1⁄2 length of horizontal process, horizontal 

process robust and straight; in lateral view postsurstylus widest 

medially with a medial, anterior, moderately wide lobe bearing 

short robust setae, medial posterior process developed, clavate 

distally; lateral aedeagal process less than 1⁄2 length of aedeagus, 

slightly curved medially in lateral view, robust; aedeagus gradually 

tapered to membranous apex; phallapodeme broadly triangular in 

lateral view, prolonged distally; hypandrium wide and shallowly 

concave in posterior view.

Type Material. The holotype male of Paralimna 

malleata is labeled “DOMINICA, W. I[,] ClarkeHallEst[ate]. 

May 20 1966[“20” handwritten] G. Steyskal/Bredin-ArchboldSmithsonian Bio.Surv.Dominica/USNM ENT 000118263 

[plastic bar code label]/HOLOTYPE ♂ Paralimna malleata

Ale-Rocha &Mathis, USNM [red].” The holotype is double 

mounted (minuten pin in a white block of plastic), is in excellent 

condition, and is deposited in the USNM. Thirty-nine paratypes 

(28♂, 11♀; INPA, USNM) bear the same locality data as the 

holotype but with different dates and collectors. DOMINICA.

Clarke Hall (15°24.5′N, 61°23.7′W; Malaise trap), 11 Jan–23 

Aug 1964, 1965, 1966, light trap, D. M. Anderson, D. Bray, 

R. J. Gagné, H. Robinson, T. J. Spilman, G. C. Steyskal, W. W. 

Wirth. Other paratypes are as follows: DOMINICA. Hampstead 

River (15°33.9′N, 61°22.3′W), 22 Mar 1989, W. N. Mathis 

(10♂; USNM). Fond Figues River (15°24.5′N, 61°18.1′W; 122 

m), 9–12 Mar 1965, W. W. Wirth (1♂; USNM). Hodges River 

(15°35.5′N, 61°20′W; mouth; swamp forest), 27 Feb 1965, 

W. W. Wirth (1♀; USNM).

Other Specimens Examined. ECUADOR. El 

Oro: Rio Grande (15 km S Zaruma; 03°42.8′S, 79°36.8′W; 

850 m), 9 Apr 1965, L. E. Peña (1♂, 1♀; CNC). Loja: San 

Pedro, Zaruma Road (03°45.8′S, 79°38.1′W; 1100 m), 9 Apr 

1965, L. E. Peña (1♀; CNC). Orellana: Coca, Napo River 

(0°27.8′S, 76°59′W; 250 m), 12–130 Apr 1965, L. E. Peña (1♀; 

CNC).

posterior medial process that are well developed, and the lateral 

aedeagal process is typical. In P. sera, the horizontal process of 

the presurstylus is widened basally, bearing long ventral setae; 

the postsurstylus is not widened medially with a weakly developed anteromedial process; the posteromedial process is undeveloped; and the lateral aedeagal process is peculiarly large and 

has striations on the lateral surface.

14. Paralimna (Paralimna) malleata, sp. nov.

Figures 66–69, 176, 203, Map 12

Diagnosis. This species is distinguished from congeners by the following combination of characters: face mostly 

dark or silvery gray but with lateral margin brown and 3 distinct brown spots, 1 on carina and 2 parallel triangular spots 

between larger facial setae, the spot on carina black for a short 

distance between bases of antennae and brown in the remaining; 

anepisternum with dorsal margin brown, central area with bases 

of numerous setulae brown, otherwise mostly gray; forefemur 

of male concave ventrally on distal half, with a posteroventral 

row of short setae at basal half and apex of femur, absent on 

concave region, slightly longer at base of femur; anteroventral 

row of curved and moderately short and slightly flattened setae; 

abdomen with brown bands more developed, extended dorsally, 

tergite 2 with basal brown band extended until posterior margin 

in middle of each side; with basoventral process large, as long as 

1⁄2 length of horizontal process.

Description. Body length 3.2–4.2 mm; body generally bicolored, dark brown dorsally, lateral surfaces gray, sometimes silvery gray.

Head: (See Figure 176) Frons chestnut, reddish brown, 

and pale brown mottled with linear yellow spots at lateral margins of ocellar triangle and in front of anterior ocellar seta, short, 

not reaching the anterior margin of frons, between inner vertical and interfrontal setae, and on fronto-orbits between proclinate fronto-orbital setae and just above antennal bases; ventral 

fronto-orbits and dorsal parafacial with 2 small dark brown to 

pale brown spots lateral to antenna, separated by yellow spot. 

Antenna brown, covered by dense paler microtomentum; basal 

flagellomere with slender elongate pale setae on margin; arista 

with 11–13 long, dorsal rays. Postcranium with 2 complete series of postocular setae. Face dark gray or silvery gray but with 

lateral margin brown and 3 distinct brown spots, 1 on carina, 

black for a short distance between bases of antennae and as long 

as length of antenna, and 2 large parallel triangular spots between larger facial setae; parafacial and gena silvery gray; gena 

with only few setae on middle and ventral margin; clypeus silvery gray. Eye rounded, about as wide as high, height about 

three times the height of gena. Gena high, height subequal or 

slightly greater than length of basal flagellomere; gena-to-eye 

ratio 0.33–0.42.

Thorax: Mesonotum yellowish gray with dark brown 

spots, 3 slender stripes between dorsocentral rows, 1 wide stripe 
4 6 • smithsonian contributions to zoology

Remarks. Externally this species is generally rather 

common but is distinguished from congeners as follows: the 

male forefemur is concave ventrally, the basal brown tergal 

bands are well developed, almost reaching the posterior margins, the basal brown band of tergite two is extended to the 

posterior margin on each side, and the basoventral process is 

notably large.

Type Locality. Neotropical: Dominica. Layou Valley, Clarke Hall Estate (15°24.5′N, 61°23.7′W).

Distribution. (Map 12) Neotropical: Ecuador (El 

Oro, Loja, Orellana), West Indies (Dominica).

Etymology. The species epithet, malleata, is of 

Latin derivation and means hammer, alluding to the hammerlike shape of the presurstylus.

FIGURES 66–69. Structures of male terminalia of Paralimna (Paralimna) malleata, sp. nov. (Dominica. Clarke Hall): (66) 

epandrium, cerci, and presurstyli, posterior aspect; (67) postsurstylus, lateral aspect; (68) aedeagus, phallapodeme, and hypandrium, lateral aspect; (69) same, ventral aspect. Scale bar = 0.1 mm. (See Figures 2–5 for labels of structures.)
number 643 • 4 7

MAP 12. Distribution map for Paralimna (Paralimna) malleata, sp. nov., and P. (Paralimna) plumbiceps Cresson.

15. Paralimna (Paralimna) meridionalis Cresson

Figures 70–73, 177–178, Map 13

Paralimna meridionalis Cresson 1916:119 [Costa Rica. Cartago: La Carpintera (9°53.9′N, 83°58.8′W); HT ♂, ANSP (6092)]; 1918:47 [review; 

Costa Rica].

Paralimna (Paralimna) meridionalis.—Cresson 1947:49 [review, Neotropical species].—Wirth 1968:15 [Neotropical catalog].—Mathis and Zatwarnicki 1995:122 [world catalog].

Diagnosis. This species is distinguished from congeners by the following characters: face with coloration variable, without distinct spots, dark gray with lateral margins dark 

brown, wholly brown or slightly grayish brown with central 

portion dark brown and becoming yellowish or silvery gray 

toward distal and lateral margins, sometimes face gray with 

3 distinct golden-brown spots, 1 on carina and 2 small spots 

between larger facial setae; mesonotum mostly brown with few 

small yellowish-gray areas or slender stripes along setal tracks 

and laterally; forefemur of male with anteroventral row of short, 

acutely pointed, curved, slightly flattened setae on distal half of 

femur; posteroventral surface with 2 series of numerous and conspicuous setae.

Description. Body length 3.2–4.5 mm; body generally bicolored, dark brown dorsally, lateral surfaces pale or dark 

gray.

Head: (See Figures 177–178) Frons brown, reddish 

brown tinged on anterior half, with linear yellow spots at lateral 

margins of ocellar triangle, in front of anterior ocellar seta, not 

reaching the anterior margin of frons, between inner vertical and 

interfrontal setae, and on fronto-orbits between proclinate frontoorbital setae and just above antennal bases; ventral fronto-orbits 

and dorsal parafacial with gray spot separated by yellowish-gray 

area. Antennal scape and pedicel dark brown, pedicel with distal 

margin paler; basal flagellomere pale brown to yellowish brown 

according to incidence of light, with slender, elongate, pale setae 

on margin; arista with 10–17 long, dorsal rays. Postcranium with 

2 complete series of postocular setae. Face with coloration variable: without distinct spots, dark gray but dark brown through 
4 8 • smithsonian contributions to zoology

handwritten].” The holotype is double mounted (minuten pin in 

a vertical rectangular card), is in good condition (thorax partly 

torn by the pin; abdomen not dissected), and is deposited in the 

ANSP (6092).

Other Specimens Examined. COLOMBIA.

Santander: Aljibes, Providencia (about 33 km SW Zaragosa; 

06°46.1′N, 73°32.7′W), 13 Oct 1970, R. W. Pinger (8♂, 6♀; 

USNM). Valle del Cauca: Buenaventura 70 km E, Anchicayá 

(03°36.9′N, 76°54.8′W; 400 m), 17–20 Feb 1970, D. M. Wood 

(1♀; CNC).

COSTA RICA. Alajuela: Peñas Blancas (10°04′N, 

84°37′W; 700 m; rain forest), 18 Aug 1986, L. Masner (3♂, 

1♀; CNC). Cartago: Cartago (9°51.4′N, 83°55.2′W), 4 Jul 

1909, P. P. Calvert (2♂, 1♀; ANSP); Cartago (near; 09°51.3′N, 

83°55.2′W), 15 Dec 1909, P. P. Calvert (2♂, 2♀; ANSP); La 

Carpintera (09°53.9′N, 83°58.8′W), 4 Dec 1909, P. P. Calvert 

(3♂, 1♀; paratypes; allotype ♀; ANSP); Pejibaye (09°49′N, 

83°42′W), 22–24 Mar 1987, W. E. Steiner (1♀; USNM); 24 Mar 

1987, at black light in cut-over forest near river (2♂; USNM); 

Turrialba (09°54.1′N, 83°41.1′W), Nov 1922, P. Schild (1♀; 

USNM). Guanacaste: La Palma (10°15.2′N, 85°03.3′W), Aug 

1923 (2♂; ANSP). Heredia: Santo Domingo, Instituto Nacional 

de Biodiversidad (INBio) (09°58.4′N, 84°5.6′W), 18 Feb 2003, 

W. N. Mathis (1♂; USNM); La Selva Biological Station 

(10°27.2′N, 84°0.2′W; trap in garden), Jan 1993, P. Hanson (14♂, 15♀; USNM). Limón: Puerto Vargas (09°43.9′N, 

82°48.9′W; beach), 27–28 Jun 2001, W. N. Mathis (4♂; 

USNM). Paraíso: Navarro (09°48.6′N, 83°52.7′W; light trap), 

Jul 1962, F. S. Blanton (1♀; USNM). Puntarenas: Monte Lirio 

(08°46.8′N, 82°49.8′W), 27 Aug 1923, R. C. Shannon (4♂, 1♀; 

ANSP); Piedras Blancas (24 km W; 08°47′N, 83°15′W; 200 m; 

Malaise trap), Dec 1990, P. Hanson (1♂; USNM); San Vito, Las 

Cruces (08°49.3′N, 82°58.3′W; 1200 m), 9 Jul 1983, B. Gill 

(1♂; CNC); San Vito, Las Alturas (08°57′N, 82′50′W, 1500 m; 

Malaise trap), Dec 1991, P. Hanson (4♂, 3♀; USNM). San José: 

Ciudad Colon (09°55′N, 84°15′W; 800 m; Malaise trap), Dec 

1989, P. Hanson (4♀; USNM); El Rodeo (09°54.6′N, 84°12.2′W, 

1860 m), 26 Jun 2001, W. N. Mathis (5♂, 1♀; USNM); Río 

Savegre (09°33′N, 83°48.5′W; 2180 m), 7–8 Aug 2001, D. and 

W. N. Mathis (1♂, 1♀; USNM); Zurquí (10°2.6N, 84°0.4W), 

19 Feb 2003, W. N Mathis (4♂; 3♀; USNM); Zurquí de Moravia (10°05′N, 84°01′W, 160 m; Malaise trap), Mar 1991, 

P. Hanson (3♂, 8♀; USNM).

ECUADOR. El Oro: El Guabo (0°38.2′S, 76°08.9′W), Dec 

1955, R. Levi-Castillo (12♂; USNM); Pasaje (6 km E; 03°18.1′S, 

79°47.1′W), 13 Jan 1978, W. N. Mathis (1♀; USNM). Cotopaxi: Latacunga (N; 00°52.2′S, 78°37.1′W; 3300 m), Dec 

1970, L. E. Peña (2♂, 2♀; INPA, USNM). Guayas: Machala 

(49.5 km NNE; 02°40′S, 79°13.4′W; 40 m), 13 Jan 1978, W. N. 

Mathis (1♂; USNM). Loja: La Toma, W of Loja (03°59′S, 

79°21′W; 1500 m), Nov 1970, L. E. Peña (1♀; MZUSP); Loja 

(03°59.4′S, 79°12.3′W; 1200 m), 26 Mar–23 Nov 1965, 1970, 

L. E. Peña (3♂; CNC, MZUSP). Napo: Limoncocha (00°24.7′S, 

76°37.5′W), 9 Mar–15 Jun 1976, 1977, P. J. Spangler, 

lateral facial setae, wholly brown or slightly grayish brown with 

central portion dark brown and becoming yellowish or silvery 

gray toward distal and lateral margins, both hardly contrasting 

with silvery-gray antennal groove; or face dark to pale gray with 

3 distinct spots, 1 pale golden-brown spot on carina, the same 

length of antenna, and 2 small ovate horizontal or triangular 

golden-brown spots between larger facial setae; parafacial and 

gena silvery gray or yellowish gray; gena with only few setae on 

middle and ventral margin; clypeus silvery or yellowish gray. Eye 

rounded, slightly higher than wide, height two or three times the 

height of gena. Gena high, height slightly longer than length of 

basal flagellomere; gena-to-eye ratio 0.30–0.62.

Thorax: Mesonotum yellowish or silvery gray with 

brown spots, 3 stripes between dorsocentral rows, complete or 

incomplete, not reaching the posterior margin of scutum, plus 

some small spots laterally, base of setae brown; postpronotum, 

notopleuron, and supra-alar areas mostly yellowish or silvery 

gray; scutellum brown dorsally, with apex blackish brown and 

silvery or yellowish gray laterally. Pleurae mostly silvery or yellowish gray, with dark gray or yellowish gray microtomentum; 

anepisternum variable, gray with stains of bases of setae brown, 

sometimes coalescing to form a large spot at middle area, with 

dorsal and ventral margin widely or narrowly brown, sometimes 

with a rectangular brown spot on dorsal margin or with dorsal 

1⁄3 and posteroventral margin with a rounded spot brown. Wing 

usually slightly brown tinged; veins brown, sometimes with crossveins r–m and dm–cu with infuscate halo and a short stump on R1

; 

costal-vein ratio 0.35–0.45; M-vein ratio 0.95–1.15. Legs blackish 

brown with silvery-gray microtomentum and a dorsal green reflection; tarsi lighter with some yellow coloration, especially ventrally; 

forefemur of male with anteroventral series of short setae, slightly 

flattened on distal half of femur; posteroventral surface with 

2 series of numerous and conspicuous setae; foretibia straight.

Abdomen: Tergites distinctly bicolored, mostly brown, 

brown bands more developed, extended dorsally, tergite 2 with 

basal brown band extended until posterior margin in middle 

of each side, otherwise silvery or dark gray. Male terminalia: 

(Figures 70–73) Presurstylus bearing short, slender setae, bifurcate, with basoventral process short, somewhat tapered ventrally, length almost 1⁄3 of horizontal process length; horizontal 

process fairly arcuate, sometimes rather straight; in lateral view 

postsurstylus widest medially with medial weakly produced 

lobe bearing small, slender, sparse setae, and a medial small 

pointed process posteriorly; lateral aedeagal process about 1⁄2 

or less than 1⁄2 length of aedeagus, thin, weakly curved in middle, rounded apically; aedeagus long, slightly constricted in the 

apical 1⁄3 and gently widened to membranous apex, sometimes 

moderately tapered to membranous apex; in lateral view phallapodeme triangular, prolonged distally; hypandrium deeply 

concave.

Type Material. The holotype male of Paralimna 

meridionalis is labeled “La Carpintera[,] Cartago[,] C[osta]

R[ica][,] 4 XII 1909/♂/TYPE Paralimna MERIDIONALIS 

E.T.Cresson,Jr. [maroon; “Paralimna MERIDIONALIS” 
number 643 • 4 9

Spangler (7♂; INPA, USNM). Zamora-Chinchipe: W Zamora 

(04°03.8′S, 78°57.9′W; 1200 m), Nov 1970, L. E. Peña (3♂; 

MZUSP).

GUYANA. Mazaruni-Potaro: Takutu Mountains (06°15′N, 

59°5′W; Malaise trap near stream in montane rainforest, Earth 

Research Expedition), 3–19 Dec 1983, P. J. Spangler, W. E 

Steiner (7♂, 4♀; USNM); Kartabo Point (06°23′N, 58°41′W), 

22–26 Dec 1983, W. E. Steiner (1♂, 1♀; USNM).

D. R. Givens, G. E. Shewell (2♂, 2♀; CNC, USNM). Orellana:

Coca, Napo River (0°27.8′S, 76°59′W; 250 m), 12–13 Apr 1965, 

L. E. Peña (1♀; CNC); Rio Tiputini (0°38.2′S, 76°8.9′W), 12–26 

Aug 1999, W. N. Mathis, A. Baptista, M. Kotrba (5♂; USNM). 

Pastaza: Río Puyo (01°29.1′S, 77°59.7′W), R. Levi-Castillo (4♂; 

USNM). Pichincha: Perucho (N; 00°10.2′N, 78°24.4′W; 2000 

m), Jan 1971, L. E. Peña (1♂; MZUSP). Tungurahua: Banos 

(11 km SE; 01°23.8′S, 78°19′W; 1660 m), 25 Jan 1976, P. J. 

FIGURES 70–73. Structures of male terminalia of Paralimna (Paralimna) meridionalis Cresson (Costa Rica. Cartago): (70) 

epandrium, cerci, and presurstyli, posterior aspect; (71) postsurstylus, lateral aspect; (72) aedeagus, phallapodeme, and hypandrium, lateral aspect; (73) same, ventral aspect. Scale bar = 0.1 mm. (See Figures 2–5 for labels of structures.)
5 0 • smithsonian contributions to zoology

(4♂; USNM); Tingo Maria (6 km S; 09°22.7′S, 75°58.4′W), 8 

Feb 1984, W. N. Mathis (1♀; USNM); Las Palmas, Río Huallaga (09°26.5′S, 75°58.3′W), 8 Feb 1984, W. N. Mathis (1♂; 

USNM).

VENEZUELA. Lara: Yacambú (10°0.7′N, 69°14.6′W; 

1200 m), 10 May 1981, H. K. Townes (16♂, 12♀; CNC). 

Zulia: El Tucuco (45 km SW of Maquiches; 09°50.7′N, 

72°48.7′W), 5–6 Jun 1976, A. S. Menke, D. Vincent (4♂, 

20♀; USNM).

Type Locality. Neotropical. Costa Rica. Cartago: La 

Carpintera (9°53.9′N, 83°58.8′W).

Distribution. (Map 13) Neotropical: Costa Rica 

(Alajuela, Cartago, Guanacaste, Heredia, Paraíso, Puntarenas, 

San José), Colombia (Santander), Ecuador (Cotopaxi, El Oro, 

Guayas, Loja, Napo, Orellana, Pastaza, Pichincha, Tungurahua, Zamora-Chinchipe), Guyana (Mazaruni-Potaro), Panamá 

(Chiriqui, Cocle, Guna Yala, Panamá, Veraguas), Peru (Huánuco), Venezuela (Lara, Zulia).

Remarks. Paralimna brunneiceps, P. fellerae, P. meridionalis, P. nigropicta, and P. punctipennis have very similar 

male terminalia, especially the presurstylus with a developed 

basoventral process that is rounded on its ventral margin. 

PANAMÁ. Chiriqui: Cerro Punta (2 km W; 08°51.1′N, 

82°35.9′W; 1700 m), 19 May–8 Jun 1977, S. Peck, H. Howden 

(1♀; CNC); Hato del Volcán (15 km NW; 08°48.8′N, 82°40.6′W; 

1200 m), 24–31 May 1977, S. Peck, H. Howden (2♂, 4♀; CNC); 

Monte Lirio (08°47.5′N, 82°49.5′W), 27 Aug 1923, R. C. Shannon (5♂, 1♀; ANSP); El Valle (08°25.9′N, 82°19.8′W), Dec 1953 

(1♂; USNM). Cocle: Playa Santa Clara (08°22.4′N, 80°06.3′W), 

2 Jul 1967, W. W. Wirth (2♂, 8♀; USNM). Guna Yala: San 

Blas, Rio Carti Grande near coast (09°27.3′N, 78°57.8′W), 2 

Mar 1985, O. S. Flint Jr., J. E. Louton (2♂; USNM). Panamá:

Canal Zone (08°48′N, 79°34′W), Dec 1946, N. L. H. Krauss 

(1♀; USNM); La Hermita (08°26.2′N, 80°02.7′W), 18 Sep 

1952 (1♀;USNM); Pedro Miguel (09°01.1′N, 79°36.7′W), 

R. C. Shannon (1♀; ANSP); Barro Colorado Island (09°09.1′N, 

79°50.8′W), 16 Jun–25 Dec 1928, 1967, 1978, A. Aiello, R. 

Silberglied, W. W. Wirth, N. E. Woodley (3♂, 6♀; USNM); 

Gamboa, Pipeline Road (09°07′N, 79°42′W; Malaise trap), Jul 

1967, W. W. Wirth (7♂, 1♀; USNM). Veraguas: Cerro Campana (08°25.9′N, 81°16.9′W), Jul 1967, W. W. Wirth (1♂, 1♀; 

USNM).

PERU. Huánuco: Tingo Maria, Río Monzon Valley 

(09°16.2′S, 76°04.2′W), 23 Dec 1954, E. L. Schlinger, E. S. Ross 

MAP 13. Distribution map for Paralimna (Paralimna) meridionalis Cresson.
number 643 • 5 1

Face bicolor, pale golden on dorsal half and lateral margins and 

becoming dark gray in the ventral half; parafacial and gena silvery 

gray; gena with only few setae on middle and ventral margin; clypeus gray. Eye rounded, about as wide as high, height about twice 

the height of gena. Gena high, height subequal to length of basal 

flagellomere; gena-to-eye ratio 0.36–0.50.

Thorax: Mesonotum yellowish gray to pale golden with 

golden to reddish-brown spots, 3 slender stripes between dorsocentral rows, the lateral ones brown and the central one reddish 

brown, plus golden brown areas laterally; postpronotum, notopleuron, and supra-alar areas mostly yellowish gray; scutellum 

wholly reddish brown with lateral margin golden. Pleurae yellowish gray to pale brown with anepisternum wholly brown or 

slightly paler in middle; anepimeron slightly brownish on ventral 

margin. Wing slightly brownish; veins pale brown; costal-vein 

ratio 0.33–0.57; M-vein ratio 0.92–1.05. Legs brown with dense 

reddish-brown microtomentum; tarsi lighter with some yellow 

coloration, especially ventrally; forefemur of male (see Figure 212) 

with anteroventral and posteroventral series of setae complete 

with 1 series of long, slender, and sparse setae, as long as width 

of femur, shortened near apex of femur; foretibia of male slightly 

arcuate.

Abdomen: Tergites distinctly bicolored, tergite 1 brown 

with lateral corner yellowish gray, tergites 2–5 brown along anterior half and medially, otherwise silvery gray dorsally, ventrolateral margins of tergites brown, sometimes silvery gray. Male 

terminalia: (Figures 74–77) Presurstylus not bifurcate, horizontal 

process slender, slightly convex dorsally, with sparse slender setae, 

usually 2; in lateral view postsurstylus widest medially with wide, 

medial, anteriorly produced lobe bearing several slender, moderately elongate setae extended until half the medial region on external surface, a thin medial lobe poorly produced posteriorly; lateral 

aedeagal process more than 1⁄2 length of aedeagus, robust, apex 

rounded; aedeagus short, narrowed apically in posterior view, 

short and broad in lateral view; phallapodeme triangular in lateral 

view; hypandrium shallowly concave.

Type Material. The lectotype female of Paralimna 

molossus Schiner (designated by Cresson 1929:192) is labeled 

“TYPE [red]/Lindig 1894 Venezuela/Paralimna molossus 

Schin[er].[handwritten]/molo[s]sus [sic; handwritten] Alte Sammlung.” The lectotype is double mounted (minute pin in a block 

of wood), is in good condition (left basal flagellomere and arista 

and hindtarsomeres 2–5 of both legs missing; vertical setae broken; abdomen not dissected), and is deposited in NMW.

The lectotype male of Paralimna bistriata Hendel, designated herein, is labeled “El Cairo.Boliv.VIII.26.Lind.D.ChacoExped [black margin]/Paralimna bistriata Hend. [handwritten] 

F. Hendel det./Type. Hendel 1930 [handwritten; red ink; label 

with a black border]/SMN/Lectotype ♂ Paralimna bistriata Hendel, designated by Ale-Rocha&Mathis [red].” The lectotype is 

double mounted (minuten pin in a rectangular card), is in excellent condition (abdomen not dissected), and is deposited in 

the SMN. We also examined two female paralectotypes from the 

same locality.

Additionally, the anteroventral margin of the male forefemur of 

P. meridionalis and P. punctipennis has simple setae that are not 

flattened. Paralimna meridionalis, however, can be easily distinguished from P. punctipennis by lacking short perpendicular 

stump veins on the second section of vein M and by the wholly 

gray mesopleuron with some small spots. Specimens of P. punctipennis have one or more short, perpendicular, stump veins on 

the second section of vein M, and its mesopleuron is distinctly 

light colored on the dorsal half and dark ventrally.

16. Paralimna (Paralimna) molossus Schiner

Figures 74–77, 179–180, 212, Map 14

Paralimna molossus Schiner 1868:242.—Cresson 1947:52 [review, Neotropical species].—Lizarralde de Grosso 1989:38 [list, Argentina].—Mathis 

and Zatwarnicki 1995:122 [world catalog].

Paralimna puncticornis variety captiosa Cresson 1916:122 [Trinidad. Montserrat; HT ♂, USNM (19685)].—Cresson 1929:192 [synonymy].

Paralimna luctans Cresson 1933:66 [Argentina. Misiones: Posadas; HT ♂, 

BMNH].—Cresson 1947:52 [synonymy].

Paralimna bistriata Hendel 1930:128 [Bolivia. El Cairo, Santa Cruz de 

la Sierra: LT ♂, SMN].—Cresson 1947:53 [list, Neotropical species].—

Wirth 1968:15 [Neotropical catalog].—Mathis and Zatwarnicki 

1995:119 [world catalog]. NEW SYNONYM.

Diagnosis. This species is distinguished from congeners by the following characters: general coloration rust; face bicolor, golden gray on dorsal half and lateral margins, becoming 

gray in the ventral half, carina sometimes golden tinged; parafacial and gena silvery gray; scutellum wholly reddish brown with 

lateral margin golden; anepisternum wholly brown or slightly 

paler in middle; pedicel with a pale golden dorsal spot; legs 

brown with dense reddish- brown microtomentum, forefemur 

of male with anteroventral and posteroventral series of setae 

complete with 1 series of long, slender, and sparse setae as long 

as width of femur, shortened near apex of femur; ventrolateral 

margins of tergites brown.

Description. Body length 3.5–4.6 mm; body generally bicolored, pale brown dorsally, lateral surfaces yellowish or 

dark gray.

Head: (See Figures 179–180) Frons golden brown to 

reddish brown with linear pale golden spots at lateral margins of 

ocellar triangle and in front of anterior ocellar seta, reaching the 

anterior margin of frons, between inner vertical and interfrontal 

seta, and on fronto-orbits between proclinate fronto-orbital setae 

and just above antennal bases; ventral fronto-orbits and dorsal 

parafacial with small inconspicuous brown and pale brown spots, 

respectively, lateral to antenna and separated by yellow spot. Antennal scape and pedicel brown, pedicel with distal margin paler 

with a pale golden dorsal spot; basal flagellomere brown to yellow, according to incidence of light, with slender, elongate, pale 

setae on margin; arista with 9 long, dorsal rays. Postcranium with 

2 complete series of postocular setae, the second series irregular. 
5 2 • smithsonian contributions to zoology

2001, W. N. Mathis (68♂, 7♀; USNM); Chulumani (8 km S; 

16°35.6′S, 68°51.2′W, 3840 m), ), 9–10 Mar 2001, A. Freidberg, 

W. N. Mathis (3♂, 3♀; USNM); Chulumani (8 km S; 16°22′S, 

67°30.4′W; 1950–2100 m), 23 Mar 2001, S. D. Gaimari (1♂; 

USNM); Chulumani (8 km S; 16°35.6′S, 68°51.2′W, 1960 m), 9 

Mar 2001, A. Freidberg (2♂, 2♀; USNM); Chulumani (2 km S; 

16°23.5′S, 67°31.8′W; 1750 m), 9–10 Mar 2001, A. Freidberg 

Other Specimens Examined. Neotropical. 

ARGENTINA. Misiones: Santa Ana (27°22.1′S, 55°34.9′W), 9 

Nov 1949, M. Aczel (5♀; USNM).

BOLIVIA. Beni: Puerto Cavinas (12°32.1′S, 66°55′W), 

Jan 1921–1922, W. Mann, Bio. Expl. (2♀; USNM). La Paz:

Coroico (16°11.30′S, 67°43.6′W) (1♂; ANSP); Apa Apa (8 

km S Chulumani; 16°22′S, 67°30.4′W; 1960 m), 9–10 Mar 

FIGURES 74–77. Structures of male terminalia of Paralimna (Paralimna) molossus Schiner (Paraguay. Pirapó): (74) epandrium, 

cerci, and presurstyli, posterior aspect; (75) postsurstylus, lateral aspect; (76) aedeagus, phallapodeme, and hypandrium, lateral 

aspect; (77) same, ventral aspect. Scale bar = 0.1 mm. (See Figures 2–5 for labels of structures.)
number 643 • 5 3

1962, L. E. Peña (3♂, 2♀; CNC). Huánuco: Cochicoto (09°13′S, 

76°12.8′W), 5 Sep 1965, J. C. Hitchcock Jr. (2♂, 2♀; USNM); 

Tingo Maria, Río Monzon Valley (09°16.2′S, 76°04.2′W), 23 

Dec 1954, E. L. Schlinger, E. S. Ross (4♂; USNM); Tingo María 

(6 km S; 09°22.7′S, 75°58.4′W), 8 Feb 1984, W. N. Mathis (4♂, 

6♀; USNM); Tingo Maria (1 km S; 09°19.6′S, 75°59.8′W), 4–6 

Feb 1984, W. N. Mathis (3♀, 2♂; USNM); La Esperanza (11 km 

N Huánuco; 09°53.6′S, 76°13.3′W) 5 Feb 1984, W. N. Mathis 

(8♂, 5♀; USNM); Las Palmas (1 km S; 09°26.9′S, 75°58.6′W), 

8 Feb 1984, W. N. Mathis (12♂, 3♀; USNM); Las Palmas, Río 

Huallaga (09°26.5′S, 75°58.3′W), 8 Feb 1984, W. N. Mathis 

(1♂; USNM); Las Palmas (1 km N; 09°26.3′S, 75°58′W; small 

waterfall), 8 Feb 1984, W. N. Mathis (2♂; USNM). Junín:

Perené (10°56.8′S, 75°13.6′W), R. C. Shannon (1♀; USNM). 

Lima: Lima (12°02.8′S, 77°03.7′W), Aug 1914, H. S. Parish 

(2♂; USNM); Lima (12°02.9′S, 77°03.7′W), 2 Feb–Aug 1914, 

1984, W. N. Mathis, H. S. Parish (15♂, 7♀; USNM); Chosica 

(11°56.3′S, 76°41.6′W), C. H. T. Townsend (1♀; USNM). 

Madre de Dios: Río Manu, Erika (near Salvación; 12°50.7′S, 

71°23.3′W; 550 m), 5–6 Sep 1988, W. N. Mathis (17♂, 1♀; 

USNM); Río Manu, Cocha Salvador (11°59.9′S, 71°13.9′W; 

300 m), 14 Sep 1988, W. N. Mathis (1♂; USNM); Río Manu, 

Pakitza (11°56.6′S, 71°16.9′W; 250 m), 9–23 Sep 1988, W. N. 

Mathis (1♀; USNM). El Conde: San Bartolomé (01°08′S, 

78°36′W), Jul 1913, C. T. Brues (3♂, 5♀; USNM). Loreto: Iquitos (12 km W; 03°48.4′S, 73°20.5′W), 16 Feb 1984, W. N. 

Mathis (1♂; USNM).

TRINIDAD AND TOBAGO. Trinidad. Montserrat Hills 

(10°25′N, 61°20′W), 4 Jul 1905, A. Busck (1♂; paratype of P. 

captiosa; USNM).

VENEZUELA. “Caife,” Jan 1943, P. Anduze (1♂, 2♀: 

USNM). Barinas: Santa Rosa (08°26.4′N, 69°42′W), Dec 1942, 

P. Anduze (1♀; 1♂; USNM). Bolívar: Gran Sabana, Akuriman 

(08°16′N, 62°45.3′W), Nov–Feb 1940 (3♂; USNM). Carabobo:

San Esteban (10°23.5′N, 67°58.8′W), 26 Nov–27 Dec 1939, P. 

Anduze (4♂, 7♀; USNM); Valle Seco (10°30.5′N, 67°35.3′W), 

Jan 1940, P. Anduze (2♂; 6♀; ANSP, USNM). Lara: Parque Nacional Yacambu (09°47.1′N, 69°35′W), 6–8 Apr 1981, L. Hollenberg, A. S. Menke (1♀; USNM).

Type Locality. Neotropical. “Venezuela.”

Distribution. (Map 14) Neotropical: Argentina 

(Misiones), Bolivia (Beni, La Paz, Santa Cruz), Brazil (Minas 

Gerais, Santa Catarina, São Paulo), Chile (Arica and Parinacota, 

Tarapaca), Colombia (Magdalena), Ecuador (Azuay, Guayas, El 

Oro), Paraguay (Guaira, Pirapó), Peru (Cuzco, El Conde, Huánuco, Junín, Lima, Loreto, Madre de Dios), Trinidad and Tobago, Venezuela (Barinas, Bolívar Carabobo, Lara).

Remarks. The male terminalia of P. molossus resemble those of P. stigmata sp. nov., especially the simple presurstylus and the robust lateral aedeagal process. The other structures 

of the male terminalia, however, are quite different. Additionally, these species can be easily distinguished by the staining on 

the body, especially on the face.

(10♂, 1♀; USNM); Mapiri (15°18.6′S, 68°13′W; 720 m), 15 

Mar 2001, S. D. Gaimari (1♂; USNM); Puente Villa (2 km E; 

16°24′S, 67°38′W; 1960 m), 11 Mar 2001, W. N. Mathis (40♂, 

1♀; USNM); Puente Villa (17 km W; 16°20.9′S, 67°49′W; 2070 

m), 11 Mar 2001, W. N. Mathis (16♂; USNM). Santa Cruz:

Pedro Lorenzo (15 km S; 18°01.40′S, 63°11.84′W, 540 m), 1 

Sep 2000, S. D. Gaimari (4♀, 1♂; USNM); 1 Nov 2000, S. D. 

Gaimari (1♀; USNM); Buena Vista Hotel Flora y Fauna (4–6 

km SSE; 17°29.49′S, 63°33.152′W; 400–500 m), 1–10 Sep 2002, 

R. Leschen (1♀; USNM); Campo Guairay (24 km S Camini; 

20°11.8′S, 63°28.6′W; 870 m) 2 Sep 2000, S. D. Gaimari (1♂; 

USNM);

BRAZIL. Minas Gerais: Passos (10 km ENE; 20°43.9′S, 

46°43.4′W), 1 Jun 1991, A. L. Norrbom, R. A. Zucchi (1♂, 

3♀; USNM); Minas Gerais: Delfinópolis, Serra da Canastra, Pousada Cachoeira Paraíso (20°20.3′S, 46°47.1′W; 

790–860 m; sweeping), 3 Jan 2007, D. Amorim, G. Ribeiro 

(1♀; MZUSP); Delfinópolis, Serra da Canastra, Pousada Cachoeira Paraíso (20°20.5′S, 46°47.5′W; 800 m; light trap), 3 

Jan 2007 (1♂; 2♀; MZUSP); 3–7 Jan 2007 (2♂, 6♀; MZUSP). 

Santa Catarina: Nova Teutônia (27°11′S, 52°23′W; 300–500 

m), Apr 1950, F. Plaumann (2♂; USNM). São Paulo: São 

Paulo (23°33.6′S, 46°35.8′W) (4♂, 4♀; ANSP); Cássia do 

Coqueiros, Municipalidade Cajuru (21°16.5′S, 47°18.3′W), 

Jan 1947, M. P. Barreto (1♀; MZUSP); Córrego Azul, Araçatuba (21°12.7′S, 50°25.5′W), Mar 1947, M. P. Barreto (1♂; 

MZUSP); Fazenda Pau D’Alho (80 km N São Paulo), 28–29 

Oct 1972, R. B. Peterson (3♂; CNC). Distrito Federal: Parque 

Nacional (15°45′S, 47°50′W), 3 Mar 1970, J. M. and B. A. 

Campbell (1♀; CNC).

CHILE. Arica and Parinacota: Arica (18°28.5′S, 

70°18.9′W), Oct 1932, F. Jaffuel (1♂; USNM). Tarapaca: Azapa 

(18°31′S, 70°11′W), 8–10 Nov 1955, L. E. Peña (2♂; CNC)

COLOMBIA. Magdalena: Aracataca (10°35.5′N, 

74°11.3′W; in heavy forest with dense undergrowth), 13 Aug 

1920 (1♂; ANSP); Ujhelyi (Sierra Nevada de Santa Marta, San 

Lorenzo; 11°07′N, 74°02′W) (1♂; ANSP).

ECUADOR. Azuay: Paute (02°46.8′S, 78°45.5′W; 1800 

m), 8 Mar 1965, L. E. Peña (3♂, 3♀; CNC). El Oro: Rio Grande 

(15 km S Zaruma; 03°42.8′S, 79°36.8′W; 850 m), 9 Apr 1965, 

L. E. Peña (2♂, 1♀; CNC). Guayas: Machala (49.5 km NNE; 

02°40′S, 79°13.4′W; 40 m), 13 Jan 1978, W. N. Mathis (1♀; 

USNM). ECUADOR. “Ecuador,” 1914, H. A. Parish (1♂, 1♀; 

USNM).

PARAGUAY. Guaira: Independencia (25°41.7′S, 

56°15.5′W), 2 Oct 1951, J. Foster (1♂, 2♀; USNM); Estancia 

Molina Cue (25°03.7′S, 56°35.8′W), Oct 1939, F. Schade (1♀; 

USNM). Pirapó: Villarica (26°51.3′S, 55°32.5′W), May–Sep 

1937, 1938, 1939, F. Schade (13♂, 12♀; USNM).

PERU. “Peru,” 1914. H. A. Parish (1♀; USNM). Cuzco: 

Paucartambo, Atalaya (Río Alto Madre de Dios; 12°53.1′S, 

71°21.6′W; 600 m), 4 Sep 1988, W. N. Mathis (11♂, 5♀; 

USNM); Quincemil (13°13.7′S, 70°45.6′W; 740 m), 13–31 Aug 
5 4 • smithsonian contributions to zoology

MAP 14. Distribution map for Paralimna (Paralimna) molossus Schiner.

17. Paralimna (Paralimna) nigropicta Cresson

Figures 78–81, 182, Map 15

Paralimna nigropicta Cresson 1916:118 [Guatemala. Puerto Barrios 

(15°43.5′N, 88°35.6′W); HT ♀, OSUC].

Paralimna (Paralimna) nigropicta.—Cresson 1947:51 [review, Neotropical 

species].—Wirth 1968:15 [Neotropical catalog].—Mathis and Zatwarnicki 1995:123 [world catalog].

Diagnosis. This species is distinguished from congeners by the following characters: face silvery gray or yellowish gray with a dark brown to black velvety quadrangular short 

spot between antennal bases, face uniformly silvery or yellowish 

gray, rarely an inconspicuous pale golden-brown stripe below 

black velvety spot on the carina and sometimes paired small pale 

golden spots between largest facial setae; ventral fronto-orbits 

with a dark brown spot lateral to antenna forming a horizontal 

dark brown band at level of antennal bases; pleurae silvery gray, 

only anepisternum with dorsal margin pale brown tinged; forefemur of male concave ventrally near apex, bearing anteroventral 

row of curved, flattened setae and a posteroventral linear patch 

of short, acutely pointed, curved or straight setae, ending at 1⁄3

distal of femur; tergite 1 brown with lateral angles silvery gray.

Description. Body length 3.1–4.6 mm; body generally bicolored, pale brown, sometimes dark brown, dorsally, 

lateral surfaces silvery, yellowish, or dark gray.

Head: (See Figure 182) Head wider than high in anterior 

view; frons pale brown to dark brown with linear pale yellow spots 

at lateral margins of ocellar triangle, in front of anterior ocellar 

seta, not reaching the anterior margin of frons, between inner vertical and interfrontal seta, and on fronto-orbits between proclinate 

fronto-orbital setae and just above antennal bases; ventral frontoorbit with a dark brown spot at level of base of antenna and dorsal 

parafacial with a dark gray spot lateral of antenna, separated from 

the dark brown spot by silvery-gray spot. Antennal scape and 

pedicel brown, pedicel with distal margin paler; basal flagellomere 

pale brown, with slender, elongate, pale setae on margin; arista 

with 10–13 long, dorsal rays. Postcranium with 2 complete series 

of postocular setae. Face uniformly silvery or yellowish gray, with 

a dark brown to black quadrangular velvety short spot between 

antennal bases, rarely an inconspicuous pale golden-brown stripe 
number 643 • 5 5

km S; 28°02.2′N, 97°52.2′W; 15 m; Nueces River), 6 Jun 2004, 

W. N. Mathis (5♂, 1♀; USNM). Kleberg: Kingsville (27°31′N, 

97°51.4′W), 7 Jun 1921, F. M. Hull (1♂; USNM). Mills: Star 

(31°28.1′N, 98°19′W), 4 Apr 1946, F. A Cowan (3♂; USNM). 

San Patricio: Mathis (28°05.7′N, 97°49.7′W), 17 Jul 1954, 

R. R. Dreisbach (1♂, 1♀; USNM); Mathis (5 km SW at Nueces 

River; 28°02.3′N, 97°51.5′W), 9 Dec 1984, W. Steiner, B. Gill, 

D. Whitehead (2♀; USNM); Mathis (6.5 km S; 28°02.8′N, 

97°51.8′W; 18 m; Nueces River), 6 Jun 2004, W. N. Mathis 

(5♂; USNM). Victoria: Victoria (28°48.3′N, 97°0.2′W), 21 Jun 

1909, J. D. Mitchell (1♀; USNM).

Neotropical. BELIZE. Stann Creek: Cockscomb Basin Wildlife Sanctuary (16°47′N, 88′30′W), 5–6 Apr 1993, W. N. Mathis 

(2♂; USNM); Maya Center: Cabbage Haul Creek (16°48′N, 

88°23′W), 3 Apr 1993, W. N. Mathis (1♂; USNM); Wee Wee 

Cay (16°45.9′N, 88°08.6′W), Jan, Nov 1987, 1988, C. Feller, 

W. N. Mathis (8♂, 10♀; USNM); Hopkins (16°52′N, 88°17′W), 

3 Apr 1993, W. N. Mathis (2♂1♀; USNM); Dangriga (16°58.2′N, 

88°13.8′W), 30 Mar–4 Apr 1988, 1993, W. N. Mathis (5♂; 

USNM); Silk Grass Creek (16°54′N, 88°26′W), 3 Apr 1993, 

W. N. Mathis (7♂; USNM); Sittee Point (16°48.5′N, 88°15′W), 

25 Apr 1987, R. Faitoute, P. J. Spangler (1♂; USNM); Sittee 

River, Possum Point Biological Station (16°52.1′N, 88°22.5′W), 

6 Nov 1987, D. and W. N. Mathis (2♂, 2♀; USNM).

COSTA RICA. Guanacaste: Liberia (10°37.8′N, 

85°26.4′W), 27 Mar 1987, J. M. Hill (1♂; USNM); Liberia (16 

km NW; Rio Ahogados; 10°45.2′N, 85°30.9′W; black light), 25 

Jul 1965, P. J. Spangler (5♂; 1♀; USNM). Heredia: La Selva 

Biological Station (10°27.2′N, 84°0.2′W; trap in garden), Jan 

1993, P. Hanson (3♂; USNM). Limón: Río Banano (09°50.5′N, 

82°56′W), 9 Nov 1909, P. P. Calvert (1♂; ANSP); Westfalia (4 

km S; 09°54.5′N, 82°59′W; beach), 27 Jun 2001, W. N. Mathis 

(2♂; 2♀; USNM); Puerto Vargas (09°43.9′N, 82°48.9′W; 

beach), 27–28 Jun 2001, W. N. Mathis (2♂ 1♀; USNM). Puntarenas: Puerto Jiminez (5 km N; 10°13′N, 83°43′W; Malaise 

trap), Nov–Dec 1990, P. Hanson (1♂, 3♀; USNM); Rincón (3 

km SW; 08°42.5′N, 83°29.2′W), Oct–Dec 1990, P. Hanson (1♀; 

USNM); San Pedrillo (08°37.2′N, 83°44.1′W), 12–15 Aug 2001, 

D. and W. N. Mathis (11♂, 1♀; USNM); Montezuma (1 km 

S; 09°39′N, 85°4.3′W), 20 Jun 2001, W. N. Mathis (1♂, 2♀; 

USNM); Malpais (09°37.6′N, 85°09.1′W; beach), 2 Jun 2001, 

W. N. Mathis (5♂, 2♀; USNM); Jacó (09°37′N, 84°38.5′W, 

beach), 13 Jun 2003, D. and W. N. Mathis (1♂; USNM).

EL SALVADOR. La Libertad: La Libertad (13°40.9′N, 

89°21.6′W), 29 Oct 1965, N. L. H. Krauss (1♂; USNM). Sonsonate: Sonzacate (13°44′N, 89°43′W), 25 Jun 1958, L. J. Bottimer (1♀; USNM).

GUATEMALA. Escuintla: Nueva Concepción (14°11′N, 

91°18.4′W; 50 ft [15 m]), 17 Aug 1963, D. Q. Cavagnaro, 

M. E. Irwin (2♂; USNM). Izabal: Puerto Barrios (15°43.5′N, 

88°35.6′W), 3–14 Mar 1905 (1♀ paratype; ANSP). Zacapa:

Gualán (15°06.9′N, 89°21.7′W), 15 Feb 1905 (1♀ paratype, 

1♀; ANSP).

on the carina below the dark brown spot and extended until middle of face, and sometimes paired small pale golden spots between 

largest facial setae; parafacial and gena silvery or yellowish gray; 

gena with only few setae on middle and ventral margin; clypeus 

silvery gray. Eye slightly higher than high, height about twice the 

height of gena. Gena high, height slightly longer than the length of 

basal flagellomere; gena-to-eye ratio 0.38–0.45.

Thorax: Mesonotum mostly pale brown with pale 

yellowish-gray areas, usually along setal tracks, 3 stripes between 

dorsocentral rows, sometimes coalesced to form a large brown 

area; postpronotum, notopleuron, and supra-alar areas gray; 

scutellum brown dorsally, with apex dark brown and pale gray 

laterally. Pleurae silvery gray, only anepisternum with dorsal margin pale brown tinged. Wing semi-hyaline; veins brown; costalvein ratio 0.38–0.39; M-vein ratio 0.97–1.15. Legs pale brown 

to brown with sparse silvery-gray microtomentum; tarsi lighter 

with some yellow coloration, especially ventrally; forefemur of 

male bearing anteroventral row of curved, flattened setae and a 

posteroventral linear patch of short, acutely pointed, curved or 

straight setae, ending at distal 1⁄3 of femur; foretibia straight.

Abdomen: Tergites distinctly bicolored; tergite 1 brown 

with lateral angles silvery gray; tergites 2–6 dark brown along 

anterior 2⁄3 and medially, otherwise silvery gray. Male terminalia: 

(Figures 78–81) Presurstylus bifurcate, with basoventral process 

somewhat developed, length about 1⁄3 of horizontal process length, 

horizontal process shallowly curved; in lateral view postsurstylus 

widest medially with medial, anterior weakly produced lobe bearing long and slender setae extended along anterior 2⁄3 of external 

surface medially and an elongate digitiform curved process anterodistally; lateral aedeagal process less than 1⁄2 length of aedeagus, 

slightly curved subapically, bluntly rounded; aedeagus gradually 

tapered to membranous apex; phallapodeme broadly triangular in 

lateral view; hypandrium somewhat deeply concave.

Type Material. The holotype female of Paralimna 

nigropicta is labeled “Pt. Barrios[,] Guatemala[,] M[ar]ch.3–

14’05/Holo-TYPE Paralimna NIGROPICTA E. T. Cresson Jr 

[maroon; “Paralimna NIGROPICTA” handwritten]/OSUC 

0319623 [bar code].” The holotype is double mounted (glued 

to a paper triangle), is in excellent condition (abdomen not dissected), and is deposited in the OSUC.

Other Specimens Examined. Nearctic. UNITED 

STATES. Texas. Aransas: Aransas National Wildlife Refuge (28°06.9′N, 96°48′W), 22 Apr 1965, R. H. Jones, W. W. 

Wirth (3♂, 4♀; USNM); 20 May 1972 (1♀; USNM); Salt Creek 

(28°17.4′N, 96°57.9′W), 5 Jun 1953 (1♀; USNM). Cameron:

Brownsville (25°54.1′N, 97°51.8′W), Jun–22 Nov 1910 (2♂, 

1♀; ANSP); Harlingen (28°02.8′N, 97°51.8′W), 23 Feb–Aug 

1945, D. E. Hardy, V. L. Wooley (3♂, 8♀; USNM). Gonzales:

Ottine, Palmetto State Park (29°35.2′N, 97°35′W), 10 Jul 1950 

(2♂, 1♀; USNM). Hidalgo: Mercedes (26°09′N, 97°54.7′W), 

7 Jun 1948 (1♀; USNM); Mission (26°13′N, 98°19.5′W), 15 

Oct 1948 (1♂; USNM); Progresso (26°05.5′N, 97°57.4′W), 1 

Jul 1938, R. H. Beamer (1♂; USNM). Jim Wells: Mathis (7.5 
5 6 • smithsonian contributions to zoology

Palenque; 17°15.3′N, 92°06.9′W), 7 May 1985, W. N. Mathis 

(2♀; USNM); El Triunfo, (49 km S Jaltenango; 15°39.4′N, 

92°48.5′W), 14 May 1985, W. N. Mathis (1♂, 1♀; USNM); Finca 

Prusia (33 km S Jaltenango; 15°49′N, 92°42′W), 10–12 May 1985, 

W. N. Mathis (3♂, 5♀; USNM); Jaltenango (21 km S; 15°54′N, 

92°40′W), 16 May 1985, W. N. Mathis (31♂, 2♀; USNM); Río 

Izapa (14°55.4′N, 92°10.7′W), 21 Apr 1983, W. N. Mathis (4♂, 

HONDURAS. Cortés: San Pedro Sula (8 km S; 15°25.7′N, 

88°01.4′W), 25–26 Sep 1995, D. and W. N. Mathis (2♂: 

USNM).

MEXICO. Campeche: Quinta Chilla (18°45.1′N, 

91°34.8′W), 20 Dec 1948, H. B. Leech (2♂; USNM). Chiapas: Bochil (4 km N; 16°59.8′N, 92°53.8′W), 4 Jun 1969, 

J. J. Teskey (1♂, 1♀; CNC); Cascadas de Agua Azul (62 km SW 

FIGURES 78–81. Structures of male terminalia of Paralimna (Paralimna) nigropicta Cresson (Belize. Stan Creek): (78) epandrium, cerci, and presurstyli, posterior aspect; (79) postsurstylus, lateral aspect; (80) aedeagus, phallapodeme, and hypandrium, lateral aspect; (81) same, ventral aspect. Scale bar = 0.1 mm. (See Figures 2–5 for labels of structures.)
number 643 • 5 7

NICARAGUA. Leon: La Cruz de La India (5 km SW, Hwy. 

26; 12°43.2′N, 86°19.86′W; 215 m), 22 Jun 2007, N. E. Woodley (2♂; USNM).

PANAMÁ. Bocas del Toro: Bananas (09°41.5′N, 

82°10′W), 22 Sep 1932 (1♂; USNM). Cocle: Playa Santa Clara 

(08°22.4′N, 80°06.3′W), 2 Jul 1967, W. W. Wirth (1♂, 1♀; 

USNM).

WEST INDIES. DOMINICA. Coulibistrie (15°27.1′N, 

61°26.8′W), 21 Mar 1989, W. N. Mathis (4♂, 2♀; USNM). 

Dublanc (15°30.9′N, 61°28.1′W), 21 Mar 1989, W. N. Mathis 

(22♂, 1♀; USNM). Hampstead River (15°33.9′N, 61°22.3′W), 

22 Mar 1989, W. N. Mathis (1♂; USNM). Layou River mouth 

(15°23.6′N, 61°25.5′W), 9 Jan–24 Mar 1965, D. F. Bray, A. B. 

Gurney, H. Robinson, W. W. Wirth (9♂, 11♀; USNM). Macoucherie (15°27.1′N, 61°26.7′W; seashore), 1–15 Feb 1965, 

W. W. Wirth (2♂, 2♀; USNM).

Type Locality. Neotropical. Guatemala. Puerto 

Barrios (15°43.5′N, 88°35.6′W).

