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smithsonian contributions to zoology • number 633

A Revision of the Genus 

Aulacigaster Macquart

(Diptera: Aulacigastridae)

Alessandra Rung and Wayne N. Mathis
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smithsonian contributions to zoology • number 633

A Revision of the Genus 

Aulacigaster Macquart

(Diptera: Aulacigastridae)

Alessandra Rung and Wayne N. Mathis

washington d.c.

2011
ABSTRACT

Rung, Alessandra, and Wayne N. Mathis. A Revision of the Genus Aulacigaster Macquart (Diptera: Aulacigastridae). Smithsonian Contributions to Zoology, number 633, x + 132 pages, 220 figures, 18 maps, 

2011.—The world’s described species of the genus Aulacigaster Macquart, now numbering 55, are revised. 

New species from the Afrotropical (2 species), Neotropical (37 species), and Oriental Regions (2 species) are 

described, and the genus is divided into seven species groups, of which six are Neotropical. Keys to the species groups and to the known species occurring in the Afrotropical, Nearctic, Neotropical, Palearctic, and 

Oriental Regions are provided. Diagnoses, detailed distributional data for species of the genus, notes on the 

biology, and illustrations (photographs and drawings) are included to facilitate species identification. A phylogenetic analysis was performed to test the monophyly of the genus Aulacigaster and to discover relationships 

between included species, hence indicating the monophyly of the species groups. The ingroup includes a total 

of 24 exemplar congeners. Outgroup sampling includes exemplars from the putative sister group, Curiosimusca. Analyses with and without successive weighting recovered a monophyletic Aulacigaster and indicated 

clades within the genus.

Cover images: left, Aulacigaster korneyevi, new species (♀); right, head of A. proxima.

Published by SMITHSONIAN INSTITUTION SCHOLARLY PRESS

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Library of Congress Cataloging-in-Publication Data

Rung, Alessandra.

A revision of the genus Aulacigaster Macquart (Diptera:Aulacigastridae) / Alessandra Rung and Wayne N. 

Mathis.

 p. cm. — (Smithsonian contributions to zoology; number 633)

Includes bibliographical references and index.

1. Aulacigaster—Classification. I. Mathis, Wayne N. II. Title. 

QL537.A9R86 2011

595.77’3—dc22

2010042836

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

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 vii

LIST OF MAPS ix

INTRODUCTION 1

Methods and Materials 2

Acknowledgments 3

SYSTEMATICS 4

Genus Aulacigaster Macquart 4

 Implicit Attributes 6

 Key to the Neotropical Species Groups and Species of 

Aulacigaster Macquart 6

 The bromeliae Group 9

 Aulacigaster bromeliae, new species 10

 Aulacigaster korneyevi, new species 12

 Aulacigaster lopesi, new species 14

 Aulacigaster serrana, new species 17

 Aulacigaster tibanae, new species 18

 Aulacigaster vespertina, new species 19

 The ecuadoriensis Group 21

 Aulacigaster aenigma, new species 21

 Aulacigaster ecuadoriensis (Hennig) 22

 Aulacigaster fastidiosa, new species 24

 Aulacigaster melanoleuca (Hennig) 27

 Aulacigaster stenoptera, new species 28

 The femorata Group 31

 Aulacigaster bella, new species 32

 Aulacigaster erwini, new species 33

 Aulacigaster femorata, new species 35

 Aulacigaster formosa, new species 37

 Aulacigaster guyana, new species 38

 Aulacigaster irwini, new species 39
i v • smithsonian contributions to zoology

 Aulacigaster lobata, new species 41

 Aulacigaster nigritibia, new species 43

 Aulacigaster peruana, new species 44

 Aulacigaster unifasciata, new species 46

 The grimaldii Group 48

 Aulacigaster angusta, new species 48

 Aulacigaster grimaldii, new species 49

 The minuta Group 52

 Aulacigaster appendiculata, new species 53

 Aulacigaster condylura, new species 54

 Aulacigaster costaricana, new species 57

 Aulacigaster curvata, new species 58

 Aulacigaster gaimarii, new species 60

 Aulacigaster minuta (Hennig) 61

 Aulacigaster nigritarsa, new species 63

The plesiomorphica Group 64

 Aulacigaster albifacies, new species 64

 Aulacigaster atriseta, new species 66

 Aulacigaster belize, new species 68

 Aulacigaster bifasciata, new species 69

 Aulacigaster conspicua, new species 71

 Aulacigaster erika, new species 73

 Aulacigaster plesiomorphica (Hennig) 75

 Aulacigaster proxima, new species 76

 Aulacigaster rufifemur, new species 78

 Aulacigaster trifasciata, new species 82

The leucopeza Group 83

 Afrotropical Species 83

 Key to the Afrotropical Species of Aulacigaster 83

 Aulacigaster africana Barraclough 84

 Aulacigaster borbonica Hilger and Kassebeer 85

 Aulacigaster freidbergi, new species 86

 Aulacigaster malawana, new species 88

 Aulacigaster perata Barraclough 89

 Nearctic Species 90

 Key to the Nearctic Species of Aulacigaster 90

 Aulacigaster mcalpinei Mathis and Freidberg 90

 Aulacigaster neoleucopeza Mathis and Freidberg 92

 Aulacigaster sabroskyi Mathis and Freidberg 98

 Neotropical Species 100

 Key to the Neotropical Species of the leucopeza

Group of Aulacigaster 100

 Aulacigaster spangleri, new species 101

 Aulacigaster L1, undescribed species 101

 Aulacigaster L2, undescribed species 103

 Palearctic Species 104

 Key to the Palearctic Species of Aulacigaster 104

 Aulacigaster afghanorum Papp 104

 Aulacigaster falcata Papp 105

 Aulacigaster leucopeza (Meigen) 106
number 633 • v

 Aulacigaster pappi Kassebeer 109

 Oriental Species 112

 Key to the Oriental Species of Aulacigaster 112

 Aulacigaster india, new species 112

 Aulacigaster srilanka, new species 113

 Aulacigaster L3, undescribed species 114

PHYLOGENETIC ANALYSIS AND DISCUSSION 115

Characters Used in the Cladistic Analysis 115

Discussion 116

FIGURES 203–220 121

APPENDIX: Character Matrix 125

REFERENCES 127

INDEX to the species of aulacigaster 131

Figures

1. Frontispiece: Aulacigaster korneyevi, new species xii

2–4. Aulacigaster bromeliae, new species (male) 11

5–8. Aulacigaster bromeliae, new species 11

9–14. Aulacigaster korneyevi, new species (male) 14

15. Aulacigaster korneyevi, new species (male) 15

16. Aulacigaster lopesi, new species (male) 16

17. Aulacigaster serrana, new species (male) 17

18–20. Aulacigaster tibanae, new species (male) 18

21–25. Aulacigaster vespertina, new species (male) 20

26–29. Aulacigaster aenigma, new species (male) 22

30–31. Aulacigaster aenigma, new species (male) 23

32–35. Aulacigaster ecuadoriensis (Hennig) 25

36–38. Aulacigaster ecuadoriensis (Hennig) (male) 26

39–45. Aulacigaster fastidiosa, new species 27

46–47. Aulacigaster fastidiosa, new species (male) 28

48–56. Aulacigaster melanoleuca (Hennig) 29

57–58. Aulacigaster melanoleuca (Hennig) (male) 30

59–61. Aulacigaster stenoptera, new species (male) 31

62–65. Aulacigaster bella, new species (male) 33

66–68. Aulacigaster bella, new species (male) 34

69. Aulacigaster erwini, new species (male) 36

70–71. Aulacigaster femorata, new species (male) 36

72–73. Aulacigaster femorata, new species (male) 37

74–76. Aulacigaster formosa, new species (male) 38

77–79. Aulacigaster guyana, new species (male) 40

80–81. Aulacigaster irwini, new species (male) 40

82–83. Aulacigaster irwini, new species (male) 41

84–85. Aulacigaster lobata, new species 42

86–87. Aulacigaster lobata, new species (male) 43

88–91. Aulacigaster nigritibia, new species (male) 45

92–93. Aulacigaster peruana, new species (male) 46

94–95. Aulacigaster peruana, new species (male) 46
viii • smithsonian contributions to zoology

96–98. Aulacigaster unifasciata, new species (male) 47

99. Aulacigaster angusta, new species (male) 49

100–104. Aulacigaster grimaldii, new species 51

105–106. Aulacigaster grimaldii, new species (male) 52

107–110. Aulacigaster appendiculata, new species (male) 54

111. Aulacigaster appendiculata, new species (male) 55

112–114. Aulacigaster condylura, new species (male) 56

115. Aulacigaster condylura, new species (male) 57

116–119. Aulacigaster costaricana, new species (male) 58

120. Aulacigaster costaricana, new species (male) 59

121–122. Aulacigaster curvata, new species (male) 59

123. Aulacigaster curvata, new species (male) 60

124–125. Aulacigaster gaimarii, new species 61

126–127. Aulacigaster gaimarii, new species (male) 61

128–129. Aulacigaster minuta (Hennig) (male) 62

130–131. Aulacigaster nigritarsa, new species (male) 63

132–133. Aulacigaster albifacies, new species (male) 65

134–135. Aulacigaster atriseta, new species (male) 67

136. Aulacigaster atriseta, new species (male) 68

137–138. Aulacigaster belize, new species (male) 70

139–140. Aulacigaster bifasciata, new species (male) 71

141–144. Aulacigaster conspicua, new species (male) 72

145–146. Aulacigaster conspicua, new species (male) 73

147. Aulacigaster erika, new species (male) 75

148–151. Aulacigaster plesiomorphica (Hennig) and A. proxima 76

152–154. Aulacigaster plesiomorphica (Hennig) (male) 77

155. Aulacigaster proxima, new species 78

156–159. Aulacigaster rufifemur, new species (male) 80

160. Aulacigaster trifasciata, new species (male) 82

161–164. Aulacigaster africana Barraclough (male) 85

165. Aulacigaster africana Barraclough (male) 86

166–167. Aulacigaster freidbergi, new species (male) 88

168–169. Aulacigaster malawana, new species (male) 89

170–171. Aulacigaster mcalpinei Mathis and Freidberg (male) 91

172–177. Aulacigaster neoleucopeza Mathis and Freidberg 94

178–182. Aulacigaster neoleucopeza Mathis and Freidberg (male) 96

183–184. Aulacigaster sabroskyi Mathis and Freidberg (male) 100

185–186. Aulacigaster L2, undescribed species (female) 103

187. Aulacigaster falcata Papp (male) 106

188–191. Aulacigaster leucopeza (Meigen) (male) 108

192–193. Aulacigaster leucopeza (Meigen) (male) 109

194–197. Aulacigaster pappi Kassebeer (male) 110

198–199. Aulacigaster pappi Kassebeer (male) 111

200. Aulacigaster india, new species (male) 113

201. Aulacigaster srilanka, new species (male) 115

202. Tree representing phylogenetic relationships between species 

groups of Aulacigaster 118

203–208. Heads of Aulacigaster species 121

209–214. Heads of Aulacigaster species 122

215–220. Heads of Aulacigaster species 123
Maps

1. Distribution of Aulacigaster bromeliae, A. lopesi, A. serrana, and 

A. tibanae 13

2. Distribution of Aulacigaster korneyevi and A. vespertina 16

3. Distribution of the ecuadoriensis species group of Aulacigaster 24

4. Distribution of Aulacigaster bella, A. erwini, A. femorata, and 

A. guyana 35

5. Distribution of Aulacigaster formosa, A. irwini, A. peruana, and 

A. unifasciata 39

6. Distribution of Aulacigaster lobata and A. nigritibia 44

7. Distribution of the grimaldii species group of Aulacigaster 50

8. Distribution of the minuta species group of Aulacigaster 56

9. Distribution of Aulacigaster albifacies and A. bifasciata 66

10. Distribution of Aulacigaster atriseta, A. belize, and 

A. plesiomorphica 69

11. Distribution of Aulacigaster conspicua and A. erika 74

12. Distribution of Aulacigaster proxima and A. trifasciata 79

13. Distribution of Aulacigaster rufifemur 81

14. Distribution of the Afrotropical species of Aulacigaster 87

15. Distribution of Aulacigaster mcalpinei 93

16. Distribution of Aulacigaster neoleucopeza and A. sabroskyi 99

17. Distribution of the Neotropical species of the leucopeza group of 

Aulacigaster 102

18. Distribution of the Oriental species of the leucopeza group of 

Aulacigaster 114
FIGURE 1. Frontispiece of Aulacigaster korneyevi, new species (Costa Rica. Heredia: Santo Domingo, ♀).
A Revision of the Genus Aulacigaster

Macquart (Diptera: Aulacigastridae)

Introduction

This revision concerns Aulacigaster Macquart, an obscure genus of Acalyptrate Diptera that until recently was the only included genus in the family 

Aulacigastridae. Until this revision, Aulacigaster included relatively few species 

that were usually associated with slime fluxes of deciduous trees (Robinson, 

1953; Cole and Streams, 1970; Davis and Zack, 1978; Ferrar, 1987; Mathis 

and Freidberg, 1994; Hilger and Kassebeer, 2000; Papp, 2008) or with the phytotelmata of bromeliads (A. R and WNM, personal observations). Most previously described species are from temperate zones; however, we document far 

greater species diversity in the subtropics and tropics (Afrotropical, Holarctic, 

Neotropical, and Oriental Regions). The genus has received increased attention 

during the last decade not only because it has revealed surprising diversity and 

species richness, but also because of challenges that its classification and characterization have posed to systematists. The primary purposes for this revision are 

to (1) describe and classify the largely unknown fauna of Aulacigaster (Mathis 

and Freidberg, 1994; Baptista, 1998a) within a phylogenetic context and (2) 

clarify some issues concerning the phylogenetic placement and characterization 

of the family.

Historical and Taxonomic Background. Aulacigaster is 

currently classified in the family Aulacigastridae, superfamily Opomyzoidea (J. F. 

McAlpine, 1989). Characterization of Aulacigastridae differs widely among authors (see below), and up to six genera have been assigned to the family in addition to the nominate genus, Aulacigaster. Duda (1924) first accorded familial 

status to Aulacigastridae, and recent authors (Mathis and Freidberg, 1994; Papp, 

1998a; Baptista, 1998b; and Hilger and Kassebeer, 2000) have adopted a monotypic characterization. A broader characterization of the family includes the genera Cyamops Melander, Stenomicra Coquillett, Planinasus Cresson, and a fossil 

Alessandra Rung, Plant Pest Diagnostics 

Branch, California Department of Food and 

Agriculture, 3294 Meadowview Road, Sacramento, California 95832-1448, USA; Wayne N.

Mathis, Department of Entomology, National 

Museum of Natural History, Smithsonian Institution, P.O. Box 37012, NHB CE-619, MRC 

169, Washington, D.C. 20013-7012, USA. 

Correspondence: mathisw@si.edu. Manuscript 

received 29 March 2010, accepted 26 July 2010.
2 • smithsonian contributions to zoology

species from Baltic amber, Protaulacigaster electrica Hennig 

(described in 1965). Hennig (1958, 1965, 1969, 1971) first 

advocated this characterization, which was followed by 

Griffiths (1972, with the exclusion of Stenomicra), Teskey 

(1987) and J. F. McAlpine (1989). D. K. McAlpine (1978, 

1983) proposed a rather different characterization that: 

(1) transferred the genera Stenomicra, Cyamops (1978), 

and Planinasus (1983) from Aulacigastridae to the family Periscelididae; (2) excluded Protoaulacigaster electra

from Aulacigastridae; and (3) described two new genera 

in the family, Ningulus (Afrotropical Region) and Nemo

(Australian Region). Freidberg (1994) proposed the family 

Neminidae for Ningulus, Nemo, and his new Afrotropical 

genus, Nemula. A result of Freidberg’s classification was a 

return to a monotypic concept for Aulacigastridae. Most 

recently, Evenhuis (1989, 1994a, 1994b) listed Protaulacigaster electrica in Aulacigaster, and Rohácˇek (1998) tentatively transferred the genus Echidnocephalodes Sabrosky 

from Anthomyzidae to Aulacigastridae. Rung et al. (2005) 

described the genus Curiosimusca in the family, with three 

species from the Oriental Region. The monophyly of Aulacigaster + Curiosimusca is supported by seven putative 

synapomorphies (Rung et al., 2005). Hennig’s original concept of Aulacigastridae (see above) has been recently resurrected in a molecular phylogeny of the Opomyzoidea, using 

the 28S ribosomal DNA and CAD (rudimentary) genes 

(Winkler et al, 2010). In that analysis (which did not include examplars of Curiosimusca), Stenomicra, Cyamops

and Planinasus grouped consistently and with moderate 

support with the genus Aulacigaster.

The genus Aulacigaster was originally described for 

A. rufitarsis Macquart (1835), a junior synonym of A. leucopeza (Meigen, 1830). The genus remained monotypic 

for more than a century following its publication, and A. 

leucopeza was believed to occur in the Palearctic (type locality) and Nearctic Regions (Malloch and McAtee, 1924; 

Christianson and Ryckman, 1955; Wirth, 1965; Teskey, 

1976; Davis and Zack, 1978). Hennig (1956) described 

the genus Schizochroa in Aulacigastridae for a new species, S. melanoleuca, from Costa Rica. Thirteen years 

later, Hennig (1969) added three more species to Schizochroa: S. plesiomorphica (Peru), S. minuta (Ecuador), and 

S. ecuadoriensis (Ecuador). D. K. McAlpine (1983) synonymized the genus Schizochroa with Aulacigaster.

Interest in the taxonomy of Aulacigaster has steadily 

escalated during the last two decades. Barraclough (1993) 

revised the Afrotropical fauna and described two new 

species, A. africana (South Africa) and A. perata (Cameroon). A year later, Mathis and Freidberg (1994) revised the Nearctic fauna, describing three new species: A. 

neoleucopeza and A. mcalpinei, which occur sympatrically 

and frequently at the same flux, and A. sabroskyi. The latter authors (op. cit.) also noted that previous records of A. 

leucopeza for the Nearctic Region were misidentifications 

of A. neoleucopeza. Papp (1998a) briefly reviewed the Palearctic fauna of Aulacigaster and described two new species found in that region: A. afghanorum (Afghanistan) 

and A. falcata (Hungary). Papp (1998a) also reported the 

presence of A. neoleucopeza in the Palearctic Region, noting that previous distributional records of A. leucopeza

need verification. Recently, Hilger and Kassebeer (2000) 

described a new species, A. borbonica, from specimens 

collected in the Afrotropical Region (island of Reunión), 

and Kassebeer (2001) proposed A. pappi from the Palearctic Region for misidentified Palearctic specimens of 

A. neoleucopeza.

Until the present study, Aulacigaster has included 

14 valid species, although a large number of undescribed 

species has been reported from the Neotropical Region 

(Mathis and Freidberg, 1994; Baptista, 1998a). Keys are 

available for the Afrotropical (Barraclough, 1993), Nearctic (Mathis and Freidberg, 1994), and Palearctic (Papp, 

1998a; Kassebeer, 2001) Regions.

Methods and Materials

We have adopted the descriptive terminology of J. F. 

McAlpine (1981), with the modifications noted below. 

We follow Sabrosky (1983) in using the term “microtomentum” rather than pruinosity. Setae posterior to the 

pseudovibrissal seta are called peristomal setae. The nomenclature for structures of the male terminalia follows 

Cumming et al. (1995) and, for larval morphology, follows Courtney et al. (2000).

Procedures. Label data of holotypes are 

quoted verbatim as they appear on the label with a slash (/) 

denoting the end of a label. Clarifying comments are included within brackets.

Because specimens are small, study and illustration 

of the male and female terminalia required use of a compound microscope. Photographs were taken with a Nikon 

Coolpix 9907 digital camera and an Auto-Montage Digital 

Imaging System7, using a Wild Photomakroskop M4007 

stereomicroscope. The photographs were enhanced, formatted, and arranged in Adobe Photoshop7.

Characters and information on their states within 

taxa were stored in DEscriptive Language for TAxonomy 

(DELTA) 7 files (Dallwitz, 1980; Pankhurst, 1991). The 

descriptions and keys were generated using the DELTA 

software package (Dallwitz, 1980; Dallwitz et al., 1993). 
number 633 • 3

Species descriptions are composite, including information 

on the holotype and other conspecific specimens. Characters used to characterize the genus and species groups 

are not repeated in the species descriptions. To generate 

such inclusive descriptions, we wrote a separate “tonath” 

DELTA file for the diagnosis of the genus Aulacigaster and 

species groups, and a tonath DELTA file for each of the six 

species groups. Species diagnostic descriptions were generated with the aid of Intkey (Dallwitz, 1980; Dallwitz et al., 

1993). They distinguish each taxon in at least one respect 

from every other taxon in the genus.

Distribution maps were made using ESRI ArcView7 

GIS 3.2 (ESRI, Redlands, Calif.). 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.

The following ratios are used in the descriptions (two 

males and two females were measured for each species, 

when available; cases to the contrary are indicated under 

the appropriate species description):

1. Scutal ratio: the greatest length of scutum divided 

by the greatest width of scutum.

2. Head ratio: the greatest length of head divided by 

the greatest height of head.

Phylogenetic Analysis. The phylogenetic 

analysis was performed to test the monophyly of the species groups proposed in this study and their relationships to 

each other. The monophyly of the genus Aulacigaster and 

its sister group relationship to Curiosimusca have been discussed in a former contribution (Rung et al., 2005). Readers 

should consult this paper for a list and discussion of putative synapomorphies. To accomplish our goals, an analysis 

including all of the component species of the genus was unnecessary. We opted, therefore, the exemplar method for 

selecting taxa (Yeates, 1995). At least two exemplars representing each species group were included in the analysis. 

If the monophyly of a group seemed doubtful, we included 

additional exemplars from that group. This procedure 

was used with the leucopeza, femorata, and ecuadoriensis

groups. Twenty-four exemplar species were included in the 

ingroup (see Appendix 1). The outgroup was selected from 

the genus Curiosimusca, the sister group of Aulacigaster.

New and traditional morphological characters were 

combined in a quantitative phylogenetic analysis. We endeavored to include characters that seemed independent 

and informative at the generic and species group levels. 

Only characters that showed obvious discrete variation 

were included. Characters were coded for the analysis as 

either binary (17 characters) or qualitative multistate (7 

characters).

Analyses were done by using PAUP 4.0 (Swofford, 

2002). The matrix was initially analyzed with a branchand-bound search, with all characters unordered and 

equally weighted. Following initial parsimony analysis, 

successive weighting (Farris, 1969) was applied by using 

Farris’ (1989) rescaled consistency index, an approach 

that gives highest weight to characters that have the least 

amount of homoplasy for the given set of trees.

Acknowledgments

Although most specimens for this study, including 

primary types, are in the National Museum of Natural 

History (USNM), numerous others were borrowed, particularly type specimens of species previously described. 

To our colleagues and their institutions listed below who 

loaned specimens, we express our sincere thanks. Without their cooperation this study could not have been 

completed. (Institutions are abbreviated for subsequent 

mention in the text.)

ABSF Archbold Biological Station, Lake Placid, 

Florida (Dr. Mark Deyrup)

AMNH American Museum of Natural History, 

New York, New York (Dr. David A. 

Grimaldi)

CAS California Academy of Sciences, San Francisco, California (Dr. Paul H. Arnaud, Jr.)

CNC Canadian National Collection, Ottawa, 

Ontario, Canada (Dr. Jeffrey M. Cumming 

and Mr. Bruce Cooper)

CSCA California State Collection of Arthropods, 

Sacramento, California (Dr. Stephen D. 

Gaimari)

DEI Deutsches Entomologisches Institut, 

Müncheberg, Germany (Dr. Frank Menzel 

and Joachim Ziegler)

EPNE Escola Politécnica Nacional, Quito, Ecuador (Mr. Adrián E. Troya)

GWMP George Washington Memorial Parkway, National Park Service, Turkey Run, McLean, 

Virginia (Mr. Brent W. Steury)

HNHM Hungarian Natural History Museum, Budapest, Hungary (Dr. László Papp)

INBio Instituto Nacional de Biodiversidad, Santo 

Domingo, Heredia, Costa Rica (Mr. Manuel Zumbado)
4 • smithsonian contributions to zoology

MHNJP Museo de Historia Natural “Javier Prado,” 

Universidade Nacional Mayor de San Marcos, Lima, Peru (Dr. Gerardo Lamas)

MNRJ Museu Nacional do Rio de Janeiro, Rio de 

Janeiro, Brazil (Dr. Márcia Souto Couri)

MZUSP Museu de Zoologia da Universidade de São 

Paulo, São Paulo, Brazil (Dr. Francisca do 

Val and Dr. Carlos Lamas)

NMSA Natal Museum, Pietermaritzburg, South 

Africa (Dr. David A. Barraclough)

TAU Tel Aviv University, Tel Aviv, Israel (Dr. 

Amnon Freidberg)

UCLA University of California, Los Angeles, California (Dr. Henry A. Hespenheide)

USU Utah State University insect collection, 

Logan, Utah (Dr. James Pitts)

USNM former United States National Museum, 

collections in the National Museum of 

Natural History, Smithsonian Institution, 

Washington, D.C.

UZMC Universitetets Zoologiske Museum, Copenhagen (Dr. Thomas Pape)

UGE University of Guelph, Guelph, Canada 

(Drs. Stephen A. Marshall and Matthias 

Buck)

ZMB Museum für Naturkunde, HumboldtUniversität, Berlin, Germany (Dr. Joachim 

Ziegler)

We are indebted to Dr. Hugo de Souza Lopez, Dr. 

Amnon Freidberg, and Ms. Rita Tibana for graciously 

providing us with their preliminary but valuable notes and 

ideas on various taxonomic aspects of Aulacigaster. They 

had earlier started research projects on this family. Dr. 

Charles Mitter supervised the dissertation research of the 

first author. His patience, support, and continuous guidance are acknowledged and appreciated. We also thank 

the members of first author’s academic committee, Drs. 

Jeffrey Shultz, Charles Delwiche, and David Hawthorne. 

We also thank Dr. Marion Kotrba, a friend and colleague 

throughout this research, who gave us invaluable information on the female genital tract of acalyptrates. The frontispiece was beautifully rendered by Mr. Young T. Sohn. The 

late Ms. Hollis B. Williams provided technical support. 

Mr. Kirk Sorensen produced the maps. Drs. Stephen D. 

Gaimari and Douglass R. Miller generally encouraged and 

supported our research on this project. For reviewing a 

draft of this paper we thank Dr. Allen L. Norrbom.

This research and much of the fieldwork was generously funded by a grant from the Partnerships for 

Enhancing Expertise in Taxonomy program (PEET), 

National Science Foundation (PEET 952-1773). We are 

grateful to PEET for the opportunity to learn and practice 

taxonomy, and for allowing the first author to open further 

the fascinating world of Diptera or true flies. Some of the 

fieldwork in Brazil was funded by the Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP), which 

we also acknowledge and thank. The Department of Entomology, University of Maryland (College Park, Maryland), 

provided the first author with graduate teaching assistantships and helped finance the later stages of this research.

Final aspects of this research, especially the preparation of the manuscript for publication, were supported by 

the Williston Fund, Department of Entomology, Smithsonian Institution.

Fieldwork in the West Indies was funded largely by 

grants from the Biodiversity Program (Biological Surveys 

and Inventories), National Museum of Natural History, 

Smithsonian Institution (Dr. Lynne R. Parenti and Dr. 

George R. Zug, former chairs).

Fieldwork in Guyana was supported by the Smithsonian Institution’s Biological Diversity of the Guiana Shield 

Program (publication number 163; Dr. Vicki A. Funk, Director; Dr. Carol L. Kelloff, Coordinator).

Fieldwork in Mongolia was generously supported by a 

grant entitled “Survey and Inventory of the Aquatic Insects 

of the Altai and Hangai Mountains’ drainages, Mongolia” 

awarded to Dr. J. Gelhaus, Dr. J. Morse, Dr. C. R. Nelson, 

United States National Science Foundation: Biodiversity Inventories Program (DEB-0743732), awarded for 2008–2011. 

We are especially indebted to the principal investigators, Dr. 

Jon K. Gelhaus in particular, for inviting us to participate in 

the program and fieldwork in the summer of 2009.

Fieldwork at Turkey Run, which is part of the George 

Washington Memorial Parkway, was supported by a grant 

from the National Park Service, and we especially thank 

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

support.

This is contribution number 876 from the Caribbean 

Coral Reef Ecosystems (CCRE), Smithsonian Institution, 

which is partly supported by a grant from the Exxon 

Corporation.

Systematics

Genus Aulacigaster Macquart

Aulacigaster Macquart, 1835:579. Type species: Aulacigaster rufitarsis

Macquart, 1835 (= Diastata leucopeza Meigen, 1830), by mono-
number 633 • 5

typy.—Schiner, 1863:269–270 [review, synonymy].—Duda, 1934:1–5 

[revision, Palearctic fauna].—Wirth, 1965:823 [Nearctic catalog].—

Papavero, 1967:91 [Neotropical catalog].—Teskey, 1976:12 [Diptera 

associated with trees]; 1987:891–894 [Manual of Nearctic Diptera].—

Sabrosky, 1977:230 [Oriental catalog].—Papp, 1984:61 [Palearctic 

catalog]; 1998a:225–234 [revision, Palearctic species]; 1998b:279–284 

[Manual of Palearctic Diptera].—Barraclough, 1993:331–342 [revision, Afrotropical species].—Mathis and Freidberg, 1994:586–591 

[Nearctic species].—Bächli, 1998:288 [checklist, Switzerland]; Bächli 

et al., 1999:119–122 [checklist, Greece, Italy, Switzerland].—Hilger 

and Kassebeer, 2000:167–172 [new species, Réunion].—Kassebeer, 

2001:23–32 [review, new species, western Palearctic Region].

Apotomella Dufour, 1846:455. Type species: Apotomella impressifrons

Dufour, 1846 [= Diastata leucopeza (Meigen, 1830)], by monotypy.—

Schiner, 1863:269 [synonymy with Aulacigaster].

Ampycophora Wahlberg, 1847:261. Type species Ampycophora tarsata

Wahlberg, 1847 [= Diastata leucopeza (Meigen, 1830)], by monotypy.—Schiner, 1863:269 [synonymy with Aulacigaster].

Sephanilla Rondani, 1874:276. Type species: Sephanilla sertulata Rondani, 

1874 [= Diastata leucopeza Meigen, 1830)], by original designation (p. 

246).—Mathis and Freidberg, 1994:585 [synonymy with Aulacigaster].

Aulacogaster Oldenberg, 1914:30 [unjustified emendation for Aulacigaster].

Schizochroa Hennig, 1956:151–152. Type species: Schizochroa melanoleuca

Hennig, 1956, by original designation.—D. K. McAlpine, 1983:58 

[synonymy with Aulacigaster].

Description. Adult: Coloration: Body predominantly dark brown to black. Cervix gray microtomentose. Frons often with anterior 1/4–1/2 yellowish in ground 

color or bearing a transverse, large, orange band. Scutum 

typically bearing microtomentum; ventral half of thoracic 

pleura in some species white to pale yellow. Wing usually 

mostly hyaline, partly infuscate with brown in many species, rarely mostly brown infuscate.

Morphology: Body robust (scutal ratio 1.10–1.35), 

or elongate (scutal ratio 1.50–1.75). 

Head: Higher than long (head ratio less than 0.9) 

or longer than high (head ratio higher than 1.1); face completely sclerotized, often protuberant, easily visible in lateral 

view; gena narrow, usually less than 1/3 the width of 1st 

flagellomere. First flagellomere round, oval or triangular; 

arista unsegmented, often rayed (appearing naked under 

low magnification in some species), zigzagged on apical 

2/3, dorsal and ventral rays alternating; basal 1/3 of arista 

naked or bearing a double row of long rays dorsally; ocellar 

seta, when present, minute; medial vertical setae typically 

slightly convergent, seta at most 3/4 length of lateral vertical seta; fronto-orbital setae 2, in some species with posterior seta medial to and almost horizontally aligned with 

anterior seta; setulae between fronto-orbital setae 1–2, one 

proclinate; intrafrontal seta absent; peristomal vestiture 

typically consisting of weaker setae posterior to pseudovibrissal seta.

Thorax: Anepisternum bearing 1–2 setae along 

posterior margin; scutellar setae 2, apical pair of setae nearly 

parallel, curved posteriorly. Cell dm flat, lacking a longitudinal crease or fold; crossvein r-m basal to middle of combined 

cell dm + bm; subcosta partially fused with vein R1

 apically.

Male abdomen and terminalia: Pregenital segment 

large, at least twice as long as tergite 5. Abdominal sternites 

slightly increasing in width posteriorly. Surstylus fused with 

epandrium; cerci separate or partially fused; subepandrial 

sclerite with caudal portion attached to cerci and proximal 

portion attached to hypandrial bridge; hypandrium bearing a lateral, anterior apodeme at base of each gonopod, 

halves of hypandrium connected posteriorly by a narrow 

bridge above phallapodeme; gonopods fused to hypandrium, forming a shovel-like structure, gonopodal halves 

connected posteroventrally, each bearing a posterior lobe or 

process and one to a few central setae; phallapodeme cuneiform with asymmetrical posterolateral projections; phallus 

short, partially sclerotized; parameres inconspicuous.

Female abdomen and terminalia: Postabdomen 

distinctly shorter than preabdomen. Three spermathecae 

present, often with ventral digitiform projections.

Immatures: Described for A. leucopeza (Robinson, 

1953) and A. africana (Papp, 2008). Egg: Oval in dorsal and lateral views, length 0.35–0.59 mm, width 0.15–

0.22 mm. Flattened, hatching strip demarcated by a flanged 

edge. Chorion covered with honeycomb reticulations and 

longitudinal ribs. Third-instar larva: Subcylindrical, amphipneustic, bearing a long, partially retractile terminal 

respiratory tube; integument white, almost transparent, 

covered with minute spicules and sensory setae, or smooth; 

cephaloskeleton slender with well-developed mandible 

bearing 2 pairs of accessory teeth; hypopharyngeal and 

tentoropharyngeal sclerites separate; 3 thoracic and 8 abdominal segments well produced; abdominal segments with 

ventral creeping pads bearing minute spicules; last abdominal segment with a small perianal pad; anterior spiracles 

small, retracted into a pair of deep, narrow pockets; posterior spiracles on apex of respiratory tube, joined at their 

base, bifurcating posteriorly. Puparium: Light brown, 

oval in shape, with prothoracic spiracle completely everted.

Distribution. Worldwide, excluding Australasian and Oceanic Regions. The Neotropical fauna is 

considerably more diverse than other regional faunas and 

herein includes 41 species, of which 37 are described in 

this paper. Of the seven species groups that we recognize 

in the genus, six are exclusively found in Neotropics, and 
6 • smithsonian contributions to zoology

based on the sampling of specimens now available, most 

Neotropical species are apparently endemic or of restricted 

distribution. The genus has not been well sampled in the 

Neotropics, however, and many more species are likely to 

be discovered there.

Biology. Information in the literature on the 

natural history of the genus Aulacigaster pertains to species 

of the leucopeza group only. Adults and immatures are peculiar to weeping wounds and sap fluxes of deciduous trees 

(Robinson, 1953; Teskey, 1976; Ferrar, 1987). A group of 

species (here named “the bromeliae group”) inhabits bromeliads in the Neotropical Region. Larvae of these species are 

fully aquatic and probably feed on decaying organic matter.

In addition to weeping wounds, some adults of Aulacigaster were collected on freshly cut logs. We also collected 

numerous adults, sometimes of more than one species, by 

sweeping between the shaded buttresses of ceba trees in 

the Neotropics. A few species have been collected on Heliconiaceae leaves.

Implicit Attributes

Unless indicated otherwise, the following attributes 

are implicit throughout the descriptions, except where the 

characters concerned are inapplicable.

Adult. Coloration: Occiput mostly subshiny 

to dull; anterior portion of frons without a microtomentose band; facial band absent. Scutum uniformly microtomentose; thoracic pleura uniformly black. Wing hyaline. 

Abdomen completely dark brown to black; pregenital segment mostly polished.

Morphology: Body robust, scutal ratio 1.1–1.35. 

Head: Vertex nearly straight between compound 

eyes; medial portion of frons not depressed; margins of eye 

closer together at ventral portion of frons than at vertex; 

1st flagellomere round; arista long, distinctly zigzagged on 

apical 2/3 and bearing short hairs, basal 1/3 of arista naked; 

face completely sclerotized; gena in lateral view a narrow 

ridge circumventing ventral margin of eye; clypeus stout, 

1.5–2.5 times longer than wide; palpus short, length about 

half that of 1st flagellomere. Cephalic setae not elongated; 

ocellar seta present; medial vertical setae present, convergent; fronto-orbital setae transversely aligned; peristomal 

vestiture consisting mostly of one strong and a few weak 

setae following pseudovibrissal seta. 

Thorax: Thoracic setae not elongated; costa of 

male not bearing relatively strong, conspicuous spines. 

Legs: Hind trochanter of male without a ventral 

tuft of setae; hind femur of male and female unmodified, 

without ventral spines; hind femur of male without a conspicuous lobe and ventral spines; hind tibia of male and 

female with a rounded, posteroventral projection, without 

apicodorsal setae.

Male abdomen and terminalia: Abdominal sternites slightly increasing in width posteriorly; sternites 3–5 

unmodified, with posterior margin straight; cerci separate; 

halves of hypandrium connected by a narrow bridge above 

phallapodeme.

Key to the Neotropical Species Groups and Species of Aulacigaster Macquart

1. Body robust, scutal ratio 1.1–1.35 [Figures 9, 64, 100, 161] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

Body elongate, scutal ratio 1.5–1.75 [Figures 40, 107] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32

2. Arista relatively long, bearing short to long rays on apical 2/3 [Figures 4, 204, 206–207, 209–214] . . . . . . . . . . . . . . . 3

Arista relatively short, appearing naked under low magnification [Figures 186, 216]; frons orange on medial portion, 

infuscate with brown laterally (Neotropical species of the leucopeza group) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .4

3. Arista distinctly zigzagged, bearing short rays on apical 2/3, naked on basal 1/3; 1st flagellomere round, not decumbent 

[Figures 206–207, 209–214]; clypeus stout, 1.5–2.5 times longer than wide [Figure 33] . . . . . . . . . . . . . . . . . . . . . . . . 6

Arista bearing alternate rays on apical portion, appearing zigzagged; basal 1/3 of arista straight, with a double row of 

dorsal rays; rays of arista long [Figure 4]; 1st flagellomere elongate, decumbent [Figures 4, 11, 21, 203–204]; clypeus 

elongate, 3.5–5.5 times longer than wide [Figure 21] (the bromeliae group) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27

4. Frons bearing a horizontal, unusually large orange band extended beyond base of anterior ocellus medially [Figure 185]; 

face in lateral view dorsoventrally projected, ventral portion of face not receded; facial band unusually wide, ~1/3 length 

of 1st flagellomere [Figure 186] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. L2, undescribed species

Frons at most with a narrow, orange-brown transverse band [as in Figures 215, 217, 219]; ventral portion of face receded 

[Figure 216] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

5. Frons yellowish-orange on central portion, merging with black along lateral margins [Figure 215]; facial band absent 

[Figures 215–216] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. spangleri, new species

Frons with a transverse, narrow, orange-brown band; facial band present . . . . . . . . . . . . . . . A. L1, undescribed species
number 633 • 7

6. Frons mostly brown, at most with anterior margin pale, yellowish [Figures 207, 209]; anepisternum mostly polished . . . . . 7

Frons with anterior 1/3–1/2 yellowish in ground color (yellowish portion of frons invested with dense microtomentum, 

silver under certain angles) [Figure 213]; anepisternum mostly dull (the plesiomorphica group) . . . . . . . . . . . . . . . . . .18

7. Abdomen with syntergite 1+2 whitish [Figure 101]; mesofrons with a delicate, transverse veil of microtomentum anterior 

to lower fronto-orbital setae, but not silver [Figure 209]; hind femur and tibia of male and female not modified, without 

conspicuous ventral spines (the grimaldii group) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

Abdomen with a whitish to yellowish region on syntergite 1+2, sometimes extending to tergite 3 [Figure 62]; hind femur 

and tibia of male and female modified, bearing tiny, conspicuous ventral spines [Figure 71]; hind femur of male often 

strong, bearing a basal lobe [Figures 66, 70–71, 74, 77, 81–82, 86, 91, 96] (the femorata group). . . . . . . . . . . . . . . . . 9

8. Face and legs mostly yellow [Figures 100, 102] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. grimaldii, new species

Face and legs mostly brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. angusta, new species

9. Fore tarsus mostly brown, only basitarsomere yellow; scutellum often very weakly raised [Figure 64]. . . . . . . . . . . . . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. bella, new species

Fore tarsus mostly yellow (at most with apical and subapical tarsomere brown); scutellum moderately raised (angle with 

scutum less than 135°) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10

10. Wing clear, at most with apex infuscate [Figures 80, 85, 93] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11

Posterior half of wing infuscate with brown. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. unifasciata, new species

11. Hind femur completely yellow [Figure 86] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12

Hind femur at least partially brown [Figures 91–92] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15

12. Hind tibia yellowish, at most very faintly brown infuscate on basal third . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13

Hind tibia brown to black on central portion, base and apex yellowish . . . . . . . . . . . . . . . . . . . . A. lobata, new species

13. Lobe on hind femur of male subrectangular, followed by a concavity . . . . . . . . . . . . . . . . . . . . . . A. erwini, new species

Lobe on hind femur of male cylindrical or triangular, not followed by a concavity [Figures 74, 81–82] . . . . . . . . . . . 14

14. Postpronotum yellow to brownish; lobe on hind femur of male approximately cylindrical [Figures 81–82]; less densely 

microtomentose flies, usually without large microtomentose regions on mesofrons and mesonotum . . . . . . . . . . . . . . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. irwini, new species

Postpronotum brown, concolorous with mesonotum; lobe on male hind femur approximately triangular [Figure 74]; 

more densely microtomentose flies, usually with a large medial microtomentose region on the mesofrons and microtomentum over mesonotum reaching anterior dorsocentral seta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. formosa, new species

15. Hind femur with dark mark on apical 1/3–1/2 [Figure 92]; fore femur yellowish . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16

Hind femur dark brown on apical 3/5 [Figure 91]; fore femur yellow basally, apex darened . . . . A. nigritibia, new species

16. Hind femur of male without a conspicuous lobe [Figure 92]; hind tibia yellowish . . . . . . . . . . . A. peruana, new species

Hind femur of male with a conspicuous lobe [Figures 70–71, 77]; hind tibia yellowish, often with brown to black mark 

on central portion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17

17. Hind femur of male with a basoventral lobe followed by a distinct concavity; lobe approximately rectangular [Figures 70, 

71]; postpronotum mostly polished . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. femorata, new species

Hind femur of male with a basoventral lobe followed by a slight concavity; lobe approximately triangular [Figure 77]; 

postpronotum microtomentose over ventral and anterior margins, otherwise polished . . . . . . . . A. guyana, new species

18. Ocellar tubercle mostly polished. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. trifasciata, new species

Ocellar tubercle dull microtomentose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19

19. Hind tibia without apical ventral seta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .20

Hind tibia with strong, dark, apical ventral seta [Figure 152] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25

20. Facial band absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21

Facial band present (sometimes very inconspicuous, delimited as a transversely wrinkled band mostly lacking microtomentum) [Figure 213] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22

21. Frons white-microtomentose stripe not bordered posteriorly by a conspicuous velvety-black stripe . . . A. erika, new species

Frons white-microtomentose stripe bordered posteriorly by a conspicuous velvety-black stripe [as in Figures 135, 213] . . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. bifasciata, new species

22. Facial band delimited as a transversely wrinkled band on dorsal portion of face, but mostly lacking microtomentum; cerci 

of male and female not fused . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. belize, new species
8 • smithsonian contributions to zoology

Facial band more distinct, gray or golden microtomentum over brown mark, on dorsal portion of face [as in Figure 142]; 

cerci of male partially fused [Figure 144] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23

23. Hind femur yellow; surstylus approximately triangular, apex strongly pointed, surface concave [Figure 159] . . . . . . . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. rufifemur, new species

Hind femur at least partially dark; surstylus not as above [Figures 132, 136, 143, 146] . . . . . . . . . . . . . . . . . . . . . . . 24

24. Vertex mostly polished, microtomentum restricted to a narrow area posterior to ocellar tubercle and, in some specimens, 

around vertical seta; ventral portion of face not bulbous, difficult to see in lateral view; surstylus approximately rectangular, in lateral view approximately “boat-shaped” [Figure 132]. . . . . . . . . . . . . . . . . . . . . . . . . . A. albifacies, new species

Vertex mostly dull microtomentose, shiny spot, when present, reaching at most halfway to eye margin; ventral portion of face 

bulbous, easily seen in lateral view [Figure 141]; surstylus keel-shaped, with one subapical seta [Figures 143, 146] . . . . . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. conspicua, new species

25. Hind trochanter of male without a ventral tuft of setae; surstylus folded ventrally to epandrium, difficult to discern [Figure 

136] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. atriseta, new species

Hind trochanter of male with a ventral tuft of setae [Figures 148, 149]; surstylus conspicuous in lateral view [Figures 154, 

155] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26

26. Hind trochanter of male with a ventral, yellowish tuft of setae not born on a projection [Figure 149]; surstylus with apex 

approximately foot-shaped, with 2 subterminal setae and 2 basal, marginal setae [Figure 155] . . . . A. proxima, new species

Hind trochanter of male with a ventral, black tuft of setae born on a projection [Figure 148]; surstylus approximately 

triangular, with a terminal seta and 2 basal, marginal setae [Figure 154] . . . . . . . . . . . . . . . A. plesiomorphica (Hennig)

27. Medial portion of frons slightly depressed from ptilinal suture to base of ocellus [Figure 19] . . . . . . . . . . . . . . . . . . . 28

Medial portion of frons deeply depressed, often velvety black [Figures 14, 22, 203] . . . . . . . . . . . . . . . . . . . . . . . . . . . 30

28. Ocellar seta absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. serrana, new species

Ocellar seta present, though minute . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29

29. Antenna infuscate on dorsal margin; medial vertical seta present [Figure 19]. . . . . . . . . . . . . . . . A. tibanae, new species

Antenna infuscate on dorsal half; medial vertical seta absent [as in Figure 22] . . . . . . . . . . . . . . . . A. lopesi, new species

30. Fronto-orbital plate completely brown to dark brown [Figure 22]; ocellar tubercle mostly polished; katepisternum mostly 

polished; medial vertical seta absent [Figure 22] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31

Fronto-orbital plate yellowish in ground color; medial vertical seta present [Figure 203]; ocellar tubercle microtomentose, 

subshiny; katepisternum mostly microtomentose (some specimens with a polished area dorsally) . . . . . . . . . . . . . . . . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. bromeliae, new species

31. Vertex mostly or entirely polished; costa of male not bearing conspicuous spines [Figure 13] . . . . . A. korneyevi, new species

Vertex with small polished spot extended from ocellus to at most 1/3 distance to eye margin; costa of male bearing a few 

tiny, but conspicuous spines [Figure 24] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. vespertina, new species

32. Thoracic pleura uniformly black (the minuta group) [Figure 107] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33

Thoracic pleura white to pale yellow on ventral half; legs white to pale yellow [Figures 40, 49] (the ecuadoriensis group)

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39

33. Wing hyaline, at most with apex very slightly infuscate [as in Figure 116]; lobe on hind femur of male, when present, not 

appearing “sac-like” [Figures 108, 128, 131] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34

Wing infuscate with brown, at least on apical 2/3 of cells r2+3 and r4+5 [Figure 114], lobe on hind femur of male appearing 

“sac-like,” with a few marginal setae on apex [Figures 112–113, 122, 124]. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37

34. Antenna not infuscate dorsally; hind tarsus with basal tarsomere brown, apical tarsomere yellowish . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. nigritarsa, new species

Antenna infuscate along dorsal margin; hind tarsus yellowish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35

35. Hind femur mostly dark brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. minuta (Hennig)

Hind femur mostly yellowish, apex brown infuscate, or yellowish-brownish, but never dark brown [Figure 108] . . . . . . 36

36. Hind femur of male with a small, inconspicuous lobe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. costaricana, new species

Hind femur of male with a basoventral lobe followed by a distinct concavity [Figure 108] . . . . . . . . . . . . . . . . . . . . . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. appendiculata, new species

37. Hind femur almost completely brown, at most with a narrow yellow area basally [Figure 121]; scutellum slightly raised, 

completely microtomentose, or dull microtomentose on posterior portion, anterior portion shiny; lobe on hind femur of 
number 633 • 9

The bromeliae Group

Figures 1–25, 203–204, Maps 1–2

Diagnosis. The bromeliae group is distinguished from other species groups by the following combination of characters: Stout, medium-sized flies, body length 

2.6–4.0 mm. Coloration and vestiture: Occiput typically 

with medial occipital sclerite polished; mesofrons mostly 

dark brown; antenna often invested with whitish, pubescent setulae or macropubescence; face polished, brown to 

black. Anepisternum often polished. Wing often bearing a 

central brown mark.

Morphology: 

Head: Typically higher than long (head ratio less 

than 0.9); vertex obviously excavated between compound 

eyes; medial portion of frons slightly to broadly depressed, 

deep. Margins of eye closer together at vertex than at level 

of antenna; 1st flagellomere oval, decumbent; arista long, 

with a double row of dorsal rays on basal 1/3, and alternate rays on apical 2/3; face in lateral view projected dorsoventrally, ventral portion of face not receded; gena in 

lateral view projected at level of anteroventral margin of 

eye, approximately triangular in shape, thereafter following 

ventral margin of eye as a narrow ridge; clypeus elongate, 

3.5–5.5 times longer than wide; ocellar seta often present, 

minute; fronto-orbital setae with posterior seta slightly internal to and almost transversely aligned with anterior seta; 

medial vertical seta, when present, nearly cruciate; peristomal area typically with 1 strong seta immediately behind 

pseudovibrissal seta, followed by 2–3 weaker setae and 

other setulae.

Thorax: Costal vein of male often with a few minute (sometimes inconspicuous) spines on the apical 1/3; 

subcosta partially fused with vein R1

 apically, terminating 

at costal vein.

Male abdomen and terminalia: Surstylus slender, 

digitiform, posteroventral or posterior; subepandrial sclerite consisting of paired, ribbon-like sclerites separate by a 

membranous, central region. Gonopods short, wide, with 

3–4 central setulae and a posterior lobe or process. Cercus 

typically bearing a few axial and apical strongly developed 

setae.

Distribution. Neotropical: Brazil, Costa Rica, 

Ecuador.

