Citation
An illustrated key for the genera of Ceratopogonidae (Diptera) of the world

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Title:
An illustrated key for the genera of Ceratopogonidae (Diptera) of the world
Creator:
Ratanaworabhan, Niphan Chanthawanich, 1939-
Publication Date:
Language:
English
Physical Description:
262 leaves : ill. ; 28 cm.

Subjects

Subjects / Keywords:
Abdomen ( jstor )
Antennas ( jstor )
Claws ( jstor )
Female animals ( jstor )
Hair ( jstor )
Legs ( jstor )
Male genitalia ( jstor )
Mandible ( jstor )
Palps ( jstor )
Species ( jstor )
Ceratopogonidae ( lcsh )
Diptera ( lcsh )
Dissertations, Academic -- Entomology and Nematology -- UF
Entomology and Nematology thesis Ph. D
Genre:
bibliography ( marcgt )
non-fiction ( marcgt )

Notes

Thesis:
Thesis (Ph. D.)--University of Florida, 1969.
Bibliography:
Includes bibliographical references (leaves 246-260).
General Note:
Typesript.
General Note:
Vita.
Statement of Responsibility:
by Niphan Chanthawanich Ratanaworabhan.

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University of Florida
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University of Florida
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AN ILLUSTRATED KEY FOR THE GENERA OF

CERATOPOGONIDAE (DIPTERA) OF THE WORLD














By

NIPHAN CHANTHAWANICH RATANAWORABHAN














A DISSERTATION PRESENTED TO THE GRADUATE COUNCIL OF
THE UNIVERSITY OF FLORIDA
IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE
DEGREE OF DOCTOR OF PHILOSOPHY












UNIVERSITY OF FLORIDA 1969















ACKNOWLEDGMENTS


The author expresses her deep gratitude to Dr. F. S. Hlanton,

chairman of her supervisory committee, for his help and encouragement in all phases of her graduate study.

She wishes to express her most sincere appreciation to Dr. W. W. Wirth for all assistance, criticism and for providing the available specimens from the U. S. National Museum for this taxonomic study.

For their criticism of the manuscript, appreciation is also

extended to committee members: Dr. M. J. Westfall, Dr. L. C. Kuitert and Dr. D. H. Habeck.

The author also expresses her thanks to the U. S. Arqr (project number D A-49-193-MD-2177) for sponsoring this program and finally, to her husband for his help, encouragement, cooperation and understanding during her graduate study.






ii
















TABLE OF CONTENTS

Page
ACIKNLEDGM NTS LIST OF FIGURES . . . . . . . . . . * * vii INTRODUCTION .......... * **** 1 IEY FOR TIH GEMERA OF CERATOPOGNIDAE . . . . . . . 4 DESCRIPTIONS ..... . . . . * 14

Genus Leptoconoos Skuse . ... . . . . . . 14

Key for the Subgenera of Leptoconops . . . . 19

Genus Atrichoporon Kieffer . . . . . . . . . . 20

Key for the Subgonera of Atrichopogon . . . 25

Genus Forcinomia laigen . . . . . . . . . . 26

Key for the Subgenera of Forciomyia . . . . . 37

Genus Dasyhelea Kieffer . . . . . . . . 40

Key for the Subgenera of Dasyhelea . . . . . 45

Genus Paradasyhelea Macfie . . . . . . . . . 46 Genus Culicoides Latreille . . . . . . . . . . 47

Key for the Subgenera of Culicoides . . . . . 58

Genus Austroconops Lirth and Lee . . . . . . . . 61 Genus Ca-otopterohela Wirth and Hubert . . . . . 62 Genus Ceratopogon Ieigen . . .. . * 63

Key for the Sib-onora of Ceratopogon . . . . . 67

Genus Alluaudorzna Kieffer ....... . . 68 iii










Genus Stilobezzia Kieffer . . .... . . 70

Key for the Subgenera of Stilobezzia . . . 75 Genus Echinohelea Macfie .. . . . 76 Genus Monohelea Kieffer . . . . . . . 77

Key for the Subgenera of Monohelea . . . . 80 Genus Scrro21va NMigon . . . a . . 81 Genus hcrurohelea I. & . ... .. . . 83 Genus Parabezzia I4aJloch . . . . . . . . 84 Genus Clinohela Xeiffer . . . . . . . . 85 Genus Tetrabezzia Kieffer . . . . . . . 87 Genus Heteron iia Say . . . . . . . . . 88 Genus PelucClMdowa Macfie . . . . . . . . 90 Genus Teurohelea Kieffer . . . . . . . . . 91 Genus Johannsenoz1ia Malloch .. . . 93 Genus Xenohelea Kieffer . . . . . . . . . 95 Genus Jenkinshelea Macfie . . . . . 97 Genus Probezzia Kieffer ... . . 98 Genus Calnttoonoon Kieffer . . . . . . . . 100 Genus Spharomias Curtis . . . . . . . . 102

Xey for the Subgenera of Sphaero'ias ... 105 Genus AMllochohelea Wirth . . . . . . . 106 Genus Nilobezzia Kieffer . . . . . . . . 107

Key for the Subsenera of Nilobezzia . . . 110 Genus Stenoxenus Coquillett . . . . . . . 111 Genus Paryphoconus Enderlein . . . . . . . 112 iv









Genus Pachyhelea irth . . . . . . . . . . 113 Genus Palpon~ria Migen . . . . . . . . . . 115

Key for the Subgenera of Palor-ria . . . . 118

Genus Bezzia Keiffer . ......... 119

Key for the Subgenera of Bezzia . . . . 122 9 Genus Phaenobezzia I aeselbarth .. .. . . 123

Genus A . . . . . . . . . . . . 124

Genus B . . . . . . . . . . . .. . 125

Genus C ... . . . . . . . . . . . 126

Genus D ........... 126

Genus E . . . . . . . . . . . . . . 127

Genus F . . . . . . . . . . . . . . 128

Genus G . . . .. . . . . . 128 GenusH . . . . . . . . . . . . . 129

Genus J .. . . . . . . . . . . 129

Genus . . . . . . . . . . . . . 130

Genus L . . . . . . . . . . . . . 130

Genus M . . . . . . . . .. 1 � � � 31 Genus N * * * ** 132 Genus 0 * * * * � � 132 Genus P * * . . . . . . . . . 132 Genus ..... 133 Genus R .. . ....... , 134

Genus S . . * * * * * * * * * * 134 Genus T ..0*. .**. . . . 135





v









Gemnus Dolichoheloa Edvards . . . . . * 135 Genus Acanthohelea 10.effer . . . . . 136 Genus Luci~ -ia de Xeillon . . . . . . . * 136 Genus Ceratobezzia Kieffer * * * * � � 136 ALPHABETICAL LIST OF GIA AND SUBGMZRA . . . . . . 137 LIT~rATURE CITED * * * * * ... ... .. 242 BIOGRAPHICAL S~TTC . . . . .... 261

































vi













LIST OF FIGURES

Fieure

1. Culicoides furens (Pocy) . . . . . . . . . . . . . . 140

2. Leptoconops (Leptoconops) stygius Skuse . . . . . . 141 3. Leptoconops (Holoconops) kerteszi K. . . . . . . . . 142 4. Lentoconops (Styloconops) albiventris (Meijere). . . 143 5. Leptocono-ps (Styloconops) spinosifrons carter. . . . 144 6. Atrichopogon (Atrichonpooon) levis (Coq.) . . . . . . 145 7. Atrichooogon (Psilokemnia) arcticus (Coq.). . . . ... 146 8. Atrichopogon (Meloehelea) meloesumans K. . . . . . . 147 9. Forcipor~yia (Forcipoyia) biounctata (L.). . . . . . 148 10. Forcipomyia (Proforcipoyia) Firthi Saunders . . . . 149 11. Forciponyia (Warniea) aeria Saunders . . . . . . . . 150 12. Forciporimia (Warmkrea) tuberculata Saunders ..... 151 13. ForcipoZria (Caloforciponyia) glauca Edwards . . . . 152 14. Forcipotria (Metaforcipomyia) pluvialis Maioch .. 153 15. Forcipoia (Thyridomyia) pal'stris Saunders . . . . 154 16. Forcinonria (Synthyridomia) colemani Wirth . . . . 155 17. Forciporyia (Trichohelea) fijiensis (Nacfie) . . . . 156 18. Forcipompyia (Lasiohelea) fairfaxensis Wirth . . . . 157 19. Forcipoqnia (Neoforciporyia) eques (Joh.) . . . . . 158 20. Forcipon~ia (Dacnoforciporria) anabaenae Chan & Saunders159 21. Forcipomyia (Phasmidohelea) fuliginosa (Mg.) . . . . 160 vii










22. Forcipomia (Phasmidohelea) sp. . . . . . . . . . 161

23. Forcipomyqia (Lepidohelea) annulatipes Macfie . . . . 162 24. * Forcipomyia (Pterobosca) fusicornis (Coq.) . . . . . 163 25. Dasyhelea (Dasyhelea) grisea (Coq.) ... . . . . . . . 164

26. Dasyhelea (Pseudoculicoides) mutabilis (Coq.) . . . . 165 27. Dasyhelea (Prokempia) cincta (Coq.) . . . . . . . . . 166

28. Paradasyhelea minuta Wirth & Lee . . . . . . . . 167 29. Culicoides (Culicoides) yukonensis Hoffman . . . . . 168 30. Culicoides (Trithecoides) flaviscutatus W. & H. . . . 169 31. Culicoides (Haemophoructus) gemellus Macfie . . . . . 170 32. Culicoides (Meijerehelea) guttifer (Mijire). . . . . 171 33. Culicoides (Hoffmania) insignis Lutz . . . . . . .. 172 34. Culicoides (Avaritia) obsoletus (Mg.). . . . . . . 173 35- Culicoides (Oecacta) furens (Poey) . . . . . . . . . 174 36. Culicoides (Drymodesmyia) copiosus R. & H. . . . . . 175 37. Culicoides (Diphaeoiria) baueri Hoffman . . . . ... 176 38. Culicoides (Anilonia) covagarciai Ortiz . . . . . . 177 39. Culicoides (Beltrannyia) crepuscularis Malloch. . .. . 178 40. Culicoides (Glaphiromyia) scopus R. & H. . . . . . . 179 41. Culicoides (Haematonyidium) paraensis (Goeldi). . . . 180 42. Culicoides (Macfiella) phlebotomus (Will.) . . . . . 181 43. Culicoides (Mataemyia) mojingaensis W. & B. . . . . . 182 44. Culicoides (Monoculicoides) variipennis (Coq.). . . . 183 45. Culicoides (Selfia) hieroglyphicus Malloch. . . . . . 184 46. Austroconops mmillani Wirth & Lee.. ....... 185 iii









47. Cartopterohelea hoogstraali W. & H. . . . . . . ... 186

48. Ceratogoon (Ceratoooon) culicoiclithora x iofnan. . 187 49. Ceratopogon (Isolea) sti is Coq . . . . . . . 188 50. Alluaudomya bella (Coq.) .... . ........ 189

51. Alluudomyla arva irth . . . . .. . . . .... 190

52. Stilobezzia (Stilobezzia) festiva K ......... 191 53. Stilobezzia (T2h1raiohelea) eleantula (Joh.) . . . . 192 54. Stilobezzia (N ostilobezzia) lutea ( och). . . . 193 55. Echrinohelea lanci irth . . . .. ... . . . 194

56. Monohelea (,onohelea) hieroglyphica K . . . . . . . 195 57. Monohelea (Schizohelea) leucoza (g.). . . . . . . 196 58. Serro ia femorata (Lg.) . . . . . . . . . . 197

59. Mcrurohelea setosa irth . . . . . . . . .... . 98
60. Parabezzia lata l. . . . . . .. . . . . 199

61. Clinohelca bLraclata (Loew) . . . . . . a . . . . . 200

62. Tetrabezzia pictiTennis (K.) . . . . . . . . . . . . 201

63. Hotoroyia fasciata Say . . . . . . ....... 202

64. Pellucidonia sp. . . .. . . ... . . . 203
65. reurohelea nigra irth . . . . . .... . . ... 204

66. Johannenorria arentata (Loew). . . . . . . . . . . 205

67. Xnohelca tonnoiri Lee . ........ . . . . . 206

68. Jonkinshelea rmnEnnis (Joh.). . . . . . . . . . . 207

69. Probezzia pallida lloch. . . . . . . . . . . . .. 20

70. agyphtcroon Sibbosus (Woied.). ....... ... . . 209
71. Sphaeromias (Shacromias) lonrennis (Loew) . . . . 210 ix