Distribution. (Map 15) Nearctic: Mexico (Morelos, 

San Luis Potosi, Sinaloa). United States (Texas). Neotropical: Belize 

3♀; USNM); Tuxtla Gutiérrez (16°44.8′N, 93°06.6′W), 2 Aug 

1969, D. Kritsch (3♂; CNC). Guerrero: Acapulco (16°51.8′N, 

99°52.9′W), 6 Aug 1954, J. G. Chillcott (3♀; CNC); Iguala near 

Río Balsas (72 km S; 17°56.1′N, 99°35.3′W), 16 Nov 1946, 

F. E. Skinner (1♂; USNM). Jalisco: Puerto Vallarta (20°36.9′N, 

105°13.8′W), 10 Mar 1983, J. R. Vockeroth (1♂, 6♀; CNC). 

Morelos: Cuernavaca (18°56.1′N, 99°13.9′W), Mar–May 

1965, N. L. H. Krauss (1♀; USNM); Cuernavaca (24 km S; 

18°50.6′N, 99°13.1′W), 15 Nov 1946, F. E. Skinner (1♂; 

1♀; USNM). Oaxaca: Tehuantepec (16°19.2′N, 95°14.5′W; 

light trap), 15–23 Jul 1964, P. J. Spangler (8♂, 15♀; USNM). 

Quintana Roo: Tulum (20°12.7′N, 87°27.9′W), 8 Jan 1992, 

J. R. Vockeroth (6♂, 2♀; CNC). San Luis Potosi: Tamazunchale (21°15.3′N, 98°47.3′W), 23 Nov 1964, F. E. Skinner (17♂, 5♀; USNM). Sinaloa: Mazatlán (23°14.2′N, 

106°24.9′W), 7 Aug 1964, J. F. McAlpine (1♀; CNC). Tabasco: Teapa (8 km SW; 17°27.2′N, 93°01.2′W), 6 May 1985, 

W. N. Mathis (13♂, 7♀; USNM). Veracruz-Llave: Fortin 

de las Flores (18°54′N, 97°W; 952 m), 2 May 1985, W. N. 

Mathis (1♂; 2♀; USNM).

MAP 15. Distribution map for Paralimna (Paralimna) nigropicta Cresson.
5 8 • smithsonian contributions to zoology

(Stann Creek, Cockscomb, Maya Center), Costa Rica (Guanacaste, 

Heredia, Limón, Puntarenas), El Salvador, Guatemala (Izabal, Escuintla, Zacapa), Mexico (Campeche, Chiapas, Guerrero, Oaxaca, 

Quintana Roo, Tabasco, Veracruz-Llave), Honduras, Nicaragua, 

Panamá (Bocas del Toro, Cocle), West Indies (Dominica).

Remarks. Although this species is generally very distinctive, its facial coloration varies from golden to silvery gray. 

Sometimes, besides the pale brown spot on the carina, there are 

paired small, rounded, faint, golden spots between the dorsal 

facial setae on specimens with a silvery gray face; also the anterior margin of the frons is dark gray on specimens with a silvery 

gray face.

Cresson (1947) wrote that this species is similar to P. 

thomae. We have also generally noted that structures of the 

male terminalia of these two species are similar, except for the 

shape of the postsurstylus. Some external characters of the body, 

however, such as the shape of the male foretibia (ventral surface 

straight in P. nigropicta and wavy in P. thomae) and posteroventral series of setae of male forefemur (short, numerous, and 

present on basal 1⁄2 or 2⁄3 of femur in P. nigropicta; in P. thomae

these setae are long, fewer in number, and restricted to the basal 

1⁄3 or 1⁄4) distinguish them. Externally the most conspicuous character to distinguish between these two species is the small, dark, 

velvety spot between the antennal bases in P. nigropicta (small, 

dark, velvety spots are lacking in P. thomae).

Specimens of P. nigropicta have also been confused with P. 

texana. The dark spot between the antennal bases, arrangement 

of posteroventral setae along the male forefemur, the shape of 

male foretibia, the wing without infuscate halos on crossveins, 

the normally developed brown bands on the abdomen, plus 

characters of the male terminalia are usually sufficient to distinguish between them.

18. Paralimna (Paralimna) pallida, sp. nov.

Figures 82–85, 183, Map 16

Diagnosis. This species is distinguished from congeners by the following combination of characters: general 

aspect of body yellowish gray with coloration washed out; scutum lighter laterally, dorsal stripes inconspicuous; palpus very 

slender; postcranium with 2 complete series of postocular setae; 

pleurae without spots, pale gray to silvery gray; forefemur of 

male with more than 1 series of setae dense, developed, and 

strong on posteroventral surface, and a series of short setae on 

anteroventral surface, not flattened; male foretibia also with conspicuous, erect numerous setae on ventral surface.

Description. Body length 4.5–5.0 mm; body generally bicolored, with faded coloration, pale brown dorsally, lateral surfaces silvery gray, sometimes yellowish gray.

Head: (See Figure 183) Frons pale brown with pale gray 

spots, 1 stripe at lateral margins of ocellar triangle in front of anterior ocellar seta extended to anterior margin of frons, between inner 

vertical and interfrontal setae, margin of eye from outer vertical seta 

to just above antennal bases; ventral fronto-orbits with a dark gray 

spot and dorsal parafacial with a pale brown spot lateral to antenna, separated by pale gray spot. Antenna pale gray; pedicel with a 

brown dorsolateral spot, with 1 dorsal, thicker seta and some short 

setae on the margin; basal flagellomere with slender, elongate, pale 

setae on margin; arista with 12–15 long, dorsal rays. Postcranium 

with 2 complete series of postocular setae. Face pale gray with a 

single slender, short, pale brown spot on carina, as long as the length 

of basal flagellomere; parafacial and gena silvery gray; gena almost 

nude, few setae only on distal margin and in middle; clypeus silvery gray. Palpus very slender. Eye rounded, about as wide as high, 

height twice the height of gena. Gena high, height slightly longer 

than the length of basal flagellomere; gena-to-eye ratio 0.44–0.46.

Thorax: Thorax with faded coloration: mesonotum pale 

gray with 3 pale brown stripes between dorsocentral rows, the 

central stripe more slender than the others, linear anteriorly and 

widened posteriorly; postpronotal lobe gray; scutellum pale gray 

with distal margin dark brown, distal ocellar pair crossing distally. 

Pleurae pale gray to silvery gray, without spots. Wing brownish tinged; veins brown to orange brown; lacking infuscation; 

costal-vein ratio 0.43–0.46; M-vein ratio 0.95–1.10. Legs with 

silvery-gray microtomentum laterally and dark gray dorsally, except basitarsomere yellow and the remaining tarsomeres become 

brown distally; forefemur of male with dense series of developed 

setae on posteroventral surface and a series of short setae on anteroventral surface, not flattened; foretibia straight also with conspicuous erect numerous setae on ventral surface.

Abdomen: Tergites bicolored but the bands are less 

defined, washed out, with pale grayish-brown bands on laterobasal half of tergites 2 and 3, plus medially on tergites 4 

and 5, otherwise gray (more yellowish dorsally and silvery gray 

laterally). Male terminalia: (Figures 82–85) Presurstylus not 

bifurcate, with horizontal process slender and slightly curved 

with short, slender, sparse setae along the length; in lateral view 

postsurstylus widest medially with a medial, anteriorly protuberant lobe bearing moderately long and slender setae and 

a pointed process posteriorly; lateral aedeagal process shortened, less than 1⁄2 length of aedeagus, robust and slightly curved 

medially in lateral view, truncated apically in posterior view; 

aedeagus rather robust and gradually narrowed toward membranous apex in dorsal view; phallapodeme broadly triangular in lateral view; hypandrium moderately deeply concave in 

posterior view.

Type Material. The holotype male of Paralimna 

pallida is labeled “BRAZIL. Amazonas: Manaus, INPA 

(03°55.8′S, 59°59.1′W, 60m), 4 May 2010[,] D. & W. N. 

Mathis/USNM ENT 00118269 [plastic bar code label]/HOLOTYPE ♂ Paralimna pallida Ale-Rocha&Mathis, INPA [red].” 

The holotype is double mounted, is in excellent condition, and is 

deposited in INPA. Five paratypes (3♂, 2♀; INPA, USNM) bear 

the same data as the holotype.

Other Specimens Examined: BRAZIL. Amazonas: Manaus, Campus Universitário (03°06′S, 59°58.5′W; 

Malaise trap), 21 Jun 1982, J. A. Rafael (1♀; INPA).
number 643 • 5 9

FIGURES 82–85. Structures of male terminalia of Paralimna (Paralimna) pallida, sp. nov. (Brazil. Amazonas): (82) epandrium, cerci, and presurstyli, posterior aspect; (83) postsurstylus, lateral aspect; (84) aedeagus, phallapodeme, and 

hypandrium, lateral aspect; (85) same, ventral aspect. Scale bar = 0.1 mm. (See Figures 2–5 for labels of structures.)

Type Locality. Neotropical. Brazil. Amazonas: 

Manaus, INPA (03°55.8′S, 59°59.1′W).

Distribution. (Map 16) Neotropical: Brazil (Amazonas).

Etymology. The species epithet, pallida, is of Latin 

derivation and means ashen or wan, alluding to the faded coloration of the body.

Remarks. The male terminalia of P. pallida sp. 

nov. resembles that of P. molossus in some respects, such as 

the simple presurstylus and the rather robust and shortened lateral aedeagal process, but it is a very easily distinguished from 

this species by its coloration (body yellowish gray with coloration washed out; scutum lighter laterally and dorsal stripes 

inconspicuous).
6 0 • smithsonian contributions to zoology

MAP 16. Distribution map for Paralimna (Paralimna) pallida, sp. nov., and P. (Paralimna) stellata, sp. nov.

19. Paralimna (Paralimna) pauca, sp. nov.

Figures 86–89, 184, 204, Map 17

Diagnosis. This species is distinguished from congeners by the following combination of characters: face dark gray 

with 3 pale golden-brown spots, 1 elongate spot on carina, the 

same length of antenna, and 2 parallel elongate rectangular spots 

between largest facial setae; pedicel with a very small, not welldefined, silvery-white spot dorsally; anepisternum mostly gray 

but dorsal and ventral margins brown mottled, bases of setae 

and setulae brown in middle; wing hyaline, crossvein dm–cu 

lacking infuscate halo; forefemur of male bearing anteroventral, 

comb-like row of not flattened, long, and numerous setae; and 

posteroventral surface of male forefemur bearing row of long 

and widely spaced setae, longest setae as long as width of femur.

Description. Body length 3.2–4.2 mm; body generally bicolored, pale brown dorsally, lateral surfaces silvery gray.

Head: (See Figure 184) Frons brown with linear yellow 

spots at lateral margins of ocellar triangle and in front of anterior 

ocellar seta extended to anterior margin of frons, between inner 

vertical and interfrontal setae, and on fronto-orbits above dorsal 

proclinate fronto-orbital seta, between proclinate fronto-orbital 

setae and just above antennal bases; ventral fronto-orbits and 

dorsal parafacial with 2 small pale brown to dark brown spots 

lateral to antenna, separated by yellow spot. Antenna brown, 

covered by yellow microtomentum; pedicel with a very small, not 

well-defined, silvery-white spot dorsally; basal flagellomere with 

slender, elongate, pale setae on margin; arista with 9–11 long, dorsal rays. Postcranium with 2 complete series of postocular setae. 

Face dark gray with 3 pale golden-brown spots, 1 elongate spot on 

carina, the same length of antenna, and 2 parallel elongate rectangular spots between largest facial setae; parafacial and gena silvery 

gray; gena with only few setae on middle and ventral margin; clypeus silvery gray. Eye slightly higher than wide, height more than 

twice the height of gena. Gena high, height slightly longer than 

length of basal flagellomere; gena-to-eye ratio 0.36–0.38.

Thorax: Mesonotum yellowish gray with pale brown 

spots, 3 wider stripes between dorsocentral rows, plus several short 

to moderately long stripes on supra-alar area; postpronotum yellowish gray; scutellum yellowish gray with base and insertion of 

setae brownish, with apex blackish brown. Pleurae silvery gray, anepisternum with dorsal and ventral brown spots and base of setae 

brown tinged in middle, small brown spots in middle sometimes 
number 643 • 6 1

clustered as larger brown area, anteroventral corner of anepimeron 

brown. Wing (see Figure 204) hyaline; veins pale brown; costalvein ratio 0.30–0.44; M-vein ratio 1.10–1.16. Legs brown with 

silvery-gray microtomentum; tarsi lighter with some yellow coloration, especially ventrally, the distal segments darkened; forefemur 

of male with anteroventral series of long, curved, numerous and 

not flattened setae, as long as width of femur, posteroventral surface with a row of long and widely spaced setae, longest setae as 

long as width of femur; foretibia of male slightly arcuate.

Abdomen: Tergites distinctly bicolored, tergite 1 dark 

brown with laterodistal corner yellow, tergites 2–5 dark brown 

along anterior 2⁄3 and medially, otherwise yellowish gray. Male terminalia: (Figures 86–89) Presurstylus not bifurcate, with horizontal 

process long, slender, and straight, slightly widened on basal half, 

with 3 long setae; in lateral view postsurstylus widest medially with 

a medial, anteriorly protuberant lobe bearing long and slender 

setae and a digitiform process posteriorly; lateral aedeagal process 

long, longer than 1⁄2 length of aedeagus, robust and slightly curved 

FIGURES 86–89. Structures of male terminalia of Paralimna (Paralimna) pauca, sp. nov. (Dominican Republic. Independéncia): (86) epandrium, cerci, and presurstyli, posterior aspect; (87) postsurstylus, lateral aspect; (88) aedeagus, phallapodeme, 

and hypandrium, lateral aspect; (89) same, ventral aspect. Scale bar = 0.1 mm. (See Figures 2–5 for labels of structures.)
6 2 • smithsonian contributions to zoology

MAP 17. Distribution map for Paralimna (Paralimna) pauca, sp. nov., and P. (Paralimna) sera Cresson.

medially in lateral view, rounded apically; aedeagus slender, hardly 

narrowed on distal 1⁄3 in posterior view; phallapodeme broadly triangular in lateral view; hypandrium deeply concave in posterior view.

Type Material. The holotype male of Paralimna pauca

is labeled “DOM[INICAN]. REP[UBLIC]. Independ[encia].: La 

Descubierta, 18°34.1′N, 71°43.8′W, 0 m, 25 Mar 1999, Wayne 

N. Mathis/USNM ENT 00089872 [plastic bar code label]/

HOLOTYPE ♂ Paralimna pauca Ale-Rocha&Mathis, USNM 

[red].” The holotype is double mounted (minuten pin in a block 

of plastic), is in excellent condition (tarsus of right midleg missing), 

and is deposited in the USNM. Two paratypes (1♂, 1♀; USNM) 

bear the same label data as the holotype. Other paratypes are as 

follows: DOMINICAN REPUBLIC. Barahona: Baoruco (Sierra 

de Neiba, Los Guineos; Rio Colorado; 18°35′N, 71°11′W), 11–12 

Aug 1990, J. Rawlins, S. Thompson (45♂, 34♀; CMP). Independencia: Los Bolos (18°37.8′N, 71°39.2′W; 1370 m), 24 Mar 1999, 

W. N. Mathis (4♂, 1♀; INPA, USNM). Pedernales: Pedernales 

(18°01.8′N, 71°44.7′W), 19–20 Mar 1999, W. N. Mathis (1♀; 

USNM); Pedernales (21 km N; Rio Mulito; 18°09.3′N, 71°45.6′W; 

270 m), 18 Mar 1999, W. N. Mathis (1♂; USNM).

Type Locality. Neotropical. Dominican Republic. 

Independencia: La Descubierta (18°34.1′N, 71°43.8′W).

Distribution. (Map 17) Neotropical: West Indies 

(Dominican Republic).

Etymology. The species epithet, pauca, is of Latin derivation and means few or little, alluding to the relatively sparse setae 

on the anteroventral and posteroventral surfaces of the forefemur.

Remarks. Externally this species is very similar to 

P. molossus, but it can be distinguished from the latter by the 

length of the setae along the male forefemur and by structures of 

the male terminalia, such as the shape of the presurstylus (singular in P. pauca sp. nov.) and the setulae of the pre- and postsurstylus that are more robust that in P. molossus.

20. Paralimna (Paralimna) pectinata Hendel

Figures 90–93, 185, 213, Map 18

Paralimna pectinata Hendel 1930:131 [Argentina. Lapango, Pilcomayo 

(25°22′S, 57°39.3′W); HT ♂, SMN].—Cresson 1947:51 [review, Neotropical species].—Wirth 1968:15 [Neotropical catalog].—Lizarralde 

de Grosso 1989:38 [list, Argentina].—Mathis and Zatwarnicki 

1995:123 [world catalog].
number 643 • 6 3

Diagnosis. This species is distinguished from congeners by the following combination of characters: face gray or 

dark gray with a single slender longitudinal brown spot on carina; pleural area distinctly lighter on dorsal half (dorsal pleurites gray, only thin band on ventral margin of anepisternum and 

anepimeron brown) and darker on ventral half (ventral pleurites 

brown); forefemur of male concave ventrally on distal half, with 

distinctly long and flattened setae on anteroventral surface and 

rows of setae on basal half of posteroventral surface; foretibia of 

male with irregular ventral surface, wavy.

Description. Body length 2.8–4.3 mm; body generally bicolored, brown dorsally and ventrally, lateral medial surfaces mostly gray, sometimes dark gray.

Head: (See Figure 185) Frons brown to reddish brown 

with linear yellowish-gray spots at lateral margins of ocellar triangle in front of anterior ocellar seta but not reaching the anterior 

margin of frons; 1 pale brown less-defined and slender stripe between inner vertical and interfrontal setae, on front-orbits above 

dorsal proclinate fronto-orbital seta, between proclinate frontoorbital setae, and just above antennal bases; ventral fronto-orbits 

with a dark brown spot and dorsal parafacial with a gray spot, 

lateral to antenna, separated by silvery-gray spot. Antennal scape 

and basal flagellomere dark gray to pale gray; pedicel brown with 

1 dorsal, thicker seta and some short setae on the margin; basal 

flagellomere with slender, elongate, pale setae on margin; arista 

with 9–11 long dorsal rays. Postcranium with 2 complete series 

of postocular setae. Face silvery gray to dark gray with a single 

linear short spot with orange-brown microtomentum on carina, 

as long as the length of basal flagellomere; parafacial and gena 

silvery gray; gena almost nude, with setae only on distal margin; 

clypeus orange brown to silvery gray. Eye rounded, about as wide 

as high, height more than twice the height of gena. Gena high, 

height slightly longer than the length of basal flagellomere; genato-eye ratio 0.40–0.44.

Thorax: Mesonotum gray to yellowish gray with brown 

spots, 3 stripes between dorsocentral rows being 2 lateral, welldefined stripes ending at level of third dorsocentral seta and 1 irregular, slender, and less-defined acrostichal stripe; a slender and 

short stripe on supra-alar area and small brown spots on base of 

setae; postpronotal lobe silvery gray; notopleuron gray with a little brown spot on dorsal margin; scutellum silvery gray with distal 

margin dark brown and basal margin pale brown. Pleural area 

distinctly lighter on dorsal half (dorsal pleurites gray, only thin 

band on ventral margin of anepisternum and anepimeron blackish brown) and darker on ventral half (ventral pleurites blackish brown). Wing semi-hyaline; veins brown, crossveins r–m and 

dm–cu with infuscate halo; costal-vein ratio 0.37–0.50; M-vein 

ratio 1.04–1.05. Legs brown to black except foretarsus lighter 

ventrally and the remaining tarsi yellow; forefemur of male (see 

Figure 213) concave ventrally on distal half, with elongate flattened setae on anteroventral surface and a dense series of setae on 

basal half of ventral surface; foretibia of male irregular on ventral 

surface, wavy.

Abdomen: Tergites distinctly bicolored, dark brown 

along anterior half and medially, otherwise yellowish gray 

dorsally and silvery gray laterally, except tergite 2 predominantly yellowish gray with medial brown line reduced and 

dorsolateral brown spot on 1⁄2 basal half. Male terminalia: (Figures 90–93) Presurstylus not bifurcate, with horizontal process 

wide basally and narrowed distally, shallowly curved; in lateral 

view postsurstylus widest medially with a medial anteriorly 

protuberant lobe bearing long and robust setae and a bulged 

process posteriorly; lateral aedeagal process about 1⁄2 length of 

aedeagus, slightly curved subapically in lateral view, bluntly 

rounded; aedeagus gradually tapered to membranous apex; 

phallapodeme broadly triangular in lateral view; hypandrium 

deeply concave.

Type Material. The holotype male of Paralimna 

pectinata Hendel is labeled “[ARGENTINA.] Lapango.Pilc.

IX.1925.Lind.D.Chaco-Exped [black margin]/Paralimna pectinata Hend. ♂ [handwritten] F. Hendel det./Type. Hendel 1930 

[handwritten; red ink; label with a black border]/SMN.” The 

holotype is double mounted (minuten pin in a paper card), is 

in excellent condition (left wing slightly torn; abdomen not dissected), and is deposited in the SMN.

Other Specimens Examined. ARGENTINA. 

Catamarca: El Pintado, S La Viña (28°03.3′S, 65°35.1′W; 

650 m), 27–28 Sep 1968, L. E. Peña (2♂; CNC); La Merced 

(28°09.9′S, 65°41.1′W; 650 m), 26 Sep 1968, L. E. Peña (1♀; 

CNC); Yunka Suma, N Aconqulja (27°23.1′S, 65°58.9′W; 

5 m), 2 Oct 1968, L. E. Peña (1♀; CNC). Jujuy: Aqua Caliente 

(NE Guemes; 24°02.7′S, 66°31.2′W; 1100 m),18–19 Oct 1968, 

L. E. Peña (1♂; CNC); Ing. Ledesma, near Gen. San Martin (23°50.1′S, 64°47.5′W; 1100 m), 10 Oct 1968, L. E. Peña 

(1♂; CNC); Mendoza (24°10.7′S, 65°17.9′W), Feb 1970, L. E. 

Peña (1♀; MZUSP). Córdoba: Ascochinga (30°57′S, 64°16′W), 

28 Feb 1971, L. E. Peña (10♂, 11♀; MZUSP). Misiones (27°S, 

54°35′W): 1927, F. and M. Edwards (1♂, 1♀; ANSP). Salta: Rio 

Juramento (60 km S Guemes; 24°49.4′S, 65°24.4′W; 650 m), 18 

Oct 1968, L. E. Peña (1♀; CNC). Tucumán: Arroyo (14 km S 

El Tala; 26°56.4′S, 65°09.7′W; 700 m), 13–14 Oct 1968, L. E. 

Peña (2♂, 2♀; CNC).

BOLIVIA: Cochabamba: Cochabamba (17°23.3′S, 

66°07′W; 2610 m), 26 Jan–25 Mar 1976, 2001, W. N. Mathis, 

L. E. Peña (15♂, 2♀; CNC, USNM).

BRAZIL: Amazonas: Sítio Vida tropical (02°51.9′S, 

59°55.9′W, 60m), 5 May 2010, D. and W. N. Mathis (15♂, 

7♀; INPA, USNM). Espírito Santo: Baixo Guandu (19°31′S, 

41°01.1′W), Sep 1970, P. C. Elias (8♂, 19♀; MZUSP). Goiás: 

Jataí (17°53.6′S, 51°42.9′W), Jan 1955, M. Carrera (1♂; 

MZUSP). Pará: Oriximiná-Obidos (01°38.7′S, 55°37′W), Nov 

1969, Expedição Permanente da Amazônia (1♀; MZUSP); Taperinha, Santarém (02°31.9′S, 54°17.7′W), Nov 1970 (1♂; 

MZUSP); Belterra, Flona Tapajós (02°36.1′S, 54°56.6′W; 100 

m; light trap), 15–16 Apr 2008, J. A. Rafael, F. F. Xavier (1♀; 

INPA); Rio Nhamundá, Igarapé Areias (01°35.2′S, 57°37.6′W; 

25 m; Malaise trap), 17–20 May 2008, J. A. Rafael and team (1♀; 

INPA); Alter do Chão, Igarapé Sonrisal (02°31.4′S, 54°57.5′W; 

light trap), 18 Apr 2008, J. A. Rafael, F. F. Xavier (2♂; INPA). 

Paraná: Bocaiúva do Sul (25°14.9′S, 49°8.9′W; 890 m), 2–4 
6 4 • smithsonian contributions to zoology

Nov 2010, D. and W. N. Mathis (1♂, 1♀; USNM); Castro 

(24°47.4′S, 50°0.3′ W; wetlands), 24–25 Dec 2009, D and W. N. 

Mathis (2♂; USNM); Curitiba, UFPR [Universidade Federal do 

Paraná], (25°26.9′S, 49°14′; 915 m), 6 Feb 2010, D. and W. N. 

Mathis (2♂; USNM); 28 Dec–15 Jan 2009, 2010, D. and W. N. 

Mathis (3♂; USNM); Matinhos (N; 25°47.4′S, 48°31.6′W; 1 m), 

30 Jan 2010, D. and W. N. Mathis (3♂; USNM); Rio Da Onça 

(25°47.4′S, 48°31.6′W; 3 m), 12 Nov 2010, D. and W. N. Mathis 

(8♂; USNM). Pernambuco: Olinda (08°0.6′S, 35°51.3′W), Oct 

1961, N. L. H. Krauss (1♂;USNM); Tapera (08°24′S, 38°05′W), 

17 Apr 1969, H. S. Lopes (1♂; MZUSP). Rio Grande do Norte:

Natal (05°47.7′S, 35°12.6′W), Jul 1955, P. Pereira (4♀, 4♂; 

MZUSP). Santa Catarina: Nova Teutônia (27°11′S, 52°23′W; 

300–500 m), 1971, F. Plaumann (1♀; MZUSP).

FIGURES 90–93. Structures of male terminalia of Paralimna (Paralimna) pectinata Hendel (Argentina. Córdoba): (90) epandrium, cerci, and presurstyli, posterior aspect; (91) postsurstylus, lateral aspect; (92) aedeagus, phallapodeme, and hypandrium, 

lateral aspect; (93) same, ventral aspect. Scale bar = 0.1 mm. (See Figures 2–5 for labels of structures.)
number 643 • 6 5

GUYANA. Conservation of Ecological Interactions and 

Biotic Associations (~40 km S Georgetown; 06°29.9′N, 

58°13.1′W), 13 Apr 1994 (1♂; USNM). Dubulay Ranch-Aramatani Creek (05°40.9′N, 57°51.5′W), 9–11 Apr 1994, W. N. 

Mathis (12♂; USNM). Kanuku Mountains, Kumu River and Falls 

(03°15.9′N, 59°43.5′W), 28–30 Apr 1995, W. N. Mathis (3♂, 

1♀; USNM). Karanambo, Rupununi River, ox bow (03°45.1′N, 

59°18.6′W), 31 Mar–2 Apr 1994, W. Mathis (32♂, 3♀; USNM). 

Kumo Pond, Lethem (~20 km S; 03°16.7′N, 59°43.5′W), 30 Apr 

1995, W. N. Mathis (1♂; USNM). Paramakatoi (04°42′N, 

59°42.8′W), 24–25 Aug 1997, W. N. Mathis (2♂; USNM). 

Pirara Ranch, Cashew Creek (03°37.3′N, 59°44.2′W), 23 Apr 

1995, W. N. Mathis (26♂, 3♀; USNM). Pirara Ranch and River 

(03°32.1′N, 59°40.5′W), 24–25 Apr 1995, W. N. Mathis (10♂; 

USNM). Wiruni River (05°46.6′N, 58°0.8′W), 11 Apr 1994, 

W. N. Mathis (1♀; USNM).

PARAGUAY. Itapua: Encarnación (27°20.4′S, 55°52′W), 

16 Jan 1927, F. and M. Edwards (1♂, 1♀; ANSP).

TRINIDAD and TOBAGO. Trinidad. Mayaro: Plaisance (5 km N; 10°16′N, 61°28′W), 23 Mar 1985, G. F. and 

J. F. Hevel (49♂, 33♀; USNM). Port of Spain: Port of Spain 

(10°40′N, 61°31′W), Jun 1953, U.S. Army, 25 Medical Detachment (1♂; USNM). St. Andrew: Lower Manzanilla (5 km S; 

10°28′N, 61°03′W), 20 Jun 1993, W. N. Mathis (1♂; USNM); 

Lower Manzanilla (12 km S; 10°24′N, 61°02′W), bridge over 

Nariva River, 20–27 Jun 1993, W. N. Mathis (2♂; USNM); 

Lower Manzanilla (16 km S; 10°22′N, 61°01′W), 20 Jun 1993, 

W. N. Mathis (6♂, 1♀; USNM); Valencia (1 km W; 10°39′N, 

61°13′W), Aripo River, 20 Jun 1993, W. N. Mathis (7♂, 1♀; 

USNM); Valencia (near; 10°39.4′N, 61°09.1′W), 23 Mar 1985, 

G. F. and J. F. Hevel (1♂; USNM). St. George: Caura River 

Valley (end of Caura Royal Road; 10°42′N, 61°21′W), 31 Mar 

1996, C. Chaboo (13♂, 15♀; AMNH).

URUGUAY. Montevideo: Montevideo (35°53′S, 

56°10.9′W), 21–22 Jan 1927, F. and M. Edwards (1♀; ANSP).

VENEZUELA: Bolívar: Guasipati (40 km N; 07°43.7′N, 

61°11.6′W; at marsh, lake), 20 Mar 1982, G. F and J. F. Hevel 

(29♂, 26♀; USNM).

Type Locality. Neotropical. Argentina. Formosa: 

Lapango, Pilcomayo (25°22′S, 57°39.3′W).

Distribution. (Map 18) Neotropical: Argentina 

(Catamarca, Córdoba, Jujuy, Misiones, Salta, Tucumán), Bolivia 

MAP 18. Distribution map for Paralimna (Paralimna) pectinata Hendel.
6 6 • smithsonian contributions to zoology

(Cochabamba), Brazil (Amazonas, Espírito Santo, Goiás, Pará, 

Paraná, Pernambuco, Rio Grande do Norte, Santa Catarina), 

Guyana, Paraguay (Itapua), Trinidad and Tobago, Uruguay 

(Montevideo), Venezuela (Bolívar).

Remarks. This species has a variable abdominal pattern; specimens from Bolivia have the coloration of abdomen 

very uniform. This species is similar to P. fellerae by the coloration of mesopleuron that is distinctly bicolored, silvery gray on 

the dorsal half and brown on the ventral half, but differs by the 

coloration of the face and the shape of the male forefemur and 

foretibia. There is 1 brown spot on the carina in P. pectinata and 

3 spots on P. fellerae; both species have flattened setae on the 

anteroventral surface of the male forefemur, but in P. pectinata

the forefemur is concave ventrally on the distal half, the flattened 

setae are longer, curved, and wider than in P. fellerae, and the 

foretibia has the ventral surface distinctly sinuous (straight in P. 

fellerae).

This species differs from P. punctipennis by the male forefemur being concave ventrally and the foretibia is sinuous ventrally 

(not sinuous in P. punctipennis). Additionally, the basoventral 

process of the presurstylus is distinctly developed and the horizontal process is slender in P. fellerae and P. punctipennis, and 

the basoventral process is undeveloped and the horizontal process is robust in P. pectinata.

21. Paralimna (Paralimna) piger Cresson

Figures 94–97, 186, Map 10

Paralimna piger Cresson 1933:65 [“Ecuador”; HT ♂, USNM (44805)]; 

1947:51 [review, Neotropical species].—Wirth 1968:15 [Neotropical 

catalog].—Mathis and Zatwarnicki 1995:123 [world catalog].

Diagnosis. This species is distinguished from congeners by the following characters: face dark gray with 3 brown 

spots, 1 on the carina and 2 longitudinal, parallel brown spots 

between largest facial setae, extending to margin of face; gena 

brown with dorsal margin silvery gray; pleurae predominantly 

brown, anepisternum brown on ventral margin, dorsal margin 

spotted by brown bases of setae, and a large central rounded 

stain, the remaining gray; propleuron, anterior and posterior 

margins of katepisternum, anatergite, and katatergite gray; 

forefemur with ventral surface slightly convex, with very long 

posteroventral setae on basal half of the femur, longer than the 

larger region of the femur, anteroventral series of flattened setae 

occupying basal 3⁄4 length of femur; lateral margin of tergites 

brown; male foretibia with ventral surface irregular, wavy.

Description. Body length 2.7–4.2 mm; body predominantly brown with slender gray stripes on scutum and small 

gray areas on pleurae and on lateral of abdomen.

Head: (See Figure 186) Frons brown with linear gray 

spots at lateral margin of ocellar triangle, in front of anterior ocellar seta and extended anteriorly discreetly to the anterior margin 

of frons, a linear inconspicuous spot between inner vertical setae 

and interfrontal seta, with a small quadrate spot just below inner 

vertical setae, and on fronto-orbits between proclinate frontorbital setae and just above antennal bases; ventral fronto-orbits 

and dorsal parafacial with 2 dark brown spots lateral to antenna, 

separated by silvery-white spot. Antennal scape and pedicel brown 

with silvery-gray microtomentum, pedicel with silvery-gray microtomentum on distal margin; basal flagellomere pale brown covered by dense pale micropilosity, longer on dorsal margin; arista 

with 10 long, dorsal rays. Postcranium with 2 complete series of 

postocular setae. Face dark gray with antennal groove whitish 

gray; lateral margins brown; with 3 brown spots, 1 longitudinal 

spot on the carina, as long as antennal length, and 2 longitudinal, parallel brown spots between largest facial setae, extending 

to margin of face; gena bicolored, brown with dorsal margin silvery gray, parafacial and clypeus silvery gray; gena with few setae 

only on middle and ventral margin. Eye rounded, about as wide as 

high, height more than three times the height of gena. Gena high, 

height subequal to length of basal flagellomere; gena-to-eye ratio 

0.27–0.30.

Thorax: Mesonotum mostly brown with longitudinal 

linear gray spots, usually along setal tracks, becoming more gray 

laterally through postpronotum and notopleuron, postalar area 

and lateral margin of scutellum gray, dorsum of scutellum brown. 

Pleurae predominantly brown, anepisternum brown on ventral 

margin and on a large central rounded stain, dorsal margin spotted by brown insertion of setae, the remaining gray; propleuron, 

anterior and posterior margins of katepisternum, anatergite, and 

katatergite gray; setae of anepisternum brown, short, slender, 

several longer setae below middle of anepisternum. Wing semihyaline; veins brown; costal-vein ratio 0.45–0.46; M-vein ratio 

1.00–1.05. Legs brown with very sparse gray microtomentum, 

except tarsomeres lighter with yellow coloration. Forefemur 

with ventral surface slightly convex, with anteroventral series of 

flattened setae, the setae longer on middle of femur and occupying basal 3⁄4 length of femur; setae of anteroventral surface very 

long on basal half of the femur, longer than the larger region of 

the femur; male foretibia with ventral surface wavy.

Abdomen: Tergites distinctly bicolored, dark brown 

along anterior half, medially, and on lateral margin, otherwise 

gray. Male terminalia: (Figures 94–97) Presurstylus bearing short, 

slender setae, not bifurcate, horizontal process almost straight; in 

lateral view postsurstylus widest medially with sparse slender setae 

in middle; lateral aedeagal process less than 1⁄2 length of aedeagus, 

slightly curved in middle and rounded apically; aedeagus slightly 

narrowed apically in anterior view; in lateral view phallapodeme 

wider on basal 2⁄3 and slender on the remainder; hypandrium shallowly concave; cerci long and narrowed, almost 4× longer than 

wide.

Type Material. The holotype male of Paralimna 

piger is labeled “ECUADOR[,] HA Parish[,] 1914/♂/TYPE 

Paralimna PIGER E. T.Cresson,Jr. [“Paralimna PIGER” handwritten; pink]/Type No 44805 U S N M [“44805” handwritten; red].” The holotype is pinned directly, is in good condition 
number 643 • 6 7

(some Lepidoptera scales; abdomen not dissected), and is deposited in the USNM (44805).

Other Specimens Examined. ECUADOR. Carchi: Los Duendes (0°28′S, 77°56′W; 2000 m), 10 Jun 1965, L. E. 

Peña (8♂, 2♀; CNC). Chimborazo: Chilicay (02°15′S, 79°06′W), 

Jul 1955, R. Levi-Castillo (7♂; USNM). Cotopaxi: Latacunga (79 

km E Quevedo; 00°52.2′S, 78°37.1′W; 3300 m), 12 May 1975, 

A. Gurney (2♂, 1♀; USNM); Quevedo (66 km E; 00°57.7′S, 

78°54.1′W), 15 Jan 1978, W. N. Mathis (2♂, 1♀; USNM); 

Tingo (0°54′S, 79°03′W; 1000 m), 5–6 Jul 1965, L. E. Peña (1♀; 

CNC). El Oro: Pasaje (6 km E; 03°18.1′S, 79°47.1′W), 13 Jan 

1978, W. N. Mathis (9♂, 6♀; USNM). Esmeraldas: Río Guayllabamba (00°29′S, 79°25.6′W; 1600 m), Jan 1971, L. E. Peña 

(2♂, 3♀; MZUSP). Guayas: El Triunfo (60 km N Guayaquil; 

FIGURES 94–97. Structures of male terminalia of Paralimna (Paralimna) piger Cresson (Ecuador): (94) epandrium, cerci, 

and presurstyli, posterior aspect; (95) postsurstylus, lateral aspect; (96) aedeagus, phallapodeme, and hypandrium, lateral 

aspect; (97) same, ventral aspect. Scale bar = 0.1 mm. (See Figures 2–5 for labels of structures.)
6 8 • smithsonian contributions to zoology

01°55.3′S, 80°0.7′W), 11 Feb 1973, M. A. Deyrup (4♂, 3♀; 

USNM); Machala (49.5 km NNE; 03°03.9′S, 79°44.8′W; 40 m), 

13 Jan 978, W. N. Mathis (29♂, 14♀; USNM); Rio Frio, Balao 

Chico (02°45.1′S, 79°44.6′W), 26–30 Apr 1965, L. E. Peña 

(2♀; CNC). Imbabura: Río Taguando, NW Ibarra (0°24.1′S, 

78°08′W; 1650–1900 m), 9 Jun 1965, L. E. Peña (72♂, 31♀; 

CNC); Yaguarcocha (3 km N Ibarra; 0°22.4′S, 78°06.1′W; 1950 

m), 8–9 Jun 1965, L. E. Peña (11♂, 5♀; CNC). Loja: Catamayo 

(03°59′S, 79°21′W; 1500 m), 24 Mar 1965, L. E. Peña (8♂, 2♀; 

CNC); Loja (S; 03°59.4′S, 79°12.3′W; 1200 m), 26 Mar 1965, 

L. E. Peña (4♂, 6♀; CNC); Loja (W; 03°58.5′S, 79°14.8′W; 2500 

m), 25 Mar 1965, L. E. Peña (1♀; CNC); San Pedro, Zaruma 

Road (03°45.8′S, 79°38.1′W; 1100 m), 9 Apr 1965, L. E. Peña 

(22♂, 25♀; CNC); Saraguro (S; 03°37.5′S, 79°14.6′W; 2900 

m), Nov 1970, L. E. Peña (5♂, 3♀; MZUSP). Los Ríos: Quevedo (11 km S; 01°09′S, 79°26.7′W), 3 Jul 1975 (1♂; USNM); 

Río Palenque (0°35′S, 79°22′W; 150 m), 22–26 Feb 1976, 

G. E. Shewell (1♂, 1♀; CNC). Manabi: Pedro Carbo (45 km NW; 

01°42.7′S, 80°24.7′W), 11 Jan 1978, W. N. Mathis (8♂, 8♀; 

USNM); Quevedo (117 km W; 01°03.9′S, 80°14.7′W; 915 m), 

1 Jul 1975, A. Langley, J. Cohen (30♂, 12♀; USNM); Quevedo 

Road, Portoviejo (01°03.3′S, 80°27.2′W; 400 m), 13 Apr 1965, 

L. E. Peña (1♂; CNC). Orellana: Coca, Napo River (0°27.8′S, 

76°59′W; 250 m), 12–130 Apr 1965, L. E. Peña (6♂, 4♀; CNC); 

Pomeya, Napo River (0°26.5′S, 76°36.6′W), 25–30 Apr 1965, 

L. E. Peña (1♂; CNC). Pichincha: Guaillabama (0°03.5′S, 

78°20.5′W; 1850 m), 7 Jun 1965, L. E. Peña (2♂, 1♀; CNC); 

Tandapi (40 km SW Quito; 0°24.9′S, 78°48′W; 1300–1500 

m), 15–21 Jun 1965, L. E. Peña (69♂, 36♀; CNC); Tinalandia 

(00°15.3′S, 79°10.4′W; 800–1000 m), 8 Feb 1983, L. Huggert, 

L. Masner (1♂, 1♀; CNC, USNM). Santa Elena: Rio Ayampe 

(01°40.3′S, 80°42.2′W), 26 Jul 1976, J. Cohen (1♂; USNM). 

Santo Domingo de los Tsáchilas: Río Toachi (4 km E of Alluriquin; 00°05.4′S, 79°12′W), 17 Apr 1977, E. R. S. Hodges 

(1♂; USNM); Santo Domingo de Los Colorados (00°15.3′S, 

79°10.4′W), 27 Feb–6 Mar 1973, M. A. Deyrup, M. R. Wheeler 

(2♂, 6♀; USNM); Santo Domingo de Los Colorados (14 km E; 

00°13.9′S, 79°05.7′W), 5 Jul 1976, J. Cohen, A. Langley (1♂; 

USNM); Santo Domingo de Los Colorados (47 km S; 00°24.5′S, 

79°18′W; 220 m), 7 Jan–29 Jul 1976, 1978, J. Cohen, W. N. 

Mathis (23♂, 20♀; USNM).

Type Locality. Neotropical. “Ecuador.”

Distribution. (Map 10). Neotropical: Ecuador 

(Carchi, Chimborazo, Cotopaxi, El Oro, Esmeraldas, Guayas, 

Imbabura, Loja, Los Ríos, Manabi, Pichincha, Santa Elena, 

Santo Domingo de los Tsáchilas).

Remarks. This species is very similar to P. puncticornis in the coloration of the face and mesopleuron but can be 

distinguished by the brown gena, the sinuous ventral surface of 

the foretibia of males, the anteroventral setae that are long and 

widely flattened, and the posteroventral setae that are very long 

(longer than width of femur) and present along only the basal 

half of the femur in P. piger (gena gray, tibia straight, anteroventral setae slightly flattened, and ventral series complete and not 

so long in P. puncticornis). The more distinctive feature of the 

male terminalia is the shape of the presurstylus and the anteromedial process that is pointed apically in P. piger, whereas it is 

larger and with a rounded margin in P. puncticornis.

22. Paralimna (Paralimna) pleurivittata

Cresson

Figures 98–101, 187, 205, Map 19

Paralimna pleurivittata Cresson 1916:122 [Peru. Lima: Callanga (12°19′S, 

75°35.6′W); HT ♂, HNHM]; 1947:53 [review, Neotropical species].—

Wirth 1968:15 [Neotropical catalog].—Mathis and Zatwarnicki 

1995:123 [world catalog].

Diagnosis. This species is distinguished from congeners by the following combination of characters: frons protuberant and almost horizontal; face long, about twice the length of 

frons, yellowish gray to golden brown in the most part, silvery 

gray on central portion and distal margin, with 2 longitudinal, 

parallel golden-brown spots between largest facial setae; eye rather 

rectangular, wider than high in lateral view; mesopleuron with a 

wide golden-brown stripe extending from prostigma over mostly 

anepisternum to wholly anepimeron, dorsal and ventral margins 

of anepisternum paler; lateroventral margin of tergites brown; 

forefemur with anteroventral row of setae, elongate on basal half 

of forefemur, short and flattened on distal half, and posteroventral 

series of elongate setae; wing slightly brown, transverse veins with 

infuscate halo; pedicel with a silvery white spot dorsally.

Description. Body length 2.5–4.6 mm; body predominantly brown dorsally, gray or dark gray ventrally, and lateral medial surfaces mostly brown.

Head: (See Figure 187) Frons protuberant and almost 

horizontal, pale brown with linear pale yellow spots at lateral 

margins of ocellar triangle, in front of anterior ocellar seta, 

reaching the anterior margin of frons, between inner vertical 

and interfrontal seta, and on fronto-orbits above dorsal proclinate fronto-orbital seta, silvery gray between proclinate frontoorbital setae and just above antennal bases; ventral fronto-orbits 

with a dark brown spot and dorsal parafacial with a pale brown 

spot lateral to antenna, separated by silvery-gray spot. Antenna 

inserted high on face; scape and pedicel red brown, pedicel with 

silvery-white spot dorsally; basal flagellomere pale brown to 

brownish golden yellow according to incidence of light, with 

slender, elongate, pale setae on margin; arista with 15 long, 

dorsal rays. Postcranium with 1 complete series of postocular 

setae. Face long, about twice the length of frons; yellowish gray 

to golden brown in the most part, silvery gray on central portion 

and distal margin, with 2 longitudinal, parallel golden-brown 

spots between largest facial setae; parafacial silvery gray and 

gena yellowish gray; gena with only few setae on middle and 

ventral margin; clypeus gray. Eye rather rectangular, slightly 

wider than high in lateral view, height twice the height of gena. 

Gena high, height slightly longer than length of basal flagellomere; gena-to-eye ratio 0.40–0.44.
number 643 • 6 9

Thorax: Mesonotum with coloration faded, mostly pale 

golden brown with yellowish gray areas, usually along setal tracks, 

2 pale golden-brown stripes between dorsocentral rows; postpronotum, notopleuron, and supra-alar areas gray; scutellum brown 

dorsally, with apex blackish brown and silvery gray laterally. Pleurae: a wide golden-brown stripe extending from prostigma over 

most of anepisternum to whole of anepimeron, dorsal and ventral 

margins of anepisternum paler; metapleuron, laterotergite, katepisternum, and meron gray. Wing (see Figure 205) faintly brown, 

transverse veins with infuscate halo; costal-vein ratio 0.46–0.51; 

M-vein ratio 1.12–1.20. Legs brown with sparse silvery-gray microtomentum; tarsi lighter with some yellow coloration, especially 

ventrally; forefemur with anteroventral series of setae, elongate 

on basal half of forefemur, short and flattened on distal half, and 

posteroventral series of elongate setae; foretibia straight.

Abdomen: Tergites distinctly bicolored, brown along anterior half, medially, and on lateroventral margin, otherwise gray. 

Male terminalia: (Figures 98–101) Presurstylus not obviously 

FIGURES 98–101. Structures of male terminalia of Paralimna (Paralimna) pleurivittata Cresson (Peru. Lima): (98) epandrium, cerci, and presurstyli, posterior aspect; (99) postsurstylus, lateral aspect; (100) aedeagus, phallapodeme, and hypandrium, lateral aspect; (101) same, ventral aspect. Scale bar = 0.1 mm. (See Figures 2–5 for labels of structures.)
7 0 • smithsonian contributions to zoology

MAP 19. Distribution map for Paralimna (Paralimna) pleurivittata Cresson.

bifurcate, just widened basoventrally, with 2 short and robust 

setae inserted over tuberculate bases, horizontal process almost 

straight, bearing 2 slender setae near apex; in lateral view postsurstylus widened medially with anteromedial lobe large with 

truncate apex bearing elongate, slender setae, setae extended over 

median line of external surface, and a short, pointed anteromedial process; lateral aedeagal process about 1⁄2 length of aedeagus, 

robust, slightly curved in middle, rounded apex; aedeagus rather 

slender, tapering gradually toward membranous apex in posterior 

view; in lateral view phallapodeme triangular, not much developed; hypandrium moderately deeply concave; cerci long, slender, 

almost 5× longer than wide.

Type Material. The holotype male of Paralimna 

pleurivittata was not examined, but we examined the male and 

female paratypes (USNM) from the same series.

Other Specimens Examined: BOLIVIA. Beni:

Ivón (Mulford Biological Expedition; 11°07.6′S, 66°07.9′W), 

Feb 1921–1922, W. M. Mann (1♀; USNM); Rurrenabaque 

(14°26.5′S, 67°31.7′W; 175 m), 10–23 Oct 1956, L. E. Peña 

(1♀; CNC).

BRAZIL. Amazonas: Ipixuna, Rio Gregório, Comunidade 

Lago Grande (07°09.9′S, 70°48.9′W; light trap), 21 May 2011, 

J. A. Rafael, J. Câmara, R. Silva. A. Somavilla (1♂; INPA). 

Pará: Rio Xingu (camp, about 60 km S Altamira; 03°39′S, 

52°22′W), 8–16 Oct 1986, O. S. Flint Jr., P. J. Spangler (3♂, 

10♂; USNM).

PERU. Lima: Callanga (12°19′S, 75°35.6′W) (1♂, 1♀; paratypes; USNM). Madre de Dios: Avispas (12°58.7′S, 70°20.8′W; 

400 m), 1–15 Oct 1962, L. E. Peña (22♂, 22♀; CNC); Río 

Manu, Erika (near Salvación; 12°50.7′S, 71°23.3′W; 550 m), 

5–6 Sep 1988, W. N. Mathis (6♂, 1♀; USNM).

Type Locality. Neotropical. Peru. Lima: Callanga 

(12°19′S, 75°35.6′W).

Distribution. (Map 19) Neotropical: Bolivia (Beni), 

Brazil (Amazonas, Pará), Peru (Lima, Madre de Dios).

Remarks. This species is very easily distinguished 

from other species of Paralimna by the set of characters presented in the diagnosis. In addition, the male terminalia have a 

presurstylus with a weakly developed basoventral process, the 

postsurstylus has an anteromedial lobe that is rather quadrate 
number 643 • 7 1

and bears strong setae, and the posteromedial process is digitiform.

23. Paralimna (Paralimna) plumbiceps Cresson

Figures 102–105, 188, Map 12

Paralimna plumbiceps Cresson 1916:110 [West Indies. “Jamaica”; HT ♂, 

ANSP (6090)].

Paralimna (Paralimna) plumbiceps.—Cresson 1947:51 [review, Neotropical 

species].—Wirth 1968:15 [Neotropical catalog].—Mathis and Zatwarnicki 1995:123 [world catalog].

Diagnosis. This species is distinguished from congeners by the following combination of characters: face mostly 

entirely gray to whitish gray, sometimes slightly yellowish or 

with an inconspicuous, rather rectangular golden-brown spot 

on carina and 2 small, rounded spots between largest facial 

setae; frons mottled, dark to whitish brown; scutellum guttate, mostly brown, especially base of setae; anepisternum and 

katepisternum mostly concolorous (anepisternum with dorsal 

margin sometimes brown), faintly yellowish gray to silvery 

gray; forefemur of male concave ventrally near apex, bearing anteroventral, comb-like row of distinctly flattened, long 

setae; posteroventral surface of male forefemur bearing row of 

long setae along most of length; foretibia of male with ventral 

surface wavy, covered by conspicuous dense elongate ventral 

setae.

Description. Body length 3.0–3.5 mm; body generally bicolored, brown dorsally, lateral surfaces dark or silvery 

gray.

Head: (See Figure 188) Frons grayish brown becoming 

dark gray anteriorly, with yellowish-gray spots poorly defined, a 

stripe at lateral margins of ocellar triangle, in front of anterior 

ocellar seta, not reaching the anterior margin of frons, between 

inner vertical and interfrontal setae, and on fronto-orbits between 

proclinate fronto-orbital setae and just above antennal bases; spots 

on ventral fronto-orbits and dorsal parafacial little evident. Antennal scape and pedicel grayish brown, pedicel with distal margin 

paler; basal flagellomere pale brown, with slender, elongate, pale 

setae on margin; arista with 9 long dorsal rays. Postcranium with 

2 series of postocular setae, the inner series irregular. Face gray 

to whitish gray with a golden-brown, sometimes inconspicuous, 

rectangular spot on the carina, the same length of antenna, and 

2 small, rounded spots between largest facial setae; parafacial and 

gena silvery gray; gena with only few setae on middle and ventral 

margin; clypeus gray. Eye rounded, about as wide as high, height 

two or three times the height of gena. Gena high, height subequal 

or slightly greater than length of basal flagellomere; gena-to-eye 

ratio 0.28–0.35.

Thorax: Mesonotum mostly brown to grayish brown 

with yellowish-gray areas, usually along setal tracks, 3 

well-defined brown stripes between dorsocentral rows; postpronotum and supra-alar areas silvery gray; scutellum brown 

basally, yellowish gray laterally and in middle, with apex 

blackish brown. Pleurae silvery gray, anepisternum with a 

small, rounded brown spot on dorsal margin and ventral margin brown tinged. Wing hyaline; veins brown; crossvein dm–

cu lacking infuscate halo; costal-vein ratio 0.40–0.46; M-vein 

ratio 1.06–1.12. Legs dark brown with sparse gray microtomentum; tarsi lighter with some yellow coloration, especially 

ventrally; forefemur of male with ventral surface concave near 

apex, anteroventral surface with a row of very long flattened 

setae on distal 3⁄4 of forefemur, posteroventral surface with 1 

row of very long and spaced setae, longer than width of femur; 

foretibia of male with ventral surface wavy, covered by conspicuous, dense, elongate setae.

Abdomen: Tergite 1 brown with a slender, small band 

on each side of anterior margin; tergites 2–5 distinctly bicolored, tergite 2 brown on anterior half and silvery gray on the 

remaining, tergites 3–5 brown along anterior 2⁄3 and medially, 

otherwise silvery gray. Male terminalia: (Figures 102–105) 

Presurstylus bearing short, slender, sparse setae, widened basally, basoventral process not distinctly developed, horizontal 

process slightly curved; in lateral view postsurstylus widened 

medially with ventral margin rounded, bearing dense, elongate, 

slender setae and an elongated, pointed anteromedial process; 

lateral aedeagal process shortened, about 1⁄3 length of aedeagus, 

thin, slightly curved in middle, rounded apex; aedeagus rather 

robust tapering gradually toward membranous apex; in lateral 

view phallapodeme triangular; hypandrium moderately deeply 

concave; cerci long, rather deeply concave on distal half of internal margin.