Biology. Species of this group are rare in collections, although specimens are conspicuous and relatively easy to collect on leaves of bromeliads that have large 

basal “cups,” where water accumulates. In Brazil, adults 

are apparently present throughout the year, although in 

male either not followed by a concavity [Figure 124], or followed by a strong concavity [Figures 121–122] . . . . . . . . 38

Hind femur mostly brown, yellow on basal 1/3–1/2; scutellum very weakly raised (making almost no angle with the scutum), dull microtomentose on lateral margins and apex, disk bright shiny; lobe on hind femur of male followed by a weak 

concavity [Figures 112–113] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. condylura, new species

38. Lobe on hind femur of the male ovoid, with apical setae only, and followed by a strong concavity [Figures 121–122] . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. curvata, new species

Lobe on hind femur of the male approximately foot-shaped, with apical and posterior marginal setae, not followed by a 

concavity [Figure 124] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. gaimarii, new species

39. First flagellomere round [Figures 26, 59] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40

First flagellomere triangular [Figures 32, 50, 206] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41

40. Face mostly white [as in Figure 33]; vertex completely dark-brown; wing mostly brown, infuscate; hind femur of male 

with lobe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. stenoptera, new species

Face white dorsally, brown on ventral 1/3–1/2 [Figure 27]; vertex with yellow mark interrupted behind ocellar tubercle 

[Figure 28]; wing mostly brown infuscate, with a transverse hyaline spot not reaching cell r2+3 [Figure 29]; hind femur of 

male without lobe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. aenigma, new species

41. Face brown on ventral 1/4–1/3 [Figure 42]; hind femur of male without lobe [Figure 43] . . . . . A. fastidiosa, new species

Face mostly white or grey, if brown infuscation present, only laterally on ventral 1/4 [Figure 33]; hind femur of male 

without [as in Figure 43] or with [Figure 36] lobe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42

42. Wing with apical 2/3 of cells r1

 and r2+3 brown infuscate [Figure 52]; hind femur of male without a conspicuous lobe [as 

in Figure 43]; female abdomen with sternites uniformly sclerotized and membranous region between tergites and sternites 

dark-brown [Figure 55] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. melanoleuca (Hennig)

Wing with apical half mostly brown; hind femur of male with a basal lobe [Figure 36]; female abdomen with sternites less 

sclerotized medially and membranous region between tergites and sternites pale [Figure 35] . . . A. ecuadoriensis (Hennig)
1 0 • smithsonian contributions to zoology

fewer numbers during the winter (Jun–Aug). Larvae are 

fully aquatic and can be easily found near the bases of the 

bromeliad leaflets, where they probably feed on decaying 

organic matter.

Discussion. Species of the bromeliae group 

can be easily identified by external characters, such as coloration and the shape of the concavity of the medial portion 

of the frons, the microtomentum coverage of the vertex, 

and the presence/absence of a medial vertical seta and ocellar seta. Additionally, characters of the male terminalia are 

very useful to distinguish the males.

The bromeliae group, which is exclusively Neotropical, includes six new species that are described here. Most 

species are from South America, but one, A. korneyevi,

occurs in Central America (Costa Rica).

Aulacigaster bromeliae, new species

Figures 2–8, 203–204, Map 1

Diagnosis. This species is distinguished from 

congeners by the following combination of characters: 

Fronto-orbital plate yellowish in ground color; medial portion of frons broadly depressed, concave.

Description. Coloration and vestiture: Vertex 

mostly densely microtomentose; ocellar tubercle scarcely 

microtomentose; frons with fronto-orbital plate yellowish in ground color, depressed portion of frons appearing 

velvety; mesofrons with a pair of silver microtomentose 

marks; antenna pale yellow to yellowish, infuscate along 

dorsal margin or infuscate on dorsal half; palpus brownish. 

Scutellum dull microtomentose; postpronotum concolorous 

with mesonotum, mostly polished; katepisternum mostly 

microtomentose, with a polished area dorsally; halter 

mostly white. Wing with a central, brown mark. Fore coxa 

yellow, fore femur yellowish, fore tibia brown, fore tarsus 

mostly brown to dark brown, yellowish on basal 1/3–1/2 

of basitarsomere or completely brown to dark brown; mid 

coxa brown to black, mid femur brown, mid tibia yellowish 

or with basal portion brown to dark brown, apical portion yellowish, mid tarsus yellowish, apical and subapical 

tarsomere brownish to dark brown (apical and subapical); 

hind coxa yellow, hind femur brown, hind tibia brown to 

dark brown, hind tarsus yellowish, apical tarsomere brown 

to black. Male abdomen subshiny to dull; female abdomen 

bright shiny, or with dense microtomentum medially, lateral margins of tergites nearly polished.

Morphology: 

Head: Figures 4, 203–204. Medial portion of 

frons depressed, deep; ocellar seta hair-like, minute; 

medial vertical seta weak, less than half length of lateral 

vertical seta.

Thorax: Figures 2–3. Acrostichal setae in 1 row; 

scutellum slightly raised (angle with scutum approximately 135°), approximately triangular, apex relatively 

pointed. Costa of male bearing relatively strong, conspicuous spines.

Male abdomen and terminalia: Figure 5. Surstylus 

about 1/3–1/2 length of basal margin of epandrium, posterior to posteroventral, digitiform; anterior process of gonopod about 2/3 length of basal margin of epandrium, well 

developed; posterior process of gonopod weakly differentiated, lobelike; cercus narrow, digitiform, bearing several 

long and well-developed setae.

Female abdomen and terminalia: Figure 6. Spermathecae with ventral digitiform projections.

Measurements: Body length 2.6–3.4 mm. Wing 

length 2.4–3.0 mm, wing width 0.9–1.0 mm.

Immatures: Figures 7–8. Egg whitish, oval in 

shape, dorsoventrally flattened; chorion delicately sculptured. Larva (3rd instar): Length 6.6–7.5 mm. Slender, subcylindrical, elongate, slightly tapering anteriorly; 

integument almost transparent, smooth; abdomen with 

2 pairs of lateral sensory hairs on segments 1–7; ventral 

creeping pads bearing spines arranged in rows as follows: a pair of rows on segments 1 and 8, 2 pairs on 

remaining segments; 8th segment bearing a respiratory 

process and also a pair of lateral fleshy lobes; anterior 

spiracle retractable into a pair of narrow pockets, and 

bearing 23–26 pairs of twig-like processes, plus additional bulbs on their apical part. Respiratory tube with 

posterior spiracle separate apically, bearing 3 long bristlelike interspiracular processes each. Cephaloskeleton (no 

figure available) well developed, large; tentopharyngeal 

sclerite fully sclerotized, with prominent ventral groove 

and a weak dorsal bridge. Mandible well developed, with 

2 pairs of accessory teeth. Puparium: Length 5.6–6.4 

mm; width 2.0–3.3 mm. Light brown, oval in shape, tapered towards apex. Posterior respiratory tube strongly 

retracted. Prothoracic spiracles extruded into a pair of 

1.1–1.2 mm long horns each with approximately 23–26 

pairs of twig-like papillae.

Type Material. The holotype male is labeled “BRAZIL. Rio de Janeiro: Ilha da Marambaia 

23°03.6¢S, 43°59.1¢W, 4 Sep 2000, D. & Wayne N. 

Mathis/HOLOTYPE ♂ Aulacigaster bromeliae Rung & 

Mathis MZUSP [red].” The holotype is double mounted 

(minuten in a block of plastic), is in excellent condition, 

and will be deposited in the MZUSP. Paratypes are as follows: same label data as the holotype (1♂, 1♀; MZUSP; 
number 633 • 1 1

FIGURES 2–4. Digital photographs of Aulacigaster 

bromeliae, new species (male) (the bromeliae group, 

Neotropical Region): (2) right wing, dorsal aspect; 

(3) enlargement of costal section, dorsal aspect; 

(4) left antenna, medial aspect. Not all to the same 

scale.

FIGURES 5–8. Illustrations of Aulacigaster bromeliae, new species (the bromeliae group, Neotropical 

Region): (5) epandrium and internal genitalic structures (male), lateral aspect; (6) spermathecae, ventral 

aspect; (7) egg, dorsal aspect; (8) first–instar larva, lateral aspect. Not all to the same scale.
1 2 • smithsonian contributions to zoology

3♂, 2♀; USNM). BRAZIL. Rio de Janerio: Rio de Janerio: 

Maricá (22°55.1¢S, 42°29.1¢W), 27 Feb 1995, A. Baptista, 

L. Lopez (3♂; MNRJ) (3♂; USNM).

Type Locality. Brazil. Rio de Janeiro: Ilha da 

Marambaia (23°03.6¢S, 43°59.1¢W).

Other Specimens Examined. BRAZIL. 

Rio de Janeiro: Guaratiba (22°58¢S, 42°48¢W), 1990, 

A. and R. Baptista 1♂, 3 ♀; MNRJ); Petrópolis, Retiro 

(22°30.6¢S, 43°11.1¢W), 10 May 1984, H. S. Lopes (2♀; 

MNRJ); Teresópolis, Parque Nacional da Serra dos Órgãos (22°27.9¢S, 42°59.6¢W), 28 Dec–Jul 1990, 1996, A. 

and R. Baptista (3♂, 1♀; MNRJ, USNM); Rio de Janeiro 

(22°54¢S, 43°14¢W; em broméia), 1 Jan–1 Sep 1934, H. S. 

Lopes (1♂, 2♀; MNRJ); 20 Oct 1991, A. and R. Baptista (3♀; MZUSP); Gávea (22°59¢S, 43°14.7¢W), 3 Mar 

1935, H. S. Lopes (1♂; MNRJ); Jardim Botânico do Rio 

de Janeiro (22°58¢S, 43°13.5¢W), 27 Sep–20 Dec 1990, 

1991, R. Tibana (4♂, 4♀; MZUSP); Recreio dos Bandeirantes (23°01.1¢S, 43°28.1¢W), 23–30 Sep 1974, R. Tibana (2♂, 2♀; MNRJ). Santa Catarina: Ilhota, Morro do 

Baú (26°54¢S, 48°49.6¢W; em bromélia), 26–27 Jan 1990, 

A. Chaves, R. Baptista (1♂, 2♀; MNRJ). São Paulo: Salesópolis (23°33.9¢S, 45°51.8¢W; em bromélia), 1–3 Mar 1992, 

A. and R. Baptista (1♂, 2♀; MZUSP).

Immature forms. Puparium. BRAZIL. Rio 

de Janeiro: Rio de Janeiro, Recreio dos Bandeirantes 

(23°01.1¢S, 43°28.1¢W), Sep 1974, R. Tibana (5, MNRJ); 

29 Aug 1978, R. Tibana (1, MNRJ). Larva. BRAZIL. 

Rio de Janerio. Rio de Janeiro, Recreio dos Bandeirantes, 

Sep 1974, R. Tibana (7, MNRJ).

Distribution. (Map 1) Neotropical: Brazil 

(Rio de Janeiro, Santa Catarina, São Paulo).

We have collected immatures and adults of this species on the following species of bromeliads: Neoregellia 

cruenta (R. Graham) L. B. Smith , Vriesea spp., and Canistrum lindenii (Regel) Mez. This species occurs in bromeliads near the maritime coast of the Atlantic Forest.

Etymology. The specific epithet, bromeliae,

refers to a plant genus of the family Bromeliaceae, in general recognition of the larval habitat of this species.

Remarks. Aulacigaster bromeliae is similar to 

A. vespertina and A. korneyevi in the shape of the medial 

portion of the frons, which is deeply depressed and has a 

pair of silvery microtomentose marks. Besides the characters given in the key and diagnosis, this species can be 

readily distinguished from the two species mentioned above 

by having more extensive microtomentose coverage on the 

ocellar tubercle and katepisternum, by the shape of the surstylus (digitiform with apex not swollen), and by the weakly 

differentiated posterior process of the male gonopod.

Aulacigaster korneyevi, new species

Figures 1, 9–15, Map 2

Diagnosis. This species is distinguished from 

congeners by the following combination of characters: Medial portion of frons broadly depressed, concave; vertex 

mostly polished.

Description. Coloration and vestiture: Vertex 

mostly polished; ocellar tubercle mostly polished; fronto-orbital plate and mesofrons completely brown to dark brown, 

depressed portion of frons appearing velvety; mesofrons 

with a pair of silver microtomentose marks; antenna pale 

yellow to yellowish, not infuscate dorsally or infuscate on 

dorsal half; palpus brownish. Scutellum dull microtomentose; postpronotum concolorous with mesonotum, mostly 

polished; katepisternum mostly polished; halter mostly 

white. Wing hyaline or bearing a central, brownish faint 

mark (more pronounced in female specimens). Fore coxa 

yellow, fore femur yellowish, fore tibia brown, fore tarsus 

completely brown to dark brown; mid coxa brown to black, 

mid femur brown, mid tibia yellowish, mid tarsus mostly or 

entirely yellowish, apical and subapical tarsomere brownish 

to dark brown; hind coxa yellow, hind femur brown, hind 

tibia brown to dark brown, hind tarsus mostly or entirely 

yellowish, apical tarsomere brown to black. Male abdomen 

subshiny to dull. Female abdomen with dense microtomentum over tergites 1–4/5, remaining tergites mostly polished.

Morphology: 

Head: Figures 11–12, 14. Medial portion of frons 

depressed, deep; ocellar seta hair-like, minute; medial vertical seta absent.

Thorax: Figures 9–10, 12–13. Acrostichal setae in 1 

row; scutellum strongly angled dorsally (angle with scutum 

less than 135°), triangular, apex strongly pointed. Costa of 

male not bearing relatively strong, conspicuous spines.

Male abdomen and terminalia: Figure 15. Surstylar 

length subequal to the length of basal margin of epandrium; 

surstylus posteroventral to posterior, slender, with apex 

swollen, and apical dorsal triangular projection; anterior 

process of gonopod about same length of basal margin of 

epandrium, well developed; posterior process of gonopod 

well differentiated, about as long as surstylus; cercus narrow, digitiform, bearing long and well developed setae.

Measurements: Body length 2.9–3.3 mm. Wing 

length 2.9–3.0 mm, wing width 0.8–1.1 mm.

Type Material. The hololtype male is labeled 

“COSTA RICA. Heredia: S[an]t[o].Domingo[,] INBIO, 

9°58.4¢N, 85°5.6¢W, 21 February 2003, W. N. Mathis/

USNM ENT 00197130 [plastic bar code label]/HOLOTYPE 
number 633 • 1 3

♂ Aulacigaster korneyevi Rung & Mathis INBio [red].” The 

holotype is double mounted (minuten in a block of plastic), 

is in excellent condition and is deposited in INBio. Paratypes are as follows: Same label data as the holotype (9♂, 

4♀; INBio, USNM). Same locality as the holotype but collected on 14 Jun 2003, D. and W. N. Mathis (5♂, 4♀; INBio, 

USMN); 29–31 Jul 2001, V. Korneyev (6♂, 6♀; INBio, 

USNM); 3 Oct 1999, M. Buck, S. Marshall (12♂, 5♀; UGE).

Other Specimens Examined. COSTA 

RICA. Heredia: Chilamate, 18–23 Aug 1988, W. J. Hanson (1♀, USU); Santo Domingo, Inbio Park (9°58.4¢S, 

84°05.5¢W; on bromeliad leaves), 19–20 Aug 2001, 

M. Buck (10♂, 10♀; UGE); Santo Domingo, Inbio Park 

(9°58.4¢S, 84°5.6¢W; on bromeliad leaves), 10 Jun–18 Jul 

2000, 2001, K. N. Barber, E. Fisher, (19♂; CSCA, UGE).

Type Locality. Costa Rica. Heredia: Santo 

Domingo (INBio Parque; 09°58.4¢N, 85°05.6¢W).

Distribution. (Map 2) Neotropical: Costa 

Rica (Heredia).

Etymology. The specific epithet, korneyevi, is 

a genitive patronym to honor and recognize the contributions of Dr. Valery A. Korneyev to dipterology. Valery also 

collected many of the paratypes.

Remarks. The central mark of the wing, though 

faint, is more conspicuous in female specimens than in the 

male specimens.

This species is similar to A. bromeliae and A. vespertina

in the shape of the mesofrons, which is deeply depressed 

and has a pair of silver microtomentose marks. It can be 

easily distinguished from the first species by the characters 

MAP 1. Distribution of Aulacigaster bromeliae, A. lopesi, A. serrana, and A. tibanae (the bromeliae group, Neotropical Region).
1 4 • smithsonian contributions to zoology

given in the key and diagnosis. Like A. vespertina, A. korneyevi has a dark fronto-orbital plate region, lacks the 

medial vertical seta, and has a well-developed posterior 

gonopodal process and similar arrangement of the gonopodal setae. Besides the characters given in the key and diagnosis, A. korneyevi differs from A. vespertina in the shape 

of the swollen apex of the surstylus, which is triangular in 

the first species and approximately round in the second.

Aulacigaster lopesi, new species

Figure 16, Map 1

Diagnosis. This species is distinguished from 

congeners by the following combination of characters: 

Fronto-orbital plate yellowish in ground color; medial portion of frons slightly depressed over a transverse, triangular 

FIGURES 9–14. Digital photographs of Aulacigaster korneyevi, new species (male) (the bromeliae group, Neotropical Region): (9) body, 

lateral aspect; (10) body, dorsal aspect; (11) head, lateral aspect; (12) head and thorax, dorsal aspect; (13) right wing, dorsal aspect; 

(14) head, frontal aspect. Not all to the same scale.
number 633 • 1 5

area from ptilinal suture to base of ocellus; medial vertical 

seta absent.

Description. Coloration and vestiture: Vertex 

mostly polished; fronto-orbital plate yellowish in ground 

color; ocellar tubercle mostly polished; mesofrons microtomentose; antenna pale yellow to yellowish, infuscate on 

dorsal half; palpus brownish. Scutellum dull microtomentose; postpronotum concolorous with mesonotum, scarcely 

microtomentose, subshiny; katepisternum mostly polished. 

Wing with a central, brown mark. Fore coxa yellow, fore 

femur yellowish, fore tibia brown, fore tarsus completely 

brown to dark brown; mid coxa brown to black, mid femur 

yellowish, mid tibia yellowish, mid tarsus yellowish, apical and subapical tarsomere brownish to dark brown; hind 

coxa yellow, hind femur yellow, hind tibia dark basally, yellow apically, hind tarsus yellowish, apical tarsomere brown 

to black. Male abdomen subshiny to dull; female abdomen 

subshiny, or with dense microtomentum medially, lateral 

margins of tergites nearly polished.

Morphology: 

Head: Medial portion of frons slightly depressed 

over a transverse, triangular area from ptilinal suture to 

base of ocellus; ocellar seta hair-like, minute; medial vertical seta absent. 

Thorax: Acrostichal setae in 1 row; scutellum 

slightly raised (angle with scutum approximately 135°), approximately triangular, apex relatively pointed. Costa of 

male bearing relatively strong, conspicuous spines.

Male abdomen and terminalia: Figure 16. Surstylar length about 1/2 length of basal margin of epandrium, 

posteroventral, digitiform; anterior process of gonopod 

about 1/3–1/2 length of basal margin of epandrium, weak; 

posterior process of gonopod weakly differentiated; cercus narrow, digitiform, bearing several long and welldeveloped setae.

Female abdomen and terminalia: Spermathecae 

with ventral, digitiform projections.

Measurements: Body length 3.3–3.5 mm. Wing 

length 3.1–3.3 mm, wing width 1.0–1.3 mm.

Type Material. The holotype male is labeled 

“Salinas (Três Picos), Nova Friburgo[,] R[io de]J[aneiro]-

BRASIL/♂ [handwritten, pencil]/XI 1991 A. BAPTISTA 

R. BAPTISTA em bromelia [date and habitat handwritten]/

HOLOTYPE ♂ Aulacigaster lopesi Rung & Mathis MNRJ 

[red].” The holotype is double mounted (glued on left side 

to a small triangle), is in moderately good condition (head 

setae damaged, left lateral vertical seta missing, right antenna broken, scutellar setae damaged and right wing missing) and is deposited in the MNRJ. Paratypes are as follows: 

Same label data as the holotype (4♂, 5♀; MNRJ, USNM).

Type Locality. Brazil. Rio de Janeiro: Nova 

Friburgo, Três Picos (Salinas; 22°16¢S, 42°32¢W).

Distribution. (Map 1) Neotropical: Brazil 

(Rio de Janeiro).

Biology. This species was collected from bromeliads in the Atlantic forest.

Etymology. The specific epithet, lopesi, is a 

genitive patronym to honor and recognize the numerous 

contributions of Dr. Hugo de Souza Lopes to entomology 

in Brazil and specifically to the systematics of the family 

Sarcophagidae.

Remarks. This species is similar to A. serrana

and A. tibanae in the shape of the mesofrons, which has a 

small, triangular depression, and the male gonopod with 

anterior process weakly differentiated. Besides the characters given in the key and diagnosis, A. lopesi can be easily 

distinguished from these two species by the position of the 

FIGURE 15. Illustration of Aulacigaster korneyevi, new species 

(male) (the bromeliae group, Neotropical Region): epandrium and 

internal genitalic structures, lateral aspect.
1 6 • smithsonian contributions to zoology

FIGURE 16. Illustration of Aulacigaster lopesi, new species (male) (the 

bromeliae group, Neotropical Region): epandrium and internal genitalic structures, lateral aspect.

MAP 2. Distribution of Aulacigaster korneyevi and A. vespertina (the bromeliae group, Neotropical Region).
number 633 • 1 7

surstylus, which arises from the posteroventral margin of 

the epandrium.

Aulacigaster serrana, new species

Figure 17, Map 1

Diagnosis. This species is distinguished from 

congeners by the following combination of characters: Medial portion of frons slightly depressed over a transverse, 

triangular area from ptilinal suture to base of ocellus; ocellar seta absent.

Description. Coloration and vestiture: Vertex 

mostly polished; ocellar tubercle mostly polished; mesofrons completely brown to dark brown or with fronto-orbital plate yellowish in ground color and a pair of silver 

microtomentose marks; antenna pale yellow to yellowish, 

infuscate along dorsal margin; palpus brownish. Scutellum 

dull microtomentose; postpronotum concolorous with mesonotum, mostly polished; katepisternum mostly polished; 

halter mostly white (infuscate basally). Wing with a central, 

brown mark. Fore coxa brown to black, fore femur brown, 

fore tibia brown, fore tarsus completely brown to dark 

brown; mid coxa brown to black; mid femur brown; mid 

tibia with basal portion brown to dark brown, apical portion yellowish; mid tarsus yellowish, apical and subapical 

tarsomeres brownish to dark brown; hind coxa yellow; hind 

femur brown; hind tibia brown to dark brown; hind tarsus 

yellowish; apical tarsomere brown to black. Male abdomen 

bright shiny; female abdomen with dense microtomentum 

medially, lateral margins of tergites nearly polished.

Morphology: 

Head: Medial portion of frons slightly depressed 

over a transverse, triangular area from ptilinal suture to 

base of ocellus; ocellar seta absent; medial vertical seta 

absent.

Thorax: Acrostichal setae in 1 row; scutellum 

strongly raised (angle with scutum less than 135°), triangular, apex strongly pointed. Costa of male bearing relatively 

strong, conspicuous spines.

Male abdomen and terminalia: Figure 17. Surstylus about 1/2 length of basal margin of epandrium, posterior, digitiform; anterior process of gonopod about 1/3–1/2 

length of basal margin of epandrium, weak; posterior 

process of gonopod weakly differentiated; cercus narrow, 

digitiform.

Measurements: Body length 3.7–3.9 mm. Wing 

length 3.8–4.0 mm, wing width 2.8–3.0 mm.

Type Material. The holotype male is labeled 

“Brazil. R[io]J[aneiro,] Teresópolis/A. & R. Baptista[,] Jul 

1996, Bromeliads/HOLOTYPE ♂ Aulacigaster serrana

Rung & Mathis MNRJ [red].” The holotype is double 

mounted (glued to a point on left side), is in fair condition 

(eyes collapsed, left lateral vertical, dorsal fronto-orbital 

and right scutellar setae missing), and is deposited in the 

MNRJ. A paratype is as follows: Same label data as the 

holotype (1♂; MNRJ; abdomen removed and preserved in 

a vial attached to the specimen).

Type Locality. Brazil. Rio de Janeiro: Teresópolis (22°26¢S, 42°59¢W).

Distribution. (Map 1) Neotropical: Brazil 

(Rio de Janeiro).

Biology. This species was collected from bromeliads in the Atlantic forest.

Etymology. The specific epithet, serrana, is 

taken from the type locality, Parque Nacional da Serra dos 

Orgãos.

Remarks. This species is similar to A. tibanae 

and A. lopesi in the shape of the mesofrons, which has a 

small, triangular depression, and in the male gonopod, 

which has a weakly differentiated posterior process. Aulacigaster lopesi can be easily distinguished from both of these 

two species by the position of the surstylus, which arises 

FIGURE 17. Illustration of Aulacigaster serrana, new species (male) 

(the bromeliae group, Neotropical Region): epandrium and internal 

genitalic structures, lateral aspect.
1 8 • smithsonian contributions to zoology

from the posteroventral margin of the epandrium. Aulacigaster serrana shares, with A. tibanae the posterior position 

of the surstylus, but both species are distinguished by the 

characters given in the key and diagnosis.

Aulacigaster tibanae, new species

Figures 18–20, Map 1

Diagnosis. This species is distinguished from 

congeners by the following combination of characters: Medial portion of frons slightly depressed over a transverse, 

triangular area from ptilinal suture to base of ocellus; medial vertical seta present.

Description. Coloration and vestiture: Vertex mostly polished; ocellar tubercle mostly polished; 

frons with mesofrons completely brown to dark brown or 

with anterior portion of fronto-orbital plate yellowish in 

ground color; mesofrons microtomentose, or with a pair 

of silver microtomentose marks; antenna pale yellow to 

yellowish, infuscate along dorsal margin; palpus brownish (invested with silver microtomentose hairs). Scutellum 

dull microtomentose; postpronotum concolorous with 

mesonotum (yellowish anteriorly), mostly polished; katepisternum mostly polished. Wing with a central, brown 

mark. Fore coxa brown to black; fore femur brown; fore 

tibia brown; fore tarsus completely brown to dark brown; 

mid coxa brown to black; mid femur brown; mid tibia 

with basal portion brown to dark brown, apical portion 

yellowish; mid tarsus yellowish; apical and subapical tarsomere brownish to dark brown; hind coxa yellow; hind 

femur brown; hind tibia brown to dark brown; hind tarsus 

yellowish; apical tarsomere brown to black. Male abdomen subshiny to dull; female abdomen bright shiny, or 

with dense microtomentum medially, lateral margins of 

tergites nearly polished.

Morphology: 

Head: Figures 18–19. Medial portion of frons 

slightly depressed over a transverse, triangular area 

from ptilinal suture to base of ocellus; ocellar seta minute; medial vertical seta weak, less than a 1/2 of lateral 

vertical seta.

Thorax: Acrostichal setae in 1 row; scutellum 

strongly raised (angle with scutum less than 135°), triangular, apex strongly pointed. Costa of male bearing relatively 

strong, conspicuous spines.

Male abdomen and terminalia: Figure 20. Surstylus about 1/2 length of basal margin of epandrium, 

posterior, slightly sickle-shaped; anterior process of gonopod about as long as basal margin of epandrium, well 

developed; posterior process of gonopod weakly differentiated; cercus narrow, digitiform, bearing several long and 

well-developed setae.

Female abdomen and terminalia: Spermathecae 

with ventral, digitiform projections.

Measurements: Body length 3.7–4.0 mm. Wing 

length 3.7–3.9, wing width 1.0–1.3 mm.

Type Material. The holotype male is labeled “BRASIL: PR [Paraná][,] Curitiba[,] Morretes/

A.R.P. Chaves[,] R.L.C. Baptista[,] 28–31 I 990 [28–31 

Jan 1990] Em Bromelia [handwritten]/HOLOTYPE ♂

Aulacigaster tibanae Rung & Mathis MNRJ [red].” The 

FIGURES 18–20. Digital photographs (not to same scale) and 

an illustration of Aulacigaster tibanae, new species (male) (the 

bromeliae group, Neotropical Region): (18) head, lateral aspect; 

(19) head, frontal aspect; (20) epandrium and internal genitalic 

structures, lateral aspect.
number 633 • 1 9

holotype is double mounted (glued to point of a small 

triangle), is in a fair condition (setae of head, scutellar 

setae, middles and right wing missing) and is deposited 

in the MNRJ. Paratypes are as follows: Same label data 

as the holotype (3♀; MNRJ, USNM). BRAZIL. Paraná: 

Morretes (25°28.6¢S, 48°50.1¢W; near ant column), 

9 Nov 1990, S. A. Marshall (1♀; UGE); Rio Mae Catira, 

Serra do Mar (25°21.8¢S, 48°52.6¢W), 29 Aug 2000, 

D. and W. N. Mathis (1♀; USMN). Santa Catarina: 

Ilhota, Morro do Baú (26°54¢S, 48°49.6¢W; em bromélia), 26–27 Jan 1990, A. Chaves, R. Baptista (2♂, 1♀; 

MNRJ, USNM).

Type Locality. Brazil. Paraná: Morretes 

(25°28.6¢S, 48°50.1¢W).

Other Specimens Examined. BRAZIL. 

Paraná: Curitiba, Morretes (25°28.6¢S, 48°50.1¢W; em 

bromélia), 28–31 Jan 1990, A. R. P. Chaves, R. L. C. 

Baptista (2♂; MNRJ). Rio de Janerio: Angra dos Reis 

(23°10¢S, 44°15¢W), 8 Sep 1931, L. T. (1♂; MNRJ). Santa 

Catarina: Ilhota, Morro do Baú (26°54¢S, 48°49.6¢W; em 

bromélia), 26–27 Jan 1990, A. R. P. Chaves, R. L. C. Baptista (2♀; MNRJ).

Distribution. (Map 1) Neotropical: Brazil 

(Rio de Janeiro, Paraná, Santa Catarina).

Biology. This species was collected on bromeliads in the Atlantic forests of Brazil.

Etymology. The specific epithet, tibanae, is a 

genitive patronym to honor and recognize Dr. Rita Tibana, 

who collected many specimens of Aulacigaster that occur in 

bromeliads in coastal regions of the State of Rio de Janeiro 

(Brazil).

Remarks. This species is similar to A. serrana

and A. lopesi in the shape of the mesofrons, which has a 

small, triangular depression, and in the male gonopod, 

with weakly differentiated posterior process. Aulacigaster 

lopesi can be easily distinguished from these two species 

by having the surstylus arising from the posterior ventral 

margin of the epandrium. Aulacigaster tibanae shares, with 

A. serrana, the posterior position of the surstylus, but both 

species are distinguished from each other by the characters 

given in the key and diagnosis.

Aulacigaster vespertina, new species

Figures 21–25, Map 2

Diagnosis. This species is distinguished from 

congeners by the following combination of characters: Medial portion of frons depressed, concave; frons with mesofrons completely brown to dark brown; vertex with small 

polished spot extended from ocellus to at most 1/3 distance 

to eye margin. 

Description. Coloration and vestiture: Vertex with small polished spot extended from ocellus to at 

most 1/3 distance to eye margin; ocellar tubercle mostly 

polished; frons with mesofrons completely brown to dark 

brown, depressed portion of frons appearing velvety; mesofrons with a pair of silver microtomentose marks; antenna pale yellow to yellowish, not infuscate dorsally to 

infuscate on dorsal half; palpus brownish (covered by silver 

microtomentum). Scutellum dull microtomentose; postpronotum concolorous with mesonotum, mostly polished; 

katepisternum mostly polished; halter mostly white. Wing 

with a central, brown mark. Fore coxa yellow, fore femur 

yellowish (tip brown), fore tibia brown, fore tarsus completely brown to dark brown; mid coxa brown to black, 

mid femur brown, mid tibia yellowish, mid tarsus yellowish, apical and subapical tarsomere brownish to dark 

brown; hind coxa yellow, hind femur brown, hind tibia 

brown to dark brown, hind tarsus yellowish, apical tarsomere brown to black. Male abdomen subshiny to dull; 

female abdomen with dense microtomentum over tergites 

1–4/5, remaining tergites mostly polished.

Morphology: 

Head: Figures 21–22. Medial portion of frons 

depressed, deep; ocellar seta hair-like; medial vertical seta 

absent.

Thorax: Figures 23–24. Acrostichal setae in 1 

row; scutellum strongly raised (angle with scutum less than 

135°), approximately triangular, apex relatively pointed. 

Costa of male bearing relatively strong, conspicuous spines.

Male abdomen and terminalia: Figure 25. Surstylar 

length subequal to the length of basal margin of epandrium; 

surstylus posterior, slender, with apex swollen, blunt; anterior process of gonopod about the same length of basal 

margin of epandrium, well developed; posterior process of 

gonopod half the length of surstylus; cercus narrow, digitiform, bearing several long and well-developed setae.

Measurements: Body length 2.8–3.3 mm. Wing 

length 2.7–2.9, wing width 0.9–1.1 mm.

Type Material. The holotype male is labeled 

“ECUADOR. P[ue]rt[o] Orellana: RioTiputini (0°38.2¢S, 

76°8.9¢W)[,] 12–26 Aug 1999, W. N. Mathis, A. Baptista, 

M. Kotrba/HOLOTYPE ♂ Aulacigaster vespertina Rung 

& Mathis USNM [red].” The holotype is double mounted 

(minuten in a block of plastic), is in excellent condition, and 

is deposited in the USNM. Paratypes are as follows: Same 

label data as the holotype (1♂, 6♀; EPNE, USNM).

Type Locality. Ecuador. Orellana: Rio Tiputini (00°38.2¢S, 76°08.9¢W).
2 0 • smithsonian contributions to zoology

FIGURES 21–25. Digital photographs (not to same scale) and an illustration of Aulacigaster vespertina, new species (male) (the 

bromeliae group, Neotropical Region): (21) head, lateral aspect; (22) head, frontal aspect; (23) thorax, dorsal aspect; (24) right wing, 

dorsal aspect; (25) epandrium and internal genitalic structures, lateral aspect.
number 633 • 2 1

Distribution. (Map 2) Neotropical: Ecuador 

(Orellana).

Etymology. The specific epithet, vespertina, is 

of Latin derivation and means “of the evening.” This species was collected in late afternoons and early evenings in 

the rain forest of Ecuadorian Amazon.

Remarks. This species is similar to A. bromeliae

and A. korneyevi in the shape of the mesofrons, which is 

deeply depressed and has a pair of silver microtomentose 

marks. It can be easily distinguished from the A. bromeliae

by the characters given in the key and diagnosis. Like A. korneyevi, A. vespertina has a dark fronto-orbital plate, lacks 

a medial vertical seta, and has a well-developed posterior 

gonopodal process, including a similar arrangement of the 

gonopodal setae. The surstylus of A. vespertina differs from 

that of A. korneyevi in the shape of the swollen apex, being 

somewhat rounded in A. vespertina but strongly triangular 

in A. korneyevi.

The ecuadoriensis Group

Figures 26–61, 205–206, Map 3

Diagnosis. The ecuadoriensis group is distinguished from other species groups by the following combination of characters: Relatively large, dorsoventrally 

flattened flies with elongate body, body length 3.0–5.2 mm. 

Coloration and vestiture: Vertex mostly polished; ocellar 

tubercle mostly polished; frons mostly brown, anterior margin pale, yellowish. Scutum mostly devoid of microtomentum (rarely microtomentose on posterior margin, between 

dorsocentral setae); pleura and legs white to yellowish on 

ventral half. Wing often partly infuscate.

Morphology: Body elongate, scutal ratio 1.5–1.75. 

Head: Typically longer than high (head ratio higher 

than 1.1); 1st flagellomere triangular or round; face at level 

of pseudovibrissal seta narrower than width of 1stst flagellomere; face typically with dorsal 2/3 flat, ventral portion 

projected, bulbous, easily visible in lateral view; ocellar seta 

absent; medial vertical seta about 3/4 length of lateral vertical seta, sometimes slightly proclinate.

Thorax: Costal vein of the male with or without 

differentiated spines. Subcosta partially fused with vein R1

apically but terminating on costal vein. Hind femur of male 

and female modified, with 2 ventral rows of spines, each 

spine bearing an apical seta; hind femur of male with or 

without a sub-basal lobe.

Male abdomen and terminalia: Surstylus in lateral 

view a wide, posterior ventral extension of epandrium or a 

short, posteroventral lobe; cerci partially fused; subepandrial 

sclerite forming a single, plate-like structure; gonopods 

short, wide, with a few central setulae and 2 posterior processes (one process weakly developed in A. stenoptera), one 

of them with 4–7 strong, stout setae.

Distribution. Neotropical: Brazil, Costa 

Rica, Ecuador, Panama, Peru.

Biology. The biology of this species group is 

mostly unknown. Aulacigaster ecuadoriensis has been collected on leaves of Heliconiacea.

Discussion. Species of the ecuadoriensis group 

can be best identified by the shape of the 1st flagellomere, 

the coloration of the wings, the presence/absence of a lobe 

on the hind femur of the male, the coloration of the face, 

and the shape of the surstylus.

The ecuadoriensis group includes five species of which 

two were described previously. The group is exclusively 

Neotropical.

Aulacigaster aenigma, new species

Figures 26–31, Map 3

Diagnosis. This species is distinguished from 

congeners by the following combination of characters: Vertex, between medial vertical setae, with a yellow mark interrupted behind ocellar tubercle.

Description. Coloration and vestiture: Vertex, between medial vertical setae, with a yellow mark 

interrupted behind ocellar tubercle; mesofrons mostly polished; antenna pale yellow to yellowish, infuscate along 

dorsal margin; face dark brown on ventral 1/3–1/2 (some 

teneral specimens with a dark brown facial band, ventral portion of face yellow to light brown); palpus white. 

Scutum microtomentose near posterior margin, between 

dorsocentral setae; scutellum dull microtomentose; postpronotum yellowish or light brown; anepisternum subshiny; katepisternum subshiny to dull; halter mostly tan. 

Wing mostly brown infuscate, with a posterior transverse 

hyaline mark not reaching cell r2+3. Abdomen with anterior 

margin white; male abdomen subshiny to dull; epandrium 

yellow on anteroventral portion; female abdomen with 

scarce microtomentum and membranous region between 

tergites and sternites pale.

Morphology: 

Head: Figures 26–28. First flagellomere round; 

peristomal area with about 5–7 pale setae following pseudovibrissal seta.

Thorax: Figure 29. Acrostichal setae in 1 row; scutellum very weakly raised (making almost no angle with 

scutum), trapezoidal, apex nearly straight, disk of scutellum 
2 2 • smithsonian contributions to zoology

slightly convex; basal scutellar seta less than 1/2 length of 

posterior seta. Costa of male not bearing relatively strong, 

conspicuous spines. Hind femur of male without a conspicuous lobe.

Male abdomen and terminalia: Figures 30–31. Surstylus a short, posteroventral lobe in lateral view, lobe approximately finger-like, tapering apically; cercus bearing 2 

well-differentiated setae, the shorter seta less than 1/2 the 

length of the longer seta; gonopods with 2 subequal posterior processes, one bearing 5–6 strong, stout, setae.

Female abdomen: Sternite one reduced to a pair of 

lateral sclerotized plates; sclerotization on sternites 2–8 obviously weaker than on respective tergites, particularly on 

medial portion. 

Measurements: Body length 4.0–4.5 mm. Wing 

length 4.0–4.5 mm, wing width 0.95–1.0 mm.

Type Material. The holotype male is labeled 

“ECU[ADOR]: Napo, El Chaco, 4.8 km W, 1750 m, 7 Nov 

1999, S. A. Marshall, debu00108197/Holotype ♂ Aulacigaster aenigma Rung & Mathis UGE [red].” The holotype is double mounted (glued to a point), is in excellent 

condition (wing tips slightly folded), and is deposited in the 

UGE. Paratypes are as follows: Same label data as the holotype (4♂, 4♀; UGE, USNM).

Type Locality. Ecuador. Napo: El Chaco (4.8 

km W; 00°20¢S, 77°49¢W).

Distribution. (Map 3) Neotropical: Ecuador 

(Napo).

Etymology. The specific epithet, aenigma, is 

of Greek derivation and means “obscure, or a mystery,” in 

reference to the unusual features displayed by this species.

Remarks. This species is most similar to A. fastidiosa, but can be easily distinguished from it and from 

other congeners by the characters given in the key and 

diagnosis.

Aulacigaster ecuadoriensis (Hennig)

Figures 32–38, 205–206, Map 3

Schizochroa ecuadoriensis Hennig, 1969: 610, 605 [figure of head and 

wing].—Papavero, 1967: 91 [Neotropical catalog].

FIGURES 26–29. Digital photographs of Aulacigaster aenigma, new species (male) (the ecuadoriensis group, Neotropical Region): 

(26) head, lateral aspect; (27) head, ventral–frontal aspect; (28) head and anterior portion of thorax, dorsal aspect; (29) right wing, 

dorsal aspect. Not all to the same scale.
number 633 • 2 3

Aulacigaster ecuadoriensis.—D. K. McAlpine, 1983: 58 [generic combination].

Diagnosis. This species is distinguished from 

congeners by the following combination of characters: First 

flagellomere triangular; lobe on hind femur of male present.

Description. Coloration and vestiture: Mesofrons mostly polished; antenna pale yellow to yellowish, infuscate along dorsal margin or infuscate on dorsal half; face 

white to yellowish (infuscate with brown below insertion of 

pseudovibrissa in some specimens); palpus white or yellowish. Scutellum dull, microtomentose on lateral margins and 

apex, medial portion of scutellum bright shiny; postpronotum light brown or concolorous with mesonotum; anepisternum subshiny; katepisternum subshiny to dull; halter 

mostly white, or mostly brown to dark brown. Wing with 

apical half mostly brown. Abdomen with anterior margin 

white; male abdomen subshiny to dull; epandrium whitish 

to pale yellow on anteroventral portion; female abdomen 

with dense microtomentum medially; lateral margins of tergites nearly polished; membranous region between tergites 

and sternites pale.

Morphology: 

Head: Figures 32–33, 205–206. First flagellomere 

triangular; peristomal area with about 4 setae following 

pseudovibrissal seta.

Thorax: Figures 34, 36. Acrostichal setae in 1 row; 

scutellum slightly raised (angle with scutum approximately 

135°), trapezoidal, apex nearly straight, disk of scutellum 

slightly convex; basal scutellar seta less than 1/2 length of 

posterior seta. Costa of male bearing relatively strong, conspicuous spines. Hind femur of male with a basal lobe not 

followed by a posterior concavity, lobe approximately triangular, setae on lobe yellow, 10–12, none especially elongate or strong.

Male abdomen and terminalia: Figures 37–38. 

Surstylus a short, posteroventral lobe in lateral view, approximately trapezoidal in shape, with posterior lateral 

margin slightly longer than anterior margin; cercus bearing 2 well-differentiated setae, the shorter seta less than 

1/2 the length of the longer seta; gonopods with 2 posterior processes, one bearing 4–5 strong, stout, setae; anterior, setae-bearing process slightly shorter than posterior, 

naked process. 

Female abdomen: Figure 35. Sternite 1 reduced to a 

pair of lateral round, small plates; sclerotization on sternites 

2–8 obviously weaker medially.

Measurements: Body length 4.5–4.8 mm. Wing 

length 4.1–4.3 mm, wing width 0.9–1.1 mm.

Type Material. The holotype male is labeled 

“Coca, Napo R[iver]., Napo, ECUADOR[,] .V.1965 [May 

1965] 250m., L. Pena/HOLOTYPUS Schizochroa ecuadoriensis Hg. [red][on reverse side “wing on slide”]/Holotype 

Schizochroa ecuadoriensis Hennig CNC No. 9931.” The 

holotype is glued directly to a pin, lacks a right wing, but 

otherwise is in good condition, and is deposited in the CNC 

(9931). Paratypes are as follows: Same label data as the holotype (1♂, 1♀; CNC (9931)).

FIGURES 30–31. Illustrations of Aulacigaster aenigma, new 

species (male) (the ecuadoriensis group, Neotropical Region): (30) gonopod, lateral aspect; (31) epandrium, lateral 

aspect.
2 4 • smithsonian contributions to zoology

Type Locality. Ecuador. Orellana: Napo River, 

Puerto Francisco de Orellana (Coca; 00°28¢S, 76°58¢W).

Other Specimens Examined. ECUADOR. 

Los Ríos: Rio Palenque (00°29¢S, 79°22¢W), 22–26 Feb 

1976, G. and M. Wood (1♂, teneral; CNC). Pichincha: Maquipucuna Biological Research Station (00°07¢N, 78°38¢W; 

river trail; Heliconia leaves; 1200 m), 26 Apr–29 Oct 1999, 

2002, M Buck, S. A. Marshall (13♂, 6♀; UGE).

Distribution. (Map 3) Neotropical: Ecuador 

(Los Ríos, Orellana, Pichincha).

Remarks. This species is similar to A. melanoleuca in having the 1st flagellomere triangular and the face 

mostly yellow. Both species can be easily distinguished with 

the characters given in the key and diagnosis.

Aulacigaster fastidiosa, new species

Figures 39–47, Map 3

Diagnosis. This species is distinguished from 

congeners by the following combination of characters: 1st 

flagellomere triangular; costa of male not bearing relatively 

strong, conspicuous spines.

Description. Coloration and vestiture: Mesofrons mostly polished; antenna pale yellow to yellowish, 

infuscate along dorsal margin; face dark brown on ventral 

1/4 to ventral 1/3 (some teneral specimens with a dark 

brown facial band, ventral portion of face yellow to light 

brown); palpus white. Scutum and scutellum mostly devoid 

MAP 3. Distribution of the ecuadoriensis species group of Aulacigaster (Neotropical Region).
number 633 • 2 5

of microtomentum; postpronotum dark brown or concolorous with mesonotum; anepisternum subshiny; katepisternum subshiny to dull; halter white to tan. Wing mostly 

brown infuscate on apical half. Abdomen with anterior 

margin white; male abdomen subshiny to dull; epandrium 

yellow on anteroventral portion; female abdomen with 

scarce microtomentum and membranous region between 

tergites and sternites pale.

Morphology: 

Head: Figures 42, 44. First flagellomere triangular; 

peristomal area with about 5–7 setae following pseudovibrissal seta.

Thorax: Figures 39–41, 43. Acrostichal setae in 

1 row; scutellum not raised (angle with scutum approximately 0°), trapezoidal, apex nearly straight, disk of scutellum slightly convex; basal scutellar seta less than 1/2 

FIGURES 32–35. Digital photographs of Aulacigaster ecuadoriensis (Hennig) (the ecuadoriensis group, Neotropical Region): (32) head of male, lateral aspect; (33) head, frontal–ventral 

aspect; (34) left wing of male, dorsal aspect; (35) female abdomen, ventral aspect. Not all to 

the same scale.
2 6 • smithsonian contributions to zoology

length of posterior seta. Costa of male not bearing relatively 

strong, conspicuous spines. Hind femur of male without a 

conspicuous lobe.

Male abdomen and terminalia: Figures 46–47. 

Surstylus a short, posteroventral lobe in lateral view, approximately triangular in shape; cercus bearing 2 well-differentiated setae, the shorter seta less than 1/2 the length of 

the longer seta; gonopods with 2 posterior processes, one 

bearing 5–6 strong, stout, setae; anterior, setae-bearing process shorter than posterior, naked process.

Female abdomen: Figure 45. Sternite one reduced 

to a pair of lateral sclerotized plates; sclerotization on sternites 2–8 obviously weaker than on respective tergites, particularly on medial portion. 

Measurements: Body length 4.0–4.7 mm. Wing 

length 3.7–4.0 mm, wing width 1.9–2.0 mm.

Type Material. The holotype male is labeled 

“ECU[ADOR]:Napo, Jatun Sacha Res., 6 km E Misahuallí, 

450 m, 1°4¢S, 77°37¢W, on foliage, 30 Apr–8 May 2002, 

M. Buck, debu00186240/Holotype ♂ Aulacigaster fastidiosa Rung & Mathis UGE [red].” The holotype is double 

mounted (glued to a point), is in excellent condition, and is 

deposited in the UGE. Paratypes are as follows: Same label 

data as the holotype (15♂, 6♀; UGE, USNM); same locality and collector as the holotype, but date of 4–7 May 2002 

(1♂, 4♀; UGE).

Type Locality. Ecuador. Napo: Masahuallí 

(6 km E; 01°04¢S, 77°37¢W; 450 m).

Distribution. (Map 3) Neotropical: Ecuador 

(Napo).

Biology. The type series was collected “on 

foliage.”

Etymology. The specific epithet, fastidiosa, is 

of Latin derivation and has reference to the delicate appearance of this species.

Remarks. This species is most similar to A. aenigma but can be easily distinguished from it and from other 

congeners by the characters given in the key and diagnosis.

FIGURES 36–38. Illustrations of Aulacigaster ecuadoriensis (Hennig) (male) (the ecuadoriensis group, 

Neotropical Region): (36) basal portion of the left hind femur, internal aspect; (37) gonopod, lateral aspect; 

(38) epandrium, lateral aspect. Not all to the same scale.
number 633 • 2 7

Aulacigaster melanoleuca (Hennig)

Figures 48–58, Map 3

Schizochroa melanoleuca Hennig, 1956:152–154 [figure of head, wing and 

male postabdomen].—Papavero, 1967:91 [Neotropical catalog].

Aulacigaster melanoleuca.—D. K. McAlpine, 1983:58 [generic combination].

Diagnosis. This species is distinguished from 

congeners by the following combination of characters: First 

flagellomere triangular; wing with apical 2/3 of cells r1

 and 

r2+3 dark brown, remaining wing light brown; female abdomen with membranous region between tergites and sternites brown to dark brown.

Description. Coloration and vestiture: Mesofrons mostly polished; antenna pale yellow to yellowish, 

infuscate along dorsal margin or infuscate on dorsal half; 

face white to yellowish (infuscate with brown below insertion of pseudovibrissa in some specimens); palpus white, 

or yellowish. Scutellum commonly dull microtomentose 

on lateral margins and apex, medial portion of scutellum 

FIGURES 39–45. Digital photographs of Aulacigaster fastidiosa, new species (the ecuadoriensis group, Neotropical Region). (39) body of 

male, dorsal aspect; (40) body of male, lateral aspect; (41) right wing of male, dorsal aspect; (42) head of male, frontal–ventral aspect; (43) hind 

femur of male, dorsal aspect; (44) head of male, lateral aspect; (45) female abdomen, ventral aspect. Not all to the same scale.
2 8 • smithsonian contributions to zoology

bright shiny; postpronotum concolorous with mesonotum; 

anepisternum subshiny; katepisternum subshiny to dull; 

halter mostly brown to dark brown (base whitish). Wing 

with apical 2/3 of cells r1 and r2+3 dark brown, remaining 

wing light brown. Abdomen with anterior margin white; 

male abdomen subshiny to dull; epandrium whitish to pale 

yellow on anteroventral portion; female abdomen with 

dense microtomentum medially, lateral margins of tergites 

nearly polished; female abdomen with membranous region 

between tergites and sternites brown to dark brown.

Morphology: 

Head: Figures 50–51. First flagellomere triangular; 

peristomal area with about 4 setae following pseudovibrissal seta.

Thorax: Figures 48–49, 52–53. Acrostichal setae 

in 1 row; scutellum slightly raised (angle with scutum approximately 135°), trapezoidal, apex nearly straight, disk 

of scutellum flat or slightly convex; basal scutellar seta less 

than 1/2 length of posterior seta. Costa of male bearing 

relatively strong, conspicuous spines. Hind femur of male 

without a conspicuous lobe.