72. Sphaerorrias (Honohelea) barkudensis Edwards. . . . . 211 73. a2llochohelea albibasis (Mall.) . . . . . . . . � 212 74. Nilobezzia schwarzli (Co.) ............ 213

75. Stenoxe ns ins niervis cfie. . . . . . . . . . . 214

76. Paryphoconus anustinls Enderlein . . . . . . . 215 77. Pachyhelea nach ymera (ll.) . . . . . . . . . . 216

78. PaTora plebeia (LoeW) . ..... ........ 217

79. Bezzia (Bezzia) setulosa (Loew). . . . .. . . . . . 218

80. Bozzia (Pseudobezzia) flavitarsis (ll.) . . . . . 219 81. Phaenobezzia pistiae (I. & M.) . . . . . . . . . . 220

82. Genus A . . . . . . . . . . . . . . . . * . . 221

83. Genus . . . . . . . . . . . . . * * * 222

84. Genus C . . . . . . . . . . . . . . . . . . 223

85. Genus D . . . . . . . . . . . . . . .. . . . . . 224

86. Genus D sp. 2 . . . . . . . . . . . . . . . . . . 225

87. Genus E . . . . . . . . . . . . . . . . . 226

88. Genus F sp. 1 . . . . . . . . . . . . . . * . * . 227

89. Genus F-sp. 2 . . . . . . . . . . . . . . . . . . . 228

90. Genus G . . . . . . . . . . . . . . . . . . * .. 229

91. Genus H . . . . . . . . . . . . . . . . . . . . 230

92. Genus J . . . . . . . . . . . . . . . . . . . . . . 231

93. Genus K . . . . . . . . . . . . . . . . . . . 232

94. Genus L . . . . . . . . . . . . . . . . . . . . . . 233

95. Genus . . . . . . . . . . . . . . . . . . . . . 234

96. Genus N .. . . . . . . . . . . . . . . . . . 235











97. Genus 0. . . ............. . . . . .. 236

98. Genus P. . . ..... ....... . .... 237

99. Genus Q . ....... .... . . .. ... .. 238

100. Genus. * . . . ......... ..... .. 239

101. Genus S ............ . .... ..... .. 240

102. Genus T .......... . .. ... .. . . . . 241

















Ceratopogonidae is a large family of minute flies, which was brought to the attention of taxonomists in 1852 by Winnertz in his monograph of the European species of the genus Ceratopogon, The contributions following Winnertz were those of Coquillett who published on American species from 1900 to 1905. He included all the species in Ceratopogon. Kieffer (1899-192k) described several species and erected many genera. Ceratopogonidae was recognized as a subfamily of Chironomidae until Malloch (1917) finally elevated them as' belonging to a distinct family. Later EdwardS(1926) published the diagnostic characters of many genera which are now being used as the basis for modern classification.

Since 1925 Macfie has been monographing groups on a worldwide basis.
The early publications on the genera of American ceratopogonids were published by Malloch (1915), Thomsoen (1935, 1937), Hoffran (1924-1939), Root and Hoffman (1937) and Johannsen (1943). Recent ones are those of Wirth (1952, 1962a) and Wirth and Stone (1956).

The important papers on European Ceratopogonidae are those of Kieffer (1925), Edwards(1926), Goetghebuer (1920), Lindner (1934) and Clastrier (1961- ).

Elsewhere, Lutz (1912-1914) and Lane (1944-1956) worked on the Neotropical genera, Carter, Ingram and Macfie (1920-1922),
1




2


on the West African, de Meillon (1931- ) on the South African, Tokunaga (1939- ) on the Japanese and the Micronesian (Tokunaga and Eurachi, 1959), and Lee (1948- ) and Skuse (1889) on the Australasian species.

A key to the genera of Ceratopogonidae was published in 1926 by Kieffer who separated the famTily into 59 genera. Other keys have been made by many authors who mainly limited their keys to the genera and the species in a certain region. Edwards (1926) published the diagnostic characters of the genera occuring in Great Britain. Johannsen (1932) constructed a key based on the collection from ,alayan Subregion with the additional genera which were recorded from the entire Oriental region. Goetghebuer (1933) published a key for the genera of the Palearctic region.

iMacfie's key (1940a) was considered to be the complete key to the known genera of that time. It was a concise key which contained 63 genera but he failed to give the generic definitions, which made the key ineffective to taxonomists. The best key to North and South American genera, Johannsen (1943), included the illustrations of rany important generic characters. Lee (1948) published a key to the genera known to occur in the Australasian region. He divided the family into 10 groups, using a cfie's groupings (1940a). He also presented a comparison of closely related genera. A key to some

American genera was published in 1952 by W1-irth, based on his studies of California ceratopogonids. A key to the North American genera wa s also presented by WKirth and Stone (1956). Recent keys to the subfamilies, genera and subgenera of the Ceratopogonidae of the %'orld by Wirth (1959c, unpublished) and the genera of the tribe Sphaeromiinae





j






3


(1962) are probably the most comprehensive ones. The latter provided some illustrations of the important characters and generic diagnoses were added.

There has been a great deal of discrepancy among the current

keys because of the confusion which is almost always present with the key and the generic diagnosis. Many previously established genera were synornymized or sometimes reduced to subgeneric rank. It is very difficult to use available keys with this added confusion. Therefore a complete and comprehensive key is needed to facilitate the classification of ceratopogonids. An attempt is made in these studies to provide the diagnostic characters of each genus with detailed illustrations of its representative species. Whenever possible the type species was selected for study. Keysto the subgenera are also presented and the important references are provided.

The following genera: Dolichohelea, Acanthohelea, Luciamia and Ceratobezzia are not included in the key for the genera of Ceratopogonidae. The specimens of these genera are not represented in the collection. The information obtained is only that published by the authors and the fragmentary information was summarized in this report.

The information on the immature stages, biology and habitats of any genus is included whenever it is available.

The specimens for this study were obtained from the U. S. National Museum.















KEY FOR THE GENERA OF CERATOPOGONIAE


1. Female antenna 13-14-segmented........ . . . . . 2

Female antenna 15-segmented . . . . . . . . . . . . . . . . . . 3


2. The r-m crossvein absent . . . . . . . . . Leptoconops (Fig. 2)

The r-m crossvein present. . . . . . . . . . . . Genus P (Fig. 98)


3. Female empodium large . . . . . . . . . . . . . . . . . . . 4

Female empodium small or vestigial . . . . . . . . . . . . . . . 5


4. Radial cells not well developed, first small or obliterated,

second small, wing with abundant flattened macrotrichia; body densely covered with hairs; male genitalia: ninth

tergum short and tapered, setose apicolateral process

present . . . . . . . . . . . . . . . Forcipomyia (Fig. 9)

Radial cells well developed, second one large, wing with

moderate stiff macrotrichia; body hairs sparse; male

genitalia: ninth tergum rounded, apicolateral process

absent . . . . . . . . . . . . . . . . .Atrichopoon (Fig. 6)


5. Antennal segments sculptured . . . . . . . . . . Dasyhelea (Fig. 25)

Antennal segments not sculptured . .. . ..00 .. .. 6 6. Radial cells both absent . . . . . . . . . . . . . . . . . . 7

Radial cells present, at least one. .............. 10


4






5


7. Palpi five-segmented . . . . . . . ... . . Genus M (Fig. 95)

Palpi two-four-segmented . . . . . . . . . . . . . . * 8


8. Palpi four-segmented; wing not broad; male antenna

14-segmented . . . . . . . . . . . Paradasyhelea (Fig. 28)

Palpi two-segmented; wing broad; male antenna

eight-segmented . . . . . . . .. . . . . . . . . * 9


9. Posterior branch of media narrowly interrupted at base,

cell M4 large, costa extended about half of wing

length . . . . . . . . . . . . . . . * . . . Genus Q (Fig. 99)

Posterior branch of media broadly interrupted,

cell X small, costa short extended about onethird of wing length .... . . . .. . Genus S (Fig. 101)


10. Media petiolate- , usually forking behind r-m crossvein . . . .11

Media sessile, forking at or before r-m crossvein . . . . . . 24 11. Claws small, equal and simple in both sexes; one or two

radial cells almost equal in size . . . . . . . . . . . 12

Claws of female usually large; one or two radial cells,

second one always much longer than the first . . . . . . . 14 12. Palpi five-segmented . . . . . . . . . . Culicoides (Fig. 29)

Palpi three-four-segmented . .... . . . .. . . . . 13


13. Palpi three-segmented; one radial cell, lower branch of

media absent; one spermatheca . . . . . . Genus N (Fig. 96)

Palpi four-segmented; two medial cells, long and broad,

lower branch of media present; two spermathecae

. . . . . . . . . . . . . . . . . . . Austroconos (Fig. 46)






6


14. Second radial cell not or little longer than the first,

one or both cells may be obliterated (in Alluaudoyria

first one usually obliterated) costa about half of

wing length . . . . . . . . . . . . . . . . .. . . . . . 15

Second radial cell much longer than the first, costa

extended to wing tip or two-thirds of wing length . . . . 17 15. Wing with one radial cell, usually with 1-20 small

dark spots and sometimes with grayish streaks along

the wing veins; two claws on each leg.Alluaudomyia (Fig. 50,51)

Wing usually with two radial cells, usually without

dark spots; one or two claws on each leg . . . . . . . . . . 16 16. All claws single, stout and slightly curved; palpi twosegmented; wing veins with only cubital stem and IA

present, Cui indistinct, microtrichia spinelike

. . . . . . . . . . . . . . . . . Camototerohelea (Fig. 47)

Two claws on each leg, equal or subequal; palpi fivesegmented; wing with lower branch of media often absent

or interrupted, without microtrichia . . Ceratopogon (Fig. 48)


17. Costa extending beyond vein R5, one or two radial

cells; claws usually equal .. . . . . . . . . . . . . . .18

Costa not extending beyond vein RS, two radial cells

present, claws equal or unequal . . . . . . .. . . 20 18. Two radial cells, first one small; palpi threesegmented . . . . . . . . . . . . . . . . . Genus 0 (Fig. 97)

One radial cell, palpi four-segmented . . . . . . . . .... 19






7


19. First tarsomere of mid leg much longer than that of the

other legs and bearing spines; Coxae of male with

few hairs; male genitalia: aedeagus arched, paramere

absent ...... . . . . . . ...... Genus R (Fig. 100)

First tarsomere of mid leg simple; coxae of male densely

covered with bristles; male genitalia: aedeagus large, paramere small and fused to form a triangular sclerite

behind aedeagus . . . . . . . . . .. Parabezzia (Fig. 60)


20. Hind femur simple, armed or unarmed, tibia not curved .... 21

Hind femur swollen in both sexes, armed ventrally with

numerous spines, tibia curved basally . . Serroryia (Fig. 58)


21. Female abdomen simple . . . . . . . . . . . . . . . . . . . 22

Female abdomen highly modified with tenth segment curved

forward and reaching the ventral side of anterior

segments . . ... ...... . . Macrurohelea (Fig. 59)


22. Male antenna plumose; femora and tibiae unarmed; last

antennal segment simple . . . . . . . . . . . . . . . . . . 23

Male antenna without plumes; almost all femora and

tibia armed with long spines; last antennal segment

usually shorter than the prece ding four segments

. . . . . . . . . . . . . . . . . . . . Echinohelea (Fig. 55)

23. Female claws all unequal; first tarsomere of mid leg

bearing subbasal spur; male genitalia: ninth tergite

rounded, aedeagus reduced to a pair of oblique lateral

sclerites, parameres with modified, well developed

submedian process . . . . . . . . . Stilobezzia (Fig. 52)






8


Female anterior four claws equal, hind claw with one long

and with or without another short one; male genitalia:

bilobed or truncate ninth tergum, aedeagus usually with

a pair of lateral sclerites and additional dorsomedian lobe,

parameres fused or paired . . . . . . Monohelea (Fig. 56)


24. Female fifth tarsomere with numerous ventral blunt

spines or batonnets . . . . . ... . . . . . . . . . 25

Female fifth tarsomere with or without sharp batonnets . . . 37 25. Female claws highly unequal; wing with two radial cells . . . 26

All female claws equal with internal or external basal

tooth; one or two radial cells . . . . . . . . . . . . 27


26. All female claws unequal; batonnets appear on basal half

of fifth tarsomere . . . . . . . . . ... Xenohelea (Fig. 67)

Claws highly unequal only on four posterior legs, each

with external basal tooth; fifth tarsomere with numerous

batonnets along the segment . . . . Johannsenonyia (Fig. 66)


27. Female abdomen with ventral sclerotized plate on

seventh segment . . . . . . . . . . . . . . . . .. . . . . 28

Female abdomen without sclerotized plate on seventh segment . . 29


28. Wing with two radial cells; anterior part of thorax

cone-shaped . . . . . . .... . . Genus H (Fig. 91)