Type Material. The holotype male of Paralimna 

plumbiceps is labeled “♂/JAMAICA/HoloTYPE 6090 [red; 

“6090” handwritten]/TYPE Paralimna PLUMBICEPS 

E.T.Cresson,Jr. [maroon; “Paralimna PLUMBICEPS” handwritten].” The holotype is double mounted (minuten pin in a block 

of cork), is in good condition (left basal flagellomere missing; 

abdomen not dissected), and is deposited in the ANSP (6090). 

Three paratypes, including the allotype (1♂, 2♀; ANSP), bear 

the same label data as the holotype.

Other Specimens Examined: WEST INDIES. 

DOMINICAN REPUBLIC. El Seibo: Pedro Sáchez (18°51.4′N, 

69°6.5′W), 26 May 1998, D. and W. N. Mathis (3♂; USNM); 

Rincón (near; 18°45.3′N, 68°55.7′W), 12 May 1995, W. N. 

Mathis (2♂; USNM).

JAMAICA. Portland: Port Antonio (18°10.5′N, 76°27′W), 

Apr 1891 (3♂; ANSP, USNM). St. Andrew: Kingston (18°01′N, 

76°47.6′W), Apr (1♀; USNM).

PUERTO RICO. Adjuntas (18°09.8′N, 66°43.2′W), 8–13 

Jun 1915 (6♂, 3♀; AMNH, ANSP).

Type Locality. Neotropical. “Jamaica.”

Distribution. (Map 12). Neotropical: West Indies 

(Dominican Republic, Jamaica, Puerto Rico).
7 2 • smithsonian contributions to zoology

Remarks. Paralimna plumbiceps is easily distinguished from congeners by the sinuous ventral surface of the 

male foretibia that bears conspicuous, dense, and elongate setae. 

The male terminalia also provides useful characters for the 

recognition of this species. The postsurstylus has a conspicuously 

elongated and pointed posteromedial lobe, the lateral aedeagal 

process is short, less than 1⁄2 the aedeagal length, and the horizontal process of the presurstylus is widened basally and gradually narrowed toward the apex (similar to P. sera but lacking the 

long, slender ventral setae).

24. Paralimna (Paralimna) puncticornis Cresson

Figures 106–109, 189, Map 20

Paralimna puncticornis Cresson 1916:121 [Costa Rica. Cartago: Peralta 

(9°58.1′N, 83°36.9′W); HT ♀, ANSP (6095)]; 1918:47 [review; Costa 

Rica].

Paralimna (Paralimna) puncticornis.—Cresson 1947:52–53 [review, Neotropical species].—Wirth 1968:15 [Neotropical catalog].—Mathis and 

Zatwarnicki 1995:124 [world catalog].

FIGURES 102–105. Structures of male terminalia of Paralimna (Paralimna) plumbiceps Cresson (Jamaica): (102) epandrium, 

cerci, and presurstyli, posterior aspect; (103) postsurstylus, lateral aspect; (104) aedeagus, phallapodeme, and hypandrium, 

lateral aspect; (105) same, ventral aspect. Scale bar = 0.1 mm. (See Figures 2–5 for labels of structures.)
number 643 • 7 3

Diagnosis. This species is distinguished from congeners by the following characters: face yellowish to golden gray 

with golden-brown lateral margin and 3 brown spots, 1 on carina and 2 longitudinal parallel also rectangular large spots between largest facial setae, occupying almost the entire width of 

face and reaching the ventral margin; pedicel with a conspicuous 

silvery-gray spot dorsally; pleurae mostly gray or yellowish gray, 

katepisternum with darkened area anteriorly and anepisternum with 3 brown, well-defined spots—1 dorsal, 1 ventral, and 

1 rounded, central spot in middle, sometimes the central spot 

is inconspicuous or the spots expand and become completely 

brown anepisternum; forefemur of the male bearing posteroventral row of widely spaced and elongate setae and anteroventral 

row of sparse setae, elongate on basal half and shortened and 

slightly flattened on distal half of forefemur, setae of anterior 

row less than half length of setae of posterior row; crossvein 

dm–cu with faintly infuscate halo.

Description. Body length 6.5–8.2 mm; body generally bicolored, dark brown dorsally and brown ventrally, lateral 

medial surfaces mostly gray.

Head: (See Figure 189) Frons reddish brown with 

frontal design well-defined; linear gray or yellowish-gray spots 

at lateral margins of ocellar triangle, in front of anterior ocellar seta, converging anteriorly, reaching the anterior margin 

of frons, between inner vertical and interfrontal setae, and on 

fronto-orbits above dorsal proclinate fronto-orbital seta, between proclinate fronto-orbital setae, and just above antennal 

bases; ventral fronto-orbits and dorsal parafacial with a dark 

gray spot lateral to antenna, separated by gray spot. Antennal scape and pedicel dark brown, pedicel with a conspicuous 

silvery-gray spot dorsally; basal flagellomere pale brown to 

brown according to incidence of light, with slender, elongate, 

pale setae on margin; arista with 12–15 long, dorsal rays. Postcranium with 2 complete series of postocular setae. Face yellowish to golden gray with golden-brown lateral margin and 

3 brown spots, 1 on the carina, same length of antenna, and 

2 longitudinal, parallel, rectangular large spots between largest facial setae, occupying almost the entire width of face and 

reaching the ventral margin; parafacial and gena gray; gena 

with only few setae on middle and ventral margin; clypeus 

brownish gray with silvery-gray distal margin. Eye rounded, 

about as wide as high, height more than three times the height 

of gena. Gena high, subequal to length of basal flagellomere; 

gena-to-eye ratio 0.26–0.28.

Thorax: Mesonotum predominantly brownish copper with small gray or yellowish gray areas or stripes along 

the setal tracks; postpronotum, notopleuron, and supra-alar 

areas yellowish gray; scutellum brown dorsally with small gray 

spots, blackish brown apex, and gray laterally. Pleurae gray or 

yellowish gray except anepisternum with 3 brown, well-delimited spots—1 dorsal, 1 ventral, and 1 rounded spot in middle, 

sometimes the central spot is inconspicuous or the spots expand and become completely brown anepisternum; anepimeron 

brown on anteroventral corner and katepisternum with a darkened area anteriorly. Wing semi-hyaline, crossvein dm–cu with 

faintly infuscate halo; veins brown; costal-vein ratio 0.38–0.45; 

M-vein ratio 0.98–1.19. Legs blackish brown with sparse silvery-gray microtomentum; tarsi yellow; forefemur of the male 

with posteroventral row of widely spaced and elongate setae, 

and anteroventral surface with a row of sparse setae, elongate 

on basal half of forefemur, shortened and slightly flattened on 

distal half of the forefemur; foretibia straight.

Abdomen: Tergites distinctly bicolored, tergite 1 dark 

brown medially and gray on lateral corners; tergites 2–5 dark 

brown along anterior 2⁄3 and medially, otherwise yellowish 

gray; lateroventral margin of tergites brown. Male terminalia: 

(Figures 106–109) Presurstylus not bifurcate, horizontal process 

long, almost straight, with 4 or more strong and long setae inserted on tubercle; in lateral view postsurstylus widest medially 

with wide, medial, anteriorly produced lobe, bearing several stout, 

elongate setae extended through medial region of external surface, 

distal half of postsurstylus also slender; lateral aedeagal process 

robust, about 1⁄2 length of aedeagus, slightly curved subapically, 

bluntly rounded; aedeagus gradually tapered toward membranous apex in posterior view; in lateral view phallapodeme wider 

on basal 2⁄3 and slender on the remainder; hypandrium shallowly 

concave; cerci elongate, about 4× longer than wide.

Type Material. The holotype female of Paralimna 

puncticornis is labeled “Peralta[,] 24III [19]10C[osta]R[ica][,] 

P P Calvert/Sweeping over muddy road/TYPE Paralimna PUNCTICORNIS E.T.Cresson,Jr. [maroon; “Paralimna PUNCTICORNIS” handwritten].” The holotype is double mounted 

(minuten pin in a vertical rectangular card), is in very good 

condition (left wing with a small tear in anal vein region, some 

setae disoriented; abdomen not dissected), and is deposited in the 

ANSP (6095).

Other Specimens Examined. BOLIVIA. La Paz:

El Choro (16°19′S, 68°03.5′W; 900 m), 7 Jan 1976, L. E. Peña 

(1♂; CNC).

COSTA RICA. Alajuela: Higuito, San Mateo (09°56.7’N, 

84°32.8’W), P. Schild (1♂, 1♀; ANSP); Peñas Blancas (10°04′N, 

84°37′W; 700 m; rain forest), 18 Aug 1986, L. Masner (1♀; 

CNC). Cartago: Turrialba (09°54.1′N, 83°41.1′W; light trap), 

Mar–4 Sep 1954, 1986, C. H. Batcherder, L. Masner (2♂, 4♀; 

CNC, USNM); Turrialba (09°54.1′N, 83°41.1′W), 15 Jul 1965, 

P. J. Spangler (5♂; USNM); La Suiza (09°51.5′N, 83°37.5′W), 

6 May 1926, P. Schild, A. L. Melander (1♂; USNM) (1♂, 

3♀; ANSP). Heredia: La Selva Field Station near Puerto Viejo 

(10°27.2′N, 84°0.2′W), Feb–28 Mar 1980, 1988, W. Mason, 

W. E. Steiner (3♂, 2♀; CNC, USNM); La Selva (10°27.2′N, 

84°0.2′W; trap in garden), Jan 1993, P. Hanson (125♂; 108♀; 

USNM); Santo Domingo (INBio Parque; 09°58.4′N, 84°5.6′W), 

14 Jun 2003, D. and W. N. Mathis ( 10♂, 12♀: USNM). 

Limón: Bribri (4 km E; 10°00′N, 83°15′W; 50 m; Malaise 

trap), Jul–Sep 1990, P. Hanson (14♀; USNM). Puntarenas:

Manuel Antonio National Park (09°23.5′N, 84°08.7′W; coastal 

rainforest), 23–28 Aug 1986, L. Masner (1♂; CNC); Puerto 

Jiminez (5 km N; 10°13′N, 83°43′W; Malaise trap), Nov–

Dec 1990, P. Hanson (6♀; USNM); Piedras Blancas (24 km 

W; 08°47′N, 83°15′W; 200 m), Dec 1990 (1♂, 10♀; USNM); 
7 4 • smithsonian contributions to zoology

Rincón (3 km SW; 08°42.5′N, 83°29.2′W), Oct–Dec 1990, P. 

Hanson (4♂, 12♀; USNM); Res. For. Golfo Dulce, Rincón (3 

km SW; 08°42.5′N, 83°29.2′W), Dec 1989–Mar 1990, P. Hanson (8♂; USNM); San Pedrillo (08°37.2′N, 83°44.1′W), 12–14 

Aug 2001, D. and W. N. Mathis (1♂, 4♀; USNM); San Vito, 

Las Cruces (08°49.3′N, 82°58.3′W; 1200 m), 9 Jul 1983, B. 

Gill (1♀; CNC). San José: Ciudad Colon (09°55′N, 84°15′W; 

800 m; Malaise trap), Dec 1989, P. Hanson (4♂, 2♀; USNM); 

Pedregoso (09°22′N, 83°43′W), D. L. Rounds (3♂, 1♀; ANSP); 

Río Paraíso (09°33.8′N, 84°7.4′W; 350-400 m), 15–17 Feb 

2003, D. and W. N. Mathis (3♂; 6♀; USNM).

ECUADOR. Loja: Loja (S; 03°59.4′S, 79°12.3′W; 1200 

m), 26 Mar 1965, L. E. Peña (1♂, 1♀; CNC). Los Ríos: Río 

Palenque (0°35′S, 79°22′W; 150 m), 22–26 Feb 1976, G. E. 

Shewell (1♀; CNC). Orellana: Coca, Napo River (0°27.8′S, 

76°59′W; 250 m), 12–13 Apr 1965, L. E. Peña (12♂, 17♀; 

FIGURES 106–109. Structures of male terminalia of Paralimna (Paralimna) puncticornis Cresson (Costa Rica. Herédia): 

(106) epandrium, cerci, and presurstyli, posterior aspect; (107) postsurstylus, lateral aspect; (108) aedeagus, phallapodeme, 

and hypandrium, lateral aspect; (109) same, ventral aspect. Scale bar = 0.1 mm. (See Figures 2–5 for labels of structures.)
number 643 • 7 5

CNC). Zamora-Chinchipe: Río Jumboé (04°04′S, 78°55.8′W; 

1200 m), 1–2 Apr 1965, L. E. Peña (1♂, 1♀; CNC).

HONDURAS. Colon: Corocito (15°46.4′N, 85°47.4′W), 3 

Apr 1924 (3♀; ANSP).

PANAMÁ. Chiriqui: El Valle (08°25.9′N, 82°19.8′W), 

Nov 1943, N. L. H. Krauss (1♂; USNM). Guna Yala: San Blas, 

Rio Carti Grande near coast (09°27.3′N, 78°57.8′W), 2 Mar 

1985, O. S. Flint Jr., J. E. Louton (1♂, 1♀; USNM). Panamá:

Barro Colorado Island (09°09.1′N, 79°50.8′W, Canal Zone), 

21 Dec 1928, C. H. Curran (1♀; ANSP); Barro Colorado Island 

(09°09.1′N, 79°50.8′W; Malaise trap), Jul 1967, W. W. Wirth 

(1♂, 1♀; USNM); Chiva Chiva, Road 8, 8 air km N Ft. Clayton 

(09°02′N, 79°35′W), 23 Jul 1978, N. E. Woodley (1♂; USNM); 

Gamboa, Pipeline Road (09°07′N, 79°42′W; Malaise trap), Jul 

1967, W. W. Wirth (4♀; USNM). Veraguas: Corozal (08°59′N, 

79°34.1′W; Canal Zone), 16 Jan 1929, C. H. Curran (1♂; ANSP).

PERU. Cuzco: Quincemil (13°13.7′S, 70°45.6′W; 740 m), 

13–31 Aug 1962, L. E. Peña (2♂, 2♀; CNC). Madre de Dios:

Avispas (12°58.7′S, 70°20.8′W; 400 m), 1–30 Oct 1962, L. E. 

Peña (1♂, 2♀; CNC).

TRINIDAD and TOBAGO. Tobago. St. John: Charlotteville (5 km S; 11°18.9′N, 60°34.5′W), Hermitage River 

and beach, 22 Apr–11 Jun 1993, 1994, D. and W. N. Mathis 

(1♂; USNM). Trinidad. St. Andrew: Valencia (1 km W; Aripo 

River; 10°39′N, 61°13′W), 20 Jun 1993, W. N. Mathis (3♂, 

1♀; USNM). St. George: Arima (8 km N; 10°41′N, 61°18′W), 

Verdant Vale (10°41′N, 61°17′W), 19 Jun 1993, W. N. Mathis 

(1♂; USNM); Caura River Valley (end of Caura River Road; 

10°42.5′N, 61°21.5′W), 31 Mar 1996, C. Chaboo (10♂, 7♀; 

AMNH).

VENEZUELA. Bolívar: Gran Sabana (N edge at Pioneer 

Monument; 08°16′N, 62°45.3′W), 21 Mar 1982, G. F. and 

J. F. Hevel (1♂; USNM). Lara: Yacambú (10°0.7′N, 69°14.6′W; 

1200 m), 10 May 1981, H. K. Townes (1♂, 4♀; CNC).

Type Locality. Neotropical. Costa Rica. Cartago: 

Peralta (9°58.1′N, 83°36.9′W).

Distribution. (Map 20) Neotropical: Bolivia (La 

Paz), Costa Rica (Alajuela, Cartago, Heredia, Limón, Puntarenas, San José), Ecuador (Loja, Los Ríos, Orellana, ZamoraChinchipe), Honduras (Colon), Panamá (Chiriqui, Guna Yala, 

MAP 20. Distribution map for Paralimna (Paralimna) puncticornis Cresson.
7 6 • smithsonian contributions to zoology

Panamá, Veraguas), Peru (Cuzco, Madre de Dios), Trinidad and 

Tobago, Venezuela (Bolívar, Lara).

Remarks. This species has facial coloration that is 

similar to P. crinita sp. nov.; however, it can be easily distinguished by characters of the male terminalia: the basoventral 

process of the presurstylus is falciform and the lateral aedeagal 

process is large in P. crinita, and the basoventral process is not 

developed and the lateral aedeagal process is normally developed in P. puncticornis. The legs also differ: the male forefemur 

has the ventral surface distinctly concave on its distal 1⁄3, and 

the male foretibia bears dense and bristling setae on the ventral 

surface in P. crinita; the ventral surface of the male forefemur is 

not concave and the male foretibia lacks dense and bristling setae 

on the ventral surface in P. puncticornis. Some specimens of P. 

molossus have large golden-brown spots on the ventral half of 

the face that can be confused with P. puncticornis, but features 

of the male terminalia are dissimilar.

25. Paralimna (Paralimna) punctipennis

(Wiedemann)

Figures 110–113, 190, 206, Map 21

Notiphila punctipennis Wiedemann 1830:590.

Paralimna punctipennis.—Cresson 1929:184 [generic combination]; 

1946:229 [review].—Wirth 1965:748 [Nearctic catalog].—Mathis and 

Zatwarnicki 1995:124 [world catalog].

Paralimna appendiculata Loew 1862a:138 [United States. “Middle 

States” and “Georgia”; ST ♂♀, MCZ (11134)].—Cresson 1929:184 

[synonymy].

Diagnosis. This species is distinguished from congeners by the following combination of characters: face silvery 

gray to yellowish gray with 3 golden-brown spots, a rectangular 

spot on the carina, and 2 small, rounded spots between largest 

facial setae; pleural area distinctly lighter on dorsal half (dorsal 

pleurites gray, only thin band on ventral margin of anepisternum 

and anepimeron brown) and darker on ventral half (ventral pleurites blackish brown); wing hyaline, second section of vein M 

with 1 to several perpendicular stumps, the stumps and remaining transversal veins infuscate; forefemur of male bearing simple 

setae on anteroventral and posteroventral surface.

Description. Body length 3.1–5.7 mm; body generally bicolored, brown dorsally and blackish brown ventrally, 

lateral medial surfaces mostly pale to yellowish gray.

Head: (See Figure 190) Frons mottled, dark to whitish 

brown with almost inconspicuous pale yellow spots at lateral margins of ocellar triangle, in front of anterior ocellar seta, not reaching 

the anterior margin of frons, between inner vertical and interfrontal seta, and on fronto-orbits between proclinate fronto-orbital 

setae and just above antennal bases; ventral fronto-orbits without 

conspicuous spot; dorsal parafacial with a gray spot lateral to antenna. Antennal scape and pedicel brown, pedicel paler on distal 

margin dorsally; basal flagellomere pale brown to brown according 

to incidence of light, with slender, elongate, pale setae on margin; 

arista with 11–13 long, dorsal rays. Postcranium with 2 complete 

series of postocular setae. Face silvery gray to yellowish gray with 3 

golden-brown spots, a rectangular spot on the carina, shorter than 

the length of antenna, and 2 small, rounded spots between largest 

facial setae; parafacial and gena silvery gray; gena with only few 

setae on middle and ventral margin; clypeus silvery to yellowish 

gray. Eye rounded, slightly higher than wide, height about twice 

the height of gena. Gena high, height subequal or slightly greater 

than length of basal flagellomere; gena-to-eye ratio 0.42–0.57.

Thorax: Mesonotum mostly grayish brown to brown, 

lateral surfaces mostly gray, sometimes silvery white, 3 brown 

stripes between dorsocentral rows; postpronotum, notopleuron, 

and supra-alar areas silvery gray; scutellum brown dorsally, with 

apex blackish brown and pale gray laterally. Pleural area distinctly 

lighter on dorsal half (dorsal pleurites gray, only thin band on 

ventral margin of anepisternum and anepimeron brown) and 

darker on ventral half (ventral pleurites blackish brown). Wing 

(see Figure 206) hyaline; veins pale brown; second section of vein 

M with 1 to several perpendicular stumps, the stumps and remaining transversal veins infuscate; costal-vein ratio 0.43–0.47; M-vein 

ratio 0.95–1.05. Legs brown with brown microtomentum; tarsi 

lighter with some yellow coloration, especially ventrally on first 

and second tarsomeres; forefemur of male with anteroventral and 

ventral series of short setae, ventral series more numerous, flattened setae absent; foretibia straight.

Abdomen: Tergites distinctly bicolored, dark brown 

along anterior 2⁄3 and medially, otherwise silvery to yellowish gray 

and lateral margins brown, except tergite 1 wholly brown. Male 

terminalia: (Figures 110–113) Presurstylus with basoventral process weakly developed, short, wide, length about 1⁄3 horizontal 

process; horizontal process long, slender with long, slender setae; 

in lateral view postsurstylus widest medially with wide, medial, 

anteriorly produced lobe bearing several stout, elongate setae extended until half the medial region on external surface, a stout 

medial short lobe posteriorly produced, distal half of postsurstylus 

large, widened in lateral view; lateral aedeagal process constricted 

in middle, about 1⁄2 length of aedeagus, curved subapically, bluntly 

rounded; aedeagus slender in posterior view; phallapodeme 

broadly triangular in lateral view; hypandrium deeply concave; 

cerci widened, almost 2× longer than wide.

Type Material. The lectotype female of Notiphila 

punctipennis Wiedemann (designated by Cresson 1929:184) is labeled “punctipennis [handwritten] Alte Sammlung/TYPE Notiphila punctipennis [“Notiphila punctipennis” handwritten; red].” 

The lectotype is pinned directly, is in good condition (mesonotum 

cracked where pin is inserted; abdomen not dissected), and is deposited in NMW.

Other Specimens Examined. UNITED STATES. 

Alabama. Lauderdale: Shoal Creek near Killen (34°51.8′N, 

87°32.3′W; 150 m), 8 Aug 1939, J. A. G. Rehn, J. W. H. Rehn 

(1♀; ANSP).

Arizona. Cochise: Douglas (31°20.9′N, 109°32.6′W), Aug, 

F. H. Snow (1♀; ANSP); San Bernardino Ranch (31°20.2′N, 

109°16.8′W), Aug, F. H. Snow (1♀; ANSP).
number 643 • 7 7

Colorado. Baca: Regnier (37°N, 102°50′W), 6–9 Jun (1♂; 

ANSP). Boulder: Longmont, Ute Creek (40°10′N, 105°06′W), 

7 Aug, R. W. Dawson (1♀; ANSP). Crowley: Olney Springs 

(38°09.8′N, 103°56.4′W), 28 May 1932, M. T. James. El Paso:

Colorado Springs (38°50′N, 104°49′W; 1570 m), Jul, E. S. 

Tucker (1♀; ANSP). Weld: Roggoo (40°10.1′N, 104°22.3′W), 

31 Aug 1938, M. T. James (1♀; ANSP).

Delaware. Kent: Dover (39°09.5′N, 75°31.5′W), 24 

Jun 1939, A. L. Melander (1♂, 3♀; ANSP, USNM). Sussex:

Delaware Seashore State Park (38°40.6′N, 75°04.1′W; beach), 

17 May 2006, D. and W. N. Mathis (1♀; USNM); Dewey Beach 

(38°41.5′N, 75°05.5′W), 27 Aug 1972, L. V. Knutson (3♂, 4♀; 

USNM); Lewes Angola Neck Park (38°40.3′N, 75°09.1′W), 21 

Sep 2007, G. A. Foster, A. M. Welch (1♀; USNM); Rehoboth 

(38°43.2′N, 75°04.6′W), 25 Jun–1 Sep 1939, 1977, A. L. Melander, W. W. Wirth (2♂, 8♀; ANSP, USNM).

District of Columbia. Anacostia (38°52′N, 76°59′W), 4 Jun 

1913, P. R. Myers (1♂; USNM); Rock Creek, Boundary Bridge 

FIGURES 110–113. Structures of male terminalia of Paralimna (Paralimna) punctipennis (Wiedemann) (United States. 

Virginia): (110) epandrium, cerci, and presurstyli, posterior aspect; (111) postsurstylus, lateral aspect; (112) aedeagus, 

phallapodeme, and hypandrium, lateral aspect; (113) same, ventral aspect. Scale bar = 0.1 mm. (See Figures 2–5 for 

labels of structures.)
7 8 • smithsonian contributions to zoology

(38°59.2′N, 77°03.2′W), 18 May 2007, D. and W. N. Mathis 

(1♂, 2♀; USNM); Rock Creek, Milkhouse Ford (38°57.9′N, 

77°02.9′W), 18 May 2007, D. and W. N. Mathis (4♂, 2♀; 

NPSRC, USNM); Rock Creek, Picnic Area 28 (38°56.7′N, 

77°03′W; 35 m), 19 May 2007, D. and W. N. Mathis (4♂, 2♀; 

USNM); Washington (38°49.8′N, 77°00.6′W), 17 Aug 1913, 

D. W. Coquillett (2♂; ANSP, USNM).

Florida. Alachua: Austin Cary Forest (29°44.6′N, 

82°12.8′W), 15 Oct 1976, G. F. Fairchild, H. V. Weems Jr. (1♂, 

3♀; FSCA); Gainesville (29°39.1′N, 82°19.5′W), 4 May–3 Oct 

1932, 1960, 1973, H. Howden, H. V. Weems Jr., R. E. Woodruff 

(2♂, 9♀; CNC, FSCA); Newmans Lake (29°39′N, 82°13′W), 27 

Feb 1948, E. D. McRae (♂, ♀; FSCA). Collier: Collier-Seminole 

State Park (28°58.5′N, 81°36.2′W), 8 Dec 1977, J. E. Rawlins 

(1♀; CMP). Duval: Jacksonville (30°17.1′N, 81°37.8′W), A. T. 

Slosson (1♀; FSCA). Escambia: Bratt (30°58′N, 87°26′W), 

1 Apr–19 Jun 1938, 1974, A. Blanton, H. V. Weems Jr. (2♀; 

FSCA, USNM). Highlands: Archbold Biological Station 

(27°10.8′N, 81°21.1′W), 11 Jul–12 Dec 1959, 1978, 1979, S. W. 

Frost, T. A. Weber, H. V. Weems Jr. (5♂, 6♀; USNM); Archbold 

Biological Station (27°11.3′N, 81°20.3′W), 2 Jan–5 Dec 1985, 

1988, 1989, 1995, 1999, M. A. Deyrup, S. Lenberger (22♂, 

35♀; ABSF); Archbold Biological Station (8 km W; 27°10.9′N, 

81°28′W), 15 Apr 1989, D. and W. N. Mathis (4♂, 6♀; USNM); 

Rte. 70, cattle pond (27°12.5′N, 81°23.3′W), 7 Feb 2000, D. 

and W. N. Mathis (1♂; USNM). Jackson: Florida Caverns State 

Park (30°48′N, 85°13′W), 14 Apr–26 May 1962, 1973, H. V. 

Weems Jr., W. W. Wirth (4♀; FSCA, USNM). Lake: Altoona (11 

km E; Clearwater Lake; 28°58.7′N, 81°33.2′W), 6 May 1986, 

W. E. Steiner (1♀; USNM). Levy: (29°25′N, 82°54′W), 8 Feb 

1953, H. V. Weems Jr. (2♂, 3♀; FSCA); Cedar Key (27 km NE; 

29°27.8′N, 82°59.2′W), 25 May 1970, D. L. Bailey (1♂, 2♀; 

USNM); (29°17.5′N, 82°45′W), 11 Aug 1953, H. V. Weems (1♂; 

USNM). Liberty: Torreya State Park (30°34.1′N, 84°56.9′W), 22 

Apr 1967, W. W. Wirth (1♂, 1♀; USNM). Marion: Salt Springs 

(29°21′N, 81°44.1′W), 14 Apr 1989, D. and W. N. Mathis (5♂, 

4♀; USNM); Salt Springs (8 km N; 29°29.7′N, 81°44.1′W), 

14 Apr 1989, D. and W. N. Mathis (1♂, 2♀; USNM); Silver 

Springs (29°13′N, 82°03.5′W), 12 Mar–2 Apr 1932, 1942, A. L. 

Melander (15♂, 12♀; ANSP, USNM). Okalossa: Holt (7.2 km 

N; 30°50′N, 86°45′W), 17 Jun 1978, L. A. Stange (1♂, 2♀; 

FSCA). Orange: Orlando (28°32.3′N, 81°22.8′W), 2 Feb–26 Jun 

1932, A. L. Melander, M. R. Wheeler (1♂, 2♀; USNM); Rock 

Springs (28°45.4′N, 81°30.1′W), 25 Feb–21 Apr 1942, 1970, 

A. L. Melander, W. W. Wirth (3♀; USNM). Osceola: Kissimmee (28°17.5′N, 81°24.5′W), 1 Feb 1932, A. L. Melander (1♀; 

USNM). Palm Beach: Lake Worth (26°37′N, 80°03.4′W; light 

trap), 9 Mar 1951, W. W. Wirth (1♂; USNM). Polk: Lakeland 

(28°02.4′N, 81°57′W), 6 May 1916, J. C. Bradley (1♂; USNM). 

Santa Rosa: Blackwater River State Park (30°42′N, 86°51.6′W), 

23 Jun 1973, W. W. Wirth (2♂, 6♀; USNM); Lake Carr (6.5 km 

W; 30°46.6′N, 87°14.2′W), 22 Jun 1974, C. R. Arnaud, H. V. 

Weems Jr. (3♂, 3♀; FSCA).

Georgia. Charlton: Billys Island (30°48.3′N, 82°20.4′W), 

Jun 1912 (2♀; ANSP). Cobb: Austell (33°48.8′N, 84°38.1′W), 

27 Aug 1910 (1♂; ANSP). Decatur: Spring Creek (30°51.3′N, 

84°35.1′W), 16–19 Jul 1912 (2♂, 3♀; ANSP). Gilmer: Alley 

(34°41.1′N, 84°28.6′W), 18 Aug 1938, D. G. Hall (3♂, 5♀; 

ANSP). Monroe: Forsyth (33°02.1′N, 83°56.3′W), 2 Jun 1970 

(1♂; CNC). Pierce: Blackshear (31°18.4′N, 82°14.5′W), 10 May 

1911 (1♂; ANSP). Rabun: Clayton (34°52.7′N, 83°24.1′W), 

18–26 May 1911, J. C. Bradley (1♀; ANSP).

Illinois. Jackson: Grand Tower (37°37.6′N, 89°29.9′W), 30 

Jun 1909 (1♀; ANSP); Murphysboro (37°45.9′N, 89°20.1′W), 

16 May 1910 (1♀; ANSP). Jo Daviess: Galena (42°25′N, 

90°25.7′W), 5 Jul 1917 (1♂; ANSP). Marion: Odin (38°37′N, 

89°03.1′W), 16 May 1910 (1♀; ANSP). Morgan: Meredosia 

(39°50′N, 90°33.6′W), 19 Aug 1917 (1♂; ANSP). Pulaski: Pulaski 

(37°12.8′N, 89°12.4′W), 28 Jun 1909 (1♀; ANSP). Union: Dongola (37°21.7′N, 89°10′W), 12 May 1917 (1♂, 1♀; ANSP). Washington: Du Bois (38°13.4′N, 89°12.7′W), 8 Aug 1917 (1♀; ANSP).

Indiana. Tippecanoe: Lafayette (40°25′N, 86°52.5′W), 27 

May–2 Aug 1915, J. M. Aldrich (2♂, 1♀; ANSP).

Kansas. Bourbon: (37°50′N, 94°50′W), 1915, R. H. Beamer 

(1♀; ANSP). Douglas: (38°54′N, 95°16′W), F. H. Snow (1♂, 

1♀; ANSP); Lawrence (38°58.3′N, 95°14.1′W), 9 Jun 1922, 

C. H. Curran (1♀; CNC). Ellsworth: (38°43.8′N, 98°13.4′W), 

12 Jul 1923, R. H. Beamer (1♂, 1♀; ANSP). Gove: (38°57′N, 

100°20′W; 857 m), F. X. Williams (1♂, 2♀; ANSP). Labette:

(37°12′N, 95°18′W), 1915, R. H. Beamer (1♀; ANSP). Miami:

(38°38′N, 94°50′W), 1915, R. H. Beamer (1♂; ANSP). Ottawa:

(38°35′N, 95°14′W), 24 Jun 1934, C. W. Sabrosky (2♂, 3♀; 

ANSP). Pottawatomie: Onaga (39°29.3′N, 96°10.2′W), 9 May 

1911 (1♂; ANSP). Riley: Manhattan (39°11′N, 96°34.3′W), 7 Jul 

1932, C. W. Sabrosky (1♀; ANSP). Reno: Medora (38°08.9′N, 

97°50.7′W), 4 Jun–Apr 1959, E. Becker, H. Howden (3♂, 1♀; 

CNC). Washington: Linn (39°40.8′N, 97°05′W), 1915, R. H. 

Beamer (1♀; ANSP).

Louisiana. Calcasieu: Lake Charles (5 mi [8 km] N; 

30°19.6′N, 93°13′W), 8 Dec 1933, C. W. Sabrosky (1♀; 

ANSP). De Soto: Logansport (31°58.5′N, 93°59.9′W), 24 Mar 

1908, E. S. Tucker (1♀; ANSP). Madison: Mound (32°20.4′N, 

91°01.4′W), 24 May 1907 (1♂; ANSP). St. Tammany: Sidell 

(30°16.5′N, 89°46.9′W), 2–6 Jul 1905 (1♂; ANSP).

Maryland. Anne Arundel: Edgewater (6 km S; 38°53′N, 

76°33′W; Smithsonian Environmental Research Center), 4 

Aug 1994, W. N. Mathis (1♂; USNM). Carroll: Eldersburg 

(39°24.2′N, 76°57′W), 2 Jun 1985, J. E. Lowry, W. E. Steiner 

(1♂; USNM). Calvert: Calvert Beach (38°27.9′N, 76°28.6′W), 

17 Sep 1989, J. M. Hill, W. E. Steiner, J. M. Swearingen (2♂, 

4♀; USNM). Cecil: Octoraro Creek (39°39.7′N, 76°08.8′W), 

17 Jun 1939, E. G. Fisher (1♀; ANSP). Garrett: Broadford Lake 

(39°24.7′N, 79°22.4′W; 750 m), 24 Aug 2006, D. and W. N. 

Mathis (4♂, 1♀; USNM); Savage River (39°30.1′N, 79°06.9′W; 

400 m), 23 Aug 2006, D. and W. N. Mathis (1♀; USNM); Savage River Reservoir (near Floyd; 39°30.7′N, 79°09.3′W; 420 

m), 23 Aug 2006, D. and W. N. Mathis (1♀; USNM). Howard:

Clarksville (39°12.4′N, 76°56.6′W), 12 Sep 1981, N. E. Woodley (1♀; USNM). Montgomery: Ashton (6.5 km SW; 39°06.5′N, 

77°01.5′W), 27 Apr–31 Oct 2000, 2001, 2002, G. F. Hevel (5♂, 
number 643 • 7 9

12♀; USNM); Bethesda (38°58.8′N, 77°06′W), 2 Aug–12 Sep 

1964, 1981, G. C. Steyskal, N. E. Woodley (3♂, 1♀; USNM); 

Cabin John (38°58.5′N, 77°09.5′W), 25 Jul 1972, G. C. Steyskal (3♂, 3♀; USNM); Clarksburg, Little Bennett Regional Park 

(39°16′N, 77°16.8′W), 9 Jul–21 Sep 1989, 1990, M. J. and 

R. Mollineaux, W. E. Steiner (15♂, 15♀; USNM); Colesville 

(39°04.8′N, 77°00.1′W), 14 Jun–31 Jul 1975, W. W. Wirth (6♂, 

11♀; USNM); Cropley (38°58.8′N, 77°13.2′W), 12 Jul 1970, L. 

V. Knutson (2♂; USNM); Forest Glen (39°00.9′N, 77°03.2′W), 

20 Aug 1966, W. W. Wirth (1♀; USNM); Pennyfield Lock, 

C&O Canal (39°03.2′N, 77°17.3′W), 5 Jul 1953, A. Stone (1♀; 

USNM); Plummers Island (38°58.2′N, 77°10.6′W), 20 May–17 

Aug 1914, 1988, 1989, A. Freidberg, F. K. Knab, R. C. Shannon, P. J. Spangler (3♂, 4♀; USNM); Takoma Park (38°58.7′N, 

77°00.4′W), 4 Aug 1987, W. E. Steiner (1♀; USNM); Wheaton (39°02.3′N, 77°03.3′W), 26 Jul 1979, A. Freidberg (8♂, 

7♀; USNM). Prince George′s: Adelphi (NW Branch Creek; 

39°00.2′N, 76°58.3′W), 27 Nov 1981, E. R. S. Hodges (8♂, 

3♀; USNM); Beltsville (39°02.1′N, 76°54.5′W), 15 Jun 1913, 

W. L. McAtee (2♂, 5♀; USNM); Bladensburg (38°56.4′N, 

76°56′W), 23 Sep–2 Oct 1914, 1915, R. C. Shannon (1♂, 5♀; 

USNM); Camp Springs (38°48.2′N, 76°54.4′W), 18 Jun 1969, 

G. F. Hevel (1♀; USNM); Cheverly (38°56′N, 76°55′W), 4–22 

Aug 1995, W. E. Steiner, J. M. Swearingen (2♂; USNM); College Park (38°58.8′N, 76°56.2′W), 25 May–5 Sep 1913, 1914, 

1977, C. T. Greene, F. K. Knab, W. N. Mathis (8♂, 7♀; USNM); 

Hyattsville (38°57.4′N, 76°56.7′W), 1 Sep 1912, J. R. Malloch 

(1♂, 1♀; ANSP, USNM); Oxon Hill (38°48.2′N, 76°59.4′W), 

30 Jun–22 Jul 1971, 1978, G. F. Hevel (2♂, 3♀; USNM); 

Patuxent Wildlife Research Center (39°03′N, 76°48.2′W), 14 

Jul 1978, W. W. Wirth (1♀; USNM). Talbot: McDaniel (Wades 

Point; 38°49.6′N, 76°17.9′W), 21 Sep 1985, W. E. Steiner (1♂, 

1♀; USNM).

Michigan. Allegan: Saugatuck (42°39.3′N, 86°12.1′W), 25 

Jun 1938, C. W. Sabrosky (1♂; ANSP). Livingston: Hamburg 

(42°26.9′N, 83°49.1′W), 21 May 1939, G. C. Steyskal (1♀; 

ANSP). Midland: Midland (43°36.8′N, 84°14.9′W), 6–7 Jun 

1936, C. W. Sabrosky (1♂, 1♀; ANSP).

Mississippi. Hinds: Jackson (32°17.9′N, 90°11.1′W), 30 

Oct 1944 (2♂; ANSP); Utica (32°06.6′N, 90°37.4′W), Aug (1♂, 

1♀; ANSP). Lafayette: Oxford (34°22′N, 89°31.1′W), Jun 1943 

(1♀; ANSP). Lowndes: Artesia (33°24.9′N, 88°38.6′W), 1 May 

1921, A. McIntosh (1♂; ANSP). Oktbbeha: Agriculture College (State College; 33°27.1′N, 88°47′W), 10 Mar–25 Sep 1921, 

1928, H. W. Allen, J. M. Langston, W. Stafford (1♂, 4♀; ANSP).

Missouri. Boone: Columbia (38°57.1′N, 92°20′W; Malaise 

trap), 26 May–5 Sep 1906, 1968, C. R. Crosby, F. D. Parker (6♂, 

21♀; ANSP, USNM). Greene: Springfield (24 km W; 37°11.7′N, 

93°26.6′W), 15–21 Jun 1982, S. E. and P. M. Miller (4♀; 

USNM); Willard (37°18.3′N, 93°25.7′W), 30 Mar (1♂; USNM). 

Howard: Boonslick Salt Spring (39°03.4′N, 92°52.5′W), 17 May 

1969, W. W. Wirth (1♂; USNM). Jackson: Atherton (39°11.2′N, 

94°18.3′W), Jul–Sep (3♂, 4♀; ANSP).

Nebraska. Dodge: Fremont (41°26′N, 96°29.9′W), 18 Jul 

1900 (1♂; ANSP).

New Jersey. Burlington: Riverton (40°0.1′N, 75°0.9′W), 19 

Sep 1909 (1♀; ANSP). Camden: Ashland (39°51.8′N, 75°0.4′W), 

13 May 1906 (1♂; ANSP). Cape May: Cape May (38°56.1′N, 

74°54.4′W), 22 Jul–24 Aug 1936, 1938, W. Stone (1♂, 1♀; 

ANSP). Mercer: Trenton (40°13′N, 74°44.6′W), 19–23 Aug 

1909 (4♂, 3♀; ANSP). Ocean: Tuckerton (16 km N; 39°53.3′N, 

74°22.8′W), 26 Sep 2003, D. and W. N. Mathis (2♂; USNM).

New Mexico. Otero: High Rolls (32°57′N, 105°50.1′W), 31 

May–2 Jun 1902 (2♂, 4♀; ANSP).

North Carolina. Columbus: Lake Waccamaw (34°18.5′N, 

78°29.7′W), 6 Jul 1985, A. Gerberich, W. Steiner (3♀; USNM). 

Dare: Kill Devil Hills (36°01.8′N, 75°40.6′W), 23 Jun 1954, K. V. 

Krombein (1♀; USNM). Graham: Lake Santeetlah (35°29.8′N, 

83°52′W; 460 m), 6 Aug 1939, J. A. G. Rehn, J. W. H. Rehn 

(1♀; ANSP). Jones: Kinston (25 km SE; 35°08.1′N, 77°28.2′W), 

23 Apr 1988, M. J. Mollineaux (1♀; USNM). Onslow (34°39′N, 

77°28′W): 25 Jun 1964, J. R. Cornell (1♂; USNM). Swain: Clingmans Dome, Great Smoky Mountains National Park (35°33.7′N, 

83°30′W), 5 Aug 1939 (1♀; ANSP). Wake: Crabtree Creek (3.2 

km NW Raleigh; 35°49.1′N, 78°37.8′W), 23 Apr 1964, G. A. Matuza (3♂, 7♀; USNM); Raleigh (35°46.3′N, 78°38.3′W), 14 Oct 

1971, P. W. Oman (1♂, 1♀; USNM). Watauga: Pisgah National 

Forest, Mortimer Recreational Area (35°25.3′N, 81°48.6′W; 460 

m), 25 May 1999, S. D. Gaimari (1♂, 3♀; USNM).

Ohio. Butler: Four-Mile Creek, near Oxford (39°28′N, 

84°36.2′W), 26 May 1979, B. A. Steinly (1♂, 1♀; USNM); Indian Creek C. P. near Oxford (39°28.5′N, 84°47.6′W), 25 May 

1976, B. A. Steinly (1♀; USNM). Erie: Sandusky (41°27′N, 

82°42.5′W), Jul 1913 (1♂; ANSP). Franklin: Columbus 

(39°57′N, 83°0.4′W), 23 May (1♀; ANSP). Fulton: Wauseon 

(41°32′N, 84°08.5′W), 23-25 Aug 1902 (3♂, 2♀; ANSP). Muskingum: Zanesville (39°56.4′N, 82°0.8′W), 24 Aug 1952, C. W. 

Sabrosky (1♂; USNM).

Oklahoma. Cherokee: Scraper (1.6 km NE; 36°0.3′N, 

94°51.9′W), 7 May 1974, G. F. and J. F. Hevel (1♂; USNM). 

Mayes: Spavinaw Recreation Area (36°23.5′N, 95°02.8′W), 

9 May 1984, G. F. and J. F. Hevel (5♂, 7♀; USNM). Woods:

Alabaster Caverns State Park, near Freedom (36°47.5′N, 

95°58.4′W), 11 May 1984, G. F. and J. F. Hevel (1♀; USNM).

Pennsylvania. Delaware: Swarthmore (39°54.1′N, 

75°21′W), 6 Jun–26 Sep 1905, 1908, 1909, E. T. Cresson 

Jr. (10♂, 24♀; ANSP). Montgomery: Lansdale (40°14.5′N, 

75°17′W), 30 May 1909, E. T. Cresson Jr. (1♀; ANSP); Narberth (3.2 km N; 40°0.5′N, 75°15.6′W), 9 Sep 1915, E. T. 

Cresson Jr. (1♀; ANSP). Pennsylvania: Fern Rock (40°02.6′N, 

75°08.4′W), 29 Jun 1905 (1♂; ANSP); Germantown (40°01.7′N, 

75°10.5′W), 12 Jun 1906 (1♀; ANSP); Manayunk (40°01.8′N, 

75°13.3′W) (1♂; ANSP).

South Carolina. Aiken: Kitchings Mill (4 km S; 33°33.2′N, 

81°28′W), 8 May 1986, W. E. Steiner (1♀; USNM). Berkeley:

Jamestown (33°17.1′N, 79°41.4′W), 2 Jul 1953, M. R. Wheeler 

(1♂; USNM). Charleston: Isle of Palms (32°48′N, 79°45′W), 

23 Jun 1957, G. C. Steyskal (1♀; USNM). Clarendon: Manning (33°41.7′N, 80°12.6′W), 29–30 May 1914, W. Stone (1♀; 

ANSP). Spartanburg: Cross Anchor (34°38.6′N, 81°51.5′W), 
8 0 • smithsonian contributions to zoology

3 Jul 1953, M. R. Wheeler (1♂; USNM); Spartanburg (34°57′N, 

81°56′W), 8 May 1961 (1♂; USNM).

South Dakota. Bennett: Martin (43°10.4′N, 101°44′W), 6 

Jul 1924 (2♂, 2♀; ANSP). Canton: Newton Hills (43°13.6′N, 

96°35′W), 24 Jun 1935, H. C. Severin (1♀; ANSP). Brookings:

Brookings (44°18.7′N, 96°48′W), 4 Jun 1928, H. C. Severin 

(1♀; ANSP). Tripp: Winner (43°22.6′N, 99°51.5′W), 3–6 Jul 

1924 (1♂, 3♀; ANSP). Union: Elk Point (42°40′N, 96°41′W), 

12 Jun 1925, H. C. Severin (1♀; ANSP).

Tennessee. Granger: Rutledge (36°16.5′N, 81°31.3′W), 

Jul 1954, M. R. Wheeler (1♀; USNM). Humphreys: Buffalo 

(34°43.5′N, 81°41′W), 19 May 1979, G. C. Steyskal (1♂; 

USNM). Shelby: Memphis (35°09′N, 90°03′W), 23 Sep 1924 

(1♀; ANSP). Wilson: Lebanon (36°12.5′N, 86°17.5′W), 9 Jun 

1946, G. C. Steyskal (1♂; USNM).

Texas. Comal: New Braunfels, Comal River (29°42.2′N, 

98°06.8′W), 24 Mar 1942, A. L. Melander (2♀; USNM). Fort 

Bend: Sugar Land (29°37.2′N, 95°38.1′W), 11 Dec 1933, C. 

W. Sabrosky (1♀; ANSP). Galveston: Dickinson (29°27.6′N, 

95°03.1′W), Jun 1929 (2♂, 3♀; ANSP). Grayson: Denison 

(33°45.3′N, 96°32.2′W), 22 Jun 1904, H. S. Barber (1♀; ANSP). 

Liberty: Liberty (30°03.5′N, 94°47.7′W), 18 Mar 1908, E. S. 

Tucker (2♀; ANSP). McLennon: Waco (31°33′N, 97°08.8′W), 

G. W. Belfrage (1♂; ANSP). San Augustine: San Augustine 

(31°31.8′N, 94°06.4′W), 13 Oct 1895 (1♀; ANSP). Travis: Austin (30°16′N, 97°44.6′W), Oct 1900 (1♂; ANSP).

Virginia. Accomack: Assateague Island, near Refuge headquarters (37°54.5′N, 75°21.6′W), 19 Sep–3 Oct 2005, 2007, 

D. and W. N. Mathis (10♂, 3♀; USNM); Assateague Island, 

wildlife loop (37°54.6′N, 75°21.1′W), 15 Jun–20 Sep 2007, D. 

and W. N. Mathis (1♂, 2♀; USNM); Chincoteague (37°55.4′N, 

75°21.2′W), 15 Jun–20 Sep 2007, D. and W. N. Mathis (3♂, 

1♀; USNM). Arlington: Arlington (38°52′N, 77°05.9′W), Jun 

1938, J. R. Malloch (1♂; USNM); 4-Mile Run (38°50.4′N, 

77°02.7′W), 17 Jun–24 Sep 1976, 1977, W. N. Mathis, H. B. Williams (17♂, 36♀; USNM); Glencarlyn (38°51.8′N, 77°7.8′W), 

17 Jun 1914, 1925, N. Banks, J. R. Malloch, W. L. McAtee 

(2♂; ANSP, USNM); Glencarlyn Park, 4-Mile Run (38°51.5′N, 

77°07.5′W), 20 Aug 1977, H. B. Williams (3♂, 1♀; USNM); 

Rosslyn (38°53.8′N, 77°4.3′W), 15 May 1913, R. C. Shannon 

(1♂, 2♀; ANSP, USNM). Chesterfield: Pocahontas State Park 

(37°23.1′N, 77°32.4′W), 11 May 2002, D. and W. N. Mathis 

(13♂; USNM). Essex: Tappahannock (37°55.8′N, 76°51.4′W; 

Rappahannock River), 18 Sep 2004, D. and W. N. Mathis 

(4♂, 1♀; USNM). Fairfax: Dead Run (38°58′N, 77°10.3′W), 

29 Jun 1915, R. C. Shannon (1♀; USNM); Dead Run (swamp; 

38°57.8′N, 77°10.3′W), 4 May 2006, D. and W. N. Mathis 

(4♂, 3♀; USNM); Dyke Marsh (38°46.1′N, 77°02.9′W), 13 Aug 

1978, W. N. Mathis (1♂, 2♀; USNM); Great Falls (38°59.9′N, 

77°15.3′W), 26 Jun–9 Jul 1913, 1920, A. L. Melander, R. C. 

Shannon (3♂, 2♀; ANSP, USNM); Great Falls (Clay Pond; 

39°00.1′N, 77°15.4′W), 12 May–17 Aug 2006, 2007, D. and 

W. N. Mathis, F. C. and B. J. Thompson (5♂, 1♀; USNM); Great 

Falls (Patowmack Canal; 39°00.1′N, 77°15.2′W), 27 Jun–25 

Jul 2006, 2007, 2008, D. and W. N. Mathis (6♂, 1♀; USNM); 

Great Falls (Potomac River; 39°0.2′N, 77°15.2′W), 4 Oct 2007, 

D. and W. N. Mathis (1♂; USNM); Great Falls (swamp trail; 

38°59.4′N, 77°15.2′W), 12 May 2006, D. and W. N. Mathis 

(3♂, 1♀; USNM); Holmes Run (38°50.9′N, 77°10.4′W; at light), 

4 Jul–6 Sep 1961, W. W. Wirth (1♂, 3♀; USNM); Lake Barcroft (38°50.9′N, 77°09.4′W), 28 May 1977, W. N. Mathis (7♂, 

8♀; USNM); Turkey Run (38°57.8′N, 77°09.4′W), 4 May–10 

Jul 2006, 2008, D. and W. N. Mathis (3♂, 1♀; USNM); Turkey Run (mouth; 38°57.9′N, 77°09.4′W), 22 May–17 Sep 2006, 

2007, 2008, D. and W. N. Mathis, H. B. Williams (26♂, 5♀; 

USNM). Giles: Mountain Lake (37°22.5′N, 80°31.5′W; 260 

m), 26 May 1962, J. G. Chillcott (2♂, 1♀; CNC). Henry: Martinsville Reservoir (36°44.7′N, 79°52.2′W), 17 May 2005, D. 

and W. N. Mathis (5♂, 2♀; USNM); Martinsville, Smith River 

(36°39.9′N, 79°53′W; 225 m), 18 May 2005, D. and W. N. 

Mathis (2♂; USNM). Isle of Wight: Zuni (6 km S at Blackwater 

River; 36°51.9′N, 76°45.9′W), 14–15 Jun 1989, W. E. Steiner 

(1♀; USNM). King George: Dahlgren Wayside (38°21.6′N, 

77°00.9′W; Potomac River), 19 May–8 Jun 2006, 2007, D. and 

W. N. Mathis (7♂, 1♀; USNM). Lancaster: Belle Isle State Park 

(Humphreys Picnic area; 37°46.4′N, 76°35.6′W; Rappahannock 

River), 28 Apr 2007, D. and W. N. Mathis (1♂, 1♀; USNM); 

Belle Isle State Park (Watch House; 37°46.4′N, 76°36.1′W; Rappahannock River), 28 Apr 2007, D. and W. N. Mathis (1♀; 

USNM). Louisa: Lake Anna (38°05.8′N, 77°53.6′W; Christopher Run Campground), 4 May 2007, D. and W. N. Mathis (6♂; 

USNM). Middlesex: Dragon Run (37°38′N, 76°41.7′W), 17 Sep 

2004, D. and W. N. Mathis (10♂, 3♀; USNM). Montgomery:

Blacksburg (37°13.8′N, 80°24.8′W; 640 m), 28 May 1962, 

J. G. Chillcott (1♀; CNC). Northumberland: Vir-Mar Beach 

(37°55.8′N, 76°18.5′W), 18 Sep 2004, D. and W. N. Mathis (4♂, 

1♀; USNM). Patrick: Meadows of Dan (36°44.2′N, 80°22.9′W), 

18 May 2005, D. and W. N. Mathis (1♂, 1♀; USNM); Woolwine (36°47.4′N, 80°16.7′W; 300 m), 17 May 2005, D. and 

W. N. Mathis (2♂, 2♀; USNM). Prince William: Prince William 

Forest Park (38°34′N, 77°22′W; Carter Pond), 13 Aug 1993, D. 

and W. N. Mathis (6♂, 4♀; USNM); Prince William Forest Park, 

South Quantico Creek (38°34′N, 77°22′W; shoreline), 10–30 Jul 

1993, D. and W. N. Mathis (16♂, 15♀; USNM). Rappahannock: Hazel River (NW Culpeper; 38°33.8′N, 78°11.6′W, 171 

m), 28 Jun–24 Jul 2008, D. and W. N. Mathis, T. Zatwarnicki 

(8♂, 1♀; USNM). Richmond: Warsaw (37°57.4′N, 76°45.6′W), 

27 Aug 2004, D. and W. N. Mathis (7♂, 3♀; USNM). Roanoke: Salem (Roanoke River; 37°16.1′N, 80°02.2′W; 300 m), 

23 Sep 2007, D. and W. N. Mathis (4♂; USNM). Smyth: Saltville (N at N Fork Holston River; 36°32.9′N, 82°36.7′W), 25 

Sep 1981, O. S. Flint Jr. (7♂, 8♀; USNM); Saltville (36°52.3′N, 

81°46.4′W; 521 m), 24 Sep 2007, D. and W. N. Mathis (2♂; 

USNM). Spotsylvania: Motts Reservoir (38°19′N, 77°33.5′W), 

15 Apr 2006, D. and W. N. Mathis (3♂, 4♀; USNM); Rappahannock River (38°18.8′N, 77°32.5′W), 14 Aug–10 Oct 2006, 

D. and W. N. Mathis (6♂; USNM). Stafford: Aquia Creek 

(38°29.1′N, 77°23.8′W), 6 Jun 2005, D. and W. N. Mathis (3♂; 

USNM); Aquia Harbour (38°27.7′N, 77°23.3′W), 15 May–21 

Jul 1988, 2000, D. and W. N. Mathis (14♂, 3♀; USNM); Aquia 
number 643 • 8 1

MAP 21. Distribution map for Paralimna (Paralimna) punctipennis (Wiedemann).