Male abdomen and terminalia: Figures 56–58. Surstylus a short, posteroventral lobe in lateral view, lobe with 

sub-basal constriction, round at apex, posterior-ventral 

margin slightly folded dorsally; cercus bearing 2 well-differentiated setae, the shorter seta less than 1/2 the length of the 

longer seta; gonopods with 2 subequal posterior processes, 

one bearing 4–5 strong, stout setae.

Female abdomen: Figures 54–55. Sternite 1 reduced to a pair of round lateral sclerotized plates and an 

anterior, thin stripe; sclerotization on sternites 2–8 not obviously weaker than on respective tergites.

Measurements: Body length 4.7–5.2 mm. Wing 

length 4.2–4.3 mm, wing width 2.2–2.5 mm.

Type Material. The holotype male is labeled 

“Costa Rica[,] Farm Hamburg am Reventazon[,] 8.3.28. [8 

Mar 1928][green]/Eing. 1928 No 54 [handwritten on underside of green label]/Holotypus [red] 1972 [handwritten 

on underside of “Holotypus” label]/Holotypus [black submargin on red label]/Coll. DEI Eberswalde. The holotype is 

double mounted, is in fair condition (the proboscis, left antenna, right wing, and abdomen beyond the first tergite are 

missing), and is deposited in DEI. Paratypes are as follows: 

Same label data as the holotype (4♂, 3♀, 2 ex. (abdomens 

missing); DEI).

Type Locality. Costa Rica. Limón: Hamburg 

Farm on the Río Reventazón (10°15¢N, 83°28¢W).

Other Specimens Examined. COSTA 

RICA. Heredia: La Selva Biological Station, Puerto 

Viejo (3 km S; 10°25¢N, 84°01¢W), 26 Mar–8 Aug, 

1980, 1982, 1984, 1986, 1987, 1992, H. A. Hespenheide (3♂, UCLA; 2♂, 2♀; USNM; 1♂, 5♀; CSCA); La 

Selva Biological Station, Puerto Viejo (3 km S; 10°25¢N, 

81°01¢W; under palm leaf), 17 Feb 2003, S. D. Gaimari 

(1♀; CSCA). Guanacaste: Estación Pitilla, Santa Cecilia 

(9 km S; 10°59.5¢N, 85°25.8¢W), C. Moranga, Jul 1991 

(3♂, 1♀; INBio). Puntarenas: Piedras Blancas (24 km W; 

08°47¢N, 83°30¢W; Malaise trap), Oct 1990, P. Hanson (2♀; USNM). San José: RioSParaíso (09°33.8¢N, 

84°07.4¢W; 350–400 m), 15–17 Feb 2003, W. N. Mathis 

(1♂; USNM).

PANAMA. Coclé: El Valle (08°36¢N, 80°08¢W), Jan 

1947, N. L. H. Krauss (1♂; USNM).

Distribution. (Map 3) Neotropical: Costa 

Rica (Guanacaste, Heredia, Limón, Puntarenas, San José), 

Panama (Coclé).

Remarks. See “Remarks” under A. ecuadoriensis.

Aulacigaster stenoptera, new species

Figures 59–61, Map 3

Diagnosis. This species is distinguished from 

congeners by the following combination of characters: Thoracic pleura white to pale yellow on ventral half; 1st flagellomere round; wing mostly brown, infuscate (cells r2+3 and 

anterior half of wing more heavily infuscate).

Description. Coloration and vestiture: Mesofrons mostly polished; antenna pale yellow to yellowish, 

FIGURES 46–47. Illustrations of Aulacigaster fastidiosa, new species 

(male) (the ecuadoriensis group, Neotropical Region). (46) gonopod, lateral aspect; (47) epandrium, lateral aspect. 
number 633 • 2 9

FIGURES 48–56. Digital photographs of Aulacigaster melanoleuca (Hennig) (the ecuadoriensis group, Neotropical Region). (48) body 

of male, dorsal aspect; (49) body, lateral aspect; (50) head of male, lateral aspect; (51) head of male, dorsal aspect; (52) right wing of 

male, dorsal aspect; (53) enlargement of costal section, dorsal aspect; (54) sternites 1 and 2 of female abdomen (after KOH cleaning), 

ventral aspect; (55) female abdomen, ventral aspect; (56) subepandrial sclerite of male, ventral aspect. Not all to the same scale.
3 0 • smithsonian contributions to zoology

not infuscate dorsally or infuscate along dorsal margin; 

face white to yellowish; palpus white. Scutellum dull microtomentose; postpronotum concolorous with mesonotum, 

mostly polished; anepisternum polished; katepisternum 

subshiny to dull; halter mostly brown to dark brown. Wing 

mostly brown, infuscate (cells r2+3 and anterior half of wing 

more heavily infuscate). Male abdomen subshiny to dull; 

female abdomen subshiny.

Morphology: 

Head: Figure 59. First flagellomere round; 2–4 

peristomal setae following pseudovibrissal setae present.

Thorax: Acrostichal setae in 1 row; scutellum 

slightly raised (angle with scutum approximately 135°), 

approximately triangular, apex relatively pointed, disk of 

scutellum slightly convex; basal scutellar seta less than 1/2 

length of posterior seta. Costal vein of the male without 

differentiated spines. Hind femur of male with a basal lobe 

not followed by a posterior concavity, lobe small, approximately triangular, setae on lobe 2–3; hind tibia with a row 

of tiny ventral spines.

Male abdomen and terminalia: Figures 60–61. Surstylus in lateral view a wide, ventral extension of the epandrium; cercus bearing 2 well-differentiated setae, the shorter 

seta less than 1/2 the length of the longer seta; gonopods 

short, wide, with a few central setulae and two posterior 

processes, none bearing strong seate, posterior process only 

weakly developed.

Measurements: Body length 3.0–3.3 mm. Wing 

length 3.0–3.1 mm, wing width 0.7–0.8 mm.

Type Material. The holotype male is labeled 

“PERU. Madre de Dios: Manu, Rio Manu, 250 m[,] Pakitza, 12°7¢S, 70°58¢W, 9–23 Sep 1988[,] Amnon Freidberg/

HOLOTYPE ♂ Aulacigaster stenoptera Rung & Mathis 

USNM [red].” The holotype is double mounted (minuten in 

a block of plastic), is in excellent condition, and is deposited 

in the USNM. Paratypes are as follows: Same label data as 

the holotype (1♂; USNM). PERU. Madre de Dios: Cocha 

Salvador (11°57’S, 72°07’W; 240 m), 14 Sep 1988, A. Freidberg, W. N. Mathis (11♂, 12♀; MHNJP, USNM); Pakitza, 

Aguajal (11°58.2¢S, 71°17¢W), 19 Sep 1988, A. Freidberg 

(8♂, 4♀; USNM).

Type Locality. Peru. Madre de Dios: Manu, 

Rio Manu, Pakitza (11°56.6¢S, 71°16.9¢W; 250 m).

Other Specimens Examined. BRAZIL. 

Rondônia: Ariquemes (62 Km SE; 10°34¢S, 62°22¢W), 

15–22 Mar 1991, W. J. Hanson, G. Bohart (1♀, USU).

PERU. Madre de Dios: Los Amigos Biological Station 

(12°34¢S, 70°06.1¢W), 2–4 Jun 2006, S. Paiero, J. Klymko 

FIGURES 57–58. Illustrations of Aulacigaster melanoleuca (Hennig) (male) (the ecuadoriensis group, 

Neotropical Region). (57) gonopod, lateral aspect; (58) epandrium, lateral aspect.
number 633 • 3 1

(1♂, 2♀; UGE); Rio Manu, Erika (near Salvación; 

12°53¢S, 71°12¢W; 550 m), 5–6 Sep 1988, A. Freidberg 

(1♀; USNM).

Distribution. (Map 3) Neotropical: Brazil 

(Rondônia), Peru (Madre de Dios).

Etymology. The specific epithet, stenoptera,

is of Greek derivation and is a combination of the Greek 

words for narrow (steno) and wing (ptera).

Remarks. This species is peculiar to the A. ecuadoriensis group and is distinguished by the rounded 1st 

flagellomere and the gonopods that lack a posterior process 

that bears setae. The shape of the surstylus is also unique 

for this species.

The femorata Group

Figures 62–98, 207–208, Maps 4–6

Diagnosis. The femorata group is distinguished from other species groups by the following 

combination of characters: Relatively small, delicate-looking flies, body length 2.0–3.5 mm. Coloration and vestiture: Frons mostly brown, anterior margin pale, yellowish; 

anterior portion of frons sometimes densely microtomentose on anterior 1/6, silver from certain angles. Scutum 

typically mostly devoid of microtomentum; anepisternum 

polished; katepisternum mostly polished. Wing often with 

apex slightly infuscate. Abdomen with a yellowish region 

on syntergite 1+2.

Morphology: 

Head: Head higher than long (head ratio less than 

0.9), or longer than high (head ratio higher than 1.1); margins of the eye nearly parallel; face slightly convex, barely 

visible in lateral view; ocellar seta typically hair-like; medial vertical setae often convergent and slightly proclinate.

Thorax: Subcosta incomplete, fused with vein R1

apically. Hind femur of male and female modified with 2 

ventral rows of spines, each spine bearing an apical seta; 

hind tibia with a row of tiny ventral spines; hind femur of 

male often bearing a sub-basal lobe.

FIGURES 59–61. Illustrations of Aulacigaster stenoptera, new species (male) (the ecuadoriensis

group, Neotropical Region). (59) head, lateral aspect; (60) gonopod, lateral aspect; (61) epandrium, 

lateral aspect. Not all to the same scale.
3 2 • smithsonian contributions to zoology

Male abdomen and terminalia: Surstylus in lateral 

view a wide, ventroposterior extension of epandrium, cerci 

often partially fused; subepandrial sclerite often forming a 

single, T-shaped structure; gonopods approximately rectangular, with a long, posterior, digitiform process. Cercus 

bearing 1 long, apical seta.

Distribution. Neotropical: Brazil, Bolivia, 

Costa Rica, Ecuador, Guyana, Paraguay, Peru, Trinidad 

and Tobago. (Even though Brazil and the island of Trinidad 

and Tobago are not listed as localities for any of the species 

described below, we have studied two females from Brazil 

(Rondônia: 62 Km SE of Ariquemes, 13–25 Apr 1992 and 

7–18 Nov 1995, W. J. Hanson) and one from Trinidad and 

Tobago (Simla Research Station, 2–15 Jun 1981, W. J. Hanson and Clemons) that belong in the femorata group, near 

A. femorata new species).

Discussion. Species of the femorata group are 

distinguished best by the shape of the basal lobe on the male 

hind femur and the coloration of the legs. For the most part, 

females can only be identified with certainty when males are 

present in the sample.

The femorata group includes seven species of which six are 

newly described here. The group is exclusively Neotropical.

Aulacigaster bella, new species

Figures 62–68, Map 4

Diagnosis. This species is distinguished from 

congeners by the following combination of characters: Abdomen with a yellowish region starting on syntergite 1+2, reaching tergite 4; fore tarsus mostly brown, basitarsomere yellow.

Description. Coloration and vestiture: Vertex 

mostly polished; ocellar tubercle mostly polished; mesofrons 

with a medial microtomentose stripe from base of antenna 

to level of anterior fronto-orbital seta; antenna pale yellow 

to yellowish, infuscate along dorsal margin; face pale yellow; palpus white. Scutum dull microtomentose on anteriormost portion (anterior 1/5), microtomentum faded on 

midline of scutum, between dorsocentral setae rows; scutellum mostly dull microtomentose; postpronotum concolorous with mesonotum, mostly polished; halter mostly white. 

Fore coxa yellow, fore femur yellowish, fore tibia yellowish, 

fore tarsus brown to dark brown, basal tarsomere yellow; 

mid coxa yellow, mid femur yellowish (tip slightly darkened in some specimens), mid tibia yellowish, mid tarsus 

mostly or entirely yellowish, apical and subapical tarsomere 

brownish to dark brown; hind coxa yellow, hind femur yellow on basal portion, apical portion dark (apical 1/3–1/2), 

hind tibia brown to black on central portion, base and apex 

yellowish, hind tarsus yellowish. Male abdomen subshiny to 

dull, pregenital segment polished anteriorly, dull posteriorly.

Morphology: 

Head: Figure 63. Face at level of pseudovibrissal 

seta narrower than width of 1st flagellomere; medial vertical seta 1/2 length of lateral vertical seta.

Thorax: Figures 62, 64, 66. Acrostichal setae in 

2 rows; scutellum very weakly raised (making almost no 

angle with scutum), approximately cylindrical, apex relatively pointed. Hind femur of male with a basoventral lobe 

followed by a slight concavity, lobe approximately cylindrical, bearing 1 apical and 2 subapical setae; lobe preceded by 

a few yellow setae.

Male abdomen and terminalia: Figures 65, 67–68. 

Surstylus about as wide as posterior process of gonopod; 

lateral margins of surstylar projection approximately parallel, slightly convergent toward apex; surstylus with subapical posterior projection, forming a roughly triangular apex; 

gonopod as in Figure 67.

Measurements: Body length 2.5–3.0 mm. Wing 

length 2.3–2.4, wing width 1.7–1.8 mm.

Type Material. The holotype male is labeled 

“C[osta]R[ica], Prov. Limon, Bribri, 4 km NE, 50 m, Dec 

1989–Mar 1990, P.Hanson, debu00100753/HOLOTYPE 

♂ Aulacigaster bella Rung & Mathis UGE [red].” The holotype is double mounted (glued to a point), is in good condition (wings slightly folded, some head setae missing or 

disoriented), and is deposited in the UGE. Paratypes are as 

follows: same locality, date and collector as the holotype 

(3♂, 1♀; UGE, USNM).

Type Locality. Costa Rica. Limón: Bribri (4 

km NE; 09°39¢N, 82°49¢W).

Other Specimens Examined. BOLIVIA. 

La Paz: Mapiri (5 km W Mapiri; 15°17.8¢S, 68°15.6¢W; 

750 m), 16 Mar 2001, W. N. Mathis (2♂, 2♀; USNM).

COSTA RICA. Alajuela: Higuito, San Mateo (09°57¢N, 

84°33¢W), Pablo Schild (2♂; USNM). Limón: Bribri 

(09°38¢N, 82°50¢W; Malaise trap), Jul–Sep 1990, P. Hanson (1♂, 1♀; USNM); Dec 1989–Mar 1990, P. Hanson 

(1♂; USNM).

ECUADOR. Orellana: Reserva Etnica Waorani, 

Onkonegare Camp (1 km S; 0°39.4¢S, 76°27.2¢W; 216 

m; lot #1549), 21 Jun 1996, T. L. Erwin (1♀; USNM). 

Pichincha: Santo Domingo de los Colorados (1.6 km W; 

0°15¢S, 79°09¢W), E. S. Ross, E. I. Schlinger, 23 Feb 1955 

(1♂; CAS). Zamora-Chinchipe: Cumbaratza (3°59.4¢S, 

71°51.8¢W; 700 m), 31 Mar 1965, L. E. Peña (1♂; CNC).

PARAGUAY. Cazaapá, San Rafael, Hermosa, Lopez 

Family Residence (26°19.3¢S, 55°44.9¢W; 90 m), 3–6 Dec 

2000, Z. H. Falin (1♀; UGE).
number 633 • 3 3

Distribution. (Map 4) Neotropical: Bolivia 

(La Paz), Costa Rica (Alajuela, Limón), Ecuador (Orellana, 

Pichincha, Zamora-Chinchipe), Paraguay.

Etymology. The specific epithet, bella, is of 

Latin derivation and means “beautiful.”

Remarks. The specimens listed under “Other 

Specimens Examined” have the scutellum more strongly 

raised than the type series. Aulacigaster bella can be easily 

distinguished from congeners by the characters given in the 

key and diagnosis. The approximately cylindrical lobe followed by a concavity and the shape of the surstylus are also 

characteristic of this species. 

Aulacigaster erwini, new species

Figures 69, Map 4

Diagnosis. This species is distinguished from 

congeners by the following combination of characters: 

Postpronotum yellow to yellowish-brownish; hind femur 

completely yellow, with a basoventral lobe followed by a 

distinct concavity, lobe approximately rectangular; hind 

tibia yellow with a narrow brown mark on apical 2/3.

Description. Coloration: Vertex mostly polished; ocellar tubercle mostly polished; mesofrons with 

a faint medial microtomentose stripe from base of antenna to level of ocellar tubercle; antenna pale yellow 

to yellowish, infuscate along dorsal margin; face yellow 

(pale); palpus yellowish. Scutum dull microtomentose on 

anteriormost portion; scutellum dull posteriorly, anterior 

portion shiny; postpronotum yellow to yellowish-brownish, microtomentose over ventral and posterior margins, 

otherwise polished; halter mostly white (base infuscate). 

Male abdomen subshiny to dull, pregenital segment polished posteriorly, dull anteriorly (slightly so). Fore coxa 

yellow, fore femur yellowish, fore tarsus mostly or entirely yellowish, apical and sometimes subapical tarsomeres brownish; mid coxa yellow, mid femur yellowish, 

mid tibia yellowish, mid tarsus mostly yellowish; hind 

coxa yellow, hind femur yellow, hind tibia yellowish, 

with a small brown mark near apical 2/3, hind tarsus 

yellowish.

FIGURES 62–65. Digital photographs of Aulacigaster bella, new species (male) (the femorata group, Neotropical Region). 

(62) thorax and abdomen, dorsal aspect; (63) head, frontal aspect; (64) body, lateral aspect; (65) subepandrial sclerite, 

ventral aspect. Not all to the same scale.
3 4 • smithsonian contributions to zoology

Morphology: 

Head: Face at level of pseudovibrissal seta narrower than width of 1st flagellomere; medial vertical seta 

1/2 length of lateral vertical seta.

Thorax: Acrostichal setae in 1 row; scutellum 

slightly raised (angle with scutum approximately 135°), approximately triangular, apex relatively pointed. Hind femur 

of male with a basoventral lobe followed by a distinct concavity, lobe approximately rectangular; setae on lobe pale 

yellow, 4–6, lobe preceded by a few yellow setae.

Male abdomen and terminalia: Figure 69. Surstylus apically approximately 2 times the width of posterior 

process of gonopod, surstylar projection constricted subbasally, apex of surstylus approximately round to footshaped; gonopod as in Figure 69.

Measurements: Body length 3.1 mm, wing length 

2.6 mm, wing width 0.6 mm.

Type Material. The holotype male is labeled 

“ECUADOR. Orellana: Res. Etnica Waorani (0°39.4¢S, 

76°27.2¢W; 216 m; lot #1089)[,] 2 Jul 1995, T. L. Erwin/

HOLOTYPE ♂ Aulacigaster erwini Rung & Mathis 

USNM [red].” The holotype is double mounted (glued to 

a point), is in good condition (apical segments of abdomen 

removed, dissected, and stored in an attached microvial), 

and is held in trust at the USNM for eventual deposit in 

Ecuador. Paratype is as follows: same locality and collector 

as the holotype, but date of 2 Oct 1996 and lot number 

1720 (1♀; USNM).

Type Locality. Ecuador. Orellana: Reserva 

Etnica Waorani, Onkonegare Camp (1 km S; 0°39.4¢S, 

76°27.2¢W; 216 m).

Distribution. (Map 4) Neotropical: Ecuador 

(Orellana).

Etymology. The specific epithet, erwini, is 

a genitive patronym to honor and recognize Dr. Terry L. 

Erwin for his numerous contributions to entomology. He 

also was the collector of the type series.

Remarks. This species is similar to A. femorata 

but can be easily distinguished from that species by the characters given in the key and diagnosis. Aulacigaster lobata is 

also very similar to and difficult to tell apart from A. erwini. 

Both species share a yellow hind femur with a rectangular 

lobe followed by a distinct concavity. The concavity on the 

hind femur of A. erwini, however, is less pronounced than 

in A. lobata. The face of A. erwini is pale yellow, and the 

postpronotum is paler in color compared to the rest of the 

mesonotum. In A. lobata, the face is yellowish-brownish 

and the postpronotum is concolorous with the mesonotum. 

Both species also differ in the coloration of the hind tibia, 

which is mostly yellow in A. erwini (at most with a small 

FIGURES 66–68. Illustrations of Aulacigaster 

bella, new species (male) (the femorata group, 

Neotropical Region). (66) left hind femur, medial 

aspect; (67) gonopod, lateral aspect; (68) epandrium, lateral aspect. Not all to the same scale.
number 633 • 3 5

brown mark near apical 2/3) and brown on the central portion in A. lobata.

Aulacigaster femorata, new species

Figures 70–73, 207–208, Map 4

Diagnosis. This species is distinguished from 

congeners by the following combination of characters: 

Abdomen with a yellowish region on syntergite 1+2; hind 

femur of male with a basoventral lobe followed by a pronounced concavity; hind femur yellow on basal portion, 

apical portion dark.

Description. Coloration and vestiture: Vertex mostly polished; ocellar tubercle mostly polished; 

mesofrons mostly polished, or with a faint medial microtomentose stripe from base of antenna to level of anterior 

fronto-orbital seta; antenna pale yellow to yellowish, infuscate along dorsal margin; face yellow; palpus white, or 

yellowish. Scutum dull microtomentose on anteriormost 

portion (anterior half); scutellum dull microtomentose 

on lateral margins and apex, medial portion of scutellum 

bright shiny; postpronotum concolorous with mesonotum, 

mostly polished (anteriormost portion and ventral margin 

microtomentose); halter mostly white (base infuscate). 

Fore coxa yellow, fore femur yellowish, fore tibia yellowish, fore tarsus yellowish, apical and sometimes subapical 

tarsomeres brown to black; mid coxa yellow, mid femur 

yellowish or dark brown to black apically, yellowish basally, mid tibia yellowish, mid tarsus mostly yellowish; 

MAP 4. Distribution of Aulacigaster bella, A. erwini, A. femorata, and A. guyana (the femorata group, Neotropical Region).
3 6 • smithsonian contributions to zoology

hind coxa yellow, hind femur yellow on basal portion, 

apical portion dark (apical 1/2 blackish), hind tibia brown 

to black on central portion, base and apex yellowish, hind 

tarsus yellowish. Pregenital segment mostly subshiny; female abdomen subshiny (some microtomentum on lateral 

margin of tergites).

Morphology: 

Head: Figures 72, 207–208. Face at level of pseudovibrissal seta narrower than width of 1st flagellomere; 

medial vertical seta 1/2 length of lateral vertical seta.

Thorax: Figures 70–71. Acrostichal setae in 1–2 

rows, bifurcating posteriorly if 2; scutellum slightly raised 

(angle with scutum approximately 135°), trapezoidal, apex 

nearly straight or approximately triangular, apex relatively 

pointed. Hind femur of male with a basoventral lobe followed by a distinct concavity, lobe approximately rectangular; setae on lobe pale yellow, 4–6, lobe preceded by a few 

yellow setae.

Male abdomen and terminalia: Figure 73. Surstylus 

apically almost 2 times the width of posterior process of 

gonopod, surstylar projection with a sub-basal constriction, 

apex with round corners; gonopod as in Figure 73. 

Measurements: Body length 3.0–3.5 mm. Wing 

length 2.0–2.5 mm, wing width 0.8–1.0 mm.

Type Material. The holotype male is labeled “PERU. Madre de Dios: Manu, Rio Manu, 250 

m[,] Pakitza, 12°7¢S, 70°58¢W, 9–23 Sep 1988[,] Amnon 

Freidberg/HOLOTYPE ♂ Aulacigaster femorata Rung & 

Mathis USNM [red].” The holotype is double mounted 

(minuten in a block of plastic), is in excellent condition, and 

is deposited in the USNM. Paratypes are as follows: Same 

label data as the holotype (24♂, 10♀; MHNJP, USNM); 

same locality and data, collected by W. N. Mathis (11♂, 

3♀; USNM).

Type Locality. Peru. Madre de Dios: Rio 

Manu, Pakitza (11°56.6¢S, 71°16.9¢W).

Distribution. (Map 4) Neotropical: Peru 

(Madre de Dios).

Etymology. The specific epithet, femorata, is 

of Latin derivation and refers to the enlarged hindfemora 

of this species.

Remarks. Aulacigaster femorata keys out next 

to A. guyana but it can be easily separated from this species 

by the characters given in the key. Two other species, A. 

lobata and A. erwini, can also be confounded with A. femorata. The coloration of the hind femur, brown on apical 2/3 

in A. femorata, will readily separate this species from the 

other two (hind femur completely yellow in A. lobata and 

A. erwini).

FIGURES 70–71. Digital photographs of Aulacigaster femorata, 

new species (male) (the femorata group, Neotropical Region). (70) 

hindlegs, posterior aspect; (71) left hind femur, medial aspect. Not 

all to the same scale.

FIGURE 69. Illustrations of Aulacigaster erwini, new species (male) 

(the femorata group, Neotropical Region): gonopod and epandriuim, lateral aspect.
number 633 • 3 7

Aulacigaster formosa, new species

Figures 74–76, Map 5

Diagnosis. This species is distinguished from 

congeners by the following combination of characters: 

Abdomen with a yellowish region on syntergite 1+2; hind 

femur of male yellow, with a basal lobe not followed by a 

posterior concavity, lobe approximately triangular.

Description. Coloration and vestiture: Vertex 

mostly polished; ocellar tubercle mostly polished; mesofrons 

with a medial microtomentose stripe from base of antenna 

to level of anterior fronto-orbital seta; antenna pale yellow 

to yellowish, not infuscate dorsally or infuscate along dorsal, apical and ventral margins (antenna infuscate in females 

only); face white; palpus white. Scutum dull microtomentose on anteriormost portion (anterior 1/5–2/3); scutellum dull posteriorly, anterior portion shiny; postpronotum 

concolorous with mesonotum, mostly polished; halter 

mostly white. Fore coxa yellow, fore femur yellowish, fore 

tibia yellowish, fore tarsus mostly yellowish; mid coxa yellow, mid femur yellowish, mid tibia yellowish, mid tarsus 

mostly yellowish; hind coxa yellow, hind femur yellow (females with basal 1/3 infuscate with brown), hind tibia yellowish, hind tarsus yellowish. Male abdomen subshiny to 

dull, pregenital segment polished posteriorly, dull anteriorly.

Morphology: 

Head: Face at level of pseudovibrissal seta narrower than width of 1st flagellomere; medial vertical seta 

1/2 length of lateral vertical seta.

Thorax: Figure 74. Acrostichal setae in 2 rows; 

scutellum slightly raised (angle with scutum approximately 

135°), approximately triangular, apex relatively pointed. 

Hind femur of male with a basal lobe not followed by a 

posterior concavity, lobe approximately triangular; setae on 

lobe pale yellow, 4–6, preceded by a few yellow setae.

FIGURES 72–73. Illustrations of Aulacigaster femorata, new species (male) (the femorata group, Neotropical 

Region). (72) head, lateral aspect; (73) epandrium and gonopod, lateral aspect.
3 8 • smithsonian contributions to zoology

Male abdomen and terminalia: Figures 75–76. Surstylus apically approximately twice the width of anterior 

process of gonopod, surstylus constricted sub-basally, apex 

approximately round with a slight beak posteroventrally; 

gonopod as in Figure 75.

Measurements: Body length 2.5–2.7 mm. Wing 

length 0.8–0.9 mm.

Type Material. The holotype male is labeled 

“ECUADOR. Prt.[Puerto] Orll[a]na: RioTiputini (0°38.2¢S, 

76°8.9¢W)[,] 12–26 Aug 1999, W. N. Mathis, A. Baptista, 

M. Kotrba/ USNM ENT 00090416 [plastic bar code label]/

HOLOTYPE ♂ Aulacigaster formosa Rung & Mathis 

USNM [red].” The holotype is double mounted (minuten in 

a block of plastic), is in excellent condition, and is deposited 

in the USNM. Paratypes are as follows: Same label data as 

the holotype (5♂, 2♀; EPNE, USNM).

Type Locality. Ecuador. Orellana: Rio Tiputini (00°38.2¢S, 76°08.9¢W).

Distribution. (Map 5) Neotropical: Ecuador 

(Orellana).

Etymology. The specific epithet, formosa, is 

of Latin derivation and means “graceful, beautiful.”

Remarks. Aulacigaster formosa is most similar 

to A. irwini and also to A. unifasciata. Besides the characters 

given in the key and diagnosis, A. formosa can be readily 

distinguished from A. irwini by the shape of the lobe on the 

hind femur, triangular (approximately cylindrical in A. irwini). Aulacigaster formosa can be readily distinguished 

from A. unifasciata by the coloration of the mostly yellow hind tibia (with a medial to sub-apical brown mark in 

A. unifasciata) and also by characters of the male terminalia.

Aulacigaster guyana, new species

Figures 77–79, Map 4

Diagnosis. This species is distinguished from 

congeners by the following combination of characters:

Abdomen with a yellowish region on syntergite 1+2; postpronotum microtomentose on dorsal and anterior margins, 

otherwise polished; hind femur yellow on basal portion, 

apical portion dark.

Description. Coloration: Vertex mostly polished; ocellar tubercle mostly polished; mesofrons with a 

medial microtomentose stripe from base of antenna to level 

of anterior fronto-orbital seta; antenna pale yellow to yellowish, infuscate along dorsal margin; face yellow (pale); 

palpus whitish. Scutum dull microtomentose on anteriormost portion; scutellum dull posteriorly, anterior portion 

shiny; postpronotum concolorous with mesonotum, mostly 

polished, microtomentum, when present, over anterior and 

ventral margins; halter mostly white (base infuscate). Fore 

coxa yellow, fore femur yellowish, fore tarsus mostly yellowish; mid coxa yellow, mid femur yellowish, mid tarsus 

mostly yellowish; hind coxa yellow; hind femur yellow basally, apical 1/2 brown (Figure 77), hind tibia yellowish, hind 

tarsus yellowish. Male abdomen subshiny to dull, pregenital 

segment polished posteriorly, dull anteriorly (slightly so).

Morphology: 

Head: Face at level of pseudovibrissal seta narrower than width of 1st flagellomere; medial vertical seta 

1/2 length of lateral vertical seta.

Thorax: Figure 77. Acrostichal setae in 2 rows; 

scutellum slightly raised (angle with scutum approximately 135°), approximately triangular, apex relatively 

pointed. Hind femur of male with a basoventral lobe followed by a shallow concavity, lobe approximately rectangular; setae on lobe pale yellow, 4–6; lobe preceded by a 

few yellow setae.

Male abdomen and terminalia: Figures 78–79. 

Surstylus broad, surstylar projection apically more than 2 

times the width of anterior process of gonopod, with a subbasal constriction, apex with corners approximately round; 

gonopod as in Figure 78.

Measurements: Body length 2.0–2.3 mm. Wing 

length 2.0–2.3, wing width 0.6–0.7 mm.

FIGURES 74–76. Illustrations of Aulacigaster formosa, new species (male) (the femorata group, Neotropical Region). (74) left hind 

femur, medial aspect; (75) gonopod, lateral aspect; (76) epandrium, 

lateral aspect. Not all to the same scale.
number 633 • 3 9

Type Material. The holotype male is labeled “GUYANA. Moco-Moco[,] Lethem(30 km east)[,] 

3°18.2¢N, 59°39.0¢W[,] 3–6 April 1994[,] Wayne N. 

Mathis/HOLOTYPE ♂ Aulacigaster guyana Rung & 

Mathis USNM [red].” The holotype is double mounted 

(minuten in a block of plastic), is in excellent condition, and 

is deposited in the USNM. Paratypes are as follows: Same 

label data as the holotype (4♂; USNM).

Type Locality. Guyana. Moco-Moco (30 km 

E Lethem (3°18.2¢N, 59°39.0¢W).

Distribution. (Map 4) Neotropical: Guyana.

Etymology. The specific epithet, guyana, is a 

noun in apposition and recognizes the country where this 

species was collected.

Remarks. The shape of the hindfemoral lobe of 

A. guyana is characteristic for this species and, in conjunction with the coloration of the hind femur and shape of the 

surstylus (broader than in the other species of the femorata

group), make this species easy to identify.

Aulacigaster irwini, new species

Figures 80–83, Map 5

Diagnosis. This species is distinguished from 

congeners by the following combination of characters: Abdomen with a yellowish region on syntergite 1+2; postpronotum yellow to yellowish-brownish; hind femur yellow; 

MAP 5. Distribution of Aulacigaster formosa, A. irwini, A. peruana, and A. unifasciata (the femorata group, Neotropical Region).
4 0 • smithsonian contributions to zoology

lobe on hind femur approximately cylindrical, not followed 

by a concavity.

Description. Coloration and vestiture: Vertex mostly polished; ocellar tubercle mostly polished; 

mesofrons mostly polished, or with a faint medial microtomentose stripe from base of antenna to level of anterior 

fronto-orbital seta; antenna pale yellow to yellowish, infuscate along dorsal margin; face white to pale yellow; palpus 

white. Scutum mostly polished, dull microtomentose on 

anteriormost portion (anterior 1/5); scutellum dull posteriorly, anterior portion shiny, or dull microtomentose on lateral margins and apex, medial portion of scutellum bright 

shiny; postpronotum yellow to yellowish-brownish, mostly 

polished; halter mostly white. Fore coxa yellow, fore femur 

yellowish or yellow basally, apex darkened, fore tibia yellowish, fore tarsus mostly yellowish; mid coxa yellow, mid 

femur yellowish, mid tibia yellowish, mid tarsus mostly yellowish; hind coxa yellow, hind femur yellow, hind tibia yellowish (infuscate with brown at base in some specimens), 

hind tarsus yellowish. Male abdomen subshiny to dull; epandrium polished anteriorly, dull posteriorly; female abdomen subshiny.

Morphology: 

Head: Face at level of pseudovibrissal seta narrower than width of 1st flagellomere; medial vertical seta 

1/2 length of lateral vertical seta.

Thorax: Figures 80–82. Acrostichal setae in 1 row, 

or 2 rows on anteriormost portion, merging into 1 row 

posteriorly; scutellum slightly raised (angle with scutum 

approximately 135°), trapezoidal, apex nearly straight. 

Hind femur of male with a basal lobe not followed by a 

posterior concavity, lobe approximately cylindrical, bearing 1 apical and 1 or 2 subapical setae; lobe preceded by 

a few yellow setae.

Male abdomen and terminalia: Figure 83. Surstylus 2–3 times the width of posterior process of gonopod; 

lateral margins of surstylar projection approximately parallel; apex of surstylus slightly expanded and approximately 

round to approximately quadrate; gonopod as in Figure 83.

Measurements: Body length 2.4–3.0 mm. Wing 

length 2.3–2.9 mm, wing width 0.6–0.8 mm.

Type Material. The holotype male is labeled “PERU: Madre de Dios: Manu, Rio Manu, 250 m[,] 

Pakitza, 12°7¢S, 70°58¢W, 9–23 Sep 1988, Amnon Freidberg/HOLOTYPE ♂ Aulacigaster irwini Rung & Mathis 

USNM [red].” The holotype is double mounted (minuten in 

a block of plastic), is in excellent condition, and is deposited 

FIGURES 77–79. Illustrations of Aulacigaster guyana, new species (male) (the femorata group, Neotropical Region). (77) left hind 

femur, medial aspect; (78) gonopod, lateral aspect; (79) epandrium, 

lateral aspect. Not all to the same scale.

FIGURES 80–81. Digital photographs of Aulacigaster irwini, new 

species (male) (the femorata group, Neotropical Region). (80) right 

wing, dorsal aspect; (81) hindlegs, posterior aspect. Not all to the 

same scale.
number 633 • 4 1

in the USNM. Paratypes are as follows: Same label data as 

the holotype (2♂, 1♀; USNM).

Type Locality. Peru. Madre de Dios: Rio 

Manu, Pakitza (11°56.6¢S, 71°16.9¢W).

Other Specimens Examined. ECUADOR. Orellana: Reserva Etnica Waorani, Onkonegare 

Camp (1 km S; 0°39.4¢S, 76°27.2¢W; 216 m; lot #998, 

1457, 1717), 7 Feb–2 Oct 1995, 1996, T. L. Erwin (2♂, 

1♀; EPNE, USNM); Tiputini Biodiversity Station (0°37.9¢S, 

76°08.6¢W; 220–250 m; lot #2085), 8 Feb 1999, T. L. 

Erwin (1♂; USNM).

Distribution. (Map 5) Neotropical: Ecuador 

(Orellana), Peru (Madre de Dios).

Etymology. The specific epithet, irwini, is a 

genitive patronym to honor and recognize Dr. Michael E. 

Irwin for his numerous contributions to dipterology and his 

support and friendship.

Remarks. The specimens from Ecuador have 

a less shiny frons and thorax than specimens from Peru. 

Aulacigaster irwini is most similar to A. formosa and also 

to A. unifasciata. Besides the characters given in the key 

and diagnosis, A. irwini can be readily distinguished from 

A. formosa by the shape of the lobe on the hind femur, 

approximately cylindrical (triangular in A. formosa). Aulacigaster irwini differs from A. unifasciata in having the hind 

tibia mostly yellow (with a medial to sub-apical brown mark 

in A. unifasciata), and the surstylus approximately rectangular (surstylus with triangular apex in A. unifasciata).

Aulacigaster lobata, new species

Figures 84–87, Map 6

Diagnosis. This species is distinguished from 

congeners by the following combination of characters: 

Postpronotum concolorous with mesonotum; hind femur 

completely yellow, with a basoventral lobe followed by a 

distinct concavity, lobe approximately rectangular.

Description. Coloration: Vertex mostly polished; ocellar tubercle mostly polished; mesofrons with a 

faint medial microtomentose stripe from base of antenna 

to level of anterior fronto-orbital seta; antenna pale yellow 

to yellowish, infuscate along dorsal margin; face yellowish 

brown; palpus yellowish. Scutum dull microtomentose on 

anteriormost portion; scutellum dull posteriorly, anterior 

portion shiny; postpronotum concolorous with mesonotum, 

FIGURES 82–83. Illustrations of Aulacigaster irwini, new species (male) (the femorata group, 

Neotropical Region). (82) left hind femur, medial aspect; (83) epandrium and gonopod, lateral 

aspect. Not all to the same scale.
4 2 • smithsonian contributions to zoology

microtomentose over ventral and posterior margins, otherwise polished; halter mostly white (base infuscate). Male 

abdomen subshiny to dull, pregenital segment polished posteriorly, dull anteriorly (slightly so). Fore coxa pale yellow, 

fore femur yellowish, fore tibia yellowish; fore tarsus mostly 

or entirely yellowish, apical and sometimes subapical tarsomeres brown to black; mid coxa pale yellow, mid femur 

yellowish, mid tibia yellowish, mid tarsus mostly yellowish, 

apical and sometimes subapical tarsomeres brown to black; 

hind coxa pale yellow, hind femur yellow (rarely with apex 

brown infuscate), hind tibia brown to black on central portion, base and apex yellowish, hind tarsus yellowish.

Morphology: 

Head: Face at level of pseudovibrissal seta narrower than width of 1st flagellomere; medial vertical seta 

1/2 length of lateral vertical seta.

Thorax: Figures 84–86. Acrostichal setae in 1 

row; scutellum slightly raised (angle with scutum approximately 135°), approximately triangular, apex relatively 

pointed. Hind femur of male with a basoventral lobe followed by a distinct concavity, lobe approximately rectangular; setae on lobe pale yellow, 4–6, lobe preceded by a 

few yellow setae.

Male abdomen and terminalia: Figure 87. Surstylus apically approximately 2 times the width of posterior 

process of gonopod, surstylar projection constricted subbasally, apex of surstylus approximately round to footshaped; gonopod as in Figure 87.

Measurements: Body length 3.1–3.5 mm. Wing 

length 2.7–2.9 mm, wing width 0.8–1.0 mm.

Type Material. The holotype male is labeled “PERU. Madre de Dios: Manu, Rio Manu, 250 m[,] 

Pakitza, 12°7¢S, 70°58¢W, 9–23 Sep 1988[,] Wayne N. Mathis/HOLOTYPE ♂ Aulacigaster lobata Rung & Mathis 

USNM [red].” The holotype is double mounted (minuten 

in a block of plastic), is in excellent condition, and is deposited in the USNM. Paratypes are as follows: Same label data 

as the holotype (2♂; USNM); same locality data but collected by A. Freidberg (7♂; USNM). PERU. Madre de Dios: 

Manu, Rio Manu, Pakitza, Aguajal (11°58.2¢S, 71°17¢W), 

19 Sep 1988, A. Freidberg, (1♂; USNM).

Type Locality. Peru. Madre de Dios: Rio 

Manu, Pakitza (11°56.6¢S, 71°16.9¢W).

Distribution. (Map 6) Neotropical: Peru 

(Madre de Dios).

Etymology. The specific epithet, lobata, is of 

Greek derivation and means rounded projection or protuberance with reference to the shape of the lobe on the 

hind femur.

Remarks. This species is similar to A. erwini

and A. femorata. Aulacigaster femorata can be readily 

distinguished by the coloration of the hind femur which is 

brown on apical 2/3. Separation from A. lobata is trickier. 

Both species share a completely yellow hind femur with a 

rectangular lobe followed by a distinct concavity. The concavity in the hind femur of A. erwini, however, is less pronounced than in A. lobata. The face of A. erwini is pale 

yellow and the postpronotum is paler in color as compared 

with the rest of the mesonotum. In A. lobata, the face is 

yellowish-brownish and the postpronotum is concolorous 

with the mesonotum. Both species differ also in the coloration of the hind tibia, which is mostly yellowish in A. erwini (at most with a small brown mark near apical 2/3) and 

brown on central portion in A. lobata.

FIGURES 84–85. Digital photographs of Aulacigaster 

lobata, new species (the femorata group, Neotropical Region). (84) right wing, dorsal aspect; (85) enlargement of 

costal section, dorsal aspect.
number 633 • 4 3

Aulacigaster nigritibia, new species

Figures 88–91, Map 6

Diagnosis. This species is distinguished from 

congeners by the following combination of characters: 

Abdomen with a yellowish region on syntergite 1+2; hind 

femur mostly dark brown, yellowish at basal 1/5–1/4.

Description. Coloration and vestiture: Vertex 

mostly polished; ocellar tubercle mostly polished; mesofrons with a medial microtomentose stripe from base of antenna to level of anterior fronto-orbital seta (stripe shaped 

as an inverted triangle); antenna pale yellow to yellowish, 

infuscate along dorsal margin; face white to yellow; palpus white. Scutum dull microtomentose on anteriormost 

portion; scutellum dull posteriorly, anterior portion shiny; 

postpronotum concolorous with mesonotum, scarcely microtomentose, subshiny; halter mostly white (infuscate at 

base). Fore coxa yellow, fore femur yellow basally, apex 

darkened, fore tibia yellowish, fore tarsus mostly or entirely yellowish, apical and sometimes subapical tarsomeres 

brown to black; mid coxa yellow, mid femur yellowish or 

dark brown to black apically, yellowish basally, mid tibia 

mostly or entirely yellowish on central portion, apex and 

basis brown, mid tarsus mostly yellowish; hind coxa yellow, 

hind femur mostly dark brown, yellowish at basal 1/5–1/4, 

hind tibia brown to dark brown, or brown to black on central portion, base and apex yellowish, hind tarsus yellowish. 

Male abdomen bright shiny (tergite 5 more uniformly microtomentose), pregenital segment polished posteriorly, dull 

anteriorly; female abdomen with dense microtomentum 

medially, lateral margins of tergites nearly polished.

Morphology: 

Head: Figure 88. Face at level of pseudovibrissal 

seta narrower than width of 1st flagellomere; medial vertical seta 1/2 length of lateral vertical seta.

Thorax: Figure 91. Acrostichal setae in 1 row, 

or 2 rows on anteriormost portion, merging into 1 row 

posteriorly; scutellum slightly raised (angle with scutum 

approximately 135°), approximately triangular, apex relatively pointed. Hind femur of male with a basoventral lobe 

followed by a slight to deep concavity, lobe approximately 

cylindrical; setae on lobe pale yellow, 2; lobe preceded by 

2–3 yellow setae.

Male abdomen and terminalia: Figures 89–90. Surstylus 1.3 times the width of posterior process of gonopod; 

lateral margins of surstylar projection sinuous, convergent 

towards apex; gonopod as in Figure 89.

Measurements: Body length 2.2 mm. Wing length 

2.1 mm, wing width 0.7 mm.

Type Material. The holotype male is labeled “PERU. Madre de Dios: Manu, Rio Manu, 250 m[,] 

Pakitza, 12°7¢S, 70°58¢W, 9–23 Sep 1988[,] Wayne N. Mathis/HOLOTYPE ♂ Aulacigaster nigritibia Rung & Mathis 

USNM [red].” The holotype is double mounted (minuten in 

a block of plastic), is in excellent condition, and is deposited 

FIGURES 86–87. Illustrations of Aulacigaster 

lobata, new species (male) (the femorata group, 

Neotropical Region). (86) left hind femur, medial 

aspect; (87) gonopod and epandrium, lateral aspect. Not all to the same scale.
4 4 • smithsonian contributions to zoology

in the USNM. The holotype is double mounted (minuten in 

a block of plastic), is in good condition (abdomen removed 

and dissected, anterior left and posterior right fronto-orbital setae as well as vertical setae broke), and is deposited 

in the USNM.

Type Locality. Peru. Madre de Dios: Manu, 

Rio Manu, Pakitza (12°7¢S, 70°58¢W; 250 m).

Distribution. (Map 6) Neotropical: Peru 

(Madre de Dios).

Etymology. The species epithet, nigritibia, is 

of Greek derivation and refers to the mostly darkened hind 

tibia.

Remarks. Besides the characters given in the 

key and diagnosis, this species is easy to separate from congeners based on the shape of the surstylus (as in Figure 90) 

and the cylindrical lobe on the hind femur, which is followed by a slight to deep concavity.

Aulacigaster peruana, new species

Figures 92–95, Map 5

Diagnosis. This species is distinguished from 

congeners by the following combination of characters: 

Abdomen with a yellowish region on syntergite 1+2; frons 

mostly brown with anterior margin pale, yellowish; hind 

femur of male without a conspicuous lobe.

Description. Coloration and vestiture: Vertex 

mostly polished; ocellar tubercle mostly polished; mesofrons 

mostly polished, or with a faint medial microtomentose 

MAP 6. Distribution of Aulacigaster lobata and A. nigritibia (the femorata group, Neotropical Region).
number 633 • 4 5

stripe from base of antenna to level of anterior fronto-orbital seta; antenna pale yellow to yellowish, infuscate along 

dorsal margin; face white to yellow; palpus white. Scutum 

dull microtomentose on anteriormost portion (anterior 

1/6–1/2); scutellum dull posteriorly, anterior portion shiny 

(completely microtomentose in some specimens); postpronotum concolorous with mesonotum, scarcely microtomentose, subshiny; halter mostly white (base infuscate). 

Fore coxa yellow, fore femur yellowish, fore tibia yellowish, 

fore tarsus mostly yellowish; mid coxa yellow, mid femur 

yellowish, mid tibia yellowish or mostly yellowish, apex 

brown infuscate (very faint in male), mid tarsus mostly 

yellowish; hind coxa yellow, hind femur yellow on basal 

portion, apical portion dark (apical 1/3–1/2), hind tibia yellowish, hind tarsus yellowish. Male abdomen subshiny to 

dull, pregenital segment polished posteriorly, dull anteriorly; female abdomen with dense microtomentum over anterior margin of tergites, posterior margin of tergites bright 

shiny (tergites 1–3 of in some specimens also with lateral 

microtomentum).

Morphology: 

Head: Face at level of pseudovibrissal seta narrower than width of 1st flagellomere; medial vertical seta 

1/2 length of lateral vertical seta.

Thorax: Figures 92–93. Acrostichal setae in 2 rows; 

scutellum slightly raised (angle with scutum approximately 

FIGURES 88–91. Illustrations of Aulacigaster nigritibia, new species (male) (the femorata group, 

Neotropical Region). (88) head, lateral aspect; (89) gonopod, lateral aspect; (90) epandrium, lateral 

aspect; (91) left hind femur, medial aspect. Not all to the same scale.
4 6 • smithsonian contributions to zoology

135°), approximately triangular, apex relatively pointed (to 

triangular). Hind femur of male without a sub-basal lobe.

Male abdomen and terminalia: Figures 94–95. Surstylus slightly angled posteriorly, surstylar projection narrowing apically, apex approximately 3 times the width of 

anterior process of gonopod; gonopod as in Figure 94.

Measurements: Body length 2.5–3.3 mm. Wing 

length 2.6–2.9 mm, wing width 0.7–0.9 mm.

Type Material. The holotype male is labeled “PERU. Madre de Dios: Manu, Rio Manu, 250 m[,] 

Pakitza, 12°7¢S, 70°58¢W, 9–23 Sep 1988[,] Wayne N. 

Mathis/HOLOTYPE ♂ Aulacigaster peruana Rung & Mathis USNM [red].” The holotype is double mounted (minuten in a block of plastic), is in excellent condition, and 

is deposited in the USNM. Paratypes are as follows: Same 

label data as the holotype (48♂, 7♀; USNM). Same locality and data, collected by A. Freidberg (22♂, 6♀; MHNJP, 

USNM).

Type Locality. Peru. Madre de Dios: Rio 

Manu, Pakitza (11°56.6¢S, 71°16.9¢W).

Other Specimens Examined. BOLIVIA. 

La Paz: S. Inicua, Rio Alto Beni (1100 m; 15°20¢W, 

67°33¢W), 15–18 Jan 1976, L. E. Peña (1♂; CNC); Mapiri 

(15°18.6¢S, 68°13¢W; 720 m), 16 Mar 2001, A. Freidberg 

(2♂, 1♀; USNM).

PERU. Madre de Dios: Rio Tambopata Reserve 

(30 km SW Puerto Maldorado; 12°50.4¢S, 69°17.4¢W), 

19 Sep–10 Oct 1984, D. A. Grimaldi (7♂, 2♀; AMNH).

Distribution. (Map 5) Neotropical: Bolivia 

(La Paz), Peru (Madre de Dios).

Etymology. The species epithet, peruana,

recognizes the country, Peru, where the type series was 

collected.

Remarks. This species is peculiar amongst congeners in the femorata group by lacking a lobe on the hind 

femur of the male. The wide surstylus, which is angled posteriorly, is also characteristic for A. peruana.

Aulacigaster unifasciata, new species

Figures 96–98, Map 5

Diagnosis. This species is distinguished from 

congeners by the following combination of characters: Abdomen with a yellowish region on syntergite 1+2; up to 

apical half of wing brown infuscate; hind femur yellow on 

basal 1/2–2/3, apical portion dark.

Description. Coloration and vestiture: Vertex 

mostly polished; ocellar tubercle mostly polished; mesofrons mostly polished, or with a pair of silver microtomentose marks; antenna pale yellow to yellowish, infuscate 

along dorsal margin; face yellow; palpus white. Scutum 

dull microtomentose on anteriormost portion; scutellum 

dull microtomentose on lateral margins and apex, medial 

portion of scutellum bright shiny; postpronotum concolorous with mesonotum, mostly polished; halter mostly white. 