Wing with one radial cell; anterior part of thorax

rounded ............... Genus K (Fig. 93)






9


29. Hind tarsus about three times longer than mid

tarsus . . . . . . . . . . . . . . Captooon (Fig. 70)

Hind tarsus simple . . . . . . . . . . . . . . . . . . . . . 30


30. Wing of female with usually broad anal lobe.Jenkinshelea (Fig. 68)

Wing with anal lobe narrow and simple . . . . . . . . . . . . 31


31. Female claws each with internal basal tooth (Genus J:

claws with internal tooth in one and external tooth

in the other claw); ventral hair tufts absent . . . . . . 32

Female claws each with external basal tooth; abdomen

with ventral hair tufts . . . . . . . . .. . . . . . . 34


32. One spermatheca; anterior tubercle -ell developed

* . . . . . . . . . . . . . . . . . . . . . Genus A (Fig. 82)

Two spermathecae; anterior tubercle poorly developed

or absent . . . . . . * * * * * * . . . . . . . . . . . .* 33


33. One radial cell; claws each with internal tooth in one and

external tooth in the other; leg unarmed . . Genus J (Fig. 92)

Two radial cells; claws each with internal basal tooth;

legs with numerous spines on ventral side of femora and

bristles on dorsal side of tibiae . .. . Sphaeromias (Fig. 71)


34. Legs armed with ventral spines on femora and dorsal spines

on tibiae; male genitalia: highly modified, with basistyle

poorly developed and fused to form a cone-shaped process

. . . . . . . .... . . . . . . . . . . Nilobezzia (Fig. 74)

Leg with femora and tibiae unarmed . . . . .... . . .35






10


35. Wing with two radial cells . . . . . . . 4allochohelea (Fig. 73)

Wing with one radial cell .. ... . . . . . . *..* 36


36. Costa very long reaching the wing tip; fourth tarsomere

bilobed; spermatheca small; mandible without serrate

margin; male genitalia: parameres fused mesally, tip

slender and separated . . . . . . . . . . Probezzia (Fig. 69)

Costa short, not reaching the wing tip; fourth tarsomere

oval; spermathecae large; mandible with serrate margin;

male genitalia: parameres separated then fused again

at the tip . . ... ....... . Genus T (Fig. 102)


37. Fifth tarsomere at least on one leg with one to five

ventral sharp spines, or abdomen cone-shaped . . . . . . . 38

Fifth tarsomere without ventral sharp spines . . . . . . . . 44 38. Abdomen cone-shaped . . . . .. . . . . . . 39

Abdomen simple, not cone-shaped . . . . . . .. . . . . . . . 40


39. Fifth tarsomere with about three ventral spines on distal

half; gland rods on seventh abdominal segment only

S. . . . . . . . . . . . . . . . . . . . Pachyhelea (Fig. 77)

Fifth tarsomere without ventral sharp spines; gland rods

present on segments five, six, and seven .Genus D (Fig. 85,86) 40. Female abdomen with gland rods . . . . . . . . . . . .. . 41

Female abdomen without gland rods . . . . . .. . . . . ... 43





11


41. Femora armed with ventral spines; first hind tarsomere

longest . . . . . . . . . . . . . . . . . . Genus B (Fig. 83)

Femora unarmed; first hind tarsomere simple . . . . . . . . . 42


42. One radial cell; tarsal segments two to five rather short;

all femora rather swollen . . . . . . . . . Genus L (Fig. 94)

Two radial cells; tarsal segments two to five simple;

four anterior femora slender, hind femora enlarged

S. . . . . . . . . . . . . . . . . . . . . Genus E (Fig. 87)


43. One spermatheca, large, pear-shaped; claws small; hind

femora slender; fifth tarsomere stout with one to two

sharp spines; two radial cells; mandible with five

large teeth . . . . .. . . . . . . . . Genus G (Fig. 90)

Two spermathecae; claws large especially of hind legs,

each with inner basal tooth; hind femora enlarged and

curved; fifth tarsomere slender, sometimes with spines;

mandible with about 11 teeth . . . . . . Genus F (Fig. 88,89) 44. Hind tarsus of female about three or more times longer

than midtarsus; claws highly unequal, single long hind

claws with barb . . . . . . . . . . . . . . . . . . . 45

Hind tarsus of female simple; all claws equal, if unequal

without any single claws . . . . . . . . . . . . . . . . . 47


45. Femora of fore legs greatly swollen and armed ventrally with

spines; fore tibiae arched along femora . Heteronwia (Fig. 63)

Femora of fore legs simple; fore tibiae not arched . . . . .46









46. Fourth tarsomeres with bifid spinose lobes on fore and

mid legs but long and cylindrical on hind legs; fore

femora armed ventrally; abdomen petiolate; wing fasciate;

fifth tarsomere of fore leg simple . . . Tetrabezzia (Fig. 62)

Fourth tarsomeres cordiform to transverse on fore and

mid legs but elongated on hind legs; fore femora

unarmed; abdomen slightly curved downward; wing not

fasciate; fifth tarsomere of fore leg swollen

. * *... .. . . . .* . .... . .Pellucido=ia (Fig. 64)


47. Fourth tarsomeres of at least four posterior legs

strongly bilobed and each lobe terminates in a

stout spine . . . . ... . . . . . . . Clinohelea (Fig. 61)

Fourth tarsomeres cordate or cylindrical . . . . . . . . . . 48 48. Antennal segments with stiff hairs; radial cell narrow . . . 49

Antennal segments without stiff hairs; radial cell simple .. 50


49. Vein M2 elbowed in female; female antennal segments three

to ten with stiff hairs; palpi four-segmented; fourth

tarsomere spoon-shaped to subcylindrical; last three

tarsomeres simple; male genitalia: parameres fused with

broad rounded tip; two spermathecae . . . Stencxenus (Fig. 75)

Vein M2 not elbowed; female antennal segments 11-15 with

stiff hairs; palpi five-segmented; fourth tarsomere cordate

to subcylindrical; last three tarsomeres relatively short;

male genitalia: parameres separated; one spermatheca

. . . . . . . . . . . Paryphoconus (Fig. 76)






13


50. Costa extended beyond R5; two radial cells; femora
unarmed . . . . . . . . . . . . . . . Neurohelea (Fig. 65)

Costa not extended beyond R5; one or two radial cells;

femora usually armed, at least on one leg . . . . . . .51


51. One radial cell . . . . . . . . . . . . . . .... . . . . 52
Two radial cell. . . . . . . . . . . . . . . . . . . . . . 53


52. Male genitalia: basistyle and dististyle reduced but
distinctly separated . . . . . . . . .Phaenobezzia (Fig. 81)

Male genitalia: basistyle and dististyle well developed

. . . . . . . . . . . . . . . . . . . . . . Bezzia (Fig. 79)


53. Hind claws longer than the claws on the first four legs;

fourth tarsomere oval to enlongated; gland rods

absent . . . . . . . . . . . . . . . . . . . Genus C (Fig. 84)

All laws equal; fourth tarsomere bilobed or cordiform;

gland rods always present . . . . . . Palpoia (Fig. 78)














DESCRIPTIONS


Genus Leptoconops Skuse

Leptoconops Skuse, 1890, Proc. Linn. Soc. N. S. Wales, 4:288;

Johannsen, 1905, Bull. N. Y. State ms., 86:94; Kieffer, 1906,

Gen. Insectorum, fasc. 42:48; 1908, Ann. rus. Nat. Hung., 6:577; 1921, Arch. Inst. Pasteur Afr. Nord, 1:107; Carter, 1921, Bull.

Ent. Res., 12:1; Johannsen, 1943, Ann. Ent. Soc. Amer., 36:777;

Lee, 1948, Proc. Linn. Soc., N. W. W. 72:332-8; Wirth, 1952, Univ.

Calif. Publs. Ent., 9:109; Smee, 1966, Aust. J. Zool., 14:993; Chanthawanich and Delfinado, 1967, J. Med. Ent., 4:294 (Typespecies: Leptoconops stygius Skuse, Mon).

Tersesthes Townsend, 1893, Psyche, 6:370 (Type-species: Tersesthes

torrens Townsend, mon.).

Centrotypus Grassi, 1901, Studi di Zool. sulla Mal. Roana, p. 118

(nomen nudum).

Mycterotyus Noe, 1905, Rend. atti Accad. Lincei, 14:114 (Type-species:

yvcterotypus bezzii Noe, desig. Carter, 1921).

Iycteromyia Lutz, 1912 (not Philippi, 1865), Mem. Inst. Oswaldo Cruz,

4:24 (lapsus for yecterotypus Noe).

Holoconops Kieffer, 1918, Ann. Nbs. Nat. Hung., 16:135 (Type-species:

Leptoconops kerteszi Kieffer, orig. desig.).

14






15


Schizoconops Kieffer, 1918, Ann. Mhs. Nat. Hung., 16:135 (Type-species:

Schizoconops indicus Kieffer, orig. desig.).

Styloconos Kieffer, 1921, Arch. Inst. Pasteur Afr. Nord, 1:107

(Type-species: Leptoconops albiventris de Meijere, orig. desig.). Protersesthes Kieffer, 1921, Arch. Inst. Pasteur Afr. Nord, 1:107

(Type-species: Tersesthes brasiliensis Lutz, orig. desig.). Nicroconops Kieffer, 1921, Arch. Inst. Pasteur Afr. Nord, 1:108

(Type species: Microconops vexans Kieffer, orig. desig.). Acanthoconops Carter, 1921, Bull. Ent. Res., 12:24 (Type-species:

Acanthoconops spinosifrons Carter, orig. desig.).


DIAGNOSIS. Eyes bare, widely separated in both sexes; vertex with bristles; frons and clypeus bare or hairy. Mandible about 16 teeth. Palpi "four-segmented"; distal two segments fused; male palpi longer than in female. Female antennae 13- or 14-segmented, the flagella segments subspherical or oval, each with few hairs and transparent sensory spines, the terminal segment elongate. Male antennae 15segmented, plumose. Thorax arched anteriorly; humeral pits present. Wings usually milky white; R separate at base, fused with costa about mid wing forming a stigma; crossvein absent; intercalary vein above vein M, conspicuous. Legs slender, hind legs longest; femora unarmed, tibiae each with stout apical spur; hind tibial comb five to seven; basitarsus with prominent ventral spines. Tarsal claws with small bristle or toothed; empodium small and branched. Abdomen with ninth segment bearing long or short lamellae or cerci, two well developed spermathecae. Male genitalia conspicuous; basistyles









with subbasal ventromesal lobe; dististyles short, small and tapering; aedeagus simple; parameres usually stout, heavily sclerotized.

Approximately 62 species have been described. Distribition: World-wide.

Keys to the larval and pupal stages of North American species are presented by Wirth and Stone (1956) and Thomsen (1937). The immature stages of the Bodega black gnat were described in detail by Smith and Lowe (1948).

EGG. The egg is unsculptured, usually banana-shaped, blunt at one end and narrower at the other, slightly curved and circular in cross section. LARVA. The head is partly sclerotized, inside the head and first thoracic region is a system of heavily chitinized rods connected with the mandibular processes. Labrum scoop-like. Antennae at the extreme tip of the head are protrusible. Abdominal segments each with an intercalary segment, giving the appearance of 23 segments in larva. PUPA. Respiratory tube two-segmented, bearing 15 spiracles apically. The head is rugose. Abdominal segments covered with sharp recurved spines, located on large tubercles. Last segment is forked, without spines or tubercles.

Adult habits and larval habitats: adult feeds on vertebrate blood, female may feed only once or as many as four times, males do not feed. Carter (1921) states that the adults are most active during the hottest parts of the day. Larvae are found in damp soil and along the seacoast in sand with organic matter.

The genus Leptoconops was erected by Skuse in 1889 for Lontoconos stygius Skuse. In 1893, Townsend set up the new genus Tersesthes and Noe (1905) proposed the genus Mvcterotypus, which are now placed as






17


synonyms of Leptoconops. In 1918 and 1921 the genus was divided into five new genera by Kieffer, these were: Holoconops, Schizoconops, Styloconops, Proteresethes and licroconops. Carter (1921) retained only the genus Letoconops, with two subgenera, Leptoconops s.s. and Holoconops, and also erecting a new genus, Acanthoconops. Acanthoconops was synonymized under Styloconops Kieffer by Kieffer (1921a) and it was reduced to a subgenus by Wirth (1952). Three subgenera of Letoconops are retained: Leptoconops, Holoconops and Styloconops.