Harbour, Aquia Creek (38°27.8′N, 77°23.1′W), 2 Sep 2006, 

D. and W. N. Mathis (3♂; USNM); Aquia Harbour, Lions Park 

(38°27′N, 77°23.3′W), 3 May–16 Nov 2003, 2004, 2005, 2006, 

2007, 2008, D. and W. N. Mathis (45♂, 6♀; USNM); Aquia 

Landing (38°23.2′N, 77°19′W), 7 May–5 Sep 2005, D. and W. N. 

Mathis (7♂, 1♀; USNM); Falmouth (38°19.2′N, 77°28.1′W; 

Rappahannock River; 9 m), 18 Apr–30 Jun 2007, 2008, D. and 

W. N. Mathis (4♂, 3♀; USNM). Westmoreland: Westmoreland State Park (38°09.7′N, 76°51.9′W), 16 Sep 1994, D. and 

W. N. Mathis (3♂; USNM). York: Maury Lake (near James River; 

37°02.5′N, 76°29.2′W), 19 Aug 2006, D. and W. N. Mathis 

(6♂, 3♀; USNM). Independent Cities: Alexandria (38°48.3′N, 

77°02.8′W), 14 Jun 1952, C. W. Sabrosky (1♂; USNM). Falls 

Church (38°52.9′N, 77°10.3′W), 4 May–3 Aug 1914, 1954, 

1968, N. Banks, G. M. Greene, R. C. Shannon, W. W. Wirth 

(16♂, 14♀; ANSP, MCZ, USNM). Fredericksburg (Alum Park; 

38°17.4′N, 77°28.9′W), 30 Apr 2007, D. and W. N. Mathis 

(5♂, 1♀; USNM). Norfolk, Hickory (10 km SSE; 36°50.8′N, 

76°17.1′W; Northwest River Park), 24–25 Aug 1985, D. Bogar, 

W. E. Steiner (1♀; USNM). Norfolk, Ocean View (36°56.6′N, 

76°14.5′W), 9 Aug, A. N. Candell (3♀; USNM). Richmond 

(37°32.6′N, 77°26.1′W), 24 Oct 1915, A. H. Sturtevant (2♂; 

USNM). Virginia Beach (36°55.9′N, 76°01.2′W), 14 Aug 1913, 

F. K. Knab (2♂, 8♀; ANSP, USNM).

West Virginia. Cavell: Culloden (38°25.2′N, 82°03.3′W), 

Jul 1954, M. R. Wheeler (1♀; USNM). Greenbrier: Charmco 

(38°00.1′N, 80°44.2′W), 6 Sep 1982, G. F. and J. F. Hevel (3♀; 

USNM). McDowell: Roderfield (37°26.7′N, 81°42.2′W; 335 m), 

25 Sep 2007, D. and W. N. Mathis (4♂; USNM). Mercer: Ceres 

(Kee Reservoir; 37°18.4′N, 81°10.4′W; 757 m), 24 Sep 2007, 

D. and W. N. Mathis (3♂; USNM). Pocahontas: Marlinton 

(Greenbriar River; 38°13.5′N, 80°05.7′W), 21 Jun 2007, D. and 

W. N. Mathis (3♂; USNM); Tea Creek (Right Fork; 38°20′N, 

80°9.9′W), 29 Jul 1982, O. S. Flint Jr., W. N. Mathis (1♂; 

USNM). Randolph: Cheat Mountain (38°36.5′N, 79°56.5′W), 

A. H. Sturtevant (1♀; USNM). Summers: Bluestone State Park 

(37°36.7′N, 80°56.1′W; 440 m), 26 Sep 2007, D. and W. N. 

Mathis (3♂; USNM); Hinton (37°41.8′N, 80°53′W; New River; 

427 m), 26 Sep 2007, D. and W. N. Mathis (2♂; USNM). Wyoming: R. D. Bailey Lake (37°35.7′N, 81°46.8′W; 324 m), 25 Sep 

2007, D. and W. N. Mathis (3♂; USNM).

Type Locality. Nearctic: “United States.”

Distribution. (Map 21) Nearctic: United States 

(Alabama, Arizona, Colorado, Delaware, District of Columbia, 
8 2 • smithsonian contributions to zoology

Florida, Georgia, Illinois, Indiana, Kansas, Louisiana, Maryland, Michigan, Mississippi, Missouri, Nebraska, New Jersey, 

New Mexico, North Carolina, Ohio, Oklahoma, Pennsylvania, 

South Carolina, South Dakota, Tennessee, Texas, Virginia, West 

Virginia).

Remarks. This species is similar to P. pectinata and 

P. fellerae in having the coloration of mesopleuron distinctly bicolored, silvery or yellowish gray on the dorsal half and brown 

on the ventral half. This species can be distinguished from these 

two species by the presence of small stump veins on vein R1

 (absent in P. pectinata and P. fellerae), coloration of the face, and 

shape of male forefemur and foretibia. In P. punctipennis, the 

face has 3 spots (1 in P. pectinata), the forefemur of males lacks 

flattened setulae (flattened setulae present in P. pectinata and P. 

fellerae), and the male forefemur and foretibia are simple (concave along ventral surface in P. pectinata). We discuss the distinguishing characters of the male terminalia of these species in the 

“Remarks” for P. pectinata.

26. Paralimna (Paralimna) sana Cresson

Figures 114–117, 191, Map 22

Paralimna sana Cresson 1929:185 [Paraguay. San Bernardino (25°16′S, 

57°19.4′W); HT ♂, ANSP (6338)].—Wirth 1968:15 [Neotropical catalog].—Mathis and Zatwarnicki 1995:125 [world catalog].

Paralimna secunda sana Cresson 1947:48 [review, Neotropical species].

Paralimna cilifera Hendel 1930:130 [Argentina. Aguarai; Bolivia. El 

Cairo, Santa Cruz de la Sierra; ST ♂♀, SMN].—Cresson 1947:53–54 

[list, Neotropical species].—Lizarralde de Grosso 1989:38 [list, Argentina].—Mathis and Zatwarnicki 1995:119 [world catalog]. NEW

SYNONYM.

Diagnosis. This species is distinguished from congeners by the following characters: face silvery gray with 1 goldenbrown, elongate, rather triangular spot on the carina, as long as the 

length of antenna, sometimes with 2 small inconspicuous, rounded 

pale brown spots between largest facial setae; anepisternum with 

dorsal and ventral margins golden brown tinged and bases of 

setae brown; forefemur of male with anteroventral row of elongate setae, as long as half the width of femur, all the same length 

and slightly flattened; posteroventral row shortened near apex of 

femur, shorter than anteroventral setae and arranged in a series.

Description. Body length 3.0–3.5 mm; body generally bicolored, brown dorsally, lateral surfaces gray, silvery, or 

yellowish gray.

Head: (See Figure 191) Frons pale brown to brown with 

linear, pale yellow spots at lateral margins of ocellar triangle, in 

front of anterior ocellar seta, not reaching the anterior margin of 

frons, between inner vertical and interfrontal setae, and on frontoorbits above dorsal proclinate fronto-orbital seta, between proclinate fronto-orbital setae, and just above antennal bases; ventral 

fronto-orbits with a dark brown spot and dorsal parafacial with 

a dark gray spot lateral to antenna, separated by silvery-gray spot. 

Antennal scape and pedicel dark brown, pedicel with silvery-gray 

margin; basal flagellomere pale brown to brown according to incidence of light, with slender, elongate, pale setae on margin; arista 

with 9–12 long, dorsal rays. Postcranium with 1 complete series or 

2 irregular series of postocular setae. Face silvery gray, sometimes 

dark gray, with 1 golden-brown elongate, rather triangular, spot 

on the carina, as long as the length of antenna, sometimes with 2 

small, inconspicuous, rounded pale brown spots between largest 

facial setae; parafacial and gena silvery gray; gena with only few 

setae on middle and ventral margin; clypeus silvery gray. Eye round, 

as wide as high, height about twice the height of gena. Gena high, 

height longer than length of basal flagellomere; gena-to-eye ratio 

0.36–0.50.

Thorax: Mesonotum mostly golden brown with silvery 

or yellowish gray areas, usually along setal tracks, 2 well-defined, 

wide brown stripes between dorsocentral rows and a acrostichal 

irregular stripe; postpronotum, notopleuron, and supra-alar areas 

silvery gray; scutellum golden brown, with apex blackish brown 

and silvery gray laterally. Pleurae silvery to yellowish gray, anepisternum with dorsal and ventral margins golden brown tinged and 

bases of setae brown. Wing hyaline; veins brown; transverse veins 

infuscate; costal-vein ratio 0.46–0.47; M-vein ratio 0.95–1.33. 

Legs brown with sparse silvery-gray microtomentum; tarsi lighter 

with some yellow coloration, darkened on distal tarsomeres; forefemur of male with anteroventral row of elongate setae, as long as 

half the width of femur, all the same length and slightly flattened; 

posteroventral row shortened near apex of femur, shorter than 

anteroventral setae and arranged in a series; foretibia straight.

Abdomen: Tergite 1 wholly brown; tergites 2–5 distinctly 

bicolored, dark brown along anterior 2⁄3 and medially, otherwise 

yellowish gray dorsally and silvery gray laterally. Male terminalia: 

(Figures 114–117) Presurstylus not bifurcate, horizontal process 

wide basally and gradually tapering toward apex, with sparse slender setae; in lateral view postsurstylus widest medially with wide, 

medial, anteriorly produced lobe bearing several slender, elongate 

setae extended until half of medial region on external surface, a 

thin medial lobe posteriorly produced; lateral aedeagal process 

about 1⁄2 length of aedeagus, almost straight, bluntly rounded; aedeagus slender in posterior view; phallapodeme broadly triangular 

in lateral view; hypandrium shallowly concave.

Type Material. The holotype male of Paralimna 

sana is labeled “[K.] Fiebrig[,] Paraguay[,] S. Bernardino/TYPE 

Paralimna SANA [“Paralimna SANA” handwritten] E. T. Cresson, Jr. 6338 [maroon].” The holotype is double mounted (minuten pin in a foam block), is in excellent condition (abdomen 

not dissected), and is deposited in the ANSP (6338).

The lectotype male of Paralimna cilifera Hendel, designated herein, is labeled “Aguarai arg. VI.26.Lind. D.ChacoExped [black margin]/Paralimna cilifera H. [handwritten] F. 

Hendel det./Type. Hendel 1930 [handwritten; red ink; label 

with a black border]/SMN/LECTOTYPE ♂ Paralimna cilifera

Hendel, designated by Ale-Rocha&Mathis [red].” The lectotype 

is double mounted (minuten pin in a rectangular card), is in 

excellent condition (abdomen not dissected), and is deposited 
number 643 • 8 3

in the SMN. In the original description, Hendel (1930:130) 

wrote that the syntype series comprised three specimens: two 

(1♂, 1♀) from Aguarai, Argentina, and one (1♂) from El Cairo, 

Bolivia. Our study of the type series indicates that the two specimens from Argentina are conspecific and that the third from 

Bolivia is another (P. molossus). The female from Aguarai is a 

paralectotype.

Other Specimens Examined. ARGENTINA. 

Catamarca: El Pintado, S La Viña (28°03.3′S, 65°35.1′W; 

650 m), 27–28 Sep 1968, L. E. Peña (18♂, 17♀; CNC). Jujuy:

Ing. Ledesma, near Gen. San Martin (23°50.1′S, 64°47.5′W; 

1100 m), 10 Oct 1968, L. E. Peña (1♂, 2♀; CNC). Misiones:

Rompland (27°28.5′S, 55°21′W), 13–14 Jan 1927, F. and M. 

Edwards (1♂, 1♀; ANSP). Salta: Escoipe (38 km SW Salta; 

25°10′S, 65°46′W; 1900 m), 8 Oct 1968, L. E. Peña (3♀; CNC). 

Tucumán: Arroyo (14 km S El Tala; 26°56.4′S, 65°09.7′W; 

700 m), 13–14 Oct 1968, L. E. Peña (4♂; CNC); San Miguel 

de Tucumán (26°48.5′S, 65°13′W), 18 Apr–4 May 1913, 1914, 

T. C. Barber, A. H. Rosenfeld (3♂; ANSP, USNM); San Pedro de 

Colalaeo (26°14′S, 65°29′W), 14 Oct 1968, L. E. Peña (3♂, 1♀; 

FIGURES 114–117. Structures of male terminalia of Paralimna (Paralimna) sana Cresson (Argentina. Missiones): (114) epandrium, cerci, and presurstyli, posterior aspect; (115) postsurstylus, lateral aspect; (116) aedeagus, phallapodeme, and hypandrium, 

lateral aspect; (117) same, ventral aspect. Scale bar = 0.1 mm. (See Figures 2–5 for labels of structures.)
8 4 • smithsonian contributions to zoology

MAP 22. Distribution map for Paralimna (Paralimna) sana Cresson.

CNC); Tucumán (30 km N; 26°20.2′S, 65°16.9′W; 700 m), 15 

Oct 1968, L. E. Peña (2♂; CNC).

BOLIVIA. Santa Cruz: Campo Guairay (24 km S Camini; 

20°11.8′S, 63°28.6′W; 870 m) 2 Sep 2000, S. D. Gaimari (13♂, 

8♀; USNM).

BRAZIL. Paraná: Santo Antonio da Platina, Fazenda Dora 

(23°17.9′S, 50°04.1′W), Jun 1965, N. Papavero (1♀; MZUSP). 

São Paulo: Araçatuba, Corrego Azul (21°05′S, 50°32′W), Mar 

1947, M. P. Barreto (1♂; MZUSP).

PARAGUAY. Asunción: Asunción (25°16.9′S, 57°38.1′W), 

7 Oct. 1980, D. C. Lowrie (1♂, 1♀; USNM); 10–12 Oct 

1980 (1♂, 1♀; USNM). Cordillera: San Bernardino (25°16′S, 

57°19.4′W), K. Fiebrig (paratype 1♀; ANSP; 1♂; USNM). Pirapó: Villarica (26°51.3′S, 55°32.6′W), Sep 1931, F. Schade (1♂; 

USNM).

Type Locality. Neotropical. Paraguay. Cordillera: 

San Bernardino (25°16′S, 57°19.4′W).

Distribution. (Map 22) Neotropical: Argentina 

(Catamarca, Jujuy, Misiones, Salta, Tucumán), Bolivia (Santa 

Cruz), Brazil (Paraná, São Paulo), Paraguay (Asunción, Cordillera, 

Pirapó).

Remarks. Cresson (1947) suggested that this species 

is a southern South American subspecies of P. secunda. However, the posteroventral setae of the male forefemur are sparse 

and arranged in a single series and the wing has infuscate halos 

on crossvein dm–cu in P. sana (the posteroventral setae are numerous and usually arranged in more than 1 series, and the wing 

lacks an infuscate halo on crossvein dm–cu in P. secunda). Additionally, the features of male terminalia are dissimilar in these 

two species, such as the shape of the presurstylus. The pattern 

of coloration of the face and body is not useful to distinguish 

between these species due to variation in both.

27. Paralimna (Paralimna) secunda Schiner

Figures 118–121, 192, Map 23

Paralimna secunda Schiner 1868:241.

Paralimna (Paralimna) secunda.—Cresson 1947:47–48 [review, Neotropical 

species].—Wirth 1968:15 [Neotropical catalog].—Mathis and Zatwarnicki 1995:125 [world catalog].
number 643 • 8 5

Diagnosis. This species is distinguished from congeners by the following combination of characters: face gray or 

yellowish gray with 3 brown spots, 1 elongate spot on carina and 

usually 2 spots of variable shapes between largest facial setae, 

rarely ventral spots are absent; pleurae gray with dorsal and ventral margins of anepisternum brown and with dotted aspect in 

the middle by small brown spots on insertion of setae; forefemur 

with conspicuous but not flattened setae on anteroventral surface, ventral and posteroventral surface with numerous robust 

setae; presurstylus lacking basoventral process, horizontal process broad; cercus fused at the apex of dorsal margin of horizontal process; lateral aedeagal process long and slender.

Description. Body length 3.6–5.5 mm; body generally bicolored, brown dorsally, lateral surfaces mostly gray.

Head: (See Figure 192) Frons mostly pale brown with 

linear, silvery-white spots at lateral margins of ocellar triangle 

and weakly extended to anterior margin of the frons, between 

inner vertical and reclinate fronto-orbital setae, and on margin 

of eye from dorsal proclinate fronto-orbital to just above antennal bases; margin of the frons with a black spot lateral to 

antenna and dorsal parafacial with a dark gray spot. Region of 

insertion of all setae blackish. Antenna with gray microtomentum on scape; pedicel with gray microtomentum on margin 

and ventrally, dorsum dark gray to blackish microtomentum, 

with 1 dorsal, thicker seta and some short setae on the margin; basal flagellomere densely covered by silvery-white microtomentum and slender, elongate, pale setae on margin; arista 

with 13 long, dorsal rays. Ocellar pair inserted at the same 

line of anterior ocellus. Postcranium with 2 complete series of 

postocular setae. Face, parafacial, and gena gray or yellowish 

gray, face with brown spots: 1 elongate spot on carina, as wide 

as the distance between lateral bases of antennae and as long 

as length of scape and pedicel together, and usually 2 spots 

of variable shapes between largest facial setae, rarely ventral 

spots are absent; gena almost nude, few setae only on ventral 

margin and in middle; clypeus densely silvery gray or darker 

gray according to incidence of light. Eye rounded, about as 

wide as high, height twice the height of gena. Gena high, height 

subequal the length of basal flagellomere; gena-to-eye ratio 

0.36–0.48.

Thorax: Mesonotum generally brown, the same color of 

the frons, with gray stripes, 2 acrostichal and 1 dorsocentral; lateral of scutum predominantly gray with brown spots; scutellum 

mostly gray with apex and base brown to blackish brown. Pleurae 

gray, dorsal and ventral margins of anepisternum brown and with 

dotted aspect in the middle by small brown spots on insertion of 

setae; anepimeron usually without spots, wholly gray. Wing: hyaline, lacking infuscation; costal-vein ratio 0.38–0.44; M-vein ratio 

0.80–1.06. Legs: coxae, trochanters, femora, and tibiae brown 

with gray microtomentum; tarsomeres yellow. Forefemur with 

conspicuous setae on anteroventral surface, but the setae are not 

flattened, ventral and posteroventral surface with numerous robust setae; midfemur with strong anteroventral setae on base of 

distal 1⁄3.

Abdomen: Tergites 3–4 distinctly bicolored, dark 

brown on anterior half and medially, remaining portion gray; 

tergite 1 gray only in sides; tergite 2 predominantly gray with 

brown lateral spot basally and a longitudinal median spot basally, not extended to apical margin of segment. Male terminalia: 

(Figures 118–121) Presurstylus not bifurcate, horizontal process 

broad, concave ventrally, cercus fused at the apex of dorsal margin of horizontal process; in lateral view postsurstylus weakly widest medially with wide, medial, anteriorly produced lobe bearing 

several slender elongate setae extended until the half of medial 

region on external surface, a small medial lobe posteriorly produced; lateral aedeagal process slender, longer than 1⁄2 of length 

of aedeagus, almost straight, bluntly rounded; aedeagus slender in 

posterior view; phallapodeme broadly triangular in lateral view; 

hypandrium deeply concave.

Type Material. The lectotype female of Paralimna 

secunda Schiner (designated by Cresson 1929:185) is labeled 

“Lindig 1864 Venezuela/secunda [handwritten] Alte Sammlung/

Paralimna secunda Schi[ner]. [handwritten]/TYPE Paralimna secunda [red; species name handwritten by E. T. Cresson, Jr.].” 

There is also a female paralectotype bearing the same locality 

data as the lectotype. The lectotype is double mounted (cut pin 

in a rectangular card), is in good condition, and is deposited in 

the NMW.

Other Specimens Examined. BELIZE. Stan 

Creek District: Sittee River, Possum Point Biological Station 

(16°52.1′N, 88°22.5′W), 22–29 Apr 1987, R. Faitoute, P. J. 

Spangler (1♂, 5♀; USNM); Kendal (16°47.4′N, 88°22.9′W), 1 

May 1987, R. Faitoute, P. J. Spangler (1♀;USNM); Maya Center 

Cabbage Haul Creek (16°48′ N, 88°22′ W), 3 Apr 1993, W. N. 

Mathis (3♂, 1♀; USNM); Silk Grass Creek (16°54′ N, 88°26′

W), 3 Apr 1993, W. N. Mathis (1♂, 1♀; USNM); Cockscomb 

Basin Wildlife Sanctuary (16°45′N, 88°30′W), 5–6 Apr 1993, 

W. N. Mathis (1♂; USNM); Dangriga (16°58′ N, 88°13′ W), 3–4 

Apr 1993, W. N. Mathis (2♂, 1♀; USNM).

BOLIVIA. Bení: Rurrenabaque (14°26.5′S, 67°31.7′W; 

175 m), 10–23 Oct 1956, L. E. Peña (1♂; CNC). La Paz: Apa 

Apa (8 km S Chulumani; 16°22′S, 67°30.4′W; 1960 m), 9–10 

Mar 2001, W. N. Mathis (5♂; 2♀; USNM); El Choro (16°19′S, 

68°03.5′W; 900 m), 7 Jan 1976, L. E. Peña (1♂; CNC); Puente 

Villa (16°28.9′S, 68°07.1′W; 1200 m), 12–20 Dec 1955, L. E. 

Peña (1♂; CNC); Puente Villa (2 km E; 16°24′S, 67°38′W; 1969 

m) 1 Mar 2001, W. N. Mathis (5♂, 3♀; USNM); Ganay (1 km E; 

15°30.2′S, 69°52.3′W; 580 m), 13 Feb 2001, S. D. Gaimari (2♂; 

USNM); Ganay (3 km E; 15°30.2′S, 67°52.3′W; 200 m), 14 Mar 

2001, W. N. Mathis (3♂, 1♀; USNM).

BRAZIL. Amazonas: Autaz Mirim, Fazenda S. Lucas 

(03°04.1′S, 59°57.1′W; Malaise trap), 16 Jul 1994, J. Vidal 

(1♀; INPA); Balawa-u (01°48.4′N, 63°47.1′W; Malaise trap), 

14 Sep 1995, L. S. Aquino (1♀; INPA); Igarapé Cabeça Branca 

(2°35.1′S, 60°01.9′W; 65 m), 8 May 2010, W. N. Mathis (1♂; 

USNM); Ipixuna, Rio Liberdade, Estirão da Preta (07°03.1′S, 

71°41.7′W; sweeping), 11–15 May 2011, J. A Rafael, J. T. Câmara, R. F. Silva, A Somavilla, R. Ale-Rocha (5♂, 5♀; INPA); 
8 6 • smithsonian contributions to zoology

Manaus, Reserva Biológica da Campina (02°30′S, 60°00′W; 

Malaise trap), 2–3 Dec 1981, J. A. Rafael, E. Binda (3♀; INPA); 

Manaus, Universidade Federal do Amazonas (Campus Universitário; 03°06′S, 59°58.5′W; Malaise trap), 7 Apr–28 Jul 1982, 

J. A. Rafael (5♂, 8♀; INPA); Manaus, INPA (03°55.8′S, 59°59.1′W, 

60m), 4 May 2010, D. and W. N. Mathis (2♂; USNM); Maués, 

Rio Abacaxis, Campina Pacamiri (04°35.8′S, 58°13.2′W; Malaise trap), 30–31 May 2008, J. A. Rafael et al. (1♀; INPA); 

Parque Nacional do Jaú, Rio Carabinani, Igarapé Ipiranga 

(02°03.6′S, 61°33.5′W; Malaise trap), 29–31 Jul 1995, J. A Rafael, J. Vidal (1♀; INPA); Reserva Ducke (26 km NE Manaus; 

02°55.8′S, 59°58.5′W; 40 m; Malaise trap), 9 Feb–16 Sep 1976, 

FIGURES 118–121. Structures of male terminalia of Paralimna (Paralimna) secunda Schiner (Venezuela. Territorio Federal 

Amazonas): (118) epandrium, cerci, and presurstyli, posterior aspect; (119) postsurstylus, lateral aspect; (120) aedeagus, phallapodeme, and hypandrium, lateral aspect; (121) same, ventral aspect. Scale bar = 0.1 mm. (See Figures 2–5 for labels of structures.)
number 643 • 8 7

1982, J. A. Rafael (7♂, 13♀; INPA); Rio Abacaxis, Flona Pau 

Rosa (01°15.2′S, 58°41.9′W; Malaise trap), 28–29 May 2008, 

J. A. Rafael et al. (1♀; INPA); Rio Gregório, Comunidade Lago 

Grande (07°10.2′S, 70°49.2′W; some from light and Malaise 

traps), 18–23 May 2011, J. A. Rafael et al. (2♂, 1♀; INPA); Rio 

Nhamundá (01°35.2′S, 57°37.5′W; 25 m; Malaise trap), 17–20 

May 2008, J. A. Rafael et al. (4♂, 12♀; INPA); Rio Nhamundá, 

Igarape Areias (01°35.2′S, 57°37.6′W; 25 m; Malaise trap), 

17–20 May 2008, J. A. Rafael et al. (6♂, 9♀; INPA); Santa 

Izabel do Rio Negro, Maturacá (00°32.5′S, 66°08.2′W), 11–13 

May 1990 (1♂; INPA). Pará: Belém (01°27.3′S, 48°30.1′W), 

Sep 1952, Th. Dobzhansky (1♂; USNM); Castanhal (01°17.8′S, 

47°55.3′W), Aug 1952, Th. Dobzhansky (1♂; USNM); Óbidos, 

Sítio Curió (01°47.1′S, 55°07.1′W; Malaise trap), 29 Aug–8 Sep 

2001, J. A. Rafael, J. Vidal (6♂, 5♀; INPA); Óbidos, Fazenda 

Pajurá (01°37.4′S, 55°23.2′W; Malaise trap), 5–11 Sep 2001, 

J. A. Rafael, J. Vidal (2♀; INPA); Óbidos, Estrada OriximináÓbidos (01°38.7′S, 55°37′W), Nov 1969, Expedição Permanente 

da Amazônia (2♂, 1♀; MZUSP); Oriximiná, Rio Trombetas 

(01°27.9′S, 56°18.6′W), Oct 1969, Expedição Permanente da 

Amazônia (3♂, 1♀; MZUSP); Rio Iriri (~100 km S Altamira; 

03°50′S, 52°40′W), 17–18 Oct 1986, O. S. Flint, P. J. Spangler (2♂; USNM); Rio Xingu (camp, about 60 km S Altamira; 

03°39′S, 52°22′W; Igarapé Jabuti; Malaise trap), 8–16 Oct 

1986, O. S. Flint, P. J. Spangler (8♂, 9♀; USNM); Taperinha, 

Santarém (02°31.9′S, 54°17.7′W), Oct–Nov 1970, Expedição 

Permanente da Amazônia (1♂; MZUSP). São Paulo: São Paulo 

(23°33.6′S, 46°35.8′W), G. G. Barbreillini (6♂, 1♀; ANSP). Tocantins: Jalapão, Mateiros, Rio Novo (10°33.4′S, 46°35.6′W; 

400 m; light trap), 31 Mar 2008, J. A. Rafael, F. F. Xavier (12♂, 

8♀; INPA).

COLOMBIA. Magdalena: Aracataca (10°35.9′N, 

74°11.3′W), 24 Aug 1920 (5♂, 1♀; ANSP); Cali (10 km W Valle 

de Cauca; 03°26.3′N, 76°33.9′W; 1630 m), Mar 1955, E. S. 

Ross, E. I. Schlinger (2♂; USNM); Sevilla (17 km W; 10°46′N, 

74°10.5′W), Mar 1955, E. S. Ross, E. I. Schlinger (1♀; USNM).

COSTA RICA. Alajuela: Alajuela (10°01′N, 84°13′W; 945 

m), 15 Sep 1909, P. P. Calvert (1♂; ANSP); Higuito, San Mateo 

(09°56.7’N, 84°32.8’W), P. Schild (1♀; ANSP); Rio Surubres 

(09°56.1′N, 84°35′W; Bonnefil farm; 245 m; sweeping), 21 Oct 

1909, P. P. Calvert (1♂, 1♀; ANSP); Turrúcares (09°57.6′N, 

84°19.2′W), 22 Dec 1909, P. P. Calvert (1♀; ANSP). Cartago:

Cachí (09°49.7′N, 83°48.7′W; stagnant pool, bank of Rio 

Reventazon), 10 Mar 1910, P. P. Calvert (1♀; ANSP); Cartago 

(9°51.4′N, 83°55.2′W), 4 Jul–12 Dec 1909, P. P. Calvert (6♂, 

1♀; ANSP); Cartago (near; 09°51.3′N, 83°55.2′W; 1525 m), 

15 Dec 1909, P. P. Calvert (1♀; ANSP); Turrialba (09°54.1′N, 

83°41.1′W), Nov 1922, P. Schild (3♂, 2♀; ANSP). Limón: Río 

Banano (09°50.5′N, 82°56′W), 9 Nov 1909, P. P. Calvert (1♂; 

ANSP). San José: San José (09°56′N, 84°05′W), Dec 1919, 1925, 

A. Alfaro (3♂, 1♀; ANSP).

ECUADOR. El Oro: Rio Grande (15 km S Zaruma; 

03°42.8′S, 79°36.8′W; 850 m), 9 Apr 1965, L. E. Peña (1♂; 

CNC). Guayas: Machala (49.5 km NNE; 02°40′S, 79°13.4′W; 

40 m), 13 Jan 1978, W. N. Mathis (9♂, 6♀; USNM). Loja: San 

Pedro-Zarumard (04°04′S, 79°12.4′W; 850–1100 m), 9 Apr 

1965, L. E. Peña (1♂; CNC). Orellana: RíoTiputini (0°38.2′S, 

76°8.9′W), 12–26 Aug 1999, W. N. Mathis, A. Baptista, M. 

Kotrba (14♂, 1♀; USNM). Sucumbios: Lago Agrio (8 km W; 

0°04.9′S, 77°0.3′W; Malaise trap), 28 Aug 1975, J. Cohen, A. 

Langley (1♂, 2♀; USNM).

EL SALVADOR. La Libertad: La Libertad (13°40.9′N, 

89°21.6′W), 29 Oct 1965, N. L. H. Krauss (1♂; USNM); 

Quegaltepeque (13°50.1′N, 89°16.1′W), 11 Feb 1965, W. D. 

Duckworth (1♂; USNM). San Salvador: Nejapa (13°48.5′N, 

89°13.7′W), 6 Jul 1959, P. A. Berry (2♂, 2♀; USNM). Santa 

Ana: Santa Ana (13°59.7′N, 89°33.1′W), Oct 1959, N. L. H. 

Krauss (1♀; USNM).

GUATEMALA. Chimaltango: Yepocapa (14°30′N, 90°57′W), 

20 Feb 1948(1♂; USNM). Escuintla: San José (13°55.7′N, 

90°49.3′W), 20 Jul 1943, D. G. Hall (2♀; USNM). Guatemala:

La Providencia (14°39′N, 90°26.8′W), 14 Apr 1926, J. M. Aldrich 

(1♂, 2♀; ANSP). Izabal: Puerto Barrios (15°43′N, 88°35′W), 3–14 

Mar 1905 (2♂, 1♀; ANSP); Piña Conchita (14°44′N, 91°43′W), 20 

Apr 1948, H. Dalmat (1♂; USNM). Zacapa: El Jicaral (14°30′N, 

89°33′W), 11 May 1926 (1♂; ANSP); Santa Lucia (14°54.6′N, 

89°31.4′W), 2 Feb 1905 (2♀; ANSP).

GUYANA. Atkinson Airport Road (06°29.9′N, 

58°15.2′W; in Teobroma cacao plantation), 2 Jun 1965, S. 

Medina, L. F. Martore (2 ♀; USNM). Bartica District, Kartabo 

(06°23′N, 58°41′W), 12 Jun 1922 (1♂; ANSP). Berbice River 

(05°40.9′N, 57°51.5′W), 9–11 Apr 1994, W. N. Mathis 

(6♂, 1♀; USNM). Conservation of Ecological Interactions 

and Biotic Associations (~40 km S Georgetown; 06°29.9′N, 

58°13.1′W), 13 Apr–28 Aug 1994, 1995, 1997 (29♂; USNM). 

Dubulay Ranch-Aramatani Creek (05°40.9′N, 57°51.5′W), 

9–11 Apr 1994, W. N. Mathis (3♂; USNM). Dubulay Ranch, 

Warniabo Creek (05°39.8′N, 57°53.4′W), 10–11 Apr 1994, 

W. N. Mathis (1♂, 1♀; USNM). Dubulay Ranch, small creek 

(05°40.9′N, 57°51.5′W), 10 Apr 1994, W. N. Mathis (17♂; 

USNM). Kaieteur Falls (05°10.7′N, 59°29.2′W; 570 m), 7 

Apr 1994, W. N. Mathis (13♂, 4♀; USNM). Kaieteur Falls 

(05°10.5′N, 59°28.9′W), 21–24 Aug 1997, W. N. Mathis (4♂, 

2♀; USNM). Karanambo (03°45.1′N, 59°18.6′W), 31 Mar 

1994, W. N. Mathis (2♂; USNM). Karanambo, Rupununi 

River (03°45.1′N, 59°18.6′W; ox bow), 2 Apr 1994, W. N. 

Mathis (1♂, 1♀; USNM). Kanuku Mountains, Moco Moco 

River (03°18.2′N, 59°38.9′W), 29 Apr 1995, W. N. Mathis 

(6♂, 4♀; USNM). Kanuku Mountains, Kumu River and Falls 

(03°15.9′N, 59°43.5′W), 28–30 Apr 1995, W. N. Mathis (6♂, 

1♀; USNM). Kartabo Point (Mazaruni; 06°23′N, 58°41′W; 

Malaise trap at edge of secondary forest and farmed field), 

22–27 Dec 1982, 1983, W. E. Steiner (3♂, 1♀; USNM). Kato, 

Chiung River (04°39.7′N, 59°50.0′W), 1 May 1995, W. N. 

Mathis (5♂, 3♀; USNM). Kumu River, Lethem (25 km SE; 

03°15.9′N, 59°43.6′W), 4–5 Apr 1994, W. N. Mathis (5♂, 3♀; 
8 8 • smithsonian contributions to zoology

USNM). Menzies Landing (near Kaieteur Falls; 05°10.1′N, 

59°29.5′W), 23 Aug 1997, W. N. Mathis (8♂; 4♀; USNM). 

Moco Moco (30 km E Lethem; 03°18.2′N, 59°39.0′W), 3–6 

Apr 1994, W. N. Mathis (11♂, 2♀; USNM). Paramakatoi (04°42′N, 59°42.8′W), 24–25 Aug 1997, W. N. Mathis 

(39♂, 6♀; USNM). Takutu Mountains (06°15′N, 59°5′W; 

Malaise trap, near stream in montane rainforest), 6–19 Dec 

1983, EARTHWATCH Research Expedition, P. J. Spangler, 

W. E. Steiner (40♂, 29♀; USNM). Wiruni River (05°46.6′N, 

58°0.8′W), 11 Apr 1994, W. N. Mathis (13♂, 4♀; USNM).

MEXICO. Chiapas: Río Izapa (14°55.4′N, 92°10.7′W), 21 

Apr 1983, W. N. Mathis (16♂, 10♀; USNM); Cacahoatán (7 

km N; 15°03.8′N, 92°09.1′W), 22 Apr 1983, W. N. Mathis (7♂, 

2♀; USNM); Union Juárez (9 km S; 15°02.2′N, 92°05.4′W), 23 

Apr 1983, W. N. Mathis (3♂; USNM). Veracruz-Llave: Ciudad 

Alemán (18°11.3′N, 96°05.6′W; 100 m), 3 May 1985, W. N. 

Mathis (8♂, 5♀; USNM); Fortin de las Flores (18°54′N, 97°W; 

952 m), 2 May 1985, W. N. Mathis (2♂; USNM); Ocotal Chico 

(18°15.6′N, 94°51.5′W; 600 m), 4–5 May 1985, W. N. Mathis 

(1♀; USNM).

PANAMÁ. Cocle: Playa Santa Clara (08°22.4′N, 

80°06.3′W), 2 Jul 1967, W. W. Wirth (9♂, 5♀; USNM). Panamá: Balboa (08°56.9′N, 79°33.9′W), Feb 1958, M. R. Wheeler 

(1♂; USNM).

PARAGUAY. Asunción: Asunción (25°16.9′S, 57°38.1′W), 

7 Oct 1980, D. C. Lowrie (1♂, 2♀; USNM). Presidente Hayes:

Toldo-cué (22°52′S, 58°36′W) (1♀; ANSP).

PERU. Cuzco: Paucartambo, Atalaya (Río Alto Madre de 

Dios; 12°53.1′S, 71°21.6′W; 600 m), 4 Sep 1988, W. N. Mathis 

(1♂, 2♀; USNM); Paucartambo, Puente San Pedro (~50 km 

NW Pìlcopata; 13°03.3′S, 71°32.8′W; 1600 m), 3 Sep 1988, 

W. N. Mathis (1♂; USNM). Loreto: Explornapo Camp on Rio 

Sucusari, 2 km upstream from Rio Napo (160 km NE Iquitos; 

03°15.4′S, 72°55.2′W; Malaise trap), 24 Jun–20 Aug 1990, B. 

Awertschenko, A. Menke (6♀; USNM); Granja UNAP, near 

Iquitos (03°42′S, 73°13.7′W), 23 Feb 1984, W. N. Mathis 

(29♂, 3♀; USNM); Iquitos (12 km W; 03°48.4′S, 73°20.5′W), 

16 Feb 1984, W. N. Mathis (13♂, 8♀; USNM); Iquitos (14 

km W; 03°52.1′S, 73°28.2′W), 16–21 Feb 1984, W. N. Mathis 

(25♂, 8♀; USNM); Iquitos (15 km W; 03°52.1′S, 73°29′W), 

16 Feb 1984, W. N. Mathis (5♂, 4♀; USNM); Iquitos (5 km 

E; 03°50.3′S, 73°11.1′W), 17 Feb 1984, W. N. Mathis (4♂; 

USNM); Rio Itaya (25 km S Iquitos; 04°11.3′S, 73°15′W), 22 

Feb 1984, W. N. Mathis (2♂; USNM); Río Momón (~25 km 

NW Iquitos; 03°40.1′S, 73°17.6′W), 17 Feb 1984, W. N. Mathis 

(1♂; USNM).

TRINIDAD AND TOBAGO. Tobago. St. John: Bloody 

Bay River (11°18′N, 60°38′W), 14 Jun 1993, W. N. Mathis (3♂; 

USNM); Charlotteville (beach; 11°19.5′N, 60°32.9′W), 16 Apr–

16 Jun 1993, 1994, D. and W. N. Mathis (24♂, 4♀; USNM); 

Charlotteville (5 km S; 11°18.9′N, 60°34.5′W), Hermitage River 

and beach, 22 Apr–11 Jun 1993, 1994, D. and W. N. Mathis 

(16♂, 1♀; USNM); Kings Bay Reservoir (11°17′N, 60°33′W), 

15 Jun 1993, W. N. Mathis (1♂; USNM); Parlatuvier (creek; 

11°17.9′N, 60°35′W), 14 Jun 1993, W. N. Mathis (9♂; USNM); 

Speyside (11°18′N, 60°32′W), 13–15 Jun 1993, W. N. Mathis 

(3♂, 1♀; USNM); Speyside (Doctor River; 1 km NW; 11°18′N, 

60°32′W), 12–13 Jun 1993, W. N. Mathis (6♂; USNM). St. 

Paul: Argyle Falls (11°15′N, 60°35′W), 21 Apr 1994, W. 

N. Mathis (2♂, 1♀; USNM); Delaford, Kings Bay (11°16′N, 

60°32.8′W), 13 Jun 1993, W. N. Mathis (5♂; USNM). Trinidad. 

St. George: Filette (1 km SE; 10°47′N, 61°21′W; Yarra River), 

25 Jun 1993; W. N. Mathis (2♂, 1♀; USNM); Marianne River 

(9 km S; 10°46′N, 61°18′W), 25 Jun 1993, W. N. Mathis (2♂, 

1♀; USNM).

VENEZUELA. Territorio Federal Amazonas: Cerro de la 

Neblina, basecamp (bank of Rio Baria; 00°50′N, 66°09.7′W; 

140 m), 27 Jan–24 Mar 1984, 1985, W. E. Steiner (49♂, 59♀; 

USNM); Cerro de la Neblina (basecamp; 00°50′N, 66°10′W), 

5–18 Feb 1985 (2♂; USNM); Cerro de la Neblina (00°51′N, 

66°10′W; 140 m), 13–19 Mar 1984, D. Davis, T. McCabe, 

W. E. Steiner (10♂, 3♀; USNM). Carabobo: Boquerón (Cumbres; 10°04′N, 67°55.4′W; frente a Bajo Seco; 1600 m), 

16.Apr 1972 (1♂;USNM); Embalse de Guataparo, W Valencia (10°10.9′N, 68°03.5′W), 13 Apr 1972, L. V. Knutson (1♂; 

USNM).

WEST INDIES. CUBA. Matanzas: Buena Ventura (7 km 

N; 22°18.1′N, 81°11.7′W), 1 May 1983, W. N. Mathis (1♂; 

USNM).

Type Locality. Neotropical. “Venezuela.”

Distribution. (Map 23) Neotropical: Belize 

(Stann Creek), Bolivia (Bení, La Paz), Brazil (Amazonas, Pará, 

São Paulo, Tocantins), Colombia (Magdalena), Ecuador (El 

Oro, Guayas, Loja, Orellana, Sucumbios), El Salvador (La 

Libertad, San Salvador, Santa Ana), Guatemala (Chimaltango, 

Escuintla, Guatemala, Iazabal, Zacapa), Guyana, Mexico 

(Chiapas, Veracruz-Llave), Panamá (Cocle, Panamá), Paraguay (Asunción, President Hayes), Peru (Cuzco, Loreto), Trinidad and Tobago, Venezuela (Amazonas, Carabodo), West 

Indies (Cuba).

Remarks. The type material of P. secunda includes 

only females and is thus a difficult species to identify. Cresson 

(1947) presented a diagnosis of the species, including males, 

that helped us in the identification of this species in addition 

to the material determined by Cresson that is in the ANSP and 

USNM. This species shows a wide range of colorational variation. Brazilian specimens have a face with a single rectangular 

spot of pale brown microtomentum as long as the length of the 

antenna; the dorsal and ventral margins of the anepisternum 

are brown, there are small brown spots at the base of the setae 

in the middle; the tarsomeres are lighter with yellowish-orange 

coloration ventrally and dorsally on the first tarsomeres; and 

tergite 2 has lateral, marginal, silvery bands that are nearly approximate medially, separated medially by a slender pale brown 

stripe. In the male terminalia of P. secunda, the presurstylus 

varies in width, but the characteristic convex dorsal margin and 

cercus inserted at the apex of the horizontal process is distinctive 

and constant.
number 643 • 8 9

MAP 23. Distribution map for Paralimna (Paralimna) secunda Schiner.

28. Paralimna (Paralimna) sera Cresson

Figures 122–125, 193, Map 17

Paralimna sera Cresson 1933:67 [West Indies. Jamaica. St. Thomas: Yallahs 

Valley, Blue Mountains; HT ♂, AMNH].

Paralimna (Paralimna) sera.—Cresson 1947:52 [review, Neotropical species].—Wirth 1968:15 [Neotropical catalog].—Mathis and Zatwarnicki 1995:125 [world catalog].

Diagnosis. This species is distinguished from congeners by the following combination of characters: face dark 

gray with a wide and short dark grayish-brown spot on carina, 

lateral margins dark gray to brown, thereafter ventrally mostly 

to entirely gray; anepisternum with dorsal and ventral margins 

brown, central area mostly gray, sometimes with bases of setulae 

brown; katepisternum mostly whitish gray to dark gray, concolorous with anepimeron; legs unusually slender, forefemur of 

male bearing anteroventral comb-like row of distinctly flattened, 

moderately long setae, concave ventrally near apex, foretibia of 

male with slightly wavy ventral surface; tergites predominantly 

dark brown with a slender gray band on posterior margin.

Description. Body length 4.0–4.2 mm; body generally bicolored, dark brown dorsally, lateral surfaces dark, 

rather bluish gray.

Head: (See Figure 193) Frons brown to pale brown, with 

linear gray spots at lateral margins of ocellar triangle, in front of 

anterior ocellar seta, reaching the anterior margin of frons, between 

inner vertical and interfrontal seta, and on fronto-orbits just above 

antennal bases; ventral fronto-orbits and dorsal parafacial without 

outstanding spots. Antennal scape and pedicel dark brown, pedicel 

with pale gray margin; basal flagellomere pale brown to brown according to incidence of light, few and short setae on margin; arista 

with 8–9 long, dorsal rays. Postcranium with 2 complete series 

of postocular setae. Face dark gray with a wide and short dark 

grayish-brown spot on carina, lateral margins dark gray to brown, 

thereafter ventrally mostly to entirely gray; parafacial and gena 

gray; gena with only few setae on middle and ventral margin; clypeus silvery gray. Eye rounded, about as wide as high, height more 

than twice the height of gena. Gena high, height slightly greater 

than length of basal flagellomere; gena-to-eye ratio 0.35–0.36.

Thorax: Mesonotum predominantly brown with gray 

spots along dorsocentral track and laterally; postpronotum and 

supra-alar areas gray; scutellum brown basally, with apex blackish 
9 0 • smithsonian contributions to zoology

brown and grayish brown laterally. Pleurae dark gray, anepisternum with dorsal and ventral margins brown, central area mostly 

gray, sometimes with bases of setulae brown; katepisternum 

mostly whitish gray to dark gray, concolorous with anepimeron. 

Wing hyaline; veins brown; costal-vein ratio 0.37–0.39; M-vein 

ratio 1.00–1.04. Legs brown with sparse silvery-gray microtomentum, tarsi lighter with some yellow coloration, especially ventrally 

and on tarsi 1–3; legs unusually slender; forefemur of male concave ventrally near apex, bearing anteroventral comb-like row of 

distinctly flattened, moderately long setae and anteroventral surface bearing sparse row of short setae along most of length; foretibia of male with slightly wavy ventral surface.

Abdomen: Tergites predominantly dark brown with a 

slender gray band on posterior margin; tergites 1 and 2 brown 

with a complete gray band; tergites 3–5 with gray band incomplete, interrupted medially, brown bands of tergites 4 and 

5 extended laterally to lateral margin of tergite. Male terminalia: (Figures 122–125) Presurstylus not bifurcate, only bulged 

FIGURES 122–125. Structures of male terminalia of Paralimna (Paralimna) sera Cresson (Jamaica. St. Andrew): (122) epandrium, cerci, and presurstyli, posterior aspect; (123) postsurstylus, lateral aspect; (124) aedeagus, phallapodeme, and hypandrium, lateral aspect; (125) same, ventral aspect. Scale bar = 0.1 mm. (See Figures 2–5 for labels of structures.)
number 643 • 9 1

basoventrally, horizontal process gradually tapering toward apex, 

shallowly curved with very long, slender setae on distal half; in 

lateral view postsurstylus not obviously wider medially, medial 

anterior lobe poorly produced, elongate setae extended over medial region of external surface, more dense and close to each other 

on medial anterior lobe; lateral aedeagal process large, longer 

than 2⁄3 length of aedeagus, widened and slightly curved in middle, 

broadly rounded distally; aedeagus slender in posterior view; phallapodeme broadly triangular in lateral view, with anteroventral 

side waved; hypandrium moderately concave.

Type Material. The holotype male of Paralimna 

sera is labeled “Yallahs Valley[,] Blue Mts.[,] Jamaica[,] Feb. 

24. 1911/♂/Am. Mus. Nat. Hist. Dept. Invert. Zool. No./TYPE 

Paralimna SERA E. T. Cresson,Jr. [“Paralimna SERA” handwritten; pink].” The holotype is pinned directly, is in good 

condition (mesonotum cracked where pin is inserted; abdomen 

not dissected), and is deposited in the AMNH.

Other Specimens Examined. JAMAICA. Portland: Green Hills (18°05.7′N, 76°43′W; 780 m), 28 Apr 2000, 

W. N. Mathis (5♂; USNM); Hollywell (18°05.5′N, 76°43.6′W; 

1170 m), 27 Apr 2000, W. N. Mathis (11♂, 1♀; USNM); Section 

(0.5 km E; 18°05.2′N, 76°43.9′W; 1020 m), 28 Apr 2000, W. N. 

Mathis (11♂, 3♀; USNM). St. Andrew: Cinchona (18°04.4′N, 

76°39.3′W), 28 Jul 1928, C. C. Gowdey (1♀; USNM); Cinchona (18°04.4′N, 76°39.3′W; 1400 m), 29 Apr 2000, W. N. 

Mathis (1♂, 1♀; USNM); Clydsdale (18°04.9′N, 76°40.2′W; 

1030 m), 29 Apr 2000, W. N. Mathis (9♂, 1♀; USNM); Hardwar Gap (18°04.2′N, 76°44′W; 1170 m), 10 Mar–8 Jun 1970, 

1989, 1996, 2000, E. Becker, T. H. Farr, H. Howden, D. and 

W. N. Mathis, B. and M. Sorensson, H. B. Williams, W. W. 

Wirth (33♂, 16♀; CNC, USNM, ZIL); Hardwar Gap (2.5 km 

E; at Fairy Glades Trail; 18°04.4′N, 76°43.8′W; 1350 m), 9 Jun 

1989, B. and M. Sorensson (1♀; ZIL); Hollywell (18°05.2′N, 

76°43.9′W; 1100 m), 28 Apr–13 Jun 1989, W. N. Mathis, B. 

and M. Sorensson (27♂, 16♀; USNM, ZIL); Newcastle Road 

(mi 17 [km 27] from Kingston; 18°04.1′N, 76°42.8′W), 3 May 

1954, T. H. Farr (1♀; USNM); Mavis Bank (near coffee factory; 18°01.4′N, 76°39.7′W; waterfall), 21–23 Apr 2000, W. N. 

Mathis (14♂, 4♀; USNM); Mavis Bank (1.7 km E; 18°02.4′N, 

77°39.5′W; 575 m), Yallahs River, 21–22 Apr–1 May 2000, 

W. N. Mathis (16♂, 2♀; USNM); Mavis Bank (4.3 km SE; 

18°01.4′N, 76°38.1′W; 480 m), Yallahs River, 22–23 Apr 2000, 

W. N. Mathis (4♂; USNM); Newcastle (6 km S; 18°04.3′N, 

76°42.6′W; 950 m; waterfall), 30 Apr 2000, W. N. Mathis (6♂, 

1♀; USNM); Second Breakfast Spring (18°03′N, 76°46′W), 3 

May–29 Aug 1954, T. H. Farr (2♂, 1♀; USNM); Silver Hill Gap 

(18°05.1′N, 76°41.1′W; 920 m), 26 Apr 2000, W. N. Mathis 

(13♂, 2♀; USNM); Silver Hill Gap (18°05.3′N, 76°43′W; 940 

m), 29 Apr 2000, W. N. Mathis (7♂; USNM). St. Thomas:

Blue Mountains, Yallahs Valley (17°52.6′N, 76°33.1′W), 24 

Feb 1911 (3♀; AMNH, ANSP, USNM); Hagley Gap (1 km E; 

18°00.1′N, 76°36.7′W), 16 May 1996, D. and W. N. Mathis, 

H. B. Williams (12♂, 20♀; USNM).

Type Locality. Neotropical. Jamaica. St. Thomas: 

Blue Mountains, Yallahs Valley (17°52.6′N, 76°33.1′W).

Distribution. (Map 17). Neotropical: West Indies 

(Jamaica).

Remarks. This species is apparently endemic to Jamaica and is similar to Paralimna (Paralimna) maculata sp. nov. 

Distinguishing characters are discussed in the “Remarks” for P. 

maculata.

29. Paralimna (Paralimna) stellata, sp. nov.

Figures 1, 126–129, 194, 207, Map 16

Diagnosis. This species is distinguished from congeners by the following combination of characters: species with 

body predominantly brown with long face and notably protuberant carina; frons brown with a wide central portion yellow 

and a large black velvety spot on ventral front orbit lateral to antenna; face reddish brown; scape and pedicel black velvety; wing 

brown with hyaline halos, crossvein dm–cu concave, curved 

to base; vein M with short perpendicular stump veins; tergites 

brown with laterodistal triangular yellow spots.

Description. Body length 3.5–4.1 mm; body generally dark colored, dark brown dorsally and lateral surfaces dark 

gray.