Wing with up to apical 1/2 brown infuscate, remaining 

wing light brown. Fore coxa yellow, fore femur yellowish, 

FIGURES 92–93. Digital photographs of Aulacigaster peruana, new 

species (male) (the femorata group, Neotropical Region). (92) left 

hind femur, medial aspect; (93) right wing, dorsal aspect. Not to the 

same scale.

FIGURES 94–95. Illustrations of Aulacigaster peruana, new species 

(male) (the femorata group, Neotropical Region). (94) gonopod, lateral aspect; (95) epandrium, lateral aspect.
number 633 • 4 7

fore tibia yellowish, fore tarsus mostly yellowish; mid coxa 

yellow, mid femur yellowish (slightly infuscate on apical 1/3 

in female specimens), mid tibia yellowish, mid tarsus mostly 

yellowish; hind coxa yellow, hind femur yellow on basal 1/2 

to basal 2/3, apical portion dark , hind tibia brown to black 

on central portion, base and apex yellowish, hind tarsus 

yellowish. Male abdomen subshiny to dull; female abdomen subshiny.

Morphology: 

Head: Face at level of pseudovibrissal seta narrower than width of 1st flagellomere; inner vertical seta 1/2 

length of outer vertical seta.

Thorax: Figure 96. Acrostichal setae 2 rows on anteriormost portion, merging into 1 row posteriorly; scutellum slightly raised (angle with scutum approximately 135°), 

trapezoidal, apex nearly straight. Hind femur of male with 

a basal lobe not followed by a posterior concavity, lobe approximately cylindrical, bearing two apical setae and one 

subapical seta; lobe preceded by a few small setae.

Male abdomen and terminalia: Figures 97–98. Surstylus more than twice the width of posterior process of 

gonopod; lateral margins of surstylar projection approximately parallel; apex of surstylus approximately triangular; 

gonopod as in Figure 97.

Measurements: Body length 2.9–3.3 mm. Wing 

length 2.5–3.1 mm, wing width 0.8–1.0 mm.

Type Material. The holotype male is labeled 

“ECUADOR. Orellana, Res. Etnica Waorani (0°39.4¢ S, 

76°27.2¢ W, 216m; lot # 1581, 26 Jun 1998, T.L.Erwin/

HOLOTYPE ♂ Aulacigaster unifasciata Rung & Mathis 

[red].” The holotype is double mounted (glued to a point), 

is in excellent condition (apical segments of abdomen removed, dissected, and stored in an attached microvial), and 

is held in trust at the USNM for eventual deposit in Ecuador. Paratypes are as follows: same locality and collector as 

the holotype, but with dates of 15 Jan–1 Oct 1994, 1995, 

1996 and lot numbers 577, 700, 1160, 1430, 1690 (1♂, 

4♀; EPNE, USNM).

FIGURES 96–98. Illustrations of Aulacigaster unifasciata, new species (male) (the femorata group, 

Neotropical Region). (96) right hind femur, medial aspect; (97) gonopod, lateral aspect; (98) epandrium, lateral aspect.
4 8 • smithsonian contributions to zoology

Type Locality. Ecuador. Orellana: Reserva 

Etnica Waorani, Onkonegare Camp (1 km S; 0°39.4¢S, 

76°27.2¢W; 216 m).

Other Specimens Examined. PERU. 

Madre de Dios: Rio Manu, Erika (near Salvación; 12°53¢S, 

71°12¢W; 550 m), 5–6 Sep 1988, A. Freidberg (1♂, 1♀; 

USNM).

Distribution. (Map 5) Neotropical: Ecuador 

(Orellana), Peru (Madre de Dios).

Remarks. This species is similar to A. irwini. 

In addition to characters given in the key and diagnosis, 

A. unifasciata can be readily distinguished from this species by the coloration of the hind tibia (with a more conspicuous central brown to black mark), and the shape of 

the surstylus.

The grimaldii Group

Figures 99–106, 209–210, Map 7

Diagnosis. The grimaldii group is distinguished from other species groups by the following combination of characters: Small flies, body length 1.88–2.80 

mm. Coloration and vestiture: Frons mostly brown, anterior margin pale, yellowish; anterior portion of frons with 

a veil of microtomentum anterior to lower fronto-orbital 

setae, but not silver. Anepisternum polished. Abdomen with 

syntergite 1+2 whitish.

Morphology: 

Head: Round, about as high as long (head ratio 

0.9–1.1); face at level of pseudovibrissal seta narrower than 

width of 1st flagellomere, in lateral view projected from 

base of antenna to level of pseudovibrissal seta, thereafter 

receded to buccal opening (males) or strongly convex on 

ventral 2/3, easily visible in lateral view (females), bulbous; 

ocellar seta typically minute; fronto-orbital setae typically 

with posterior seta only slightly internal to anterior seta.

Thorax: Subcosta partially fused with vein R1

 apically but terminating in costa.

Male abdomen and terminalia: Surstylus often arising from posteroventral margin of epandrium; subepandrial 

sclerite Y-shaped. Gonopods short, wide, with a few central 

setulae and a posterior process. Cerci partially fused, each 

cercus bearing 1 very long, posteriorly porrect seta.

Distribution. Neotropical: Bolivia, Colombia, Costa Rica, Dominican Republic, Dominica, Ecuador, 

Guyana, Panama, Peru, Venezuela.

Discussion. Species of the grimaldii group can 

be best identified by the coloration of the face and legs, and 

the shape of the surstylus and other structures of the male 

internal terminalia.

The grimaldii group includes two species, none described previously.

Aulacigaster angusta, new species

Figure 99, Map 7

Diagnosis. This species is distinguished from 

congeners by the following combination of characters: 

Abdomen with syntergite 1+2 white. Frons mostly brown, 

with anterior margin pale yellowish; face brown.

Description. Coloration and vestiture: Vertex 

mostly polished; ocellar tubercle mostly microtomentose; 

antenna pale yellow to yellowish, infuscate along dorsal 

margin; face brown; palpus white. Scutum strongly microtomentose medially, polished between pleural region and 

dorsocentral row of setae, posterior of postpronotum; scutellum dull microtomentose; postpronotum pale brown, or 

concolorous with mesonotum, scarcely microtomentose, 

subshiny; katepisternum mostly polished; halter mostly 

white. Fore coxa mostly yellow, brown basally; fore femur 

mostly brown, fore tibia mostly brown, fore tarsus mostly 

brown, with apical and subapical tarsomere infuscate with 

brown; mid coxa yellow, mid femur mostly brown (tip and 

base yellowish), mid tibia yellowish, mid tarsus yellowish, 

apical and subapical tarsomere brownish to dark brown; 

hind coxa yellow, hind femur mostly brown (tip and base 

yellowish), hind tibia brown basally, yellowish apically, 

hind tarsus yellowish, apical tarsomere brown to black. 

Male abdomen bright shiny.

Morphology: 

Head: Head round, about as high as long (head 

ratio 1.1); medial vertical seta 1/2 length of lateral vertical 

seta; fronto-orbital setae with posterior seta slightly internal 

to and almost transversely aligned with anterior seta; peristomal vestiture consisting mostly of weak setae following 

pseudovibrissal seta (unusually thin); 1 strong peristomal 

seta present, following pseudovibrissal seta.

Thorax: Acrostichal setae in 2 rows; scutellum 

slightly raised (angle with scutum approximately 135°), 

approximately triangular, apex relatively pointed, disk of 

scutellum slightly convex; basal scutellar seta 1/2 length of 

apical seta.

Male abdomen and terminalia: Figure 99. Surstylus 

short (shorter than in A. grimaldii), with lateral margins 

subparalell, apex approximately round. Anterior process of 

gonopod making a right angle with posterior process.
number 633 • 4 9

Measurements: Body length 2.5–2.8 mm. Wing 

length 2.3 mm, wing width 0.9 mm.

Type Material. The holotype male is labeled “COSTA RICA: Puntarenas, Las Alturas,20 km NE 

San Vito de Hava, 1500 m.[,] 20/III/91 [20 Mar 1991], 

Grimaldi&Stark[,] sweeping tree trunks./HOLOTYPE ♂

Aulacigaster angusta Rung & Mathis AMNH[red].” The 

holotype is double mounted (glued to a point), is in excellent condition, and is deposited in the AMNH.

Type Locality. Costa Rica. Puntarenas: 

Las Alturas (20 km NE San Vito de Hava; 08°56.4¢S, 

82°50.1¢W).

Distribution. (Map 7) Neotropical: Costa 

Rica (Puntarenas).

Etymology. The species epithet, angusta, is of 

Latin derivation and means narrow in reference to the narrow face of males.

Remarks. This species can be easily identified 

from the only other species in this group, A. grimaldii, by 

the characters given in the key. Aulacigaster angusta also 

differs from that species by having the ocellar tubercle 

mostly microtomentose.

Aulacigaster grimaldii, new species

Figures 100–106, 209–210, Map 7

Diagnosis. This species is distinguished from 

congeners by the following combination of characters: 

Abdomen with syntergite 1+2 white. Frons mostly brown, 

with anterior margin pale yellowish; face yellowish.

Description. Coloration and vestiture: Vertex 

mostly polished; ocellar tubercle mostly polished; antenna 

pale yellow to yellowish, infuscate along dorsal margin; 

face yellow; palpus white. Scutum strongly microtomentose 

medially, polished between pleural region and dorsocentral 

row of setae, posterior of postpronotum; scutellum dull microtomentose; postpronotum pale brown, or concolorous 

with mesonotum, scarcely microtomentose, subshiny; katepisternum mostly polished; halter mostly white. Fore coxa 

yellow, fore femur yellowish, with apex brown infuscate in 

some specimens; fore tibia mostly brown (base yellowish), 

fore tarsus with basal tarsomere yellowish, remainder tarsomeres infuscate with brown, or brown to black (apical 

and subapical); mid coxa yellow, mid femur yellowish (tip 

darkened), mid tibia yellowish, mid tarsus yellowish, apical and subapical tarsomere brownish to dark brown; hind 

coxa yellow, hind femur yellow on basal portion, apical 

portion dark (1/2), hind tibia yellowish, hind tarsus yellowish, apical tarsomere brown to black. Male abdomen bright 

shiny; female abdomen subshiny.

Morphology: 

Head: Figures 103–104, 209–210. Head round, 

about as high as long (head ratio 0.9–1.1); medial vertical 

seta 1/2 length of lateral vertical seta; fronto-orbital setae 

with posterior seta slightly internal to and almost transversely aligned with anterior seta; peristomal vestiture consisting mostly of weak setae following pseudovibrissal setal 

seta (unusually thin); 1 strong peristomal seta present, following pseudovibrissal seta.

Thorax: Figures 100–102. Acrostichal setae in 2 

rows; scutellum slightly raised (angle with scutum approximately 135°), approximately triangular, apex relatively 

pointed, disk of scutellum slightly convex; basal scutellar 

seta 1/2 length of apical seta.

Male abdomen and terminalia: Figures 105–106. 

Surstylus longer than in A. angusta, with margins slightly 

converging toward round apex. Anterior process of 

gonopod making a wide angle with posterior gonopodal 

process.

Female abdomen: Sternites 1–5 appearing divided, 

often weakly sclerotized.

FIGURE 99. llustrations of Aulacigaster angusta, new species (male) 

(the grimaldii group, Neotropical Region): epandrium and gonopod, 

lateral aspect. 
5 0 • smithsonian contributions to zoology

Measurements: Body length 1.8–2.2 mm. Wing 

length 1.8–2.2 mm. Wing width 0.6–0.7 mm.

Type Material. The holotype male is labeled “VENEZUELA: Amazonas[,] Rio Mavaca Camp 

65°06¢W–2°2¢N[,] 150 m, 16-27/III/89 [16–27 Mar 1989]/

Phipps-FUDCI Expedition by Amer. Mus. Nat. Hist. D. A. 

Grimaldi, coll./HOLOTYPE ♂ Aulacigaster grimaldi Rung 

& Mathis AMNH[red].” The holotype is double mounted 

(glued to a point), is in excellent condition, and is deposited 

in the AMNH. Paratypes are as follows: same label data 

as the holotype (1♂, 2♀; AMNH, USNM). VENEZUELA. 

Maracay: Rancho Grande Biological Station (Portochuelo 

Pass; 10°02.8¢N, 67°33¢W; 1400 m), 27 Feb 1995, S. A. 

Marshall (1♀; UGE).

Type Locality. Venezuela. Amazonas: Rio 

Mavaca Camp (02°02¢N, 65°06¢W).

Other Specimens Examined. BOLIVIA. La 

Paz: Arroyo Tuhiri (5 Km W Mapiri; 15°17.8¢S, 68°15.6¢W; 

750 m), 19 Mar 2001, S. D. Gaimari (1♂; USNM).

COLOMBIA. Valle: Río Raposo (03°40¢S, 75°05¢W; 

light trap), Aug–Oct 1964, V. H. Lee (2♀; USNM).

COSTA RICA. Puntarenas: Peninsula de Osa, Rincón 

(08°42¢N, 83°29¢W; sea level; sweeping in palm-mangrove 

forest), 24 Mar 1991, D. A. Grimaldi, J. Stark (2♂; 

USNM); Peninsula de Osa, Rincón (08°42¢N, 83°29¢W; 

Malaise trap), Oct–Dec 1990, P. Hanson (2♀; USNM).

DOMINICA. Pont Cassé (15°22.8¢N, 61°20.8¢W, 

12–14 Oct 1964, P. J. Spangler (1♂; USNM).

MAP 7. Distribution of the grimaldii species group of Aulacigaster (Neotropical Region).
number 633 • 5 1

DOMINICAN REPUBLIC. Peravia: San José de Ocoa 

(30.6 km N; 18°33.8¢N, 70°30¢W; 915 m; near stream), 

30 Jul 1991, D. A. Grimaldi, J. Stark (1♂; AMNH).

ECUADOR. Orellana: Reserva Etnica Waorani, 

Onkonegare Camp (1 km S; 0°39.4¢S, 76°27.2¢W; 216 m; 

lots 1407, 1578, 1733, 2071), 4 Feb–3 Oct 1996, 1999, 

T. L. Erwin (3♂, 1♀; EPNE, USNM).

GUYANA. Kumo River (25 km SE Lethem in Kanuku 

Mountains; 03°15.9¢N, 59°43.6¢W), 3–6 Apr 1994, W. N. 

Mathis (1♂; USNM).

FIGURES 100–104. Digital photographs of Aulacigaster grimaldii, new species (the grimaldii group, 

Neotropical Region). (100) body of male, lateral aspect; (101) body of male, dorsal aspect; (102) body 

of female, lateral aspect; (103) head of male, lateral aspect; (104) head of female, lateral aspect. Not all 

to the same scale.
5 2 • smithsonian contributions to zoology

PANAMA. Canal Zone: Barro Colorado Island 

(08°19¢N, 82°38¢W; on fresh-cut log), 15–30 Jun 1986, 

D. A. Grimaldi (1♂; AMNH).

PERU. Madre de Dios: Rio Manu, Pakitza (11°56.6¢S, 

71°16.9¢W; 250 m), 9–23 Sep 1988, A. Freidberg (1♂; 

USNM); Rio Tambopata Reserve (30 km SW Puerto Maldonado; 12°50.4¢S, 69°17.4¢W; tropical moist forest) 

19 Sep–10 Oct 1984, D. A. Grimaldi (1♀; AMNH).

VENEZUELA. Táchira: Pregonero, Presa Las Cuevas 

(8°01.5¢N, 71°45.8¢W; 650 m; rain forest; Malaise trap), 

9–31 Jul 1989, S. and J. Peck (1♀; UGE).

Distribution. (Map 7) Neotropical: Bolivia 

(La Paz), Colombia, Costa Rica (Puntarenas), Dominica, 

Dominican Republic (Peravia), Ecuador (Orellana), Guyana, Panama (Canal Zone), Peru (Madre de Dios), Venezuela (Amazonas, Táchira).

Etymology. The specific epithet, grimaldii, is 

a genitive patronym to honor and recognize Dr. David A. 

Grimaldi for his numerous contributions to dipterology and 

his friendship.

Remarks. This species can be easily identified 

from the only other species in this group, A. angusta, by 

the characters given in the key and also by having a mostly 

polished ocellar tubercle.

The minuta Group

Figures 107–131, 211–212, Map 8

Diagnosis. The minuta group is distinguished 

from other species groups by the following combination of 

characters: Relatively small, slender, delicate looking, dorsoventrally flattened flies with elongate body, body length 

2.3–3.6 mm. Coloration and vestiture: Vertex mostly 

polished; ocellar tubercle mostly polished; frons typically 

mostly brown, anterior margin pale, yellowish, sometimes 

invested with dense microtomentum. Scutum mostly devoid 

of microtomentum; anepisternum polished; katepisternum 

mostly polished. Wing generally hyaline, infuscate with 

brown in some species.

Morphology: Body elongate, scutal ratio scutal ratio 

1.5–1.75. 

Head: Head typically longer than high (head ratio 

higher than 1.1); face slightly convex, barely visible in 

FIGUREs 105–106. Illustrations of Aulacigaster grimaldii, new species (male) (the grimaldii group, Neotropical Region). (105) subepandrial sclerite, ventral aspect; 

(106) epandrium and internal genitalic structures (male), 

lateral aspect. Not all to the same scale.
number 633 • 5 3

lateral view; face at level of pseudovibrissal seta slightly 

narrower than width of 1st flagellomere; ocellar seta absent; medial vertical setae typically convergent and slightly 

proclinate.

Thorax: Subcosta incomplete, fused with vein R1

apically. Hind femur of male and female modified, with 2 

ventral rows of spines, each spine bearing an apical seta; 

hind tibia with a row of tiny ventral spines. Hind femur of 

male often with a lobe.

Male abdomen and terminalia: Sternites 3–5 generally unmodified. Surstylus in lateral view a wide, ventroposterior extension of epandrium; cerci separate or partially 

fused; subepandrial sclerite forming a single, T-shaped 

structure; gonopods approximately rectangular, with a 

posterior, digitiform process and ventral, wide sclerotized 

plates; cercus bearing 1 very long, ventral seta and 1–3 longer setulae.

Distribution. Neotropical: Bolivia, Costa 

Rica, Ecuador, Panama, Peru.

Biology. The biology of species belonging 

to this group is mostly unknown. One male of A. appendiculata was collected on cacao foliar nectaries. Specimens 

of A. gaimarii and A. minuta were collected on leaves of 

Heliconiaceae.

Discussion. Species of the minuta group can 

be best identified by the shape of the basal lobe on the male 

hind femur and coloration of the legs: as with the femorata

group, the shape of the surstylus and internal male terminalia do not provide many characters that are useful in species 

identification.

The minuta group includes four species of which one, 

A. minuta (Hennig), was previously described. The group 

is exclusively Neotropical.

Aulacigaster appendiculata, new species

Figures 107–111, Map 8

Diagnosis. This species is distinguished from 

congeners by the following combination of characters: 

Body elongate, scutal ratio 1.5–1.75; thoracic pleura uniformly black; wing hyaline (at most with tip infuscate with 

brown); hind femur of male with a distinct basoventral lobe 

followed by a distinct concavity.

Description. Coloration and vestiture: Mesofrons mostly polished, or with a faint medial microtomentose stripe from base of antenna to level of anterior 

fronto-orbital seta; antenna pale yellow to yellowish, infuscate along dorsal margin; face brownish; antenna pale 

yellow to yellowish, infuscate along dorsal margin; palpus 

white. Scutum mostly devoid of microtomentum; scutellum 

dull posteriorly, anterior portion shiny; postpronotum concolorous with mesonotum, mostly polished; halter mostly 

yellowish. Fore coxa yellow, fore femur yellowish, fore tibia 

yellowish, fore tarsus mostly yellowish; mid coxa yellow, 

mid femur yellowish, mid tibia yellowish, mid tarsus mostly 

yellowish; hind coxa yellow, hind femur mostly yellowish, 

apex brown infuscate, hind tibia brown to black on central portion, base and apex yellowish, hind tarsus yellowish, apical and sometimes subapical tarsomeres brown to 

black. Male abdomen subshiny to dull; female abdomen 

with dense microtomentum over syntergite 1+2, and anterior margin of remaining segments.

Morphology: 

Head: Figures 109–110. Medial vertical seta 1/2 

length of lateral vertical seta; peristomal area with about 

3–5 setae following pseudovibrissal seta.

Thorax: Figures107–108. Acrostichal setae in 1 

row; scutellum slightly raised (angle with scutum approximately 135°), approximately triangular, apex relatively 

pointed, disk of scutellum flat; basal scutellar seta less than 

1/2 length of posterior seta. Hind femur of male with a 

basoventral lobe followed by a distinct concavity, lobe approximately rectangular, long (about 1/3 the length of basal 

tarsomere), bearing a few apical and subapical setae and 

preceded by a few weak setae

Male abdomen and terminalia: Figure 111. Surstylus at most 3 times the width of posterior process of 

gonopod; lateral margins of surstylar projection slightly 

convergent toward apex, with a subapical constriction; 

apex of surstylus roughly round; gonopod as in Figure 111.

Measurements: Body length 2.5–2.8 mm. Wing 

length 2.2–2.4 mm, wing width 0.6 mm.

Type Material. The holotype male is labeled 

“COSTA RICA: Heredia Pr.: La Selva Biol.[ogical] Sta.

[tion] 3 Km S P[uer]to. Viejo 10°26¢N 84°01¢W/18 vi 1991 

[18 Jun 1991] H. A. Hespenheide/HOLOTYPE ♂ Aulacigaster appendiculata Rung & Mathis USNM [red].” The 

holotype is double mounted (glued to a paper point), is in 

excellent condition, and is deposited in USNM. Paratypes 

are as follows: Same label data as the holotype but with 

dates as follows: 2 May 1990, (1♂; UCLA); 8 Jul 1987 

(at foliar nectaries of Byttenaria aculeata Jacq., plant # ?, 

cacao), H. A. Hespenheide (1♂; CAS).

Type Locality. Costa Rica. Heredia: La 

Selva Biological Station (3 km S Puerto Viejo; 10°26¢N, 

84°01¢W).

Other Specimens Examined. COSTA 

RICA. Heredia: La Selva Biological Station (3 km S Puerto 

Viejo; 10°26¢N, 84°01¢W; at foliar nectaries of Byttenaria 
5 4 • smithsonian contributions to zoology

aculeata Jacq., plant # ?, cacao), 8–26 Jul 1982, H. A. Hespenheide (5♂, 14♀; CSCA).

Distribution. (Map 8) Neotropical: Costa 

Rica (Heredia).

Etymology. The specific epithet, appendiculata, is of Latin derivation and means “little appendage” 

with reference to the lobe on the hind femur.

Remarks. This species is closest to and shares a 

similar geographical distribution with A. costaricana. Males 

of both species can be readily distinguished by the characters 

given in the key and diagnosis. Females of A. appendiculata

and A. costaricana cannot be distinguished with certainty 

when males from the same series are not available.

Aulacigaster condylura, new species

Figures 112–115, Map 8

Diagnosis. This species is distinguished from 

congeners by the following combination of characters: Body 

FIGUREs 107–110. Digital photographs of Aulacigaster appendiculata, new species (male) (the minuta group, Neotropical Region). (107) body, lateral aspect; (108) left hind femur, medial aspect; (109) head, frontal aspect; (110) head, 

lateral aspect. Not all to the same scale.
number 633 • 5 5

elongate, scutal ratio 1.5–1.75; pleura uniformly black; scutellum dull microtomentose on lateral margins and apex, 

medial portion of scutellum bright shiny.

Description. Coloration and vestiture: Mesofrons mostly polished; antenna pale yellow to yellowish, 

infuscate along dorsal margin in some specimens; face whitish or yellowish; palpus white. Scutum mostly devoid of 

microtomentum; scutellum dull microtomentose on lateral 

margins and apex, medial portion shiny; postpronotum concolorous with mesonotum, mostly polished; halter mostly 

white. Wing mostly brown, infuscate. Fore coxa yellow, 

fore femur yellowish, fore tibia yellowish, fore tarsus mostly 

yellowish; mid coxa yellow, mid femur yellowish or dark 

brown to black apically, yellowish basally, mid tibia yellowish, mid tarsus mostly yellowish; hind coxa yellow, hind 

femur mostly brown, yellow on basal 1/3 to basal 1/2, hind 

tibia brown to black on central portion, base and apex yellowish, hind tarsus yellowish. Male abdomen subshiny to 

dull, pregenital segment polished posteriorly, dull anteriorly; 

female abdomen with dense microtomentum over syntergite 

1+2, and anterior margin of remaining segments.

Morphology: 

Head: Medial vertical seta 1/2 length of lateral vertical seta; 1–2 peristomal setae present, following pseudovibrissal seta.

Thorax: Figures 112–114. Acrostichal setae in 1 

row; scutellum very weakly raised (making almost no angle 

with scutum), approximately triangular, apex relatively 

pointed, disk of scutellum flat or slightly convex; basal scutellar seta less than 1/2 length of posterior seta. Hind femur 

of male with a basoventral lobe followed by a shallow concavity, lobe roughly ovoid, bearing about 6 apical marginal 

setae and proceeded by a few yellow setae.

Male abdomen and terminalia: Figure 115. Sternite 

3 with posterior margin and medial portion only weakly 

sclerotized, posterior margin with a strong concavity, sternite appearing medially divided; sternite 4 modified, posterior margin with a strong concavity with a medial weakly 

sclerotized area; sternite 5 with a central, more membranous 

area, posterior margin with a strong concavity. Surstylus at 

most 3 times the width of posterior process of gonopod; 

lateral margins of surstylar projection approximately parallel, slightly convergent toward apex, with a subapical constriction; apex of surstylus roughly round; gonopod as in 

Figure115.

Measurements: Body length 2.3–2.6 mm. Wing 

length 2.3–2.6 mm, wing width 3.5–4.0 mm.

Type Material. The holotype male is labeled 

“ECU[ADOR]:Napo, Jatun Sacha Res., 6 km E Misahuallí, 

450 m, 1°4¢S, 77°37¢W, on foliage, 30 Apr–8 May 2002, 

M. Buck, debu00186183/Holotype ♂ Aulacigaster condylura Rung & Mathis UGE [red].” The holotype is double 

mounted (glued to a point), is in excellent condition, and is 

deposited in the UGE. Paratypes are as follows: Same label 

data as the holotype (31♂, 12♀; UGE); same locality and 

collector as the holotype, but date of 4–7 May 2002 (22♂, 

28♀; UGE).

Type Locality. Ecuador. Napo: Masahuallí (6 

km E; 01°04¢S, 77°37¢W; 450 m).

Distribution. (Map 8) Neotropical. Ecuador 

(Napo).

Biology. The type series was collected “on 

foliage.”

Etymology. The specific epithet, condylura, is 

of Latin derivation and means “knob, prominence or enlarged end”, in reference to the lobe on the hind femur of 

the male.

Remarks. Aulacigaster condylura is very similar to A. curvata and A. gaimarii. Even though males of 

FIGURE 111. Illustrations of Aulacigaster appendiculata, new species (male) (the minuta group, Neotropical Region): epandrium and 

gonopod, lateral aspect.
5 6 • smithsonian contributions to zoology

FIGUREs 112–114. Digital photographs of Aulacigaster condylura, new species (male) (the 

minuta group, Neotropical Region). (112) enlargement of anterior portion of hind femur, 

ventral aspect; (113) hind femur, ventral aspect; 

(114) right wing, dorsal aspect. Not all to the 

same scale.

MAP 8. Distribution of the minuta species group of Aulacigaster (Neotropical Region).
number 633 • 5 7

these three species can be easily separated from each other 

with the characters given in the key and diagnosis, female 

identification is more difficult. Females of A. condylura can 

be for most part separated from females of the other two 

species by the less extensive dark coloration of the hind 

femur, and the flat scutellum. If the know distribution of 

these three species reflects their real distribution, they can 

also be separated based on country of origin. 

Aulacigaster costaricana, new species

Figures 116–120, Map 8

Diagnosis. This species is distinguished from 

congeners by the following combination of characters: 

Body elongate, scutal ratio 1.5–1.75; thoracic pleura uniformly black; hind femur of male with a small, inconspicuous, approximately cylindrical lobe.

Description. Coloration and vestiture: Mesofrons mostly polished, or with a faint medial microtomentose stripe from base of antenna to level of anterior 

fronto-orbital seta; antenna pale yellow to yellowish, 

infuscate along dorsal margin; face brownish; palpus white, 

or yellowish. Scutum mostly devoid of microtomentum; 

scutellum dull posteriorly, anterior portion shiny; postpronotum concolorous with mesonotum, mostly polished; 

halter mostly yellowish. Fore coxa yellow, fore femur yellowish, fore tibia yellowish, fore tarsus mostly yellowish; 

mid coxa yellow, mid femur yellowish, mid tibia yellowish 

(mid femur and mid tibia infuscate apically in some specimens), mid tarsus mostly yellowish; hind coxa yellow, hind 

femur mostly yellowish, apex brown infuscate, hind tibia 

brown to dark brown, yellowish at basal fifth or brown 

to black on central portion, base and apex yellowish, hind 

tarsus yellowish. Male abdomen subshiny to dull; female 

abdomen with dense microtomentum over syntergite 1+2, 

and anterior margin of remaining segments.

Morphology: 

Head: Medial vertical seta 1/2 the length of lateral 

vertical seta; peristomal area with about 3–5 setae following pseudovibrissal seta.

Thorax: Figure 116. Acrostichal setae in 1 row; 

scutellum slightly raised (angle with scutum approximately 

135°), approximately triangular, apex relatively pointed, 

disk of scutellum flat; basal scutellar seta less than 1/2 

length of posterior seta. Hind femur of male with a small, 

inconspicuous, approximately cylindrical lobe; hind tibia 

with a row of tiny ventral spines.

Male abdomen and terminalia: Figures 117–120. 

Sternite 4 modified, posterior margin with a medial weakly 

sclerotized area; sternite 5 with a central, more membranous area, posterior margin appearing receded. Surstylus 

at most 3 times the width of posterior process of gonopod; lateral margins of surstylar projection convergent toward apex, with a subapical constriction; apex of surstylus 

roughly round; gonopod as in Figure 120.

Measurements: Body length 3.1–3.5 mm. Wing 

length 2.4–2.9 mm, wing width 0.6–0.9 mm.

Type Material. The holotype male is labeled 

“COSTA RICA: Heredia[,] Pr.: La Selva Biol.[ogical] Sta.

[tion] 3 Km S Pto. Viejo 10°26¢N 84°01¢W/18. vi 1991 [18 

Jun 1991] H. A. Hespenheide/HOLOTYPE ♂ Aulacigaster 

costaricana Rung & Mathis USNM [red].” The holotype is 

double mounted (minuten in a block of plastic), is in excellent condition, and is deposited in USNM. Paratypes are 

as follows: Same label data as the holotype (2♂; UCLA, 

USNM); same locality data but different collector (P. Hanson) and date (Jan 1993) (1♂; USNM). COSTA RICA. 

Cartago: La Suiza (09°51.5¢N, 83°37.5¢W), P. Schild (A. L. 

Melander Collection) (3♂; USNM).

Type Locality. Costa Rica. Heredia: La 

Selva Biological Station (3 km S Puerto Viejo; 10°26¢N, 

84°01¢W).

FIGURE 115. Illustrations of Aulacigaster condylura, new species 

(male) (the minuta group, Neotropical Region): epandrium and 

gonopod, lateral aspect.
5 8 • smithsonian contributions to zoology

Other Specimens Examined. COSTA 

RICA. Limón: Amubri (09°31¢N, 82°58¢W), Talamanca, 

26 Jul 1975, W. J. Hanson (1♂, 1♀; USU).

Distribution. (Map 8) Neotropical. Costa 

Rica (Cartago, Heredia, Limón).

Etymology. The specific epithet, costaricana,

refers to the country, Costa Rica, where the type series was 

collected.

Remarks. See “Remarks” under A. appendiculata.

Aulacigaster curvata, new species

Figures 121–123, Map 8

Diagnosis. This species is distinguished from 

congeners by the following combination of characters: 

Body elongate, scutal ratio 1.5–1.75; pleura uniformly 

black; wing infuscate with brown; lobe on hind femur of 

male followed by a deep concavity.

Description. Coloration and vestiture: Mesofrons mostly polished, antenna pale yellow to yellowish, 

infuscate along dorsal margin in some specimens; face whitish; palpus white. Scutum mostly devoid of microtomentum; 

scutellum mostly dull, or dull posteriorly, anterior portion 

shiny; postpronotum concolorous with mesonotum, mostly 

polished; halter mostly white. Wing mostly brown, infuscate. Fore coxa yellow, fore femur yellowish, infuscate with 

brown apically in some females, fore tibia yellowish, fore 

tarsus mostly yellowish; mid coxa yellow, mid femur yellowish or dark brown to black dorsally on apical 2/3, yellowish basally, mid tibia yellowish (partly to completely 

infuscate with brown in some male specimens), mid tarsus 

mostly yellowish; hind coxa yellow, hind femur completely 

brown, or mostly brown, yellow on basal 1/5 to basal 1/4, 

FIGURES 116–119. Digital photographs of Aulacigaster costaricana, new species (male) (the minuta group, 

Neotropical Region). (116) right wing, dorsal aspect; (117) subepandrial sclerite, ventral aspect; (118) epandrium and gonopod, ventral aspect; (119) abdomen, dorsal aspect. Not all to the same scale.
number 633 • 5 9

hind tibia brown to black on central portion, base and apex 

yellowish, hind tarsus yellowish. Male abdomen subshiny 

to dull, pregenital segment polished posteriorly, dull anteriorly; female abdomen with dense microtomentum over 

syntergite 1+2, and anterior margin of remaining segments.

Morphology: 

Head: Medial vertical seta 1/2 length of lateral vertical seta; 1–2 peristomal seta present, following pseudovibrissal seta.

Thorax: Figures 121–122. Acrostichal setae in 1 

row; scutellum slightly raised (angle with scutum approximately 135°), approximately triangular, apex relatively 

pointed, disk of scutellum flat or slightly convex; basal scutellar seta less than 1/2 length of posterior seta. Hind femur 

of male with a basoventral lobe followed by a strong concavity, lobe roughly ovoid, bearing about 6 apical marginal 

setae and proceeded by a few yellow setae.

Male abdomen and terminalia: Figure 123. Sternite 

3 with posterior margin and medial portion only weakly 

sclerotized, posterior margin with a strong concavity, sternite appearing medially divided; sternite 4 modified, posterior margin with a strong concavity; sternite 5 with a central, 

more membranous area, posterior margin with a strong 

concavity. Surstylus at least 3 times the width of posterior 

process of gonopod; lateral margins of surstylar projection 

convergent toward apex, with a subapical constriction; apex 

of surstylus roughly round; gonopod as in Figure 123.

Measurements: Body length 3–3.4 mm. Wing 

length 2–2.4 mm, wing width 3.5–3.9 mm.

Type Material. The holotype male is labeled 

“ECU[ADOR]:Napo, Jatun Sacha Res., 6 km E Misahuallí, 

450 m, 1°4¢S, 77°37¢W, on foliage, 30 Apr–8 May 2002, 

M. Buck, debu00186306/Holotype ♂ Aulacigaster curvata Rung & Mathis UGE [red].” The holotype is double 

mounted (glued to a point), is in excellent condition, and is 

deposited in the UGE. Paratypes are as follows: Same label 

data as the holotype (9♂, 20♀; UGE, USNM); same locality and collector as the holotype, but date of 4–7 May 2002 

(13♂, 14♀; UGE, USNM); same locality but collected by O. 

Lonsdale, 2–7 May 2002 (1♂, 2♀; UGE); same locality, 350 

m, Feb 1983, M.J. Sharkey (1♂; UGE).

Type Locality. Ecuador. Napo: Masahuallí 

(6 km E; 01°04¢S, 77°37¢W; 450 m).

Other Specimens Examined. ECUADOR. Pichincha: Maquipucuna Biological Research Station 

FIGURE 120. Illustrations of Aulacigaster costaricana, new species (male) (the minuta group, Neotropical Region): epandrium and 

gonopod, lateral aspect.

FIGURES 121–122. Digital photographs of Aulacigaster curvata, 

new species (male) (the minuta group, Neotropical Region). (121) 

hind femur, ventral aspect; (122) enlargement of anterior portion of 

hind femur, ventral aspect. Not to the same scale.
6 0 • smithsonian contributions to zoology

(00°07¢N, 78°38¢W; river trail; Heliconia leaves; 1200 m), 26 

Apr–29 Oct 1999, 2002, M Buck, S. A. Marshall (3♀; UGE).

PERU. Madre de Dios: Los Amigos Biological Station (12°34¢S, 70°06.1¢W), 2–14 Jun 2006, S. Paiero, 

J. Klymko (3♂, 1♀; UGE).

Distribution. (Map 8) Neotropical: Ecuador 

(Napo, Pichincha), Peru (Madre de Dios).

Biology. The type series was collected “on 

foliage.”

Etymology. The specific epithet, curvata, is of 

Latin derivation and means “curved, bent,” in reference to 

the strong curvature on the male hind femur following the 

lobe.

Remarks. See above under A. condylura.

Aulacigaster gaimarii, new species

Figures 124–127, Map 8

Diagnosis. This species is distinguished from 

congeners by the following combination of characters: 

Body elongate, scutal ratio 1.5–1.75; thoracic pleura uniformly black; lobe on hind femur of male approximately 

foot-shaped, with apical and posterior marginal setae, not 

followed by a concavity.

Description. Coloration and vestiture: Mesofrons mostly polished; antenna pale yellow to yellowish, infuscate along dorsal margin in some specimens; face 

pale yellow, often with a brownish area posteriorly; palpus 

white. Scutum mostly devoid of microtomentum; scutellum dull posteriorly, anterior portion shiny; postpronotum 

concolorous with mesonotum, mostly polished; halter 

mostly white. Wing mostly brown, infuscate. Fore coxa yellow, fore femur yellowish, fore tibia yellowish, fore tarsus 

mostly yellowish; mid coxa yellow, mid femur yellowish or 

dark brown to black apically, yellowish basally, mid tibia 

yellowish (partly to completely infuscate with brown in 

some female specimens), mid tarsus mostly yellowish; hind 

coxa yellow, hind femur completely to mostly brown, yellow on basal 1/4, hind tibia brown to black on central portion, base and apex yellowish, hind tarsus yellowish. Male 

abdomen subshiny to dull, pregenital segment polished 

posteriorly, dull anteriorly; female abdomen with dense microtomentum over syntergite 1+2, and anterior margin of 

remaining segments.

Morphology: 

Head: medial vertical seta 1/2 length of lateral vertical seta; 1 peristomal seta present, following pseudovibrissal seta.

Thorax: Figures 124–125. Acrostichal setae in 1 

row; scutellum slightly raised (angle with scutum approximately 135°), approximately triangular, apex relatively 

pointed, disk of scutellum flat or slightly convex; basal 

scutellar seta less than 1/2 length of posterior seta. Hind 

femur of male with a basoventral lobe not followed by a 

distinct concavity, lobe roughly foot-shaped, bearing apical and posterior marginal setae and proceeded by a few 

yellow setae.

Male abdomen and terminalia: Figures 126–127. 

Sternite 3 with posterior margin and medial portion only 

weakly sclerotized, posterior margin receded, sternite appearing medially divided; sternite 4 modified, posterior 

margin with a medial weakly sclerotized area; sternite 5 

with a central, more membranous area, posterior margin 

appearing receded. Surstylus at most 2 times the width of 

posterior process of gonopod; lateral margins of surstylar 

projection convergent toward apex, with a subapical constriction; apex of surstylus roughly round; gonopod as in 

Figure 126.

Measurements: Body length 2.7–3.1 mm. Wing 

length 2.4–2.7 mm, wing width 0.6–0.7 mm.

FIGURE 123. Illustrations of Aulacigaster curvata, new species 

(male) (the minuta group, Neotropical Region): epandrium and 

gonopod, lateral aspect.
number 633 • 6 1

Type Material. The holotype male is labeled 

“BOLIVIA: Depto. La Paz[,] 5 km W Mapiri, Arroyo 

Tuhiri[,] 750 m, 15°17.8¢S 68°15.6¢W[,] 19-iii-2001 [19 

Mar 2001], S. D. Gaimari/HOLOTYPE ♂ Aulacigaster 

gaimarii Rung & Mathis USNM [red].” The holotype is 

double mounted (minuten in a block of plastic), is in excellent condition, and is deposited in the USNM. Paratypes are 

as follows: Same label data as the holotype (5♂, USNM); 

same locality and collector data as the holotype but with 

dates of 17 Mar 2001 (1♀; USNM), 18 Mar 2001 (6♂, 8♀; 

USNM); same locality, 16 Mar 2001, A. Freidberg (15♂, 

7♀; USNM).

Type Locality. Bolivia. La Paz: Mapiri (5 km 

W; Arroyo Tuhiri; 15°17.8¢S 68°15.6¢W).

Other Specimens Examined. BOLIVIA. 

La Paz: Arroyo Tuhiri, W. Mapiri (15°17.5¢S 68°15.5¢W), 

10 Apr 2001, S. A. Marshall (5♂, 3♀; UGE).

Distribution. (Map 8) Neotropical. Bolivia 

(La Paz).

Biology. The type series was collected on 

leaves of plants belonging to the family Heliconiacea (S. D. 

Gaimari, personal communication).

Etymology. The specific epithet, gaimarii, is a 

genitive patronym to honor and recognize Dr. Stephen D. 

Gaimari, who collected this species and many other interesting Acalyptrate Diptera in Bolivia.

Remarks. See above under A. condylura.

Aulacigaster minuta (Hennig)

Figures 128–129, Map 8

Schizochroa minuta Hennig, 1969: 609, 605 [figure of wing].—Papavero, 

1967:91 [Neotropical catalog].

Aulacigaster minuta.—D. K. McAlpine, 1983:58 [generic combination].

Diagnosis. This species is distinguished from 

congeners by the following combination of characters: 

FIGURES 124–125. Digital photographs of Aulacigaster gaimarii, new species (the minuta group, 

Neotropical Region). (124) right hind femur of 

male, medial aspect; (125) right wing, dorsal aspect. 

Not to the same scale.

FIGURES 126–127. Illustrations of Aulacigaster gaimarii, new species (male) (the minuta group, Neotropical Region). (126) gonopod, 

lateral aspect; (127) epandrium, lateral aspect.
6 2 • smithsonian contributions to zoology

Body elongate, scutal ratio 1.50–1.75; hind femur mostly 

dark brown, yellowish at basal 1/5–1/4; lobe on hind femur 

shaped as an inverted “M.”

Description. Coloration and vestiture: Mesofrons mostly polished, or with a faint medial microtomentose stripe from base of antenna to level of anterior 

fronto-orbital seta (females without stripe); antenna pale 

yellow to yellowish, in some specimens infuscate along 

dorsal margin; face yellowish to light brown; palpus white. 

Scutum mostly devoid of microtomentum; scutellum dull 

posteriorly, anterior portion shiny (dull along apical margin); postpronotum concolorous with mesonotum, mostly 

polished; halter mostly white, or mostly brown to dark 

brown. Fore coxa yellow, fore femur yellowish, fore tibia 

yellowish, fore tarsus mostly yellowish; mid coxa yellow, 

mid femur yellowish or dark brown to black apically, yellowish basally, mid tibia yellowish or brownish, mid tarsus mostly yellowish; hind coxa yellow, hind femur mostly 

dark brown, yellowish at basal 1/5–1/4, hind tibia yellow 

basally, dark apically or brown to black on central portion, 

base and apex yellowish, hind tarsus yellowish. Male abdomen subshiny to dull (mostly on lateral margin of tergites); 

female abdomen with dense microtomentum over lateral 

margins of syntergite 1+2, remaining segments shiny.

Morphology: 

Head: Medial vertical seta 1/2 length of lateral vertical seta; peristomal area with 2–4 setae following pseudovibrissal seta.

Thorax: Figure 128. Acrostichal setae in 1 row; 

scutellum slightly raised (angle with scutum approximately 

135°), approximately triangular, apex relatively pointed, 

disk of scutellum flat; basal scutellar seta less than 1/2 

length of posterior seta. Hind femur modified, with 2 ventral rows of spines, each spine bearing an apical seta; hind 

femur of male with a basal lobe not followed by a posterior 

concavity, lobe approximately shaped as an inverted “M”, 

with one seta on each pointed projection; setae on lobe yellowish to brown, lobe preceded by a 2–3 weak setae.

Male abdomen and terminalia: Figure 129. Surstylus at most 3 times the width of posterior process of 

gonopod; lateral margins of surstylar projection convergent 

toward apex, with a subapical constriction; apex of surstylus roughly round; gonopod as in Figure129.

Measurements: Body length 2.7–3 mm. Wing 

length 2.2–2.3 mm. Wing width 0.5–0.6 mm.

Type Material. The holotype female is labeled 

“Coca, Napo R[iver]., Napo, ECUADOR, 25.30.IV.65, 

250 m., L. Peña/Schizochroa minuta Hg. HOLOTYPUS 

wing on slide [red]/HOLOTYPE Schizochroa minuta Hennig CNC No. 9930 [red].” The holotype is glued directly 

to the pin, is in good condition (left wing is missing), and is 

deposited in the CNC (9930).

Type Locality. Ecuador. Orellana: Napo River, 

Puerto Francisco de Orellana (Coca; 00°28¢S, 76°58¢W).

Other Specimens Examined. PANAMA. 

Canal Zone: Corozal (08°59¢N, 79°34¢W), 19 Jan 1929, 

C. H. Curran (1♂; AMNH); Erwin Island, 17 Jul 1923, 

R. C. Shannon (1♂; USNM); Madden Forest (09°06¢N, 

79°37¢W), 7–16 Mar 1961, S. B. Pipkin (3♂, 5♀; USNM); 

Paraíso (09°01.5¢N, 79°37.5¢W), 28 Jan–11 Feb 1911, 

A. Busck (2♂, 1♀; USNM); Piña Area (09°15.4¢N, 

79°57.9¢W), 17 May 1960, W. J. Hanson (1♂; USU); Pipeline road (09°08.6¢N, 79°43.6¢W), 22 Mar 1982, W. J. 

Hanson (1♀; USU); Barro Colorado Island (09°09.3¢N, 

79°50.8¢W), 18 Jul 1924, N. Banks (2♂, 3♀; AMNH). Darien: El Real (08°08¢N, 77°43¢W; ex. Heliconia curhispatha), 28 Nov 1963, S. B. Pipkin (1♂; USNM).

FIGURES 128–129. Illustrations of Aulacigaster minuta (Hennig) 

(male) (the minuta group, Neotropical Region). (128) basal portion 

of the left hind femur, medial aspect; (129) epandrium and gonopod, 

lateral aspect. Not to the same scale.
number 633 • 6 3

PERU. Loreto: Iquitos (03°44.9¢S, 73°14.8¢W), Mar–

Apr 1931, R. C. Shannon (4♀; USNM).

Distribution. (Map 8) Neotropical. Ecuador 

(Orellana), Panama (Canal Zone, Darien), Peru (Loreto).

Remarks. This species is similar to A. nigritarsa

but is distinguished from that species by its completely yellow antenna, the coloration of the hind femur (yellow on 

basal 1/5) and hind tarsus (completely yellow), and by the 

shape of the lobe on the hind femur of the male (shaped as 

an inverted “M”, and bearing apical setae).

Aulacigaster nigritarsa, new species

Figures 130–131, Map 8

Diagnosis. This species is distinguished from 

congeners by the following combination of characters: 

Body elongate, scutal ratio 1.50–1.75; hind tibia entirely 

brown to dark brown.

Description. Coloration and vestiture: Mesofrons with a faint medial microtomentose stripe from base 

of antenna to level of anterior fronto-orbital seta; antenna 

pale yellow to yellowish, not infuscate dorsally; face brownish; palpus white. Scutum mostly devoid of microtomentum; scutellum dull microtomentose, or dull posteriorly, 

anterior portion shiny; postpronotum concolorous with 

mesonotum, mostly polished; halter mostly yellowish. Fore 

coxa yellow, fore femur yellowish, fore tibia yellowish, fore 

tarsus mostly yellowish; mid coxa yellow, mid femur yellowish, mid tibia brownish or with basal portion brown 

to dark brown, apical portion yellowish, mid tarsus mostly 

yellowish; hind coxa yellow, hind femur brown, hind tibia 

brown to dark brown, hind tarsus with basal tarsomere 

brown, apical tarsomere yellowish. Male abdomen subshiny to dull, pregenital segment mostly subshiny.

Morphology: 

Head: Medial vertical seta thin, about 1/2 of lateral 

vertical seta; peristomal area with about 2 setae following 

pseudovibrissal seta.

Thorax: Figure 131. Acrostichal setae in 1 row; 

scutellum slightly raised (angle with scutum approximately 

135°), approximately triangular, apex relatively pointed, 

disk of scutellum flat; basal scutellar seta less than 1/2 length 

of posterior seta. Hind femur modified, with 2 ventral rows 

of spines, each spine bearing an apical seta; hind femur of 

male with a basal lobe not followed by a posterior concavity, lobe approximately cylindrical approximately triangular, bearing an apical seta; lobe preceded by 5–6 weak setae

Male abdomen and terminalia: Figure 130. Sternite 

5 with a central, more membranous area, posterior margin 

appearing receded surstylus at most 3 times the width of 

posterior process of gonopod; lateral margins of surstylar 

projection convergent toward apex, without a conspicuous 

subapical constriction; apex of surstylus roughly round.

Measurements: Body length 3.2–3.6 mm. Wing 

length 2.4–2.6 mm, wing width 1.6–1.7 mm.

FIGURES 130–131. Illustrations of Aulacigaster nigritarsa, new 

species (male) (the minuta group, Neotropical Region). (130) epandrium, lateral aspect; (131) left hind femur, medial aspect. Not to 

the same scale.
6 4 • smithsonian contributions to zoology

Type Material. The holotype male is labeled 

“Higuito[,] San Mateo[,]C[osta]R[ica]/PabloSchild Coll/

HOLOTYPE ♂ Aulacigaster nigritarsa Rung & Mathis 

USNM [red].” The holotype is double mounted (minuten 

in a narrow, rectangular block of pith), is in fair condition 

(setae of head and mesonotum broken, right wing slightly 

folded, right midleg is missing), and is deposited in the 

USNM. Paratypes are as follows: Same label data as the 

holotype (4♂, 4♀; USNM).

Type Locality. Costa Rica. Alajuela: Higuito, 

San Mateo (09°56¢N, 84°31¢W).

Other Specimens Examined. COSTA 

RICA. Puntarenas: Cerro de Oro (08°39¢N, 83°29¢W; 200 

m), B. Gamboa, 5–9 May 1995 (1♀; USNM); Peninsula de 

Osa, Rincón (08°42¢N, 83°29¢W; Malaise trap), Oct–Dec 

1990, P. Hanson (1♀; USNM); Peninsula de Osa, Rincón 

(08°42.2¢N, 83°30.8¢W; pan traps, grasses under canopy), 

10–11 Aug 2001, K. N. Barger (1♀; UGE). San José: San 

Carlos, Riosparaíso Reserve, Pecarí Station (16 km NE 

Quepos; 09°33¢532N, 84°7¢322W; 400m), 15 Feb 2003, 

S. A. Marshall (1♀; UGE).

Distribution. (Map 8) Neotropical: Costa 

Rica (Alajuela, Puntarenas).