This genus is different from most of the ceratopogonids and

readily recognized by the following characters: Palpus four-segmented. Antennae of female with 13 or 14 segments. Wings milky white; crossvein r-m absent; intercalary vein above vein M, prominent. Legs, basitarsi usually armed with ventral spines. Lamellae or cerci of female usually prominent. Male genitalia, basistyle with subbasal lobe; dististyle with terminal appendage.


Subgenus Leptoconops s. str.

Subgenus type: Leptoconops stydius Skuse, mon.

Distinguished from the others by having toothed tarsal claws;

sensory area of the third palpal segment large occupying almost the entire length. Female antennae 14-segmented. The lamellae of female long slender.

Illustration: Fig. 2 Leptoconops (Leptoconops) stygius Skuse.






18


Subgenus Holoconops Kieffer, 1918 Subgenus type: Leptoconops kerteszi Kieffer, orig. des.

This subgenus is characterized by having simple tarsal claws and subapical pit on third palpal segment. Female antennae 13-segmented. Female genitalia with moderately long, broad lamellae. Male genitalia with large, sclerotized parameres having lateral projections; dististyle bifid.

Illustration: Fig. 3 Leptoconos (Holoconops) kerteszi K.


Subgenus Styloconops Kieffer, 1921 Subgenus type: Leptoconops albiventris de Meijere, orig. des.

This subgenus is distinguished from the others by the following characters: Head with numerous spines or short bristles on frons and vertex; female antennae 14-segmented; palpus with sensory pit centrally or subbasally located. Tarsal claws with small basal tooth. Female genitalia with very short lamellae; parameres of male varied; the aedeagus fairly large.

Illustrations: Fig, 4 Leptoconops (Styloconops) albiventris (Meijere) and Fig. 5 Leptoconops (Styloconops)spinosifrons Carter.






19









Key for the Subgenera of Leptoconops


1. Female antenna with 13 segments . . . . . . . Holoconops (Fig. 3)

Female antenna with 14 segments ... . . *.. ... .2


2. Female genitalia with lamella very long and slender; head

with frons bare, few bristles on vertex; palpal sensory

area occupying almost entire length of the third

segment (Fig. 2B) . . . . . . . . . .. Leptoconops (Fig. 2)

Female genitalia with lamella very short (Fig. 5H);

frons and vertex with numerous spines or short

bristles (Fig. 4C, 5C); palpus with sensory pit subbasally

or centrally located (Fig. 4A, 5A) . . Styloconops (Fig. 4,5)






20


Genus Atrchooon effer

Atrichopogon Kieffer, 1906, Ann. Soc. Sci. Brux., 30: ?; 1906,

Gen. Insectorum, faso. 42:53; Coquillett, 1910, Proc. U. S. Nat.

9us., 37:512; Kieffer, 1913, Rec. Ind. Mus., 9:174; 1919, Bull.
Soc. Ent. France, p. 193; 1919, Ann, Mus. Nat. Hung., 17:20;

Goetghebuer, 1920, Mem. Mus, Roy, Hist. Nat. Belg., 8:29; Carter,

Ingram, and Macfie, 1921, Ann. Trop Med. & Parasit., 15:321;

Kieffer, 1924, Arch. Inst. Pasteur d' Algerie, 2:396; 1924, Bull.

Soc. R. Ent. Egypte, p248; 1925, Faune de France, fasc. 11,

P. 51; Edwards, 1926, Trans. Ent. Soc. London, 74:399; Johannsen,
1931, Arch. f. Hydrobiol., 9:414; Ingram and Macfie, 1931,

Dipt. Pat. & S. Chile, pt. 2, fasc. 4, p. 172; Macfie, 1932,

Ann. Trop. Med. Parasit., 26:31; 1934, Ann. Trop. Med. & Parasit., 28:184; 1939, Rev. Ent., 10:173; Tokunaga, 1940, Tenthredo, 3:109;

Macfie, 1940, Ann. Trop. Med. & Parasit., 34:15; Zilahi-Sebess, 1940, Folia. Ent. Hungar. 5:34; Johannsen, 1943, Ann. Ent. Soc.

Amer., 36:775; Macfie, 1947, Proc. R. Ent. Soc. Lon. (B), 16:72;

Wirth, 1956, Proc. Ent. Soc. Wash.,, 58:16; E~en and Saunders,

1958, Canad. J. Zool., 36:719; Tokunaga, 1959, Pacif. Ins., 1:179;

Tokunaga and Murachi, 1959, Ins. Micron., 12(3):116; Remn, 1959, Ent. Obozr., 38:683; Remm, 1961, Ent. Obozr., 40:925; Tokunaga,

1962, Pacif. Ins., 4:154. (Type-species: Ceratopogon levs Coquillett as exiles Coquillett, by desig. Coquillett, 1910). Didymophleps Weyenbergh, 1863, Stett, Ent. Zeit., 44:108 (Typespeciess Didymophleps hortorum Weyenbergh, mon.).









Kempia Kieffer, 1913, Rec. Ind. Mus., 9:162 (As subgenus of Dasyhelea)

(Type-species: Dasyhelea (Kempia) calcuttensis Kieffer, by ?

desig.).

Gyinohelea Kieffer, 1921, Arch. Inst. Pasteur Afr. Nord, 1:115

(Type-species: ?).

LophomVidium Cordero, 1929, An. Mus. Hist. Nat. Montevideo, 3:94

(Type-species: Lophomyidium uruuayense Cordero, mon.).

Psilokempia Enderlein, Tierwelt Mitteleur., 6(XVI):49 (Type-species:

Kempia appendiculata Goetghebuer, mon.).

DIAGNOSIS. Body stout, moderately hairy. Eyes pubescent or bare. Antennae 15-segmented, segments 4-10 globular or transverse, last five segments elongate, usually last segment terminates with nipple or style. Mesonotum bare or with few hairs. Legs slender; fore femora setose anteriorly. Wings moderately covered with macrotrichia, finer than in Forcipomyia and always lesser in male; costa long, about two-thirds of wing length, second radial cell well developed; intercalary fork distinct; median fork with short stem. Female abdomen usually bearing abdominal armature on sixth or seventh steral processes absent; aedeagus large, usually trilobed; parameres small, usually hook-shaped, or absent.

Apprarcimately 296 species have been described. Distribution: World-wide.

The biology and immature stages of Atrichopogon were studied by Johannsen (1952) and Ewen and Saunders (1958). The life history and






22


all stages of Atrichoporon levis were described in Boesel and Snyder (1944). Keys were provided by Thomsen (1937) and Wirth and Stone (1956) for North American species.

EGG. Atrichopogon levis egg is smooth, subshining, black; elongate and slightly curved. Most of the species lay eggs in masses, in aquatic situations.

LARVA. Body usually appears flattened as the sides of the segments often protrude into lateral processes. Head Iypognathous, antennae spur-like. Head capsule commonly bears eleven pairs of setae; abdominal segments one to seven with six pairs of setae which is similar to metathorax. The mesothorax has two lateral setae and three on the prothorax. Prothorax with bilobed, pseudopod bearing backward-curved hooklets arranged in two transverse rows. The ninth abdominal segment is cylindrical, covered dorsally by an unpaired sclerite. Posterior segment with a pair of protrusible anal papillae and the anal pseudopod. PUPA. Body broad anteriorly and tapering posteriorly. Respiratory horn variable in shape, appearing boot-shaped, flattened, or cylindrical. Thorax with six pairs of cuticular processes on the dorsal side. Posterior median point of mesothorax never overlaps the first abdominal segment. Abdomen with branched or setaceous projections on first five segments. Segment nine terminates with a pair of diverging

and tapering processes and covered with small stout spinules directed forward. Male genital sheaths are ventral and protrusible. Adult habits and larval habitats: Adult habits of Atrichoporon levis are given by Boesel and Snyder (1944). Males and females have been found flying in broad daylight above moist mud covered with algae.






23


Some were collected along streams and stagnant water. The species is abundant in the short grasses of parks, lawns or roadsides. Males usually swarm during sunset or other times of equivalent light. Larvae occur chiefly in shaded localities where mold, bacteria and algae are present in areas along the streams covering with algae, or wood and soil covering with moss. Most species are aquatic.

The adults of this genus are difficult to identify or even to

separate distinctly into subgenera. As pointed out by Edwards (1926), Macfie (1939c), Nielsen (1951) and IWirth (1952, 1956), the adults offer no good, clear-cut character or combination . of characters. The male genitalia might be very useful if there were no variation in appearance caused by protrusion or retraction. The form of the ventral armature of the female abdomen and the spermathecae might make a more satisfactory key, if an intensive study was made of the above characters.

Ewen and Saunders (1958) based their taxonomy mostly on the immature stages, which show excellent and accurate separations.

Atrichopogon is a well defined genus. It resembles Forciporvia by having a large empodium, but can be distinguished from ForciponTa adults by the following characters: second radial cell long, macrotrichia few or absent, microtrichia conspicuous and the nature of the fringe on the posterior margin of wing.


Subgenus Atrichopogon s. str.

Proboscis normal, straight. Eyes separated. Female antenna, segments 3-10 oval, segments 11-15 cylindrical and elongated. Spermatheca one.

Illustration: Fig. 6 Atrichopogon (Atrichopogon) levis (Coq.)






24


Subgenus Psilokempia Enderlein, 1936

Subgenus type: Kempia appendiculata End. (orig. desig.)

Enderlein (1936) erected genus Psilokempia for Kempia appendiculata End. It was reduced to subgenus by Remm (1959).

This subgenus is distinguished from the others by the following points: proboscis not curved, antennae with last five segments together about as long as segments 3-10 combined, palpi stout; macrotrichia of wing few or absent; one spermatheca; does not parasitize insects.

Illustration: Fig. 7 Atrichopogon (Psilokempia) arcticus (Coq) Subgenus Meloehelea Wirth, 1956

Subgenus type: Atrichopogon meloesugans Kieffer (orig. desig.)

Wirth (1956c) erected this subgenus based on specimens that

attack the bettles of family Meloidae. This group is characterized by having an unusually upcurved proboscis, palpi slender, antennae segments 11-15 together about twice as long as segments 3-10 combined; two spermathecae; macrotrichia abundant on wing; hind basitarsus about

2.5 times as long as the second segment. Illustration: Fig. 8 Atrichopogon (Meloehelea) meloesugans K.






25







Key for the Subgenera of Atrichopoon

1. Proboscis upcurved; two spermathecae (Fig. 8L).Meloehelea (Fig. 8)

Proboscis straight; one spermatheca ........... . . * . 2

2. Antennal segments 3-10 rounded, 11-15 subspherical (Fig. 7A);
female palpi with short segments (Fig. 70); eyes contiguous

(Fig. 7F). . . . . . . . . . . . . . . Psilokempia (Fig. 7)



Antennal segments 3-10 oval, 11-15 cylindrical and elongated

(Fig. 6A); female palpi with slender segments (Fig. 60);

eyes separated (Fig. 6F) . . . . . . . . Atrichopogon (Fig. 6)






26


Genus Forcipomyia Meigen

Forcipomyia (Megerle MS) Meigen, 1818, Syst. Beschr. Eur. Zweifl.

Ins., 1:59; Kieffer, 1906, Gen. Insectorum, fasc, 42:52; 1913,

Rec, Ind, Mus., 9:166; Ialloch, 1915, Bull. Ill. State Lab. Nat.

Hist., 10:311; Ingram and Macfie, 1924, Ann. Trop. Ned. & Parasit., 18:541; Edwards, 1926, Trans. Ent. Soc. London, 74:393; Johannsen,

1931, Arch. f. Hydrobiol., 9:407; Ingram and Yacfie, 1931, Dipt.

Pat. & S. Chile, pt. 2, fasc. 4, pp. 156,177; Macfie, 1932,

Tijdschr. Ent., 75:279; 1934, Ann. Trop. Med. & Parasit., 28:178;

Goetghebuer, 1935, Rev. Zool. Bot. Afr., 27:145; Macfie, 1939,

Rev. Ent., 10:142; 1940, Ann. Trop. Med. & Parasit., 34:16;

Tokunaga, 1940, Tenthredo, 3:61; Zilahi-Sebess, 1940, Folia.

Ent. Hungar. 5:21;.Johannsen, 1943, Ann. Ent. Soc. Amer., 36:777;

Harant and Huttel W. & N., 1951, Ann. Parasit. Hum. Comp.,

26:471; Saunders, 1956, Canad. J. Zool., 34:657; Tokunaga and

Murachi, 1959, Ins. Micron., 12 (3):144; Tokunaga, 1959, Pacif.

Ins., 1:256; Dessart, 1961, Bull. Ann. Soc. R. Ent. Belg., 97:

315; Remm, 1962, Loodusuur. Seltsi Aastar., 54:168; Tokunaga, 1962, Pacif. Ins., 4:169; Dessart, 1963, Mem. Inst. Sci. Nat.