Head: (See Figure 194) Frons stained reddish brown and 

yellow: brown region extends from interfrontal seta to inner and 

outer vertical setae reaching the lateral margin of frons, yellow 

region extends from between inner vertical setae toward anterior 

fronto-orbital seta reaching the lateral margin of frons forming a 

triangular stain, slender longitudinal stripes visible on yellow region depending of incidence of light, the more conspicuous brown 

stripe extends from anterior ocellus to anterior margin of frons, 

margin anterolateral of frons with a wide black velvety spot lateral to antenna; dorsal parafacial with 1 elongate pale brown spot 

lateral to antenna, separated from black velvety dorsal frontoorbital spot by a yellow spot. Antennal scape and pedicel black 

velvety; basal flagellomere brown, covered by yellowish-orange 

microtomentum; basal flagellomere with slender, elongate, pale 

setae on margin; arista with 13 long, dorsal rays. Postcranium 

with 2 complete series of postocular setae, the second series irregular. Face long, in anterior view twice height of frons; reddish 

brown with a yellowish-gray small spot in middle; parafacial yellow and gena reddish brown; carina notably protuberant; gena 

with only few setae on middle and ventral margin; clypeus reddish brown. Eye rounded, about as wide as high, height more 

than twice the height of gena. Gena high, height slightly greater 

than length of basal flagellomere; gena-to-eye ratio 0.36–0.40.

Thorax: Mesonotum yellow with brown spots, 3 stripes 

between dorsocentral rows, plus several short stripes laterally; 

postpronotum and supra-alar areas yellow; scutellum brown dorsally with a slender longitudinal yellow stripe, with apex blackish 

brown and yellow otherwise. Pleurae stained with brown and yellow, anepisternum brown with small area yellow surrounding the 

ventral anepisternal setae, anepimeron brown on ventral half and 

yellow above, katepisternum brown with margins yellow, the remaining yellow. Wing (see Figure 207) brown with hyaline halos; 
9 2 • smithsonian contributions to zoology

costal-vein ratio 0.37–0.39; M-vein ratio 1.00–1.04; veins brown, 

crossvein dm–cu concave, curved to base; vein M with short perpendicular stump veins. Legs blackish brown with dense reddishbrown microtomentum; tarsi lighter with some yellow coloration, 

especially ventrally, the distal tarsomeres brown; forefemur of 

male with anteroventral series of short and slightly flattened setae, 

ventral surface with a comb of very short and slender setae; foretibia of male straight.

Abdomen: Tergites brown with laterodistal triangular yellow spots, lateroventral margin brown. Male terminalia: 

(Figures 126–129) Presurstylus not bifurcate, horizontal process 

somewhat robust, shallowly curved with very long, slender setae 

along the length; in lateral view postsurstylus widest medially with 

a medial, anteriorly protuberant, apically truncate lobe bearing 

long and slender setae and a rounded process posteriorly; lateral 

aedeagal process about 1⁄2 length of aedeagus, slender and mostly 

FIGURES 126–129. Structures of male terminalia of Paralimna (Paralimna) stellata, sp. nov. (Ecuador. Pastaza): (126) epandrium, cerci, and presurstyli, posterior aspect; (127) postsurstylus, lateral aspect; (128) aedeagus, phallapodeme, and hypandrium, lateral aspect; (129) same, ventral aspect. Scale bar = 0.1 mm. (See Figures 2–5 for labels of structures.)
number 643 • 9 3

straight in lateral view; moderately robust in posterior view and 

slender in lateral view; phallapodeme broadly triangular in lateral 

view; hypandrium deeply concave.

Type Material. The holotype male of Paralimna 

stellata is labeled “ECUADOR. Past[aza]. Puyo (22 Kms. 

E) 2 February 1976[,] Spangler, et al./ECUADOR-PEACE 

CORPS-SMITHSONIAN INSTITUTION AQUATIC INSECT 

SURVEY/USNM ENT 00118288 [plastic bar code label]/HOLOTYPE ♂ Paralimna stellata Ale-Rocha&Mathis, USNM 

[red].” The holotype is double mounted (glued to a paper triangle), is in excellent condition, and is deposited in the USNM. 

One paratype (♂; USNM) bears the same label data as the holotype. Other paratypes are as follows: ECUADOR. Pastaza:

Puyo (01°29′S, 78°0.1′W), 30 Jan 1976, P. J. Spangler et al. (1♀; 

USNM); Puyo (21 km E; 01°17.9′S, 77°52.9′W), 14 Jul 1976, 

J. Cohen (1♀; USNM). Sucumbios: Lago Agrio (Nueva Loja; 

00°05.1′S, 76°53′W), 16 Apr 1976, A. Langley (1♂; USNM); 

Lago Agrio (3 km NE; 00°06.4′S, 76°52.6′W; at pozo number 

3), 17 May 1975, A. Gurney (1♀; USNM).

Other Specimens Examined. ECUADOR.

Napo: Puerto Misahuallí (01°02′S, 78°40.2′W; 350 m), Feb 1993, 

M. Sharkey (1♂; CNC). Orellana: Coca, Napo River (0°27.8′S, 

76°59′W; 250 m), 12–13 Apr 1965, L. E. Peña (1♂; CNC); Río 

Tiputini (0°38.2′S, 76°08.9′W), 12–26 Aug 1999, W. N. Mathis, 

A. Baptista, M. Kotrba (4♂, 2♀; USNM). Pastaza: Puyo, (3–4 km 

N; 01°28.1′S, 77°59.7′W; 953 m), 9 Feb 1955 (2♂, 1♀;USNM).

PERU. Huánuco: Tingo María (09°17.7′S, 75°59.8′W; 

800 m), 5–9 Apr 1962, L. E. Peña (1♂; CNC). Junín: Perené 

(10°56.8′S, 75°13.6′W), R. C. Shannon (2♂, 1♀; USNM). Loreto: Explornado Camp on Rio Sucusari, 2 km upstream from 

Rio Napo (160 km NE Iquitos; Malaise trap), 24 Jun–20 Jul 

1990, B. Awertschenko, A. Menke (1♀; USNM). San Martin:

Tarapoto (06°29′S, 76°22′W), Sept 1955, Th. Dobzhansky, C. 

Pavan (1♂; USNM).

Type Locality. Neotropical. Ecuador. Pastaza: Puyo 

(22 km E; 01°17.9′S, 77°52.9′W).

Distribution. (Map 16) Neotropical: Ecuador 

(Napo, Orellana, Pastaza, Sucumbios), Peru (Huánuco, Junín, 

Loreto, San Martin).

Etymology. The species epithet, stellata, is of Latin 

derivation and means starred or spotted, alluding to the spotted 

aspect of the body and wing.

Remarks. This species forms a monophyletic group 

that also includes P. guttata sp. nov. and P. fulgifrons sp. nov. 

The differences between these species are discussed in the “Remarks” for P. guttata sp. nov.

30. Paralimna (Paralimna) stigmata, sp. nov.

Figures 130–133, 195, 208, Map 24

Diagnosis. This species is distinguished from congeners by the following combination of characters: face brown to 

dark brown with lateral margin silvery gray to pale gray and an 

inverted U-shaped, wide, gray to pale gray band on middle of 

face; pleurae predominantly brown with dorsal portion of proepimeron, posterodorsal corner and a small semicircular anterior medial spot on anepisternum, dorsal half of anepimeron, 

laterotergite, and anterodorsal portion of katepisternum gray; 

forefemur of male without outstanding setae.

Description. Body length 3.3–3.7 mm; body generally bicolored, dark brown dorsally, lateral surfaces predominantly brown with some small areas gray.

Head: (See Figure 195) Frons brown with gray spots, a 

slender stripe at lateral margins of ocellar triangle, in front of anterior ocellar seta but not reaching the anterior margin of frons, 

between inner vertical and interfrontal setae, and on front-orbits 

above dorsal proclinate fronto-orbital seta, between proclinate 

fronto-orbital setae, and just above antennal bases; ventral frontoorbits and dorsal parafacial with 2 brown spots lateral to antenna, 

separated by gray spot. Antennal scape and pedicel dark brown, 

pedicel with dorsal distal margin silvery gray, with 1 dorsal, 

thicker seta and some short setae on the margin; basal flagellomere 

gray with slender, elongate pale brown setae on margin; arista 

with 12–13 long dorsal rays. Postcranium with 2 series of postocular setae, the posterior series is somewhat irregular. Face brown 

to dark brown with lateral margin silvery gray to pale gray and an 

inverted U-shaped, wide, gray to pale gray band on middle of face; 

parafacial and gena silvery gray to pale gray; gena with setae only 

on distal margin; clypeus gray. Eye round, as wide as high, height 

more than twice the height of gena. Gena high, height subequal 

or slightly greater than length of basal flagellomere; gena-to-eye 

ratio 0.30–0.36.

Thorax: Mesonotum pale gray with brown spots, 

3 stripes between dorsocentral rows being 2 lateral, well-defined 

stripes and 1 irregular, slender, and less-defined acrostichal stripe; 

1 slender short stripe on supra-alar area and several small spots in 

the base of setae; postpronotal lobe gray; notopleuron gray with 

a little brown spot on dorsal margin; scutellum pale gray, slightly 

tinged of brown dorsally, with distal margin dark brown. Pleurae 

predominantly brown with dorsal portion of proepimeron, dorsal 

posterior corner and a small anterior medial spot on anepisternum, dorsal half of anepimeron, laterotergite, and anterodorsal 

portion of katepisternum gray. Wing (see Figure 208) pale brown 

tinged or semi-hyaline; veins brown; crossvein dm–cu slightly infuscate; costal-vein ratio 0.45–0.46; M-vein ratio 1.09–1.17. Legs 

brown except basitarsus lighter, yellowish tinged; forefemur of 

male without outstanding setae; foretibia straight.

Abdomen: Tergites distinctly bicolored, dark brown along 

anterior 2⁄3 and medially, the remaining yellowish gray dorsally and 

silvery gray laterally, except tergite 2 predominantly yellowish gray 

with only a dorsolateral small brown spot on basal half; lateroventral margin of tergites brown. Male terminalia: (Figures 130–133) 

Presurstylus not bifurcate, with horizontal process slender and 

slightly wavy; in lateral view postsurstylus widest medially with a 

medial, anteriorly protuberant lobe bearing rather short and slender setae and a robust digitiform process posteriorly; lateral aedeagal process about 1⁄2 length of aedeagus, robust and slightly curved 
9 4 • smithsonian contributions to zoology

subapically in lateral view, bluntly rounded; aedeagus rather robust, rounded on membranous apex; phallapodeme broadly triangular in lateral view; hypandrium shallowly concave.

Type Material. The holotype male of Paralimna 

stigmata is labeled “BRAZIL. Amazonas: Manaus, UFAM 

[Univeridade Federal do Amazonas] (03°05.9′S, 59°58.2′W, 50 

m), 7 May 2010/D. & W. N. Mathis/USNM ENT 00118266 

[plastic bar code label]/HOLOTYPE ♂ Paralimna stigmata AleRocha&Mathis, INPA [red].” The holotype is double mounted 

(minuten pin in a white block of plastic), is in excellent condition, and is deposited in INPA. Six paratypes (6♂; INPA, USNM) 

bear the same label data as the holotype. Other paratypes are 

as follows: BRAZIL. Amazonas: Manaus, Reserva Ducke (km 

24; 02°55.8′S, 59°58.5′W; Malaise trap), 21 Aug 1990, J. Vidal 

(1♂; INPA); Manaus, INPA (03°55.8′S, 59°59.2′W; 60 m), 4 

May 2010, D. and W. N. Mathis (8♂; INPA, USNM); Ipixuna, 

FIGURES 130–133. Structures of male terminalia of Paralimna (Paralimna) stigmata, sp. nov. (Brazil. Amazonas): (130) 

epandrium, cerci, and presurstyli, posterior aspect; (131) postsurstylus, lateral aspect; (132) aedeagus, phallapodeme, and 

hypandrium, lateral aspect; (133) same, ventral aspect. Scale bar = 0.1 mm. (See Figures 2–5 for labels of structures.)
number 643 • 9 5

MAP 24. Distribution map for Paralimna (Paralimna) stigmata, sp. nov.

Rio Gregório, Comunidade Lago Grande (07°09.9′S, 70°48.9′W; 

sweeping), 18–23 May 2011, J. A. Rafael; J. T. Camara, R. F. 

Silva, A. Somavilla, R. Ale-Rocha (1♀; INPA).

ECUADOR. Orellana: Río Tiputini (0°38.2′S, 76°8.9′W), 

12–26 Aug 1999, W. N. Mathis, A. Baptista, M. Kotrba (10♂, 

3♀; INPA, USNM).

GUYANA. Kanuku Mountains, Kumu River and Falls 

(03°15.9′N, 59°43.5′W), 28–30 Apr 1995, W. N. Mathis 

(6♂, 3♀; USNM); Kumu River Lethem (25 km SE; 03°15.9′N, 

59°46.6′W), 4–5 Apr 1994, W. N. Mathis (14♂; 5♀; USNM); 

Moco-Moco (30 km E Lethem; 03°18.2′N, 59°39.0′W), 3–6 

Apr 1994, W. N. Mathis (11♂, 4♀; INPA, USNM); Paramakatoi (04°42′N, 59°42.8′W), 24–25 Aug 1997, W. N. Mathis (1♂; 

USNM).

VENEZUELA. Territorio Federal Amazonas: Cerro de La 

Neblina (basecamp; 00°51′N, 66°10′W; 140 m; Malaise trap), 

20–24 Mar 1984, O. S. Flint, J. Louton (1♀; USNM). Carabobo: Valley Seco (10°28′N, 68°01′W), Jan 1940, P. Adams (1♂; 

ANSP).

Other Specimens Examined. ECUADOR.

Orellana: Coca, Napo River (0°27.8′S, 76°59′W; 250 m), 

12–13 Apr 1965, L. E. Peña (2♂; CNC); Pomeya, Napo River 

(0°26.5′S, 76°36.6′W), 25–30 Apr 1965, L. E. Peña (1♂, 1♀; 

CNC). Zamora-Chinchipe: Cumbaratza (03°59.5′S, 78°51.9′W; 

700 m), 30–31 Mar 1965, L. E. Peña (2♂; CNC); Río Jumboé 

(04°04′S, 78°55.8′W; 1200 m), 1–2 Apr 1965, L. E. Peña (3♂, 

2♀; CNC); Timbara (04°01.9′S, 78°54.1′W; 1400 m), 4 Apr 

1965, L. E. Peña (5♂; CNC); Yacuambi (03°35.7′S, 78°55.7′W; 

1000 m), 4 Apr 1965, L. E. Peña (1♂; CNC).

VENEZUELA. Bolívar: Gran Sabana, Akuriman (08°16′N, 

62°45.3′W), Nov–Dec 1940, P. Anduze (2♂; USNM).

Type Locality. Neotropical. Brazil. Amazonas: Manaus, Universidade Federal do Amazonas (03°05.9′S, 

59°58.2′W; 50 m).

Distribution. (Map 24) Neotropical: Brazil (Amazonas), Ecuador (Orellana, Zamora-Chinchipe), Guyana, Venezuela (Territorio Federal Amazonas, Bolívar, Carabobo).

Etymology. The species epithet, stigmata, is of Latin 

derivation and means mark or bank, alluding to the inverted 

U-shaped golden spot on the face.

Remarks. The presurstylus of this species is a horizontal, simple process that is rather sinuous, similar to that of 
9 6 • smithsonian contributions to zoology

P. molossus and P. stellata sp. nov., but it can be distinguished 

from these species by the brown face that has an inverted Ushaped, wide, gray to pale gray band in the middle of the face. 

The similarity of the male terminalia of P. molossus and P. stigmata sp. nov. is discussed in the “Remarks” for P. molossus.

31. Paralimna (Paralimna) taurus Cresson

Figures 134–137, Map 25

Paralimna taurus Cresson 1916:123 [Brazil. Mato Grosso do Sul: Corumbá (19°0.6′S, 57°39.3′W); HT ♂, ANSP (6096)].—Wirth 

1968:15 [Neotropical catalog].—Mathis and Zatwarnicki 1995: 25 

[world catalog].

Paralimna secunda taurus Cresson 1947:48 [review, Neotropical species].

Paralimna sticta Hendel 1930:127 [Argentina. San José; HT ♂, SMN]. Cresson 1947:53 [list, Neotropical species].—Wirth 1968:15 [Neotropical 

catalog].—Mathis and Zatwarnicki 1995:125 [world catalog]. NEW

SYNONYM.

Diagnosis. This species is distinguished from congeners by the following combination of characters: face gray with 

a faint, short pale brown spot on carina, half the length of antenna; face long in anterior view, about twice height of frons; 

frons horizontal; mesopleuron and legs densely silvery gray; 

anepisternum with dorsal margin brown tinged, with a dotted 

aspect in the remaining by bases of setae brown; base of scutal 

setae brown giving a dotted aspect to scutum; forefemur of male 

with ventral and anteroventral series of short setae, the anteroventral setae discreetly flattened and shortened toward the apex 

of the femur.

Description. Body length 3.5–5.5 mm; body generally bicolored, pale brown dorsally, lateral surfaces densely silvery gray.

Head: Frons horizontal, pale brown to brown with linear, gray, faded spots at lateral margins of ocellar triangle, in 

front of anterior ocellar seta, not reaching the anterior margin of 

frons, between inner vertical and interfrontal setae, and on frontoorbits, between proclinate fronto-orbital setae and just above 

antennal bases; ventral fronto-orbits with a dark gray spot and 

dorsal parafacial spot inconspicuous. Antennal scape and pedicel 

dark brown; basal flagellomere pale brown with slender, elongate, pale setae on margin; arista with 12–13 long, dorsal rays. 

Postcranium with 2 complete series of postocular setae. Face 

long in anterior view, about twice height of frons, gray with a 

faint, short pale brown spot on carina, half of the length of antenna; parafacial and gena silvery gray; gena with only few setae 

on middle and ventral margin; clypeus silvery gray. Eye rounded, 

about as wide as high. Gena high, height subequal to length of 

basal flagellomere.

Thorax: Mesonotum gray with pale brown spots, 3 poorly 

defined wide stripes between dorsocentral rows, plus some short 

brown stripes on intra- and supra-alar areas, and base of setae 

brown giving a dotted aspect to scutum; postpronotum and supra-alar areas silvery gray; scutellum brown basally, with apex 

blackish brown and yellowish gray laterally and in middle. Pleurae densely silvery gray, anepisternum with dorsal margin slightly 

brown tinged, with a dotted aspect in the remaining by bases of 

setae brown. Wing hyaline; veins brown. Legs somewhat slender, pale brown with dense silvery-gray microtomentum; tarsi 

lighter with some yellow coloration, darkened on distal tarsomeres; forefemur of male with posteroventral and anteroventral 

complete series of short setae, the anteroventral setae discreetly 

flattened and shortened toward the apex of the femur; foretibia 

straight.

Abdomen: Tergites distinctly bicolored; tergite 1 brown 

in the most part with laterodistal corner silvery gray; tergites 2–5 

brown along anterior half and medially, otherwise silvery gray, 

except tergite 2 predominantly silvery gray with medial brown 

line reduced and dorsolateral brown spot on basal 1⁄2–1⁄3. Male 

terminalia: (Figures 134–137) Presurstylus not bifurcate, only 

bulged basoventrally, horizontal process rather robust, shallowly curved with long, slender setae; in lateral view postsurstylus widest medially, with medial, anterior lobe greatly produced, 

bearing short and stout setae extended over medial region until 

near posterior margin, a medial developed projection on posterior margin; lateral aedeagal process thin, about 1⁄3 length of 

aedeagus, almost straight; aedeagus slender in posterior view; 

phallapodeme broadly triangular in lateral view; hypandrium 

deeply concave.

Type Material. The holotype male of Paralimna 

taurus is labeled “May (above the cork block)/Corumbá/♂/TYPE 

Paralimna TAURUS E. T. Cresson, Jr. [maroon; “Paralimna 

TAURUS” handwritten].” The holotype is double mounted (pin 

in a rectangular block of cork, is in excellent condition (wings 

slightly torn), and is deposited in the ANSP (6096). Two paratypes (1♂, 1♀; ANSP), including the allotype, bear the same 

label data as the holotype.

The lectotype male of Paralimna sticta Hendel, designated 

herein, is labeled “San José,N-Arg X.25.Lindner. D.ChacoExped [black margin]/Paralimna sticta Hend. [handwritten] 

F. Hendel det./Type. Hendel 1930 [handwritten; red ink; label 

with a black border]/SMN/Lectotype ♂ Paralimna sticta Hendel, designated by Ale-Rocha&Mathis [red].” The lectotype 

is double mounted (minuten pin in a rectangular card), is in 

good condition (right antenna and some setae missing, wing 

slightly torn; abdomen not dissected), and is deposited in the 

SMN. We also examined two male paralectotypes from the 

same locality.

Other Specimens Examined. ARGENTINA. 

Formosa: Lapango, Pilcomayo (25°22′S, 57°39.3′W), Sep 1925, 

E. Lindner (1♀; SMN).

PARAGUAY. Cordillera: San Bernardino (25°16′S, 

57°19.4′W), K. Fiebrig (1♀; ANSP).

Type Locality. Neotropical. Brazil. Mato Grosso do 

Sul: Corumbá (19°0.6′S, 57°39.3′W).
number 643 • 9 7

Distribution. (Map 25) Neotropical: Argentina 

(Formosa), Brazil (Mato Grosso do Sul), Paraguay (Cordillera).

Remarks. Like P. crinita sp. nov. and P. pleurivittata,

this species has an almost horizontal frons that is slightly projected above the antennal bases. In P. taurus, however, the eye 

is rounded, the pedicel lacks a dorsal, silvery-white spot (somewhat rectangular eye, wider than high in P. pleurivittata; pedicel 

with a dorsal silvery white spot in P. pleurivittata and P. crinita

sp. nov.), and the ventral surface of the male forefemur is not 

concave (concave near apex in P. crinita sp. nov.). Structures of 

the male terminalia of these species are quite different and are 

useful to distinguish between them.

FIGURES 134–137. Structures of male terminalia of Paralimna (Paralimna) taurus Cresson (Brazil. Mato Grosso do Sul): (134) 

epandrium, cerci, and presurstyli, posterior aspect; (135) postsurstylus, lateral aspect; (136) aedeagus, phallapodeme, and hypandrium, lateral aspect; (137) same, ventral aspect. Scale bar = 0.1 mm. (See Figures 2–5 for labels of structures.)
9 8 • smithsonian contributions to zoology

MAP 25. Distribution map for Paralimna (Paralimna) taurus Cresson.

32. Paralimna (Paralimna) texana Cresson

Figures 138–141, 196, Map 26

Paralimna texana Cresson 1915:69 [USA. Texas. Travis: Austin (30°16′N, 

97°44.6′W); HT ♂, USNM]; 1946:229 [review].—Wirth 1965:748 

[Nearctic catalog]; 1968:15 [Neotropical catalog].—Mathis and Zatwarnicki 1995:125–126 [world catalog].

Diagnosis. This species is distinguished from congeners by the following combination of characters: general aspect 

of body densely whitish to yellowish gray; face silvery to yellowish gray with 3 golden-brown spots, 1 slender linear spot on 

carina and 2 paired, small, rounded, very weak golden-brown 

spots between largest facial seta; mesonotum predominantly silvery to yellowish gray with 2 pale brown stripes between dorsocentral rows and few small pale brown spots on intra- and 

postalar areas; forefemur of male with ventral surface concave, 

anteroventral row with long and conspicuous yellow setae, flattened on distal 2⁄3 of forefemur, basal 2⁄3 of posteroventral surface 

with row of long and robust setae, sometime discontinuous, longer than width of femur; foretibia slightly arcuate with ventral 

surface concave in middle; tergite 2 whitish gray for the most 

part with a pared small brown spot on 1⁄4 anterior, tergites 3–4 

with a narrow band on 1⁄3 anterior and a slender, medial, faded 

line golden brown; tergite 5 in male wholly whitish gray and 

in the female with a narrow band on 1⁄4 anterior and a slender, 

medial, faded line golden brown; insertion of abdominal setae 

brown, abdomen with mottled appearance.

Description. Body length 3.5–4.7 mm; body generally pale colored, pale brown dorsally, lateral surfaces whitish 

gray, sometimes yellowish gray.

Head: (See Figure 196) Frons golden brownish gray, 

more gray on anterior margin, with silvery spots at lateral margins 

of ocellar triangle, in front of anterior ocellar seta, not reaching 

the anterior margin of frons, between inner vertical and interfrontal setae, and on fronto-orbits above dorsal proclinate fronto-orbital seta, between proclinate fronto-orbital setae, and just above 

antennal bases; ventral fronto-orbits with a dark grayish-brown 

spot and dorsal parafacial with a dark gray spot lateral to antenna, separated by silvery-gray spot. Antennal scape and pedicel brown, with golden-gray microtomentum; basal flagellomere 

pale brown to brown according to incidence of light, with slender, 

elongate, pale setae on margin; arista with 7–9 long, dorsal rays. 
number 643 • 9 9

Postcranium with 2 complete series of postocular setae. Face silvery to yellowish gray with 3 golden-brown spots, 1 slender linear 

spot on carina, shorter than the length of antenna, and 2 paired, 

small, rounded, very weak golden-brown spots between largest facial seta; parafacial and gena silvery gray; gena with only few setae 

on middle and ventral margin; clypeus silvery gray with yellowish 

gray distal margin. Eye rounded, about as wide as high, height 

more than twice the height of gena. Gena high, height more than 

twice the length of basal flagellomere; gena-to-eye ratio 0.35–0.41.

Thorax: Mesonotum predominantly silvery to yellowish 

gray with 2 wide pale brown stripes between dorsocentral rows, 

plus few small pale brown spots on intra- and supra-alar areas; 

postpronotum, notopleuron, and supra-alar areas silvery to yellowish gray; scutellum brown dorsally, with apex dark brown and 

silvery gray laterally. Pleurae with sparse silvery-gray microtomentum, anepisternum with bases of setae pale brown, without distinct 

spot. Wing hyaline; veins pale brown; crossveins r–m and dm–cu 

with smooth gray infuscate halo; costal-vein ratio 0.35–0.38; Mvein ratio 0.97–1.12. Legs pale brown with sparse whitish- to 

yellowish-gray microtomentum; tarsi lighter with some yellow coloration, especially ventrally; forefemur with ventral surface concave, anteroventral series of long and conspicuous yellow setae, 

flattened on distal 2⁄3 of forefemur, basal 2⁄3 of posteroventral surface 

with a row of long and robust setae, longer than width of femur; 

foretibia with ventral surface concave in middle, slightly arcuate.

Abdomen: Tergites distinctly bicolored; tergite 1 with 

central region brown with laterodistal corners whitish gray, tergite 

2 whitish gray for the most part with a paired, small brown spots 

on anterior margin, tergites 3–4 with a narrow band on anterior 

1⁄3 and a slender, medial, faded line golden brown; tergite 5 in male 

wholly whitish gray and in the female with a narrow band on anterior 1⁄4 and a slender, medial, faded, golden brown line; insertion 

of abdominal setae brown, abdomen with mottled appearance. 

Male terminalia: (Figures 138–141) Presurstylus bifurcate, basoventral process short, rather triangular, about 1⁄3 length horizontal process, with dense brush of setae on apex, horizontal process 

rather robust, shallowly curved; in lateral view postsurstylus widest medially, with medial, anteriorly produced, pointed lobe bearing elongate and robust setae, some small, slender setae over the 

medial region, a medial developed projection pointing downward 

on posterior margin; lateral aedeagal process thin, about 2⁄3 length 

of aedeagus in lateral view; aedeagus slender, gradually tapered to 

membranous apex in posterior view; phallapodeme broadly triangular in lateral view; hypandrium rather deeply concave.

Type Material. The holotype male of Paralimna 

texana is labeled “Austin Oct. [19]00 Tex[as.]/♂/Holo-TYPE 

Paralimna TEXANA E. T. Cresson, Jr [maroon; “Paralimna 

TEXANA” handwritten]/ALMelander Collection 1961/Type No 

70634 U S N M [red; “70634” handwritten].” The holotype is 

directly pinned, is in very good condition (left wing partially torn; 

abdomen not dissected), and is deposited in the USNM (70634).

Other Specimens Examined. Nearctic. MEXICO. Aguascalientes: Aguacalientes (21°52.9′N, 102°17.3′W), 

3–5 Aug 1963, P. J. Spangler (1♂, 1♀; USNM). Durango:

Durango (24°01.7′N, 104°39.2′W), 3 Aug 1972, R. Mangan, 

T. P. Sluss (1♂, 1♀; USNM). Morelos: Cuernavaca (18°56.1′N, 

99°13.9′W), Jul 1965, N. L. H. Krauss (1♀; USNM).

UNITED STATES. Arizona. Cochise: Herb Martyr 

(31°52.4′N, 109°14.1′W; 2133 m), 7 Jun–1 Jul 1969, 1973, 

W. N. Mathis, S. L. Wood (3♂, 4♀; USNM); John Hands Park 

(31°52.6′N, 109°13.4′W), 12 Sep 1958, H. V. Weems (1♀; 

USNM); Pinery Canyon (31°56′N, 109°16.3′W; 1825 m), 21 

Jun 1919 (1♂, 1♀; ANSP); Portal (8 km SW; Southwest Research 

Station; 31°53′N, 109°12.3′W; 1545 m), 5–25 Sep 1965, C. W. 

Sabrosky (3♂, 3♀; USNM); Sunnyside Canyon (31°20.4′N, 

110°29.5′W), 8–12 Jul 1940, D. G. Hall (2♂, 2♀; ANSP).

Florida. Marion: Salt Springs (29°21′N, 81°44.1′W), 14 

Apr 1989, D. and W. N. Mathis (5♂, 4♀; USNM); Salt Springs 

(8 km N; 29°29.7′N, 81°44.1′W), 14 Apr 1989, D. and W. N. 

Mathis (1♂, 2♀; USNM).

Nebraska. Cherry: Big Alkali (42°38.3′N, 100°36.7′W), 2 

Jun 1969, W. W. Wirth (1♂; USNM); Snake River (42°40.5′N, 

100°51′W; river margin), 2 Jun 1969, W. W. Wirth (3♂; USNM). 

Lancaster: Lincoln (40°48.6′N, 96°40.9′W; salt lake), 19 Jun 

1969, W. W. Wirth (1♂, 3♀; USNM).

Nevada. Clark: Las Vegas (36°10.5′N, 114°30′W; 600 m), 

15–20 May 2004, D. and W. N. Mathis (1♂, 1♀; USNM); Las 

Vegas Springs (36°10.4′N, 115°11′W), 11 May 2001, D. and 

W. N. Mathis (3♂, 8♀; USNM).

New Mexico. Arriba: Espanola (35°59.5′N, 106°04.9′W), 

11 Aug 1953, W. B. Heed (1♂, 1♀; USNM). Colfax: Cimarron (36°30.4′N, 104°55.5′W; river margin), 26 May 1969, 

W. W. Wirth (7♂, 4♀; USNM). Grant: Mimbres River (NM Hwy. 

61 & Royal John Mine Road; 32°43.8′N, 107°52′W; 1665 m), 21 

Aug 2009, W. N. Mathis (1♀; USNM). Otero: Cloudcroft (4 km 

E; 32°57.7′N, 105°41.6′W), 28 Jun 1973, W. N. Mathis (17♂, 

29♀; USNM). Sandoval: La Cueva (Junction of Hwys. 126 & 4; 

35°52′N, 106°38.4′W; 2342 m), 6–7 Aug 2007, D. and W. N. 

Mathis (4♂; USNM); Valles Caldera National Preserve (E Fork 

Jemez River; 35°51′N, 106°29.5′W; 2580 m), 6 Aug 2007, D. 

and W. N. Mathis (14♂, 1♀; USNM); Valles Caldera National 

Preserve, Rincon de los Soldados (stock pond 2; 35°54.6′N, 

106°25.3′W; 2727 m), 14 Jun–6 Aug 2008, 2011, D. and W. 

N. Mathis (1♂, 1♀; USNM); Valles Caldera National Preserve, 

Rincon de los Soldados (stock pond 1; 35°52.7′N, 106°26.2′W; 

2610 m), 6 Aug 2008, D. and W. N. Mathis (2♂; USNM); Valles 

Caldera National Preserve, San Antonio Creek (35°58.3′N, 

106°34.7′W; 2564 m), 5 Aug 2008, D. and W. N. Mathis (1♂; 

USNM); Valles Caldera National Preserve, Union Building (1.6 

km N; pond; 35°53.3′N, 106°34.9′W; 2666 m), 5 Aug 2008, D. 

and W. N. Mathis (1♂; USNM). San Miguel: Montezuma (0.6 km 

NE; 35°39.3′N, 105°17.4′W), 26 Jun 1973, W. N. Mathis (44♂, 

71♀; USNM). Socorro: Socorro (34°03.5′N, 106°53.5′W), 1916, 

S. W. Williston (1♀; ANSP). Union: Des Moines (36°45.7′N, 

103°50′W), 18 Jun 1910, F. C. Bishopp (1♂; ANSP).

Oklahoma. Alfalfa: Great Salt Plains (36°45′N, 98°15.1′W), 

22 May 1969, W. W. Wirth (1♂; USNM). Garfield (36°26.5′N, 

97°52.2′W): 9 Jul 1977, J. F. Reinert (1♀; USNM). Marshall:

Texoma Lake (53°33.7′N, 96°40.5′W), 15 Jul 1954, J. G. Chillcott 

(1♀; CNC).
100 • smithsonian contributions to zoology

South Dakota. Mellette: Little White River (43°33.7′N, 

100°44.5′W; river margin), 4 Jun 1969, W. W. Wirth (1♀; 

USNM). Yankton: Gavins Point (42°51.7′N, 97°33.5′W), 17 Jun 

1969, W. W. Wirth (1♀; USNM).

Texas. Aransas: Aransas National Wildlife Refuge 

(28°06.9′N, 96°48′W), 20 May 1972, W. W. Wirth (2♀; 

USNM); Salt Creek (28°15.4′N, 96°56.5′W), 5 Jun 1953 

(1♀; USNM). Blanco: Davis Ranch (NW Blanco; 30°27′N, 

98°32.9′W), 26 Apr 1959, J. F. McAlpine (12♂, 8♀; CNC); 

Miller Creek (30°15.2′N, 98°31.7′W; 410 m), 3 Jun 2004, 

W. N. Mathis (4♂; USNM). Brewster: Big Bend National Park, 

Boquillas Crossing (29°11.4′N, 102°56.8′W; 550 m), 8 May 

1959, J. F. McAlpine (28♂, 21♀; CNC); Big Bend National 

Park, Pine Canyon (29°15.3′N, 103°12.6′W; 1225–1830 m), 

4 May 1959, J. F. McAlpine (3♂, 1♀; CNC); Big Bend National Park, Chisos Basin (29°16.5′N, 103°18.1′W), 29 May 

1959, J. F. McAlpine (1♂, 1♀; CNC). Gillespie: Spring Creek 

(17 km E Harper; 30°18.2′N, 99°03.4′W; 580 m), 3 Jun 2004, 

FIGURES 138–141. Structures of male terminalia of Paralimna (Paralimna) texana Cresson (United States. Texas): (138) 

epandrium, cerci, and presurstyli, posterior aspect; (139) postsurstylus, lateral aspect; (140) aedeagus, phallapodeme, 

and hypandrium, lateral aspect; (141) same, ventral aspect. Scale bar = 0.1 mm. (See Figures 2–5 for labels of structures.)
number 643 • 101

W. N. Mathis (3♂; USNM). Gonzales: Guadalupe River 

(29°29.8′N, 97°27.3′W), 22 Apr 1956, W. W. Wirth (3♂, 

1♀; USNM). Jeff Davis: Fort Davis (30°55.3′N, 103°53.7′W; 

1525 m), 1 Jun 1959, J. F. McAlpine (1♀; CNC). Jim Wells:

Mathis (7.5 km S; Nueces River; 28°02.2′N, 97°52.2′W; 

15 m), 6 Jun 2004, W. N. Mathis (12♂, 1♀; USNM). Kerr:

Adam Wilson’s Cave (6.5 km SE Hwy. 39, 60 km SSW Kerrville; 29°55.6′N, 99°14.2′W), 31 Mar 1959, J. F. McAlpine 

(2♀; CNC); Kerrville (30°02.8′N, 99°08.4′W), 1 Apr 1959, 

J. F. McAlpine, J. R. Vockeroth (4♂, 14♀; CNC). Kimble:

Junction (South Llano River; 30°29.6′N, 99°45.1′W; 510 m), 

4 Jun 2004, W. N. Mathis (9♂, 2♀; USNM); Llano River 

(30°27′N, 99°49′W), 23 May 1972, W. W. Wirth (6♂, 1♀; 

USNM). Kinney: Del Rio (30 km E; Pinto Creek; 29°20′N, 

100°31.6′W), 5 Jun 2004, W. N. Mathis (7♂, 4♀; USNM). 

Real: Leakey, Rio Frio (29°37.6′N, 98°40.4′W), 23 May 

1972, W. W. Wirth (8♂, 4♀; USNM). San Patricio: Mathis 

(6.5 km S; Nueces River; 28°02.8′N, 97°51.8′W; 18 m), 6 

Jun 2004, W. N. Mathis (12♂; USNM); Mathis (5 km SW at 

Nueces River; 28°02.4′N, 97°51.5′W), 9 Dec 1984, B. Gill, 

W. E. Steiner, D. Whitehead (22♂, 35♀; USNM); Sinton (12 

km NE; Welder Wildlife Refuge; 28°06.1′N, 97°22′W), 8 

Dec 1984, B. Gill, W. E. Steiner, D. Whitehead (16♂, 19♀; 

USNM). Terrell: Lozier Canyon (29°48.1′N, 101°48.2′W), 

14 Jun 1992, R. W. Baumann, L. Liu (6♂, 12♀; BYU). Travis: Austin (Zilker Park; 30°15.8′N, 97°46.3′W), 2 Jun 2004, 

W. N. Mathis (5♂, 1♀; USNM); South Austin (Slaughter Park; 

30°11.2′N, 97°51.6′W; 235 m), 3 Jun 2004, W. N. Mathis 

(6♂; USNM); Austin (paratype 1♀; ANSP); 6–20 Oct 1901 

(paratype 2♂; ANSP); 7 Oct 1889 (2♀; paratypes; ANSP, 

USNM); 6 Oct 1901 (paratype 1♂; ANSP); 2 Mar 1900 (1♂; 

paratype; ANSP, 1♀ USNM). Val Verde: Comstock (16 km 

N; Pecos River; 29°53.7′N, 101°09.1′W), 28 Apr–4 Jun 1959, 

2004, W. N. Mathis, J. F. McAlpine (7♂, 8♀; CNC, USNM); 

Devils River (29°56.2′N, 100°56.6′W; 550 m), 26 Apr 1959, 

J. F. McAlpine (2♂, 1♀; CNC).

Utah. San Juan: Johnson Creek meadow (22 km N Blanding: 37°47.5′N, 109°30.5′W; 2384 m), 25 Aug 2014, D. and 

W. N. Mathis (1♂; USNM).

Type Locality. Nearctic: United States. Texas. Travis: Austin (30°16′N, 97°44.6′W).

Distribution. (Map 26) Nearctic: Mexico (Aguacalientes, Durango, Morelos). United States (Arizona, Florida, Nebraska, 

Nevada, New Mexico, Oklahoma, South Dakota, Texas, Utah).

MAP 26. Distribution map for Paralimna (Paralimna) texana Cresson.
102 • smithsonian contributions to zoology

Remarks. This species is similar to and has been confused with P. thomae but can be distinguished from the latter by 

the coloration of the frons, scutum, and abdomen; the infuscation on the crossveins; the setulae of the male forefemur; and the 

structures of the male terminalia. Additional details are in the 

“Remarks” for P. thomae.

33. Paralimna (Paralimna) thomae

(Wiedemann)

Figures 142–145, 181, Map 27

Ephydra thomae Wiedemann 1830:593.

Paralimna thomae.—Cogan and Wirth 1977:335 [generic combination].

Paralimna multipunctata Williston 1896:390.—Grossbeck 1912:378 [types 

in AMNH].—Wirth 1956:17 [review, Bahamas].—Woodruff et al. 

1998:68 [list, Grenada]. NEW SYNONYM.

Paralimna (Paralimna) multipunctata.—Cresson 1946:229 [review]; 

1947:50 [review, Neotropical species].—Wirth 1965:748 [Nearctic catalog]; 1968:15 [Neotropical catalog].—Mathis and Edmiston 

1991:832 [review of Williston’s St. Vincent species].—Mathis and 

Zatwarnicki 1995:122 [world catalog].—Mathis 1997:54 [review, 

Belize].

Paralimna ciliata Cresson 1916:111 [Costa Rica. Cartago: Cartago; HT ♂, 

ANSP (6091)]; 1918:46 [review; Costa Rica]; 1946:229 [synonymy]; 

1947:50.—Thaxter 1917:671–675 [parasites: Stigmatomyces curvirostris Thaxter, S. paralimnae Thaxter (Laboulbeniaceae)]; 1917:676; 

1931:145–155 [parasites: S. jamaicensis Thaxter, S. gracilior Thaxter, 

S. recurvatus Thaxter, S. rostratus Thaxter (Laboulbeniaceae)]; Curran 

1928:60 [list, Puerto Rico, St. Thomas].

Diagnosis. This species is distinguished from congeners by the following combination of characters: body predominantly gray, dorsum, especially frons and mesonotum, mostly 

grayish brown to brown, lateral surfaces mostly gray, sometimes 

silvery white; frons mottled, dark to whitish brown; scutellum 

guttate, mostly brown, especially base of setae; anepisternum 

and katepisternum mostly concolorous (anepisternum with dorsal margin usually brownish), faintly yellowish gray to silvery 

gray; wing mostly hyaline; crossvein dm–cu lacking infuscate 

halo; forefemur of male bearing anteroventral comb-like row 

of distinctly flattened, long setae; and posteroventral surface of 

male forefemur bearing patch of long setae at basal 1⁄3; lateral 

aedeagal process about 1⁄2 length of aedeagus, robust, curved in 

middle, rounded apex.

Description. Body length 2.3–3.6 mm; body generally gray, some pale brown spots dorsally, lateral surfaces yellowish gray, sometimes whitish gray.

Head: (See Figure 181) Frons pale brown with linear 

pale yellow spots at lateral margins of ocellar triangle and in 

front of anterior ocellar seta, short, not reaching the anterior 

margin of frons, between inner vertical and interfrontal setae, 

and on fronto-orbits above dorsal proclinate fronto-orbital seta, 

between proclinate fronto-orbital setae, and just above antennal 

bases; ventral fronto-orbits and dorsal parafacial with 2 small, 

inconspicuous pale brown spots lateral to antenna, separated by 

yellow spot. Antenna pale brown, covered by dense yellow microtomentum; basal flagellomere with slender, elongate, pale setae 

on margin; arista with 11 long, dorsal rays. Postcranium with 2 

complete series of postocular setae, the second series irregular. 

Face yellowish gray to golden with a single pale golden-brown 

spot on the carina, shorter than the length of antenna; parafacial 

silvery gray and gena yellowish gray; gena with only few setae on 

middle and ventral margin; clypeus yellowish gray with silvery 

gray distal margin. Eye rounded, slightly higher than wide, height 

about twice the height of gena. Gena high, height subequal to 

half the length of basal flagellomere; gena-to-eye ratio 0.36–0.47.

Thorax: Mesonotum mostly pale brown with yellowishgray areas, usually along setal tracks, 2 slender, widely separated, 

pale brown stripes between dorsocentral rows, plus 1 short stripe 

on supra-alar area; insertion of setae brown giving a spotted aspect 

to scutum; postpronotum, notopleuron, and supra-alar areas yellowish gray; scutellum brown basally, with apex blackish brown 

and yellowish gray otherwise. Pleurae yellowish gray, anepisternum usually with dorsal margin brownish, sometimes without 

spots. Wing hyaline; veins pale brown, crossveins without infuscate halos; costal-vein ratio 0.40–0.42; M-vein ratio 0.97–1.05. 

Legs blackish brown with dense silvery-gray to yellowish-gray microtomentum; tarsi lighter with some yellow coloration, especially 

ventrally; forefemur of male with anteroventral series of long and 

wide flattened setae, as long as width of femur, anteroventral surface with a comb of long and robust setae near base; foretibia of 

male arcuate, with a groove ventrally.

Abdomen: Tergites distinctly bicolored, dark brown 

along anterior 2⁄3 and medially, otherwise yellowish gray, posterior 

margin of brown band convex on both sides and with a conspicuous medial stripe extended to posterior margin. Male terminalia: 

(Figures 142–145) Presurstylus bearing somewhat elongate, slender setae, bifurcate, with basoventral process weakly developed, 

length more than 1⁄3 of horizontal process length; horizontal process slightly curved; in lateral view postsurstylus robust, widest 

medially with medial, anteriorly produced, somewhat pointed 

lobe bearing sparse, slender, short setae, distal portion of postsurstylus widened; lateral aedeagal process about 1⁄2 length of aedeagus, robust, curved in middle, rounded apex; aedeagus rather 

robust tapering gradually toward membranous apex; in lateral 

view phallapodeme triangular, prolonged distally; hypandrium 

moderately deeply concave.

Type Material. The lectotype of Ephydra thomae

Wiedemann, designated herein, is labeled “E[phydra]. thomae 

Wied. Isle S. Thomas [H. B.] Hornbeck/LECTOTYPE ♂ Ephydra thomae Wied. det. B. H. Cogan 1978[“8” handwritten; 

this lectotype designation was never published]./Paralimna 

thomae ♂ (Wied.) det. B. H. Cogan 197 ./Mus. Westerm[ann]. 

[black submargin across top and left side of label]/TYPE [red]/

ZMUC 00025029/LECTOTYPE ♂ Ephydra thomae Wiedemann, designated by Ale-Rocha&Mathis [red].” The lectotype 
number 643 • 103

is directly pinned, is in good condition (abdomen not dissected), 

and is deposited in the ZMUC. We also examined a male paralectotype, designated herein and also from St. Thomas.

The lectotype male of Paralimna multipunctata (designated 

by Mathis and Edmiston 1991:832) is labeled “Co-type [circular 

label with a yellow border]/Leeward side St. Vincent, W.I. H. H. 

Smith./W.Indies. 1907-66./Paralimna multipunctata Will [handwritten, faint red ink] /LECTOTYPE Paralimna multipunctata 

Will. ♂ By W. N. Mathis [handwritten except for “LECTOTYPE” and “By”, black submarginal border].” The lectotype 

is double mounted (pin in a rectangular piece of cardboard), is 

in good condition, and is deposited in the BMNH. There are 

also 10 paralectotypes as follows: BMNH (2♂, 2♀, 1ex), KU 

(1♂, 1♀), AMNH, (3♀, 20329a, 20327, 20327a). In the original description, Williston indicated that there were “numerous 

specimens” in the type series.

FIGURES 142–145. Structures of male terminalia of Paralimna (Paralimna) thomae (Wiedemann) (Grenada. St. George): (142) epandrium, cerci, and presurstyli, posterior aspect; (143) postsurstylus, lateral aspect; (144) aedeagus, phallapodeme, and hypandrium, 

lateral aspect; (145) same, ventral aspect. Scale bar = 0.1 mm. (See Figures 2–5 for labels of structures.)
104 • smithsonian contributions to zoology

Other Specimens Examined: Nearctic: MEXICO. Colima: Colima (19°14.4′N, 103°43.3′W), Sep 1965, 

N. L. H. Krauss (1♂; USNM). México: Ixtapan de La Sal 

(19°26.9′N, 99°14.7′W; 1676 m), 9 Aug 1954, J. G. Chillcott 

(2♂; CNC). Morelos: Cuernavaca (18°56′N, 99°14′W), Jul 

1965, N. L. H. Krauss (1♂; USNM); Cuernavaca (12 km E; 

18°58.1′N, 99°07.7′W; 1310 m), 14 Aug 1954, J. G. Chillcott 

(11♂, 8♀; CNC). San Luis Potosí: Xilitla (16 km E; 21°26.3′N, 

98°55.7′W; 305 m), 25 Aug 1954, J. G. Chillcott (1♂, 3♀; 

CNC); Xilitla (21°23′N, 98°59.5′W; 425 m), 25 Aug 1954, 

J. G. Chillcott (2♂; CNC). Tamaulipas: Morelos Antiguo 

(22°33.1′N, 99°05.1′W; 305 m), 21 Jul 1954, J. G. Chillcott 

(8♂, 2♀; CNC).

UNITED STATES. Arizona: Cochise: Cochise (32°06.8′N, 

109°55.3′W): 13 Jul 1919, W. Stone (1♀; ANSP).

California. Imperial: Calipatria (33°07.5′N, 115°30.8′W), 

13 Nov 1921, E. R. Kalmback (1♀; ANSP). Riverside: Riverside 

(33°57.2′N, 117°23.8′W), 7 Oct 1934, A. L. Melander (2♂, 4♀; 

ANSP). San Bernardino: Death Valley, Furnace Creek (36°27.5′N, 

116°52.2′W), 23 Apr 1935, A. L. Melander (1♀, ANSP).

Florida. Highlands: Lake Istokpoga, Windy Point Park 

(27°18.4′N, 81°18.9′W; 18 m), 15–16 Jan 2009, D. and 

W. N. Mathis (3♂, 1♀; USNM). Miami-Dade: Florida City 

(25°26.9′N, 80°28.8′W), 13 Jan 2007, D. and W. N. Mathis 

(2♂; USNM); Royal Palm Park Ranger Station (25°22.9′N, 

80°36.6′W), 28–29 Jan 1933, 1939, A. L. Melander (1♂, 1♀; 

ANSP); Miami (25°46′N, 80°12′W), 2–20 Dec 1912, F. Knab 

(2♂; ANSP).

Texas: Bexar: San Antonio (29°29.4′N, 98°29.6′W), 

Nov 1942, E. S. Ross (1♂; USNM). Cameron: Brownsville 

(25°56.5′N, 97°29.7′W), 30 Jun 1938, R. L. Sailer (1♂; USNM); 

Brownsville (N; 25°59.8′N, 97°29.7′W), 18 Jul 1933, R. Nabours, C. W. Sabrosky (1♀; ANSP).

Neotropical: BAHAMAS. Crooked Island: Landrail Point 

(22°48.2′N, 74°20.2′W), 5 Mar 1953, E. B. Hayden (1♂; 

AMNH). Eleuthera Island: Hatchet Bay (near Alicetown; 

25°21.1′N, 76°29.3′W) 2 Apr 1952, E. B. Hayden (1♀; AMNH). 

Exuma Cays: Stainard Creek (24°50.6′N, 77°53.4′W), 13 Jan 

1953, E. B. Hayden (1♀; AMNH). Grand Bahama Island: West 

End (26°41′N, 78°58′W), 24 Jun 1987, W. E. Steiner, R. Mollineaux (3♂; USNM). Long Island: Deadman′s Cay (23°10.8′N, 

75°06.7′W), 11 Mar 1954, E. B. Hayden (1♂, 8♀; AMNH). San 

Salvador: near Cockburn Town (24°03.1′N, 74°31.6′W), 18 

Mar 1953, L. Giovannoli (1♀; ANSP).

BELIZE. Corazal: Corozal (18°23.9′N, 88°23.3′W), 30 Jul 

1967 (1♂; USNM). Stann Creek: Dangriga (16°58′N, 88°13′W), 

3–4 Apr 1993, W. N. Mathis (2♂; USNM); S Sittee River, Possum Point Biological Station (16°52.1′N, 88°22.5′W), 6 Nov 

1987, D. and W. N. Mathis (2♂, 2♀; USNM); Sittee River, Possum Point Biological Station (16°52.1′N, 88°22.5′W), 25 Apr 

1987, R. Faitoute, P. J. Spangler (5♂, 5♀; USNM); 29 Apr 1987, 

R. Faitoute, P. J. Spangler, collected on river bank at Possum 

Point Biological Station (1♂; USNM); Mullins River (17 km N 

Dangriga; 17°06.2′N, 88°17.8′W), 29 Mar 1988, W. N. Mathis 

(1♀; USNM).

BOLIVIA. La Paz: Tajihui (15°40.8′S, 67°41.7′W; 590 m), 

12 Mar 2001, W. N. Mathis (2♂; USNM); Guanay (15°29.8′ S, 

67°52.7′W; 460 m), 13 Mar 2001, W. N. Mathis (19♂; USNM); 

Guanay (3 km E; 15°30.2′S, 67°52.3′W; 480 m), 13 Mar 2001, 

W. N. Mathis (8♂; USNM); Guanay (3 km E; 15°30.2′S, 

67°52.3′W; 500 m), 14 Mar 2001, W. N. Mathis (8♂; USNM); 

Guanay (1 km E; 15°30.2′S, 67°52.3′W; 580 m), 13 Mar 2001, 

S. D. Gaimari (1♂, USNM); Mapiri (5 km W; 15°18.6′S, 

68°13′W; 720 m), 15 Mar 2001, S. D. Gaimari (1♂; USNM); 

Mapiri (5 km W; 15°18.6′S, 68°13′W; 750 m), 16 Mar 2001, 

W. N. Mathis (4♂; USNM).

COLOMBIA. Tolima: Armero (04°58′N, 74°54′W; Malaise trap), 30 Jan–5 Feb 1977, E. L. Peyton (3♂;USNM).

COSTA RICA. Cartago: Cartago (9°51.4′N, 83°55.2′W), 

4 Jul 1909, P. P. Calvert (1 ♀; ANSP); 21 Nov 1909 (1♂, 1♀; 

ANSP); Cartago (near; 09°51.3′N, 83°55.2′W), 15 Dec 1909, 

P. P. Calvert (1♂, 2♀; ANSP). Guanacaste: Santa Cruz (14 km 

S; La Casona; 10°10.4′N, 85°35.7′W; 180 m), 23 Jun 2001, 

W. N. Mathis (5♂, 1♀; USNM); Cañas (10°25.6′N, 85°05.7′W), 

13 Jul 1966, S. L. Wood (1♂; BYU). Heredia: Santo Domingo 

(INBio Parque; 9°58.4′N, 84°5.6′W), 23 Jun 2001, W. N. 