Etymology. The specific epithet, nigritarsa, is 

of Latin derivation and refers to the black coloration of the 

tarsus.

Remarks. See “Remarks” under A. minuta.

The plesiomorphica Group

Figures 132–160, 213–214, Maps 9–13

Diagnosis. The plesiomorphica group is distinguished from other species groups by the following combination of characters: Stout, medium-sized flies, body length 

2.4–3.5 mm. Coloration and vestiture: Frons with anterior 

1/3–1/2 yellowish in ground color; anterior portion of frons 

densely microtomentose, microtomentum forming a with 

a wide, lunate, silvery microtomentose stripe; facial band 

often present. Anepisternum mostly dull microtomentose. 

Abdomen in a few cases with a large, white to yellowish 

region on syntergite 1+2.

Morphology: 

Head: Head higher than long (head ratio less than 

0.9); face strongly convex on ventral 2/3, easily visible in 

lateral view; ocellar seta minute; fronto-orbital setae with 

posterior seta slightly internal to and almost transversely 

aligned with anterior seta.

Thorax: Subcosta partially fused with vein R1

 apically but terminating on costa.

Male abdomen and terminalia: Surstylus a short, 

posteroventral lobe in lateral view; cerci often partially 

fused; subepandrial sclerite typically forming a single, platelike structure; gonopods often with 2 posterior processes; 

one bearing 4–5 strong, stout setae.

Distribution. Neotropical: Belize, Brazil, 

Costa Rica, Ecuador, Guyana, Panama, Peru, Venezuela.

Biology. There is sparse biological information 

on species of this group. Exemplars collected by us in the 

rainforest were collected on tree trunks and aerial portions 

of large Ceiba trees with no visible, wound-exuding sap 

fluxes.

Discussion. Species of the plesiomorphica

group are distinguished by the presence/absence of a facial 

band, the shape of the face, and coloration of the legs. In 

some cases, species can only be identified with certainty by 

the shape of the surstylus and other structural characters of 

the male, such as the shape of sternite 5, presence/absence 

of setae on the hind trochanter and hind tibia.

The plesiomorphica group includes nine species, and 

of these only one was previously described. The group is 

exclusively Neotropical. 

Aulacigaster albifacies, new species

Figures 132–133, Map 9

Diagnosis. This species is distinguished from 

congeners by the following combination of characters: 

Body robust, scutal ratio 1.1–1.35; katepisternum subshiny 

to dull, microtomentose; face, in lateral view, barely visible, 

not appearing bulbous; scutellum triangular, apex strongly 

pointed.

Description. Coloration and vestiture: Vertex mostly polished, with a densely microtomentose stripe 

posterior to ocellar tubercle, with a shiny spot extended 

from ocellus to 2/3 distance to eye margin; ocellar tubercle dull microtomentose; frontal white-microtomentose 

stripe bordered posteriorly by a conspicuous velvety-black 

stripe; antenna pale yellow to yellowish, infuscate along 

dorsal margin; face white to yellow; facial band present, distinct, gray or golden microtomentose over brown 

mark, on dorsal portion of face; palpus white. Scutum 

mostly completely microtomentose (specimens from Venezuela) or polished on anteriormost portion, opposite to 

postcranium (specimens from Guyana and Peru); scutellum dull microtomentose; postpronotum concolorous 

with mesonotum, scarcely microtomentose, subshiny; 

anepisternum mostly dull (dorsoposterior portion more 

shiny in some specimens); katepisternum subshiny to dull; 
number 633 • 6 5

halter mostly white. Fore coxa brown to black, fore femur 

brown, fore tibia yellowish or brown, fore tarsus yellowish, apical and sometimes subapical tarsomeres brown 

to black; mid coxa brown to black, mid femur brown or 

brown to black, apex yellowish, mid tibia yellowish or 

brownish, mid tarsus mostly or entirely yellowish, apical 

and subapical tarsomere brownish to dark brown; hind 

coxa brown, hind femur brown or brown to black, apex 

yellowish, hind tibia yellowish-brownish or brown to dark 

brown, hind tarsus yellowish, apical tarsomere brown to 

black. Abdomen completely dark brown to black or with 

a large, white to yellowish region on syntergite 1+2, male 

abdomen bright shiny, pregenital segment mostly subshiny; female abdomen with dense microtomentum medially, lateral margins of tergites nearly polished.

Morphology: 

Head: Head higher than long (head ratio less than 

0.9); face at level of pseudovibrissal seta approximately the 

same width as the 1st flagellomere, not appearing bulbous; 

medial vertical seta 1/2–3/4 length of lateral vertical seta; 

1 strong peristomal seta present, following pseudovibrissal 

seta.

Thorax: Acrostichal setae in 2 rows; scutellum 

strongly raised (angle with scutum less than 135°), triangular, apex strongly pointed, disk of scutellum flat; basal 

scutellar seta 1/2 length of posterior seta.

Male abdomen and terminalia: Figures 132–133. 

Sternite 5 with a central, more membranous area, posterior 

margin appearing receded, and bearing a tongue-like medial projection. Surstylus approximately rectangular, from 

lateral view “boat-shaped” and bearing one apical and one 

subapical setae; anterior process of the gonopod almost as 

long and as large as the surstylus; gonopod as in Figure 

132; cerci partially fused, each cercus bearing 1 very long, 

ventral seta and 1–3 longer setulae.

Measurements: Body length 2.7–3.0 mm. Wing 

length 2.7–2.9 mm, wing width 0.8–0.9 mm.

Type Material. The holotype male is labeled “ECUADOR. Prt.[Puerto]. Orellana: RioTiputini 

(0°38.2¢S, 76°8.9¢W)[,] 12–26 Aug 1999, W. N. Mathis, 

A. Baptista, M. Kotrba/USNM ENT 00090363 [plastic bar 

code label]/HOLOTYPE ♂ Aulacigaster albifacies Rung 

& Mathis USNM [red].” The holotype is double mounted 

(minuten in a block of plastic), is in excellent condition, and 

is deposited in the USNM. Paratypes are as follows: Same 

label data as the holotype (7♂; EPNE, USNM).

Type Locality. Ecuador. Orellana: Rio Tiputini (00°38.2¢S, 76°08.9¢W).

Other Specimens Examined. GUYANA. 

Moco Moco River (30 km E Lethem in Kanuku Mountains; 

03°18.2¢N, 59°39.0¢W), 3–29 Apr 1994, 1995, W. N. 

Mathis (5♂, 1♀; USNM).

PERU. Madre de Dios: Manu, Rio Manu, Pakitza, 

Aguajal (11°58.2¢S, 71°17¢W), 19 Sep 1988, A. Freidberg, 

(2♂, 1♀; USNM).

VENEZUELA. Táchira: Pregonero, Presa Las Cuevas 

(8°01.5¢N, 71°45.8¢W; 650 m; rain forest; Malaise trap), 

9–31 Jul 1989, S. and J. Peck (1♂, 1♀; UGE).

Distribution. (Map 9) Neotropical: Ecuador (Orellana), Guyana, Peru (Madre de Dios), Venezuela 

(Táchira).

Biology. This species was collected while 

sweeping trunks of Ceiba trees (Bombacaceae: Ceiba sp.) 

in the rainforest of Ecuador. The tree trunk had no visible 

wound exuding sap flux.

Etymology. The specific epithet, albifacies, is 

of Latin derivation and refers to the white coloration of the 

facial band.

FIGURES 132–133. Illustrations of Aulacigaster albifacies, new species (male) (the plesiomorphica group, Neotropical Region). (132) 

epandrium and gonopod, lateral aspect; (133) sternite 5, ventral aspect. Not all to the same scale.
6 6 • smithsonian contributions to zoology

Remarks. Specimens collected in Venezuela have the scutum mostly completely microtomentose, 

whereas specimens collected in Guyana and Peru have a 

polished region on the anteriormost portion of scutum, opposite to postcranium. We did not observe any other difference that would justify description of another species. This 

species shares with A. trifasciata the presence of a tonguelike medial projection on the posterior margin of the male 

fifth sternite and a predominantly polished vertex. The surstylus is also very similar in both species. Specimens of A. 

albifacies are stouter and darker than those of A. trifasciata, 

have the femora and tibiae mostly black, and a microtomentose ocellar tubercle. Specimens of A. trifasciata are 

smaller, have the femora and tibiae mostly yellow, and have 

a polished ocellar tubercle.

Aulacigaster atriseta, new species

Figure 134–136, Map 10

Diagnosis. This species is distinguished from 

congeners by the following combination of characters: 

Frons with anterior 1/3–1/2 yellowish in ground color; hind 

tibia with strong, dark, apicodorsal setae; hind trochanter 

of male without a ventral tuff of setae.

Description. Coloration and vestiture: Vertex mostly densely microtomentose, with a shiny spot extended from ocellus to 2/3 distance to eye margin to with 

a shiny spot extended from ocellus at most 1/2 way to 

eye margin; ocellar tubercle dull microtomentose; frontal 

white-microtomentose stripe bordered posteriorly by a 

MAP 9. Distribution of Aulacigaster albifacies and A. bifasciata (the plesiomorphica group, Neotropical Region).
number 633 • 6 7

conspicuous velvety-black stripe; antenna pale yellow to 

yellowish, infuscate along dorsal margin; face yellow anteriorly, brown to black on posterior portion; facial band 

present (region of facial band “wrinkled and matt), delimited as a transversely wrinkled band on dorsal portion of 

face, mostly lacking microtomentum; palpus white. Scutellum dull microtomentose; postpronotum concolorous with 

mesonotum, dull microtomentose; anepisternum mostly 

dull; katepisternum subshiny to dull; halter mostly white. 

Fore coxa brown to black, fore femur brown to black, 

apex yellowish, fore tibia yellowish (brown infuscate basally), fore tarsus mostly yellowish to yellowish, apical and 

sometimes subapical tarsomeres brown to black; mid coxa 

brown to black, mid femur brown to black, apex yellowish, 

mid tibia yellowish, mid tarsus mostly yellowish; hind coxa 

brown, hind femur brown to black, apex yellowish, hind 

tibia yellowish or dark basally, yellow apically, hind tarsus 

yellowish. Male abdomen bright shiny, pregenital segment 

polished posteriorly, dull anteriorly; female abdomen with 

dense microtomentum medially, lateral margins of tergites 

nearly polished.

Morphology: 

Head: Figures 134–135. Head round, about as 

high as long (head ratio 0.9–1.1), or higher than long (head 

ratio less than 0.9); face at level of pseudovibrissal seta approximately the same width as the 1st flagellomere, not appearing “bulbous”; medial vertical seta 1/2–3/4 length of 

lateral vertical seta; 1 strong peristomal seta present, following pseudovibrissal seta.

Thorax: Figure 134. Acrostichal setae in 1 row; 

scutellum strongly raised (angle with scutum less than 

135°), approximately triangular, apex relatively pointed, 

disk of scutellum slightly convex; basal scutellar seta 1/2 

length of posterior seta. Hind tibia with strong, dark, apicodorsal setae.

Male abdomen and terminalia: Figure 136. Surstylus not well differentiated from folded portion of epandrium, difficult to discern in lateral view; gonopod as in 

Figure 136; cerci bearing 1 very long, ventral seta and 1–3 

longer setulae.

Measurements: Body length 2.9 mm, wing length 

2.9 mm, wing width 0.9 mm.

Type Material. The holotype male is labeled 

“PERU. Madre de Dios: [Rio] Manu, Erika (near Salvacion), 550 m, 5–6 Sept1988,W. N. Mathis/HOLOTYPE ♂

Aulacigaster atriseta Rung & Mathis USNM [red].” The 

holotype is double mounted (minuten in a block of plastic), is in good condition (ventral right fronto-orbital seta 

broken), and is deposited in the USNM. A paratype is as 

follows: PERU. Madre de Dios: Manu, Rio Manu, Pakitza 

(11°56.6¢S, 71°16.9¢W; 250 m), 9–23 Sep 1988, W. N. Mathis (1♂; USNM).

Type Locality. Peru. Madre de Dios: Rio 

Manu, Erika (near Salvación; 12°53¢S, 71°12¢W).

Distribution. (Map 10) Neotropical: Peru 

(Madre de Dios).

Etymology. The specific epithet, atriseta, is 

of Latin derivation and is a combination of atra, meaning 

black, and seta and refers to the black, dorsoapical setae on 

the hindfemora.

Remarks. This species shares with A. proxima

and A. plesiomorphica the presence of black, dorsoapical 

setae on the hind femur, but it can be easily distinguished 

from these two species by the absence of well-developed 

setae on the hind trochanter of the male and by the absence 

of a conspicuous surstylus.

FIGURES 134–135. Digital photographs of Aulacigaster atriseta, 

new species (male) (the plesiomorphica group, Neotropical Region). 

(134) head and mesonotum of male, lateral aspect; (135) head, frontal aspect. Not all to the same scale.
6 8 • smithsonian contributions to zoology

Aulacigaster belize, new species

Figures 137–138, Map 10

Diagnosis. This species is distinguished from 

congeners by the following combination of characters: 

Frons with anterior 1/3–1/2 yellowish in ground color; 

face not appearing bulbous; hind tibia without apicodorsal 

setae; surstylus more or less folded below the epandrium, 

difficult to see in lateral view.

Description. Coloration and vestiture: Vertex 

mostly densely microtomentose, with a shiny spot extended 

from ocellus at most 1/2 way to eye margin or with a shiny 

spot extended from ocellus to 2/3 distance to eye margin; 

ocellar tubercle dull microtomentose; frontal white-microtomentose stripe bordered posteriorly by a conspicuous 

velvety-black stripe; antenna pale yellow to yellowish, infuscate along dorsal margin; face yellow anteriorly, brown 

to black on posterior portion; facial band present, delimited 

as a transversely wrinkled band on dorsal portion of face, 

mostly lacking microtomentum; palpus white. Scutellum 

dull microtomentose; postpronotum concolorous with mesonotum, dull microtomentose; anepisternum mostly dull; 

katepisternum subshiny to dull; halter mostly white. Fore 

coxa brown to black, fore femur brown to black, apex yellowish, fore tibia yellowish (brown infuscate basally), fore 

tarsus yellowish, apical and sometimes subapical tarsomeres brown to black; mid coxa brown to black, mid femur 

brown to black, apex yellowish, mid tibia yellowish (brown 

infuscate on basal portion), mid tarsus mostly yellowish; 

hind coxa brown, femur brown to black, apex yellowish, 

hind tibia yellowish or dark basally, yellow apically, hind 

tarsus yellowish. Male abdomen bright shiny, pregenital 

segment polished posteriorly, dull anteriorly; female abdomen with dense microtomentum medially, lateral margins 

of tergites nearly polished.

Morphology: 

Head: Head round, about as high as long (head 

ratio 0.9 to 1.1) to higher than long (head ratio less than 

0.9); face at level of pseudovibrissal seta wider than width 

of 1st flagellomere or approximately the same width as the 

1st flagellomere, not appearing bulbous; medial vertical 

seta 1/2–3/4 length of lateral vertical seta; 1 strong peristomal seta present, following pseudovibrissal seta.

Thorax: Acrostichal setae in 1 row; scutellum 

slightly raised (angle with scutum approximately 135°), 

semicircular, disk of scutellum slightly convex; basal scutellar seta 1/2 length of posterior seta.

Male abdomen and terminalia: Figures 137–138. 

Sternite 5 with a central, more membranous area, posterior 

margin appearing receded. Surstylus more or less folded 

below the epandrium, difficult to see in lateral view, approximately triangular, with apex pointing dorsally towards cerci, bearing one medial and one strong apical setae; 

gonopod as in Figure 137; cerci separate, narrow, digitiform, each cercus bearing 1 very long, ventral seta and 1–3 

longer setulae.

Measurements: Body length 2.8–3.0 mm. Wing 

length 2.5–2.7 mm, wing width 0.7–0.9 mm.

Type Material. The holotype male is labeled 

“BELIZE. Stann Creek District: Cockscomb Basin Wldlfe. 

Sanct. (16°47¢N, 88°30¢W)[,] 5–6 Apr 1993, W. Mathis/

HOLOTYPE ♂ Aulacigaster belize Rung & Mathis USNM 

[red].” The holotype is double mounted (minuten in a block 

of plastic), is in excellent condition, and is deposited in the 

USNM. Paratypes are as follows: Same label data as the 

holotype (10♂, 6♀; USNM).

Type Locality. Belize. Stann Creek District: 

Cockscomb Basin Wildlife Sanctuary (16°47¢N, 88°30¢W).

Other Specimens Examined. BRAZIL. 

Rondônia: Ariquemes (62 Km SE; 10°34¢S, 62°22¢W), 

7–20 Nov 1994, 1995, W. J. Hanson (1♂, 1♀; USU).

FIGURE 136. Illustrations of Aulacigaster atriseta, new species 

(male) (the plesiomorphica group, Neotropical Region): pregenital 

segment, epandrium and gonopod, lateral aspect.
number 633 • 6 9

COSTA RICA. Puntarenas: Peninsula de Osa, Rincón (08°42¢N, 83°29¢W; sea level; sweeping in palmmangrove forest), 24 Mar 1991, D. A. Grimaldi, J. Stark 

(3♂; AMNH).

Distribution. (Map 10) Neotropical: Brazil 

(Rondônia), Belize, Costa Rica (Puntarenas).

Etymology. The specific epithet, belize, refers 

to the country of Belize, where the type locality is located 

and is a noun in apposition.

Remarks. This species is not easily confounded 

with congeners and can be reliably separated from them 

with the characters given in the key and diagnosis. It is also 

the only species of the genus known from Belize.

Aulacigaster bifasciata, new species

Figures 139–140, Map 9

Diagnosis. This species is distinguished from 

congeners by the following combination of characters: 

Frons with anterior 1/3–1/2 yellowish in ground color; face 

bulbous; facial band absent; frons white-microtomentose 

stripe bordered posteriorly by a conspicuous velvety-black 

stripe.

Description. Coloration and vestiture: Vertex 

mostly densely microtomentose, with a shiny spot extended 

from ocellus at most 1/2 way to eye margin; ocellar tubercle 

MAP 10. Distribution of Aulacigaster atriseta, A. belize, and A. plesiomorphica (the plesiomorphica group, Neotropical Region).
7 0 • smithsonian contributions to zoology

dull microtomentose; frontal white-microtomentose stripe 

bordered posteriorly by a conspicuous velvety-black stripe; 

antenna pale yellow to yellowish, infuscate along dorsal 

margin or infuscate on dorsal half; face brownish; palpus 

white, or yellowish. Scutellum dull microtomentose; postpronotum concolorous with mesonotum, mostly polished; 

anepisternum mostly dull; katepisternum subshiny to dull; 

halter mostly white. Fore coxa brown to black, fore femur 

brown, fore tibia mostly or entirely yellowish-brownish, 

fore tarsus mostly yellowish; mid coxa brown to black, mid 

femur brown; mid tibia yellowish to brownish, mid tarsus 

mostly yellowish; hind coxa brown, hind femur brown, 

hind tibia yellowish-brownish or brown to black on central 

portion, base and apex yellowish, hind tarsus yellowish. 

Male abdomen bright shiny, pregenital segment polished 

posteriorly, dull anteriorly; female abdomen with dense 

microtomentum medially, lateral margins of tergites nearly 

polished.

Morphology: 

Head: Head higher than long (head ratio less than 

0.9); face at level of pseudovibrissal seta wider than width 

of 1st flagellomere, bulbous; medial vertical seta 1/2–3/4 

length of lateral vertical seta; 1 strong peristomal seta present, following pseudovibrissal seta.

Thorax: Acrostichal setae 2 rows on anteriormost 

portion, merging into 1 row posteriorly; scutellum slightly 

raised (angle with scutum approximately 135°), semicircular, disk of scutellum strongly convex; basal scutellar seta 

3/4 length of posterior seta.

Male abdomen and terminalia: Figures 139–140. 

Surstylus a short, posteroventral lobe, in lateral view somewhat inconspicuous, approximately beak-shaped; cercus 

bearing 1 very long, posteriorly porrect setulae; gonopod 

as in Figure 139.

Measurements: Body length 2.9–3.1 mm. Wing 

length 2.5–2.7 mm, wing width 0.8–1.0 mm.

Type Material. The holotype male is labeled “PERU: Madre de Dios: Manu, Rio Manu, 250 m[,] 

Pakitza, 12°7¢S, 70°58¢W, 9–23 Sep 1988, Wayne N. 

Mathis/HOLOTYPE ♂ Aulacigaster bifaciata Rung & 

Mathis USNM [red].” The holotype is double mounted 

(minuten in a block of plastic), is in excellent condition, and 

is deposited in the USNM. Paratypes are as follows: Same 

label data as the holotype (4♂, 5♀; USNM).

Type Locality. Peru. Madre de Dios: Rio 

Manu, Pakitza (11°56.6¢S, 71°16.9¢W).

Other Specimens Examined. COSTA 

RICA. Guanacaste: Estación Santa Rosa (10°50.1¢N, 

FIGURES 137–138. Illustrations of Aulacigaster belize, new species (male) (the plesiomorphica group, Neotropical 

Region). (137) gonopod, lateral aspect; (138) epandrium, lateral aspect.
number 633 • 7 1

85°36.8¢W; 300 m), 11 Feb 1996, S. A. Marshall (1♀; 

UGE). Puntarenas: Peninsula de Osa, Rincón (08°42¢N, 

83°29¢W; sea level, sweeping in palm-mangrove forest), 24 

Mar 1991, D. A. Grimaldi, J. Stark (1♂; AMNH).

GUYANA. Moco Moco (30 km E Lethem in Kanuku 

Mountains; 03°18.2¢N, 59°39.0¢W), 3–6 Apr 1994, W. N. 

Mathis (1♀; USNM).

Distribution. (Map 9) Neotropical: Costa Rica 

(Guanacaste, Puntarenas), Guyana, Peru (Madre de Dios).

Etymology. The specific epithet, bifasciata, is 

of Latin derivation and is a combination of the prefix “bi-”, 

meaning two, and the Latin word for band (fascia) with reference to the black and silvery stripes toward the anterior 

margin of the frons.

Remarks. This species is not easily confounded 

with congeners and can be reliably separated from them 

with the characters given in the key and diagnosis.

Aulacigaster conspicua, new species

Figures 141–146, Map 11

Diagnosis. This species is distinguished from 

congeners by the following combination of characters: 

Frons with anterior 1/3–1/2 yellowish in ground color; face 

bulbous; facial band present; hind femur brown to black on 

central portion, apex and base yellowish.

Description. Coloration and vestiture: Vertex 

mostly densely microtomentose, without a shiny spot or 

with a shiny spot extended from ocellus at most 1/2 way 

to eye margin; ocellar tubercle dull microtomentose; frontal white-microtomentose stripe not bordered posteriorly by 

a conspicuous velvety-black stripe; antenna pale yellow to 

yellowish, infuscate along dorsal margin; face yellow anteriorly, brown to black on posterior portion; facial band present, distinct, gray or golden microtomentose over brown 

mark, on dorsal portion of face; palpus white. Scutellum 

dull microtomentose; postpronotum yellow to yellowishbrownish or concolorous with mesonotum, scarcely microtomentose, subshiny; katepisternum subshiny to dull; halter 

mostly white. Fore coxa brown to black, fore femur brown 

to black, apex yellowish, fore tibia yellowish, fore tarsus 

mostly or entirely yellowish, apical and sometimes subapical 

tarsomeres brown to black; mid coxa brown to black, mid 

femur brown to black, apex yellowish, mid tibia yellowish, 

mid tarsus mostly or entirely yellowish, apical and subapical 

tarsomere brownish to dark brown; hind coxa brown, hind 

femur brown to black on central portion, apex and base 

FIGURES 139–140. Illustrations of Aulacigaster bifasciata, new species (male) (the plesiomorphica group, Neotropical Region). (139) gonopod, lateral aspect; (140) epandrium, lateral aspect.
7 2 • smithsonian contributions to zoology

yellowish, hind tibia yellowish or brown to black on central 

portion, base and apex yellowish, hind tarsus mostly or entirely yellowish, apical tarsomere brown to black. Male abdomen bright shiny, pregenital segment polished posteriorly, 

dull anteriorly; female abdomen with dense microtomentum 

medially, lateral margins of tergites nearly polished.

Morphology: 

Head: Figures 141–142. Head higher than long 

(head ratio less than 0.9); face at level of pseudovibrissal 

seta wider than width of 1st flagellomere, bulbous; medial 

vertical seta 1/2–3/4 length of lateral vertical seta; 1 strong 

peristomal seta present, following pseudovibrissal seta.

Thorax: Acrostichal setae in 1 row; scutellum 

slightly raised (angle with scutum approximately 135°), 

semicircular, disk of scutellum slightly convex; basal scutellar seta 1/2 length of posterior seta.

Male abdomen and terminalia: Figures 143–146. 

Sternite 5 with a central, more membranous area, posterior 

margin appearing receded. Surstylus keel-shaped, with an 

apical and 1–2 basal marginal, strong setae; gonopod as in 

Figure 145; cerci partially fused, each cercus bearing 1 very 

long, posteriorly porrect setula.

Measurements: Body length 3.1–3.2 mm. Wing 

length 2.5–2.7 mm, wing width 0.8–0.9 mm.

Type Material. The holotype male is labeled 

“COSTA RICA: Puntarenas, Peninsula de Osa, Rincon[,] 

24 March 1991, sea level[,] Grimaldi & Stark, sweeping 

in palm-mangrove forest/HOLOTYPE ♂ Aulacigaster conspicua Rung & Mathis AMNH [red].” The holotype is 

double mounted (minuten in a block of plastic), is in good 

condition (vertical, left fronto-orbital, scutal and scutellar 

setae partially broken), and is deposited in the AMNH. 

Paratypes are as follows: Same label data as the holotype 

(1♂, 2♀; AMNH, USNM).

Type Locality. Costa Rica. Puntarenas: Peninsula de Osa, Rincón (08°42¢N, 83°29¢W).

Other Specimens Examined. COSTA 

RICA. Puntarenas: Rincón (08°42¢N, 83°29¢W; Malaise 

trap), Oct–Dec 1990, P. Hanson (5♀; USNM).

PANAMA. Canal Zone: Barro Colorado Island 

(09°09.3¢N, 79°50.8¢W), 10 Jan 1929, C. H. Curran (1♂, 

1♀; AMNH).

Distribution. (Map 11) Neotropical: Costa 

Rica (Puntarenas), Panama (Canal Zone).

FIGURES 141–144. Digital photographs of Aulacigaster conspicua, new species (male) (the plesiomorphica

group, Neotropical Region). (141) head, lateral aspect; 

(142) head, frontal aspect; (143) epandrium, lateral aspect; (144) subepandrial sclerite, ventral aspect. Not all 

to the same scale.
number 633 • 7 3

Etymology. The specific epithet, conspicua, is 

of Latin derivation and means “prominent” with reference 

to the conspicuous habitus, allowing for easy identification 

of this species.

Remarks. The four females from Puntarenas 

(Costa Rica), which were caught in a Malaise trap, are 

apparently this species but the legs and face are much 

lighter in the coloration. Aulacigaster conspicua is not easily confounded with congeners and can be reliably separated from them with the characters given in the key and 

diagnosis.

Aulacigaster erika, new species

Figures 147, Map 11

Diagnosis. This species is distinguished from 

congeners by the following combination of characters: 

Frons with anterior 1/3–1/2 yellowish in ground color; 

frons white-microtomentose stripe bordered posteriorly by 

a posterior, velvety-black stripe; facial band absent.

Description. Coloration: Vertex mostly 

densely microtomentose, with a shiny spot extended from 

ocellus to 2/3 distance to eye margin; ocellar tubercle dull 

microtomentose; frontal white-microtomentose stripe not 

bordered posteriorly by a posterior, velvet black stripe; 

antenna pale yellow to yellowish, infuscate along dorsal 

margin; face brownish; palpus white, or yellowish. Scutellum dull microtomentose; postpronotum concolorous with 

mesonotum; katepisternum subshiny to dull; halter mostly 

white (base dark). Fore coxa brown to black, fore femur 

brown to black, apex yellowish; fore tibia yellowish, fore 

tarsus mostly yellowish; mid coxa brown to black, mid 

femur brown to black, tip yellowish; mid tibia yellowish to 

brownish; mid tarsus mostly yellowish; hind coxa brown to 

black, hind femur brown to black, tip yellowish; hind tibia 

yellowish-brownish or brown to black on central portion, 

base and apex yellowish, hind tarsus yellowish. Male abdomen bright shiny, pregenital segment polished posteriorly, 

dull anteriorly; female abdomen with dense microtomentum medially, lateral margins of tergites nearly polished.

Morphology: 

Head: Head higher than long (head ratio less than 

0.9); face at level of pseudovibrissal seta wider than width 

of 1st flagellomere, not appearing bulbous; medial vertical 

seta 1/2–3/4 length of lateral vertical seta; 1 strong peristomal seta present, following pseudovibrissal seta.

Thorax: Acrostichal setae in 1 row; scutellum 

slightly raised (angle with scutum approximately 135°), 

approximately triangular, apex relatively pointed, disk of 

FIGURES 145–146. Illustrations of Aulacigaster conspicua, new species (male) (the plesiomorphica

group, Neotropical Region). (145) gonopod, lateral aspect; (146) epandrium, lateral aspect.
7 4 • smithsonian contributions to zoology

scutellum slightly convex; basal scutellar seta 1/2 length of 

posterior seta.

Male abdomen and terminalia: Figure 147. Sternite 

5 with a central, more membranous area, posterior margin 

appearing receded. Surstylus approximately nose-shaped, 

apex slightly pointed; gonopod as in Figure 147; cerci partially fused, each cercus bearing 1 very long, posteriorly 

porrect setulae.

Measurements: Body length 2.4–2.5 mm. Wing 

length 2.3–2.4 mm, wing width 0.8–0.9 mm.

Type Material. The holotype male is labeled 

“PERU. Madre de Dios: Manu, Erika (near Salvacion), 

550 m, 5–6 Sept 1988,W.N.Mathis/HOLOTYPE ♂ Aulacigaster erika Rung & Mathis USNM [red].” The holotype 

is double mounted (minuten in a block of plastic), is in 

excellent condition, and is deposited in the USNM. Paratypes are as follows: Same label data as the holotype (3♂, 

7♀; USNM); same locality data but with A. Freidberg (9♂, 

1♀; USNM). PERU. Madre de Dios: Pakitza (11°56.6¢S, 

71°16.9¢W; 250 m), 9–23 Sep 1988, A. Freidberg, W. N. 

Mathis (12♂, 7♀; USNM).

Type Locality. Peru. Madre de Dios: Rio 

Manu, Rio Manu, Erika (near Salvación; 12°53¢S, 71°12¢W; 

550 m).

Other Specimens Examined. ECUADOR. 

Orellana: Rio Tiputini (00°38.2¢S, 76°08.9¢W), 12–26 

Aug 1999, W. N. Mathis, A. Baptista, M. Kotrba (5♂, 2♀; 

EPNE, USNM).

MAP 11. Distribution of Aulacigaster conspicua and A. erika (the plesiomorphica group, Neotropical Region).
number 633 • 7 5

Distribution. (Map 11) Neotropical: Ecuador (Orellana), Peru (Madre de Dios).

Biology. This species was collected while sweeping the aerial portion of prop roots of a large Ceiba tree.

Etymology. The specific epithet, erika, refers 

to the type locality, Erika, and is a noun in apposition.

Remarks. This species is not easily confounded 

with congeners and can be reliably separated from them 

with the characters given in the key and diagnosis.

Aulacigaster plesiomorphica (Hennig)

Figures 148, 152–154, Map 10

Schizochroa plesiomorphica Hennig 1969: 608, 605 [figure of head and 

wing].—Papavero 1967:91 [Neotropical catalog].

Aulacigaster plesiomorphica.—D. K. McAlpine 1983:58 [generic combination].

Diagnosis. This species is distinguished from 

congeners by the following combination of characters: 

Hind trochanter of male with a ventral, black tuff of setae 

born on a projection.

Description. Coloration and vestiture: Vertex 

mostly densely microtomentose, with a shiny spot extended 

from ocellus at most 1/2 way to eye margin; ocellar tubercle 

dull microtomentose; frontal white-microtomentose stripe 

bordered posteriorly by a conspicuous velvety-black stripe; 

antenna pale yellow to yellowish, infuscate on dorsal half; 

face brown to black; facial band present, delimited as a 

transversely wrinkled band on dorsal portion of face, mostly 

lacking microtomentum; palpus white. Scutellum dull microtomentose; postpronotum concolorous with mesonotum; anepisternum mostly dull; katepisternum subshiny to 

dull; halter mostly white. Fore coxa brown to black, fore 

femur brown or brown to black, apex yellowish, fore tibia 

yellowish, fore tarsus mostly yellowish; mid coxa brown to 

black, mid femur brown or brown to black, apex yellowish, mid tibia yellowish, mid tarsus mostly yellowish; hind 

coxa brown to black, hind femur brown or brown to black, 

apex yellowish, hind tibia yellowish or dark basally, yellow 

apically, hind tarsus yellowish. Male abdomen bright shiny, 

pregenital segment mostly subshiny; female abdomen with 

dense microtomentum medially, lateral margins of tergites 

nearly polished.

Morphology: 

Head: Head higher than long (head ratio less than 

0.9); face not appearing “bulbous”; medial vertical seta 

1/2–3/4 length of lateral vertical seta; 1 strong peristomal 

seta present, following pseudovibrissal seta.

Thorax: Figures 148, 152. Acrostichal setae in 1 

row; scutellum strongly raised (angle with scutum more 

than 135°), approximately triangular, apex relatively 

pointed, disk of scutellum strongly convex; basal scutellar 

seta 1/2 length of posterior seta. Hind trochanter of male 

with a ventral, black tuff of setae born on a projection; hind 

tibia with strong, dark, apicodorsal setae.

Male abdomen and terminalia: Figures 153–154. 

Surstylus more or less folded below the epandrium, difficult to see in lateral view, approximately triangular, with 1 

apical and 2 basal, marginal, strong setae; gonopod as in 

Figure 153; cerci separate.

Measurements: Body length 2.7–3.2 mm. Wing 

length 2.6–2.7mm, wing width 1.0 mm.

Type Material. The holotype is labeled 

“Quincemil [.] Cuzco, PERU[,]. 13–31.VIII.’62 [,] L. Pena. 

780 m./Schizochroa plesiomorphica Hg. [handwritten] 

Holotypus [red]/HOLOTYPE Schizochroa plesiomorphica Hennig CNC No. 9929 [red]/CNC [handwritten].” The holotype is glued directly to the pin, is in 

FIGURE 147. Illustrations of Aulacigaster erika, new species (male) 

(the plesiomorphica group, Neotropical Region): epandrium and 

gonopod, lateral aspect.
7 6 • smithsonian contributions to zoology

good condition (teneral, slightly dirty), and is deposited in 

the CNC (9929). Paratypes are as follows: same locality 

and collector as the holotype, 15–30 Dec 1962 (1♂, CNC); 

1–15 Nov 1962 (1♀, CNC).

Type Locality. Peru. Cuzco: Quincemil 

(13°13.7¢S, 70°45.6¢W).

Other Specimens Examined. PERU. 

Madre de Dios: Rio Manu, Erika (near Salvación; 12°53¢S, 

71°12¢W; 550 m), 5–6 Sep 1988, A. Freidberg (1♂; USNM).

Distribution. (Map 10) Neotropical: Peru 

(Cuzco, Madre de Dios).

Remarks. The holotype is teneral, and description of coloration is based on the two conspecific paratypes. 

Two specimens in the paratype series (1♂, 1♀; CNC) represent another species (A. rufifemur, see below). Aulacigaster 

plesimorphica share with males of A. ariseta and A.proxima

the presence of black, dorsoapical setae on the hind femur. 

Aulacigaster atriseta can be easily distinguished from the 

other two species by the absence of well-developed setae 

on the hind trochanter of the male and by the absence of 

a conspicuous surstylus. Males of A. plesiomorphica have 

a black tuft of thick sete on the hind trochanter, whereas 

males of A. proxima have a yellowish tuft of thinner setae. 

Females of these two species cannot be reliably separated.

Aulacigaster proxima, new species

Figures 149–151, 155, 213–214, Map 12

Diagnosis. This species is distinguished from 

congeners by the following combination of characters: 

Hind trochanter of male with a ventral, yellowish tuff of 

setae not born on a projection.

Description. Coloration: Vertex mostly 

densely microtomentose, with a shiny spot extended from 

ocellus to at most 1/2 way to eye margin, with a shiny spot 

FIGURES 148–151. Digital photographs of aulacigastrids (the plesiomorphica group, Neotropical Region). (148) Aulacigaster plesiomorphica

(Hennig), hind trochanter of male, posterior aspect; (149) Aulacigaster proxima, hind trochanter of male, posterior aspect; (150) enlargement 

of costal section, dorsal aspect; (151) right wing, dorsal aspect. Not all to the same scale.
number 633 • 7 7

extended from ocellus to 2/3 distance to eye margin; ocellar 

tubercle dull microtomentose; frontal white-microtomentose 

stripe bordered posteriorly by a conspicuous velvety-black 

stripe; antenna pale yellow to yellowish, infuscate along dorsal margin or infuscate on dorsal half; face yellow anteriorly, 

brown to black on posterior portion (from level of pseudovibrissa to base of antenna); facial band present, delimited as a 

transversely wrinkled band on dorsal portion of face, mostly 

lacking microtomentum; palpus white. Scutellum dull microtomentose; postpronotum concolorous with mesonotum, 

scarcely microtomentose, subshiny to dull microtomentose; anepisternum mostly dull; katepisternum subshiny to 

dull; halter mostly white. Fore coxa brown to black, fore 

femur brown to black, apex yellowish, fore tibia yellowish, 

fore tarsus mostly yellowish; mid coxa brown to black, mid 

femur brown to black, apex yellowish, mid tibia yellowish, 

mid tarsus mostly yellowish; hind coxa brown, hind femur 

brown to black, apex yellowish, hind tibia yellowish or dark 

basally, yellow apically, hind tarsus yellowish. Male abdomen bright shiny, pregenital segment polished posteriorly, 

dull anteriorly; female abdomen with dense microtomentum 

medially, lateral margins of tergites nearly polished.

Morphology: 

Head: Figures 213–214. Head round, about as high 

as long (head ratio 0.9–1.1) to higher than long (head ratio 

less than 0.9); face at level of pseudovibrissal seta narrower 

than width of 1st flagellomere, not appearing bulbous; medial vertical seta 1/2 length of lateral vertical seta; 1 strong 

peristomal seta present, following pseudovibrissal seta.

Thorax: Figures 149–151. Acrostichal setae in 1 

row; scutellum strongly raised (angle with scutum more 

than 135°), approximately triangular, apex relatively 

pointed, disk of scutellum flat; basal scutellar seta 1/2 

length of posterior seta (but thinner). Hind trochanter of 

male with a ventral, yellowish tuff of setae not borne on a 

projection; hind tibia with strong, dark, apicodorsal setae.

Male abdomen and terminalia: Figure 155. Sternite 

4 with posterior margin and medial portion only weakly 

sclerotized, posterior margin receded; sternite 5 with a 

central, more membranous area, posterior margin appearing receded. Surstylus more or less folded below the 

epandrium, difficult to see in lateral view, approximately 

foot-shaped, with 2 subapical and 2 basal, marginal, 

strong setae; gonopod as in Figure 155; cerci separate, 

FIGURES 152–154. Illustrations of Aulacigaster plesiomorphica (Hennig) (male) (the plesiomorphica group, Neotropical Region). (152) apex 

of hind tibia, anterodorsal aspect; (153) gonopod, lateral aspect; (154) epandrium, lateral aspect. Not all to the same scale. 
7 8 • smithsonian contributions to zoology

narrow, digitiform, each cercus bearing 1 very long, ventral 

seta and 1–3 longer setulae.

Measurements: Body length 3.10–3.53 mm. Wing 

length 2.7–3.2 mm, wing width 0.9–1.0 mm.

Type Material. The holotype male is labeled “PERU. Madre de Dios: Manu, Rio Manu, 250 m[,] 

Pakitza, 12°7¢S, 70°58¢W, 9–23 Sep 1988[,] Amnon 

Freidberg/HOLOTYPE ♂ Aulacigaster proxima Rung & 

Mathis USNM [red].” The holotype is double mounted 

(minuten in a block of plastic), is in good condition (left 

posterior dorsocentral seta missing), and is deposited in the 

USNM. Paratypes are as follows: Same label data as the 

holotype (2♂; USNM); same label as the holotype but collected by W. N. Mathis (11♂; USNM).

Type Locality. Peru. Madre de Dios: Rio 

Manu, Pakitza (11°56.6¢S, 71°16.9¢W).

Other Specimens Examined. ECUADOR. 

Orellana: Rio Tiputini (00°38.2¢S, 76°08.9¢W), 12–26 Aug 

1999, W. N. Mathis, A. Baptista, M. Kotrba (3♂; EPNE, 

USNM).

Distribution. (Map 12) Neotropical: Ecuador (Orellana), Peru (Madre de Dios).

Etymology. The specific epithet, proxima, is 

of Latin derivation and means near, referring to the fact 

that it is very similar to A. plesiomorphica. See “Remarks” 

under A. plesiomorphica for separating between these two 

species.

Aulacigaster rufifemur, new species

Figures 156–159, Map 13

Diagnosis. This species is distinguished from 

congeners by the following combination of characters: 

Frons with anterior 1/3–1/2 yellowish in ground color; vertex with a shiny spot extended from ocellus 1/2 way to eye 

margin; hind femur yellow.

Description. Coloration: Vertex mostly 

densely microtomentose, with a shiny spot extended from 

ocellus at most 1/2 way to eye margin; ocellar tubercle dull 

microtomentose; frontal white-microtomentose stripe bordered posteriorly by a conspicuous velvety-black stripe; 

antenna pale yellow to yellowish, infuscate along dorsal 

margin or infuscate on dorsal half; face yellow anteriorly, 

FIGURE 155. Illustrations of Aulacigaster proxima, new species (the plesiomorphica group, Neotropical Region): epandrium and internal genitalic structures (male), lateral aspect.
number 633 • 7 9

brown to black on posterior portion; facial band present, 

distinct, gray or golden microtomentose over brown mark, 

on dorsal portion of face; palpus white. Scutellum dull microtomentose; postpronotum concolorous with mesonotum; anepisternum mostly dull; katepisternum subshiny to 

dull; halter mostly white. Fore coxa brown to black, fore 

femur brown to black, up to apical third yellowish, fore 

tibia yellowish, fore tarsus yellowish, apical and sometimes 

subapical tarsomeres brown to black; mid coxa brown to 

black, mid femur brown, with apex yellowish, mid tibia 

yellowish, mid tarsus yellowish, apical and subapical tarsomere brownish to dark brown; hind coxa yellowish to 

brownish, hind femur yellow, hind tibia yellowish, hind tarsus yellowish, apical tarsomere brown to black. Male abdomen bright shiny, pregenital segment polished posteriorly, 

dull anteriorly; female abdomen with dense microtomentum medially, lateral margins of tergites nearly polished.

Morphology: 

Head: Figure 158, Head higher than long (head 

ratio less than 0.9); face at level of pseudovibrissal seta wider 

than width of 1st flagellomere, not appearing bulbous; medial vertical seta 1/2–3/4 length of lateral vertical seta; 3–4 

peristomal seta present, following pseudovibrissal seta.

Thorax: Figure 156. Acrostichal setae in 1 row, or 

two rows anteriorly, convergent into one row posteriorly; 

scutellum slightly raised (angle with scutum approximately 

135°), semicircular, disk of scutellum strongly convex; basal 

scutellar seta 1/2 length of posterior seta (at most).

Male abdomen and terminalia: Figures 157, 

159. Sternite 5 with a central, more membranous area, 

MAP 12. Distribution of Aulacigaster proxima and A. trifasciata (the plesiomorphica group, Neotropical Region).
8 0 • smithsonian contributions to zoology

posterior margin appearing receded. Surstylus approximately triangular in lateral view, apex strongly pointed, 

surface concave; gonopod as in Figure 159; cerci partially 

fused, each cercus bearing 1 very long, posteriorly porrect 

setulae.

Measurements: Body length 2.9–3.0 mm. Wing 

length 2.7–3.2 mm, wing length 2.2–2.6, wing width 0.9–

1.0 mm.

Type Material. The holotype male is labeled 

“PERU. Madre de Dios: Manu, Rio Manu, 250 m[,] Pakitza, 

FIGURES 156–159. Digital photographs and illustrations of Aulacigaster rufifemur, new species (male) (the plesiomorphica group, Neotropical Region). (156) right wing, dorsal aspect; (157) subepandrial sclerite, ventral aspect; (158) head, lateral aspect; (159) epandrium and 

gonopod, lateral aspect. Not all to the same scale.
number 633 • 8 1

12°7¢S, 70°58¢W, 9–23 Sep 1988[,] Amnon Freidberg/HOLOTYPE ♂ Aulacigaster rufifemur Rung & Mathis USNM 

[red].” The holotype is double mounted (minuten in a block 

of plastic), is in excellent condition, and is deposited in the 

USNM. Paratypes are as follows: Same label data as the 

holotype (29♂, 16♀; USNM); same locality data as the holotype but collected by W. N. Mathis (9♂, 4♀; USNM).

Type Locality. Peru. Madre de Dios: Rio 

Manu, Pakitza (11°56.6¢S, 71°16.9¢W).

Other Specimens Examined. BOLIVIA. 

La Paz: S Inicua River (15°20¢S, 67°33¢W; 1100 m), Alto 

Beni, 15–18 Jan 1976, L. E. Peña (1♀; CNC).

BRAZIL. Rondônia: Ariquemes (62 Km SE; 10°34¢S, 

62°22¢W), 22 Oct–16 Nov 1996, 1997, W. J. Hanson 

(2♀, USU).

ECUADOR. Orellana: Rio Tiputini (00°38.2¢S, 

76°08.9¢W), 12–26 Aug 1999, W. N. Mathis, A. Baptista, 

M. Kotrba. (11♂, 3♀; EPNE, USNM).

PERU. Cuzco: Quincemil (13°13.7¢S, 70°45.6¢W; 780 

m), 13–31 Aug 1962, L. Peña (1♀; labeled as a “paratype” 

of A. plesiomorphica CNC). Madre de Dios: Avispas 

(12°58.7¢S, 70°20.8¢W; 400 m), 20–30 Sep 1962, L. Peña 

(1♂; labeled as a “paratype” of A. plesiomorphica CNC); 

Rio Manu, Erika (near Salvación; 12°53¢S, 71°12¢W; 550 

m), 5–6 Sep 1988, A. Freidberg (1♀; USNM); Rio Manu, 

Cocha Salvador (11°57¢S, 72°07¢W; 240 m), 14 Sep 1988, 

W. N. Mathis (1♀; USNM).

Distribution. (Map 13) Neotropical: Brazil 

(Rondônia), Bolivia (La Paz), Ecuador (Orellana), Peru 

(Madre de Dios).

MAP 13. Distribution of Aulacigaster rufifemur (the plesiomorphica group, Neotropical Region).
8 2 • smithsonian contributions to zoology

Etymology. The specific epithet, rufifemur, is 

of Latin derivation and is a combination of the Latin words 

for red (rufus) and thigh (femur). This character makes this 

species unique and very easy to separate from congeners in 

the plesiomorphica group.

Aulacigaster trifasciata, new species

Figure 160, Map 12

Diagnosis. This species is distinguished from 

congeners by the following combination of characters: 

Ocellar tubercle mostly polished; abdomen with a large, 

white to yellowish region on syntergite 1+2.

Description. Coloration: Vertex mostly 

densely microtomentose, with a shiny spot extended from 

ocellus to 2/3 distance to eye margin; ocellar tubercle mostly 

polished; frontal white-microtomentose stripe not bordered 

posteriorly by a velvety-black stripe; antenna pale yellow to 

yellowish, infuscate along dorsal margin; face white; facial 

band present, distinct, gray or golden microtomentose over 

brown mark, on dorsal portion of face; palpus white. Scutellum dull microtomentose; postpronotum concolorous with 

mesonotum, dull microtomentose; anepisternum mostly 

dull; katepisternum subshiny to dull; halter mostly white. 

Fore coxa yellow, fore femur yellowish, fore tibia yellowish, fore tarsus yellowish, apical and sometimes subapical 

tarsomeres brown to black; mid coxa yellow, mid femur yellowish (tip infuscate in some specimens), mid tibia yellowish, mid tarsus yellowish, apical and subapical tarsomere 

brownish to dark brown; hind coxa yellow, hind femur yellow on basal portion, apical portion dark (apical 1/3 infuscate to black), hind tibia yellowish, hind tarsus yellowish, 

apical tarsomere brown to black. Abdomen with a large, 

white to yellowish region on syntergite 1+2, male abdomen 

bright shiny; female abdomen with dense microtomentum 

medially, lateral margins of tergites nearly polished.

Morphology: 

Head: Head higher than long (head ratio less than 

0.9); face not appearing bulbous; medial vertical seta 1/2–

3/4 length of lateral vertical seta; 1 strong peristomal seta 

present, following pseudovibrissal seta.

Thorax: Acrostichal setae in 1 row; scutellum 

slightly raised (angle with scutum approximately 135°), 

approximately triangular, apex relatively pointed, disk of 

scutellum slightly convex; basal scutellar seta 1/2 length of 

posterior seta. Hind femur modified, with 2 ventral rows 

of spines, each spine bearing an apical seta; hind tibia with 

a row of tiny ventral spines; hind tibia without ventroapical projections.

Male abdomen and terminalia: Figure 160. Sternite 5 with a central, more membranous area, posterior 

margin appearing receded, and bearing a tongue-like medial projection. Surstylus from lateral view “boat-shaped” 

and bearing one apical seta; gonopod as in Figure 160; 

cerci partially fused, each cercus bearing 1 very long, ventral seta and 1–3 longer setulae.

Measurements: Body length 2.5–2.7 mm, wing 

length 2.4–2.7 mm, wing width 0.8–0.9 mm.

Type Material. The holotype male is labeled 

“ECUADOR. P[ue]rt[o] Orellana: RioTiputini (0°38.2¢S, 

76°8.9¢W)[,] 12–26 Aug 1999, W. N. Mathis, A. Baptista, M. Kotrba/HOLOTYPE ♂ Aulacigaster trifasciata

Rung & Mathis USNM [red].” The holotype is double 

mounted (minuten in a block of plastic), is in excellent 

condition, and is deposited in the USNM. Paratypes are 

as follows: Same label data as the holotype (54♂, 19♀; 

EPNE, USNM).

Type Locality. Ecuador. Orellana: Rio Tiputini 

(00°38.2¢S, 76°08.9¢W).

Other Specimens Examined. COSTA RICA. 

Puntarenas: Peninsula de Osa, Rincón (08°42¢N, 83°29¢W; 

FIGURE 160. Illustrations of Aulacigaster trifasciata, new species 

(male) (the plesiomorphica group, Neotropical Region): epandrium 

and gonopod, lateral aspect. Not all to the same scale.
number 633 • 8 3

sea level; sweeping in palm-mangrove forest), 24 Mar 

1991, D. A. Grimaldi, J. Stark (1♂; AMNH).