Belg. (2) fasc. 71:13. (Type-species: Ceratopogon ambiguus

Meigen = Tioula bipunctata Linnaeus by desig. Coquillett, 1910) Tipula (part) Linnaeus, 1767, Syst. Nat., ed. 12, 2:978. Ceratopogon (part) YNigen, 1804, Klass., 1:31. Labidomyia Stephens, 1829, Cat. Brit. Ins., 2:239 (Type-species:

Tipula bipunctata Linnaeus, by desig. Westwood, 1840).






27


Tetraphora Philippi, 1865, Verh. Zool.-Bot. Ges. Wien, 15:630 (Typespecies: Tetraphora fusca Philippi, mon.).

Prohelea Kieffer, 1912, Spolia Zeylanica, 8:1 (Type-species: 7). Euforciporgia Nalloch, 1915, Bull. Ill. State Lab. Nat. Hist., 11:312

(Type-species: Euforcipomyia hirtipennis Malloch, by orig. desig.) Lepidohelea Kieffer, 1917, Ann. MNs. Nat. Hung., 15:364 (Type-species:

Ceratopogon chrysolophus Kieffer, mon.).

MIcrohelea Kieffer, 1917, Ann. AMs. Nat. Hung., 15:364 (Type-species:

Atrichopogon tropicus Kieffer, mon.).

Apelma Kieffer, 1919 (not Billberg, 1820), Ann. IMs. Nat. Hung.,

17:64 (Type-species: ApemaA auronitens Kieffer, 7 desig. by

Macfie, 1940).

Trichohelea Goetghebuer, 1920, Mem. Mns. Roy. Hist. Nat. Belg., 8:18

(Type-species: Trichohelea tonnoiri Goetghebuer, mon.).

Thyridonmia Saunders, 1925, Parasit., 17:268 (Type-species: Thvridog iapalustris Saunders, orig. desig.).

Phasmidohelea Mayer, 1937, Arb. Morph. Tax. Ent. Berlin-Dahlem,

4:233 (Type-species: Phasmidohelea crudelis Mayer, orig. desig.).

DIAGNOSIS. Body stout, densely covered with hairs. Eyes bare. Antennae 15-segmented, segments 3-10 pear-shaped to vase-shaped in female,. with dense verticils, segments 11-15 usually cylindrical to elongate, last segment often ending in a nipple; in male, segments 12-14 swollen at base, segment 12 very long, and segments 12-15 each with a single verticil. Humeral pits absent, mesonotum densely covered with hair and a few bristles. All legs and abdomen, especially






28


in male, with long bristles; feinora large, empodium well developed, usually as long as claws (absent in male of Apelma = Trichohelea). Wings with flattened macrotrichia abundant; costa ending about the middle of wing; first radial cell small or almost obliterated, second cell small but distinct; intercalary fork present; median fork with short stem, usually faint. Male genitalia not inverted; aedeagus large, shield-shaped; parameresusually present, slender and tapering, often fused at base or connected by transverse rod or plate.

Approximately 507 species have been described. Distribution: World-wide.

Biology and early stages of Forcipontia have been studied in

detail by quite a number of taxonomists. In 1923, Frew described the larval and pupal stage of F. piceus: Saunders (1924, 1925) published the most extensive papers on the immature stages of Apelma and Thyridomyia; de Meillon (1931) on one species from Transvaal; Mayer (1933) on F. comis Tokunaga (1934) on F. crinume; Lane (194V on F. argenteola and F. inornatipennis. Saunders (1956) had done the most extensive study in distinguishing the subgenera of Forciponia using characters of all stages. Recent papers on the immature stages and biology of Forciponia were published by Saunders (1959) on some species of Proforcipoyia and Thyridonlia, Saunders (1963) on subgenus Lasiohelea; Chan and Saunders (1965) on F. (Dacnoforciponyia) anabaenae; and Chan and Le Roux (1965) on F. (Neoforciponwia) eques and F. saundersi. Keys to early stages of North American species were presented by Thomsen (1937) and Wirth and Stone (1956).






29


EGG. Mayer (1934) stated that eggs of F. pcea are found in irregular masses and those of F, pllida in garlands. LARVA. The larvae of this genus exhibit a considerable range of variation of structure throughout the subgenera. Most of them have elongate body, distinctly segmented, each segment as long as wide and bearing either dome-shaped projections ending in hooks, or spearshaped, or fringed setae. Head hypognathous; they have eversible prothoracic and anal pseudopods.

PUPA. The pupa are less variable and may be confused with Atrichopogon. The posterior three segments remain tightly attached to the larval exuviae. Respiratory trumpets short, knob-like, bearing spiracles only at apex.

Adult habits and larval habitats: Saunders (1959) found many species of the subgenera Proforciponyia and TTyridonyia as pollinators of cacao. Chan and Saunders (1965) stated that F. (D.) anabaenae females are diurnal man-biters, prefering the shaded parts of the body. The females of Pterobosca group have specialized habit as ectoparasite of dragonflies. Females of eques (Johannsen) and related species suck the blood from the wing veins of lacewing flies, butterflies and other insects (Wirth and Stone, 1956). Larvae are mostly terrestrial, the gregarious larvae breeding under tree bark, in manure or moist decaying vegetable matter, in ant nests, and the like. F. calcarata (Coquillett) larvae were found along the rocky margin of a warm mineral spring at Alum Rocky Park in Santa Clara County, California (Wirth, 1952).






30


This genus is very large and shows a wide range of variation among the species. Many attempts have been made to subdivide the genus into subgenera. Saunders' system seems to be the most acceptable. He set up eight subgenera based on larval and pupal characters in addition to adult characters (Saunders, 1956). Tokunaga and Murachi (1959) mentioned using the following main characters to separate the subgenera: structure of male hypopygium; number of spernathecae; development of female antennae; value of hind tarsal ratio; segmentation of maxillary palp; wing venation, and development of scales and colored markings.

Members of this genus are readily recognized by having abundant macrotrichia on wing, radial cells very small and empodium ell developed as long as claws.


Subgenus Forcipmyia, s. str.

It is characterized by having two spermathecae; palpi usually five-segmented, third segmented with distinct sensory pit. Female antennal segments 3-10 flask-shaped, not compressed. Wing with dense macrotrichia, costa short, first radial cell small or absent, second one short. Male genitalia with parameres separated, pointed tip; aedeagus shield-shape, rather variable. Illustration: Fig. 9 Forciponyia (Forcipomyia) bipunctata (L.)


Subgenus Proforcipomyia Saunders, 1956

Subgenus type: Forcipomyia wirthi Saunders (orig. desig.)

Saunders (1956) proposed the new subgenus Proforciporrivia with

wirthi Saunders as type, to substitute for the uncertain identification






31


of hirtipennis of subgenus Euforcipomyia Malloch 1915. Since Nlloch's subgenus had been placed in miscellaneous species by Johannsen, also 'irth misplaced the species calcarata (Coquillett) in this subgenus.

Members of this subgenus can be differentiated from the others

by the following characters: wing densely covered with slender macrotrichia; costa about half of wing length; palpi five-segmented, sometimes the last two segments incompletely fused; first and second radial cells small or absent; spermathecae two. Male genitalia: paraneres 7- or V-shaped, arched, arising from basistylar apodemes; aedeagus shield-shaped.

Illustration: Fig. 10 Forcipomyia (Proforcipomyia) wirthi Saunders.


Subgenus Warikea Saunders, 1956

Subgenus type: Forcipomyia bicolor Saunders (orig. desig.)

Saunders (1956) believed this new subgenus to be a "young" offshoot from the parent stock of the subgenus Forcipomyia and that it had not had time to evolve ary great variety of species. Larval habitat is very uniform, all species but one have been found breeding in the leaf axils of water-holding plants in the Caribbean area. The larvae live above the water line and moved further up to pupate. The pupa has the best specific characters. The pupa cements itself to the leaf surface by the penultimate segment. The larval exuviae can usually be found stretched out on the leaf a few millimeters behind the pupa.

The distinctive characters of this subgenus are: female antenna

long, segments 3-10 with hyaline sensory hairs, segment 11 rather short; wing with thin microtrichia, second radial cell long. One spermatheca.






32


Male genitalia paramerea with' hooked tips. Illustrations Fig. 11 Forcipoyia (Warmkea) aeria Saunders and Fig. 12 Forciponria (Warmkea) tuberculata Saunders.


Subgenus Caloforciporyia Saunders, 1956 Subgenus types Forcipopyia caerulea Saunders (orig. desig.)

This subgenus is represented by two species from Brazil

(Saunders, 1956). The larvae have subcutaneous green or blue pigment granules in the fat body. Female antenna with hyaline sensory hairs. Segment 11 of the male antenna longest. Wing with elongated second radical cell. Male genitalia: parameres V-shaped; ninth sternite transverse.

Illustration: Fig. 13 Forciponia (CaloforcipoMyia) lauca dwards


Subgerns Metaforcipozia Saunders, 1956

Subgenus types Forcipopyia cerifera Saunders (orig. desig.)

A Brazilian species, cerifera Saunders was designated as a type of the subgenus (Saunders, 1956). The characteristics which differentiate it from other subgenera are: female antennae with basal segments flask-shaped, each with a hyaline hair; two spermathecae; male genitalia: parameres fused, basistylar apodemes curved and joined; larvae bearing wax-covered globules of liquid. Illustrations Fig. 14 Forciponyia (Metaforcip yia) pluvialis Malloch.






33


Subgenus Thyridorwia Saunders, 1925 Subgenus type: Thyridomyia palustris Saunders (orig. desig.)

It is characterized by having the basal antennal segments compressed in female; macrotrichia of wing not flattened, second radial cell subtriangular in female; one spermatheca; male genitalia with complicated aedeagus, lateral sclerites chitinized; parameres forning a pair of subtriangular internal plates reduced to small slender sclerites of well developed and connected caudal external lobes absent. Illustration: Fig. 15 Forcipokia (Thyridoyia) palustris Saunders.


Subgenus Synthyridormia Saunders, 1956

Subgenus type: Lasiohelea acidicola Tokunaga (orig. desig.)

Members of this subgenus share the common characteristic of male

genitalia with club-shaped parameres extending backward from variously shaped basal roots. Larva with well developed antennal flagellum. Body usually without scales. Spermatheca one. Illustration: Fig. 16 Forcipomyia (Synthyridoyia) colemani


Subgenus Trichohelea Goetghebuer, 1921

Trichohelea was adopted to replace Kieffer's genus Apelma, which was occupied previously by a genus of beetles. The important characteristics of this group are: empodium of male absent or vestigial. Wing with long second radial cell, thinly covered with macrotrichia. Male genitalia: parameres H-shaped with cross bar between bases of coxites and paired forward apodemal projections and backward external caudal lobes arising from bases of coxites; aedeagus large and shieldshaped.

Illustration: Fig. 17 Forcipoiyia (Trichohelea) fijiensis (acfie)






34


Subgenus Lasiohelea Kieffer, 1921 Subgenus type: Atrichopogon pilosipennis Kieffer (orig. desig.) = velox (Winnertz)

Tokunaga (1959) reduced Lasiohelea to a subgenus of Forcipomia. The following characteristics differentiate this subgenus from the others: wing with distinct bare areas adjoining veins, second radial cell long and very narrow; male genitalia with aedeagus subdivided into a pair of recurved sclerites, each bearing apical hook-like process; paramere simple, arch-like.

Illustration: Fig. 18 Forciponyia (Lasiohelea) fairfaxensis Wirth


Subgenus Neoforcipomnia Tokunaga & Murachi, 1959

Subgenus type: Forcipomyia pectinunguis de Meijere (Tokunaga & Mrachi, 1959:200-202)

This subgenus was erected by Tokunaga (1959) for species which are closely related to Lasiohelea but the male genitalia have arch-like parameres and spoon-shaped caudal lobes arising from coxite joints, coxites at lateral ends of paramere arms. The other characters are: basal antennal segment subspherical, distal segments elongated; wing densely covered with slender macrotrichia; two spermathecae. Illustration: Fig. 19 Forciomyia (Neoforcipomyia) eques (Joh.)


Subgenus Dacnoforcipomyia Chan and Saunders, 1965 Subgenus type: Forcipomia anabaenae C. & S. (Orig. desig.)

Chan and Saunders (1965) erected this subgenus based on the larval stages and the male. Members of this subgenus are characterized by the following points: larval head with one pair of small,






35


sharp, forward-pointed prominences on vertex; antennae short, stout; prothoracic pseudopod long, armed distally with two stout black hooks; anal pseudopod with only six stout black hooks; cauda short, rounded. Female antenna with basal five segments compressed, each with a pair of hyaline setae. One spermatheca. Male genitalia: basistyles slender; dististyles straight, uptapered, equal in length to basistyles, basistylar apodemes joined; parameres absent; aedeagus with two parts.