Mathis (1♂; USNM). Puntarenas: Jacó (5 km E; 9°34.7′N, 

84°35.6′W), 10 Jun 2003, D. and W. N. Mathis (8♂, 1♀; 

USNM); Parryta (Playa Palo Seco; 9°29.4′N, 84°19.5′W), 12 

Jun 2003, D. and W. N. Mathis (1♀; USNM); Cabuya (9°37′N, 

85°4.8′W), Rio Lajas, 20 Jun 2001, W. N. Mathis (1♂; USNM); 

Montezuma (1 km S; 9°39′N, 85°4.3′W), 20 Jun 2001, W. N. 

Mathis (1♂; USNM).

ECUADOR. Chimborazo: Huigra (02°17′S, 78°59′W), 

Jul 1955, R. Levi-Castillo (1♀, USNM). El Oro: El Guabo 

(03°13′S, 79°47.7′W), Dec 1955, R. Levi-Castillo (2♂, 1♀; 

USNM); Puerto Bolívar (03°16′S, 79°60′W), Dec 1955, R. LeviCastillo (1♂, 2♀; USNM). Guayas: Balao (02°55′S, 79°49′W), 

Dec 1955, R. Levi-Castillo (1♂, 4♀; USNM); Taura (02°18.6′S, 

79°43.9′W), Dec 1955, R. Levi-Castillo (7♂, 8♀; USNM). Loja:

Catamayo (03°59′S, 79°21′W), Dec 1955, R. Levi-Castillo (1♀; 

USNM). Manabí: Pichincha (01°02.7′S, 79°49.2′W), Aug 1955, 

R. Levi-Castillo (1♂; USNM). Pastaza: Río Puyo (01°29.1′S, 

77°59.7′W), R. Levi-Castillo (1♂; USNM).

EL SALVADOR. Sonzacate (13°44′N, 89°43′W), 25 Jun 

1958, L. J. Bottimer (1♂; USNM).

GUATEMALA. Escuintla: San José (13°55.7′N, 

90°49.3′W), 20 Jul 1943, D. G. Hall (1♂; USNM). Izabal: Cayuga (15°31.9′N, 88°41.8′W), Oct 1915 (1♂; ANSP). Zacapa:

Gualán (15°06.9′N, 89°21.7′W), 15 Feb 1905 (1♂, 2♀; ANSP).

GUYANA. Georgetown (06°48.6′N, 58°08.6′W), 20–29 

Aug 1997, W. N. Mathis (1♂; USNM). Karanambo, Rupununi River (03°45.1′N, 59°18.6′W; ox bow), 2 Apr 1994, W. 

Mathis (11♂, 1♀; USNM). Karanambo, Maricuba (03°45.1′N, 

59°18.6′W; pond), 31 Mar–2 Apr 1994, W. N. Mathis (15♂, 

5♀; USNM). Mahaica (06°42.8′N, 57°55.5′W), 22 Apr 1995, 

W. N. Mathis (4♂, 1♀; USNM). Mahaica (3 km W; 06°43.5′N, 

57°56.6′W), 14 Apr 1994, W. N. Mathis (4♂, 1♀; USNM). 

Pirara Ranch, Cashew Creek (03°37.3′N, 59°44.2′W), 23 Apr 

1995, W. N. Mathis (12♂, 2♀; USNM). Pirara Ranch and River 
number 643 • 105

(03°32.1′N, 59°40.5′W), 24–25 Apr 1995, W. N. Mathis (4♂, 

1♀; USNM).

HONDURAS. Cortés: Omoa (16°47.8′N, 87°58.4′W), 26 

Sep 1995, D. and W. N. Mathis (10♂; USNM); San Pedro Sula 

(8 km S; 15°25.7′N, 88°01.4′W), 25–26 Sep 1995, D. and W. N. 

Mathis (10♂; USNM).

MEXICO. Chiapas: Río Izapa (14°55.4′N, 92°10.7′W), 

21 Apr 1983, W. N. Mathis (1♀; USNM); Puente Macalapa 

(16°30′N, 92°30′W; light trap), 22 May 1964, F. S. Blanton (3♂, 

5♀; USNM); Tuxtla Gutiérrez (16°44.8′N, 93°06.6′W), 2 Aug 

1969, D. Kritsch (1♀; CNC). Guerrero: Acapulco (16°51.8′N, 

99°52.9′W), 6 Aug 1954, J. G. Chillcott (1♂; CNC). Oaxaca: 

Tehuantepec (16°19′N, 95°14.5′W), 15–16 Jul 1964, P. J. 

Spangler (1♂; USNM). Tabasco: Teapa (8 km SW; 17°27.2′N, 

93°01.2′W), 6 May 1985, W. N. Mathis (6♂, 1♀; USNM). Veracruz-Llave: Fortin de las Flores (18°54′N, 97°W; 952 m), 2 

May 1985, W. N. Mathis (17♂, 6♀; USNM); Puente Nacional 

(19°19.7′N, 96°29′W), 23–24 Jul 1965, O. S. Flint Jr., A. Ortiz 

(1♀; USNM).

NICARAGUA. León: La Cruz de la India (5 km SW, on 

Hwy. 26; 12°43.2′N, 86°19.8′W; 215 m), 22 Jun 2007, N. E. 

Woodley (7♂, 11♀; USNM).

PANAMÁ. Cocle: Playa Santa Clara (08°22.4′N, 

80°06.3′W), 2 Jul 1967, W. W. Wirth (3♂, 2 ♀; USNM). Colón:

Mojingas Swamp, Fort Sherman (09°21.8′N, 79°57.2′W), Jan 

1953 (1♂; USNM). Darién: El Real (08°07′N, 79°44′W), 19 Mar 

1953, F. S. Blanton (1♀; USNM). Herera: Chitre (07°57.9′N, 

80°25.3′W), Jul 1981, N. L. H. Krauss (1♀; USNM).

PERU. Lima: Cocachacra (18 km E Chosica; 11°55.3′S, 

76°32′W), 11 Feb 1984, W. N. Mathis (1♂; USNM); Lima, Laguna de Villa (12°03.3′S, 77°03′W), 30 Aug 1988, W. N. Mathis 

(1♂; USNM). Loreto: Indiana (15 km NE; 03°30′S, 73°02.7′W), 

20 Feb 1984, W. N. Mathis (3♂; USNM). Madre de Dios: Río 

Manu, Pakitza (11°56.6′S, 71°16.9′W; 250 m), 9–23 Sep 1988, 

W. N. Mathis (1♂, 1♀; USNM).

TRINIDAD and TOBAGO. Tobago. St. John: Charlotteville (beach; 11°19.5′N, 60°32.9′W), 16 Apr–16 Jun 1993, 

1994, D. and W. N. Mathis (16♂; USNM); Speyside (Doctor 

River; 11°18.2′N, 60°31′W), 19 Apr 1994, D. and W. N. Mathis 

(3♂, 2♀; USNM); Speyside (Doctor River; 1 km NW; 11°18′N, 

60°32′W), 12–13 Jun 1993, W. N. Mathis (2♂; USNM); Charlotteville (5 km S; 11°18.9′N, 60°34.5′W), Hermitage River and 

beach, 22 Apr–11 Jun 1993, 1994, D. and W. N. Mathis (6♂, 1♀; 

USNM). St. Paul: Delaford, Kings Bay (11°16′N, 60°32.8′W), 

13 Jun 1993, W. N. Mathis (8♂; USNM); Kendall (11°14.3′N, 

60°35.7′W), 21 Apr 1994, W. N. Mathis (9♂; USNM). Trinidad. Caroni: Caroni (3 km W; 10°36.9′N, 61°24.7′W), 22 Mar 

1985, G. F. and J. F. Hevel (13♂, 10♀; USNM). St. Andrew:

Ventral Manzanilla (5 km S; 10°28′N, 61°03′W), 20 Jun 1993, 

W. N. Mathis (5♂, 1♀; USNM).

VENEZUELA. Carabobo: Valle Seco (10°30.5′N, 

67°35.3′W), Jan 1940, P. Anduze (1♀; ANSP). Lara: Barquisimeto (S Medina; 10°03.8′N, 69°19.3′W), 7 Jun 1965, I. 

Gaud, L. F. Martorel (3♂; USNM). Zulia: Rosario (10°01.8′N, 

71°13.8′W), 14 Jun 1976, A. S. Menke, D. Vincent (1♂; USNM).

WEST INDIES. BARBADOS. Christ Church: Graeme Hall 

Swamp (13°04.2′N, 59°34.7′W), 31 Aug–2 Sep 1997, W. N. 

Mathis (5♂; USNM); Rockley Beach (13°04.3′N, 59°35.2′W), 21 

May–11 Sep 1996, 1997, D. and W. N. Mathis, H. B. Williams 

(27♂, 2♀; USNM). St. Andrew: Bawdens Ponds (13°14.8′N, 

59°34.9′W), 2 Sep 1997, W. N. Mathis (1♂; USNM); Baxters (13°13.2′N, 59°34.1′W), 21 May 1996, D. and W. N. 

Mathis, H. B. Williams (1♂; USNM); Belleplaine (13°14.8′N, 

59°33.6′W), 21 May–1 Sep 1996, 1997, D. and W. N. Mathis, 

H. B. Williams (11♂, 2♀; USNM); Long Pond (13°15.1′N, 

59°33.3′W), 1 Sep 1997, W. N. Mathis (5♂; USNM). St. Joseph: Joes River (13°12.8′N, 59°32.3′W), 10 Sep 1996, W. N. 

Mathis (5♂, 1♀; USNM). St. Peter: Six Men’s Bay (13°16.5′N, 

59°38.8′W), 22 May–12 Sep 1996, D. and W. N. Mathis, H. B. 

Williams (10♂, 1♀; USNM). St. Philip: Gemswick (13°05′N, 

59°28.5′W), 31 Aug 1997, W. N. Mathis (4♂; USNM). St. 

Thomas: Farmers (13°12.8′N, 59°35.5′W), 2 Sep 1997, W. N. 

Mathis (5♂; USNM).

BRITISH VIRGIN ISLANDS. Guano Island (18°27.9′S, 

64°34.3′W; 0–80 m; North Bay, Coccoloba forest, sea level; 

Malaise trap), 13–26 Jul 1986, S. E. Miller, M. G. Pogue (1♂: 

USNM).

CUBA. Cienfuegos: Soledad, Jardín Botánico (22°7.5′N, 

80°19.2′W), Jan–13 Dec 1927, 1994, C. T. and B. B. Brues, 

W. N. Mathis (9♂, 4♀; ANSP, USNM); Topes de Collantes (5 

km WNW; 21°56.5′N, 80°2.3′W; 600 m), 11 Dec 1994, W. N. 

Mathis (1♀; USNM). Guantánamo: Guantánamo (20°08.6′N, 

75°12.5′W), 10 Nov 1914, H. Skinner (7♂, 4♀; ANSP). Havana: Havana (beach; 23°5.8′N, 82°27.7′W), 2–14 Dec 

1994, W. N. Mathis (11♂; USNM); Ojo de Agua (22°54.6′N, 

82°29.1′W), 8 Dec 1994, W. N. Mathis (21♂, 4♀; USNM); 

Playa Jibacoa (57 km E Havana; 23°08.9′N, 81°51′W), 26 

Apr 1983, W. N. Mathis (2♀; USNM). Holguín: Holguín (N; 

20°55.5′N, 76°15.8′W), Feb 1992, M. von Tschirnhaus (2♂, 8♀; 

USNM). Mayabeque: Palmarejo (67 km E Havana; 23°08.5′N, 

81°40.6′W), 26 Apr 1983, W. N. Mathis (20♂, 1♀; USNM); 

Puerto Escondido (18°19.6′N, 71°35′W; 1370 m), 26 Apr 1983, 

W. N. Mathis (11♂, 10♀; USNM). Matanzas: Buena Ventura 

(7 km N; 22°18.1′N, 81°11.7′W), 1 May 1983, W. N. Mathis 

(9♂, 4♀; USNM); Palpite (1 km NE; 22°20.2′N, 81°10.8′W), 2 

May 1983, W. N. Mathis (1♀; USNM) . Pinar del Rio: Pinar del 

Rio (10 km S; 22°22.7′N, 83°40.3′W), 12–23 Sep 1913 (1♂, 1♀; 

AMNH); Soroa (22°47.7′N, 83°W), 27 Apr–6 Dec 1983, 1994, 

W. N. Mathis (6♂, 6♀; USNM); Soroa (2 km NW; 22°48.6′N, 

83°1.0′W), 4–5 Dec 1994, W. N. Mathis (2♂, 2♀; USNM); 

Soroa (2 km E; 22°47.7′N, 83°W), 29 Apr 1983, W. N. Mathis 

(11♂, 5♀; USNM). Sancti Spiritus: Playa Ancón (21°44.1′N, 

79°59.9′W), 12 Dec 1994, W. N. Mathis (6♂, 5♀; USNM); 

Topes de Collantes (21°54.4′N, 80°01.4′W; 670 m), 9–11 Dec 

1994, W. N. Mathis (4♂; USNM).

DOMINICA. Cabrits Swamp (15°35′N, 61°29′W), 10 

May–19 Jun 1965, 1991, D.R. Davis, D. and W. N. Mathis, 

(10♂, 7♀; USNM). Coulibistrie (15°27.1′N, 61°26.8′W), 21 

Mar 1989, W. N. Mathis (2♂; USNM). Dublanc (15°30.9′N, 

61°28.1′W), 21 Mar 1989, W. N. Mathis (4♂, 1♀; USNM). 
106 • smithsonian contributions to zoology

Hampstead River (15°33.9′N, 61°22.3′W), 22 Mar 1989, 

W. N. Mathis (1♂; USNM). Soufrière Bay (15°13.9′N, 

61°21.6′W), 24 Mar 1989, A. Freidberg (1♂; USNM). Toucari 

(15°36.6′N, 61°27.8′W), 21 Mar 1989, W. N. Mathis (5♂, 3♀; 

USNM).

DOMINICAN REPUBLIC. Azua: Padre Las Casas (8 

km NE; Rio Las Cuevas; 18°46′N, 70°53′W; 580 m), 7 Aug–4 

Oct 1990, 1991, J. Rawlins, S. Thompson (3♂, 4♀; CMP); 

near Pueblo Viejo (18°24.8′N, 70°44.7′W), 19 May 1998, 

D. and W. N. Mathis (5♂, 3♀; USNM). Barahona: Baoruco 

(18°04.6′N, 71°05.5′W; beach and river), 19 May 1998, D. and 

W. N. Mathis (6♂; USNM); Barahona (18°12′N, 71°05.3′W), 

25 Mar 1999, W. N. Mathis (3♂; USNM); Cabral (canals E of 

Cabral; 18°15.2′N, 71°13.4′W), 16 May 1995, W. N. Mathis 

(1♂; USNM); Cortico (La Mina; 18°06.7′N, 71°13.4′W; 1300 

m), 23 Mar 1999, W. N. Mathis (2♂; USNM); Los Patos 

(17°57.6′N, 71°10.9′W), 15 May 1995, W. N. Mathis (5♂; 

USNM); Ojeda (17°58.2′N, 71°10.6′W), 22 Mar 1999, W. N. 

Mathis (4♂; USNM); Paraíso (6 km NW; Rio Nizao; 18°02′N, 

71°12′W; 170 m), 25–26 Jul 1990, J. Rawlins, S. Thompson, 

C. Young (66♂, 41♀; CMP). Dajabon: Loma de Cabrera (10 

km S; 19°20′N, 71°37′W; 650 m), 12 Jul 1992, R. Davidson, J. 

Rawlins, S. Thompson, C. Young (1♂; CMP). El Seibo: Pedro 

Sáchez (18°51.4′N, 69°6.5′W), 26 May 1998, D. and W. N. 

Mathis (3♂; USNM); Rincón (near; 18°45.3′N, 68°55.7′W), 

12 May 1995, W. N. Mathis (7♂; USNM). Hato Mayor: El 

Valle (9.5 km W; 18°59′N, 69°29′W; 170 m), 6 Jul 1992, R. 

Davidson, J. Rawlins, S. Thompson, C. Young (1♀; CMP). 

Independencia: Duvergé (2 km S; 18°22′N, 71°31.4′W), 24 

Mar 1999, W. N. Mathis (2♂, 1♀; USNM); Mella (canal W; 

18°21.5′N, 71°25.7′W), 20 May 1998, D. and W. N. Mathis 

(3♂; USNM); Puerto Escondido (18°19.6′N, 71°35′W; 1370 

m), 24 Mar 1999, W. N. Mathis (1♂; USNM). La Altagarcia: Rio Chavon (mouth; 18°23.9′N, 68°53.7′W), 29 Feb 

1992, D. A. Grimaldi, J. Stark (4♂, 9♀; AMNH). La Vega:

Constanza (9 km SW; 18°52′N, 70°43′W; road to Pinar Bonito; 1433 m), 29 Jul 1991, D. A. Grimaldi, J. Stark (1♀; 

AMNH); Constanza (13.7 km SE; 18°51.5′N, 70°41.9′W; 

1505 m), 7 May 1995, W. N. Mathis (1♀; USNM); Constanza 

(15.2 km SE; 18°49.5′N, 70°41.4′W; 1580 m), 7 May 1995, 

W. N. Mathis (1♀; USNM); El Rio (9.5 km E; 19°0.9′N, 

70°33.5′W; 980 m), 6 May 1995 (10♂, 2♀; USNM); Jarabacoa (1–2 km S; 19°06.9′N, 70°37′W; 520 m), 8–21 May 

1995, 1998, D. and W. N. Mathis (14♂, 2♀; USNM); Rio 

Camu (3.5 km NW La Vega; 19°13.7′N, 70°35.2′W; 100 m), 

10 May 1995, W. N. Mathis (6♂, 1♀; USNM); Rio Camu 

(3.5 km NW La Vega; 19°13.8′N, 70°35.2′W; 100 m), 18 

May 1998, D. and W. N. Mathis (9♂; USNM); Salto Baiguate (near Jarabacoa; 19°05.7′N, 70°37′W, 570m), 16 May 

1998, D. and W. N. Mathis (1♂; USNM). Pedernales: Alcoa 

Road (km 30; N Cabo Rojo; 18°07.3′N, 71°35.8′W; 1080 

m), 20 Mar 1999, W. N. Mathis (5♂, 1♀; USNM); Cabo Rojo 

(30 km N; 18°07′N, 71°39′W; 1070 m), 23–24 Jul 1990, J. 

Rawlins, S. Thompson, C. Young (1♀; CMP); Lago Oviedo 

(N shore; 17°47′N, 71°22.5′W), 15 May 1995, W. N. Mathis 

(1♂; USNM); Pedernales (18°01.8′N, 71°44.7′W), 19–20 

Mar 1999, W. N. Mathis (3♂; USNM); Pedernales (14 km E; 

17°57.9′N, 71°38.4′W; cenotes), 20 Mar 1999, W. N. Mathis 

(1♂; USNM); Pedernales (19 km N; 18°09.2′N, 71°44.8′W; 

230 m), 20 Mar 1999, W. N. Mathis (4♂, 1♀; USNM); Rio 

Mulito (13 km N Pedernales; 18°09′N, 71°46′W; 230 m), 17 

Jul 1992, R. Davidson, J. Rawlins, S. Thompson, C. Young 

(2♂, 3♀; CMP); Rio Mulito (21 km N Pedernales; 18°09.3′N, 

71°45.6′W; 270 m), 18–20 Mar 1999, W. N. Mathis (9♂; 

USNM). Peravia: Los Ranchitos (2 km E; 10 km SSE San José 

de Ocoa; 18°28′N, 70°28′W; 90 m), 8 Oct 1991, R. Davidson, 

J. Rawlins, S. Thompson, C. Young (3♂; CMP); Rio Ocoa 

(San José de Ocoa; 18°31.7′N, 70°30.4′W), 21 May 1998, D. 

and W. N. Mathis (3♂; USNM). Puerto Plata: Rio Camu (14 

km E Puerto Plata; 19°11.9′N, 70°37.4′W), 17 May 1995, 

W. N. Mathis (2♂; USNM); Rio Camu (14 km E Puerto Plata; 

19°41.9′N, 70°37.5′W), 23 May 1998, D. and W. N. Mathis 

(1♂; USNM); Rio Pérez (near Imbert; 19°44.1′N, 70°50.2′W), 

24 May 1998, D. and W. N. Mathis (7♂; USNM). San Cristobal: Nigua (18°22.4′N, 70°03′W), 27 May 1998, D. and 

W. N. Mathis (4♂; USNM).

GRAND CAYMAN. Airport (2 km E; 19°17.1′N, 81°21′W), 

26 Apr 1994, W. N. Mathis (5♂, 1♀; USNM). Airport (W end; 

19°17.4′N, 81°22.2′W), 26 Apr 1994, W. N. Mathis (1♂, 1♀; 

USNM). The Common (22°8′N, 81°23.9′W), 27 Apr 1994, 

W. N. Mathis (1♂; USNM).

GRENADA. St. Andrew: Balthazar (12°07.7′N, 

61°39.3′W), 2 Sep 1997, W. N. Mathis (4♂; USNM); Grand 

Étang (lake; 12°05.6′N, 61°41.7′W), 14 Sep 1997, W. N. Mathis 

(7♂; USNM). St. George: Airport (Point Salines; 11°59.9′N, 

61°46.1′W), 15 Sep 1996, W. N. Mathis (1♀; USNM); Beauséjour River (12°05.6′N, 61°44.3′W), 15 Sep 1996, W. N. Mathis 

(1♂; USNM); Grand Anse (12°01.3′N, 61°45.6′W), 15 Sep 1996, 

W. N. Mathis (7♂, 1♀; USNM); Airport (Point Salines; 12°05′N, 

61°46.9′W), 11–12 Sep 1997, W. N. Mathis (5♂; USNM); Point 

Salines Airport (W end; 12°00.3′N, 61°47.7′W), 12–19 Sep 1996, 

1997, W. N. Mathis (11♂, 1♀; USNM); St. George (12°05.5′N, 

61°44.9′W; meadow at ditch), 4 Jan 1986, U. Gardenfors (12♂, 

9♀; ZIL); True Blue Beach (11°59.9′N, 61°46.1′W), 15 Sep 

1996, W. N. Mathis (7♂, 1♀; USNM). St. Patrick: Bathway 

Beach (12°12.6′N, 61°36.7′W), 13–20 Sep 1996, 1997, W. N. 

Mathis (16♂; USNM); Levera Bay (12°13.6′N, 61°36.6′W), 18 

Sep 1996, W. N. Mathis (3♂, 1♂; USNM).

GUADELOUPE. Lamentin. Ravine Chaude (16°14′N, 

61°39.8′W), 25–30 Nov 2001, M. Martinez (1♂, 1♀; USNM).

HAITI. La Morinière (18°35.5′N, 72°11.4′W), 1–5 Mar 

1922 (1♂; AMNH). Pont Beudet (18°36.2′N, 72°13.3′W), 3–4 

Mar 1922 (1♂; AMNH).

JAMAICA. (1♂; ANSP). Clarendon: Farquhars Beach 

(17°50.9′N, 77°22.8′W), 9 May 1996, D. and W. N. 

Mathis, H. B. Williams (5♂; USNM); Grantham (18°09.3′N, 

77°23.8′W; 340 m), 16 Apr 2000, W. N. Mathis (7♂; USNM); 

Rest (3.5 km N; 17°54.1′N, 77°21.1′W), 9 May 1996, D. and 

W. N. Mathis, H. B. Williams (7♂; USNM); Rest (3 km N; 

17°54.3′N, 77°21.4′W), 15 Apr 2000, W. N. Mathis (4♂; 
number 643 • 107

USNM); Toll Gate (7.7 km S; 17°58′N, 77°22.3′W), 9 May 

1996, D. and W. N. Mathis, H. B. Williams (3♂; USNM). 

Manchester: Alligator Pond (17°52.1′N, 77°33.9′W), 8 May 

1996, D. and W. N. Mathis, H. B. Williams (5♂; USNM); 

near Clandon (18°09′N, 77°28.3′W), 8 May 1996, D. and 

W. N. Mathis, H. B. Williams (2♂, 2♀; USNM); Mandeville (18°03.5′N, 77°31.9′W), 7–13 May 1996, D. and W. N. 

Mathis, H. B. Williams (3♂; USNM); near Mandeville 

(18°03.5′N, 77°31.9′W), 15–18 Apr 2000, W. N. Mathis (1♂, 

3♀; USNM); near Warwick (17°54.1′N, 77°25.5′W), 7 May 

1996, D. and W. N. Mathis, H. B. Williams (13♂, 1♀; USNM). 

Portland: Berridale (18°06.5′N, 76°20′W), Rio Grande River, 

25 Apr 2000, W. N. Mathis (5♂, 1♀; USNM); Crystal Springs 

(18°12.5′N, 76°37.9′W), 18 May 1996, D. and W. N. Mathis, 

H. B. Williams (2♂; USNM); Long Bay (2.3 km W; 18°06.5′N, 

76°20′W), 24 Apr 2000, W. N. Mathis (5♂, 1♀; USNM); 

Reach (4 km N; 18°03.6′N, 76°20.4′W), 15 May 1996, D. and 

W. N. Mathis, H. B. Williams (3♂, 1♀; USNM). St. Andrew:

Cinchona (18°04.4′N, 76°39.3′W; 1400 m), 29 Apr 2000, 

W. N. Mathis (1♂; USNM); Clydsdale (18°04.9′N, 76°40.2′W; 

1030 m), 29 Apr 2000, W. N. Mathis (1♂; USNM); Irish 

Town (8 km SW; 18°03.9′N, 76°44.5′W), 7 Dec 1975, G. F. 

Hevel (2♂, 2♀; USNM); Mavis Bank (1.7 km E; Yallahs River; 

18°02.4′N, 77°39.5′W; 575 m), 21 Apr–1 May 2000, W. N. 

Mathis (10♂, 1♀; USNM); Mavis Bank (4.3 km SE; Yallahs 

River; 18°01.4′N, 76°38.1′W; 480 m), 22–23 Apr 2000, 

W. N. Mathis (1♂; USNM). St. Ann: Runaway Bay (18°27.4′N, 

77°19.6′W), Feb 1969, W. W. Wirth (6♂, 5♀; USNM). St. 

Elizabeth: Black River (18°01.4′N, 77°51.1′W), 11 May 

1996, D. and W. N. Mathis, H. B. Williams (1♂; USNM); 

Elim (18°07.1′N, 77°40.6′W), 10 May 1996, D. and W. N. 

Mathis, H. B. Williams (3♂; USNM). St. Mary: Port Maria (1 

km W; 18°22.8′N, 76°53.6′W), 18 May 1996, D. and W. N. 

Mathis, H. B. Williams (3♂; USNM). St. Thomas: Bath Fountain Spring (17°57.6′N, 76°21.3′W), 15 May 1996, D. and 

W. N. Mathis, H. B. Williams (1♀; USNM); Bath River, Bath 

(17°56.8′N, 76°21.6′W), 16 May 1996, D. and W. N. Mathis, 

H. B. Williams (5♂; USNM); Hagley Gap (1 km E; 18°00.1′N, 

76°36.7′W), 16 May 1996, D. and W. N. Mathis, H. B. Williams (6♂, 2♀; USNM); Mt. Lebanus (17°58.2′N, 76°32.7′W), 

16 May 1996, D. and W. N. Mathis, H. B. Williams (2♂; 

USNM); Yallahs River (mouth; 17°53′N, 76°35.6′W), 14 

May 1996, D. and W. N. Mathis, H. B. Williams (11♂, 2♀; 

USNM). Westmoreland: Negril Beach (18°16.4′N, 78°21.2′W; 

mangrove, rocky shore), 12 Mar 1970, W. W. Wirth (1♂, 1♀; 

USNM); Savanna-La-Mar (18°13′N, 78°08′W), 13 Mar 1970, 

W. W. Wirth (12♂, 3♀; USNM).

NAVASSA ISLAND. Ruins near Lulu Bay (18°23.75′N, 

75°01.07′W; 22 m), 29 Jul 1998, W. E. Steiner, J. M. Swearingen (1♂, 3♀; USNM).

PUERTO RICO. Aibonito (18°08.1′N, 66°15.6′W), 

14–17 Jul 1914 (1♂; AMNH). Coamo Springs (18°02.4′N, 

66°22.4′W), 17–19 Jul 1914 (3♂, 3♀; AMNH). Fajardo, Las 

Croabas (Seven Seas Beach; 18°23′N, 65°37′W), 17 Feb 1996, 

W. E. Steiner, J. M. Swearingen (33♂, 10♀; USNM). Jayuya (2 

km E; Rio Saliente; 18°12.8′N, 66°33.9′W), 22 Sep 1995, D. and 

W. N. Mathis (1♂, 1♀; USNM). Lajas (17°59.5′N, 67°02.8′W), 

Sep–Nov 1960, R. Cotte (4♂; USNM). Guayama area (17°59′N, 

66°07′W), 3 Apr 1956, T. W. Fischer (1♂; USNM). Maricao 

(4 km WNW; 18°10.7′N, 66°59.6′W), 21 Sep 1995, D. and 

W. N. Mathis (1♂; USNM). Playa de Guayanilla (18°0.4′N, 

66°46.1′W), 19 Sep 1995, D. and W. N. Mathis (7♂; USNM). 

Ponce (road through Bo. Arus; 18°0′N, 66°32′W), 26 Feb 1965, 

S. Medina Gaud (1♂; USNM). Salinas (17°57.5′N, 66°15.6′W), 

19 Feb 1964, S. Medina, I. Gaud, R. Gaspar (1♂; USNM). San 

Juan (18°28′N, 66°06′W), 14 Jan–14 Feb 1914, 1946, G. M. 

Young (1♂; 1♀; AMNH, ANSP).

ST. LUCIA. Black Bay (13°44.1′N, 60°58.7′W), 26 Feb–1 

Mar 1978, S. A. Marshall (1♂; CNC). Castries (5 km S; 13°59′N, 

60°00′W), 16 Jun 1991, D. and W. N. Mathis (1♀; USNM). 

Dauphin Boguis (1.6 km S Marquis; 14°01′N, 60°55′W), 17 Jun 

1991, D. and W. N. Mathis (4♂; USNM). Micoud (13°49′N, 

60°54′W), 15 Jun 1991, D. and W. N. Mathis (3♂, 1♀; USNM). 

Sulphur Spring (13°50′N, 61°03′W), 14 Jun 1991, D. and W. N. 

Mathis (3♂, 2♀; USNM).

ST. THOMAS. (18°20.5′N, 64°56′W) 8 Feb (1♂, 1♀; 

ANSP).

ST. VINCENT. Charlotte: Owia Salt Pond (13°22.5′N, 

61°08.5′W), 6 Sep 1997, W. N. Mathis (6♂; USNM); Peruvian 

Vale (13°10.7′N, 61°08.7′W), 6–8 Sep 1997, W. N. Mathis (6♂; 

USNM); South Rivers (13°14.6′N, 61°09.3′W), 8 Sep 1997, 

W. N. Mathis (1♂; USNM); Spring (13°11.1′N, 61°08.5′W), 

6 Sep 1997, W. N. Mathis (4♂, 1♀; USNM); Yambou River 

(13°09.8′N, 61°08.7′W), 8–10 Sep 1997, W. N. Mathis (7♂; 

USNM). St. Andrew: Buccament Bay (near beach; 13°11′N, 

61°16′W), 25–28 Mar 1989, W. N. Mathis (20♂, 5♀; USNM); 

Camden Park (13°10.2′N, 61°14.7′W), 25 Mar–4 Sep 1989, 

1997, A. Freidberg, W. N. Mathis (10♂; USNM); Layou 

(13°12′N, 61°17′W), 8 Jun 1991, D. and W. N. Mathis (1♂, 

3♀; USNM). St. George: Yambou Head (13°10.4′N, 61°09′W), 

27 Mar 1989, W. N. Mathis (1♂, 2♀; USNM). St. Patrick: Cumberland Bay (13°16′N, 61°16′W), 28 Mar–15 Sep 1989, 1997, 

W. N. Mathis (9♂; USNM); Ferret (13°12.5′N, 61°15.7′W), 5 Sep 

1997, W. N. Mathis (10♂; USNM); Palmiste Park (13°12.7′N, 

61°14.9′W), 5 Sep 1997, W. N. Mathis (4♂, 1♀; USNM); Wallilabou (beach; 13°15′N, 61°16′W), 27 Mar 1989, W. N. Mathis 

(4♂, 1♀; USNM). St. Philip: Gemswick (13°05′N, 59°35.5′W), 

31 Aug 1997, W. N. Mathis (4♂; USNM). St. Thomas: Farmers (13°12.8′N, 59°35.5′W), 2 Sep 1997, W. N. Mathis (5♂; 

USNM).

VIRGIN ISLANDS. St. Croix: (17°44′N, 64°46′W), 4–6 

Mar 1925 (1♂, 2♀; AMNH). St. Thomas: (18°21′N, 64°56′W), 

23 Feb–2 Aug 1925, A. Busck (14♂, 16♀; AMNH).

Type Locality. Neotropical. St. Thomas (18°21′N, 

64°56′W).

Distribution. (Map 27) Nearctic: Mexico (Colima, 

México, Morelos, San Luis Potosí, Tamaulipas). United States 

(Arizona, California, Florida, Texas). Neotropical: Bahamas, Belize (Corazal, Stann Creek), Bolivia, Colombia, Costa Rica (Cartago, Guanacaste, Heredia, Puntarenas), Ecuador (Chimborazo, 
108 • smithsonian contributions to zoology

MAP 27. Distribution map for Paralimna (Paralimna) thomae (Wiedemann).

El Oro, Guayas, Loja, Manabí, Pastaza), El Salvador, Guatemala 

(Escuintla, Izabal, Zacapa), Guatemala (Escuintla), Guyana, 

Honduras (Cortés), Mexico (Chiapas, Cuernavaca, Oaxaca, 

Tabasco, Veracruz-Llave), Nicaragua, Panamá (Cocle, Herera), 

Peru (Lima, Loreto, Madre de Dios), Trinidad and Tobago, Venezuela (Carabobo, Lara, Zulia), West Indies (Barbados, British 

Virgin Islands, Cuba, Dominica, Dominican Republic, Grand 

Cayman, Grenada, Guadeloupe, Haiti, Jamaica, Navassa, Puerto 

Rico, St. Croix, St. Lucia, St. Thomas, St. Vincent, Trinidad).

Remarks. In the New World, this is the most common and most widespread species of Paralimna and was one of 

the first species of this genus to be described (Williston 1896). 

This species can be confused with P. texana, which has similar yellowish-gray coloration of the face and body. This species 

is distinguished from P. texana by the coloration of the frons 

(brown in P. thomae, orange in P. texana), pattern of the coloration on the thorax (with 2 brown stripes, mostly brown in P. 

thomae, stripes very weak, not distinct, mostly silvery- to yellowish-gray in P. texana), abdomen (brown band on the basal half in 

P. thomae, reduced to basal 1⁄3 or less, frequently tergite 5 without brown band in P. texana), gray infuscation on transverse 

veins (absent versus present). Definitive characters, however, to 

distinguish these species are those of the male forefemur (ventral 

row of setae short, present only on base of femur versus long, 

frequently extended until apex of femur) and differences in structures of the male terminalia (basoventral process of presurstylus 

short and widened in P. thomae, more slender and tapered distally in P. texana). Distinguishing between females of P. thomae 

and P. texana can be very difficult.

Although P. thomae (as P. multipunctata) was previously 

reported from Brazil (Cresson 1947), our examination of specimens from Brazil (São Paulo: São Paulo, Barbrillini collector) 

reveal them to be P. secunda.

This species was listed in recent catalogs (Cogan and Wirth, 

1977:335; Mathis and Zatwarnicki, 1995:126) as being from 

the Oriental Region, specifically from Saint Thomas Island in the 

Indian Ocean. The lectotype and paralectotype, however, were 

collected by Dr. H. B. Hornbeck (see label data), who was a 

medical doctor from 1825 to 1844 in the Lesser Antilles, where 

he also made collections of insects that were mostly sent to Denmark. Wiedemann studied some of Hornbeck’s specimens, including the type series of this species, which is the most common 
number 643 • 109

and widespread species in the West Indies. Being relatively common and widespread, it is not surprising that this species was 

described three times. We thank Tadeusz Zatwarnicki for bringing the syntypes of P. thomae to our attention and suggesting its 

probable conspecificity with P. multipunctata.

34. Paralimna (Paralimna) velutina, sp. nov.

Figures 146–149, 197, 209, Map 28

Diagnosis. This species is distinguished from congeners by the following combination of characters: anterior margin 

of the frons shiny gray mediolaterally and 1 central, small shiny 

gray spot extended until near anterior ocellus, ventral frontoorbits with a large, dark purple, rounded spot and dorsal parafacial with a dark gray spot lateral to antenna; face yellowish or 

silvery gray, antennal groove golden or gray, carina with a shiny 

silvery or gray spot between antennal insertion followed by a 

golden spot below and 2 small, rounded, diffuse golden stains 

between largest facial setae; antenna black velvety with a whitish 

dorsal spot on pedicel, forefemur of male without outstanding 

setae.

Description. Body length 3.5–4.1 mm; body generally bicolored, dark brown dorsally and brown ventrally, lateral 

medial surfaces mostly gray.

Head: (See Figure 197) Frons brown, orange tinged, 

with yellow-gray or silvery-gray spots, 1 stripe at lateral margins 

of ocellar triangle, in front of anterior ocellar seta, not reaching 

the anterior margin of the frons, 1 quite inconspicuous stripe between inner vertical and interfrontal setae and on fronto-orbits 

above dorsal proclinate front orbital seta, between proclinate 

fronto-orbital setae, and just above antennal bases; anterior margin of the frons shiny gray mediolaterally and 1 central, small 

shiny gray spot extended until near anterior ocellus, according of 

incidence of light, ventral fronto-orbits with a large, dark purple, 

rounded spot, and dorsal parafacial with a dark gray spot lateral to antenna. Antenna black velvety (basal flagellomere with 

reddish-brown microtomentum according to incidence of light); 

pedicel with a dorsal whitish-gray spot, 1 dorsal, thicker seta and 

some short setae on the margin; basal flagellomere with slender, 

elongate, pale setae on margin; arista with 13 long, dorsal rays. 

Postcranium with 1 complete series of postocular setae. Face 

with yellowish or silvery-gray microtomentum, antennal groove 

golden or gray, carina with a shiny silvery or gray spot between 

antennal insertion followed by a golden spot below and 2 small, 

rounded, diffuse golden stains between largest facial setae; parafacial silvery gray and gena pale gray, gena almost nude, few 

setae only on distal margin and in middle; clypeus silvery gray. 

Eye round, as wide as high, height more than twice the height of 

gena. Gena exceptionally high, height longer than the length of 

basal flagellomere; gena-to-eye ratio 0.41–0.42.

Thorax: Mesonotum pale gray to yellowish gray with 

brown spots, 2 well-defined longitudinal stripes between dorsocentral rows, short, ending between third and fourth dorsocentral 

setae, and 1 central, less-defined stripe more slender than the 

others, 1 stripe laterally between first dorsocentral seta and intrapostalar seta; postpronotal, notopleuron, and supra-alar area 

gray; scutellum gray with distal margin dark brown and basal 

margin brown. Pleurae silvery gray except anepisternum with 

dorsal and ventral margins pale, brown tinged, anepimeron with 

ventral half brown and katepisternum brown on anterior half. 

Wing (see Figure 209) slightly brown tinged; veins pale brown; 

costal-vein ratio 0.47–0.53; M-vein ratio 0.96–1.00. Legs with 

silvery-gray microtomentum laterally and brown dorsally, except basitarsus yellow and the remaining tarsomeres become 

brown distally; forefemur of male without outstanding setae; 

foretibia straight.

Abdomen: Tergites distinctly bicolored, dark brown 

along anterior 2⁄3 and medially, otherwise yellowish gray dorsally and silvery gray laterodorsally, except tergite 2 yellowish 

gray medially; lateroventral margin of tergites brown. Male 

terminalia: (Figures 146–149) Presurstylus not bifurcate, horizontal process slightly enlarged basally and slightly curved; in 

lateral view postsurstylus widest medially with a medial, anteriorly protuberant, pointed lobe bearing rather short and slender 

setae and a thin digitiform process posteriorly; lateral aedeagal 

process about 1⁄2 length of aedeagus, robust and slightly curved 

in lateral view, rounded distally; aedeagus rather robust in 

posterior view, rounded on membranous apex; phallapodeme 

broadly triangular in lateral view; hypandrium moderately 

deeply concave.

Type Material. The holotype male is labeled 

“BRAZIL. Para: Rio Xingu Camp (52°22′W, 3°39′S)[,] ~60 km 

S. Altamira[,] 8–16 Oct 1986[,] P. Spangler & O. Flint/Igarape 

Jabuti[,] malaise trap[,] day collection/USNM ENT 00118270 

[plastic bar code label]/HOLOTYPE ♂ Paralimna velutina AleRocha&Mathis, USNM [red].” The holotype is double mounted 

(minuten pin in a red block of plastic), is in excellent condition, 

and is deposited in the USNM. Five paratypes (3♂, 2♀; INPA, 

USNM) bear the same label data as the holotype. Other paratypes are as follows: BRAZIL. Pará: Obidos, Igarapé Curucambá (01°50.1′S, 55°29.4′W; Malaise trap), 01–08 Sep 2001, 

J. A. Rafael, J. Vidal (1♀; INPA); Rio Iriri, Camp (03°50′S, 

52°40′W), Altamira (~60 km S), 17–18 Oct 1986, O. S. Flint Jr., 

P. J. Spangler (1♂; USNM).

GUYANA. Essequibo River, Moraballi Creek (06°12.3′N, 

58°33.9′W), 3 Sep–14 Oct 1929 (2♀; BMNH); Mazaruni 

(06°25.2′N, 58°37.7′W; high forest), 20 Aug 1937, Richards, 

Smarts, B.M 1937-776 (1♀; BMNH); Mazaruni-Potaro District, 

Takutu Mountains (06°15′N, 59°5′W; Malaise trap near stream 

in montane rainforest, Earth Research Expedition), 3–19 Dec 

1983, P. J. Spangler, W. E Steiner (5♂, 1♀; INPA, USNM); Paramakatoi (04°42′N, 59°42.8′W), 24–25 Aug 1997, W. N. Mathis 

(34♂, 6♀; INPA, USNM); Rockstone (05°58.6′N, 58°31.4′W), 

9 Jul 1011 (1♂; AMNH).

VENEZUELA. Barinas: Santa Rosa (08°36.6′N, 70°14.7′W), 

Feb–Dec 1942, 1943, P. Anduze (2♂; USNM). Merida: Tovar 

(08°19.7′N, 71°45.6′W), 7 Aug 1943, P. Anduze (4♂; USNM). 
110 • smithsonian contributions to zoology

Zulia: El Tucuco (45km SW Machiques; 09°50.7′N, 72°48.7′W), 

5–6 Jun 1976, A. S. Menke, D. Vincent (1♂, 3♀; USNM).

Type Locality. Neotropical: Brazil. Pará: Rio 

Xingu (camp; 52°22′W, 3°39′S).

Distribution. (Map 28) Neotropical: Brazil (Pará, 

Roraima), Guyana, Venezuela (Barinas, Merida, Zulia).

Etymology. The species epithet, velutina, is of Latin 

derivation and means velvety, alluding to the velvety aspect of 

the antennae.

Remarks. This species is easily distinguishable from 

congeners by the color of the frons and face in addition to characters of the male terminalia.

FIGURES 146–149. Structures of male terminalia of Paralimna (Paralimna) velutina, sp. nov. (Brazil. Pará): (146) epandrium, cerci, and presurstyli, posterior aspect; (147) postsurstylus, lateral aspect; (148) aedeagus, phallapodeme, and 

hypandrium, lateral aspect; (149) same, ventral aspect. Scale bar = 0.1 mm. (See Figures 2–5 for labels of structures.)
number 643 • 111

Subgenus Phaiosterna Cresson

Phaiosterna Cresson 1916:104. Type species: Paralimna decipiens

Loew 1878, original designation.—Wirth 1968:16 [Neotropical 

catalog].—Mathis and Zatwarnicki 1995:126–127 [world catalog]; 

2002:68–88 [revision].

Diagnosis. This subgenus is distinguished from other 

congeneric subgenera by the following combination of characters: small to moderately sized shore flies, body length 1.85–3.85 

mm; body usually with darkened infuscate uniform coloration, 

with rather metallic subshining or shining aspect.

Head: Ocelli arranged in equilateral triangle. Basal flagellomere broadly rounded apically, usually with parallel sides, 

length at most 1.5× width; arista bearing 6–8 long hairs along 

dorsal surface. Face short, nearly flat, carina and antennal groove 

not evident. Eye with oblique orientation, ovate, higher than wide. 

Gena short, about 1⁄4 eye height; wholly densely setulose.

Thorax: Mesonotum and pleural area mostly unicolorous and grayish brown to black; dorsal surface of scutellum 

gently convex. Wing with crossvein and veins and rest of wing 

essentially unicolorous. Forefemur of male lacking comb-like 

row of modified setae on anteroventral surface; posteroventral 

series of setae short.

Abdomen: Lacking distinctive fasciate pattern, tergites 

with infuscate uniform coloration. First sternite fully microtomentose (shiny with microtomentose edges in Neotropical 

species). Male terminalia: Presurstylus L-shaped, basoventral 

process developed, horizontal process connected dorsally; aedeagus wide with pointed apex; lateral aedeagal process subequal in length or longer than aedeagus and usually wide, not 

arched ventrally in lateral view.

Distribution. The subgenus Phaiosterna occurs in 

Afrotropical, Nearctic, and Neotropical Regions.

Discussion. The cladistic analysis of the species 

group of Paralimna shows that the subgenus Phaiosterna is a 

monophyletic group supported by following synapomorphies: 

scutum unicolored, dark; aedeagus wide in dorsal view, not 

tubular; and lateral aedeagal process so long as the aedeagus. 

The Neotropical species of the subgenus Phaiosterna constitute 

a clade with the first sternite being shiny in the middle and with 

microtomentose edges.

MAP 28. Distribution map for Paralimna (Paralimna) velutina, sp. nov.
112 • smithsonian contributions to zoology

35. Paralimna (Phaiosterna) decipiens Loew

Figures 150–153, Map 29

Paralimna decipiens Loew 1878:195 [United States. “Texas”; ST ♂, 

MCZ (11136)].—Osten Sacken 1878:201 [Nearctic catalog].—

Becker 1896:270 [list, Texas].—Coquillett 1900:259 [list, Puerto 

Rico].—Aldrich 1905:624 [Nearctic catalog].—Jones 1906:178–

179 [key, catalog].—Thaxter 1917:672–675 [parasites: Stigmatomyces rostratus Thaxter, S. paralimnae Thaxter]; 1931:145–146 

[parasite: S. gracilior Thaxter].—Wirth and Stone 1956:470 [list, 

California].

Paralimna (Phaiosterna) decipiens.—Cresson 1916:105 [subgeneric 

combination], 108 [revision, figure of head]; 1918:45 [revision]; 

1946:230 [review, Nearctic Region]; 1947:54–55 [review].—Wirth 

1965:748 [Nearctic catalog]; 1968:16 [Neotropical catalog].—Cole 

1969:396 [list, western United States].—Mathis 1995:633–635 [list, 

Galápagos Islands]; 1997:59–60 [list, Belize, figures of male terminalia].—Mathis and Zatwarnicki 1995:126 [world catalog]; 2002:72–

76 [revision].

Diagnosis. This species is distinguished from congeners, especially those of the subgenus Phaiosterna, by the following combination of characters: coloration variable but tending to 

be more tan to grayish brown than other congeners, moderately 

microtomentose, at most subshiny; small- to medium-sized shore 

flies, body length 1.85–3.60 mm; generally dark, gray brown to 

black species, moderately densely microtomentose, dorsum appearing more or less uniform in color, dull to subshiny, especially mesonotum.

Head: Generally gray brown to black. Basal flagellomere 

bearing short, generally inconspicuous fringe of whitish setulae 

along dorsum and dorsal portion of apex, length setulae far less 

than half height of basal flagellomere. Face gray to grayish brown, 

microtomentose, appearing dull, subdued. Eye-to-gena ratio 

0.23–0.29.

Thorax: Generally gray brown to black, legs darker, 

black with some grayish microtomentum; mesonotum moderately 

densely gray to grayish-brown microtomentose, appearing dull to 

subshiny, frequently with brown, short to long stripes between 

dorsocentral setae, especially anteriorly. Costal-vein ratio 0.41–

0.50; M-vein ratio 0.94–0.97. Ventral apex of foretibia lacking 

numerous, long, slender setulae; posterior surface of forebasitarsus lacking scattered long, slender setulae.

Abdomen: Slightly to distinctly lighter in color than mesonotum, mostly brownish gray to brownish black; tergites usually 

fasciate, with anterior portion of tergites darker gray to brownish black, less microtomentose, posterior portion more densely 

microtomentose, grayer. Male terminalia: (Figures 150–153) 

Presurstylus bifurcate, basoventral process about half the length 

of horizontal process, rather triangular, horizontal process connected medially, forming a shallowly arched band at ventral 

margin of cercus, with few setae; in lateral view postsurstylus 

slender, not widest medially, medial anterior process not developed, bearing few setae, medial posterior process lacking; lateral 

aedeagal process longer than aedeagus, wide throughout length, 

broadly rounded apically, triangular in lateral view, held close 

to lateral margins of aedeagus; aedeagus wide in posterior view, 

short, broadly rounded apically, thereafter forming a short, narrow point; phallapodeme, in lateral view, with anterior margin 

rounded; hypandrium deeply invaginated, pocket-like; cercus 

acutely pointed dorsomedially and ventrally.

Specimens Examined. Nearctic: UNITED STATES. 

Arizona. Coconino: Bill Williams Fork (35°12′N, 112°12′W), 

F. H. Snow (6♂, 6♀; ANSP). Pima: Kits Peak (near, 32°01.9′N, 

111°32′W; 1100 m), Baboquivari Mountains, 7–9 Aug 1916 (2♀; 

AMNH).

Arkansas. Garland: Hot Springs (34°30.2′N, 93°03.3′W), 

24 Jun, H. S. Barber (1♂, 1♀; ANSP).

California. Del Norte: Crescent City (41°45.4′N, 

124°12.1′W), 19 Apr 1908, M. C. Van Duzee (1♀; AMNH). Imperial: Calipatria (33°07.5′N, 115°30.8′W), 13 Nov 1921, E. R. 

Kalmbach (1♂, 2♀; ANSP); Salton Sea (33°28.6′N, 116°05.1′W), 

24 Nov 1921, E. R. Kalmbach (1♂, 1♀; ANSP). San Diego: Palm 

Canyon (33°13.2′N, 116°20.1′W), 7 Nov 1934, A. L. Melander 

(2♂, 1♀; ANSP).

Florida. Brevard: Merritt Island National Wildlife Refuge (28°32.4′N, 80°40.3′W), 29 Mar 1984, B. A. Foote (2♂, 

4♀; CMP). Henry: Clewiston (26°45.3′N, 80°56′W), 20 Jan 

1938, A. L. Melander (1♂; ANSP). Highlands: Archbold Biological Station (27°12.6′N, 81°20.9′W), 7 Feb 2000, D. and 

W. N. Mathis (7♂, 1♀; USNM); Archbold Biological Station 

(27°10.8′N, 81°21.1′W), 9 Feb 2000, D. and W. N. Mathis (6♂; 

USNM); Archbold Biological Station (27°11.3′N, 81°20.3′W), 

Key to New World Species of the Subgenus Phaiosterna

1. Ventral apex of foretibia of male bearing numerous long, slender setulae; forebasitarsus of male bearing long setulae along 

posterior surface (Neotropical) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. longiseta Mathis and Zatwarnicki

- Foreleg of male lacking long, slender setulae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2

2. Abdomen subshiny to shiny, sparsely microtomentose; aedeagus in posterior view higher than wide, narrow (Nearctic, 

Neotropical). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. obscura Williston

- Abdomen appearing dull, microtomentose, tannish gray; aedeagus in posterior view wider than high; wide (Nearctic, 

Neotropical). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. decipiens Loew
number 643 • 113

FIGURES 150–153. Structures of male terminalia of Paralimna (Phaiosterna) decipiens Loew (Brazil. Pará): 

(150) epandrium, cerci, and presurstyli, posterior aspect; (151) postsurstylus, lateral aspect; (152) aedeagus, 

phallapodeme, and hypandrium, lateral aspect; (153) same, ventral aspect. Scale bar = 0.1 mm. (See Figures 

2–5 for labels of structures.)

15 Jan–2 Nov 1989, M. A. Deyrup (6♂, 16♀; ABSF); Lake Istokpoga, Windy Point Park (27°18.4′N, 81°18.9′W; 18 m), 

15–16 Jan 2009, D. and W. N. Mathis (21♂; USNM); Rte. 70, 

cattle pond (27°12.5′N, 81°23.3′W), 7 Feb 2000, D. and W. N. 

Mathis (6♂, 1♀; USNM). Manatee: Bradentown (27°29.9′N, 

82°34.5′W), Mar, M. C. Van Duzee (1♂; ANSP). Marion: Salt 

Springs (29°21′N, 81°44.1′W), 14 Apr 1989, D. and W. N. 