PANAMA. Canal Zone: Barro Colorado Island 

(09°09.3¢N, 79°50.8¢W), 10 Jan 1929, C. H. Curran (2♂; 

AMNH).

PERU. Madre de Dios: Manu, Rio Manu, Pakitza 

(11°56.6¢S, 71°16.9¢W); 250 m, 9–23 Sep 1988, A. Freidberg, W. N. Mathis (8♂, 4♀; USNM).

Distribution. (Map 12) Neotropical: Costa Rica 

(Puntarenas), Ecuador (Orellana), Panama (Canal Zone), 

Peru (Madre de Dios).

Etymology. The specific epithet, trifasciata, is of 

Latin derivation and is a combination of the prefix “tri-”, 

meaning three, and the Latin word for band (fascia) in 

reference to the three bands on eyes of live specimens.

Remarks. See “Remarks” under A. albifacies.

The leucopeza Group

Figures 161–201, 215–220, Maps 14–18

Diagnosis. The leucopeza group is distinguished from other species groups by the following combination of characters: Stout, small to medium-sized flies, 

body length 1.6–3.5 mm. Coloration and vestiture: Frons 

typically bearing a transverse, orange to orange-brown 

stripe, often followed by a brown stripe or shadow; anterior 

portion of frons densely microtomentose, with a narrow, 

anterior, silver stripe merging with brown ventrally; facial 

band often present. Wing infuscate with brown in known 

Afrotropical species. Abdomen in a few species with a large, 

white to yellowish region on syntergite 1+2.

Morphology: 

Head: Head typically higher than long (head ratio 

less than 0.9); arista short, with very inconspicuous, alternate 

hairs (arista appearing naked at low magnification) or short, 

with tiny, dense, alternate hairs, arista appearing straight; 

face in lateral view with dorsal half concave, ventral portion 

bulbous; gena in most species in lateral view approximately 

1/7–1/2 the width of 1st flagellomere; ocellar seta typically 

minute; fronto-orbital setae typically with posterior seta internal to and almost horizontally aligned with anterior seta.

Thorax: Subcosta partially fused with vein R1

 apically but terminating in costa.

Male abdomen and terminalia: Surstylus often arising from posteroventral margin of epandrium; cerci often 

broad, plate-like; subepandrial sclerite forming a single, 

plate-like structure. Gonopods short, wide, with a few central setae and a posterior lobe or process.

Distribution. Worldwide, excluding the Australian and Oceanic Regions.

Discussion. Species of the leucopeza group are 

best identified by the coloration and shape of the ventral portion of the frons, the presence and extent of microtomentum 

posterior to the fronto-orbital setae, the coloration of legs, 

and the shape of the surstylus and male internal terminalia.

The leucopeza group includes 15 species of which 

nine were described previously. Thus far, only species of 

the leucopeza group occur outside the Neotropical Region. 

Mathis and Freidberg (1994) revised the Nearctic Aulacigaster, and provided a key for the species known to occur 

in that region. Papp (1998a) published a key to the Palearctic species. Barraclough (1993) and Hilger and Kassebeer 

(2000) treated the Afrotropical species of the genus.

Afrotropical Species

Discussion. The Afrotropical fauna of the 

leucopeza group presently comprises three described and 

three previously undescribed species, two of which are described here. One species, Aulacigaster perata Barraclough, 

is known only from a single female.

Key to the Afrotropical Species of Aulacigaster

1. Anterior dorsocentral seta lacking. Wing hyaline . . . . . . . . . . . . . . . . . . A. sp., undescribed species (Barraclough, 1993)

Anterior dorsocentral seta present. Wing faintly brown infuscate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2. Palpus short, length about half that of 1st flagellomere . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

Palpus long, subequal to 1st flagellomere . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. africana Barraclough

3. Abdomen completely dark brown to black. Coxae yellowish to brownish. . . . . . . . . . . . . . . . . . . . A. perata Barraclough

Abdomen with syntergite 1+2 pale. Coxae brown to dark brown .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

4. Vertex with a shiny spot extended from ocellus at most 1/2 way to eye margin (fig. 1b; Hilger and Kassebeer, 2000:169). 

Posterodorsal process of gonopod, in lateral view, shorter than surstylus, approximately triangular (fig. 2c; Hilger and 

Kassebeer, 2000:170) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. borbonica Hilger and Kassebeer

Vertex with a shiny spot extended from ocellus to at least 2/3 of eye margin. Posterodorsal process of gonopod, in lateral 

view, almost as long as surstylus, variously shaped [Figures 166, 168] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
8 4 • smithsonian contributions to zoology

5. Surstylus shorter, about 0.2–0.3 times the length of ventral margin of epandrium [Figure 169]; posterior process of gonopod digitiform, at least 3 times wider than surstylus, bearing one central seta [Figure 168] . . . . . . . . . . . . . . . . . . . . . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. malawana, new species

Surstylus longer, about 0.3–0.4 times the length of ventral margin of epandrium [Figure 167]; posterior process of gonopod with lateral margins convergent apically, pointed, at least 1/2 the width of surstylus on its narrowest point, devoid of 

setae [Figure 166] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. freidbergi, new species

Aulacigaster africana Barraclough

Figures 161–165, Map 14

Aulacigaster africana Barraclough, 1993:34–38 [figures of head, wing and 

male terminalia].—Duxbury and Barraclough, 1994:35 [Figure 4].—

Papp, 2008:227–232 [description, illustrations of female terminalia and 

immature stages].

Diagnosis. This species is distinguished from 

congeners by the following character: Palpus long, subequal 

in length to third antennal segment.

Description. Coloration and vestiture: Vertex 

mostly densely microtomentose, with a shiny spot extended 

from ocellus at most 2/3–3/4 to eye margin; frons bearing 

a transverse, large, orange band, merging into dark brown 

ventrally; silver stripe on frons approximately straight; antenna yellow, infuscate with brown on dorsal 1/2; face yellow in ground color; facial band present, sharply defined, 

transverse, on protruding, ventral portion of face; palpus 

white. Scutum dull microtomentose anteriorly, devoid of 

microtomentum posterior of transverse suture; scutellum 

mostly devoid of microtomentum, bright shiny; postpronotum concolorous with mesonotum, microtomentose over 

ventral and posterior margins, otherwise polished; anepisternum mostly microtomentose, with a basal silver stripe extended posteriorly to anepimeron; katepisternum subshiny 

to dull; halter mostly white. Wing membrane very faintly 

infuscate, darker along costal margin and apex (not as pronounced as suggested in fig. 2; Barraclough, 1993:35). Fore 

coxa brown to black, fore femur brown, apex yellowish; 

fore tibia yellowish or brown, fore tarsus yellowish, apical 

and sometimes subapical tarsomeres brown to black; mid 

coxa brown to black, apex yellowish, mid femur brown to 

black, apex yellowish, mid tibia yellowish or brownish, mid 

tarsus yellowish, apical and subapical tarsomere brownish 

to dark brown; hind coxa brown, apex yellowish, hind 

femur brown to black, apex yellowish, hind tibia yellowish or brown to dark brown, hind tarsus yellowish, apical 

tarsomere brown to black. Male abdomen subshiny to dull.

Morphology: 

Head: Figures 162–164. Head higher than long 

(head ratio less than 0.9); face in lateral view with dorsal 

half concave, ventral portion bulbous; gena in lateral view 

1/5–1/6 the width of 1st flagellomere; palpus long, subequal 

in length to 1st flagellomere; setae on head and mesonotum 

unusually long (posterior fronto-orbital seta longer than 1/2 

length of eye in lateral view); medial vertical seta 3/4 length 

of lateral vertical seta; 1 well-developed peristomal seta and 

several setulae following pseudovibrissal setae present.

Thorax: Figure 161. Acrostichal setae in 2 rows.

Male abdomen and terminalia: Figure 165. Surstylus arising from ventral margin of epandrium, slender, 

slightly sickle-shaped, relatively long (approx. 1/2 length 

of ventral margin of epandrium); cerci broader at base, 

tapering toward round apex, with a few prominent apical 

setae; gonopod with 1 medial seta and several setae preceding posterior dorsal process, subrectangular in lateral 

view (approx. 0.7 times as high as long); posterior process 

of gonopod digitiform (slightly shorter than surstylus), 

not particularly wide (slightly wider than base of surstylus), bearing a few marginal ventral setae (figs. 3–4; Barraclough, 1993:35).

Measurements: (from one paratype male) Body 

length 2.8 mm. Wing length 3.0 mm, wing width 1.0 mm.

Type Material. The holotype male is labeled 

“S. AFRICA: Natal #41[,] Royal Natal Nat. Park[,] 28°43¢S: 

28°56¢E[,] 1540 m[,] 2.ix.1992, Indig. Forest Near Tendele 

and Tugela[,] Rock ledge, Whittington/HOLOTYPE ♂ Aulacigaster africana BARRACLOUGH [red border].” The 

holotype is deposited in NMSA (597). Paratype as follows: 

same label as the holotype, but collected by D. A. Barraclough (1♂; NMSA).

Type Locality. South Africa. KwaZulu-Natal: 

Royal Natal National Park (28°43¢S, 28°56¢E).

Other Specimens Examined. South Africa. Natal: Royal Natal Nat. Park (28°43¢S, 28°56¢E; 

Indig Forest near Tendele and Tugela, Rock ledge; 1540 m), 

2 Sep 1992, A. E. Whittington (1♂; NMSA).

Distribution. (Map 14) Afrotropical: South 

Africa (Eastern Cape, KwaZulu-Natal).

Remarks. The female is not known for this 

species. The description presented here is based on a male 

paratype and a conspecific male, both collected in the same 

locality as the holotype. According with Barraclough (1993), 

A. africana was collected from a shaded sandstone ledge.
number 633 • 8 5

Aulacigaster borbonica Hilger and Kassebeer

Map 14

Aulacigaster borbonica Hilger and Kassebeer, 2000:167–172 [figures of male 

terminalia, head and thorax].

Diagnosis. This species is distinguished from 

congeners by the following combination of characters: 

Abdomen with syntergite 1+2 whitish; mid tarsus brown, 

basal 3 tarsomeres yellow; thoracic pleural region with a 

stripe of white microtomentum beginning on fore coxa and 

ending beneath halter.

Description. Coloration and vestiture: Vertex 

mostly densely microtomentose, with a shiny spot extended 

from ocellus at most 1/2 way to eye margin; frons bearing 

a transverse, large, orange band, frontal orange band merging into dark brown ventrally; silver stripe on frons approximately straight; antenna brownish; face brownish; facial 

band present, yellow (Hilger and Kassebeer 2000:168), 

sharply defined, transverse, on protruding, ventral portion 

of face; palpus white. Scutum uniformly microtomentose; 

FIGURES 161–164. Digital photographs of Aulacigaster africana Barraclough (male) (the leucopeza group, 

Afrotropical Region). (161) body, lateral aspect; (162) head, lateral aspect; (163) head, frontal aspect; (164) 

head, dorsal aspect. Not all to the same scale.
8 6 • smithsonian contributions to zoology

postpronotum concolorous with mesonotum, more strongly 

microtomentose on lateral and posterior margins, subshiny; 

thoracic pleural region with a stripe of white microtomentum beginning on fore coxa and ending beneath the halter; 

halter mostly white. Wing mostly brown, infuscate. Fore 

coxa brown to black, fore femur brown, fore tibia brown, 

fore tarsus brown to dark brown, basal 2 tarsomeres white 

to yellow; mid coxa brown to black, mid femur brown, mid 

tibia brownish, mid tarsus brown, basal 3 tarsomeres white 

to yellow; hind coxa brown, hind femur brown, hind tibia 

brown to dark brown, hind tarsus brown, basal 3 tarsomeres white to yellow. Abdomen with dorsal portion of 

tergite 1 white.

Morphology: 

Head: Face in lateral view with dorsal half concave, ventral portion bulbous; gena in lateral view approximately 1/3–1/2 the width of 1st flagellomere; medial 

vertical seta 3/4 length of lateral vertical seta; peristomal 

area with about 1 strong and several weaker setae following 

pseudovibrissal seta (fig. 1a; Hilger and Kassebeer 2000).

Thorax: Acrostichal setae in 2 rows.

Male abdomen and terminalia: Surstylus arising 

from posteroventral margin of epandrium, slender, digitiform, relatively long (approx. 0.4 times the length of ventral margin of epandrium; Figure 2a; Hilger and Kassebeer, 

2000:170) with round and slightly enlarged apex; subepandrial sclerite with lateral arms broad and curved posteriorly (fig. 2b; Hilger and Kassebeer, 2000:170); cerci fused 

on basal 1/2, broader at base, with almost parallel margins and with a few prominent apical setae; gonopod with 

one medial seta and three setae preceding posterior process, sub-rectangular in lateral view (approx. height twice 

length), with a short pointed process posterodorsally (less 

than 1⁄2 the length of surstylus; fig. 2c; Hilger and Kassebeer, 

2000:170).

Measurements: Body length 2.0–2.8 mm. Wing 

length 2.0–2.3 mm, wing width not available in original 

description.

Type Material. The holotype male is labeled 

(quoted from Hilger and Kassebeer, 2000:168E) “La Reunión: 

Salazie, Mare à Poule d’Eau, 490 m, 21°03¢72S 55°31¢292E, 

on slime flux of Casuarina cunninghamiana, 2.3.1999, leg. 

C. F. Kassebeer (CFK) & S. Hilger, in Coll. CFK.”

Type Locality. La Reunión. Salazie (21°03¢S, 

55°30.4¢E).

Distribution. (Map 14) Afrotropical: 

Reunión.

Biology. According to Hilger and Kassebeer 

(2000), A. borbonica was collected from a slime flux of the 

Horsetail Tree (Casuarina cunninghamiana Miq.), originally from Australia and introduced to Réunion.

Remarks. The description herein presented is 

based on Hilger and Kassebeer (2000). We were not able 

to analyze the type specimens because they are deposited 

in the Kassebeer’s personal collection and he has not responded to our requests to send the material. 

Aulacigaster freidbergi, new species

Figures 166–167, Map 14

Diagnosis. This species is distinguished from 

congeners by the following combination of characters: Abdomen with syntergite 1+2 whitish; gena in lateral view 1/7 

width of 1st flagellomere; posterior process of gonopod with 

lateral margins convergent apically, pointed, at least 1/2 

width of surstylus on its narrowest point, devoid of setae.

Description. Coloration and vestiture: Vertex 

mostly densely microtomentose, with a shiny spot extended 

from ocellus to near eye margin (2/3 or more to eye margin); 

frons bearing a transverse, large, orange band, merging into 

dark brown ventrally; silver stripe on frons approximately 

straight; antenna yellow, infuscate with brown dorsally; face 

yellow in ground color; facial band present, sharply defined, 

transverse, over brown mark on protruding portion of face; 

FIGURE 165. Illustration of Aulacigaster africana Barraclough 

(male) (the leucopeza group, Afrotropical Region): gonopod, lateral 

aspect.
number 633 • 8 7

palpus white. Scutum with weakly defined microtomentose 

stripes; scutellum dull, microtomentose; postpronotum 

concolorous with mesonotum, scarcely microtomentose, 

subshiny; anepisternum mostly microtomentose with a 

semi-lunate, ventral polished area; katepisternum mostly 

polished; halter mostly white. Wing faintly brown infuscate 

in cells r1 and r2+3. Fore coxa brown to black, fore femur 

brown, fore tibia brown, fore tarsus brown to dark brown, 

basal tarsomere yellow; mid coxa brown to black, mid 

femur brown, mid tibia brownish, mid tarsus yellowish, 

apical and subapical tarsomere brown; hind coxa brown, 

hind femur brown, hind tibia brown to dark brown, hind 

tarsus yellowish, apical and subapical tarsomere brown. 

Abdomen with syntergite 1+2 white.

Morphology: 

Head: Head round (head ratio 1); face at level of 

pseudovibrissa wider than width of 1st flagellomere, in lateral view with dorsal half concave, ventral portion bulbous; 

gena in lateral view approximately 1/7 the width of 1st flagellomere; ocellar seta between 1/3–1/2 the length of medial 

vertical seta; medial vertical seta 3/4 length of lateral vertical seta; peristomal area with about 1 strong and 4 weaker 

setae following pseudovibrissal seta.

Thorax: Acrostichal setae in 2 rows. 

Male abdomen and terminalia: Figures 166–167. 

Surstylus arising from posteroventral margin of epandrium, 

slender, digitiform, relatively long (approximately 0.3–0.4 

times the length of ventral margin of epandrium), slightly 

MAP 14. Distribution of the Afrotropical species of Aulacigaster (the leucopeza group).
8 8 • smithsonian contributions to zoology

bulged medially, apex round; subepandrial sclerite with 

cerci fused on basal 1/2, lateral arms curved posteriorly and 

blunt apically; gonopod sub-rectangular in lateral view (approx. two times higher than long), with a pronounced dorsal “hunch” and with 1 medial seta and 3 setae preceding 

posterior process; posterior dorsal process of gonopod long 

(subequal in length to surstylus), with dorsal and ventral 

margins convergent to pointed apex.

Measurements: Body length 2.50–2.75 mm. Wing 

length 2.5–2.9 mm, wing width 0.9–1.1 mm.

Type Material. The holotype male is labeled “ZAMBIA: Nyika National Park[,] Chowo Forest[,] 

28.ix.1998[,] F. KAPLAN & A. FREIDBERG/HOLOTYPE 

♂ Aulacigaster freidbergi Rung & Mathis TAU [red].” The 

holotype is double mounted (minuten in a block of plastic), is in good condition (vertical, scutellar and mesonotal 

setae broken, wings slightly folded), and is deposited in the 

TAU.” Paratypes are as follows: same label data as the holotype (2♀; TAU, USNM).

Type Locality. Zambia. Nyika National Park 

(10°40¢S, 33°38¢E).

Distribution. (Map 14) Afrotropical: Zambia.

Etymology. The specific epithet, freidbergi, is 

a genitive patronym to honor and recognize the contributions of Dr. Amnon Freidberg to dipterology and in particular to this project. Amnon not only collected the type series 

of this species but numerous other specimens that gave genesis to this study.

Aulacigaster malawana, new species

Figures 168–169, Map 14

Diagnosis. This species is distinguished from 

congeners by the following combination of characters: Abdomen with syntergite 1+2 whitish; gonopod subrectangular, with one medial seta and three setae preceding posterior 

process; posterior process of gonopod digitiform, at least 3 

times wider than surstylus, bearing one central seta.

Description. Coloration and vestiture: Vertex 

mostly densely microtomentose, with a shiny spot extended 

from ocellus to near eye margin (2/3 or more to eye margin); 

frons bearing a transverse, large, orange band, merging into 

dark brown ventrally; silver stripe on frons approximately 

straight; antenna yellow, infuscate with brown dorsally; 

face yellow in ground color; facial band present, sharply defined, transverse, over brown mark on protruding portion 

of face; palpus white. Scutum with weakly defined microtomentose stripes; postpronotum concolorous with mesonotum, scarcely microtomentose, subshiny; anepisternum 

mostly microtomentose; katepisternum mostly polished; 

halter mostly white. Wing faintly brown infuscate in cells r1

and r2+3. Fore coxa brown to black, fore femur brown, fore 

tibia brown, fore tarsus brown to dark brown, basal tarsomere yellow; mid coxa brown to black, mid femur brown, 

mid tibia brownish, mid tarsus yellowish, apical and subapical tarsomere brown; hind coxa brown, hind femur 

FIGURES 166–167. Illustrations of Aulacigaster freidbergi, new species (male) (the leucopeza

group, Afrotropical Region). (166) gonopod, lateral aspect; (167) epandrium, lateral aspect.
number 633 • 8 9

brown, hind tibia brown to dark brown, hind tarsus yellowish, apical and subapical tarsomere brown. Abdomen 

with syntergite 1+2 white.

Morphology: 

Head: Head round (head ratio 1); face at level of 

pseudovibrissa wider than width of 1st flagellomere, in lateral view with dorsal half concave, ventral portion bulbous; 

gena in lateral view approximately 1/7 the width of 1st flagellomere; ocellar seta between 1/3–1/2 the length of medial 

vertical seta medial vertical seta 3/4 length of lateral vertical seta; peristomal area with about 1 strong and 4 weaker 

setae following pseudovibrissal seta.

Thorax: Acrostichal setae in 2 rows.

Male abdomen and terminalia: Figures 168–169. 

Surstylus arising from posteroventral margin of epandrium, 

slender, digitiform, relatively long (approximately 0.2–0.3 

times the length of ventral margin of epandrium), apex 

round; subepandrial sclerite with lateral arms curved posteriorly and blunt apically; cerci fused on basal 1/2; gonopod 

with 1 medial seta and 3 setae preceding posterior process, 

sub-rectangular in lateral view (approx. 2 times higher 

than long); posterior process of gonopod digitiform, long 

(slightly longer than surstylus) and wide (at least 3 times 

wider than surstylus), bearing one central seta.

Measurements: Body length 2.2–2.5 mm. Wing 

length 2.5–2.7 mm, wing width 0.9–1.1 mm.

Type Material. The holotype male is labeled 

“MALAWI: North Nyika National Park[,] Zovo-Chipolo 

Forest[,] 25-16.ix.1998, F. KAPLAN & A. FREIDBERG/

HOLOTYPE ♂ Aulacigaster malawana Rung & Mathis 

TAU [red].” The holotype is double mounted (minuten in 

a block of plastic), is in good condition (vertical, scutellar 

and mesonotal setae broken, wings slightly folded), and is 

deposited in the TAU.” Paratypes are as follows: same label 

data as the holotype (5♂, 4♀; TAU, USNM).

Type Locality. Malawi. North Malawi, Nyika 

National Park, Zovo-Chipolo Forest (10°48¢S, 33°48¢E).

Distribution. (Map 14) Afrotropical: Malawi.

Etymology. The specific epithet, malawana,

refers to the country where the type series was collected.

Aulacigaster perata Barraclough

Map 14

Aulacigaster perata Barraclough 1993:39–40.

Diagnosis. This species is distinguished from 

congeners by the following combination of characters: Abdomen completely brown; gena in lateral view, immediately 

below ventral margin of eye, approximately 1/5 width of 

1st flagellomere.

Description. Coloration and vestiture: Vertex 

mostly densely microtomentose, with a shiny spot extended 

from ocellus at most 1/2–2/3 to eye margin; frons bearing a 

transverse, large, orange band, frontal orange band merging 

into dark brown ventrally; silver stripe on frons; antenna pale 

yellow to brownish; face yellow in ground color; facial band 

present, sharply defined, over brown mark on protruding 

portion of face; palpus white. Scutum dull microtomentose, 

devoid of microtomentum on anteriormost portion; scutellum mostly microtomentose; postpronotum concolorous 

with mesonotum, uniformly microtomentose anepisternum 

mostly dull; katepisternum mostly polished; halter mostly 

yellowish, or mostly brown to dark brown (knob darker). 

Wing mostly brown, infuscate, not noticeably darker along 

costal margin and apex. Fore coxa brownish (“yellowbrown to medium brown”: Barraclough, 1993:39), fore 

femur brown, fore tibia brown, fore tarsus yellowish, apical 

and sometimes subapical tarsomeres brown to black; mid 

coxa yellowish to brownish (“paler, entirely yellow-brown: 

Barraclough, 1993:39); mid femur brown, mid tibia brownish, mid tarsus yellowish, apical and subapical tarsomere 

brownish to dark brown; hind coxa brownish (“paler, entirely yellow-brown: Barraclough, 1993:39), hind femur 

brown, hind tibia brown to dark brown, hind tarsus yellowish apical and subapical tarsomere brownish to dark brown.

Morphology: 

Head: Head higher than long (head ratio less than 

0.9); face in lateral view with dorsal half concave, ventral 

portion bulbous; gena in lateral view approximately 1/4–

1/5 the width of 1st flagellomere; medial vertical seta 3/4 

FIGURES 168–169. Illustrations of Aulacigaster malawana, new 

species (male) (the leucopeza group, Afrotropical Region). (168) 

gonopod, lateral aspect; (169) epandrium, lateral aspect.
9 0 • smithsonian contributions to zoology

length of lateral vertical seta; peristomal area with about 1 

strong and 4 weaker setae following pseudovibrissal seta.

Type Material. The holotype female is labeled “CAMEROON[.] 50Km E. Bamenda[,] off Rt.N11 

Bamballang Area[,] 21.XI.1987[,] FINI KAPLAN/HOLOTYPE ♀ Aulacigaster perata BARRACLOUGH.” The holotype is deposited in TAU.

Type Locality. Cameroon. Nord-Ouest: 

Bamenda (50 km E), Bamballang Area (off Rt. N11; 

05°53¢N, 10°23¢E).

Distribution. (Map 14) Afrotropical: 

Cameroon.

Remarks. The description herein presented is 

based on Barraclough (1993). The male is not known for 

this species. According with Barraclough (1993), A. perata

was collected at approximately 1500 m altitude.

Nearctic Species

Discussion. The Nearctic fauna includes 

three described species. One species, A. neoleucopeza

Mathis and Freidberg, was also reported from the Palearctic Region (see below) by Papp (1998a), but this record was contested by Kassebeer (2001) (see Palearctic 

Species). Mathis and Freidberg (1994) provided a key to 

the Nearctic species, which is repeated below with minor 

modifications, plus distributional data, descriptions, and 

illustrations.

Key to the Nearctic Species of Aulacigaster

1. Species larger (wing length averaging 2.6 mm), darker, and shinier; shiny spot on vertex small, at most extended from 

ocellus 1/2 way to eye margin, sometimes indistinct (fig.17; Mathis and Freidberg, 1994:594); anepisternum with ventral 

1/3 shiny, bare of microtomentum, bordered dorsally by dense gray microtomentum; katepisternum, coxae, femora, and 

abdomen strongly shiny, lacking microtomentum, although fore femur sparsely microtomentose anteriorly; acrostichal 

setulae in 2 rows along entire length (fig. 18; Mathis and Freidberg, 1994:594); 1st flagellomere usually blackened on 

dorsal 2/3. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. mcalpinei Mathis and Freidberg

Smaller and duller species (wing length averaging 2.35 mm); shiny spot on vertex larger and distinct, typically extended 

from ocellus more than 1/2 way distance to eye margin (fig. 3; Mathis and Freidberg, 1994:587); anepisternum usually entirely microtomentose, rarely with a bare, shiny stripe ventrally; katepisternum, coxae, femora, and abdomen not strongly 

shiny, at least partially covered with microtomentum; acrostichal setulae in a single medial row anteriorly, row bifurcate 

at posterior 1/2–2/5 (fig.15; Mathis and Freidberg, 1994:587); 1st flagellomere with only dorsal 1/4 darkened . . . . . . 2

2. Surstylus moderately long (0.3–0.5 times the length of basal margin of epandrium) [Figures 173, 175, 177, 179, 182]; 

gonopod in lateral view with a lobe-like posterodorsal process [Figures 172, 174, 176, 178] . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. neoleucopeza Mathis and Freidberg

Surstylus short (less than 0.2 times the length of basal margin of epandrium; Figure 184); gonopod in lateral view with a 

distinctly pointed process posterodorsally [Figure 183] . . . . . . . . . . . . . . . . . . . . . . . A. sabroskyi Mathis and Freidberg

Aulacigaster mcalpinei Mathis and Freidberg

Figures 170–171, Map 15

Aulacigaster mcalpinei Mathis and Freidberg, 1994:593–597 [illustration of 

head, thorax, male terminalia and spermatheca].

Diagnosis. This species is distinguished from 

congeners by the following combination of characters: Vertex with a shiny spot extended from ocellus at most 1/2 way 

to eye margin; surstylus broad, beak-like, wider than width 

of apical portion of phallapodeme.

Description. Coloration and vestiture: Vertex mostly densely microtomentose, with a shiny spot 

extended from ocellus at most 1/2 way to eye margin; ocellar tubercle dull microtomentose; frons bearing a transverse, large, orange band, frontal orange band merging into 

dark brown ventrally; silver stripe on frons approximately 

straight; antenna pale yellow to yellowish, infuscate along 

dorsal margin (to dorsal 2/3); face yellow, with a transverse brown mark on dorsal portion; facial band present, 

sharply defined, transverse, on protruding, ventral portion 

of face; palpus white. Scutum with weakly defined microtomentose stripes; scutellum dull microtomentose; postpronotum concolorous with mesonotum; anepisternum with 

ventral 1/3 shiny, bare of microtomentum, bordered dorsally by dense gray microtomentum; katepisternum mostly 

polished; halter mostly white (knob mostly dark brown 
number 633 • 9 1

anteriorly). Fore coxa brown to black, fore femur brown 

to black, fore tibia yellowish, fore tarsus yellowish, apical 

and sometimes subapical tarsomeres brown to black; mid 

coxa brown to black, mid femur brown to black, mid tibia 

brown to black, mid tarsus yellowish, apical and subapical tarsomere brownish to dark brown; hind coxa brown 

to black, hind femur brown to black, hind tibia brown to 

black, hind tarsus yellowish, apical tarsomere brown to 

black. Male abdomen subshiny to dull; female abdomen 

with dense microtomentum medially, lateral margins of 

tergites nearly polished.

Morphology: 

Head: Head higher than long (head ratio less than 

0.9); face at level of pseudovibrissa wider than width of 

1st flagellomere, in lateral view with dorsal half concave, 

ventral portion bulbous; gena in lateral view approximately 

1/3–1/2 the width of 1st flagellomere; medial vertical seta 

3/4 length of lateral vertical seta; 1 long and 3–4 shorter 

peristomal setae following pseudovibrissal setae present.

Thorax: Acrostichal setae in 2 rows; scutellum 

slightly raised (angle with scutum approximately 135°), 

trapezoidal, apex nearly straight, disk of scutellum slightly 

convex; basal scutellar seta 1/2 length of posterior seta.

Male abdomen and terminalia: Figures 170–171. 

Surstylus arising from posteroventral margin of epandrium, 

broad basally, shaped as a beak, relatively long; subepandrial sclerite with cerci mostly fused, lateral arms curved 

posteriorly and pointed apically (fig. 22; Mathis and Freidberg, 1994:595); gonopod (fig. 21; Mathis and Freidberg, 

1994:595) in lateral view with pointed process preceded by 

a few small setae, shallow ventrally (about as long as high), 

approximately rectangular in lateral view.

Female abdomen and terminalia: Spermathecae 

with ventral digitiform projections.

Measurements: Body length 2.5–2.6 mm. Wing 

length 2.5–2.6 mm, wing width 0.9 mm.

Type Material. The holotype male is labeled “USA. M[arylan]D: Montgomery Co., Potomac[,] 

23 Jul–23 Aug 1989[,] Amnon Freidberg[,] slime flux on 

oak/HOLOTYPE ♂ Aulacigaster mcalpinei W. N. Mathis 

& Freidberg USNM [red; “♂ Aulacigaster mcalpinei” and 

“Freidberg” handwritten].” Paratypes: Same locality data 

as the holotype (17♂, 15♀; TAU, USNM). Connecticut. 

Fairfield: Redding, 31 Mar 1929, A. L. Melander (4♂, 4♀; 

USNM). New Hampshire. Hillsborough: Hollis (Beaver 

Brook Association), 20 Apr 1980, N. E. Woodley (14♂, 

8♀; NEW, USNM). Virginia. Fairfax: Dead Run (38°58¢N, 

77°10.3¢W; at sap tree, at sap sugar maple), 8 Mar–28 Jul 

1914, 1915, 1916, 1925, R. C. and E. M. Shannon (17♂, 

21♀; USNM).

Type Locality. United States. Maryland. 

Montgomery: Potomac (39°01.1¢N, 77°12.5¢W).

Other Specimens Examined. CANADA. 

Ontario: Ottawa (45°25¢N, 75°45¢W), 27 Apr–1 May 

1955, 1983, D. G. F. Cobb, J. R. Vockeroth (3♂, 2♀; CNC; 

Ottawa, Rockcliffe, McKay Lake (45°27.1¢N, 75°40.3¢W), 

27–28 Apr 1955, 1983, D. G. F. Cobb, J. F. McAlpine (2♂, 

5♀; CNC).

FIGURES 170–171. Illustrations of Aulacigaster mcalpinei Mathis and Freidberg (male) (the leucopeza group, Nearctic Region). (170) gonopod, lateral aspect; (171) epandrium, lateral aspect.
9 2 • smithsonian contributions to zoology

UNITED STATES. Arkansas. Garland: Hot Springs 

National Park (34°31¢N, 93°03.2¢W), 29 Aug–13 Sep 

1943, B. C. Marshall (2♀; USNM). Logan: Magazine 

Mountain (35°10¢N, 93°38.7¢W; oak sap flux), 6 Jul 

1992, D. A. Grimaldi (10♂, 6♀; AMNH, USNM).

District of Columbia. Washington (38°49.8¢N, 

77°00.6¢W), 3 Mar–4 Aug 1927, H. S. Barber (1♂, 1♀; 

USNM).

Florida. Alachua: Gainesville (29°39.1¢N, 82°19.5¢W), 

22 May 1957, H. V. Weems (3♂; USNM). Highlands: 

Archbold Biological Station (27°11.3¢N, 81°20.3¢W; at 

sap flow from ambrosia beetle hole on dying Quercus laurifolia), 27 May 2002, M. Deyrup (6♂; ABSF).

Illinois. Cook: Chicago (41°47.4¢N, 87°36¢W) (1♀; 

USNM). Macoupin: Carlinville (39°16.8¢N, 89°52.9¢W), 

Robertson (1♂; USNM).

Indiana. Dubois: Ireland (38°24.9¢N, 86°60¢W; in 

trap), 17 Jul 1965, O. Mund (1♂; USNM).

Maryland. Montgomery: Colesville (39°04.8¢N, 

77°00.1¢W; Malaise trap), 26 Jun 1977, W. W. Wirth 

(1♀; USNM); Glen Echo (38°58.1¢N, 77°08.4¢W), 23 

Jul–6 Aug 1922, J. R. Malloch (1♂, 2♀; USNM); Plummers Island (38°58.2¢N, 77°10.6¢W), 25 Mar–18 Aug 

1912, 1914, 1916, W. L. McAtee, R. C. Shannon (5♂, 

3♀; USNM). Prince Georges: College Park (38°58.8¢N, 

76°56.2¢W), 29 Jun 1933, C. T. Greene (1♂; USNM).

Massachusetts. Middlesex: Cambridge (42°22.5¢N, 

71°06.3¢W), 7–16 Jul 1981, N. E. Woodley (3♀; USNM). 

Norfolk: Boston (42°21.5¢N, 71°03.6¢W; Arnold Arboretum), 18 Apr 1980, N. E. Woodley (3♂, 1♀; USNM).

Michigan. Livingston: Edwin S. George Reserve 

(42°27¢N, 83°01¢W), 16 Apr 1950, K. Bohnsack (1♀; 

USNM). Washtenaw: Ann Arbor (42°16.3¢N, 83°43.6¢W), 

30 Apr 1936, G. C. Steyskal (1♀; USNM). Wayne: Detroit 

(42°19.9¢N, 83°02.8¢W), 29 Aug 1943, G. C. Steyskal 

(1♀; USNM); Grosse Isle (42°07.7¢N, 83°08.7¢W), 2 Apr 

1957, G. C. Steyskal (1♀; USNM).

Minnesota. Olmstead (43°53.6¢N, 92°10.2¢W): 

C. Ainslio (1♀; CNC).

New Jersey. Bergen: Ridgewood (40°58.8¢N, 

74°07¢W), Apr 1987, A. Soll (2♀; AMNH). Mercer: Princeton (40°20.9¢N, 74°39.6¢W), 14 Apr 1916 (1♀; USNM).

New York. Broome: Chenango Valley State Park 

(42°12.8¢N, 75°50.5¢W), 15 Apr–16 May 1982, D. A. 

Grimaldi (1♂; AMNH). Tomkins: Ithaca (42°26.4¢N, 

76°29.8¢W), 15 Apr 1913, H. Morrison (2♀; USNM).

North Carolina. Mecklenburg: Highway 51 (1.6 km 

W of Rt. 16, near Matthews; 35°07¢N, 80°43.4¢W; Magnolia cone and flower of Magnolia grandiflora), 12–23 

May 1979, 1981, A. G. Wheeler, Jr. (1♂; PDA).

Pennsylvania. Dauphin: Harrisburg (40°16.4¢N, 

76°53.1¢W; sap flux on Ulmus), 15 Apr–19 Jul 1982, 

1985, K. R. Valley (2♂, 1♀; PDA). Schuylkill: Interstate 

81 (4.2 km S at exit 31, Rt. 433; 40°32.1¢N, 76°25.7¢W; 

sap flux on Betula), 29 Apr 1986, K. R. Valley (1♂; PDA).

Virginia. Fairfax: Mt. Vernon (38°42.5¢N, 

77°05.2¢W), 8 Jul–22 Aug 1956, 1976, P. H. Arnaud, Jr. 

(2♂; CAS, USNM). Alexandria, Maywood (38°53.8¢N, 

77°06.3¢W; McAtee’s residence; at oak sap), 4 May 1916, 

W. L. McAtee (1♂; USNM). Falls Church, Holmes Run 

(38°50.9¢N, 77°10.4¢W; at light), 14 Jun–23 Aug 1961, 

W. W. Wirth (1♂, 1♀; USNM). Scott Run, Stubblefield 

Falls (38°58.1¢N, 77°12.2¢W; on Pinus virginiana), J. R. 

Malloch (1♂, 1♀; USNM). Page: Big Meadows (38°31¢N, 

78°26.4¢W), 15 Jun 1941, A. L. Melander (1♀; USNM).

Distribution. (Map 15) Eastern North 

America: Canada (Ontario), United States (Arkansas, Connecticut, District of Columbia, Florida, Illinois, Indiana, 

Massachusetts, Maryland, Michigan, Minnesota, New 

Hampshire, New York, New Jersey, North Carolina, Pennsylvania, Virginia).

Biology. This species has been collected on tree 

wounds and fluxes of the following genera of deciduous 

trees: Acer, Pinus, and Quercus.

Aulacigaster neoleucopeza Mathis and 

Freidberg

Figures 172–182, Map 16

Aulacigaster neoleucopeza Mathis and Freidberg, 1994:586–591 [illustration 

of head, thorax, and male terminalia].—Papp, 1998a:229, 230–232 

[comparison with A. leucopeza, illustrations of head and male terminalia], 233 [key to Palearctic species of Aulacigaster]; 1998b:280–284 

[Manual of Palearctic Diptera].—Bächli et al., 1999:121 [checklist, 

Switzerland].

Diagnosis. This species is distinguished from 

congeners by the following combination of characters: 

Posterior fronto-orbital seta slightly internal to and almost horizontally aligned with anterior seta; scutum with 

weakly defined microtomentose stripes; surstylus moderately long, about 1/3 length of basal margin of epandrium 

in lateral view.

Description. Coloration and vestiture: Vertex 

mostly densely microtomentose, with a shiny spot extended 

from ocellus to 1/2–3/4 distance to eye margin; ocellar 

tubercle dull microtomentose; frons bearing a transverse, 

large, orange band merging into dark brown ventrally; silver stripe on frons approximately straight; antenna pale 
number 633 • 9 3

yellow to yellowish, infuscate along dorsal margin; face 

yellow, with a transverse brown mark on dorsal portion; 

facial band present, sharply defined, transverse, on protruding, ventral portion of face; palpus white. Scutum with 

weakly defined microtomentose stripes; scutellum dull 

microtomentose; postpronotum usually concolorous with 

mesonotum (lighter than mesonotum in more teneral specimens), dull microtomentose; anepisternum mostly dull, 

often with a semi-lunate, ventral, strongly microtomentose 

stripe; katepisternum typically with anterior portion dull 

microtomentose, and posterior portion mostly polished 

(katepisternum mostly covered with microtomentum in 

some specimens); halter mostly white (knob and base with 

brownish marks). Fore coxa brown to black, fore femur 

brown to black, apex often yellowish; fore tibia brown to 

black; fore tarsus yellowish, apical and sometimes subapical tarsomeres brown to black; mid coxa brown to black, 

mid femur brown to black, apex often yellowish; mid tarsus 

yellowish, apical and subapical tarsomere brownish to dark 

brown; hind coxa brown to black, hind femur brown to 

black, apex often yellowish; hind tarsus yellowish, apical 

tarsomere brown to black (legs dark yellow to pale brown 

in more teneral specimens). Male abdomen subshiny to 

dull; female abdomen with dense microtomentum medially, 

lateral margins of tergites nearly polished.

Morphology: 

Head: Head higher than long (head ratio less than 

0.9); face at level of pseudovibrissa wider than width of 

1st flagellomere, in lateral view with dorsal half concave, 

ventral portion bulbous; gena in lateral view approximately 

MAP 15. Distribution of Aulacigaster mcalpinei (the leucopeza group, Nearctic Region).
9 4 • smithsonian contributions to zoology

FIGURES 172–177. Illustrations of the lateral aspect of the gonopod (left hand side) and epandrium 

(right hand side) of males of Aulacigaster neoleucopeza Mathis and Freidberg (the leucopeza group, 

Nearctic Region) from various localities. (172) paratype, gonopod, lateral aspect (Montgomery 

County, Maryland); (173) paratype, epandrium, lateral aspect (Montgomery County, Maryland); 

(174) gonopod, lateral aspect (El Dorado County, California); (175) epandrium, lateral aspect (El 

Dorado County, California); (176) gonopod, lateral aspect (Sacramento County, California); (177) 

epandrium, lateral aspect (Sacramento County, California). Not all to the same scale.
number 633 • 9 5

1/3–1/2 the width of 1st flagellomere; medial vertical seta 

3/4 length of lateral vertical seta; 1 long and 3–4 shorter 

peristomal setae following pseudovibrissal setae present.

Thorax: Acrostichal setae in 1 row, bifurcating 

posteriorly; scutellum slightly raised (angle with scutum approximately 135°), trapezoidal, apex nearly straight, disk 

of scutellum slightly convex; basal scutellar seta less than 

1/2 length of posterior seta or 1/2 length of posterior seta.

Male abdomen and terminalia: Figures 172–182. 

Surstylus arising from posteroventral margin of epandrium, 

slender, digitiform, relatively long (0.3–0.5 times length of 

basal margin of epandrium), tapered toward a round to 

slightly blunt apex; surstylus in lateral view with or without 

apex slightly elevated in relation to the basal margin of epandrium; subepandrial sclerite with lateral arms well sclerotized, curved posterolaterally, with slight recurved apices; 

cerci generally with deep, narrow cleft, fused only on basal 

1/4 (fig. 9, Mathis and Freidberg, 1994: 588); gonopod 

with posterodorsal lobe bearing basal elongate setae, shallow ventrally (about 0.8–0.9 times as long as high), approximately rectangular to tongue-shaped in lateral view.

Female abdomen and terminalia: Spermathecae 

with ventral digitiform projections.

Measurements: Body length 1.63–3.0 mm. Wing 

length 2.0–3.1 mm, wing width 0.8–1.0 mm.

Type Material. The holotype male is labeled 

“USA. M[arylan]D: Montgomery Co., Potomac[,] 23 

Jul–23 Aug 1989[,] Amnon Freidberg[,] slime flux on oak/

HOLOTYPE ♂ Aulacigaster neoleucopeza W. N. Mathis 

& Freidberg USNM [red; “♂ Aulacigaster neoleucopeza” 

and “Freidberg” handwritten].” Paratypes: Same locality 

data as the holotype (22♂, 9♀; TAU, USNM). UNITED 

STATES. Maryland. Prince Georges: Camp Springs 

(38°48.2¢N, 76°54.4¢W), 1 Apr 1979, G. F. Hevel (16♂, 7♀; 

USNM). Virginia. Stafford: Stafford (6.4 km N; 38°32.8¢N, 

77°24.5¢W), 14 Aug 1990, W. N. Mathis (3♂; USNM).

Type Locality. United States. Maryland. 

Montgomery: Potomac (39°01.1¢N, 77°12.5¢W).

Other Specimens Examined. CANADA. 

Alberta. Edmonton (53°33¢N, 113°30¢W), 29 Apr 1924, 

O. Bryant (6♂, 4♀; CAS, USNM); Elkwater (49°42¢N, 

110°16¢W), 29 May 1955, J. R. Vockeroth (1♀; CNC).

British Columbia. Remo (11.3 km SW Terrace; 

54°29.4¢N, 128°43.7¢W; flowers of Heracleum), 6–13 Jun 

1960, J. G. Chillcott (1♂, 1♀; CNC).

Manitoba. Ninette (49°25.1.1¢N, 99°38¢W; flood 

plain community; bleeding stumps of Acer negundo), 

5 May 1958, J. F. McAlpine (2♂, 2♀; CNC).

New Brunswick. Kouchibouguac National Park 

(46°51¢N, 64°58¢W), 17 May 1977, B. Copper (1♀; CNC).

Ontario. Carp (45°21.8¢N, 76°02¢W; bleeding maple 

stump), 22 Apr 1954, J. F. McAlpine (2♂; CNC); Maynooth (45°12.8¢N, 77°57¢W), 22 Jun 1953, J. F. McAlpine (1♂, 1♀; CNC); Midland (44°45¢N, 79°53.4¢W), 20 

Aug 1955, J. G. Chillcott (1♀; CNC); Ottawa (45°25¢N, 

75°42¢W), 18 Apr–18 Jun 1951, 1983, D. G. F. Cobb, J. F. 

McAlpine, J. R. Vockeroth (4♂, 4♀; CNC).

Quebec. Duncan Lake near Rupert (75°46¢N, 

47°34¢W), 31 Jul 1971, J. F. McAlpine (1♀; CNC); La 

Ferme (46°6.8¢N, 76°03¢W), 11 Jul 1913, A. Robert (1♀; 

CNC); Old Chelsea (45°30¢N, 75°49¢W), 16 Apr–3 May 

1939, 1958, G. S. Walley, G. E. Shewell, J. R. Vockeroth 

(1♂, 2♀; CNC).

Saskatchewan. Maple Creek (49°55¢N, 109°29¢W), 

28 May 1955, J. R. Vockeroth (1♂; CNC).

MEXICO. Baja California: Tecate (32°34¢N, 

116°38¢W; ex Populus ooze), 16 Jun 1964, R. Duke (1♂, 

1♀; USNM).

Baja California Sur: San José del Cabo (23°09¢N, 

109°42¢W; reared from cottonwood ooze), 6 Nov 1953, 

R. E. Ryckman, C. C. Lindt (4♂, 1♀; USNM).

UNITED STATES. California. Calaveras: Calaveras 

Big Trees State Parks (38°15¢N, 120°15¢W; 4219 feet; banana trap), 8–15 Jul 2005, A. R. Cline (6♂, 9♀; CAS). 

El Dorado: Blodgett Forest (38°53.2¢N, 120°38.9¢W; 

banana bait), 30 Apr 1981, H. Spleth (2♂; CAD). Kern: 

Kern Canyon (35°27.6¢N, 118°46.7¢W), Apr 1934, A. H. 

Sturtevant (1♂, 2♀; USNM); Rosamond (34°51.9¢N, 

118°09.8¢W), 17 Oct 1956, A. H. Sturtevant (2♀; USNM); 

Tupman (6.4 km E; 35°17.9¢N, 119°21¢W; Kern River; 

Populus slime flux), 1 Jul 1984, E. M. Fisher (1♂, 1♀; 

CAS). Los Angeles: Glendora (34°08.2¢N, 117°51.9¢W), 

15 Mar 1929, A. H. Sturtevant (1♀; USNM); Mount Wilson, San Gabriel Mountains (34°17.3¢N, 117°38.8¢W; 

1675 m), 1 Oct 1980, E. Fisher (1♀; CSA). Napa: Moskowite Corner (5.5 km NW; 38°27.5¢N, 122°12.3¢W; 

200 m), Capell Creek (38°45.9¢N, 122°12.6¢W), 4 Sep 

1977, P. H. Arnaud, Jr. (1♂, 1♀; CAS). Orange: San Juan 

Canyon (33°33.3¢N, 117°33.1¢W), 22 Jul 1953, A. H. 

Sturtevant (2♀; USNM). Riverside: Mecca, Cottonwood 

Spring (33°34.3¢N, 116°04.6¢W), May 1939, A. H. 

Sturtevant (2♀; USNM). Sacramento: Sacramento River 

(pocket area; 38°18.6¢N, 121°39.2¢W; ex. Populus), 20 

Jun 2008, A. Rung, M. Hauser (9♂, 3♀; CAS). San Bernardino: Redlands (35°03.3¢N, 117°10.9¢W), 27 May 

1923, F. R. Cole (1♂, 1♀; USNM); South Fork Santa Ana 

River (34°10.3¢N, 116°49.8¢W), 2 Jul 1968, P. H. Arnaud, Jr. (1♀; CAS); 29 Palms (34°08.1¢N, 116°03.3¢W; 

Oasis Nature Area), 13 Jan 1953, P. H. Arnaud, Jr. (1♂; 

USNM). San Diego: Alpine (32°50.1¢N, 116°46¢W; ex. 
9 6 • smithsonian contributions to zoology

FIGURES 178–182. Illustrations of Aulacigaster neoleucopeza Mathis and Freidberg (male) (the leucopeza group, Nearctic Region) from two localities. Figures 178–181 (Calaveras County, California): (178) internal male genitalia, lateral 

aspect; (179) epandrium, lateral aspect; (180) internal male genitalia, ventral aspect; (181) internal male genitalia, posterior aspect. Figure 182 (Grafton County, New Hampshire): epandrium, lateral aspect. Not all to the same scale.
number 633 • 9 7

Rhus ovata), 12 May 1959, Moffet (1♀; USNM); Palomar Mountain (33°19.4¢N, 116°52.7¢W; 1615 m), 23 Jul 

1953, A. H. Sturtevant (3♂, 1♀; USNM). Santa Clara: 

Palo Alto (37°26.7¢N, 122°09.6¢W), 14 Jun–17 Jul 

1921, A. H. Sturtevant (4♂; USNM); Stanford University 

(37°25.6¢N, 122°10.2¢W), 7 May 1961, P. H. Arnaud, Jr. 

(1♀; CAS).

Connecticut. Fairfield: Redding (41°18.1¢N, 

73°23¢W), 31 Mar 1929, A. L. Melander (1♀; USNM).

District of Columbia. Washington (38°49.8¢N, 

77°00.6¢W). 19 Jun–17 Aug, 1913, 1944 (2♂, 1♀; 

USNM).

Idaho. Bear Lake: Montpelier (3.2 km E; 42°19.3¢N, 

111°17.9¢W), 15 Aug 1981, P. H. Arnaud, Jr. (1♂; CAS).

Illinois. Champaign: Champaign (40°07¢N, 

88°14.6¢W; bleeding elm), 14 Oct 1956, J. F. McAlpine 

(2♂; CNC); Urbana (40°06.6¢N, 88°12.4¢W; at light), 

20 Apr 1957, J. F. McAlpine (1♂; CNC). Crawford: Flat 

Rock (38°54.1¢N, 87°40.3¢W), 1915 (1♂, 3♀; USNM).