Illustration: Fig. 20 Forcio yia (Dacnoforcipomyia) anabaenae Chan and Saunders


Subgenus Phasmidohelea Mayer, 1937 Subgenus type: Phasmidohelea crudelis Mayer (orig. desig.)

Mayer (1937) proposed the genus Phasmidohelea for the three

species which attack phasmids. Wirth (1956c) reduced to subgenus of Forcipomyia based on the following characters which resembles ForcipoTyia: palpi with third segment swollen to apex; basitarsus short with strong spines; empodium large; two spermathecae. They are differentiated from the other subgenera by having a tick-like abdomen in female and mouthparts short, stout and highly sclerotized. Illustrations: Fig. 21 Forcipomyia (Phasmidohelea) fuliginosa Mg. and Fig. 22 Forcipomyia (Phasmidohelea) sp.


Subgenus Lepidohelea Kieffer, 1917 Subgenus type: Ceratooogon chrysolophus K. (orig. desig.)

Members of this subgenus are characterized by having foursegmented palpi; two spermathecae; male genitalia with parameres






36

separated; hind tibial comb present. Illustration: Fig. 23 Forcioomyia (Lepidohelea) annulatipes Macfie


Subgenus Pterobosca Macfie, 1932 Subgenus type: Ceratopogon aeschnosuga Meijere (acfie 1940 a:16)

It differs from subgenus Forcipomyia by having six elongated terminal antennal segments; empodium of female large and broad, adapted for clinging; wing with moderate macrotrichia but without scale-like hairs; spermatheca one or two. Illustration: Fig. 24 Forcipoxvmia (Pterobosca) fusicornis (Coq.)






37


Key for the Subgenera of ForcipoMia

1. Antennae with last six segments elongated (Fig. 24A);

legs with empodium very broad, modified for clinging (Fig. 24H) (female mainly ectoparasite on Odonata or

rarely on lacewing flies) . . . . . . . . Pterobosca (Fig. 24)

Antennae with last five segments elongated; empodium

large but normal . . . . . . . . . . . . . . . . . . . . 2


2. Two spermathecae .... . . . . . . . . . . . . . . . . . . . . . 3

One spermatheca . . . . . . . . . . . . . . . . 9


3. Last five segments of antenna combined not longer than the

preceding eight segments combined . . . . . . . . . . . . . 4

Last five segments of antenna combined much longer than the

preceding eight segments combined . . . . . . . . . . . . . 6


4. Sensory pit deep (Fig. 9C); antenna with segments almost

similar in shape and size (Fig. 9A) . . . Forcipoia (Fig. 9)

Sensoria scattering, without distinct pit; antenna with

last five segments long, each segment more than three

times longer than wide .... . .. . . * . . * . . 5


5. First tarsomere short or about the same size as the second

segment (Fig. 14G); palpi four-segmented (Fig. 14C);

male parameres fused midline, truncate anteriorly

(Fig. 14K) . . . . . . . . . Metaforcipornyia (Fig. 14)

First tarsomere longer than the second segment (Fig. 13J);

palpi five-segmented (Fig. 13C, D); male parameres V-shaped,
fused in the middle (Fig. 1314) . . . Caloforciporyia (Fig. 13)






38

6. Basitarsus very short about half the size of the second

segment (Fig. 21K, 22G) . . . . . . Phasmidohelea (Fig. 21,22)

Basitarsus simple, not shorter than the second segment . . . . 7


7. Antennal segments 3-10 oval to flask-shape (Fig. IOA)

. . . . . . . . . . . . . . . . . . Proforcipomyia (Fig. 10)

Antennal segments 3-10 depressed to transverse . . . . . . . . 8


8. Antennal segments three to seven depressed, 8-10 slightly

rounded (Fig. 17A); paramres H-shaped with cross bar and paired forward apodemal projections and backward external

caudal lobes (Fig. 17.0). . . . . . . . Trichohelea (Fig. 17)

Antennal segments 3-10 slightly transverse to oval (Fig. 19A);

parameres fused arch-like, arising from joint of basistyles (Fig. 19-0) . . . . . . . . Neoforcipomyia (Fig. 19)


9. Palpi four-segmented .... . .... .. .. ... . . 10

Palpi five-segmented . . . ..0. . . . . . .. .. 11


10. Parameres separated (Fig. 23-0); antennal segments similar

in size and flask-shaped (Fig. 23A) . . Lepidohelea (Fig. 23)

Parameres fused basally (Fig. 12D); antenna with last five

segments very long about four times as long as broad, segments 3-10 oval (Fig. 11A) . . . . . . . . Warnkea (Fig. 11,12)


11. Last five antenna segments combined not longer than all

the preceding eight segments combined . . . . . . . . . . . 12

Last five antennal segments combined much longer than the

preceding eight segments combined . . . . . . . . . 13






39


12. Male genitalia: parameres with basal apodeme forming a pair

of subtriangular internal plates (Fig. 15K)

S. . . . . . . * . . . . . . * . . . Thyridoyia (Fig. 15)

Male genitalia: parameres club-shaped, arise from basal

apodemes (Fig. 16K) . . . . . . . . Synthyridomyia (Fig. 16)


13. Third palpal segment broad without distal neck; sensory

pit large (Fig. 20B); antenna with first five segments compressed, bearing a pair of hyaline setae (Fig. 20A);

parameres absent, aedeagus separated, each with median

rounded lobe, anterior arms straight, posterior tip

blunt (Fig. 201) . . . . . . . . . Dacnoforcipotmyia (Fig. 20)

Third palpal segment slender or slightly swollen (Fig. 18C);

antennal segment 3-10 oval, 11-15 elongated (Fig. 18A); parameres slender joined mesally, arch-like (Fig. 18N),

aedeagus subdivided into pair of recurved sclerites, each

with apical hook-like process (Fig. 18.0).Lasiohelea (Fig. 18)








Genus Daasyhelea Kieffer

Dasyela Kiettffer, 1911, Bull. Soc. Hist. Nat. Metz, 27:5; 1913,

Rec. Ind. Ms, 9:179, 1913, Ann. Mus . Nat. Hung., 17:48; 1919, Bull. Soc. Ent. France, :192; Goetghebuer, 1920, Mem. M s. Roy.

Hist. Nat. Belg., 8:40; Carter, Ingram, and Macfie, 1921, Ann.

Trop. Med. & Parasit., 16:178; Kieffer, 1924, Bull. Soc. R. Ent.

Egypte :253; Edwards, 1926, Trans. Ent. Soc. London, 74:401;

Thienemann, 1926, Mitt. Geogr. Ges. Lubeck, 31:104; Ingram and

Macfie, 1931, Dipt. Pat. & S. Chile, pt. 2, faso. 4, :177;

Sebess V. Zilahi, 1931, Arch. Itydrobiol. 23:310; Goetghebuer,

1935, Rev. Zool. Bot. Afr., 27:164; Tokunaga, 1940, Tenthredo,

3:116; Zilahi-Sebess, 1940, Folia Ent. Hungar., 5:44; Johannsen,
1948, Ann. Ent. Soc. Amer., 36:778; Tokunaga & Murachi, 1959,

Inst. Micron. 12(3):238; Rem, 1962, Tartu Riik. 0lik. Toim, 120:108; Tokunaga, 1962, Pacit. Ins. 4:189. (Type-species:

Dasyhelea halophila Kieffer. (orig. desig.)

Prokempia Kieffer, 1913, Rec. Ind. aMs., 9:163, 179 (Type-species:

not desig., two orig. incl. spp., Dasyhelea (Prokempia) ornaticornis Kieffer and D. (Prokempia) longicornis Kieffer).

Pseudoculicoides Malloch, 1915, Bull. Ill. State Lab. Nat. Hist.,

10:309 (Type-speciess Ceratopogon mutabilis Coquillett, orig.

desig.).

Tetrahelea Kieffer, 1925, Arch. Inst. Pasteur d'Algerie, 3:422

(Type-species: Culicoides insignicornis Kieffer, orig. desig.). Cryptoscena Enderlein, 1936, Tierwelt Mitteleur., 6:51 (Type-speciess

Ceratopoon palustris Meigen, mon.).



















: cn-ti s rith short :ulccence. Ev::.ior aerentn.l cn rn

: : : :u y h,:S : y-v.:u .. .l :.a.ice aCtn.a r:e : nctal aot s: i:;.':. s clong ~. 1:.-::1 :- zy p::.p lo.ndor, first so: i .......-ly s "pra' -, only or cn..s viosi'ble. :.ral pits sibpsent,

L-* -,i h,!2. a Q -1.;O. 0 z-annozt::1 c;V:v z it: ht:.:C oc Clrdo; tcarsacL zc*:o*r ipl

e::.:o 0:.1 c C:: I1; a:.;zc:111c-! z:::D3 . _ -Til th rcrotrichia over aI or pat of cOf:' :rc; ccAt:. Cin :bo 1l:i of t-ng length; first radial cell obiteratcz, a :cc or.d -:i coil n.ll Pth s>ua ea c: n;







SO- ra . :-m. - e--i-: - l "- :;i and tap -ir u -i -- -t.e


nl.a 3 i-C**Or .1 :- : 1 -; :: r -cal,



d. n ; n .c









-- .. ....,...*, ..L, "* ...'. ' "





;bo53 to aC S-1_. . , O
'. .~"9 Y






42


The following descriptions are taken from Thomsen (1937), Wirth and Stone (1956) and Wirth (1965): EGG. Dasyhelea eggs are horseshoe-shaped, the masses are small, detected by microscope.

LARVA. Head oval. ~outhparts hypognathous. Eyes crescent-shaped wi-th heavily pigmented. Mandibles usually three-toothed. Hypopharynx heavily sclerotized, composed of dorsal and ventral parts, with posterior portions bearing teeth. Body segments short and thick, colored by pigmented fat-bodies beneath the hypodermis. Protrusible proleg on last segment bearing hooks. Retractile filamentous anal gills posterior to prolegs.

PUPA. Anal segment with pair of setigerous protuberances in addition to apicolateral processes. Opercula, lacking long setae, with or

without hooks in some species.

Adult habits and larval habitats: adults visit flowers, feeding on nectar. The immature stages are semiaquatic. Larvae are found in the blanket algae of ponds or slow streams, algae on dripping banks, and the fermenting sap of tree ulcers. The larvae cannot swim but travel by crawling over the substrate. Most members of the genus spin tubular cases in the last instar.

This genus vaserected by Kieffer in 1911 for the Indian species, Dasyhelea halophila. It has characteristics intermediate between Forcipomyia and Culicoides. The wing venation resembles Forciportia but tarsi lacks empodium as in Culicoides, however, it can be distinguished from the above genera by the following points: eyes pubescent; palpi four-segmented; wing vein R1 and R. fused forming






43


a single vein; humeral pits absent; male genitalia: parameres often asymmetrical with paired basal arms and single median lobe.

Dasyhelea is rather a difficult genus. The males are comparatively easier to identify than females. Tokunaga and Murachi (1959) suggested the use of the female subgenital plate for identification.


Subgenus Dasyhelea, s. str.

Wing with one radial cell usually slightly longer than broad,

rarely square or missing. Scutum pruinose grayish or bluish. Female antenna with flagella segments basally striped, segment 15 with long apical style. Spermatheca one. Legs usually banded. Male genitalia: ninth sternite not modified, tergite with long apicolateral processes; hook or lobes on inner side of basistyles. Illustration: Fig. 25 Dasyhelea (Dasyhelea) grisea (Coq.)


Subgenus Dicryptoscena End., 1936 Wing with two radial cells, both longer than broad, more or less

equal in length. Palpal segments slender. Female antenna with flagella segments not basally striped, segment 15 with apical style. Spermatheca one, subspherical, with short duct. Male genitalia: ninth tergite with apicolateral processes very short.


Subgenus Pseudoculicoides Malloch, 1915 Subgenus type: Ceratopogon mutabilis Coq.

Wing with second radial cell square or slightly longer than broad. Female antenna with flagella segments not striped, without apical style. Scutum usually dull, black; with or without pruinose gray pattern and rarely shining. Spermatheca one. Male genitalia:









ninth sternite produced caudad in the middle, joined with aedeagus; parameres symmetrical or asymmetrical. Illustration: Fig. 26 Dasyhelea (Pseudoculicoides) mutabilis Coq.


Subgenus Prokempia Kieffer, 1913

Wing with square second radial cell, macrotrichia scanty.