Mathis (12♂, 6♀; USNM); Silver Springs (29°13′N, 82°03.5′W), 

2 Apr 1932, A. L. Melander (1♂; ANSP). Miami-Dade: Biscayne 

Bay (25°33.9′N, 80°13′W), A. T. Slosson (1♀; AMNH); Everglades National Park (25°22.9′N, 80°36.6′W), 20 Mar 1967, 

B. A. Foote (1♀; CMP); Florida City (25°26.9′N, 80°28.8′W), 13 

Jan 2007, D. and W. N. Mathis (3♂; USNM); Miami (25°46′N, 

80°12′W), 23 Feb 1912, F. Knab (1♂; ANSP); Royal Palm Park 

Ranger Station (25°22.9′N, 80°36.6′W), 28–29 Jan 1933, 1939, 

A. L. Melander (1♂, 2♀; ANSP). Monroe: Key West (beach; 

24°32.8′N, 81°47.7′W), 13 Jan 2007, D. and W. N. Mathis (1♂; 

USNM); Key West (Willie Ward Park; 24°32.9′N, 81°47.9′W), 
114 • smithsonian contributions to zoology

12 Jan–11 Feb 2000, 2007, D. and W. N. Mathis (3♀; USNM); 

Marathon Key (24°41.6′N, 81°05′W), 11 Feb 2000, D. and 

W. N. Mathis (1♂; USNM). Orange: Orlando (28°32.3′N, 

81°22.8′W), 2 Feb 1932, A. L. Melander (2♂, 2♀; ANSP). Palm 

Beach: Lake Worth (26°36.9′N, 80°03.4′W), A. T. Slosson (1♂, 

1♀; AMNH). Pinellas: St. Petersburg (27°46.2′N, 82°40.7′W), 

15 Feb 1924, E. T. Cresson Jr. (3♀; ANSP). Polk: Lakeland 

(28°02.4′N, 81°57′W), 10 Nov 1911 (1♀; AMNH). St. Johns:

Crescent Beach (1.6 km W; 29°46.1′N, 81°15.2′W), 26 Mar–10 

Apr 1984, B. A. Foote (1♀; CMP).

Georgia. Charlton: Okefenokee Swamp, Billy′s Island 

(30°48.3′N, 82°20.5′W), Jun 1912 (1♀; ANSP).

Louisiana. St. Charles: New Orleans (29°57′N, 90°04.3′W), 

1 Jan 1932, A. L. Melander (1♂; ANSP).

Mississippi. Harrison: Pass Christian (30°18.9′N, 

89°14.9′W), 6 Jul 1917; J. M. Aldrich (1♂, 1♀; ANSP). Oktibbeha: Mississippi State (30°18.9′N, 89°14.9′W), 30 Oct 1925 

(1♂; ANSP).

Nevada. Clark: Las Vegas Wash (36°05.2′N, 114°58.8′W), 

10–11 May 2001, D. and W. N. Mathis (1♂; USNM).

New Jersey. Atlantic: Absecon (39°24.9′N, 74°29.6′W), 26 

Sep 2003, D. and W. N. Mathis (4♂, 1♀; USNM). Cape May:

Cape May (38°57.6′N, 74°55.7′W), 9 Sep 1932, W. Stone (1♀; 

ANSP).

New Mexico. Grant: Bear Canyon Lake (NM Hwy. 35; 

32°53.1′N, 108°W), 2 Aug 2008, D. and W. N. Mathis (1♂, 

1♀; USNM).

North Carolina. Carteret: Bogue Island (34°41′N, 76°52′W), 

17 Oct 1974, G. C. Steyskal (3♀; USNM). Columbus: Lake 

Waccamaw (34°18.5′N, 78°29.7′W), 6 Jul 1985, A. Gerberich, 

W. Steiner (1♀; USNM). Dare: Duck (36°10′N, 75°45.3′W; bay 

shore), 12 Sep 1969, W. W. Wirth (3♂, 3♀; USNM). Morehead:

Fort Macon (34°41.8′N, 76°40.7′W), 29 Jul 1941, A. L. Melander (1♀; USNM).

Oklahoma. Carter: Ardmore (37°10.5′N, 97°08.6′W), 11 

Apr 1907, F. C. Bishopp (1♂, 1♀; ANSP).

South Carolina. Beaufort: Fripp Island (32°18.9′N, 

80°29′W), 26 Sep 1973, G. C. Steyskal (1♀; USNM).

Texas. Aransas: Aransas National Wildlife Refuge 

(28°06.9′N, 96°48′W), 20 May 1972, W. W. Wirth (1♂, 1♀; 

USNM); Salt Creek (28°15.4′N, 96°56.5′W), 5 Jun 1953 (1♀; 

USNM). Cameron: Brownsville (25°54.1′N, 97°29.8′W), 2–13 

Jan 1928 (1♀; AMNH); Brownsville (25°54.1′N, 97°29.8′W; 

pond shore), 29 Nov 1910 (1♀; ANSP); Brownsville (10 km N; 

26°04.6′N, 97°34.6′W), 20 Jul 1933, C. W. Sabrosky (1♂, 1♀; 

ANSP). Galveston: Dickinson (29°27.66′N, 85°03.1′W), Jun 

1929 (2♂, 2♀; ANSP). San Augustine: Rayburn Park (31°04′N, 

94°05′W), 15 May 1993, D. and W. N. Mathis (1♂; USNM). 

San Patricio: Mathis (5 km SW at Nueces River; 28°02.4′N, 

97°51.5′W), 9 Dec 1984, B. Gill, W. E. Steiner, D. Whitehead 

(17♂, 13♀; USNM); Sinton (12 km NE; Welder Wildlife Refuge; 28°06.1′N, 97°22′W), 8 Dec 1984, B. Gill, W. E. Steiner, 

D. Whitehead (5♂, 7♀; USNM). Travis: Austin (30°15.8′N, 

97°46.3′W), 7–20 Oct 1899, 1901 (1♂, 4♀; AMNH, ANSP).

Neotropical. BAHAMAS. North Andros: Andros Town 

(24°43.3′N, 77°47.4′W), 27–28 Feb 1966, O. L. Cartwright (1♂, 

2♀; USNM); Stainard Creek (24°50.6′N, 77°53.4′W), 13 Jan 

1953, E. B. Hayden (1♂, 3♀; AMNH). Central Andros: Mangrove Cay (24°16.4′N, 77°40.2′W), 26 Apr 1953, E. B. Hayden 

(1♂, 1♀; AMNH). Rum Cay: near Port Nelson (23°39.3′N, 

74°51′W), 16 Mar 1953, E. B. Hayden, L. Giovannoli (1♂, 

2♀; AMNH). San Salvador: near Cockburn Town (24°03.1′N, 

74°31.6′W), 18 Mar 1953, L. Giovannoli (2♀; AMNH). 

Turks and Caicos Islands: South Caicos Island (21°30.5′N, 

71°31.1′W), 11 Feb 1953, E. B. Hayden (2♂; AMNH).

BRAZIL. Amazonas: Manaus, Mercado Municipal 

(03°08.4′S, 60°01.4′W), 30 Aug 2001, R. Ale-Rocha, E. F. Soares 

(3♀; INPA); INPA (03°05.9′S, 59°59.1′W; 60 m), 4 May 2010, 

D. and W. N. Mathis (6♂; USNM); Reserva Ducke (26 km NE 

Manaus; 02°55.8′S, 59°58.5′W; 40 m), 5 May 2010, D. and W. 

N. Mathis (1♂; USNM); Itacoatiara (03°08.6′S, 58°26.6′W), Oct 

1969 (2♂, 3♀; MZUSP); km 30 Manaus-Itacoatiara, “CEPLAC,” 

7 May 1976 (1♂, 1♀; MZUSP); Parintins (02°37.7′S, 56°44.2′W), 

Oct 1969 (6♂; MZUSP, INPA). Pará: Santarém (02°26.4′S, 

54°41.9′W), Sep 1969, Expedição Permanente da Amazônia (3♂, 

3♀; MZUSP); Oct–Nov 1969, 1970 (18♂, 20♀; MZUSP, INPA); 

Ilha Cará-Açu (01°48′S, 53°15′W), Nov 1969 (1♂; MZUSP). 

Paraná: Antonina (25°28′S, 48°41.3′W; 13 m), 3 Feb 2010, D. 

and W. N. Mathis (2♂; DZUP, USNM); Antonina (25°27.1′S, 

48°41.1′W; beach; Ponta da Pita), 3 Feb 2010, D. and W. N. 

Mathis (1♂; DZUP, USNM); Curitiba, Universidade Federal 

do Paraná, Reserva Biológica (25°26.9′S, 49°14′W; 915 m), 30 

Dec–6 Feb 2009, 2010, D. and W. N. Mathis (2♂; USNM); Matinhos (N; 25°46.4′S, 48°30.8′W; 1 m; beach/estuary), 30 Jan–25 

Mar 2010, D. and W. N. Mathis (5♂; DZUP, USNM); Paranaguá 

(25°30.8′S, 48°29.9′W; 3 m), 23 Jan 2010, D. and W. N. Mathis 

(6♂; DZUP, USNM); Parque Igauçu (25°33.4′S, 49°13.6′W; 880 

m), 16–22 Jan 2010, D. and W. N. Mathis (3♂; DZUP, USNM). 

Santa Catarina: Barra Velha (26°38′S, 48°41.1′W; beach), 9 Feb 

2010, D. and W. N. Mathis (1♂; DZUP, USNM). São Paulo:

Ubatuba, Praia Puruba (23°21′S, 44°55.6′W; beach), 29 Mar 

2010, D. and W. N. Mathis (1♂; USNM).

TRINIDAD and TOBAGO. Tobago. St. John: Charlotteville (beach; 11°19.5′N, 60°32.9′W), 16 Apr–16 Jun 1993, 

1994, D. and W. N. Mathis (20♂, 3♀; USNM). St. Patrick:

Pigeon Point (beach; 11°9.7′N, 60°50′W), 19 Apr 1994, D. and 

W. N. Mathis (1♂; USNM). Trinidad. St. George: Arima 

(8 km N; 10°41′N, 61°18′W; Verdant Vale), 19 Jun 1993, 

W. N. Mathis (7♂, 3♀; USNM); Mount St. Benedict (10°39′N, 

61°24′W; creek near base), 19 Jun 1993, W. N. Mathis (5♂, 

3♀; USNM).

WEST INDIES. BARBADOS. Christ Church: Rockley 

Beach (13°04.3′N, 59°35.2′W), 11 Sep 1996, W. N. Mathis (1♀; 

USNM). St. Andrew: Belleplaine (13°14.8′N, 59°33.6′W), 21 

May 1996, D. and W. N. Mathis, H. B. Williams (1♂; USNM); 

Long Pond (13°15.1′N, 59°33.3′W), 10–11 Sep 1996, W. N. 

Mathis (1♂; USNM). St. Peter: Six Men’s Bay (13°16.5′N, 

59°38.8′W), 12 Sep 1996, W. N. Mathis (1♂; USNM).
number 643 • 115

BRITISH WEST INDIES. Sombrero (17°35.2′N, 

63°25.6′W; wet, shaded rocks), 9–13 Nov 1999, M. A. Ivie, J. B. 

Runyon (1♂; MSU).

CUBA. Havana: Havana (beach; 23°5.8′N, 82°27.7′W), 

2–14 Dec 1994, W. N. Mathis (14♂; USNM); Ojo de Aqua 

(22°54.6′N, 82°29.1′W), 8 Dec 1994, W. N. Mathis (1♂, 1♀; 

USNM). Matanzas: Palpite (2 km NE; 22°21.1′N, 81°10.5′W), 

30 Apr 1983, W. N. Mathis (1♀; USNM); Playa Larga (1 km 

E; 22°16.9′N, 81°11.7′W), 2 May 1983, W. N. Mathis (1♀; 

USNM). Mayabeque: Palmarejo (67 km E Havana; 23°08.5′N, 

81°40.6′W), 26 Apr 1983, W. N. Mathis (8♂, 1♀; USNM); 

Puerto Escondido (23°08.6′N, 81°43.6′W), 26 Apr 1983, 

W. N. Mathis (1♂, 2♀; USNM). Pinar del Rio: Soroa (22°47.7′N, 

83°W), 4–6 Dec 1994, W. N. Mathis (9♂, 1♀; USNM); Soroa (2 

km E; 22°47.7′N, 83°W), 29 Apr 1983, W. N. Mathis (9♂, 4♀; 

USNM). Sancti Spiritus: Playa Ancón (21°44.1′N, 79°59.9′W), 

12 Dec 1994, W. N. Mathis (6♂, 3♀; USNM); Topes de Collantes (21°54.4′N, 80°01.4′W; 670 m), 9–11 Dec 1994, W. N. 

Mathis (7♂, 4♀; USNM).

DOMINICA. Coulibistrie (15°27.1′N, 61°26.8′W), 21 Mar 

1989, W. N. Mathis (1♂; USNM). Soufrière Bay (15°13.9′N, 

61°21.6′W), 24 Mar 1989, A. Freidberg, W. N. Mathis (7♂, 1♀; 

USNM).

DOMINICAN REPUBLIC. Azua: near Pueblo Viejo 

(18°24.8′N, 70°44.7′W), 19 May 1998, D. and W. N. Mathis 

(5♂; USNM); Puerto Viejo (18°20.9′N, 70°50.4′W), 14 May 

1995, W. N. Mathis (11♂; USNM). Barahona: Barahona 

(18°12′N, 71°5.3′W), 25 Mar–20 May 1998, 1999, D. and 

W. N. Mathis (9♂; USNM); La Cienaga (3 km SW; 18°02′N, 

71°07′W; 35 m), 2 Oct 1991, C. Young, R. Davidson, S. Thompson, J. Rawlins (1♂; CMP); Los Patos (17°57.6′N, 71°10.9′W), 

15 May 1995, W. N. Mathis (3♂; USNM); Ojeda (17°58.2′N, 

71°10.6′W), 22 Mar 1999, W. N. Mathis (1♂; USNM); Paraíso 

(5 km N; 18°01.5′N, 71°11.6′W; 150 m), 21 Mar 1999, 

W. N. Mathis (1♀; USNM). Dajabon: Loma de Cabrera (10 

km S; 19°20′N, 71°37′W; 650 m), 12 Jul 1992, R. Davidson, 

J. Rawlins, S. Thompson, C. Young (1♂; CMP). El Seibo: Pedro 

Sáchez (18°51.4′N, 69°6.5′W), 26 May 1998, D. and W. N. 

Mathis (3♂; USNM); Rincón (near; 18°45.3′N, 68°55.7′W), 

12 May 1995, W. N. Mathis (1♂; USNM). Hato Mayor: Hato 

Mayor (5.5 km E; 18°46.4′N, 69°12.5′W), 26 May 1998, D. and 

W. N. Mathis (1♂; USNM). Independencia: Duvergé (2 km S; 

18°22′N, 71°31.4′W), 24 Mar 1999, W. N. Mathis (1♂; USNM); 

Mella (canal W; 18°21.5′N, 71°25.7′W), 20 May 1998, D. and 

W. N. Mathis (6♂; USNM); near Jimani (18°32.6′N, 71°50.3′W), 

20 May 1998, D. and W. N. Mathis (2♂; USNM); La Descubierta (18°34.1′N, 71°43.8′W; 0 m), 25 Mar 1999, W. N. Mathis 

(2♂; USNM); Puerto Escondido (18°19.6′N, 71°35′W; 1370 

m), 24 Mar 1999, W. N. Mathis (1♀; USNM). La Romana:

Isla Saona, Mano Juan (18°08.1′N, 68°44.5′W), 13 May 1995, 

W. N. Mathis (2♂, 1♀; USNM). La Vega: Jarabacoa (1–2 km 

S; 19°06.9′N, 70°37′W; 520 m), 8–21 May 1995, 1998, D. and 

W. N. Mathis (3♂; USNM); Rio Camu (3.5 km NW La Vega; 

19°13.7′N, 70°35.2′W; 100 m), 10 May 1995, W. N. Mathis 

(1♂; USNM); Rio Camu (3.5 km NW La Vega; 19°13.8′N, 

70°35.2′W; 100 m), 18 May 1998, D. and W. N. Mathis (7♂, 

1♀; USNM). Pedernales: Alcoa Road (km 30; N Cabo Rojo; 

18°07.3′N, 71°35.8′W; 1080 m), 20 Mar 1999, W. N. Mathis 

(1♀; USNM); Lago Oviedo (N shore; 17°47′N, 71°22.5′W), 

15 May 1995, W. N. Mathis (1♂, 1♀; USNM); Pedernales 

(18°01.8′N, 71°44.7′W), 19–20 Mar 1999, W. N. Mathis (7♂; 

USNM); Pedernales (14 km E; 17°57.9′N, 71°38.4′W; cenotes), 

20 Mar 1999, W. N. Mathis (5♂; USNM); Rio Mulito (13 km 

N Pedernales; 18°09′N, 71°46′W; 230 m), 17 Jul 1992, R. Davidson, J. Rawlins, S. Thompson, C. Young (1♀; CMP); Rio 

Mulito (21 km N Pedernales; 18°09.3′N, 71°45.6′W; 270 m), 

18–20 Mar 1999, W. N. Mathis (8♂, 2♀; USNM). Peravia: Rio 

Ocoa (San José Ocoa; 18°31.7′N, 70°30.4′W), 21 May 1998, 

D. and W. N. Mathis (3♂; USNM). Puerto Plata: Rio Camu 

(14 km E Puerto Plata; 19°11.9′N, 70°37.4′W), 17 May 1995, 

W. N. Mathis (4♂; USNM). San Cristobal: Nigua (18°22.4′N, 

70°03′W), 27 May 1998, D. and W. N. Mathis (5♂; USNM); 

Rio Haina (18°25.9′N, 70°00.4′W), 27 May 1998, D. and W. N. 

Mathis (1♂; USNM).

GRAND CAYMAN. Airport (W end; 19°17.4′N, 

81°22.2′W), 26 Apr 1994, W. N. Mathis (5♂, 2♀; USNM). Airport (2 km E; 19°17.1′N, 81°21′W), 26 Apr 1994, W. N. Mathis 

(6♂; USNM). Frank Sound Road (19°18.9′N, 81°10.9′W), 28 

Apr 1994, W. N. Mathis (8♂; USNM). Governor Gore Bird 

Sanctuary (19°16.7′N, 81°18.5′W), 25 Apr 1994, W. N. Mathis 

(10♂; USNM). Spotts (19°16.5′N, 81°19.1′W), 25 Apr 1994, 

W. N. Mathis (5♂, 3♀; USNM).

GRENADA. St. Patrick: Bathway Beach (12°12.6′N, 

61°36.7′W), 13 Sep 1997, W. N. Mathis (1♂, 4♀; USNM).

JAMAICA. Claredon: Grantham (18°09.3′N, 77°23.8′W; 

340 m), 16 Apr 2000, W. N. Mathis (8♂; USNM). Hanover:

The Great River (18°26.8′N, 77°59.5′W), 2 Jun 1920 (1♂; 

AMNH). Manchester: Alligator Pond (17°52.1′N, 77°33.9′W), 

8 May 1996, D. and W. N. Mathis, H. B. Williams (5♂; USNM); 

near Clandon (18°09′N, 77°28.3′W), 8 May 1996, D. and W. N. 

Mathis, H. B. Williams (1♂; USNM); near Warwick (17°54.1′N, 

77°25.5′W), 7 May 1996, D. and W. N. Mathis, H. B. Williams (1♂; USNM). Portland: Berridale (18°06.5′N, 76°20′W), 

Rio Grande River, 25 Apr 2000, W. N. Mathis (1♂; USNM); 

Reach (4 km N; 18°03.6′N, 76°20.4′W), 15 May 1996, D. and 

W. N. Mathis, H. B. Williams (1♀; USNM). St. Andrew: Hope 

Botanical Garden (18°01.4′N, 76°45′W; 198 m), 4–5 Dec 1919 

(1♂; AMNH); Irish Town (8 km SW; 18°03.5′N, 76°43.3′W), 

7 Dec 1975, G. F. Hevel (1♂; USNM); Mavis Bank (1.7 km 

E; 18°02.4′N, 77°39.5′W; 575 m), Yallahs River, 21 Apr–1 

May 2000, W. N. Mathis (4♂; USNM). St. Elizabeth: Brae 

River (2 km S; 18°04.2′N, 77°39.5′W), 10 May 1996, D. and 

W. N. Mathis, H. B. Williams (5♂; USNM); Elim (18°07.1′N, 

77°40.6′W), 10 May 1996, D. and W. N. Mathis, H. B. Williams (1♂; USNM); near Port Kaiser (17°52.3′N, 77°34.9′W), 8 

May 1996, D. and W. N. Mathis, H. B. Williams (1♂; USNM); 

Wally Wash Great Pond (17°57.9′N, 77°48.6′W), 19 Apr 2000, 

W. N. Mathis (5♂, 1♀; USNM). St. Thomas: Bath Fountain 
116 • smithsonian contributions to zoology

Spring (17°57.6′N, 76°21.3′W), 15 May 1996, D. and W. N. 

Mathis, H. B. Williams (1♂; USNM); Mt. Lebanus (17°58.2′N, 

76°32.7′W), 16 May 1996, D. and W. N. Mathis, H. B. Williams 

(2♂; USNM). Apr 1891 (2♂, 2♀; ANSP).

NAVASSA ISLAND. Ruins near Lulu Bay (18°23.75′N, 

75°01.07′W; 22 m), 29 Jul 1998, W. E. Steiner, J. M. Swearingen (2♂, 3♀; USNM).

PUERTO RICO. Adjuntas (18°09.8′N, 66°43.2′W), 22 

Sep 1995, D. and W. N. Mathis (2♂; USNM). Arecibo (beach; 

18°28.7′N, 66°42′W), 23 Sep 1995, D. and W. N. Mathis 

(5♂, 1♀; USNM). Playa de Guayanilla (18°0.4′N, 66°46.1′W), 

19 Sep 1995, D. and W. N. Mathis (5♂; USNM). Punta Jacinto 

(near Guanica; 17°57′N, 66°52.6′W), 20 Sep 1995, D. and 

W. N. Mathis (1♂; USNM).

ST. LUCIA. Dauphin Boguis (1.6 km S Marquis; 14°01′N, 

60°55′W), 17 Jun 1991, D. and W. N. Mathis (4♂; USNM). Micoud (13°49′N, 60°54′W), 15 Jun 1991, D. and W. N. Mathis 

(3♂; USNM). Soufrière (beach; 13°51′N, 16°54′W), 11–12 Jun 

1991, D. and W. N. Mathis (5♂, 1♀; USNM). Sulphur Spring 

(13°50′N, 61°03′W), 14 Jun 1991, D. and W. N. Mathis (4♂, 

3♀; USNM).

ST. VINCENT. Charlotte: Owia Salt Pond (13°22.5′N, 

61°08.5′W), 6 Sep 1997, W. N. Mathis (1♂, 1♀; USNM); Peruvian Vale (13°10.7′N, 61°08.7′W), 6–8 Sep 1997, W. N. 

Mathis (6♂; USNM). St. Andrew: Camden Park (13°10.2′N, 

61°14.7′W), 25 Mar–4 Sep 1989, 1997, W. N. Mathis (20♂, 

5♀; USNM); Layou (13°12′N, 61°17′W), 8 Jun 1991, D. and 

W. N. Mathis (10♂, 3♀; USNM). St. Patrick: Cumberland 

Bay (13°16′N, 61°16′W), 28 Mar 1989, A. Freidberg, W. N. 

Mathis (1♂; USNM); Wallilabou (beach; 13°15′N, 61°16′W), 

27 Mar–8 Jun 1989, 1991, D. and W. N. Mathis (3♂, 1♀; 

USNM).

Distribution. (Map 29) Nearctic: United States 

(Alabama, Arizona, Arkansas, California, Delaware, Florida, 

Georgia, Indiana, Michigan, Mississippi, Louisiana, Nevada, 

New Jersey, New Mexico, North Carolina, South Carolina, 

Texas, Virginia). Neotropical: Bahamas, Brazil (Amazonas, 

Pará, Paraná, Santa Catarina, São Paulo), Trinidad and Tobago, West Indies (Barbados, British West Indies, Cuba, Dominica, Dominican Republic, Grand Cayman, Grenada, Jamaica, 

Navassa Island, Puerto Rico, St. Lucia, St. Vincent, Virgin 

Islands).

MAP 29. Distribution map for Paralimna (Phaiosterna) decipiens Loew.
number 643 • 117

Remarks. Mathis and Zatwarnicki (2002) revised the 

New World species of the subgenus Phaiosterna and observed 

that this species and P. obscura are very similar in appearance. 

Externally it is difficult to distinguish between them due to variation in coloration and microtomentum; coloration from dark to 

pale in both species, although there is a tendency for specimens 

of P. decipiens to be paler and duller (more microtomentose) 

but with considerable variation, sometimes overlapping with 

P. obscura. Accurate identification may require examination 

of the aedeagus and gonites, which are very distinctive: the aedeagus in P. obscura is narrower from dorsal view, slightly longer than wide, and the distal end of the sclerotized portion is 

distinctly pointed; the aedeagus in P. decipiens is much broader 

than long in a dorsal view, somewhat rectangular with the distal angles rounded, and in lateral view the distal aspect of the 

sclerotized portion is slightly concave and with the actual apex 

of the aedeagus as a pointed process that is extended from an anteroventral angle (Mathis and Zatwarnicki 2002). Additionally, 

whereas in P. decipiens the hypandrium is shallow and broad 

(approximately 2× wider than high) and the phallapodeme is 

poorly developed, in P. obscura the hypandrium is deep (as wide 

as high) and tapering in apical 2⁄3, in anterior or posterior view, 

and the phallapodeme is strongly projected ventrally in middle.

36. Paralimna (Phaiosterna) longiseta Mathis 

and Zatwarnicki

Figures 154–157, Map 30

Paralimna (Phaiosterna) longiseta Mathis and Zatwarnicki 2002:81 [West 

Indies. Dominican Republic. Azua: near Pueblo Viejo (18°24.8′N, 

70°44.7′W); HT ♂; USNM].

Diagnosis. This species is distinguished from congeners, especially those of the subgenus Phaiosterna, by the following combination of characters: moderately small to medium-sized 

shore flies, body length 2.35–3.85 mm; generally dark, yellowish 

gray to brown species, moderately densely to densely microtomentose, dorsum appearing relatively uniform in color, dull to 

slightly subshiny, especially mesonotum.

Head: Generally grayish brown to golden brown. Basal 

flagellomere bearing short, generally inconspicuous fringe of whitish setulae along dorsum and dorsal portion of apex, length setulae far less than half height of basal flagellomere. Face yellowish 

gray to golden brown, microtomentose, appearing dull, subdued. 

Eye-to-gena ratio 0.39–0.40.

Thorax: Generally grayish yellow to golden brown, legs 

darker, black with some grayish microtomentum; mesonotum 

moderately densely grayish-yellow to golden-brown microtomentose, appearing dull to very slightly subshiny, infrequently with 

darker brown, short to long stripes between dorsocentral setae. 

Costal-vein ratio 0.40–0.43; M-vein ratio 0.86–0.96. Ventral apex 

of foretibia bearing numerous, long, slender setulae; posterior surface of forebasitarsus bearing scattered long, slender setulae.

Abdomen: Slightly lighter in color than mesonotum, mostly 

grayish to yellowish brown; tergites uniformly colored or anterior 

portion of tergites very fasciate, darker gray to brown, less microtomentose. Male terminalia: (Figures 154–157) Presurstylus bifurcate, basoventral process large, about the same length of horizontal 

process, rather truncate ventrally, at margin horizontal process connected medially, forming a shallowly arched band at ventral margin of cercus, with few setae; in lateral view postsurstylus slender, 

not distinctly wide medially, medial anterior process not developed, 

bearing few setae, medial posterior process lacking; lateral aedeagal process the same length of aedeagus, narrow, curved medially, 

acutely pointed apically; aedeagus wide, rather ovate in posterior 

view, short, narrowed at apex; phallapodeme, in lateral view, with 

anterior margin rounded; hypandrium deeply invaginated, pocketlike; cercus acutely pointed dorsomedially, medial margin sinuous, 

ventral margin with shallow, narrow indentation, slightly produced 

ventrolaterally; hypandrium deeply invaginated, pocket-like.

Specimens Examined. BRAZIL. Amazonas: Parintins (02°37.7′S, 56°44.2′W), Oct 1969, Expedição Permanente 

da Amazônia (13♂, 11♀; MZUSP, INPA). Pará: Santarém 

(02°26.4′S, 54°41.9′W), Sep–Oct 1969, 1970 (6♂, 3♀; (MZUSP); 

Ilha Cará-açu, mun. Almeirim (01°31.2′S, 52°34.9′W), Nov 

1969 (2♂, 1♀; MZUSP); Lago Jacaré, Rio Trombetas (01°27.9′S, 

56°18.6′W), Oct 1969 (1♂; MZUSP).

HONDURAS. Cortés: Omoa (15°47.8′N, 87°58.4′W), 26 

Sep 1995, D. and W. N. Mathis (3♂, 1♀; USNM); San Pedro 

Sula (8 km S; 15°25.7′N, 88°01.4′W), 25–26 Sep 1995, D. and 

W. N. Mathis (2♂; USNM).

WEST INDIES. DOMINICAN REPUBLIC. Azua: near 

Pueblo Viejo (18°24.8′N, 70°44.7′W), 19 May 1998, D. and 

W. N. Mathis (14♂, 8♀; USNM). La Vega: El Rio (9.5 km 

E; 19°0.9′N, 70°33.5′W; 980 m), 6 May 1995, W. N. Mathis 

(2♂; USNM); El Rio (9.5 km E; 19°0.7′N, 70°33.6′W; 980 m), 

24 May 1998, D. and W. N. Mathis (1♂; USNM); Rio Camu 

(3.5 km NW La Vega; 19°13.7′N, 70°35.2′W; 100 m), 10 May 

1995, W. N. Mathis (6♂, 1♀; USNM); Rio Camu (3.5 km 

NW La Vega; 19°13.8′N, 70°35.2′W; 100 m), 18 May 1998, 

D. and W. N. Mathis (6♂, 4♀; USNM). Peravia: Rio Ocoa 

(San José Ocoa; 18°31.7′N, 70°30.4′W), 21 May 1998, D. and 

W. N. Mathis (5♂, 1♀; USNM). Puerto Plata: Rio Camu (14 km 

E Puerto Plata; 19°41.9′N, 70°37.5′W), 23 May 1998, D. and 

W. N. Mathis (6♂; USNM); Rio Pérez (near Imbert; 19°44.1′N, 

70°50.2′W), 24 May 1998, D. and W. N. Mathis (2♂; USNM).

JAMAICA. Portland: Berridale (18°06.5′N, 76°20′W), Rio 

Grande River, 25 Apr 2000, W. N. Mathis (5♂, 1♀; USNM). 

St. Andrew: Mavis Bank (1.7 km E; 18°02.4′N, 77°39.5′W; 575 

m), Yallahs River, 21–22 Apr–1 May 2000, W. N. Mathis (14♂, 

2♀; USNM); Wag Water River, 25 Feb 1969, W. W. Wirth (1♂; 

USNM); Mavis Bank (4.3 km SE; 18°01.4′N, 76°38.1′W; 480 m); 

Yallahs River, 22–23 Apr 2000, W. N. Mathis (6♂; USNM). St. 

Thomas: Yallahs River (mouth; 17°53′N, 76°35.6′W), 14 May 

1996, D. and W. N. Mathis, H. B. Williams (8♂, 1♀; USNM).

PUERTO RICO. Rio Hoconuco (18°7.6′N, 67°2.6′W), 20 

Sep 1995, D. and W. N. Mathis (1♂; USNM).
118 • smithsonian contributions to zoology

Distribution. (Map 30) Neotropical: Brazil (Amazonas, Pará), Honduras (Cortés), West Indies (Dominican Republic, Jamaica, Puerto Rico).

Remarks. This species is similar to P. decipiens but is 

distinguished as follows: the 3 brown stripes on the scutum are fused 

on distal half of the scutum; the phallus is round with a sharp large 

protuberance on the medial dorsal surface (usually perceptible on 

undissected specimens). This species is distinguished from P. obscura

by the grayish coloration of the body, by the lack of distinctive dorsal bands on the tergites, and by characters of the male terminalia.

FIGURES 154–157. Structures of male terminalia of Paralimna (Phaiosterna) longiseta Mathis and Zatwarnicki (Brazil. 

Amazonas): (154) epandrium, cerci, and presurstyli, posterior aspect; (155) postsurstylus, lateral aspect; (156) aedeagus, 

phallapodeme, and hypandrium, lateral aspect; (157) same, ventral aspect. Scale bar = 0.1 mm. (See Figures 2–5 for labels 

of structures.)
number 643 • 119

MAP 30. Distribution map for Paralimna (Phaiosterna) longiseta Mathis and Zatwarnicki.

37. Paralimna (Phaiosterna) obscura Williston

Figures 158–161, Map 31

Paralimna obscura Williston 1896:391 [West Indies. St. Vincent; LT ♂ (designated by Mathis and Edmiston, 1991:832–834), BMNH].—Woodruff 

et al. 1998:68 [list, Grenada].

Paralimna (Phaiosterna) obscura.—Cresson 1918:45 [revision]; 1947:54 

[review].—Wirth 1968:16 [Neotropical catalog].—Mathis and Edmiston 1991:832–834 [review of Williston′s St. Vincent species].—Wirth 

1965:748 [Nearctic catalog]; 1968:16 [Neotropical catalog].—Woodley and Hilburn 1994:31 [list, Bermuda].—Mathis and Zatwarnicki 

1995:127 [world catalog]; 2002:83–88 [revision].—Mathis 1997:56–

57, 59 [review, Belize, figures].

Diagnosis. This species is distinguished from congeners, especially those of the subgenus Phaiosterna, by the following combination of characters: specimens tending to be dark 

brown to black, shiny, sparsely microtomentose; moderately 

small- to medium-sized shore flies, body length 2.25–3.50 mm; 

generally dark, gray brown to black species, moderately densely 

microtomentose, dorsum appearing relatively uniform in color, 

dull to subshiny, especially mesonotum.

Head: Generally gray brown to black. Basal flagellomere 

bearing short, generally inconspicuous fringe of whitish setulae 

along dorsum and dorsal portion of apex, length setulae far less 

than half height of basal flagellomere. Face gray to grayish brown, 

microtomentose, appearing dull, subdued. Eye-to-gena ratio 

0.24–0.27.

Thorax: Generally gray brown to black, legs darker, 

black with some grayish microtomentum; mesonotum moderately densely gray brown to brownish black, microtomentose, 

appearing dull to subshiny, frequently with lighter brown, short 

to long stripes between dorsocentral setae, especially anteriorly. 

Costal-vein ratio 0.48–0.53; M-vein ratio 0.95–1.0. Ventral 

apex of foretibia lacking numerous, long, slender setulae; posterior surface of forebasitarsus lacking scattered, long, slender 

setulae.

Abdomen: Slightly to distinctly lighter in color than mesonotum, mostly brownish gray to brownish black; tergites usually 
120 • smithsonian contributions to zoology

fasciate, with anterior portion of tergites darker gray to brownish black, less microtomentose, posterior portion more densely 

microtomentose, grayer. Male terminalia: (Figures 158–161) 

Presurstylus bifurcate, basoventral process small, rounded ventrally, horizontal process connected medially, forming a shallowly 

arched band at ventral margin of cercus, with few setae; in lateral 

view postsurstylus robust, with anterior margin swollen medially, 

bearing several setulae, medial anterior process not developed, 

bearing few setae; lateral aedeagal process the same length of aedeagus, triangular, acutely pointed apically; aedeagus somewhat 

triangular in lateral view, with a long, narrow, medial process apically; phallapodeme, in lateral view, with keel asymmetrical and 

only moderately produced, with a shallow indentation toward 

ventral margin; hypandrium very deeply invaginated, pocket-like; 

FIGURES 158–161. Structures of male terminalia of Paralimna (Phaiosterna) obscura Williston (Brazil. Amazonas): (158) 

epandrium, cerci, and presurstyli, posterior aspect; (159) postsurstylus, lateral aspect; (160) aedeagus, phallapodeme, and 

hypandrium, lateral aspect; (161) same, ventral aspect. Scale bar = 0.1 mm. (See Figures 2–5 for labels of structures.)
number 643 • 121

cercus with medial margins deeply sinuous, concave dorsally, 

acutely pointed dorsomedially, ventral margin produced ventromedially in posterior view.

Specimens Examined. Nearctic. BERMUDA.

(32°18′N, 64°47′W), 29 Jul 1905, T. Kincaid (1♀; ANSP). Devonshire: Cloverdale (32°18′N, 64°44′W), 29 May 1991, W. N. 

Mathis (5♂, 2♀; USNM); Devonshire Bay (32°18′N, 64°44′W), 

29 May 1991, W. N. Mathis (1♂, 1♀; USNM); Devonshire 

Marsh (32°18′N, 64°45′W), 30 May 1991, W. N. Mathis (14♂, 

1♀; USNM). Paget: Hungry Bay (beach; 32°17′N, 64°45′W), 2 

Jun 1991, W. N. Mathis (5♂; USNM); Paget Marsh (32°17′N, 

64°47′W), 2 Jun 1991, W. N. Mathis (8♂; USNM). Smith’s:

Spittal Pond (32°19′N, 64°43′W), 29 May 1991, W. N. Mathis 

(4♂; USNM). Warwick: Warwick Pond (32°16′N, 64°48′W), 3 

Jun 1991, W. N. Mathis (6♂, 3♀; USNM).

MEXICO. San Luis Potosí: Guerrero (23°33.6′N, 

100°41.1′W), 3 Aug 1933, C. W. Sabrosky (1♂, 1♀; ANSP).

UNITED STATES. Florida. Highlands: Lorida (8 km SE) 

at Istokpoga Canal (27°12.5′N, 80°55′W), 1 Mar 1984, A. G. 

Gerberich, J. E. Lowry, W. E. Steiner (1♂, 1♀; USNM). Levy:

Cedar Key (27 km NE; 29°17.7′N, 83°02′W), 1 Jun 1970, 

D. L. Bailey (1♀; USNM). Monroe: Big Pine Key, Long Beach 

(24°38.1′N, 81°21.6′W), 11 Feb 2000, D. and W. N. Mathis 

(2♂, 3♀; USNM). Pinellas: Clearwater (27°58′N, 82°48′W), 15 

Nov 1976, G. C. Steyskal (1♂; USNM).

Neotropical: BAHAMAS. San Salvador: Near Cockburn 

Town (24°03.1′N, 74°31.6′W), 18 Mar 1953, E. B. Hayden 

(1♂; ANSP).

BELIZE. Belize: Lighthouse Reef: Half Moon Cay 

(17°12′N, 87°31′W), Apr 1993, W. N. Mathis (1♂, 3♀; USNM). 

Stann Creek: Bread and Butter Cay (16°45′N, 88°09′W), Mar 

1988, W. N. Mathis (1♀; USNM); Man of War Cay (17°13′N, 

87°54′W), Nov 1987, D. and W. N. Mathis (10♂, 1♀; USNM); 

Stewart Cay (16°46′N, 88°09′W), Mar 1988, W. N. Mathis (1♂, 

1♀; USNM); Twin Cays (Aanderaa Flats; 16°50′N, 88°06′W), 

Mar 1988, W. N. Mathis (3♂, 6♀; USNM); Wee Wee Cay 

(16°45.9′N, 88°08.6′W), Jan–Nov 1987–1988, C. Feller, D. and 

W. N. Mathis (9♂, 1♀; USNM).

BOLIVIA. La Paz: Guanay (9 km NW; 15°29.5′S, 

67°53.8′W; 840 m), 13 Mar 2001, W. N. Mathis (7♂; USNM); 

Guanay (3 km E; 15°30.2′S, 67°52.3′W; 500 m), 14 Mar 2001, 

W. N. Mathis (3♂; USNM); Guanay (15°18.6′S, 68°13′W; 720 

m), 15 Mar 2001, W. N. Mathis (1♂; USNM); Guanay (22 km 

SE; 15°17.8′S, 68°15.6′W; 540 m), 17 Mar 2001, W. N. Mathis 

(1♀, 1♀; USNM); Guanay (15°29.8′S, 67°52.7′W; 460 m), 13 

Mar 2001, W. N. Mathis (4♂; USNM).

BRAZIL. Amazonas: Parintins (02°37.7′S, 56°44.2′W), 

Oct 1969, Expedição Permanente da Amazônia (2♀; MZUSP); 

Itacoatiara (03°08.6′S, 58°26.6′W), Nov 1969 (1♂, 1♀; INPA); 

Manaus (03°06.4′S, 60°01.6′W), 11 Jun 1976 (1♂, 2♀; INPA), 

25 Jun 1976, Paraluppi (1♀; INPA); Mercado Municipal 

(03°08.4′S, 60°01.4′W), 30 Aug 2001, R. Ale-Rocha, E. F. Soares 

(1♂, 3♀; INPA); Reserva Ducke (26 km NE Manaus; 02°55.8′S, 

59°58.5′W), 4 Jun 1976, I. S. Gorayeb (1♂; INPA); 12 May 1977 

(1♀; INPA), 12 May [19]77, E. Castellón B. (1♀; INPA); Feira 

do Produtor (03°02.2′S, 59°56.4′W), 27 Aug 2001, R. Ale-Rocha, 

E. F. Soares (1♀: INPA); Ipixuna, Rio Liberdade, Estirão da Preta 

(07°2146.7S, 71°5207.1W; sweeping), 11–15 May 2011, J. A Rafael, J. T. Câmara, R. F. Silva, A. Somavilla, R. Ale-Rocha (1♂, 4♀; 

INPA). Mato Grosso do Sul: Corumbá, Passo do Lontra-B.E.P. 

(19°34.3′S, 57°01′W; sweep net), alojamento (Brejo), 5 Aug 2011, 

C. Lamas, S. Nihei (1♀; MZUSP). Pará: Paciência, Rio Nhamundá 

(01°58.5′S, 56°52.1′W), 16 Jan 1968 (3♂, 2♀; MZUSP); Santarém 

(02°26.4′S, 54°41.9′W; Expedição Permanente da Amazônia), Sep 

1969, Expedição Permanente da Amazônia (4♂, 2♀; MZUSP), 

Nov 1969 (2♂, 4♀; MZUSP), Oct 1970 (8♂, 13♀; MZUSP); 

Oriximiná, Rio Trombetas (01°27.9′S, 56°18.6′W), Sep 1969 (2♂; 

MZUSP); Oct 1969 (1♂; MZUSP); Belém, Porto do Serviço Especial de Saúde Pública (01°28′S, 48°30′W), Sep 1969 (1♀; MZUSP); 

Estrada Oriximiná-Óbidos (01°38.7′S, 55°37′W), Nov 1969 (9♂, 

8♀; MZUSP); Rio Iriri, Camp (03°50′S, 52°40′W; Malaise trap), 

Altamira (~100 km S), 17–18 Oct 1986, O. S. Flint Jr., P. J. 

Spangler (1♀; USNM). Paraná: Antonina (25°28′S, 48°41.3′W; 

13 m), 3 Feb 2010, D. and W. N. Mathis (1♂; DZUP, USNM); 

Antonina (25°28.4′S, 48°40.9′W; beach/mangal), 3–4 Feb 2010, 

D. and W. N. Mathis (4♂; DZUP, USNM); Antonina (25°27.1′S, 

48°41.1′W; beach; Ponta da Pita), 3–15 Feb 2010, D. and 

W. N. Mathis (3♂; DZUP, USNM); Castro (Parque Lacustre; 

24°47.4′S, 50°0.3′W; 990 m), 22–31 Dec 2009, D. and W. N. 

Mathis (2♂, 3♀; DZUP, USNM); Castro (8 km N; 24°45.3′S, 

49°58.9′W; 1010 m), 1 Jan 2010, D. and W. N. Mathis (1♂; 

DZUP, USNM); Matinhos (N.; 25°46.4′S, 48°30.8′W; 1 m; 

beach/estuary), 30 Jan–25 Mar 2010, D. and W. N. Mathis (4♂; 

DZUP, USNM); Paranaguá (25°30.8′S, 48°29.9′W; 3 m), 23 Jan 

2010, D. and W. N. Mathis (4♂; DZUP, USNM); Parque Igauçu 

(25°33.4′S, 49°13.6′W; 880 m), 22 Jan–11 Feb 2010, D. and 

W. N. Mathis (8♂, 1♀; DZUP, USNM). Rio de Janeiro: Recreio 

dos Bandeirantes (23°01′S, 43°28′W), 24 Apr 1946, S. Lopes 

(2♂; MZUSP); Gávea (22°59.2′S, 43°14.7′W), H. Souza Lopes 

(1♂; MNRJ [Museu Nacional da Universidade Federal do Rio 

de Janeiro, Rio de Janeiro, Brazil (M. Couri)]); Ilha da Marambaia (23°03.6′S, 43°59.1′W), 4 Sep 2000, W. N. Mathis (1♂, 

1♀; DZUP, USNM). Rio Grande do Norte: Natal (05°47.7′S, 

35°12.7′W), Jul 1955, P. Ferreira (1♀; MZUSP). Santa Catarina:

Barra Velha (26°38′S, 48°40.9′W; beach), 29 Apr 2010, D. and 

W. N. Mathis (6♂; DZUP, USNM). São Paulo: Juquiá (24°19.3′S, 

47°38.2′W), May 1947, E. Rabello (4♀; MZUSP); Ubatuba, Praia 

do Estaleiro (23°20.5′S, 44°53′W; beach), 29–30 Mar 2010, D. 

and W. N. Mathis (3♂, 1♀; DZUP, USNM).

COSTA RICA. Guanacaste: Filadelfia (Río Tempisque; 

10°26.7′N, 85°32.9′W), 18 Jan 1910, P. P. Calvert (1♂; ANSP); 

Santa Cruz (?10°15.7′N, 85°35.1′W), 30 Jan 1910, P. P. Calvert 

(2♂; ANSP).

ECUADOR. Orellana: Rio Tiputini Biodiversity Station 

(0°38.2′S, 76°8.9′W), 12–26 Aug 1999, A. Baptista, M. Kotrba, 

W. N. Mathis (16♂, 4♀; USNM).
122 • smithsonian contributions to zoology

GUYANA. Conservation of Ecological Interactions and 

Biotic Associations [CEIBA] (~40 km S Georgetown; 6°29.9′N, 

58°13.1′W), 13 Apr 1994, W. N. Mathis (4♂; USNM); Dubulay 

Ranch, Aramatani Creek (5°40.9′N, 57°51.5′W), 9–11 Apr 1994, 

W. N. Mathis (5♂, 4♀; USNM); Dubulay Ranch, Berbice River 

(5°40.9′N, 57°51.5′W), 9–11 Apr 1994, W. N. Mathis (2♂, 1♀; 

USNM); Hope Beach (6°44.7′N, 57°57.3′W), 14–22 Apr 1994, 

1995, W. N. Mathis (9♂, 2♀; USNM); Karanambo (3°45.1′N, 

59°18.6′W), 31 Mar 1994, W. N. Mathis (12♂, 3♀; USNM); 

Karanambo, Maricuba (pond; 3°45.1′N, 59°18.6′W), 31 Mar–2 

Apr 1994, W. N. Mathis (8♂, 1♀; USNM); Karanambo, Rupununi River (ox bow; 3°45.1′N, 59°18.6′W), 2 Apr 1994, 

W. N. Mathis (2♂, 2♀; USNM); Kumo Pond (~20 km S Lethem; 

3°16.7′N, 59°43.5′W), 30 Apr 1995, W. N. Mathis (4♂, 3♀; 

USNM); Kumu River and Falls (25 km SE Lethem in Kanuku 

Mountains; 3°15.9′N, 59°43.6′W), 4–30 Apr 1994, 1995, W. N. 

Mathis (1♀; USNM); Mahaica (3 km W; 6°43.5′N, 57°56.6′W), 

14 Apr 1994, W. N. Mathis (4♂, 1♀; USNM); Moco-Moco (30 

km E Lethem in Kanuku Mountains; 3°18.2′N, 59°39.0′W), 29 

Apr 1995, W. N. Mathis (1♂; USNM); Moreru (marsh near; 

03°54.8′N, 59°32.7′W), 26 Apr 1995, W. N. Mathis (2♂, 1♀; 

USNM); Pirara Ranch, Cashew Creek (3°37.3′N, 59°44.2′W), 

23 Apr 1995, W. N. Mathis (7♂, 1♀; USNM); Pirara Ranch, 

Cashew Lake (3°36.7′N, 59°40.5′W), 23–27 Apr 1995, W. N. 

Mathis (13♂, 1♀; USNM); Pirara Ranch and River (03°32.1′N, 

59°40.5′W), 24–25 Apr 1995, W. N. Mathis (10♂, 2♀; USNM); 

Wiruni River (5°46.6′N, 58°01′W), 11 Apr 1994, W. N. Mathis 

(1♂; USNM).

HONDURAS. Cortés: Omoa (15°47.8′N, 87°58.4′W), 26 

Sep 1995, D. and W. N. Mathis (2♂; USNM); San Pedro Sula (8 

km S; 15°25.7′N, 88°01.4′W), 25–26 Sep 1995, D. and W. N. 

Mathis (5♂; USNM).

PERU. Loreto: Granja UNAP, near Iquitos (03°42′S, 

73°13.7′W), 23 Feb 1984, W. N. Mathis (1♂; USNM).

TRINIDAD and TOBAGO. Tobago. St. John: Bloody Bay 

River (11°18′N, 60°38′W), 14 Jun 1993, W. N. Mathis (2♂; 

USNM); Charlotteville (beach; 11°19.5′N, 60°32.9′W), 16 Apr–

16 Jun 1993, 1994, D. and W. N. Mathis (18♂; USNM); Charlotteville (5 km S; 11°18.9′N, 60°34.5′W), Hermitage River and 

beach, 10–11 Jun 1993, W. N. Mathis (10♂, 1♀; USNM); Parlatuvier (creek; 11°17.9′N, 60°35′W), 14 Jun 1993, W. N. Mathis 

(3♂; USNM); Speyside (11°18′N, 60°32′W), 13–15 Jun 1993, 

W. N. Mathis (5♂; USNM); Speyside (Doctor River; 11°18.2′N, 

60°31′W), 19 Apr 1994, D. and W. N. Mathis (7♂, 1♀; USNM); 

Speyside (Doctor River; 1 km NW; 11°18′N, 60°32′W), 12–13 

Jun 1993, W. N. Mathis (5♂; USNM). St. Patrick: Pigeon 

Point (beach; 11°9.7′N, 60°50′W), 19 Apr 1994, D. and W. N. 

Mathis (8♂; USNM). St. Paul: Delaford, Kings Bay (11°16′N, 

60°32.8′W), 13 Jun 1993, W. N. Mathis (5♂; USNM); Kendall (11°14.3′N, 60°35.7′W), 21 Apr 1994, W. N. Mathis (3♂; 

USNM). Trinidad. St. Andrew: Lower Manzanilla (5 km S; 

10°28′N, 61°03′W), 20 Jun 1993, W. N. Mathis (9♂; USNM); 

Lower Manzanilla (16 km S; 10°22′N, 61°01′W), 20 Jun 1993, 

W. N. Mathis (6♂, 1♀; USNM). St. George: Chacachacare Island (10°41′N, 61°46′W), saline pond margin, 29 Jun 1993, 

W. N. Mathis (4♀; USNM); Filette (1 km SE; 10°47′N, 61°21′W), 

Yarra River, 25 Jun 1993; W. N. Mathis (1♂; USNM). St. 

Patrick: Pitch Lake (10°14′N, 61°38′W), 24 Jun 1993, W. N. 

Mathis (2♀; USNM). St. Thomas: (10°32′N, 61°25′W), 2 Aug, 

A. Busck (1♀; ANSP).

WEST INDIES. BARBADOS. Christ Church: Graeme 

Hall Swamp (13°04.2′N, 59°34.7′W), 21–22 May–12 Sep 

1996, 1997, D. and W. N. Mathis, H. B. Williams (22♂; 

USNM); Oistins (13°03.9′N, 59°32.7′W), 22 May 1996, 

D. and W. N. Mathis, H. B. Williams (4♂; USNM); Rockley 

Beach (13°04.3′N, 59°35.2′W), 21 May–11 Sep 1996, 1997, 

D. and W. N. Mathis, H. B. Williams (15♂, 1♀; USNM). St. 

Andrew: Bawdens Ponds (13°14.8′N, 59°34.9′W), 2 Sep 1997, 

W. N. Mathis (1♂; USNM); Baxters (13°13.2′N, 59°34.1′W), 

21 May–11 Sep 1996, D. and W. N. Mathis, H. B. Williams 

(10♂, 1♀; USNM); Belleplaine (13°14.8′N, 59°33.6′W), 21 

May 1996, D. and W. N. Mathis, H. B. Williams (6♂; USNM); 

Long Pond (13°15.1′N, 59°33.3′W), 21 May–11 Sep 1996, 

1997, D. and W. N. Mathis, H. B. Williams (22♂; USNM). St. 

Joseph: Joes River (13°12.8′N, 59°32.3′W), 1–10 Sep 1996, 

1997, W. N. Mathis (5♂, 1♀; USNM). St. Peter: Six Men’s 

Bay (13°16.5′N, 59°38.8′W), 22 May–12 Sep 1996, D. and 

W. N. Mathis, H. B. Williams (7♂; USNM). St. Philip: Gemswick (13°05′N, 59°28.5′W), 31 Aug 1997, W. N. Mathis (8♂; 

USNM). St. Thomas: Farmers (13°12.8′N, 59°35.5′W), 2 Sep 

1997, W. N. Mathis (4♂; USNM).

CUBA. Guantánamo: Baracoa (road to; 20°21′N, 

74°30.1′W), 28 Feb 1992, M. von Tschirnhaus (2♂, 19♀; 

USNM). Havana: Havana (23°07′N, 82°23′W), Baker (2♀; 

ANSP); San Antonio de los Baños (22°54.9′N, 82°29.3′W), 8 

Dec 1994, W. N. Mathis (1♂; USNM); Ojo de Aqua (22°54.6′N, 

82°29.1′W), 8 Dec 1994, W. N. Mathis (8♂, 3♀; USNM). 

Holguín: Santa Lucia (3 km N; 21°03.3′N, 76°59.7′W), 22 

Feb 1992, M. von Tschirnhaus (1♂, 1♀; USNM). Matanzas:

Buena Ventura (7 km N; 22°18.1′N, 81°11.7′W), 1 May 1983, 

W. N. Mathis (8♂, 3♀; USNM); Palpite (1 km NE; 22°20.2′N, 

81°10.8′W), 2 May 1983, W. N. Mathis (24♂, 3♀; USNM); Palpite (2 km NE; 22°21.1′N, 81°10.5′W), 30 Apr 1983, W. N. 