Indiana. Tippecanoe: Lafayette (40°25¢N, 86°52.5¢W), 

26 Jul 1916, J. M. Aldrich (1♀; USNM).

Iowa. Story: Ames (42°02.1¢N, 93°37.2¢W), 1 

Apr 1918 (1♀; USNM). Webster: Lehigh (42°21.6¢N, 

94°03.1¢W), 15 Apr 1955, D. M. Norris (1♀; USNM).

Kansas. Douglas: Lawrence (38°58.3¢N, 95°14.1¢W), 

22 Jun 1922, C. H. Curran (1♂, 2♀; CNC).

Maryland. Montgomery: Bethesda (38°58.8¢N, 

77°06¢W), 27 Jul 1965, G. C. Steyskal (4♂, 3♀; USNM); 

Colesville (39°04.8¢N, 77°00.1¢W), 14 Jun 1977, W. W. 

Wirth (1♀; USNM); Glen Echo (38°58.1¢N, 77°08.4¢W), 

6 Aug 1922, J. R. Malloch (1♂; USNM); Plummers Island 

(38°58.2¢N, 77°10.6¢W), 3 Aug 1912 (8♂, 5♀; USNM). 

Prince Georges: Marlboro, May 1913, H. S. Barber (1♀; 

USNM).

Massachusetts. Barnstable: Woods Hole (41°31.5¢N, 

70°40.3¢W), Jul–20 Sep 1922, A. H. Sturtevant (7♂, 8♀; 

USNM). Dukes: Naushon Island (41°29¢N, 70°45.5¢W), 

30 Jul 1922, A. H. Sturtevant (1♀; USNM). Middlesex: 

Cambridge (42°22.5¢N, 71°06.3¢W), 15 Jul 1981, N. E. 

Woodley (1♀; USNM).

Michigan. Wayne: Detroit (42°19.9¢N, 83°02.8¢W), 

13 Apr–18 Jul 1938, 1942, 1943, G. C. Steyskal (4♂, 

2♀; USNM); Grosse Isle (42°07.7¢N, 83°08.7¢W), 11 Jul 

1948, G. C. Steyskal (1♀; USNM).

Minnesota. Clearwater: Itasca State Park (47°11.9¢N, 

95°12.1¢W), 15 Jul 1952 (1♂, 1♀; AMNH).

Nevada. Washoe: Verdi, Crystal Peak Park (39°30.9¢N, 

119°59.7¢W; 1480 m), 28 Jul 1973, P. H. Arnaud, Jr. (1♂; 

CAS).

New Hampshire. Grafton: Hanover (43°42.1¢N, 

72°17.4¢W), 31 Aug 1916, A. H. Sturtevant (1♂; USNM). 

Hillsborough: Nashua, Long Hill (42°43¢N; 71°27.1¢W; 

20 m; Malaise trap), 27 Jun 2004, S. D. and A. V. Gaimari 

(1♀; CAS).

New Jersey. Morris: Morristown (40°47.8¢N, 

74°28.9¢W), 9 Apr 1922, A. H. Sturtevant (2♀; USNM).

New Mexico. Otero: Cloudcroft (32°57.4¢N, 

104°44.5¢W; Pinus ponderosa), E. J. Hay (1♂; USNM); 

Lincoln National Forest, Sacramento Mountains 

(32°50¢N, 105°48¢W; 1795 m; Malaise trap), 13–15 Jun 

2007, S. D. Gaimari (1♂; CAS).

New York. Broome: Chenango Valley State Park 

(42°12.8¢N, 75°50.5¢W), 15 Apr–6 May 1982, D. A. 

Grimaldi (1♂, 2♀; AMNH). Tompkins: Brown Farm, Caroline Center (1 km S; maple sap flux), 19 Apr 1986 (1♀; 

AMNH); Ithaca (42°26.6¢N, 76°30.2¢W), 15 Apr 1913, 

H. Morrison (2♀; USNM); Trumansburg (42°32.5¢N, 

76°40¢W; maple sap flux), 16–22 Jun 1983, D. A. Grimaldi 

(1♂; AMNH).

Oregon. Jackson: Trail (42°38.9¢N, 122°48.6¢W), 20 

Aug 1951, A. H. Sturtevant (2♂; USNM). Union: Lick 

Creek (45°07.8¢N, 117°37.8¢W; 45 km SE Union; 1305 

m; sap flow), 23 Apr 1977, R. S. Zack (1♂; USNM). Wallowa: Elgin (45°35¢N, 117°55¢W; 35.4 km N; Alnus seep), 

20 May 1977, E. S. Davis (1♀; USNM).

Pennsylvania. Dauphin: Harrisburg (40°16.4¢N, 

76°53.1¢W; sap flux on Ulmus), 15 Apr–19 Jul, K. R. Valley (3♂; PDA). Philadelphia: Roxborough (40°02.6¢N, 

75°13.3¢W), 30 Apr 1910 (2♂; USNM).

South Dakota. Brule: Chamberlain (43°48.6¢N, 

99°19.8¢W), 25 Jun 1948, A. H. Sturtevant (1♂, 3♀; 

USNM).

Utah. Cache: Logan Canyon, Bunchgrass Creek 

(41°53.5¢N, 111°33.9¢W), 8–19 Aug 1986, W. J. Hanson, G. Knowlton (1♀; USU); Turner Campground 

(41°53.1¢N, 111°34.4¢W), 10–20 Jun 1985, W. J. Hanson (3♂, 4♀; USU), Mendon Cold Spring (41°42.6¢N, 

111°58.7¢W), Jun–Jul 1977, W. J. Hanson, G. Knowlton (2♂, 9♀; USU); West Hodges Canyon (41°55¢N, 

111°25¢W; Malaise trap), 6–13 Jun 1980, Jul 1978 (1♂, 

3♀; USU). Garfield: Capitol Reef National Park, Sulphur Creek (38°17.5¢N, 111°15¢W; Malaise trap), 28 Jul 

1978, Lindhal (2♀; USU). Juab: Mountain Nebo, Red 

Creek (39°45.7¢N, 111°42.7¢W; Malaise trap), 4–19 Aug 

1977 (1♂, 1♀; USU). Kane: Chamb Ranch (37°22.7¢N, 

112°51.3¢W), 24 Jul 1981, C. R. Nelson (1♀; USU). Utah: 

Provo (40°14.7¢N, 111°38.7¢W; Malaise trap), Jun–

Jul 1985 (2♀; USU). Uintah: Bonanza (SW; 40°01.3¢N, 
9 8 • smithsonian contributions to zoology

109°10.6¢W; 1525–1710 m; TIOS R24E), 15 May 1981 

(1♀; USU). Washington: Beaver Dam Wash, Lytle Ranch 

Pres. (38°53.7¢N, 113°55.3¢W), 6–19 Aug 1987, W. J. 

Hanson (2♀; USU); Leeds Canyon (37°13¢N, 113°24¢W), 

13–16 Sep 1984, W. J. Hanson, Joussef (1♂; USU); Zion 

National Park (37°18¢N, 113°03¢W), 7 Jun 1934, A. H. 

Sturtevant (3♂, 1♀, 1ex; USNM).

Vermont. Orange: Fairlee (43°54.5¢N, 72°08.6¢W), 

5 Aug 1956, A. H. Sturtevant (1♂; USNM). Windsor: 

Mad Brook Farm, E Charleston (44°50.6¢N, 71°59.4¢W), 

15–25 Jul 1982, D. A. Grimaldi (2♂; AMNH).

Virginia. Fairfax: Dead Run (38°58¢N, 77°10.3¢W), 

R. C. Shannon (8♂, 9♀; USNM); Mt. Vernon (38°42.5¢N, 

77°05.2¢W; on tree trunk), 22 Aug 1976, P. H. Arnaud, Jr. 

(1♀; CAS); Turkey Run (mouth; 38°57.9¢N, 77°09.4¢W), 

4 May 2006, D. and W. N. Mathis (2♂; GWMP, USNM).

Washington. Walla Walla: Walla Walla, Mill Creek 

(46°02.3¢N, 118°28.7¢W), 2–6 Jul 1922, V. N. Argo, A. L. 

Melander (16♂, 8♀; USNM).

Wisconsin. Dane (43°04.8¢N, 89°23.2¢W): 29 Apr 

1951, R. H. Jones (1♂; USNM).

Distribution. (Map 16) Nearctic: Canada 

(Alberta, British Columbia, Manitoba, New Brunswick, 

Ontario, Quebec, Saskatchewan), Mexico (Baja California, 

Baja California Sur), United States (California, Connecticut, District of Columbia, Idaho, Illinois, Indiana, Iowa, 

Kansas, Massachusetts, Maryland, Michigan, Minnesota, 

Nevada, New Hampshire, New Jersey, New Mexico, New 

York, Oregon, Pennsylvania, South Dakota, Utah, Vermont, Virginia, Washington, Wisconsin). 

Biology. This species has been collected on 

wounds and fluxes from the following genera of deciduous 

trees: Acer, Alnus, Pinus, Platanus, Populus, Quercus, and 

Salix. Davis and Zack (1978) reared adults from larvae that 

were in seeps of a wounded Douglas fir (Pseudotsuga menziesii (Mirbel) Franco). Bächli et al. (1999) collected this 

species in Europe using beer/wine traps.

Remarks. Mathis and Freidberg’s illustration 

of the internal male terminalia of this species (1994:588, 

fig. 8) does not show a “ventral gonal bridge,” which the 

authors correctly recognized as being present and described 

as “straight or slightly indented posteriorly.” The “ventral 

gonal bridge” is an anteroventral connection between the 2 

halves of the hypandrium + gonopod and is present in all 

aulacigastrids. Slight changes in the position of the internal 

male structures, however, may conceal this character. Moreover, the “ventral gonal bridge” may look slightly different, 

depending on the condition of the preparation of the male 

terminalia. For this reason, we have not chosen to describe 

this feature here, as did Mathis and Freidberg (1994), but 

we do provide illustrations of the gonopod from a position 

that shows the gonopodal bridge (Figures 172, 174, 176).

This widely occurring species has the widest range of 

variation among congeners, and it is possible that this species represents a species complex. Even though we have 

found some variation in the size and shape of the shiny 

spot adjacent to the ocellar tubercle, extent of microtomentum over the katepisternum, darkness of the legs and 

postpronotum, slight variations in the size and shape of 

the surstylus, shape of the gonopod and posterodorsal 

gonopodal lobe, and the length of the cleft between the 

cerci, we have chosen a conservative approach and have 

not recognized additional species. The observed variation 

does not show any consistency that would allow for clear 

separation between forms. Perhaps with use of molecular 

evidence or information from behavior, immature stages 

or the female internal genital tract, additional species will 

be recognized.

Aulacigaster sabroskyi Mathis and Freidberg

Figures 183–184, Map 16

Aulacigaster sabroskyi Mathis and Freidberg, 1994:591–593 [illustration of 

male terminalia and spermathecae].

Diagnosis. This species is distinguished from 

congeners by the following combination of characters: Posterior fronto-orbital seta slightly internal to and almost horizontally aligned with anterior seta; katepisternum mostly 

microtomentose, with a shiny area posteriorly, below the 

posterior katepisternal seta; surstylus short, less than 0.2 

times length of basal portion of epandrium in lateral view.

Description. Coloration and vestiture: Vertex 

mostly densely microtomentose, with a shiny spot extended 

from ocellus to 2/3 distance to eye margin; ocellar tubercle 

dull microtomentose; frons bearing a transverse, large, orange band, frontal orange band merging into dark brown 

ventrally; silver stripe on frons approximately straight; antenna pale yellow to yellowish, infuscate along dorsal margin; face yellow, with a transverse brown mark on dorsal 

portion; facial band present, sharply defined, transverse, 

on protruding, ventral portion of face; palpus white. Scutum with weakly defined microtomentose stripes; scutellum 

dull microtomentose; postpronotum concolorous with mesonotum, dull microtomentose; anepisternum mostly dull; 

katepisternum mostly microtomentose, with a shiny area 

posteriorly, below the posterior katepisternal seta; halter 

mostly white. Fore coxa brown to black, fore femur brown 

to black, fore tibia brown to black; fore tarsus yellowish, 
number 633 • 9 9

apical and sometimes subapical tarsomeres brown to black; 

mid coxa brown to black, mid femur brown to black, mid 

tibia brown to black, mid tarsus yellowish, apical and subapical tarsomere brownish to dark brown; hind coxa brown 

to black, hind femur brown to black, hind tibia brown to 

black, hind tarsus yellowish, apical tarsomere brown to 

black. Male abdomen subshiny to dull, pregenital segment 

polished posteriorly, dull anteriorly; female abdomen with 

dense microtomentum medially, lateral margins of tergites 

nearly polished.

Head: Head higher than long (head ratio less than 

0.9); face at level of pseudovibrissa wider than width of 

1st flagellomere, in lateral view with dorsal half concave, 

ventral portion bulbous; gena in lateral view approximately 

1/3–1/2 the width of 1st flagellomere; medial vertical seta 

3/4 length of lateral vertical seta; 1 long and 3–4 shorter 

peristomal setae following pseudovibrissal setae present.

Thorax: Acrostichal setae in 1 row, bifurcating 

posteriorly; scutellum slightly raised (angle with scutum approximately 135°), trapezoidal, apex nearly straight, disk 

of scutellum slightly convex; basal scutellar seta 1/2 length 

of posterior seta.

Male abdomen and terminalia: Figures 183–184. 

Surstylus arising from posteroventral margin of epandrium, 

slender, digitiform, relatively short; subepandrial sclerite 

(fig. 13; Mathis and Freidberg, 1994:591) with lateral arms 

narrow, well sclerotized, more or less parallel sided, curved 

evenly posteriorly; cerci fused on basal 1/2; gonopod (fig. 

MAP 16. Distribution of Aulacigaster neoleucopeza and A. sabroskyi (the leucopeza group, Nearctic Region).
100 • smithsonian contributions to zoology

12; Mathis and Freidberg, 1994: 591) with a small pointed 

process posterodorsally, approximately rectangular in lateral view.

Measurements: Body length 2.0–2.4 mm. Wing 

length 2.0–2.4 mm, wing width 0.7 mm.

Type Material. The holotype male is labeled 

“ARIZ.: Portal 4800 ft. vi-5–13 Sept.[crossed out] 1967 

[“vi-5-” and “7” of 1967 handwritten]/CWSabrosky collector/At bleeding sap, cottonwood/HOLOTYPE ♂ Aulacigaster sabroskyi W.N.Mathis&Freidberg USNM [red; “♂

Aulacigaster sabroskyi” and “Freidberg” handwritten].” 

Paratypes: MEXICO. Chihuahua: Camargo (27°40¢N, 

105°10¢W; wound in poplar tree), 30 May 1964, J. F. McAlpine (1♀; CNC). UNITED STATES. Arizona. Cochise: 

Portal (31°53¢N, 109°12.2¢W; at bleeding sap, cottonwood; 

1463 m), 5 Jun 1967, C. W. Sabrosky (8♂, 2♀; USNM, 

TAU); Portal (31°53¢N, 109°12.2¢W; slime flux on tree), 17 

May 1973, C. W. Sabrosky (5♂, 2♀; USNM); Portal, Turkey 

Creek (31°59.9¢N, 109°45.1¢W; 1950 m), 18 May 1973, 

C. W. Sabrosky (1♂; USNM). Gila: Carrizo (33°59.6¢N, 

110°17.3¢W), 26 Sep 1954, A. H. Sturtevant (1♂; USNM).

New Mexico. Catron: Whitewater Canyon 

(33°21.6¢N, 108°03.9¢W; ulcer on Sycamore tree), 1 Jun 

1972, W. W. Wirth (3♂, 3♀; USNM).

Type Locality. United States. Arizona. Cochise: Portal (31°54.5¢N, 109°08.8¢W).

Distribution. (Map 16) Nearctic: Mexico 

(Chihuahua), Unites States (Arizona, New Mexico).

Neotropical Species

Discussion. The Neotropical fauna of the leucopeza group comprises three species of which all are described here.

FIGURES 183–184. Illustrations of Aulacigaster sabroskyi Mathis and Freidberg (male) (the leucopeza group, Nearctic Region). (183) gonopod, lateral aspect; (184) epandrium, lateral aspect.

Key to the Neotropical Species of the leucopeza Group of Aulacigaster

1. Frons bearing a horizontal, unusually large, orange band extended beyond base of anterior ocellus medially [Figure 185]; 

face in lateral view dorsoventrally projected, ventral portion of face not receded; facial band unusually wide, ~1/3 length 

of 1st flagellomere [Figure 186] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. L2, undescribed species

Frons at most with a narrower, orange-brown transverse band [as in Figure 217]; ventral portion of face receded [Figure 

216] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2. Frons yellowish-orange on central portion, merging with black along lateral margins; facial band absent [Figures 215–

216]; cephalic and thoracic setae not unusually long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. spangleri, new species

Frons with a transverse, narrow, orange-brown band; facial band present; cephalic and thoracic setae relatively long

 .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. L1, undescribed species
number 633 • 101

Aulacigaster spangleri, new species

Figures 215–216, Map 17

Diagnosis. This species is distinguished from 

congeners by the following combination of characters: 

Frons yellowish-orange on central portion, merging with 

black along lateral margins; arista with inconspicuous, alternate hairs, appearing naked.

Description. Coloration and vestiture: Vertex 

mostly polished, with a densely microtomentose stripe posterior to ocellar tubercle; ocellar tubercle dull microtomentose; frons yellowish-orange centrally, merging with black 

along lateral margins; silver stripe on frons approximately 

semi-lunate; antenna pale yellow to yellowish, not infuscate 

dorsally; face brownish; facial band absent; palpus white. 

Scutellum dull microtomentose; postpronotum concolorous with mesonotum, scarcely microtomentose, subshiny; 

anepisternum polished; katepisternum mostly polished; 

halter mostly white. Fore coxa brown to black, fore femur 

brown, fore tibia brown, fore tarsus yellowish, apical and 

sometimes subapical tarsomeres brown to black; mid coxa 

brown to black, mid femur brown (base and apex yellowish), mid tibia yellowish, mid tarsus mostly yellowish; hind 

coxa yellow, hind femur brown to black on central portion, 

apex and base yellowish (base yellowish), hind tibia brown 

to dark brown, hind tarsus yellowish, apical tarsomere 

brown to black. Abdomen with a large, white to yellowish 

region on syntergite 1+2, male abdomen subshiny to dull; 

female abdomen with dense microtomentum over anterior 

margin of tergites, posterior margin of tergites bright shiny.

Morphology: 

Head: Figures 215–216. Head higher than long 

(head ratio less than 0.9); face at level of pseudovibrissal 

seta wider than width of 1st flagellomere, in lateral view 

with dorsal half concave, ventral portion bulbous; gena in 

lateral view approximately 1/3–1/2 the width of 1st flagellomere; peristomal area with 1 long and about 4 shorter 

setae following pseudovibrissal seta.

Thorax: Acrostichal setae in 2 rows; scutellum 

slightly raised (angle with scutum approximately 135°), 

trapezoidal, apex nearly straight, disk of scutellum slightly 

convex; basal scutellar seta 1/2 length of posterior seta. Costal vein of male bearing a few tiny, but conspicuous spines.

Male abdomen and terminalia: Surstylus arising 

from posteroventral margin of epandrium, slender, approximately triangular, bearing setae (with a long terminal seta), 

relatively long; cerci broad, plate-like, bearing short setae, 

none especially elongate (observations made without dissecting abdomen).

Measurements: Body length 2.2 mm. Wing length 

2.3 mm, wing width 0.8 mm.

Type Material. The holotype male is labeled 

“COSTA RICA[.] Turrialba[,] VII-15-19-65 [15–19 Jul 

1965] P. J. Spangler/HOLOTYPE ♂ Aulacigaster spangleri Rung & Mathis USNM [red].” The holotype is double mounted (minuten in a block of polyporus), is in good 

condition (vertical, scutellar and mesonotal setae broken, 

wings slightly folded), and is deposited in the USNM. A 

paratype is as follows: Same label data as the holotype 

(1♀, USNM).

Type Locality. Costa Rica. Cartago: Turrialba (09°54¢N, 83°41¢W).

Distribution. (Map 17) Costa Rica (Cartago).

Etymology. The specific epithet, spangleri,

is a genitive patronym to honor and recognize the friendship and numerous contributions of Dr. Paul J. Spangler to 

entomology.

Remarks. The holotype is precariously pinned, 

and removal of the abdomen would probably break what 

remains of this fragile specimen. Thus, we have opted not to 

dissect this specimen.

Aulacigaster L1, undescribed species

Map 17

Diagnosis. This species is distinguished from 

congeners by the following combination of characters: 

Frons with a dark, orange, narrow stripe infuscated with 

brown; cephalic and thoracic setae relatively long.

Description. Coloration: Vertex mostly polished (microtomentose around vertical setae); ocellar tubercle dull microtomentose; frons with a transverse, narrow, 

orange-brownish stripe; silver stripe on frons approximately 

straight, bright shiny; antenna pale yellow to yellowish, not 

infuscate dorsally; face yellow, with a transverse brown 

mark on dorsal portion; facial band present, sharply defined, transverse, on protruding, ventral portion of face; 

palpus white. Scutellum dull microtomentose; postpronotum concolorous with mesonotum, dull microtomentose; 

anepisternum polished; katepisternum mostly polished; 

halter mostly white. Fore coxa brown to black, fore femur 

brown, fore tibia brown, fore tarsus yellowish, apical and 

sometimes subapical tarsomeres brown to black; mid coxa 

brown to black, mid femur brown on central portion, apex 

and base yellowish, mid tibia yellowish-brownish, mid tarsus yellowish, apical and subapical tarsomere brownish to 

dark brown; hind coxa brown, hind femur brown to black 

on central portion, apex and base yellowish, hind tibia 
102 • smithsonian contributions to zoology

yellowish, hind tarsus yellowish, apical tarsomere brown to 

black. Male abdomen subshiny to dull.

Morphology: 

Head: Head higher than long (head ratio less than 

0.9); face at level of pseudovibrissal seta wider than width 

of 1st flagellomere, in lateral view with dorsal half concave, 

ventral portion bulbous; gena in lateral view approximately 

1/3–1/2 the width of 1st flagellomere; cephalic and thoracic 

setae relatively long; ocellar seta very thin, but long, 1/3 

length of fronto-orbital setae; medial vertical seta 3/4 length 

of lateral vertical seta; peristomal area with about 3 setae 

following pseudovibrissal seta.

Thorax: Acrostichal setae in 2 rows; scutellum 

slightly raised (angle with scutum approximately 135°), 

approximately triangular, apex relatively pointed, disk of 

scutellum slightly convex; basal scutellar seta 3/4 length of 

posterior seta.

Male abdomen and terminalia: Cerci bearing short 

setae, none especially elongate.

Measurements: Body length 3.5 mm. Wing length 

3.1 mm, wing width 1.3 mm.

Material Examined. COLOMBIA. Nariño, 

Laguna, La Cocha (01°05¢N, 77°09¢W), 26 Sep 1971, Gebohart (1♂; USNM).

Distribution (Map 17) Neotropical: Colombia (Nariño).

Remarks. We have decided not to name this 

species because there is only one specimen available and the 

MAP 17. Distribution of the Neotropical species of the leucopeza group of Aulacigaster (Nearctic Region).
number 633 • 103

surstylus was damaged during dissection of the epandrium. 

Therefore, this important structure cannot be described.

Aulacigaster L2, undescribed species

Figures 185–186, Map 17

Diagnosis. This species is distinguished from 

congeners by the following combination of characters: 

Arista short, with very inconspicuous, alternate hairs 

(arista appearing naked at low magnification); face in lateral view dorsoventrally projected, ventral portion of face 

not receded.

Description. Coloration and vestiture: Vertex mostly polished, with a densely microtomentose spot 

posterior to ocellar tubercle; ocellar tubercle dull microtomentose; frons bearing a transverse, unusually large, 

orange band extended beyond anterior ocellus medially, 

frontal orange band merging into dark brown ventrally; 

silver stripe on frons approximately straight, bright shiny; 

antenna pale yellow to yellowish, infuscate along dorsal 

margin; face yellow dorsally, brown ventrally; facial band 

present, unusually broad (about 1/3 length of 1st flagellomere), sharply defined, transverse, on protruding, ventral 

portion of face; palpus white. Scutum strongly microtomentose medially, polished between pleural region and 

dorsocentral row of setae, posterior of postpronotum; 

scutellum dull microtomentose; postpronotum concolorous with mesonotum, dull microtomentose; anepisternum 

mostly dull; katepisternum with anterior portion dull microtomentose, and posterior portion mostly polished; halter mostly white (knob with dark mark). Fore coxa brown 

to black, fore femur brown, fore tibia yellowish-brownish, 

fore tarsus yellowish, apical and sometimes subapical tarsomeres brown to black; mid coxa brown to black, mid 

femur brown to black, apex yellowish, mid tibia yellowishbrownish, mid tarsus yellowish, apical and subapical tarsomere brownish to dark brown; hind coxa brown, hind 

femur brown to black, apex yellowish, hind tibia brown 

to dark brown (tip yellowish), hind tarsus yellowish, apical tarsomere brown to black. Female abdomen with dense 

microtomentum medially, lateral margins of tergites nearly 

polished (tergite 6 mostly polished).

Morphology: 

Head: Figures 185–186. Head round, about as 

high as long (head ratio 0.9–1.1); face at level of pseudovibrissal seta wider than width of 1st flagellomere, in lateral view projected dorsoventrally, ventral portion of face 

not receded; ocellar seta very thin, but 2/3 of fronto-orbital 

setae; medial vertical seta 3/4 length of lateral vertical seta.

Thorax: Acrostichal setae in 2 rows; scutellum 

strongly raised (angle with scutum less than 135°), triangular, apex strongly pointed; basal scutellar seta 1/2 length of 

posterior seta.

Specimens Examined. PERU. Madre de 

Dios: Rio Manu, Pakitza (11°56.6¢S, 71°16.9¢W; 250 m), 

9–23 Sep 1988, A. Freidberg (1♀; USNM).

Measurements: Body length 2.2 mm. Wing length 

2.0 mm, wing width 0.7 mm.

Distribution. (Map 17) Neotropical: Peru 

(Madre de Dios).

Remarks. In addition to the characters cited in 

the diagnosis, this species is peculiar among the Neotropical species of the leucopeza group in having a wide, orange 

band on the anterior portion of frons. This species was not 

named because it is represented by a single female.

FIGURES 185–186. Photographs of Aulacigaster L2, undescribed species (female) (the leucopeza

group, Neotropical Region). (185) head, frontal aspect; (186) head, lateral aspect.
104 • smithsonian contributions to zoology

Palearctic Species

Discussion. The Palearctic fauna includes four 

named species, all in the leucopeza group. Papp (1998a:233) 

provided a key to the Palearctic species of Aulacigaster that 

included four species, one of which, A. neoleucopeza Mathis 

and Papp, was later recognized by Kassebeer (2001) as a 

distinct species, A. pappi. Kassebeer (2001:30) provided a 

key to the species from Germany.

Key to the Palearctic Species of Aulacigaster

1. Frontal orange band higher than 1/2 distance of lunule to anterior ocellus. Postpronotum yellow to pale brown, lighter in 

color than rest of mesonotum. Epandrium densely setose. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

Frontal orange band shorter than 1/2 distance of lunule to anterior ocellus. Postpronotum concolorous with mesonotum. 

Epandrium not densely setose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

2. Frons posterior of fronto-orbital setae completely shiny. Surstylus digitiform (fig.1; Papp, 1998a: 227) . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. afghanorum Papp

Frons posterior of fronto-orbital setae at least partly microtomentose, with a shiny spot extended from ocellus to near eye 

margin (shiny spot separated from eye margin by a distance less than the width of one ocellus). Surstylus sickle-shaped 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. falcata Papp

3. Shiny spot on vertex extended from ocellar tubercle to near eye margin (shiny spot separated from eye margin by a distance that is approximately equal to the width of one ocellus). Surstylus shorter, ~0.2 times the length of ventral margin 

of epandrium, tapered to a blunt apex (fig. 2d; Kassebeer, 2001:31) . . . . . . . . . . . . . . . . . . . . . . . . . . A. pappi Kassebeer

Shiny spot on vertex smaller, extended from ocellus to 1/2 distance to eye margin. Surstylus longer, nearly 0.3–0.4 times 

the length of ventral margin of epandrium, slightly tapered to a rounded apex. . . . . . . . . . . . . . . . A. leucopeza Meigen

Aulacigaster afghanorum Papp

Aulacigaster afghanorum Papp, 1998a:226, 227–229 [figures of male terminalia]; 1998b:279–284 [Manual of Palearctic Diptera].

Diagnosis. This species is distinguished from 

congeners by the following combination of characters: 

Frons shiny, microtomentum absent posterior of frontoorbital setae; abdomen completely dark brown to black; 

posterior fronto-orbital seta slightly medial to and almost 

horizontally aligned with anterior seta; surstylus slender, 

approximately digitiform, making a wide angle with anterior margin of epandrium.

Description. Coloration and vestiture: Vertex 

sparsely microtomentose, frons posterior of fronto-orbital 

setae completely shiny; ocellar tubercle dull microtomentose; frons bearing a transverse, large, orange band, frontal 

orange band higher than 1/2 distance of lunule to anterior 

ocellus, merging into dark brown ventrally; silver stripe on 

frons approximately straight; antenna pale yellow to yellowish, not infuscate dorsally; face yellow, with a transverse 

brown mark on dorsal portion; facial band present, sharply 

defined, transverse, on protruding, ventral portion of face; 

palpus yellowish. Scutellum dull microtomentose; postpronotum yellow to yellowish-brownish, dull microtomentose; 

anepisternum mostly dull; katepisternum subshiny to dull; 

halter mostly yellowish. Fore coxa brown to black, fore 

femur brown or brown to black, apex yellowish, fore tibia 

brown, fore tarsus brown to dark brown, basal tarsomere 

yellow; mid coxa brown to black, mid femur brown, mid 

tibia brownish, mid tarsus yellowish, apical and subapical 

tarsomere brownish to dark brown; hind coxa brown, hind 

femur brown, hind tibia brown to dark brown, hind tarsus 

yellowish, apical tarsomere brown to black. Male abdomen 

subshiny to dull, pregenital segment mostly polished. Epandrium mostly dark brown to black.

Morphology: 

Head: Higher than long (head ratio less than 0.9); 

face at level of pseudovibrissa wider than width of 1st 

flagellomere, in lateral view with dorsal half concave, ventral portion bulbous; gena in lateral view approximately 

1/3–1/2 the width of 1st flagellomere; ocellar seta minute; 

medial vertical seta 3/4 the length of lateral vertical seta; 

fronto-orbital setae with posterior seta internal to and almost horizontally aligned with anterior seta; peristomal 

vestiture consisting of relatively well-developed setae; 

peristomal area with about 3–4 setae following pseudovibrissal seta.

Thorax: Acrostichal setae in 2 rows; scutellum 

slightly raised (angle with scutum approximately 135°), 

trapezoidal, apex nearly straight, disk of scutellum slightly 

convex; basal scutellar seta 1/2 the length of posterior seta.

Male abdomen and terminalia: Epandrium densely 

setose. Surstylus arising from posteroventral margin of 
number 633 • 105

epandrium, slender, digitiform, relatively long (approx. 

0.4 times the length of ventral margin of epandrium) (fig. 

1; Papp, 1998a:227); subepandrial sclerite (fig. 9; Papp 

1998a:229) with lateral arms spread sideways, with a beakshaped apex; cerci broad, parallel-sided, with apex setose 

and lacking well-differentiated setae; gonopod (fig. 3; Papp, 

1998a:227) with a small pointed process posterodorsally, 

preceded by several elongate setae, shallow ventrally (about 

0.7 times as long as high), approximately rectangular to 

trapezoid in lateral view.

Measurements: Body length 1.63–2.65 mm. Wing 

length 1.95–2.75 mm, wing width 0.78–1.10 mm.

Type Material. The holotype male is labeled 

“Afganistan[,] Kabul, Aliabad/ University Park, 1800 m[,] 

21.4.1974/No. 49[,] L. Papp/Aulacigaster leucopeza Meig. 

[handwritten] det. L. Papp 1986/Partypus Aulacigaster afghanorum L. Papp [white label with red borders, species 

name handwritten].” The specimen is double mounted (minuten in cardboard), is in excellent condition, and is deposited in HNHM.

Type Locality. Afghanistan. Kabul: Aliabad, 

University Park (34°31¢N, 69°08¢E).

Distribution. Palearctic: Afghanistan 

(Kabul).

Biology. This species was collected from oozing sap of Populus alba L. and Morus alba L. (Papp, 1998a).

Aulacigaster falcata Papp

Figure 187

Aulacigaster falcata Papp, 1998a:226–229 [figures of male terminalia]; 1998b:279–284 [Manual of Palearctic Diptera].—Bächli et al., 

1999:120 [checklist, Greece, Italy, Switzerland].—Kassebeer, 2001: 

24–25 [biology, locality data from Germany].

Diagnosis. This species is distinguished from 

congeners by the following combination of characters: 

Posterior fronto-orbital seta slightly internal to and almost 

horizontally aligned with anterior seta; surstylus slender, 

approximately sickle-shaped.

Description. Coloration and vestiture: Vertex 

mostly densely microtomentose, with a shiny spot extended 

from ocellus to near eye margin, leaving only a narrow microtomentose stripe parallel to eye margin; ocellar tubercle 

dull microtomentose; frons bearing a transverse, large, orange band, frontal orange band higher than 1/2 distance 

of lunule to anterior ocellus, merging into dark brown 

ventrally; silver stripe on frons approximately straight; antenna pale yellow to yellowish, infuscate along dorsal margin; face yellow, with a transverse brown mark on dorsal 

portion; facial band present, sharply defined, transverse, on 

protruding, ventral portion of face; palpus white. Scutellum 

dull microtomentose; postpronotum yellow to yellowishbrownish, dull microtomentose; anepisternum mostly dull; 

katepisternum with anterior portion dull microtomentose, 

and posterior portion mostly polished; halter mostly white, 

or mostly yellowish. Fore coxa brown to black, fore femur 

brown, fore tibia brown, fore tarsus brown to dark brown, 

basal tarsomere yellow (1–3); mid coxa brown to black, 

mid femur brown, mid tibia brownish, mid tarsus yellowish, apical and subapical tarsomere brownish to dark 

brown; hind coxa brown, hind femur brown, hind tibia 

brown to dark brown, hind tarsus yellowish, apical tarsomere brown to black. Male abdomen subshiny to dull, 

pregenital segment mostly subshiny; female abdomen with 

dense microtomentum medially, lateral margins of tergites 

nearly polished.

Morphology: 

Head: Head higher than long (head ratio less than 

0.9); face at level of pseudovibrissa wider than width of 

1st flagellomere, in lateral view with dorsal half concave, 

ventral portion bulbous; gena in lateral view approximately 1/3–1/2 the width of 1st flagellomere; ocellar seta 

minute; medial vertical seta 3/4 the length of lateral vertical seta; fronto-orbital setae with posterior seta internal to 

and almost horizontally aligned with anterior seta; setulae 

between fronto-orbital setae 1, proclinate; peristomal vestiture consisting of relatively well-developed setae; peristomal area with about 4 setae following pseudovibrissal seta.

Thorax: Acrostichal setae in 2 rows; scutellum 

slightly raised (angle with scutum approximately 135°), 

trapezoidal, apex nearly straight, disk of scutellum slightly 

convex; basal scutellar seta 1/2 the length of posterior seta.

Male abdomen and terminalia: Figure 187. Epandrium densely setose. Surstylus arising from posteroventral 

margin of epandrium, slender, digitiform, relatively long (approx. 0.6 times the length of ventral margin of epandrium) 

(fig. 2; Papp 1998a:227); subepandrial sclerite (fig. 10; Papp 

1998a:229) with lateral arms spread sideways, tapering toward pointed apex; cerci broad, parallel-sided to slightly 

divergent from each other, with apex setose and lacking 

well-differentiated setae; gonopod (fig. 4; Papp 1998a:227) 

with a small pointed process posterodorsally, preceded by 

several elongate setae, shallow ventrally (about 0.7 times as 

long as high), approximately trapezoid in lateral view.

Biology. Adults of this species have been recorded from hornbeam (Carpinus sp), oak trees (Quercus

sp) (1998a), and Morus sp (1998a). It has also been collected in beer/wine traps (Kassebeer, 2001).

Measurements: Body length 1.92–2.5 mm. Wing 

length 2.17–2.88 mm, wing width 0.80–1.07 mm.
106 • smithsonian contributions to zoology

Type Material Although we did not examine 

the holotype of this species, we did examine a topotypical 

male paratype that is reported to have the same label data 

as the holotype. This male paratype is labeled “KUP. Kunfehértó 1988. VII. 15. [handwritten]/leg.Papp L./Paratypus ♂

Aulacigaster falcata L. Papp [all except “Paratypus” handwritten; red border].” The paratype, and presumably the 

holotype, is double mounted (minuten in a flat rectangular 

card) and is deposited in HNHM.

Type Locality. Hungary. Bács-Kiskun: 

Kiskunsági National Park, Kunfehértó (47°02¢N, 18°58¢E).

Other Specimens Examined. HUNGARY. 

Bács-Kiskun: Kiskunsági National Park, Kunfehértó 

(47°02¢N, 18°58¢E), 15 Jul 1982, L. Papp (1♂; paratype; 

USNM). Budapest. Pestszeõntlorinc, Halmi-erdõ, 11 Jul 

1998, L. Papp, “tölgifák sebeirõl” (1♂, 1♀; USNM).

Distribution. Palearctic: Germany, Greece, 

Hungary, Italy, Switzerland.

Aulacigaster leucopeza (Meigen)

Figures 188–193, 217–218

Diastata leucopeza Meigen, 1830:100.

Aulacigaster leucopeza.—Oldenberg, 1914:30 [generic combination, as “Aulacogaster” = unjustified emendation].—Duda, 1924:176 [discussion]; 

1934:3–4 [revision].—Robinson, 1953:77–83 [description of biology 

and immature stages].—Chandler, 1987:40 [interpretation and usage 

as a synonym of A. rufitarsis Macquart].—Ferrar, 1987:79–81, 581 

[biology, breeding habits and immature stages].—Teskey, 1987:891, 

894 [illustrations, manual Nearctic Diptera].—Mathis and Freidberg, 

1994:583 [locality data].—Bächli, 1997:33 [faunistic and host record].—Papp, 1998a:228, 231–233 [illustrations and key to Palearctic 

species]; 1998b:279–284 [Manual of Palearctic Diptera].—Bächli et al.,

1999:120–121 [checklist, Switzerland].—Kassebeer, 2001:25–26 [biology, locality data from Germany].—Papp, 2008:229–232 [comparison 

of female terminalia and immature stages with A. africana].

Aulacigaster rufitarsis Macquart, 1835:580. Type locality: Liège (Belgium).—

Schiner, 1863:270 [review, synonymy]; 1864:56 [list of Palearctic species].—Becker, 1905:216 [Palearctic catalog].

Diastata diadema Meigen, 1838:379. Type locality: Lüetticher Gegend [near 

Liège] (Belgium).

Apotomella impressifrons Dufour, 1846:460. Type locality: France.—

Schiner, 1863:270 [synonymy with A. rufitarsis].

Ampycophora tarsata Wahlberg, 1847:261. Type locality: Sweden.—Schiner, 

1863:270 [synonymy with A. rufitarsis].

Sephanilla sertulata Rondani, 1874:268.—Mathis and Freidberg, 1994:585 

[synonymy with D. leucopeza Meigen].

Diagnosis. This species is distinguished from 

congeners by the following combination of characters: Posterior fronto-orbital seta slightly medial to and almost horizontally aligned with anterior seta; vertex mostly densely 

microtomentose, with a shiny spot extended from ocellus 

at most 1/2 way to eye margin; surstylus digitiform, relatively long (nearly 0.3 times length of ventral margin of 

epandrium).

Description. Coloration and vestiture: Vertex 

mostly densely microtomentose, with a shiny spot extended 

from ocellus at most 1/2 way to eye margin (usually not 

more than 1/3–1/2 way to eye margin); ocellar tubercle dull 

microtomentose; frons bearing a transverse, large, orange 

band, frontal orange band shorter than 1/2 distance of lunule to anterior ocellus, merging into dark brown ventrally; 

silver stripe on frons approximately straight; antenna pale 

yellow to yellowish, infuscate along dorsal margin or infuscate on dorsal half; face yellow, with a transverse brown 

mark on dorsal portion; facial band present, sharply defined, transverse, on protruding, ventral portion of face; 

palpus white. Scutum with faintly defined microtomentose stripes; scutellum dull microtomentose; postpronotum concolorous with mesonotum, dull microtomentose; 

anepisternum mostly dull; katepisternum mostly microtomentose on anterior half and on a narrow stripe over 

katepisternal setae, halter mostly white. Fore coxa brown 

to black, fore femur brown to black, apex yellowish, fore 

tibia brown, fore tarsus yellowish, apical and sometimes 

FIGURE 187. Illustrations of Aulacigaster falcata Papp (male) (the 

leucopeza group, Palearctic Region): epandrium, lateral aspect.
number 633 • 107

subapical tarsomeres brown to black; mid coxa brown to 

black, mid femur brown to black, apex yellowish, mid tibia 

completely brown or yellowish on central portion, apex 

and basis brown, mid tarsus yellowish, apical and subapical tarsomere brownish to dark brown; hind coxa brown, 

hind femur brown to black, apex yellowish, hind tibia completely brown to dark brown, or with basal and central 

portion yellowish, hind tarsus yellowish, apical tarsomere 

brown to black. Male abdomen subshiny to dull; female 

abdomen with dense microtomentum medially, lateral margins of tergites nearly polished.

Morphology: 

Head: Figures 189–191, 217–218. Head higher 

than long (head ratio less than 0.9); face at level of pseudovibrissa wider than width of 1st flagellomere, in lateral 

view with dorsal half concave, ventral portion bulbous; 

gena in lateral view approximately 1/3–1/2 the width of 

1st flagellomere; medial vertical seta 3/4 length of lateral 

vertical seta; 3–4 peristomal setae following pseudovibrissal 

setae present.

Thorax: Figures 188–189. Acrostichal setae in 2 

rows, these often convergent anteriorly; scutellum slightly 

raised (angle with scutum approximately 135°), trapezoidal, apex nearly straight, disk of scutellum slightly convex; 

basal scutellar seta 1/2 length of posterior seta.

Male abdomen and terminalia: Figures 192–193. 

Epandrium not densely setose. Surstylus arising from 

posteroventral margin of epandrium, slender, digitiform, 

relatively long (0.3–0.4 times length of ventral margin of 

epandrium), tapering toward a rounded apex; subepandrial 

sclerite (fig. 9; Papp 1998a:229) with lateral arms spread 

sideways, tapering toward pointed apex; cerci broad, bulging medially, with apex setose and lacking prominent setae; 

gonopod with a small pointed process posterodorsally, preceded by several elongate setae, shallow ventrally (about 

0.9 times as long as high), approximately rectangular in 

lateral view.

Type Material. Meigen’s description of Diastata leucopeza indicates a type series of a single specimen, 

gender unknown, that had a broken antenna and that is 

now presumably lost. Becker (1902) did not find the type, 

and our queries and searches did not result in its discovery 

(see remarks below).

Type Locality. Unknown, presumably Germany (Baumhauer’s collection).

Other Specimens Examined. AUSTRIA. 

J. Egger (in Loew’s collection (labeled as “Aulacigaster rufitarsis Macquart”) (1♀; ZMB).

DENMARK. Copenhagen: Dyrehaven (55°39.9¢N, 

12°32.6¢E), 3 May–6 Jun 1917, 1918, W. Lundbeck (9♂, 

3♀; UZMC).

GERMANY. Berlin: Dahlem (52°27¢N, 13°17¢E), 

25 May 1959, Kirchberg (1♂, 1♀; ZMB); Zehlendorf 

(52°26¢N, 13°16¢’E), 3 Jun 195?, Kirchberg (1♂, 1♀; 

ZMB). Brandenburg: Frankfurt. Oder (52°20.4¢N, 

14°31.8¢E), M. P. Riedel (labeled as “Aulacigaster leucopeza Meigen”. The female labeled as “Aulacigaster (rufitarsis) leucopeza”, P. P. Riedel Det.) (4♂,1♀; ZMB). 

Saxony-Anhalt: Dessau (51°49.8¢N, 12°13.2¢E), 8 Jul 

1924, E. Heidenreich (labeled as “Aulacigaster leucopeza

Meigen”, M. P. Riedel 1924 Det.) (1♂; ZMB).

HUNGARY. Mátra Mountains (47°55.1¢N, 

19°53.7¢E), 24 Aug 1996, A. Freidberg (1♀; USNM).

MONGOLIA. Khovd: Dund Tsenkher Gol 

(47°20.4¢N, 91°51.8¢E; 1745m), 17 July 2009, W.N. Mathis ((2♂, 1♀; USNM).

POLAND. Rauden OS (53°37¢N, 19°05¢E), 4–21 Jul 

1932, L. Duda (6♂; ZMB). Wrocław (51°05¢N, 17°00¢E), 

07 Apr 1982, T. Zatwarnicki (2♀; CAS). Wustung bei 

Habelschwerdt (50°17.9¢N, 16°38.5¢E), 16 May–25 Jun 

1924, L. Duda (6♂; ZMB).

SWEDEN: Lund (55°42¢N, 13°12¢E), 29 Oct 1954, 

J. R. Vockeroth (1♀; CNC).

Distribution. Palearctic: Austria, Denmark, 

France, Germany, Hungary, Israel, Italy, Mongolia, Poland, 

Spain, Sweden, Switzerland, United Kingdom.

Biology. This species has been recorded from 

the following hosts: Aesculus sp. (Kassebeer, 2001), Betula

sp. (Kassebeer, 2001), Populus sp. (Kassebeer, 2001), Quercus sp. (Kassebeer, 2001) and Ulmus sp. (Kassebeer, 2001). 

It has also been collected by beer/wine traps (Bächli et al.,

1999; Kassebeer, 2001).

Remarks. Until the mid-1990s, A. leucopeza

was the only known species of Aulacigaster from the Palaearctic Region. Papp 1998a then described A. afghanorum

and A. falcata and reported the presence of A. neoleucopeza

(later named A. pappi by Kassebbeer (2001) from that region and noted that previous locality records of A. leucopeza needed verification (Papp, 1998a:230). A few years 

later, after collecting and studying museum specimens from 

Germany, Kassebeer (2001) reported the presence of A. falcata (previously known only from Croatia) and A. pappi

from that country, in addition to A. leucopeza. In this study, 

we have only had access to a limited sample of the European 

fauna of Aulacigastridae. We agree with Papp (op. cit.) that 

a more extensive study of specimens from the Palaearctic 

Region is necessary to clarify the distributional ranges of 

the various species concerned. Moreover, the identity of the 

European species, particularly A. pappi (see remarks under 

A. pappi), needs further investigation, including location 

and study of any extant primary types and possibly involving DNA sequence data. The junior synonyms previously 
108 • smithsonian contributions to zoology

FIGURES 188–191. Digital photographs of Aulacigaster leucopeza (Meigen) (male) (the leucopeza group, Palearctic Region). 

(188) body, lateral aspect; (189) head and mesonotum, dorsal aspect; (190) head, frontal aspect; (191) head, lateral aspect. Not 

all to the same scale.
number 633 • 109

noted, for example, may represent older names for species 

Papp described. For these reasons and not wanting to promulgate unverified and potentially false information, we 

have not produced distribution maps for species occurring 

in the Palaearctic Region.

Aulacigaster pappi Kassebeer

Figures 194–199

Aulacigaster neoleucopeza of authors, not Mathis and Freidberg [misidentification].—Papp, 1998a:230–232 [discussion, list, Hungary].—Bächli et 

al., 1999:121 [list, Switzerland].

Aulacigaster pappi Kassebeer, 2001:26 [description, list, France, Switzerland].

Diagnosis. This species is distinguished from 

congeners by the following combination of characters: 

Posterior fronto-orbital seta slightly medial to and almost 

horizontally aligned with anterior seta; shiny spot adjacent to ocellar tubercle extended from ocellus beyond halfway to eye margin; surstylus digitiform, tapering towards 

blunt apex, relatively short (approx. 0.2 times the length 

of ventral margin of epandrium), in lateral view with 

apex slightly elevated in relation to the ventral margin of 

epandrium.

Description. Coloration and vestiture: Vertex 

mostly densely microtomentose, with a shiny spot extended 

from ocellus to near eye margin (shiny spot separated from 

eye margin by a distance approximately equal to width of 

one ocellus); ocellar tubercle dull microtomentose; frons 

bearing a transverse, large, orange band, frontal orange band 

shorter than 1/2 distance of lunule to anterior ocellus, merging into dark brown ventrally; silver stripe on frons approximately straight; antenna pale yellow to yellowish, infuscate 

along dorsal margin or infuscate on dorsal half; face yellow, 

with a transverse brown mark on dorsal portion; facial band 

present, sharply defined, transverse, on protruding, ventral 

portion of face; palpus white. Scutum with weakly defined 

microtomentose stripes; scutellum dull microtomentose; 

postpronotum concolorous with mesonotum, dull microtomentose; anepisternum mostly dull; katepisternum mostly 

microtomentose on anterior half and on a narrow stripe over 

katepisternal setae; halter mostly white. Fore coxa brown 

to black, fore femur brown to black, fore tibia brown, fore 

tarsus yellowish, apical and sometimes subapical tarsomeres 

brown to black; mid coxa brown to black, mid femur brown 

to black, mid tibia completely brown or yellowish on central 

portion, mid tarsus yellowish, apical and subapical tarsomere brownish to dark brown; hind coxa brown, hind femur 

brown to black, hind tibia completely brown to dark brown, 

or with basal and central portion yellowish, hind tarsus 

FIGURES 192–193. Illustrations of Aulacigaster leucopeza (Meigen) (male) (the leucopeza group, 

Palearctic Region). (192) gonopod, lateral aspect; (193) epandrium, lateral aspect.
110 • smithsonian contributions to zoology

yellowish, apical tarsomere brown to black. Male abdomen 

subshiny to dull; female abdomen with dense microtomentum medially, lateral margins of tergites nearly polished.

Morphology: 

Head: Figures 195–197. Head higher than long 

(head ratio less than 0.9); face at level of pseudovibrissa 

wider than width of 1st flagellomere, in lateral view with 

dorsal half concave, ventral portion bulbous; gena in lateral 

view approximately 1/3–1/2 the width of 1st flagellomere; 

medial vertical seta 3/4 length of lateral vertical seta; peristomal area with one long and 3–4 shorter setae following 

pseudovibrissal setae.

Thorax: Figures 194, 196. Acrostichal setae in 

2 rows, these convergent anteriorly at level of anterior 

FIGURES 194–197. Digital photographs of Aulacigaster pappi Kassebeer (male) (the leucopeza group, Palearctic Region). (194) body of male, lateral aspect; (195) head, dorsal aspect; (196) head and anterior portion of mesonotum, 

lateral aspect; (197) head, frontal aspect. Not all to the same scale.
number 633 • 111

dorsocentral seta; scutellum slightly raised (angle with scutum approximately 135°), trapezoidal, apex nearly straight, 

disk of scutellum slightly convex; basal scutellar seta 1/2 

length of posterior seta.