Female antenna with flagella segments not striped, no apical style. Scutum usually shining, without pruinose pattern. Spermatheca one. MKle genitalia: aedeagus irregular, H-shaped, separated from caudal border of ninth sternite; latter with two long processes; parameres asymmetrical.

Illustration: Fig. 27 Dasyhelea (Prokempia) cincta (Coq.)






4.


Key for the Subgenera of Dasvhelea


1. Two radial cells present on wing. . . . . . . . .. Dicryptoscena

One radial cell on wing .... . . . . . . . . . . . . . . 2


2. Female antenna with flagella segments basally striped,

last sogment with long apical style (Fig. 25A).

Second radial cell rarely square (Fig. 25C,D)

. . . . . . . . . . . . . . . . . . . . . Dasyhelea (Fig. 25)

Female antenna with flagella segments not striped,

without apical style (Fig. 26A, 27A). Second

radial cell square (Fig. 26E, 27D) . . . . . . . . . . 3


3. Scutum usually dull, black, with or without pruinose

gray pattern . . . . . . . . . Pseudoculicoides (Fig. 26)

Scutum usually shining, without pruinose pattern"

. .. . . . . . . . Prokemoia (Fig. 27)






46


Genus Paradasyhelea Macfie


Paradasyhelea Macfie, 1940, Ann. Trop. Med. Parasit, 34:17; Wirth

and Lee, 1959, Bull. Brooklyn Ent. Soc., 54:114; Wirth and

Blanton (in press). (1ype-species:Dasyhelea brevitalpis

Ingram and Macfie orig. desig.).

DIAGNOSIS. Eyes widely separated, covered densely with fine hairs. Clypeus hairy. Proboscis short about half as long as the length of the head, vestigial mouth parts. Palpi four-segmented, the largest segment bearing long modified sensory hairs on distal portion, the shallow, small sensory pit indistinct. Antenna 15-segmented. First segment with a row of hairs, third to fifteenth segment uniform in shape. Male antenna plumose. Humeral pits present. Legs slender, tibial comb with four spines, claws small, equal, with bifid tip in male; vestigial hair-like empodium. Wing with short costa, radial cells obliterated, r-m cross vein oblique almost in line with a single radial vein, intercalary fork well developed; anal angle obtuse; macrotrichia abundant. One spermatheca. Male genitalia with ninth sternum excavated posteriorly; ninth tergum with well developed apicolateral processes; basistyles and dististyles simple; aedeagus arched, median part small; parameres appear only as a sclerotized rod connecting the two ventral roots, middle portion broken.

Approximately three species have been described. Distribution: South America and Australia. PUPA. Elongate conical as in Culicoides and Dasyhelea, respiratory trumpet stalked, flattened, mid portion wrinkled transversely and bearing papillae around apex. Tubercles of cephalothorax with spines,









abdominal tubercles each with fine hair; apicolateral processes ending in a blade-like spine pointed almost perpendicular to the body.

Macfie (1940a) proposed this genus for Dasyhelea brevipalpis

Ingram and Macfie, which he differentiated from Dasyhelea and Forci. pomyia, as discussed by Ingram and Macfie (1931). The important characteristic is the unsculptured antennal segments; not binodose in segments 12-14 of male.

Paradasyhelea is closely related to Forcipormia, Dasyhelea, and Culicoides. It resembles Forcipomvia in having abundant macrotrichea on wing; scutellum hairy; eyes well separated; antennae with sensory pegs scattered. It is similar to Dasyhelea by the following points: palpi four-segmented; eyes hairy; proboscis and mouthparts poorly developed; and the shape of apicolateral processes of ninth tergite in male. Like Culicoides, it has prominent humeral pits; antennal sensory pits and aedeagus Y-shaped.

Illustration: Fig. 28 Paradasyhelea minuta Wirth and Lee Genus Culicoides Latreille

Culicoides Latreille, 1808, Gen. Crust. et Ins., 4:251; Johannsen,

1905, Bull. N. Y. State Mis., 86:101; Kieffer, 1906, Gen.

Insectorum, fasc. 42:53; Johannsen, 1908, Bull. N. Y. State Mus.,

124:267; Kieffer, 1913, Rec. Ind. YMs., 9:162; 1919, Ann. Mus.

Nat. Hung., 17:31; Goetghebuer, 1920, EMm. Mus. Roy. Hist. Nat.

Belg., 8:48; Carter, Ingram and Macfie, 1920, Ann. Trop. Med.

Parasit., 14:212; Hoffman, 1925, Amer. Hyg., 5:274; Edwards,

1926, Trans. Ent. Soc. London, 74:403; Johannsen, 1931, Arch.

f. Hydrobiol., Suppl., 9:428; Mhkerji, 1931, Ind. J. Med. Res.,






48



18:1054, 1056; 1931, Nature, 127:339; Pomerantzen, 1932, Nag. Parasit. Leningrad, 3:183; Virmer, 1932, Sborn. Ent. Odd. Narod. Mus. Praze 10:132; Goetghebuer, 1933, Flieg. Pal. Reg. 13a:37;

1935, Rev. Zool. Bot. Afr., 27:170; Tokunaga, 1937, Tenthredo, 1:273; Costa Lima, 1937, Mem. Inst. Oswaldo Cruz, 32:411; Root and Hoffman, 1937, Amer. J. Hyg., 25:156; Macfie, 1937, Proc. Roy. Ent. Soc. London (B), 6:112; Causey, 1938, Amer. J. Hyg., 27:399; Xono and Takahasi, 1940, Ins. Mats. 14:69; Tokunaga, 1940, Tenthredo, 3:141; Takahasi, 1941, Ins. Matsum. 15:80; Zilahi-Sebess, 1941, Folia Ent. Hung., 6:32; Fox, 1942, Puerto Rico J. Pub. Health and Trop. Med., 17:412; James, 1943, Pan-Pacif. Ent., 19:148; Johannsen, 1943, Ann. Ent. Soc. Amer., 36:779; Colaco, 1946, Ana.Inst. Xed. Trop. Lisbon, 3:222; Fox, 1946, Ann. Ent. Soc. Amer., 39:248; Barbosa, 1947, Anv.Soc. Biol. Pernambuco, 7:3; Hill, 1947, Ann. Trop. Med. Parasit., 41:55; Macfie, 1948, Ann. Trop. Med. Parasit., 42:67; Fiedler, 1951, Onderstepoort J. Vet. Res., 25:10; irth, 1951, Ann. Ent. Soc. Amer. 44:76; Downes and Kettle, 1952, Proc. R. Ent. Soc. Lond. (B), 21:77; Goetghebuer, 1952, Biol. Jaarb., 19:186; Gutsevich, 1952, Parazit. Sborn. Zool. Inst. Akad. Nauk SSSR, 14:79; Kettle and Lawson, 1952, Bull. Ent. Res., 43:428; Lee and Reye, 1953, Proc. Linn. Soc. N.S.W., 77:373; Ortiz, 1953, Rev. Sanid. Asist. Soc. 18:?; Foote and Pratt, 1954, Publ. Hlth. Pbnogr., Washington No. 18:10; Fox, 1955, J. Agr. Univ. P. R., 39:219; Kettle and Lawsen, 1955, Proc. R. Ent. Soc. Lond. (B) 24:46; Ortiz and Leon, 1955,









Bol. Inf., Cient. Nac. 7, No. 67:566; Arnaud; 1956, lNcroentomology, 21:134; Jones, 1956, ProC.Ent. Soc. Wash., 58:25; Le-wis, 1956, Canad. Ent., 88:551; Forattini, 1957, Agr. Fac. Hig. Univ. S. Paulo, 11:194; Khalaf, 1957, Amer. Midl. Nat. 58:187; Wirth and Blanton, 1956, Proc. Ent. Soc. WI-ash., 58:308; Wirth, 1958, J. Kansas Ent. Soc., 31:85; Do Prada and Gil Collado, 1959, Med. Trop. Z drid, 34:423; Ortiz, 1959, Rev. Sanid., Asist. Soc., 24:355; Sen and Das Gupta, 1959, Ann. Ent. Soc. Amer., 52:629; Tokunaga, 1959, Pacif. Ins., 1:204; Tokunaga and Mhrachi, 1959, Ins. Micron., 12, No. 3:322; Wirth and Blanton, 1959, Proc. U. S. Nat. as., 109:262; Wirth and Hubert, 1959, Pacif. Ins., 1:5; Campbell and Pelham-Clinton, 1960, Proc. Roy. Soc. Edinb. (B) 67:196; Gutsevich, 1960, Opred. Faune SSSR, 72:44; Wirth and Hubert, 1960, Ann. Ent. Soc. Amer., 53:648; Callot and Kremer, 1961, Ann. Parasit. Hum. Comp. 36:686; Delfinado, 1961, Fieldiana (Zool.), 33:636; Hubert and Wirth, 1961, Proc. Ent. Soc. Wash., 63:236; Jones, 1961, Ann. Ent. Soc. Amer., 54:741; Khalaf, 1961, Beitr. Ent. 11:450; Nagaty and Morsey, 1961, J. Arab. Vet. Med. Ass., 21:364; Pais Caoiro, 1961, Estud. Ens. Docum. Jta. Inv. Ultramar., No. 86:46; WIirth and Hubert, 1961, Pacif. Ins. 3:12; Galukhova, 1962, Trudy. Zool. Inst. Akad. Nauk SSSR, 31:218; Tokunaga, 1962, Pacif. Ins. 4:203, 458; Wirth and Hubert, 1962, Ann. Ent. Soc. Amer., 55:184; Jamnback and Wirth, 1963, Ann. Ent. Soc. Amer., 56:186; Tokunaga, 1963, Plant Prot. Bull. 5:120; Wirth and Blanton, 1967, Fla. Ent. 50(3):209. (Type-species: Culex pulicaris Linnaeus, as

Culicoides punctata Latreille, orig. desig.)







50


Cecacta Poey, 1851, Mem. Hist. Nat. Isla Cuba, 1:236.(Type-species:

Oecacta furens Poey, ron.).

Psychophaena Philippi, 1865, Verh. Zool.-Bot. Ges. Wien, 15:628

(Type-species: Psychophaena pictipennis Philippi, mon.).

Haematorpidium Goeldi, 1905, em. Mls. Goeldi, 4:137 (Type-species:

Haematomyidiun -araense Goeldi, orig. desig.).

Cotocripus Brethes, 1912, Ann. Mus. Nac. Buenos Aires, 22:451

(Type-species: Cotocrious caridei Brethes, mon.).

Haemophoructus acfie, 1925, Bull. Ent. Res., 15:349 (Type-species:

Haemophoructus maculipennis Macfie, mon.).

Synhelea Kieffer, 1925, Arch. Inst. Pasteur d'Algerie, 3:423

(Type-species: ?).

Prosapelma Kieffer, 1925, Arch. Inst. Pasteur d'Algerie, 3:417

(Type-species: Prosapelma cinerea Kieffer, orig. desig.).

Hoffmania Fox, 1948, Proc. Biol. Soc. Wash., 61:21 (Type-species:

Culicoides inamollae Fox and Hoffman, orig. desig.).

DIAGNOSIS. Body slender, with moderate hairs. Eyes usually bare. Female antennae with segments 3-10 oval or rounded and usually much shorter than the last five segments which are usually cylindrical in shape. Male antennae plumose, three apical segments long. Mesonotum usually with pattern, covered with short hairs and usually with few bristles; humeral pits well developed. Legs and tarsi simple; claws of both sexes tiny but equal; empodium vestigial. Wings densely covered with microtrichia; macrotrichia always present near wing tip; wings often with spotted pattern; costa usually to two-thirds wing






51

length; two radial cells, almost equal; media petiolate with parallel branches. Male genitalia with ninth sternite short, with mesal excavation; ninth tergite usually with apicolateral processes present; basistyle often with both dorsal and ventral root developed; dististyle long and curved; aedeagus variable, usually cone-shaped with anterior lateral arms; parameres usually separate with sharp point or irregular

shape.

Approximately 1000 species have been described. Distribution: World-wide.

The immature stages of Culicoides have been studied by several

authors. The American species were described by the following authors: Painter (1926), Dove, Hall and Hull (1932), Thomsen (1937), Fox (1942), Smith and Lowe (1948), Williams (1951), Wirth (1952), Jamnback, W'all and Collins (1958), Jones (1961). On the European species, the studies had been done by: Rieth (1922), Thienemann (1928), Lenz (1934), layer (1934) and Jobling (1953); descriptions of the British species have been given by: Hill (1947), Downes (1950), Lawson (1951) and Kettle and Lawson (1952).

Culicoides immature stages were reported; from Africa by Goetghebuer (1933), Carter, Ingram and Macfie (1920), and Ingram and hacfie (1921); from Brazil by Lutz (1912), Forattini, Rabello and Pattoli (1956, 1960); from India by Patton (1913) and Patel (1921); from French Guiana by Forattini and Rabello (1956); and from Japan by Tokunaga (1937).