Mathis (13♂, 7♀; USNM). Mayabeque: Palmarejo (67 km E 

Havana; 23°08.5′N, 81°40.6′W), 26 Apr 1983, W. N. Mathis 

(5♂, 4♀; USNM); Puerto Escondido (23°08.6′N, 81°43.6′W), 

26 Apr 1983, W. N. Mathis (1♂, 1♀; USNM). Pinar del Rio:

Pinar del Rio (S; 22°25.6′N, 83°41.3′W), 12–23 Sep 1913 

(3♂, 2♀; AMNH, ANSP); San Diego de los Baños (22°38.8′N, 

83°21.9′W), Aug (1♀; ANSP); Soroa (22°47.7′N, 83°W), 27 

Apr–6 Dec 1983, 1994, W. N. Mathis (13♂, 3♀; USNM); Soroa 

(2 km NW; 22°48.6′N, 83°1.0′W), 4–5 Dec 1994, W. N. Mathis 

(2♀; USNM); Soroa (2 km E; 22°47.7′N, 83°W), 29 Apr 1983, 

W. N. Mathis (3♂, 2♀; USNM). Cienfuegos: Soledad, Jardín 

Botánico (22°7.5′N, 80°19.2′W), Jan–Feb 1927, B. B. and C. T. 

Brues (1♂, 1♀; ANSP).
number 643 • 123

DOMINICA. Cabrits Swamp (15°35′N, 61°29′W), 22 

Mar 1989, W. N. Mathis (1♂; USNM). Layou River mouth 

(15°23.6′N, 61°25.5′W), 9 Jan–24 Mar 1965, W. W. Wirth (1♂, 

3♀; TZ). Soufrière Bay (15°13.9′N, 61°21.6′W), 24 Mar 1989, 

A. Freidberg, W. N. Mathis (1♀; USNM).

DOMINICAN REPUBLIC. Azua: near Pueblo Viejo 

(18°24.8′N, 70°44.7′W), 19 May 1998, D. and W. N. Mathis 

(6♂, 4♀; USNM). Barahona: Barahona (18°12′N, 71°05.3′W), 

25 Mar–20 May 1998, 1999, D. and W. N. Mathis (2♂; 

USNM); Cabral (canals E of Cabral; 18°15.2′N, 71°13.4′W), 16 

May 1995, W. N. Mathis (2♂, 2♀; USNM); Ojeda (17°58.2′N, 

71°10.6′W), 22 Mar 1999, W. N. Mathis (2♂; USNM). Dajabon: Loma de Cabrera (10 km S; 19°20′N, 71°37′W; 650 m), 

12 Jul 1992, R. Davidson, J. Rawlins, S. Thompson, C. Young 

(1♂; CMP). Distrito Nacional: Santo Domingo (Jardín Botánico; 

18°29.9′N, 69°56.9′W), 26 Mar 1999, W. N. Mathis (3♂; 

USNM). El Seibo: El Seibo (5 km E; 18°44.73′N, 68°59.2′W; 

120 m), 12 May 1995, W. N. Mathis (2♂; USNM); Pedro Sáchez 

(18°51.4′N, 69°6.5′W), 26 May 1998, D. and W. N. Mathis (2♂; 

USNM); Rincón (near; 18°45.3′N, 68°55.7′W), 12 May 1995, 

W. N. Mathis (6♂, 1♀; USNM). Hato Mayor: Hato Mayor (5.5 

km E; 18°46.4′N, 69°12.5′W), 26 May 1998, D. and W. N. 

Mathis (5♂, 2♀; USNM); Parque Los Haitises (near Cueva de 

Arena; 19°04′N, 69°28′W; 10 m), 7–9 Jul 1992, R. Davidson, 

J. Rawlins, S. Thompson, C. Young (2♀; CMP). La Vega: El Rio 

(9.5 km E; 19°0.7′N, 70°33.6′W; 980 m), 24 May 1998, D. and 

W. N. Mathis (1♂; USNM); Jarabacoa (1–2 km S; 19°06.9′N, 

70°37′W; 520 m), 8–21 May 1995, W. N. Mathis (1♀; USNM); 

Rio Camu (3.5 km NW La Vega; 19°13.7′N, 70°35.2′W; 100 

m), 10 May 1995, W. N. Mathis (5♂, 1♀; USNM); Rio Camu 

(3.5 km NW La Vega; 19°13.8′N, 70°35.2′W; 100 m), 18 May 

1998, D. and W. N. Mathis (2♂, 1♀; USNM). Pedernales:

Lago Oviedo (N shore; 17°47′N, 71°22.5′W), 15 May 1995, 

W. N. Mathis (1♂; USNM); Rio Mulito (21 km N Pedernales; 

18°09.3′N, 71°45.6′W; 270 m), 18–20 Mar 1999, W. N. Mathis 

(2♂; USNM). Peravia: Rio Ocoa (San José Ocoa; 18°31.7′N, 

70°30.4′W), 21 May 1998, D. and W. N. Mathis (2♂; USNM). 

Puerto Plata: Rio Camu (14 km E Puerto Plata; 19°11.9′N, 

70°37.4′W), 17 May 1995, W. N. Mathis (1♂; USNM). San 

Cristobal: Nigua (18°22.4′N, 70°03′W), 27 May 1998, D. and 

W. N. Mathis (5♂; USNM).

GRENADA. St. Andrew: Balthazar (12°07.7′N, 

61°39.3′W), 19 Sep 1996, W. N. Mathis (1♂; USNM); Grand 

Étang (lake; 12°05.6′N, 61°41.7′W), 14–20 Sep 1996, 1997, 

W. N. Mathis (8♂; USNM); Pearls Airport (12°08.7′N, 61°36.6′W), 

15–17 Sep 1996, 1997, W. N. Mathis (12♂, 1♀; USNM). St. 

George: Airport (Point Salines; 11°59.9′N, 61°46.1′W), 15 

Sep 1996, W. N. Mathis (5♂, 2♀; USNM); Beauséjour Bay 

(12°05.5′N, 61°44.9′W), 21 Sep 1996, W. N. Mathis (1♂; 

USNM); Grand Anse (12°01.3′N, 61°45.6′W), 15 Sep 1996, 

W. N. Mathis (7♂; USNM); Point Salines Airport (W end; 12°0.3′N, 

61°47.7′W), 12–19 Sep 1996, 1997, W. N. Mathis (7♂; USNM); 

Airport (Point Salines; 12°05′N, 61°46.9′W), 11–12 Sep 1997, 

W. N. Mathis (5♂; USNM); True Blue Beach (11°59.9′N, 

61°46.1′W), 15 Sep 1996, W. N. Mathis (7♂; USNM). St. John:

Palmiste (12°08.7′N, 61°44.4′W), 21 Sep 1996, W. N. Mathis 

(4♂; USNM); Palmiste Lake (12°08.3′N, 61°44′W), 19 Sep 

1996, W. N. Mathis (9♂; USNM). St. Patrick: Bathway Beach 

(12°12.6′N, 61°36.7′W), 13–20 Sep 1996, 1997, W. N. Mathis 

(19♂; USNM); Levera Bay (12°13.6′N, 61°36.6′W), 18 Sep 

1996, W. N. Mathis (6♂; USNM).

JAMAICA. Clarendon: Portland Cottage (17°45.4′N, 

77°11′W), 13 May 1996, D. and W. N. Mathis, H. B. Williams (3♂; USNM); Rest (3.5 km N; 17°54.1′N, 77°21.1′W), 

9 May 1996, D. and W. N. Mathis, H. B. Williams (2♂, 1♀; 

USNM); Rest (3 km N; 17°54.3′N, 77°21.4′W), 15 Apr 2000, 

W. N. Mathis (2♂, 1♀; USNM); Salt River (4 km N; 17°52.1′N, 

77°09.5′W), 13 May 1996, D. and W. N. Mathis, H. B. Williams 

(1♂; USNM); Toll Gate (7.7 km S; 17°58′N, 77°22.3′W), 9 May 

1996, D. and W. N. Mathis, H. B. Williams (7♂; USNM). Manchester: Alligator Pond (17°52.1′N, 77°33.9′W), 8 May 1996, 

D. and W. N. Mathis, H. B. Williams (7♂; USNM); Battersea, 

Feb 1910, R. Thaxter (2♀; ANSP); near Mandeville (18°03.5′N, 

77°31.9′W), 15–18 Apr 2000, W. N. Mathis (1♀; USNM); near 

Warwick (17°54.1′N, 77°25.5′W), 7 May 1996, D. and W. N. 

Mathis, H. B. Williams (7♂, 1♀; USNM). Portland: Berridale 

(18°06.5′N, 76°20′W), Rio Grande River, 25 Apr 2000, W. N. 

Mathis (1♂, 2♀; USNM). St. Andrew: Mavis Bank (1.7 km E; 

18°02.4′N, 77°39.5′W; 575 m), Yallahs River, 21–22 Apr–1 

May 2000, W. N. Mathis (4♂, 1♀; USNM). St. Elizabeth: Brae 

River (18°05.2′N, 77°39.3′W), 10 May 1996, D. and W. N. 

Mathis, H. B. Williams (3♂, 2♀; USNM); Brae River (2 km S; 

18°04.2′N, 77°39.5′W), 10 May 1996, D. and W. N. Mathis, 

H. B. Williams (3♂; USNM); Elim (18°07.1′N, 77°40.6′W), 

10 May 1996, D. and W. N. Mathis, H. B. Williams (1♂, 1♀; 

USNM); Elim (18°07.1′N, 77°40.5′W), 10 Apr 2000, W. N. 

Mathis (2♂; USNM); near Port Kaiser (17°52.3′N, 77°34.9′W), 

8 May 1996, D. and W. N. Mathis, H. B. Williams (2♂; 

USNM); Wally Wash Great Pond (17°57.9′N, 77°48.6′W), 19 

Apr 2000, W. N. Mathis (1♂, 1♀; USNM); Ys Falls (18°09.3′N, 

77°49.5′W), 17–18 Apr 2000, W. N. Mathis (1♀; USNM). St. 

Thomas: Bath River, Bath (17°56.8′N, 76°21.6′W), 16 May 

1996, D. and W. N. Mathis, H. B. Williams (2♂; USNM); Yallahs River (mouth; 17°53′N, 76°35.6′W), 14 May 1996, D. and 

W. N. Mathis, H. B. Williams (5♂; USNM). Westmoreland:

Negril Beach (18°16.4′N, 78°21.2′W; mangrove, rocky shore), 

12 Mar 1970, W. W. Wirth (4♂, 1♀; USNM); Negril (8 km E; 

18°16.7′N, 78°18.1′W; freshwater marsh), 13 Mar 1970, W. W. 

Wirth (5♂, 5♀; USNM); Savanna-La-Mar (18°13′N, 78°08′W), 

13 Mar 1970, W. W. Wirth (7♂, 2♀; USNM).

PUERTO RICO. Ponce (18°01′N, 66°34′W; Mercedita 

sugarcane mill, mud bank, rum distillery waste ditch), 2 May 

1965, S. M. Gaud, J. G. Tuduri; (7♂, 18♀; USNM); Arecibo 

(beach; 18°28.7′N, 66°42′W), 23 Sep 1995, D. and W. N. Mathis 

(1♂; USNM). Fajardo, Las Croabas (Seven Seas Beach; 18°23′N, 

65°37′W), 17 Feb 1996, W. E. Steiner, J. M. Swearingen (8♂, 
124 • smithsonian contributions to zoology

6♀; USNM). Maricao (4 km WNW; 18°10.7′N, 66°59.6′W), 

21 Sep 1995, D. and W. N. Mathis (5♂, 1♀; USNM). Playa de 

Guayanilla (18°0.4′N, 66°46.1′W), 19 Sep 1995, D. and W. N. 

Mathis (6♂; USNM). Punta Jacinto (near Guanica; 17°57′N, 

66°52.6′W), 20 Sep 1995, D. and W. N. Mathis (2♂, 1♀; 

USNM). Toa-Baja (18°26.7′N, 66°15.3′W), 17 Jan 1915, G. 

Garb (1♀; ANSP).

ST. LUCIA. Castries (5 km S; 13°59′N, 60°00′W), 16 Jun 

1991, D. and W. N. Mathis (3♂; USNM). Dauphin Boguis 

(1.6 km S Marquis; 14°01′N, 60°55′W), 17 Jun 1991, D. and 

W. N. Mathis (5♂; USNM). Soufrière (beach; 13°51′N, 16°54′W), 

11–12 Jun 1991, D. and W. N. Mathis (3♂; USNM). Sulphur 

Spring (13°50′N, 61°03′W), 14 Jun 1991, D. and W. N. Mathis 

(6♂; USNM).

ST. VINCENT. St. Vincent (2♂; CU; “Lot. 662 Sub. 75”; 

one of the males lacks its head. The determination label, which 

is handwritten, reads “Paralimna brunnea Will. [cursive] mss. 

may be obscura”). Charlotte: Montreal (13°12′N, 61°11′W), 

26 Mar 1989, A. Freidberg (1♂, 1♀; USNM); Peruvian Vale 

(13°10.7′N, 61°08.7′W), 6–8 Sep 1997, W. N. Mathis (1♂; 

USNM); Spring (13°11.1′N, 61°08.5′W), 6 Sep 1997, W. N. 

Mathis (5♂; USNM); Yambou River (13°09.8′N, 61°08.7′W), 

8–10 Sep 1997, W. N. Mathis (7♂; USNM). St. Andrew: Buccament Bay (near beach; 13°11′N, 61°16′W), 25–28 Mar–8 Jun 

1989, 1991, D. and W. N. Mathis (10♂, 6♀; USNM); Camden 

Park (13°10.2′N, 61°14.7′W), 25 Mar–4 Sep 1989, 1997, W. N. 

Mathis (11♂, 4♀; USNM); Layou (13°12′N, 61°17′W), 8 Jun 

1991, D. and W. N. Mathis (4♂; USNM). St. David: Richmond 

Beach, 28 Mar 1989, W. N. Mathis (2♂; USNM). St. George:

Yambou Head, 27 Mar 1989, W. N. Mathis (5♂, 2♀; USNM). 

St. Patrick: Cumberland Bay (13°16′N, 61°16′W), 8–10 Jun–15 

Sep 1991, 1997, D. and W. N. Mathis (14♂; USNM); Hermitage (13°15′N, 61°12.9′W), 9 Sep 1997, W. N. Mathis (2♂; 

USNM); Palmiste Park (13°12.7′N, 61°14.9′W), 5 Sep 1997, W. 

N. Mathis (3♂; USNM); Wallilabou (beach; 13°15′N, 61°16′W), 

8 Jun 1991, D. and W. N. Mathis (6♂, 1♀; USNM).

Distribution. (Map 31) Nearctic: Bermuda, Mexico (San Luis Potosí), United States (California, Florida, Texas). 

Neotropical: Bahamas, Belize (Belize, Stann Creek), Bolivia (La 

Paz), Brazil (Amazonas, Mato Grosso do Sul, Pará, Paraná, 

Rio de Janeiro, Rio Grande do Norte, Santa Catarina, São 

Paulo), Costa Rica (Guanacaste), Honduras (Cortés), Ecuador 

MAP 31. Distribution map for Paralimna (Phaiosterna) obscura Williston.
number 643 • 125

(Orellana), Guyana, Paraguay, Peru (Loreto), Trinidad and Tobago, West Indies (Barbados, Cuba, Dominica, Dominican Republic, Grenada, Jamaica, Puerto Rico, St. Lucia, St. Vincent, 

Virgin Islands).

Remarks. On Wee Wee Cay (Belize. Stann Creek), 

all specimens were collected from a large muddy area with a 

covering mat of blue-green algae. This species and Paralimna decipiens Loew are closely related and are very similar in external 

appearance. A discussion of the distinguishing characteristics of 

these two species is presented in the “Remarks” for P. decipiens. 

Both P. decipiens and P. obscura frequently occur sympatrically 

(Mathis and Zatwarnicki 2002).

Paralimna (Coganolimna), subgen. nov.

Type species Paralimna limbata Loew 1862b:13.

Diagnosis. This subgenus is distinguished from other 

congeneric subgenera by the following combination of characters: 

small to moderately sized shore flies, body length 2.7–3.5 mm; 

body vittate, broad brown striped on thorax with darkened infuscate coloration, yellowish to brownish gray between brown 

stripes, subshining aspect.

Head: Ocelli arranged in equilateral triangle; pseudopostvertical pair thin, not developed. Basal flagellomere narrowed toward the apex, elongated, slender, dorsal margin slightly concave, 

and length at most 1.7× width, dorsal setae scarcely developed; 

arista bearing 12–14 long hairs along dorsal surface. Face nearly 

flat, carina and antennal groove not evident. Eye slightly rounded, 

higher than wide. Gena short, about 1⁄4 eye height.

Thorax: Mesonotum and pleurae striped, yellowish to 

brownish gray between brown stripes. Wing hyaline with variable 

pattern, with strong infuscation on veins or cells fully brown spotted. Forefemur of male lacking comb-like row of modified setae 

on anteroventral surface.

Abdomen: With fasciate pattern. Sternite 1 fully microtomentose. Male terminalia: Epandrium as wide as high; cercus well 

developed, bearing numerous setulae; presurstylus variable, horizontal process well developed or not, presurstylus not connected 

dorsally; aedeagus wide with pointed apex; aedeagus and lateral 

process subequal in length; lateral aedeagal process usually wide, 

not arched ventrally in lateral view.

Distribution. The Paralimna (Coganolimna) subg. 

nov. occurs in the Afrotropical Region and includes the species 

previously included in former limbata species group.

Discussion. The cladistic analysis of the species 

group of Paralimna showed that the Paralimna (Coganolimna)

subg. nov. is a monophyletic group supported by following synapomorphies: basal flagellomere elongated, slender with dorsal 

margin slightly concave, scutum and pleural area striped, with defined bands. Paralimna (Coganolimna) subg. nov. included species very similar to species of subgenus Phaiosterna, where they 

have been previously placed (Mathis and Zatwarnicki, 1995). 

The subgenus Phaiosterna, however, is a very well-supported 

clade with homogeneous species sharing a very homogeneous 

and characteristic male terminalia. In Paralimna (Coganolimna) 

subg. nov. the male terminalia is very variable between species 

and differs considerably from that of Phaiosterna, and Mathis 

and Zatwarnicki (2002) allocated these species in the genus 

Paralimna as part of the limbata group.

PHYLOGENETIC CONSIDERATIONS

In this study, we have re-examined the evidence, all morphological characters, in an attempt to discover more definitively 

the phylogenetic placement of Paralimna and related taxa, especially those taxa that have been treated as subgenera within 

Paralimna. We asked the basic question, what are the phylogenetic relationships (hypothetical) among taxa closely related to 

or within Paralimna? As background, we first present a summary of the evidence and relationships at the tribal level (within 

Hydrelliinae) and then proceed to the evidence and our analysis 

of it for taxa within Dryxini.

The tribe Dryxini is one of five tribes placed in the subfamily Hydrelliinae (Mathis and Zatwarnicki, 1995, 2002). The 

sister group relationship of Dryxini with Notiphilini is corroborated by five synapomorphies (Mathis and Zatwarnicki, 2002): 

(1) ventral anepisternal seta elongate, twice length of dorsal seta 

(secondarily reduced in a few taxa); (2) midtibia with prominent, 

erect, extensor setae along the dorsal surface; (3) only reclinate 

fronto-orbital seta well developed, proclinate seta(e) reduced or 

lacking; (4) abdominal tergites fasciate (secondarily reduced in 

some taxa); and (5) subepandrial plate reduced.

Dryxini are distinguished from Notiphilini and the tribe’s 

monophyly is established by the following characters (synapomorphies are noted by an asterisk [*]) (Mathis and Zatwarnicki, 

2002): *gena high (secondarily short in some species); *face 

wide, transversely arched, and generally projected anteriorly 

(the face in Notiphila is comparatively narrower and much flatter); dorsocentral setae 4 (1+3; secondarily reduced in some genera of Dryxini, whereas in Notiphilini there are 3 setae [1+2], a 

synapomorphy for Notiphilini); costa elongate, extended to vein 

M (the costa is short, extended only to vein R4+5 in Notiphilini, 

a synapomorphy for Notiphilini); male terminalia with surstylus divided into a presurstylus (clasper) and postsurstylus (surstylus) (postsurstylus reduced or fused with epandrium with 

only presurstylus present in Notiphilini, a synapomorphy for 

Notiphilini); *presurstylus with apex angulate and bifurcate; 

*pre- and postgonite reduced or lacking (the structure remaining may represent a fused and/or reduced pre- and postgonite); 

and *hypandrium connected basally with postsurstylus, not 

with epandrium.

With the phylogenetic background for further study of the 

tribe Dryxini within the subfamily Hydrelliinae established and 

the monophyly of Dryxini documented, we now proceed with 

the cladistic analysis and resultant relationships among lineages 

within the genus Paralimna, the most species-rich genus within 
126 • smithsonian contributions to zoology

Dryxini, but with a few explanatory remarks first. In the presentation on relationships within Paralimna that follows, the 

characters used in the analysis are noted first. Each character is 

immediately followed by a discussion to explain its states and 

to provide perspective and any qualifying comments about that 

character. After presentation of the information on character 

evidence, a hypothesis of the cladistic relationships is presented 

and briefly discussed. The cladogram (see Figure 215) is the 

primary mode to convey relationships, and the discussion is to 

supplement the cladogram and is intended only to complement 

the latter. In the discussion of character data, a “0” indicates 

the state of the outgroup; a “1” or “2” or greater indicates the 

derived states. All characters were treated as nonadditive. The 

numbers used for characters in the presentation are the same as 

those on the cladogram, and the sequence is the same as noted 

in the character matrix (Table 1). The genera Afrolimna Cogan, 

Oedenops Becker, and Papuama Mathis and Zatwarnicki were 

the outgroups in our cladistic analysis.

Characters Used in the Analysis

1. Facial height: (0) short, about 1.5× the length of frons; 

(1) long, at least 2× the length of frons.

2. Ventral margin of eye: (0) rounded or almost truncate, eye 

rather rounded; (1) distinctly tapered, eye ovate; (2) reniform.

3. Silvery white spot on dorsum of pedicel: (0) absent; 

(1) present.

4. Insertion of ocellar pair of setae: (0) before the anterior ocellus; (1) lateral to the anterior ocellus.

5. Shape of the facial carina: (0) depressed, antennal groove indistinct; (1) high, antennal groove shallow; (2) very elevated 

or projected, antennal groove deep.

6. Shape of the frons: (0) horizontal or nearly so, antenna almost aligned with the dorsal region of the eye; (1) convex, 

antenna inserted beneath the dorsal line of the eye or well 

above half the height of the eye in lateral view.

7. Coloration of the frons: (0) lacking characteristic pattern of 

spots; (1) with characteristic pattern of spots, brown with 

linear yellow spots at lateral margins of ocellar triangle and 

in front of anterior ocellar seta, between medial vertical and 

interfrontal setae, and on fronto-orbits between proclinate 

fronto-orbital setae and just above antennal bases.

8. Aspect of pseudopostocellar setae: (0) undeveloped, thin, 

poorly differentiated, as long as or slightly longer than adjacent ocellar setae; (1) distinctly more robust and at least 

twice length of adjacent ocellar setae.

9. Genal setae: (0) gena partially setose, post-cranial series not 

mixing with genal setae; (1) completely setulose, post-cranial 

series extended to gena.

10. Genal height: (0) height greater than length of basal flagellomere; (1) height less than or equal to the length of basal 

flagellomere.

11. Post-cranial setae: (0) arranged in a single series; (1) arranged 

in two series.

12. Dorsal setulae of basal flagellomere: (0) short and usually sparsely setulose; (1) long setulae, usually numerous, sometimes irregular and longer on base of basal 

flagellomere.

13. Shape of basal flagellomere: (0) parallel sided, rounded at the 

apex; (1) tapered toward the apex.

14. Length of the basal flagellomere: (0) short, stout, dorsal margin straight or slightly convex; (1) elongated, slender, dorsal 

margin slightly concave.

15. General aspect of coloration of body: (0) matte; (1) metallic.

16. General scutal coloration: (0) dorsum dark and lighter on 

sides, usually 2 or 3 light stripes dorsally; (1) unicolorous, 

dark; (2) striped body dorsally and laterally, with defined 

bands; (3) unicolorous, pale with some poorly defined stains 

gently darkened.

17. Coloration of mesopleuron: (0) homogeneously colored 

(light or dark); (1) predominantly gray, with or without 

spots; (2) predominately brown, dorsal and ventral portions 

predominantly brown (katepisternum completely brown); 

(3) distinctly lighter on dorsal half (dorsal pleurites gray, 

only thin band on ventral margin of anepisternum and anepimeron brown) and darker on ventral half (ventral pleurites brown); (4) horizontally striped.

18. Crossvein dm–cu: (0) straight; (1) concave toward the base 

of the wing.

19. Coloration of the wing: (0) hyaline or semi-hyaline; (1) predominantly brown with hyaline halos or predominantly hyaline with small brown spots.

20. Anteroventral setae of the male forefemur: (0) long and conspicuously flattened; (1) long or short, with blunt apex, but 

not conspicuously flattened; (2) setae simple, with sharpened 

apex, not differentiated from adjacent setae.

21. Posteroventral setae of the male forefemur: (0) differentiated from adjacent setae; (1) not differentiated from adjacent 

setae.

22. Ventral surface of the male foretibia: (0) not modified, 

straight or slightly arcuate; (1) sinuous, with a groove along 

ventral surface.

23. Ventral surface of the male forefemur: (0) normal, gradually 

tapered toward the apex; (1) distinctly concave at distal 1⁄2 

or 2⁄3.

24. Coloration of tergites: (0) distinctly bicolored with a dark 

basal band and distal margin pale; (1) homogeneously 

colored.

25. Sternite 1: (0) fully microtomentose; (1) bare in the middle 

but margins microtomentose; (2) bare.

26. Dark shiny stain on the posteroventral margin of tergite 5: 

(0) absent; (1) present.

27. Shape of horizontal process of presurstylus: (0) disconnected 

dorsally; (1) connected dorsally.

28. Shape of postsurstylus in lateral view: (0) not widest medially, anterior and posterior process undeveloped; (1) widest 

medially with an anteriorly protuberant lobe and a variable 

process posteriorly in the middle.
number 643 • 127

29. Shape of aedeagus in dorsal view: (0) tubular, slightly flattened laterally; (1) wide, not tubular.

30. Lateral aedeagal process: (0) longer than 1⁄2 length of the 

aedeagus; (1) as long as the aedeagus; (2) 1⁄2 the length or 

slightly shorter than the aedeagus; (3) very short, about 1⁄6

aedeagal length.

31. Shape of the lateral aedeagal process: (0) curved, robust, 

comma shaped; (1) triangular, wide; (2) long and slender.

32. Basoventral process of presurstylus: (0) greatly developed, 

large, robust, horizontal process very short or absent; 

(1) basoventral process undeveloped; (2) developed, with 

varied shapes, horizontal process well developed.

33. Appendages of the aedeagus: (0) lacking appendages; (1) with 

lateral aedeagal process separated from the aedeagus; 

(2) with lateral aedeagal process fused with the aedeagus.

34. Setae at apex of postsurstylus: (0) absent; (1) 2 setae near 

apex.

35. Position of the gonite relative to the postsurstylus: (0) positioned at apical 1⁄4–1⁄3 of the postsurstylus; (1) positioned at 

apex of the postsurstylus.

36. Horizontal process of surstylus: (0) absent; (1) short and robust; (2) long and slender.

Analysis, Results, and Discussion

The heuristic analysis of the data matrix (Table 1), using 

the multiple TBR + TBR option of NONA (Goloboff, 1999), 

generated 100 cladograms from the analysis of the 36 unordered 

characters and 67 taxa. The cladograms have a length of 127 

steps and consistency and retention indices of 0.40 and 0.81, 

respectively. From that pool of cladograms we produced a strict 

consensus, collapsing 21 nodes (Figure 214). The WinClada software (Nixon, 2002) was used to map the characters on the cladograms by means of the slow optimization option. The following 

discussion of the distribution of character states is graphically 

portrayed on the first of the 100 obtained trees (Figure 215).

In summary, and as indicated on the cladogram (Figure 215), 

the genus Paralimna is a monophyletic lineage supported by the 

following synapomorphies (included in parentheses are character numbers and in some cases, annotations): post-cranial setae 

arranged in two series (arranged in one series in P. reticulata, 

P. monstruosa, and in a group of species within Paralimna

(Paralimna) sensu stricto formed by P. confluens, P. cressoni, P. 

curta, P. fulgifrons, P. guttata, and P. stellata) (111

); basoventral 

process of surstylus developed and quite variable, especially the 

horizontal process well developed (a very homoplastic character) (322

); lateral aedeagal process fused with the aedeagus (332); 

postsurstylus with two setae present near apex (341

); gonite positioned at apex of postsurstylus (351

); and horizontal process of 

surstylus long and slender (362

).

Paralimna is closely related to Afrolimna, which shares the 

following characters: ocellar pair of setae lateral to the anterior 

ocellus (a very homoplastic characters) (41

); gena completely 

setulose (91

) (gena partially setose in P. reticulata, P. picta, 

P. millepuncta, P. monstruosa, and in the major group formed 

by all Neotropical species and some Afrotropical species); basal 

flagellomere with long setae on dorsal margin (121

) (in P. limbata

and in the subgenus Phaiosterna, except P. fusca, the dorsal setae 

of basal flagellomere are short); basal flagellomere narrowed 

toward the apex (131

) (with parallel sides, rounded at the apex 

in the group formed by P. bicolor, P. longiseta, P. decipiens, and 

P. obscura).

The topology of the lineages within Paralimna showed three 

distinct monophyletic clades including the limbata species group, 

the subgenus Phaiosterna, and the largest clade formed by subgenus Paralimna sensu stricto (excluding the limbata species 

group). Although not all species of Paralimna have been included 

in the analysis, it is evident that the subgenus Paralimna sensu 

lato (including the limbata group) is not monophyletic as currently known. The analysis does demonstrate, however, that the 

subgenus Paralimna sensu stricto (without the limbata group) 

is a monophyletic group that is supported by two unambiguous synapomorphies—frons with characteristic pattern of spots 

(71

) and mesopleuron predominantly gray with or without spots 

(171

) (this character is secondarily modified within the subgenus 

to pleural area predominantly brown [172

], and mesopleuron 

distinctly lighter on dorsal half with dorsal pleurites predominantly gray, darker on ventral half with ventral pleurites brown 

[173

])—and one homoplasious synapomorphy—face long, at 

least twice the length of frons (shared with limbata group; this 

character is secondarily modified within the subgenus to short, 

about 1.5× the length of frons, in P. boensis, a big group formed 

by one Afrotropical species, P. hirticornis, and several Neotropical species inside Paralimna (Paralimna) sensu stricto) (11

).

Cogan (1968) recognized seven species groups among Afrotropical species of the subgenus Paralimna: the limbata group 

(3 species), the nigripes group (12 species), the nidor group 

(3 species), the arabica group (8 species), the confluens group 

(6 species), the albonotata group (1 species), and the wirthi

group (1 species). All species groups proposed by Cogan (1968) 

were represented in the analysis (at least two species of each 

species group) plus we sampled species from all zoogeographical regions. None of the groups proposed by Cogan (1968) was 

confirmed except for the limbata group, which is supported by 

three synapomorphies: basal flagellomere elongated and slender with dorsal margin concave (141

); general coloration of the 

body striped dorsally and laterally, with defined bands (162

); and 

pleural area horizontally striped (174

); and two homoplasious 

synapomorphies: face long, at least twice the length of frons (11

) 

and posteroventral setae of the male forefemur not differentiated 

from adjacent setae (211

).

The subgenus Phaiosterna emerged as a monophyletic 

group supported by six synapomorphies: scutum unicolorous, 

dark (161

); aedeagus wide in dorsal view, not tubular (291); lateral aedeagal process as long as the aedeagus (301

); coloration of 

body metallic (151

); tergites homogeneously colored (241); and 

horizontal process of presurstylus connected dorsally (271

) (the 

last three are homoplasious characters). A clade is formed by the 
128 • smithsonian contributions to zoology

TABLE 1. Matrix of characters and taxa used in the cladistic analysis of Paralimna. The genera Afrolimna Cogan (A. keiseri), Oedenops

Becker (O. nudus), and Papuama Mathis and Zatwarnicki (P. ismayi) were used as outgroups. Character numbers and character states 

are explained in the text.

Characters 1 1 1 1 1 1 1 1 1 1 2 2 2 2 2 2 2 2 2 2 3 3 3 3 3 3 3

Taxa 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6

O. nudus 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 3 0 0 0 2 1 0 0 1 2 0 0 0 0 - - 0 0 0 0 0

P. ismayi 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 2 0 0 0 0 2 0 0 0 0 - - 0 0 0 0 0

A. keiser 0 2 0 1 1 1 0 0 1 1 0 1 1 0 0 0 0 0 0 2 0 0 0 0 2 0 0 0 0 - 1 1 1 0 0 1

P. adunca 0 0 0 0 1 1 1 1 0 0 1 1 1 0 0 0 1 0 0 1 0 0 0 0 2 0 0 1 0 2 0 2 2 1 1 2

P. albonotata 1 0 1 0 2 0 1 1 0 0 1 1 1 0 0 0 1 0 0 1 1 0 0 0 2 0 0 0 0 2 0 2 2 1 1 2

P. argyrostoma 0 0 1 0 1 1 1 1 0 0 1 1 1 0 0 0 1 0 0 1 0 0 0 0 2 0 0 1 0 2 0 1 2 1 1 2

P. approximata 1 0 0 1 1 1 1 0 1 0 1 1 1 0 0 0 3 0 1 2 1 0 0 0 2 0 ? 1 ? 2 0 ? 2 1 1 ?

P. aurantia 0 0 0 0 1 1 1 1 0 0 1 1 1 0 0 0 1 0 0 1 0 0 1 0 2 0 0 1 0 2 0 1 2 1 1 2

P. bicolor 0 0 0 1 0 1 0 0 1 1 1 0 0 0 1 1 0 0 0 2 0 0 0 1 0 0 1 0 1 1 1 2 2 1 1 2

P. boensis 0 0 0 0 1 1 1 1 1 0 1 1 1 0 0 0 2 0 0 0 1 0 0 0 2 0 0 0 0 2 2 2 2 1 1 2

P. bruneiceps 0 0 0 1 1 1 1 1 0 0 1 1 1 0 0 0 1 0 0 1 0 0 0 0 2 0 0 1 0 2 0 2 2 1 1 2

P. castanea 0 0 0 1 1 1 1 1 0 0 1 1 1 0 0 0 1 0 0 1 0 0 0 0 2 0 0 1 0 2 0 2 2 1 1 2

P. concors 1 0 1 0 1 1 1 1 0 0 1 1 1 0 0 0 1 0 0 1 1 0 0 0 2 0 ? 1 0 ? ? 1 2 1 1 2

P. confluens 1 0 1 1 2 1 1 0 0 0 0 1 1 0 0 0 1 0 1 0 1 0 0 0 2 0 1 1 0 0 0 1 2 1 1 2

P. cressoni 1 0 0 1 2 0 1 0 0 0 0 1 1 0 0 0 1 0 1 1 1 0 0 0 2 0 1 1 0 2 0 1 2 1 1 2

P. crinita 1 0 1 0 2 0 1 1 0 0 1 1 1 0 0 0 1 0 0 1 0 0 0 0 1 0 0 1 0 2 0 2 2 1 1 2

P. curta 1 0 1 1 2 1 1 0 0 0 0 1 1 0 0 0 1 0 0 1 1 0 0 0 2 0 0 1 0 3 0 1 2 1 1 2

P. decipiens 0 0 0 1 0 1 0 0 1 1 1 0 0 0 1 1 0 0 0 2 0 0 0 1 0 1 1 0 1 1 1 2 2 1 1 2

P. ecuadoriensis 0 0 0 1 1 1 1 1 0 0 1 1 1 0 0 0 1 0 0 1 0 0 1 0 2 0 0 1 0 2 0 1 2 1 1 2

P. flexineuris 0 0 0 0 1 1 1 1 0 0 1 1 1 0 0 0 1 0 0 0 0 1 1 0 2 0 0 1 0 2 0 2 2 1 1 2

P. fellerae 0 0 0 0 1 1 1 1 0 0 1 1 1 0 0 0 3 0 0 0 0 0 0 0 2 0 0 1 0 2 0 2 2 1 1 2

P. fulgifrons 1 0 0 1 2 0 1 0 0 0 0 1 1 0 0 0 1 1 1 1 1 0 0 0 1 0 0 1 0 2 0 1 2 1 1 2

P. fusca 0 0 0 1 0 1 0 0 1 1 1 1 1 0 1 1 0 0 0 2 0 1 0 1 0 0 1 0 1 1 0 2 2 1 1 2

P. guttata 1 0 0 1 2 0 1 0 0 0 0 1 1 0 0 0 1 1 1 1 1 0 0 0 1 0 0 1 0 2 0 1 2 1 1 2

P. hirticornis 0 0 1 1 1 1 1 1 0 0 1 1 1 0 0 0 1 0 0 1 1 0 0 0 2 0 0 1 0 2 0 1 2 1 1 2

P. limbata 1 0 0 1 0 1 0 0 1 1 1 0 1 1 0 2 4 0 0 2 1 0 0 0 0 0 0 0 0 2 1 2 2 1 1 2

P. lineata 0 0 0 1 0 1 0 0 1 1 1 0 1 0 1 1 0 0 0 2 0 0 0 1 0 0 1 0 1 1 0 2 2 1 1 2

P. longiseta 0 0 0 1 0 1 0 0 1 1 1 0 0 0 1 1 0 0 0 2 0 0 0 1 0 1 1 0 1 1 1 2 2 1 1 2

P. maculata 0 0 0 0 1 1 1 1 0 0 1 1 1 0 0 0 1 0 0 1 1 0 1 0 2 0 0 1 0 2 0 1 2 1 1 2

P. malleata 0 0 0 0 1 1 1 1 0 0 1 1 1 0 0 0 1 0 0 1 0 0 1 0 2 0 0 1 0 2 0 2 2 1 1 2

P. meridionalis 0 0 0 1 1 1 1 1 0 0 1 1 1 0 0 0 1 0 0 1 0 0 1 0 2 0 0 1 0 2 0 2 2 1 1 2

P. millepuncta 1 1 0 0 1 1 1 ? 0 0 1 1 1 0 0 0 1 0 0 2 0 0 0 0 2 0 0 0 0 - 2 2 2 1 1 2

P. molossus 0 0 0 0 1 1 1 1 0 0 1 1 1 0 0 0 2 0 0 1 0 0 1 0 2 0 0 1 0 2 0 1 2 1 1 2

P. monstruosa 1 1 0 0 0 1 1 0 0 1 0 1 1 0 0 0 2 0 0 2 0 0 0 0 1 0 0 0 0 0 2 2 2 1 1 2

P.nebulosa 1 0 0 0 1 1 1 1 1 0 1 1 1 0 0 0 1 0 0 2 1 0 0 0 2 0 0 0 0 2 2 2 2 1 1 2

P. nidor 1 0 0 1 1 1 1 1 1 1 1 1 1 0 0 0 2 0 0 0 1 1 0 0 2 0 0 0 0 2 2 2 2 1 1 2

P. nigripes 1 0 0 1 1 1 1 1 1 0 1 1 1 0 0 0 2 0 0 0 1 1 0 0 2 0 0 0 0 2 2 2 2 1 1 2

P. nigropicta 0 0 0 0 1 1 1 1 0 0 1 1 1 0 0 0 1 0 0 0 0 0 1 0 2 0 0 1 0 2 0 2 2 1 1 2

P. nubifer 1 1 0 0 1 1 1 1 1 0 1 1 1 0 0 0 1 0 0 0 1 0 0 0 2 0 0 1 0 2 0 2 2 1 1 2

P. obscura 0 0 0 1 0 1 0 0 1 1 1 0 0 0 1 1 0 0 0 2 0 0 0 1 0 1 1 0 1 1 1 2 2 1 1 2

P. ornatifrons 1 0 1 0 2 0 1 1 0 0 1 1 1 0 0 0 1 0 0 1 1 0 0 0 2 0 0 0 0 2 0 2 2 1 1 2

P. pallida 0 0 0 0 1 1 1 1 0 0 1 1 1 0 0 0 1 0 0 1 0 0 1 0 2 0 0 1 0 2 0 1 2 1 1 2

P. pauca 0 0 0 1 1 1 1 1 0 0 1 1 1 0 0 0 1 0 0 1 0 0 1 0 2 0 0 1 0 2 0 1 2 1 1 2

P. pectinata 0 0 0 0 1 1 1 1 0 0 1 1 1 0 0 0 3 0 0 0 0 1 1 0 2 0 0 1 0 2 0 1 2 1 1 2

P. piger 0 0 0 0 1 1 1 1 0 0 1 1 1 0 0 0 1 0 0 0 0 1 1 0 2 0 0 1 0 2 0 1 2 1 1 2
number 643 • 129

Characters 1 1 1 1 1 1 1 1 1 1 2 2 2 2 2 2 2 2 2 2 3 3 3 3 3 3 3

Taxa 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6

P. pleurivittata 1 0 1 0 2 0 1 1 0 0 1 1 1 0 0 0 1 0 0 1 0 0 0 0 2 0 0 1 0 2 0 1 2 1 1 2

P. picta 1 0 0 1 1 1 1 0 0 0 1 1 1 0 0 0 1 0 0 2 0 0 0 0 2 0 0 0 0 2 0 2 2 1 1 2

P. plumbiceps 0 0 0 0 1 1 1 1 0 0 1 1 1 0 0 0 1 0 0 0 0 1 1 0 2 0 0 1 0 2 0 1 2 1 1 2

P. poecila 1 0 0 0 1 0 1 1 1 0 1 1 1 0 0 0 2 0 0 2 1 0 0 0 2 0 0 0 0 2 2 2 2 1 1 2

P. puncticollis 1 0 0 1 1 1 1 1 1 0 1 1 1 0 0 0 2 0 0 0 1 1 0 0 2 0 0 0 0 2 2 2 2 1 1 2

P. puncticornis 0 0 1 1 1 1 1 1 0 0 1 1 1 0 0 0 1 0 0 1 0 0 0 0 2 0 0 1 0 2 0 1 2 1 1 2

P. punctipennis 0 0 0 0 1 1 1 1 0 0 1 1 1 0 0 0 3 0 0 1 0 0 0 0 2 0 0 1 0 2 0 2 2 1 1 2

P. pupulata 1 0 0 1 1 1 1 1 1 0 1 1 1 0 1 0 1 0 0 2 1 0 0 0 2 0 0 1 0 2 0 2 2 1 1 2

P. quadrifascia 1 0 1 0 1 1 1 1 1 0 1 1 1 0 0 0 1 0 0 0 1 1 0 0 2 0 0 0 0 2 0 2 2 1 1 2

P. reticulata 1 0 0 1 0 1 0 0 0 1 0 1 1 1 0 2 4 0 1 2 1 0 0 0 0 0 0 0 0 ? 0 0 2 1 1 2

P. sana 0 0 0 0 1 1 1 1 0 0 1 1 1 0 0 0 1 0 0 1 0 0 0 0 2 0 0 1 0 2 0 1 2 1 1 2

P. secunda 0 0 0 0 1 1 1 1 0 0 1 1 1 0 0 0 1 0 0 1 0 0 0 0 2 0 0 1 0 0 0 1 2 1 1 2

P. sera 0 0 0 0 1 1 1 1 0 0 1 1 1 0 0 0 1 0 0 0 0 0 1 0 2 0 0 1 0 0 0 1 2 1 1 2

P. sinensis 1 0 1 0 2 0 1 1 0 0 1 1 1 0 0 0 1 0 0 1 1 0 0 0 2 0 0 0 0 2 0 2 2 1 1 2

P. sponsa 1 0 0 1 1 1 1 1 1 0 1 1 1 0 0 0 2 0 0 0 1 1 0 0 2 0 0 0 0 2 2 2 2 1 1 2

P. stellata 1 0 0 1 2 0 1 0 0 0 0 1 1 0 0 0 1 1 1 1 1 0 0 0 1 0 0 1 0 2 0 1 2 1 1 2

P. stigmata 0 0 0 0 1 1 1 1 0 0 1 1 1 0 0 0 1 0 0 1 0 0 0 0 2 0 0 1 0 2 0 1 2 1 1 2

P. taurus 1 0 0 1 2 0 1 1 0 0 1 1 1 0 0 0 1 0 0 1 0 0 0 0 2 0 0 1 0 2 0 2 2 1 1 2

P. multipunctata 0 0 0 0 1 1 1 1 0 0 1 1 1 0 0 0 1 0 0 0 0 1 1 0 2 0 0 1 0 2 0 2 2 1 1 2

P. texana 0 0 0 0 1 1 1 1 0 0 1 1 1 0 0 0 1 0 0 0 0 1 1 0 2 0 0 1 0 2 0 2 2 1 1 2

P. whirti 0 0 0 0 1 1 1 1 0 0 1 1 1 0 0 0 1 0 0 1 0 0 0 0 2 0 0 1 0 2 0 2 2 1 1 2

P. velutina 0 0 1 1 1 1 1 1 0 0 1 1 1 0 0 0 3 0 0 1 0 0 0 0 2 0 0 1 0 2 0 1 2 1 1 2

three Neotropical species of Phaiosterna, and the monophyletic 

group is supported by one synapomorphy: posteroventral margin of tergite 5 with dark shiny stain (261

).

The analysis strongly corroborates the monophyly of 

the genus Paralimna, as stated previously (Zatwarnicki and 

Mathis, 2002), and also shows the close relationship between 

Paralimna and Afrolimna. Finally, and based on these results, 

we recognize and propose another subgenus, Paralimna 

(Coganolimna), subg. nov., comprising the former limbata species group.

TABLE 1. (Continued)
130 • smithsonian contributions to zoology

FIGURES 162–215

FIGURES 162–165. Photographs of heads, anterior aspect (all scale bars = 0.5 mm): (162) Paralimna adunca, sp. nov. (Mexico. 

Morelos); (163) Paralimna argyrostoma Cresson (Costa Rica. Limón); (164) Paralimna aurantia, sp. nov. (Bolivia. La Paz); 

(165) Paralimna brunneiceps Cresson (Costa Rica. Guanacaste).
number 643 • 131

FIGURES 166–169. Photographs of heads, anterior aspect (scale bars = 0.5 mm): (166) Paralimna castanea, sp. nov. (Trinidad and 

Tobago. Tobago); (167) Paralimna crinita, sp. nov. (Costa Rica. Puntarenas); (168) Paralimna curta, sp. nov. (Brazil. Amazonas); (169) 

Paralimna ecuadoriensis, sp. nov. (Ecuador. Manabí).
132 • smithsonian contributions to zoology

FIGURES 170–173. Photographs of heads, anterior aspect (scale bars = 0.5 mm): (170) Paralimna fellerae Mathis (Belize. Stann 

Creek); (171) Paralimna flexineuris Cresson (Ecuador. El Oro); (172) Paralimna fulgifrons, sp. nov. (Peru. Loreto); (173) Paralimna 

guttata, sp. nov. (Brazil. Amazonas).
number 643 • 133

FIGURES 174–177. Photographs of heads, anterior aspect (scale bars = 0.5 mm): (174) Paralimna guttata, sp. nov. (Brazil. Amazonas); 

(175) Paralimna maculata, sp. nov. (Dominican Republic. Monseñor-Nouel); (176) Paralimna malleata, sp. nov. (Dominica. Layou Valley, Clarke Hall Estate); (177) Paralimna meridionalis Cresson (Costa Rica. San José).
134 • smithsonian contributions to zoology

FIGURES 178–181. Photographs of heads, anterior aspect (scale bars = 0.5 mm): (178) Paralimna meridionalis Cresson; (179) Paralimna 

molossus Schiner (Venezuela. Valle Seco); (180) Paralimna molossus Schiner (Bolivia. La Paz); (181) Paralimna thomae (Wiedemann).
number 643 • 135

FIGURES 182–185. Photographs of heads, anterior aspect (scale bars = 0.5 mm): (182) Paralimna nigropicta Cresson (Costa Rica. Puntarenas); (183) Paralimna pallida, sp. nov. (Brazil. Amazonas); (184) Paralimna pauca, sp. nov. (Dominican Republic. Independéncia); 

(185) Paralimna pectinata Hendel (Guyana. Karanambo).
136 • smithsonian contributions to zoology

FIGURES 186–189. Photographs of heads, anterior aspect (scale bars = 0.5 mm): (186) Paralimna piger Cresson (Ecuador. Guayas); 

(187) Paralimna pleurivittata Cresson (Peru. Madre de Dios); (188) Paralimna plumbiceps Cresson (Puerto Rico. Adjuntas); (189) 

Paralimna puncticornis Cresson (Costa Rica. Heredia).
number 643 • 137

FIGURES 190–193. Photographs of heads, anterior aspect (scale bars = 0.5 mm): (190) Paralimna punctipennis (Wiedemann) (USA. 

Virginia, Stafford); (191) Paralimna sana Cresson (Bolivia. Santa Cruz); (192) Paralimna secunda Schiner (Territorio Federal Amazonas); (193) Paralimna sera Cresson (Jamaica. St. Andrew).
138 • smithsonian contributions to zoology

FIGURES 194–197. Photographs of heads, anterior aspect (scale bars = 0.5 mm): (194) Paralimna stellata, sp. nov. (Ecuador. Pastaza); (195) 

Paralimna stigmata, sp. nov. (Brazil. Amazonas); (196) Paralimna texana Cresson (USA. Texas); (197) Paralimna velutina, sp. nov. (Guyana. 

Paramakatoi).
number 643 • 139

FIGURES 198–203. Photographs of wings, dorsal aspect: (198) Paralimna castanea, sp. nov. (Trinidad and Tobago. Tobago: St. John); (199) 

Paralimna crinita, sp. nov. (Costa Rica. Puntarenas); (200) Paralimna fulgifrons, sp. nov. (Brazil. Amazonas); (201) Paralimna guttata, sp. nov. 

(Brazil. Amazonas); (202) Paralimna guttata, sp. nov. (Brazil. Amazonas); (203) Paralimna malleata, sp. nov. (Dominica. Clarke Hall).
140 • smithsonian contributions to zoology

FIGURES 204–209. Photographs of wings, dorsal aspect: (204) Paralimna pauca, sp. nov. (Dominican Republic. Independéncia); (205) 

Paralimna pleurivittata Cresson (Peru. Madre de Dios); (206) Paralimna punctipennis (Wiedemann) (USA. Virginia); (207) Paralimna stellata, 

sp. nov. (Ecuador. Pastaza); (208) Paralimna stigmata, sp. nov. (Brazil. Amazonas); (209) Paralimna velutina, sp. nov. (Brazil. Pará).
number 643 • 141

FIGURES 210–213. Photographs of legs, anterior aspect (scale bars = 0.5 mm): (210) Paralimna brunneiceps Cresson (Costa Rica. San 

José); (211) Paralimna flexineuris Cresson (Ecuador. Manabí); (212) Paralimna molossus Schiner (Bolivia. La Paz); (213) Paralimna 

pectinata Hendel (Bolivia. Cochabama).
142 • smithsonian contributions to zoology

FIGURE 214. Consensus cladogram, with bootstrap values, of hypothetical relationships among species of Paralimna.
number 643 • 143

FIGURE 215. One of 100 trees, obtained from the analysis of the unordered 36 characters and 67 taxa, chosen to represent the hypothetical relationships among species of Paralimna. Consistency index = 0.40; retention index = 0.81; 127 steps; slow optimization used. Empty 

squares represent homoplasious synapomophies; filled squares represent non-homoplasious synapomophies.

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adunca, 9, 10–12, 130

Afrolimna, 126, 129

angustus, 5

appendiculata, 4–5, 76

argyrostoma, 6, 13–15, 130

aurantia, 9, 14–18, 130

bicolor, 127

bistriata, 51

boensis, 127

brunneiceps, 8, 18–20, 33, 50, 130, 141

captiosa, 51

castanea, 9, 20–23, 27, 131, 139

ciliata, 102

cilifera, 82

Coganolimna, 125

confluens, 127

cressoni, 127

crinita, 8, 13, 22–25, 97, 131, 139

curta, 8, 21, 25–27, 127, 131

curvirostris, 102

decipiens, 1, 111–118, 125, 127

ecuadoriensis, 9, 27–28, 131

fellerae, 7, 20, 28–33, 50, 66, 82, 132

flexineuris, 6, 13, 33–36, 132, 141

fulgifrons, 6, 36–38, 42, 93, 127, 132, 139

gracilior, 102, 112

guttata, 6, 37, 38–42, 93, 127, 132, 133, 139

hirticornis, 127

jamaicensis, 102

Index
148 • smithsonian contributions to zoology

limbata, 125, 127

longiseta, 112, 117–119, 127

luctans, 51

maculata, 7, 42–45, 91, 133

malleata, 9, 45–47, 133, 139

meridionalis, 9, 26, 33, 47–51, 

133, 134

millepuncta, 127

molossus, 1, 10, 51–55, 62, 96, 134, 

141

monstruosa, 127

multipunctata, 102, 109

nigropicta, 6, 20, 33, 50, 54–58, 135

obscura, 112, 117, 119–125, 127

Oedenops, 126

pallida, 8, 58–60, 135

Papuama, 126

Paralimna, 1–143

paralimnae, 102, 112

pauca, 10, 60–62, 135, 140

pectinata, 7, 62–66, 82, 135, 

141

Phaiosterna, 2, 5, 111–125, 

129

picta, 127

piger, 7, 41, 66–68, 136

pleurivittata, 8, 68–71, 97, 136, 

140

plumbiceps, 7, 47, 71–72, 136

puncticollis, 5

puncticornis, 10, 68, 72–76, 136

punctipennis, 1, 4, 5, 7, 33, 50–51, 66, 

76–82, 137, 140

recurvatus, 102

reticulata, 127

rostratus, 102, 112

sana, 9, 82–84, 137

secunda, 1, 8, 84–89, 137

sera, 7, 44, 62, 89–91, 137

stellata, 6, 37, 42, 60, 91–93, 96, 127, 

138, 140

sticta, 96

stigmata, 8, 53, 93–96, 138, 140

Stigmatomyces, 102, 112

taurus, 8, 96–98

texana, 7, 36, 58, 98–102, 108, 138

thomae, 1, 7, 18, 35, 58, 102–109, 134

velutina, 8, 109–111, 138, 140
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