Male abdomen and terminalia: Figures 198–199. 

Epandrium not densely setose. Surstylus arising from ventral margin of epandrium, slender, digitiform, tapering towards blunt apex, relatively short (approx. 0.2 times the 

length of ventral margin of epandrium); surstylus in lateral 

view with apex slightly elevated in relation to the ventral 

margin of epandrium (fig. 2d; Kassebeer, 2001:31); gonopod with several strong setae that are as long as the surstylus; subepandrial sclerite with lateral arms narrow, well 

sclerotized, curved evenly posteriorly; cerci completely separated, more developed ventrally, approximately tongueshaped in lateral view.

Type Material. The holotype male is labeled 

(quoted from Kassebeer’s paper) “Deutschland: Brandenburg: [7.6]–5.7.2001, Brüsenwalde, BF, leg. G. Möller, Coll. 

C. F. Kassebeer.”

Type Locality. Germany: Brandenburg, Brüsenwalde (53°13¢N, 13°28¢E).

Other Specimens Examined. HUNGARY. Nógrád: Szendehely, Keskeny-bükki p.v. tölgyerdo ̋k 

(47°51.3¢N, 19°06.3¢E), Jun 1996, L. Papp (2♂; CSCA).

Distribution. Palearctic. France, Germany, 

Hungary, Switzerland.

Biology. This species has been recorded from 

Quercus sp. (Kassebeer, 2001). It has also been collected by 

beer/wine traps (Kassebeer, 2001).

Remarks. The description provided here is 

a combination of the diagnosis published by Kassebeer 

(2001) and study of two specimens from Hungary previously identified by L. Papp as A. neoleucopeza. Kassebeer 

(2001) concluded that reports of A. neoleucopeza occurring 

in Europe (Papp, 1998a) were based on misidentifications 

and proposed the name of A. pappi for these specimens. 

Kassebeer’s discovery of specimens collected by Thalhammer over 100 years ago (DEI) also invalidates the idea of 

a recent introduction of A. neoleucopeza into the Palearctic Region. While we are accepting Kassebeer’s proposals, 

being the most recent treatment of this species, we have 

not had access to the specimens that Kassebeer examined 

and cannot confirm his findings. With the exception of 

three females deposited in the Muséum d‘histoire naturelle, 

Genève, and one male deposited in DEI, the material studied by Kassebeer is entirely held in two private collections: 

Kassebeer’s and C. Bächli’s. The only male available from 

institutional collections (DEI) was borrowed by Kassebeer 

some years ago and has not yet been returned.

FIGURES 198–199. Illustrations of Aulacigaster pappi Kassebeer (male) (the leucopeza group, Palearctic Region). (198) gonopod, lateral aspect; (199) epandrium, lateral 

aspect.
112 • smithsonian contributions to zoology

Aulacigaster india, new species

Figures 200, 219–220, Map 18

Diagnosis. This species is distinguished from 

congeners by the following combination of characters: 

Arista short, with tiny, dense, alternate hairs on apical 2/3, 

basal 1/3 of arista naked; scutum polished on anteriormost 

portion, opposite to postcranium.

Description. Coloration and vestiture: Vertex 

mostly densely microtomentose, with a shiny spot extended 

from ocellus to 2/3 distance to eye margin; ocellar tubercle 

dull microtomentose; frons bearing a transverse, large, orange band, frontal orange band merging into dark brown 

ventrally (dark brown region interrupted in some specimens); silver stripe on frons approximately straight, weak; 

antenna pale yellow to yellowish, infuscate along dorsal 

margin; face yellow, with a transverse brown mark on dorsal portion; facial band present, sharply defined, transverse, 

on protruding, ventral portion of face; palpus white. Scutum 

mostly uniformly microtomentose, polished on anteriormost 

portion, opposite postcranium; scutellum dull microtomentose; postpronotum concolorous with mesonotum, dull 

microtomentose; anepisternum mostly dull; katepisternum 

with anterior portion dull microtomentose, and posterior 

portion mostly polished; halter mostly white, or mostly yellowish (knob with brown area). Fore coxa brown to black, 

fore femur brown (tip yellowish); fore tibia yellowish (tip 

and sometimes basis dark), fore tarsus yellowish, apical 

and sometimes subapical tarsomeres brown to black (apical and subapical); mid coxa brown to black, mid femur 

brown to black, apex yellowish, mid tibia yellowish on central portion, apex and basis brown (tip, and sometimes basis 

brown), mid tarsus yellowish-brownish; hind femur brown 

to black, apex yellowish, hind tibia mostly yellow, base and 

apex infuscate with brown, hind tarsus yellowish, apical tarsomere brown to black (apical and subapical).

Morphology: 

Head: Figures 219–220. Head higher than long 

(head ratio less than 0.9); face at level of pseudovibrissa 

wider than width of 1st flagellomere, in lateral view with 

dorsal half concave, ventral portion bulbous; gena in lateral 

view approximately 1/3–1/2 the width of 1st flagellomere; 

medial vertical seta 3/4 length of lateral vertical seta; peristomal area with about 1 long and 3–4 smaller setae following pseudovibrissal seta.

Thorax: Acrostichal setae in 2 rows; scutellum 

slightly raised (angle with scutum approximately 135°), 

The specimens we studied from Hungary are generally darker than specimens of A. neoleucopeza and A. leucopeza, two species with which A. pappi could be easily 

confused. In the latter species, the apices of the coxae are 

not pale brown to yellow as in the former two species. The 

orange band on the frons is slightly darker, and the microtomentose band on the lower frons sits on a brown mark 

that is wider than that present in Palearctic and Nearctic 

species. The facial band on the lower face of A. pappi is 

weakly microtomentose and tends to be more yellow than 

in other temperate species of the leucopeza group. Aulacigaster pappi is more easily confused with and may be 

conspecific with A. neoleucopeza. The shape of the gonopod and posterodorsal gonopodal process, including the 

arrangement of the setae preceding the process, is within 

the range of variation found for A. neoleucopeza. Even 

though both species can be separated by the characters 

given herein, it is possible that analysis of additional material of A. pappi will reveal intermediate forms, making the 

possibility of this distinction tenuous.

Oriental Species

Discussion. The presence of undescribed 

Aulacigaster in the Oriental Region was previously documented (Sabrosky, 1977). Two species are described herein, 

and one other is reported. All species are classified in the 

leucopeza group.

Key to the Oriental Species of Aulacigaster

1. Vertex mostly microtomentose; facial band absent; scutum with weakly defined microtomentose stripes (Nepal) . . . . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. L3, undescribed species

Vertex with a shiny area between posterior ocellus and margin of eye; facial band present [Figure 219]; scutum without 

obvious stripes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2. Shiny spot of vertex small, extended from ocellus at most 1/2 way to eye margin; scutum uniformly microtomentose (SriLanka) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. srilanka, new species

Shiny spot of vertex large, extended from ocellus 2/3 to eye margin [Figure 219]; scutum mostly uniformly microtomentose, but polished opposite to postcranium (India) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. india, new species
number 633 • 113

trapezoidal, apex nearly straight, disk of scutellum strongly 

convex; basal scutellar seta 1/2 length of posterior seta.

Male abdomen and terminalia: Figure 200. Surstylus arising from posteroventral to ventral margin of epandrium, slender, digitiform, relatively long (approx. 0.5 times 

the length of ventral margin of epandrium), margins almost 

parallel, apex round; cerci broad, with apex bearing a few 

preeminent setae; gonopod sub-quadrate in lateral view 

(approximately as high as long); posterior medial process 

of gonopod approximately digitiform, wide (at least 3 times 

wider than surstylus) bearing ca 11 strong setae.

Measurements: Body length 2.4–2.7 mm. Wing 

length 2.4–2.7 mm, wing width 1.0 mm.

Type Material. The holotype male is labeled 

“INDIA: Meghalaya[,] Mawphlang (20 km W. Shillong)[,] 

21Ap[ril] 1980[,] A[.]Freidberg/HOLOTYPE ♂ Aulacigaster india Rung & Mathis USNM [red].” The holotype is 

double mounted (minuten in a block of plastic), is in excellent condition (abdomen removed and dissected with 

structures in an attached microvial), and is deposited in the 

USNM. Paratypes are as follows: Same label data as the 

holotype (3♀; USNM).

Type Locality. India. Meghalaya: Shillong 

(20 km W, road from Shillong to Mawphlang; 25°34¢N, 

91°45¢E).

Distribution. (Map 18) India (Meghalaya).

Etymology. The specific epithet, india, refers 

to the country where the type series was collected. The 

name is a noun in apposition.

Aulacigaster srilanka, new species

Figure 201, Map 18

Diagnosis. This species is distinguished from 

congeners by the following combination of characters: 

Gena very narrow, in lateral view approximately 1/5 the 

width of 1st flagellomere below ventral margin of eye; surstylus very reduced, inconspicuous.

Description. Coloration: Vertex mostly 

densely microtomentose, with a shiny spot extended from 

ocellus at most 1/2 way to eye margin; ocellar tubercle dull 

microtomentose; frons bearing a transverse, large, orange 

band; silver stripe on frons approximately straight, bright 

shiny; antenna pale yellow to yellowish, infuscate along dorsal margin; face yellow, with a transverse brown mark on 

dorsal portion; facial band present, sharply defined, transverse, on protruding, ventral portion of face; palpus white. 

Scutellum dull microtomentose; postpronotum concolorous with mesonotum, dull microtomentose; anepisternum 

mostly dull; katepisternum mostly microtomentose, with 

a shiny area posteriorly, below the posterior katepisternal 

seta; halter mostly brown to dark brown. Fore coxa brown 

to black; mid coxa brown to black, mid femur brown, mid 

tarsus yellowish, apical and subapical tarsomere brownish 

to dark brown; mid coxa brown, mid femur brown, mid 

tarsus yellowish; hindlegs missing in the holotype. Male abdomen bright shiny.

Thorax: Acrostichal setae in 2 rows; scutellum 

slightly raised (angle with scutum approximately 135°), 

trapezoidal, apex nearly straight, disk of scutellum slightly 

convex; basal scutellar seta less than 1/2 length of posterior seta.

Male abdomen and terminalia: Figure 201. Posteroventral margin of epandrium bearing strong setae 

surstylus reduced, inconspicuous; posterodorsal process 

of gonopod wide, digitiform, bearing approximately 11 

strong setae.

Measurements: Body length 2.7 mm. Wing length 

2.4 mm, wing width 0.7 mm.

Type Material. The holotype male is labeled “SRI LANKA. Dist. Kandy[,] Udawattakele Sanct.[,] 

FIGURE 200. Illustrations of Aulacigaster india, new species (male) 

(the leucopeza group, Oriental Region): epandrium and gonopod, 

lateral aspect.
114 • smithsonian contributions to zoology

22-III-1981 [22 Mar 1981]/K.V. Krombein[,], T. Wijesinhe[,] 

L. Weeratunge/HOLOTYPE ♂ Aulacigaster srilanka Rung 

& Mathis USNM [red].” The holotype is double mounted 

(minuten in a block of polyporus), is in good condition 

(abdomen removed and dissected with structures in an attached microvial), and is deposited in the USNM.

Type Locality. Sri Lanka. Central: Kandy, 

Udawattakele Sanctuary (07°18¢N, 80°39¢E).

Distribution. (Map 18) Oriental: Sri Lanka.

Etymology. The specific epithet, srilanka, refers to the country where the type series was collected. The 

name is a noun in apposition.

Aulacigaster L3, undescribed species

Map 18

Diagnosis. This species is distinguished from 

congeners by the following combination of characters: 

Arista short, with very inconspicuous, alternate hairs (arista 

appearing naked at low magnification); facial band absent; 

frons bearing a transverse, large, orange band.

Description. Coloration: Vertex mostly 

densely microtomentose, without a shiny spot (a small 

dot lateral to posterior ocellus is however present); ocellar 

MAP 18. Distribution of the Oriental species of the leucopeza group of Aulacigaster.
number 633 • 115

tubercle dull microtomentose; frons bearing a transverse, 

large, orange band, frontal orange band with a blackish 

mark basal to ocellar tubercle, and a pair of blackish marks 

on ventrolateral corners, above ptilinal fissure; anterior 

portion of frons densely microtomentose, with a faint, narrow, anterior silvery stripe, not followed by a black stripe 

or shadow; antenna pale yellow to yellowish, infuscate on 

dorsal half or infuscate along dorsal margin; face yellow 

anteriorly, brown to black on posterior portion; palpus 

white. Scutum with weakly defined microtomentose stripes; 

scutellum dull microtomentose; postpronotum yellow to 

yellowish-brownish, dull microtomentose; anepisternum 

mostly dull (with a silver, transverse, ill-defined stripe); katepisternum mostly polished, or with anterior portion dull 

microtomentose, and posterior portion mostly polished; 

halter mostly white (base slightly infuscate). Fore coxa 

brown to black, fore femur brown to black, apex yellowish, fore tarsus yellowish, apical and sometimes subapical 

tarsomeres brown to black; mid coxa brown to black, mid 

femur brown to black, apex yellowish, mid tarsus yellowish, apical and subapical tarsomere brownish to dark 

brown; hind femur brown to black, apex yellowish, hind 

tarsus yellowish, apical tarsomere brown to black. Female 

abdomen subshiny.

Morphology: 

Head: Head round, about as high as long (head 

ratio 0.9–1.1); face at level of pseudovibrissa wider than 

width of 1st flagellomere, in lateral view with dorsal half 

concave, ventral portion bulbous; gena in lateral view approximately 1/3–1/2 the width of 1st flagellomere; medial 

vertical seta 3/4 length of lateral vertical seta; peristomal 

area with about 4 setae following pseudovibrissal seta.

Thorax: Acrostichal setae in 1 row, bifurcating 

posteriorly; scutellum slightly raised (angle with scutum approximately 135°), trapezoidal, apex nearly straight, disk 

of scutellum slightly convex; basal scutellar seta 1/2 length 

of posterior seta.

Measurements: Body length 2.7–2.8 mm. Wing 

length 2.4–2.5 mm, wing width 0.9–1.0 mm.

Specimens Examined. NEPAL. Canadian 

Nepal Expedition (28°00¢N, 85°00¢E; Malaise trap), 26 

May 1967 (2♀; CNC).

Distribution. (Map 18) Oriental: Nepal.

Remarks. This species is not described because it 

is represented by two females only.

Phylogenetic Analysis and 

Discussion

Characters Used in the Cladistic Analysis

1. A transverse, sharp orange band on ventral frons: 

1. absent; 2. present.

2. A transverse, silver microtomentose stripe on 

ventral frons: 1. absent; 2. present as a straight 

microtomentose stripe covering ventral 1/4; 3. 

present as a semi-lunate microtomentose band 

covering ventral 1/3–1/2.

3. Transverse, silver facial band on ventral portion 

of face, extended to gena: 1. absent; 2. present.

4. Microtomentum coverage of scutum: 1. uniformly 

covering most of scutum; 2. present only between 

dorsocentral rows of setae; 3. present only on anterior 1/3–1/2 of scuttum; 4. nearly absent. 

5. Coloration of thoracic pleural region: 1. uniformly black; 2. white to pale yellow on ventral 

half.

6. Coloration of abdominal tergites 1+2: 1. without 

a yellowish region dorsally; 2. with a yellowish 

FIGURE 201. Illustrations of Aulacigaster srilanka, new species 

(male) (the leucopeza group, Oriental Region): epandrium and 

gonopod, lateral aspect.
116 • smithsonian contributions to zoology

region dorsally that sometimes extends over tergite 3.

7. Shape of body: 1. robust, scutal ratio 1.1–1.35; 2. 

elongate, scutal ratio 1.5–1.75.

8. Shape of vertex, between compound eyes: 1. 

nearly straight; 2. excavated.

9. Margins of eye: 1. not encroaching on the posterior portion of the frons; 2. encroaching on the 

posterior portion of the frons.

10. Clypeus (proportions): 1. stout, 1.5 to 2.5 times 

longer than wide; 2. elongate, 3.5–5.5 times longer than wide.

11. First flagellomere (shape): 1. round; 2. oval, decumbent; 3. triangular.

12. Arista (shape): 1. naked; 2. with very inconspicuous, alternate hairs (arista appearing naked at 

low magnification); 3. distinctly zigzagged on 

apical 2/3 and bearing short hairs; 4. distinctly 

zigzagged on apical 2/3 and bearing long hairs.

13. Dimorphism in the shape of the face: 1. absent; 

2. present.

14. Ocellar seta: 1. absent; 2. present.

15. Development of ocellar seta: 1. weak; 2. strong, 

seta-like.

16. Subcosta: 1. complete, fused with vein R1

 apically, 

and reaching costal vein; 2. incomplete, fused 

with vein R1 apically, not reaching costal vein.

17. Length of vein R2+3: 1. long, ending at or near tip 

of the wing; 2. short, ending on basal 2/3 of the 

wing.

18. Hind femur: 1. without a basal lobe; 2. with a 

ventral basal lobe.

19. Hind femur: 1. not modified, and without ventral protrusions bearing setae; 2. modified, with 

two ventral rows of protrusions, each protrusion 

bearing an apical seta.

20. Mesonotal setulae: 1. not organized in rows; 2. 

organized in rows.

21. Position of surstylus: 1. posterior to postero-ventral; 2. ventral.

22. Setae on gonopodal process: 1. absent or hairlike; 2. well developed, long; 3. strongly developed, long and thick.

23. Ventral portion of gonopod: 1. not modified; 

2. distinguished from medial portion, forming 

a weakly sclerotized ventral plate; 3. forming a 

strongly sclerotized, black ventral plate.

24. Number of anterior gonopodal processes: 1. one; 

2. two.

Discussion

The 17 optimal trees (Figure 202) resulting from an 

exhaustive search have 39 steps, a consistency-index of 

0.82, and retention index of 0.93. Successive weighing did 

not change the number of optimal trees or their topologies. After the third iteration of character weighing, the resulting 17 trees had 30.60 steps, CI = 0.89, and RI = 0.95.

The main clades supported by this analysis are the 

following. First, the genus Aulacigaster, in our rooting, is 

monophyletic. Second, all of the species groups of Aulacigaster represented by more than one exemplar were recovered, indicating that the groups proposed in this revision 

are natural groups. The most striking differences in the 

topologies of the best trees obtained concern the relationships between the plesiomorphica and bromeliae groups 

and the remaining species groups of Neotropical Aulacigaster, which remain largely unresolved. The relationships between species of the widespread leucopeza group, 

which lie beyond the scope of this analysis (see “Methods 

and Materials”), also remain unresolved and account for 

most of the conflicting topologies. For discussions of phylogenetic distribution of character states below, refer to 

Figure 202, which corresponds to PAUP tree number 1. 

This tree was optimized using accelerated transformation 

(ACTRAN).

In this analysis, the genus Aulacigaster is supported 

by three unambiguous synapomorphies: arista distinctly 

zigzagged on apical 2/3 and bearing short hairs (character 

12, state 3; Figures 51, 59); ocellar seta reduced, weak 

(character 15, state 1; Figure 11); and mesonotal setulae 

organized in rows (character 20, state 2; Figures 1, 12). 

In the sister-group Curiosimusca the arista is straight and 

naked, the ocellar seta is strongly developed, and the mesonotal setulae are randomly distributed rather than organized in rows. Apart from the zigzagged arista with short 

ray-like hairs, all characters supporting the monophyly 

of Aulacigaster in this analysis are found in presumably 

closely related families such as Periscelididae sensu lato

(including Stenomicra, Cyamops and Planinasus) and Anthomyzidae. The monophyly of Aulacigaster and its sistergroup relationship to Curiosimusca have been discussed 

elsewhere (Rung et al., 2005) and the reader should consult that contribution for additional synapomorphies and 

discussion about them.

The widely distributed leucopeza group is supported 

by two unambiguous synapomorphies: the presence of an 

orange band on ventral frons (character 1, state 2; Figure 

217), a non-homoplasious synapomorphy; and the presence 
number 633 • 117

of a silver microtomentose stripe on the ventral portion of 

the frons (character 2, state 2; Figure 217). Both characters 

are absent in Curiosimusca and closely related families. 

Species of the grimaldii group have a transverse microtomentose stripe on the lower frons that is similar to that 

found in the leucopeza group, a convergence according to 

this analysis (Figure 209). In species of the plesiomorphica

group, a wide, semi-lunate band covering a larger portion 

of the lower (ventral) frons is present (Figure 213). Because 

we believe that this condition is distinct from that found 

in the leucopeza group, we have coded it as a separate 

state (state 3). Two species of the leucopeza group, both 

Neotropical in distribution (A. L1, undescribed species 

and A. spangleri), have the frontal orange band partially 

obliterated laterally by black infuscation (Figure 215), a 

putative synapomorphy for these two species (not coded). 

These two closely related species also share a nearly naked 

arista (character 12, state 2; Figure 216).

The exclusively Neotropical groups of Aulacigaster

are united by two unambiguous synapomorphies: face 

without a sharp silver band on the ventral portion that 

extends to the gena (character 3, state 1; Figure 203); and 

vein R2+3 short, not reaching the tip of the wing (character 

17, state 2; Figure 2). Species of both Curiosimusca and 

the leucopeza group of Aulacigaster have a sharp and distinctive facial band that extends posteriorly to the gena 

(Figure 219), and vein R2+3 reaching nearly to apex of the 

wing (Figure 188). Both synapomorphies correspond to 

widespread conditions in presumably closely related families, and the second synapomorphy is reversed within the 

ecuadoriensis group (see below), which has a long vein 

R2+3 also nearly reaching the wing apex (Figure 52). Several species of the plesiomorphica group have a short, 

sometimes inconspicuous facial band located between the 

pseudovibrissae (Figure 213) and differing in appearance 

and location from the facial band present in the leucopeza

group and Curiosimusca.

The bromeliae group of Aulacigaster is well characterized by five unambiguous synapomorphies. Besides 

their unique natural history traits (see in introduction and 

under “the bromeliae group” above), species of this group 

share the following synapomorphies: an excavated vertex 

(character 8, state 2; Figure 203); eyes encroaching on the 

dorsal portion of the frons (character 9, state 2; Figure 

203); an elongated clypeus (character 10, state 2; Figure 

21); an oval, decumbent 1st flagellomere (character 11, 

state 2; Figure 4); and an arista with long rays (character 

12, state 4; Figure 4). The oval 1st flagellomere with an 

arista bearing long rays is a widespread condition in the 

Periscelididae sensu lato and also occurs in the Anthomyzidae. The remaining characters are apparently unique to 

the bromeliae group.

The monophyly of the plesiomorphica group is supported by two synapomorphies: (1) a strong, sharp, semilunate microtomentose band on the ventral portion of the 

frons (character 2, state 3; Figure 213), unique to this 

group; (2) the presence of unusually strong setae on one 

posterior process of the gonopod (character 22, state 3; 

Figure 132). The last character is also present in species 

of the ecuadoriensis group but this is hypothesized in 

this analysis to be convergent. Characters 22 and 24 may 

be dependent (linked), because generally whenever two 

processes are present on the gonopod, at least one bears 

strong setae. There are a few exceptions to this rule, however. For example, A. belize, in the plesiomorphica group 

(Figure 137), has only one posterior gonopodal process, 

and it has strong setae; also, A. india (Figure 200), A. 

srilanka (Figure 201), and A. L3 (not coded), species of 

the leucopeza group, have relatively well-developed, long 

setae on the only posterior process of their gonopod. 

Until there is evidence showing that characters 22 and 

24 are liked, we have chosen to code them as separate 

characteres.

The remaining groups of Aulacigaster (the grimaldii, 

ecuadoriensis, femorata, and minuta groups) together 

share one unambiguous (albeit homoplasious) synapomorphy, reduction in the subcosta (character 16, state 2; 

Figure 85). In Curiosimusca, as well as in species of the 

leucopeza, bromeliae, and plesimorphica groups, the subcosta is partially fused with vein R1

, but still reaches the 

Costa (state 2; Figures 3, 150). Character state 1 (subcosta 

incomplete) is also present in the ecuadoriensis group, but 

as the result of reversal based on our analysis. A subcostal 

vein partially fused with vein R1 is present in presumably 

related families, such as the Anthomyzidae and Opomyzidae, but whether the Sc vein is complete or incomplete to 

the Costa in these families was not ascertained by us, and 

would not change our results. The groups grimaldii, ecuadoriensis, femorata, and minuta also share a reduction 

in the microtomentum coverage of the scutum (character 

4, states 2–4), but the optimization of the corresponding 

states are ambiguous in our cladogram.

The two known species of the grimaldii group form a 

monophyletic group supported by two unambiguous synapomorphies: the presence of a microtomentose stripe on 

the lower frons (character 2, state 2; Figure 209), a condition similar to but not homologous with the condition 

found in the leucopeza group; and sexual dimorphism of 
118 • smithsonian contributions to zoology

Curiosimusca khooi

C.orientalis

A.angusta

A.grimaldii

A.bromeliae

A.korneyevi

A.plesiomorphica

A.trifasciata

A.ecuadoriensis

A.melanololeuca

A.stenoptera

A.bella

A.femorata

A.lobata

A.appendiculata

A.minuta

Aulacigaster L1

A.spangleri

A.india

A.srilanka

A.neoleucopeza

A.mcalpinei

A.fredbergi

A.malawana

A.falcata

A.leucopeza

18

1

16

1

23

3

The leucopeza 

 group

The plesiomorphica 

 group

The bromeliae

 group

The grimaldii 

 group

The femorata

 group

The minuta group

 The 

ecuadoriensis

 group

NE

 

NE

NA

AT

PA

OR

22

3

2

3

24

2

14

1

7

2

3 1 1 3

11 17 21 22

2 2

6 23

2 2 2

18 19 21

20

2

15

1

12

3

2

2

1

2

2

12

1

3

3

1

17

2

8 9 10 11 12

2 2 2 2 4

16

2

2

2

13

2

5

2
number 633 • 119

the face (character 13, state 2; Figures 103–104). An additional synapomorphy for these two species that is present 

only under delayed transformation optimization (DELTRAN optimization) is the microtomentose coverage of 

the scutum, present only between the dorsocentral rows of 

setae (character 4, state 2).

Certain modifications of the hindlegs support the 

monophyly of a clade comprising the ecuadoriensis, 

femorata, and minuta groups. In species of this clade, the 

hind femur of males and females is enlarged and has two 

ventral rows of well-defined protrusions, each bearing an 

apical seta (character 19, state 2; Figure 66). The male 

hind femur also bears a basal lobe with setae (character 

18, state 2) that in some species is preceded by a basal 

concavity (Figure 66). The lobe is reduced in some species, 

as in A. melanoleuca (Figure 43). Such unique modifications of the hind femur are not present in closely related 

groups outside Aulacigastridae. A third synapomorphy 

for this assembly is in the ventral position of the surstylus 

(character 21, state 2; Figure 68). In other groups of Aulacigaster, the surstylus is located posteroventrally. This 

synapomorphy is reversed in A. ecuadoriensis + A. melanoleuca, being also present in two species not represented 

in our analysis (A. aenigma and A. fastidiosa). The latter have a lobe-like, posteroventral surstylus (Figure 38), 

a condition similar to that found in the plesiomorphica

group (Figure 143).

Species of the femorata group are united by two unambiguous synapomorphies: the presence of a white to 

yellowish region on tergites 1+2 that sometimes extends 

over tergite 3 (character 6, state 2; Figure 62); and the ventral portion of the gonopod partly separated from the medial portion, forming a weakly differentiated ventral plate 

(character 23, state 2; Figure 67). This last character state 

seems at least superficially to be homologous with that 

found in the minuta group (state 3; Figure 111), whose 

species have a strongly sclerotized ventral plate, but the 

results of our analysis suggest that these structures have 

developed independently. Coding character 23 as ordered 

to force one step between states 2 and 3 does not change 

the results of this analysis. An additional synapomorphy 

for the femorata group that is present under DELTRAN 

optimization only is the microtomentose coverage of the 

scutum, present only on the anterior portion (character 

4, state 3).

Species belonging in the minuta and the ecuadoriensis

groups have an elongated body (character 7, state 2; Figures 40, 107) and show the loss of the ocellar seta (character 14, state 1). Under DELTRAN optimization, they 

share an additional synapomorphy, the almost complete 

lack of microtomentum on the scutum (character 4, state 

4). Both groups have other similarities, apparently correlated with a compressed body: head longer than high, 

face not protrudent, wing unusually longer than wide, etc. 

To which extent a complete reduction in the ocellar setae 

and compression of the body are correlated in these two 

groups cannot be determined with certainty for the present. We know, however, that reduction of the ocellar seta 

is widespread in closely related groups that do not have a 

dorsoventrally compressed body, and dorsoventrally compressed species of Neurochatetidae have long ocellar setae. 

Therefore, we have decided to treat these two characters 

as independent in this analysis.

Species of the ecuadoriensis group are unique among 

Aulacigaster in the coloration of the thoracic pleural region, which is white on the ventral half (character 5, state 

2; Figure 40). This state is absent in related families as 

well as in Curiosimusca and other species of Aulacigaster. Other synapomorphies for this group are the presence 

of two posterior processes on the gonopod (character 24, 

state 2; Figure 30), a convergence with the plesiomorphica

group, and the subcostal vein reaching the costal margin (character 16, state 1; Figure 53), which is a reversal. Within the ecuadoriensis group, A. ecuadoriensis and 

A. melanoleuca are sister-groups. They share at least one 

non-homoplasious synapomorphy: a triangular 1st flagellomere (character 11, state 3; Figure 206). Other synapomorphies shared by these two species are reversals, as the 

long vein R2+3 (character 17, state 1; Figure 41), or convergences with character states found in the plesiomorphica

group, such as the position of the surstylus (character 21, 

states 1) and the presence of strongly developed, long and 

thick setae on the posterior process of the gonopod (character 22, state 3; Figure 30).

The minuta group is unique for having a dark brown 

structure on the ventral portion of the gonopods (character 

FIGURE 202. (Facing page) One of 17 trees obtained with branch-and-bound search, unordered characters, chosen to represent the phylogenetic relationships between species groups of Aulacigaster (39 steps, consistency index = 0.82 and retention index = 0.93). Only unambiguous 

synapomorphies are represented. Empty squares represent homoplasious synapomorphies; filled circles represent non-homoplasious synapomorphies. Branches that are not supported by any characters are shown as collapsed, and a filled star shows where branches collapse on the 

strict consensus cladogram.
120 • smithsonian contributions to zoology

23, state 3; Figure 111). Species of this group are also unusually small and delicate, a condition quite different from 

that found in the ecuadoriesis group.

Even though this contribution has greatly increased 

the number of known species of Aulacigaster and expanded our knowledge about the distribution of the 

genus worldwide, we believe that much more collecting 

is needed to reveal the distribution patterns of the various species groups, particularly in the Neotropical Region. 

Most species reported in this contribution come from Central America and the northern portion of South America, 

giving the impression that the species groups to which 

they belong are primarily distributed in these regions. For 

example, species of the relatively rare minuta group are 

only known from Central America and one South American country, Bolivia. Out of the seven Neotropical species 

groups recognized in this study, only one, the bromeliae

group, is reported from the southern part of South America, and except for a few scattered specimens collected in 

Rondônia (in the frontier with Bolivia), the ecuadoriensis, the femorata, and the plesiomorphica groups are unknown from Brazil or from anywhere south of Bolivia. 

These distribution patterns, rather than reflecting true biogeographic history and ecological constraints, are probably a function of sampling error and collecting biases, the 

lack of good acalyptrate collections in museums of several 

countries in South America, and difficulties obtaining collecting and export permits from countries such as Brazil.
number 633 • 121

FIGURES 203–220

FIGURES 203–208. Digital photographs of heads of Aulacigaster

species. (203) A. bromeliae, frontal aspect; (204) A. bromeliae, lateral 

aspect; (205) A. ecuadoriensis, frontal aspect; (206) A. ecuadoriensis, 

lateral aspect; (207) A. femorata, frontal aspect; (208) A. femorata, 

lateral aspect.
122 • smithsonian contributions to zoology

FIGURES 209–214. Digital photographs of heads of Aulacigaster species. (209) A. grimaldii, frontal aspect 

of male; (210) A. grimaldii, lateral aspect; (211) A. minuta, frontal aspect; (212) A. minuta, lateral aspect; 

(213) A. proxima, frontal aspect; (214) A. proxima, lateral aspect.
number 633 • 123

FIGURES 215–220. Digital photographs of heads of Aulacigaster species. (215) 

A. spangleri, frontal aspect; (216) A. spangleri, lateral aspect; (217) A. leucopeza, 

frontal aspect; (218) A. leucopeza, lateral aspect; (219) A. india, frontal aspect; (220) 

A. india, lateral aspect.

Displayed here is a Nexus file (including 

character matrix and PAUP commands) 

used to generate the phylogenetic analysis 

in PAUP.

Appendix: Character Matrix

#NEXUS

BEGIN DATA;

DIMENSIONS NTAX=26 NCHAR=24;

FORMAT SYMBOLS= “0 1 2 3 4 5” MISSING=? GAP=- ;

MATRIX 

 123456789012345678901234

C.khooi 112111111111122111111111

C.orientalis 112111111111122111111111

A.angusta 121211111113221221121111

A.grimaldi 121211111113221221121111

A.bromeliae 111111122224121121121111

A.korneyevi 111111122224121121121111

A.plesiomorphica 131111111113121121121311

A.trifasciata 131111111113121121121311

A.ecuadoriensis 11142121113311-112221312

A.melanololeuca 11142121113311-111221312

A.stenoptera 11142121111311-122222112

A.bella 111312111113121222222121

A.femorata 111312111113121222222121

A.lobata 111312111113121222222121

A.appendiculata 11141121111311-222222131

A.minuta 11141121111311-222222131

A.L1 22211111111212111112??11

A.spangleri 22111111111212111112??11

A.india 222111111113121111121211

A.srilanka 22211111111?12111112-211

A.neoleucopeza 222111111113121111121-11

A.macalpinei 222111111113121111121111

A.fredbergii 222111111113121111121111

A.malawana 222111111113121111121111

A.falcata 222111111113121111121111

A.leucopeza 222111111113121111121111

;

END;

Set AUTOCLOSE;

LOG file = aulaciphylo.log;

bandb;

savetrees file=bandb.tre;

contree;

References

Bächli, G. 1997. Die Arten der Tanypezidae, Dryomyzidae, Periscelididae, Acartophtalmidae, Aulacigastridae and Stenomicridae in der Schweiz (Diptera). Mitteilungen 

der Entomologischen Gesellschaft Basel, 47(1):29–34.

———. 1998. “Aulacisgaster.” In Diptera–Checklist, ed. B. Merz, G. Bächli, J.-P. Haenni, 

and Y. Gonseth, pp. 288, 369. Fauna Helvetica 1. Basel: Schweizerische Entomologische Gesellschaft, Société Entomologique Suisse.

Bächli, G., L. Papp, and S. Vanin. 1999. New Records of Aulacigastridae and Drosophilidae (Diptera) from Switzerland, Italy and Greece. Mitteilungen der Schweizerishcen 

entomologischen Gesellshaft, 72:119–122.

Baptista, A. R. 1998a. “Systematics of the Family Aulacigastridae.” In Fourth International Congress of Dipterology, Abstracts, 6–13 September 1998, Oxford, UK, ed. 

J. W. Ismay.

———. 1998b. “Preliminary Cladistic Analysis of the Family Aulacigastridae sensu lato

(Diptera, Cyclorrhapha).” In 17th Meeting of the Willi Hennig Society, Abstracts, 

São Paulo, Brazil, 21–25 September 1998. São Paulo, Brazil: Universidade de São 

Paulo.

Barraclough, D. A. 1993. The Afrotropical Species of Aulacigaster Macquart (Diptera: 

Aulacigastridae: Aulacigastrinae). Annals of the Natal Museum, 34(1):31–42.

Becker, T. 1902. Die Meigen’schen Typen der sogen. Muscidae acalypterae (Muscaria 

holometopa) in Paris und Wien. Zeitschrift für systematische Hymenopterologie und 

Dipterologie 2(5):209–256, 289–349.

———. 1905. “Aulacigaster.” In Katalog der paläarktischen Dipteren, ed. T. Becker, 

M. Bezzi, K. Kertész, and P. Stein, p. 216. Hódmezövásárhely, Budapest: G. Wesselényi.

Chandler, P. J. 1987. The Families Diastatidae and Campichoetidae (Diptera, Drosophiloidea) with a Revision of Palaearctic and Nepalese Species of Diastata Meigen. 

Entomologica Scandinavica, 18(1):1–50.

Christianson, C. P., and R. E. Ryckman. 1955. First Report of Aulacigaster leucopeza

(Mg.) from Baja California, Mexico; California, New Mexico and Idaho. Bulletin of 

the Brooklyn Entomological Society, 50:17.

Cole, E. J., and F. A. Streams. 1970. Insects Emerging from Brown Slime Fluxes in Southern New England. The Canadian Entomologist, 102:321–333.

Courtney, G. W., B. J. Sinclair, and R. Meier. 2000. “Morphology and Terminology of 

Diptera Larvae.” In Contributions to a Manual of Palaearctic Diptera, ed. L. Papp 

and B. Darvas, pp. 85–161. Budapest: Science Herald. 

Cumming, J. M., B. J. Sinclair, and D. M. Wood. 1995. Homology and Phylogenetic Implications of Male Genitalia in Diptera—Eremoneura. Entomologica Scandinavica,

26(2):121–149.
128 • smithsonian contributions to zoology

Dallwitz, M. J. 1980. A General System for Coding Taxonomic 

Descriptions. Taxon, 29:41–46.

Dallwitz, M. J., T. A. Paine, and E. J. Zurcher. 1993. User’s 

Guide to the DELTA System: A General System for Processing Taxonomic Descriptions, 4th ed. http://biodiversity.uno.

edu/delta/ (accessed 20 Jul 2000).

Davis, E. J., and R. S. Zack. 1978. New Host Records and Notes 

on the Dipterous Family Aulacigastridae. The Pan-Pacific 

Entomologist, 54:129–130.

Duda, O. 1924. Beitrag zur Systematik der Drosophiliden unter 

besonderer Berücksichtigung der palärktischen u. orientalischen Arten (Dipteren). Archiv für Naturgeschichte,

90:172–234.

———. 1934. “Periseelidae, Astiidae, Aulacogastridae Curtonotidae.” In Die Fliegen der paläarktischen Region, ed. 

E. Lindner, pp. 1–5. Stuttgart: Schweizerbart.

Dufour, L. 1846. Histoire des métamorphoses de l’Aulacigaster

rufitarsis, et critiques sur ce genre de Muscides Acalyptérées (1). Annales Société de la Entomologique de France,

4:455–463.

Duxbury, K. J., and D. A. Barraclough. 1994. Rarely Encountered Diptera Families in Southern Africa: An Introductory 

Conservation Perspective. Annals of the Natal Museum,

35:25–43.

Evenhuis, N. L. 1989. “Family Aulacigastridae.” In Catalog of 

the Diptera of the Australasian and Oceanian Regions, ed. 

N. L. Evenhuis, p. 549. Honolulu and Leiden: B. P. Bishop 

Museum and E. J. Brill.

———. 1994a. Catalogue of the Fossil Flies of the World (Insecta: Diptera). Leiden: Backhuys Publishers.

———. 1994b. Catalogue of the Fossil Flies of the World (Insecta: Diptera). http://hbs.bishopmuseum.org/fossilcat (accessed 27 April 2009).

Farris, J. S. 1969. A successive approximations approach to 

character weighting. Systematic Zoology, 18:374–385.

———.1986. A Successive Approximations Approach to Character Weighting. Systematic Zoology, 26:269–276.

———. 1989. The Retention Index and the Rescaled Consistency Index. Cladistics, 5:417–419. 

Ferrar, P. 1987. A Guide to the Breeding Habits and Immature 

Stages of Diptera Cyclorrhapha. Entomonograph, 8(1):1–

478; (2):479–907.

Freidberg, A. 1994. A New Palearctic Species of Xenasteia

Hardy (Diptera: Xenasteiidae). Israel Journal of Entomology, 28:133–137.

Griffiths, G. C. D. 1972. The Phylogenetic Classification of Diptera Cyclorrhapha with Special Reference to the Structure 

of the Male Postabdomen. Series Entomologica, Vol. 8. The 

Hague: W. Junk.

Hennig, W. 1956. Neue neotropische Acalyptrata aus dem 

Deutschen Entomologischen Institut (Diptera: Acalyptrata). 

Beiträge zur Entomologie, 6:146–154.

———. Die Familien der Diptera Schizophora und ihre phylogenetischen Verwandtschaftsbeziehungen. Beiträge zur Entomologie, 8:505–688.

———. 1965. Die Acalyptraten der Baltischer Bernsteins. Stuttgarter Beiträge zur Naturkunde, 145:1–215.

———. 1969. Neue Gattungen und Arten der Acalyptratae. The 

Canadian Entomologist, 101(6):589–633.

———. 1971. Neue Untersuchungen uber die Familien der Diptera Schizophora (Diptera: Cyclorrhapha). Stuttgarter Beiträge zur Naturkunde, 226:1–76.

Hilger, S., and C. F. Kassebeer. 2000. A New Species of Aulacigaster Macquart 1835 (Diptera, Aulacigastridae) from 

Réunion. Dipteron, 3(2):167–172.

Kassebeer, C. 2001. The einheimischen Arten der Gattung Aulacigaster Macquart, 1835 (Diptera, Aulacigastridae). Dipteron, 4(1):23–32.

Macquart, J. 1835. Histoire naturelle des insectes. Diptères. Tome 

deuxi-me. Ouvrage accompagn. de planches. Paris: Roret.

Malloch, J. R., and W. L. McAtee. 1924. Flies of the Family Drosophilidae of the District of Columbia Region, with Keys to 

the Genera, and Other Notes, of Broader Application. Proceedings of the Biological Society of Washington, 37:25–42.

Mathis, W. N., and A. Freidberg. 1994. A Review of North 

American Aulacigaster Macquart (Diptera: Aulacigastridae). Proceedings of the Entomological Society of Washington, 96(4):583–598.

Mathis, W. N., and L. Papp. 1992. A New Genus of Periscelididae (Diptera) from the Neotropics. Proceedings of the Biological Society of Washington, 105(2):366–372.

McAlpine, D. K. 1978. Description and Biology of a New Genus 

of Flies Related to Anthoclusia and Representing a New 

Family (Diptera, Schizophora, Neurochaetidae). Annals of 

the Natal Museum, 23(2):273–295.

———. 1983. A New Subfamily of Aulacigastridae (Diptera: 

Schizophora), with a Discussion of Aulacigastrid Classification. Australian Journal of Zoology, 31:55–78.

McAlpine, J. F. 1981. “Morphology and Terminology—Adults.” 

In Manual of Nearctic Diptera, ed. J. F. McAlpine, B. V. 

Peterson, G. E. Shewell, H. J. Teskey, J. R. Vockeroth, and 

D. M. Wood, pp. 9–63. Ottawa: Research Branch, Agriculture Canada.

———. 1989. “Phylogeny and Classification of the Muscomorpha.” In Manual of Nearctic Diptera, ed. J. F. McAlpine, 

B. V. Peterson, G. E. Shewell, H. J. Teskey, J. R. Vockeroth, and D. M. Wood, pp. 1397–1518. Ottawa: Research 

Branch, Agriculture Canada.

Meigen, J. W. 1830. Systematische Beschreibung der bekannten 

europäischen zweiflügeligen Insekten. Volume 6. Hamm: 

Schulz-Wundermann.

———. 1838. Systematische Beschreibung der bekannten europäischen zweiflügeligen Insekten. Vol. 7, oder Supplementband. Hamm: Schulz-Wundermann.

Oldenberg, L. 1914. Beitrag zur Kenntnis der europäischen Drosophiliden (Dipt.). Archiv für Naturgeschichte, 80(2):1–42.

Pankhurst, R. J. 1991. Practical Taxonomic Computing. Cambridge: Cambridge University Press.

Papavero, N. 1967. “Family Aulacigastridae.” In A Catalogue of 

the Diptera of the Americas South of the United States, ed. 

N. Papavero, pp. 1–2. São Paulo: Departamento de Zoologia, Secretaria de Agricultura.

Papp, L. 1984. “Family Aulacigastridae.” In Catalogue of Palaearctic Diptera, ed. A. Soós and L. Papp, pp. 60–61. New 

York: Elsevier.

———. 1998a. The Palaearctic Species of Aulacigaster Macquart 

(Diptera, Aulacigastridae). Acta Zoologica Academiae Scientiarum Hungaricae, 43(3):225–234.
number 633 • 129

———. 1998b. “Family Aulacigastridae.” In Contributions to 

a Manual of Palaearctic Diptera (With Special Reference 

to Flies of Economic Importance), Higher Brachycera, ed.

L. Papp and Bela Darvas, Vol. 3, pp. 279–284. Budapest: 

Science Herald.

———. 2008. Description of the Immature Stages and the Adult 

Female of Aulacigaster africana, the First Known for the Afrotropical Aulacigastridae (Diptera: Schizophora). African 

Invertebrates, 49(2):227–232.

Robinson, I. 1953. The Postembryonic Stages in the Life Cycle 

of Aulacigaster leucopeza (Meigen) (Diptera, Cyclorrhapha: 

Aulacigasteridae). Proceedings of the Royal Entomological 

Society of London (A), 28:77–84.

Rohácˇek, J. 1998. Taxonomic Limits, Phylogeny and Higher 

Classification of Anthomyzidae (Diptera), with Special Regard to Fossil Record. European Journal of Entomology,

95:141–177.

Rondani, C. 1874. Species Italicae ordinis Dipterorum (Muscaria Rndn.) collectae et observatae. Stirps XXII Loncheinae Rndn. Bullettino della Società Entomologica Italiana,

6:243–274.

Rung, A., W. N. Mathis, and L. Papp. 2005. Curiosimusca, gen. 

nov., and Three New Species in the Family Aulacigastridae 

from the Oriental Region (Diptera: Opomyzoidea). Zootaxa, 1009:21–36.

Sabrosky, C. W. 1977. “Family Aulacigastridae.” In A Catalog 

of the Diptera of the Oriental Region. Suborder Cyclorrhapha (Excluding Division Aschiza), ed. M. D. Delfinado and 

D. E. Hardy, p. 230. Honolulu: University Press of Hawaii.

———. 1983. A Synopsis of the World Species of Desmometopa

Loew (Diptera, Milichiidae). Contributions of the American 

Entomological Institute, 19(8):1–69.

Schiner, I. R. 1863. “Die Fliegen (Diptera).” In Fauna Austriaca,

Vol. 2, pp. 81–288. Vienna: Carl Gerold’s Sohn.

———. 1864. Catalogus systematicus dipterorum Europae. Vindobonae [Vienna]: Societatis zoologico-botanicae.

Swofford, D. L. 2002. PAUP*. Phylogenetic Analysis Using Parsimony (*and Other Methods). Sinauer Associates, Sunderland, Massachusetts.

Teskey, H. J. 1976. Diptera Larvae Associated with Trees in 

North America. Memoirs of the Entomological Society of 

Canada, 100:1–53.

———. 1987. “Family Aulacigastridae.” In Manual of Nearctic 

Diptera, ed. J. F. McAlpine, pp. 891–894. Ottawa: Research 

Branch, Agriculture Canada.

Wahlberg, P. F. 1847. Tvänne nya dipter-genera af agromyzidernas familj. Öfversigt af Kongl. Vetenskaps-Akademiens Förhandlingar, 4:259–263.

Winkler, I. S., A. Rung, and S. J.Scheffer. 2010. Hennig’s Orphans Revisited: Testing Morphological Hypotheses in the 

“Opomyzoidea.” Molecular Phylogenetics and Evolution

54:746–762.

Wirth, W. W. 1965. “Aulacigastridae.” In A Catalog of the Diptera North of Mexico, ed. A. Stone, C. W. Sabrosky, W. W. 

Wirth, R. H. Foote, and J. R. Coulson, p. 823. Washington, 

D.C.: U.S. Department of Agriculture.

Yeates, D. K. 1995. Groundplans and Exemplars: Paths to the 

Tree of Life. Cladistics, 11:343–57.

Index to the Species of Aulacigaster

aenigma, 9, 21–24, 26, 119

afghanorum, 2, 104–105, 107

africana, 2, 5, 83, 84–85, 106

albifacies, 8, 64–66, 83 

angusta, 7, 48–49, 52

appendiculata, 8, 53–54

atriseta, 8, 66–68, 76

belize, 7, 68–69, 117

bella, 7, 32–33

bifasciata, 7, 66, 69–71

borbonica, 2, 83, 85–86

bromeliae, 8, 10–12, 21, 117

bromeliae group, 6, 9–10, 116, 117, 120

condylura, 9, 54–57

conspicua, 8, 71– 73

costaricana, 8, 54, 57–58

curvata, 9, 55, 58–60

ecuadoriensis, 2, 3, 9, 21, 22–24, 119

ecuadoriensis group, 8, 21, 117

erika, 7, 73–75

erwini, 7, 33–35, 36, 42

falcata, 2, 104, 105–106, 107

fastidiosa, 9, 22, 24–26, 119

femorata, 7, 32, 34, 35–36, 42

femorata group, 7, 31–32, 119

formosa, 7, 37–38, 41

freidbergi, 84, 86–88

gaimarii, 9, 53, 55, 60–61

grimaldii, 7, 48, 49–52 

grimaldii group, 7, 48, 117

guyana, 7, 36, 38–39
132 • smithsonian contributions to zoology

india, 112–113, 117

irwini, 7, 38, 39–41, 48

korneyevi, 8, 10, 12–14, 21 

L1, undescribed species, 6, 100, 

101–103, 117

L2, undescribed species, 6, 100, 103–104

L3, undescribed species, 112, 114–115, 

117

leucopeza (Meigen), 2, 5, 6, 90, 104, 

106–109 

leucopeza group, 6, 83, 100, 104, 112, 

116, 117

lobata, 7, 34, 35, 36, 41–42,

lopesi, 8, 14–17, 17, 19

malawana, 84, 88–89

mcalpinei, 2, 90–92 

melanoleuca, 2, 9, 24, 27–28, 119

minuta, 2, 8, 53,61–63

minuta group, 8, 52–53, 117, 119, 120

neoleucopeza, 2, 90, 92–98, 104, 107, 

109, 111, 112

nigritarsa, 8, 63–64 

nigritibia, 7, 43–44

pappi, 2, 104, 107, 109–112

perata, 2, 83, 89–90

peruana, 7, 44–46

plesiomorphica, 2, 8, 67, 75–76, 78, 81

plesiomorphica group, 7, 64, 117

proxima, 8, 67, 76–78

rufifemur, 8, 76, 78–82

sabroskyi, 2, 90, 98–100

serrana, 8, 15, 17–18

spangleri, 100, 101, 117

srilanka, 112, 113–114, 117

stenoptera, 9, 21, 28–31

tibanae, 8, 15, 17, 18–19

trifasciata, 7,66, 82–83

unifasciata, 7, 38, 41, 46–48

vespertina, 8, 12, 13, 14, 19–21
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