Particularly detailed descriptions have been given by Carter, Ingram and Yacfie (1920), Hill (1947), Kettle and Lawson (1952) and Jones (1961).






52


Breeding places of Culicoides were reported by Lutz (1912),

Rieth (1922), Thorpe (1927), Jobling (1929), Mayer (1934), Buckley (1938), Wirth, 0. M., W. W. irth and F. S. Blanton (1968). EGGS. Eggs are elongated and slightly curved. They are deposited in masses.

LARVA. Larvae are eel-like form, without prolegs. The body segments are short and thick cylinders, each segment carrying a pair of lateral setae. The last segment bears three pairs of anal setae, short and light in color. The head is well sclerotized with prognathous mouthparts. Mandibles are curved. Pharyngeal skeleton has combs. PUPA. The pupa is well sclerotized. It has dark brown color and is free from larval exuvium. Respiratory organ is elongated, tubular, and constricted at the base. The operculum is either vith or without spines. The anal segment has one pair of apicolateral spines. Adult habits and larval habitats: The adults are generally most active at dawn and dusk (Kettle, 1962). Their flight and other activities are strongly influenced by temperature and wind and other conditions. The females require a blood meal, but males feed only on nectar or plant juices. Each species has a range of vertebrate hosts on which it will feed. Feeding may occur anytime, but some species are crepuscular. The resting places are usually in damp cool situations, such as algae, the base of trees, moss, grass or dead leaves (Kettle, 1962).

Larvae may be found in a wide variety of moist environments,

such as ponds, creeks, lakes, tree holes, slime-covered bark, rotting forest litter, rotting banana stalks. In the recent paper, Wirth, O. M., W. W. Wirth and F. S. Blanton (1968) reported finding two





53


habitats of Pana a Culicoicdes. These habitats were rotting spadics of the Panama hat palm, Carludovica nalnata Ruiz and Pavon and older de-a7ing in florescences of the "beefsteak heliconia" Kelicoria arise Eooker f.

The majority of the species of this genus may be recognized by havirn spotted wings which is absent in some species. Yalloch (1915) was the first to show the sensory pits as a distinct generic character. These pits do not appear in ee, the nearest genus to Culicoides. They are present in Ceratooon, Monohelea and some Stilobezza which can be differentiated from Cuicoides since the females have large cla7s or modified legs.

This genus comprises a large number of species widely distributed over the world. Cany attempts have been made to divide Culicoidcs into suogenera. The important references are Vargas (1960) w-ho separated the subgenera based on wing patterns and male genitalia and Fox (1955) who emphasized only the male genitalia.


Subgenus Culicoides s. str.

Cell R5 with a marginal dark spot shaped like an hour-glass;

cubital fork in dark area; second radial cell in light spot. %Kle genitalia: basistyles short and strong pilosity at the inner margin; pararmeres with short haIrs at tip; aedeagus V-shaped. Illustration: Fig. 29 Culicoides (Culicoides) yLkonensis Foffran


Subgenus Trithecoides 7,irth and Hubert, 1959

Subgenus type: Culicoides flaviscutatus W. & H., 1959 (orig. design )

Characters as follow: three well developed spermathecae; second






54


radial cell long; wing with two pale areas, over r-m and over apex of second radial cell; large pale area on the basal part of wing across extreme wing tip. le genitalia with ventral root reduced; aedeagus blunt tip.

Illustration: Fig. 30 Culicoides (Trithecoides) flaviscutatus W. & H.


Subgenus Haemophoructus Macfie, 1925

Subgenus type: Haemophoructus maculipennis M'acfie (orig. desig.)

Female wing with a single radial cell.

Illustration: Fig. 31 Culicoides (Haemohoructus) gemallus 'acfie


Subgenus Meijerehelea Wirth and Hubert, 1961 Subgenus type: Ceratopogon guttifer Yeijere (orig. desig.)

One spermatheca; wing with costal pale spot in cell R5 immediately distal to second radial cell and a round pale spot at extreme apex of cell. Male genitalia: aedeagus usually flaring at tip; parameres with large basal knob directed laterad. Illustration: Fig. 32 Culicoides (NMeijerehelea) Ruttifer (Mleijere)


Subgenus Hoffmania Fox, 1947

Subgenus type: Culicoides inamollae Fox and Hoffman (orig. desig.)= insignis Lutz, 1913.

Pale spot on vein M2; cubital fork in light area; macrotrichia abundant; r-m crossvein dark with large light spot. Male genitalia: aedeagus bottle-shaped, parameres fused in some species. Illustration: Fig. 33 Culicoides (Hoffmania) insignis Lutz






55


Subgenus Avaritia Fox, 1955

Subgenus type: Ceratooopon obsoletus eigen (orig. desig.)

Wing marking faint; macrotrichia scanty. Yale genitalia: ninth tergite without apicolateral processes; ventral root of basistyle very long.

Illustration: Fig. 3J Culicoides (Avaritia) obsoletus (Mg.) Subgenus Oecacta Poey, 1851

Subgenus type: Oecacta furens Poey (mon.)

Second radial cell in dark spot; clear spot on vein M2 absent; three pale spots on Mi. Yale genitalia: ventral root with retrorse tooth; parameres with spinose tip; aedeagus triangular. Illustration: Fig. 35 Culicoides (Oecacta) furens (Poey)


Subgenus Drymodesnyia Vargas, 1960 Subgenus type: Culicoides copiosus Root and Hoffman, 1937 (orig. desig.)

Wing hairy; second radial cell within dark area; cell R5 with two pale spots, one is about the middle of cell. Male genitalia: basistyles with ventral hook, dististyles hook-shaped, paramere with sharp tip.

Illustration: Fig. 36 Culicoides (Drymodesnyia) copiosus R. & H.


Subgenus Diphaeomyia Vargas, 1960 Subgenus type: Culicoides baueri Hoffman, 1925 (orig. desig.)

Male genitalia with ventral root long, curved, rounded apically with a small notch at base; parameres with many short spines. Illustration: Fig. 37 Culicoides (Diphaeoywia) baueri Hoffman






56


Subgenus Anilonia Vargas, 1960 Subgenus type: Culicoides covagarciai Ortiz, 1950

Wing with darkest spot occurring basal to second radial cell. Male genitalia with inner margin of basistyle spinose; aedeagus bottle-shaped; parameres fused or closely appressed with a terinal brush.

Illustration: Fig. 38 Culicoides (Anilonyia) covagarciai Ortiz


Subgenus Beltrany'ia Vargas, 1953 Subgenus type: Culicoides crepuscularis Ialloch, 1915 (orig. desig.)

Wing dark with pale markings and mrry long hairs. Male genitalia: aedaegus V- or Y-shaped, truncate, broad tip.

Illustration: Fig. 39 Culicoides (Beltranyia) crepuscularis Mlloch


Subgenus Glaphiromyia Vargas, 1960 Subgenus type: Culicoides scopus Root and Hoffman, 1937 (orig. desig.)

Wing with pale spot on vein M1 and M2; r-m crossvein dark in some species. Male genitalia: ventral root long and sharp; paraneres with terminal hairs; aedeagus arch-shaped. Illustration: Fig. 40 Culicoides (Glaphiromyia) scopus R. & H.


Subgenus Haemato myidium Goeldi, 1905 Subgenus type: Culicoides paraensis (Goeldi, 1905)

Crossvein r-m usually dark but clear basally. Male genitalia:

ventral root well developed with retrorse tooth; parameres with spinoso tips; aedeagus rounded apically. Illustration: Fig. 41 Culicoides (Haematomyidium) paraensis (Goeldi)






57


Subgenus Macfiella Fox, 1955

Subgenus type: Ceratonogon phlebotomus Williston

Second radial cell in- a dark spot. Mle genitalia with ninth tergite rounded apically; basal root long; dorsal root somewhat boot-shaped.

Illustration: Fig. 42 Culicoides (Mcfiella) phlebotoms (Will.)


Subgenus Mataen~la Vargas, 1960 Subgenus type: Culicoides mojingaensis Wirth and Blanton, 1953 (orig. desig.)

Vein M, usually without pale spot; TM and M2 with long light bands. Ventral root of male genitalia prominent; parameres sharp pointed.

Illustration: Fig. 43 Culicoides (Mataemyia) mojingaensis W. & B.


Subgenus Monoculicoides Khalaf, 1954

Subgenus type: Ceratopogon nubeculosus Meigen (orig. desig.)

Wing with second radial cell in a dark spot. One spermatheca. Male genitalia with paramere fused forming a plate and usually bifid distally; aedeagus deeply bifid; dorsal root often boot-shaped. Illustration: Fig. 44 Culicoides (Monoculicoides) variioennis (Coq


Subgenus Selfia Khalaf, 1954

Subgenus type: Culicoides hieroglyphicus Malloch (Orig. desig.)

Wing dark without pale marking. Spermathecae not apparent.

hale genitalia with parameres fused. Aedeagus with two sclerotized pieces often fused apically.

Illustration: Fig. 45 Culicoides (Selfia) hieroalv-phicus 31lloch






58


Key for the Subgenera of Culicoides

1. Spermathecae unsclerotized, not apparent; wing dark

without pale daring . . . . . . . . . . . . Selfia (Fig. 45)

Spermathecae sclerotized; wing usually with distinct
pale spots .. ... ........ . ... . 2


2. One spermatheca . . . . . . . . . . . . . . . . . . . . . . . 3

Two to three spermathecae ....... ....... . . 5


3. Parameres separated . . . . . . . . . . ... . . . . . . . 4

Parameres fused to form a plate usually bifid distally

with two apical points (Fig. 44.0) . . lonoculicoides (Fig. 44)


4. Paramere with large basal knob directed laterad (Fig. 32P),

stem short, rather stout simple tip without distal

fringing spines. . . . . . . . . . . Ieijerehelea (Fig. 32)

Paramere with small base, tapering, directed basally,

and pointed tip. (Fig. 39*0) . . . . . . Beltranria (Fig. 39)


5. Three well developed spermathecae (Fig. 30M).Trithecoides (Fig. 30)

Two well developed spermathecae ............... 6


6. Female wing with a single radial cell (Fig. 31E) . Haemo-hoructus (Fig. 31)

Female wing with two radial cells, almost equal in size . . . 7 7. Ventral root absent or poorly developed . . . . . . . . . . . 8

Ventral root large and vwll developed . . . . . . . . . . . . 10






59

8. Base of paramere sharply bent and close together, tip

with short hairs (Fig. 29L); wing without pale

spot on vein l2 . . . . . . . . . . . . . Culicoides (Fig. 29)

Base of paramere gradually bent; wing with pale spot

on vein iM2 ..... ........ . ... . 9


9. Wing with cubital fork in a dark area (Fig. 38E)

* . . . . . . . . . . . . . . . . . . . . Anilomyia (Fig. 38)

Wing with cubital fork in a light area (Fig. 33E)

* . . . . . . . . . . . . . . . . . . . Hoffnania (Fig. 33)


10. Second radial cell included in a pale area (Fig. 34E)

. .. . . . . . . . . . Avaritia (Fig. 34)

Second radial cell included in a dark area . . . . . . . . 11 11. Middle of vein M2 not straddled by pale spot ...... .. 12

Middle of vein MN straddled by pale spot . ... . . . . ... 13


12. Vein M1 and M2 within long dark bands (Fig. 41E); ventral

root with retrorse tooth (Fig. 41N). .Haematomyidium (Fig. 41)

Vein M1 and M2 within long pale bands (Fig. 43D);

ventral root prominent . . . . . . . . . hataemyia (Fig. 43)


13. Vein M and M2 within dark bands . . . . . . . . . . . . . . 14

Vein M, and M2 within pale bands . . . . . . .... . 15


14. Third palpal segment with sensory pit scattering, without

a t:iue pit (Fig. 42C); aedeagus without a prominent arch,

apical portion long, truncate distally, basally with a

long, curved hook-like pointed processes (Fig. 42N)
. acfiella (Fig. 42)





60


Third palpal segment with a true and large sensory pit

(Fig. 40C); aedeagus arch with long stem (Fig. 40N)

. . . . . . * . . . . * * * * * . . GlaphiromTia (Fig. 40)

15. Aedeagus with small, tapering tip, with short process on arch
(Fig. 37N) . . . . . . . . . . . . . . . Dirhaeomyia (Fig. 37)

Aedeagus with broad or truncate tip, without process

on arch . * * * * * * * * * * * 16


16. Base of paramere bent perpendicularly, sharp tip, not

spinose (Fig. 36-0) . . . . . . . . Dymodesmyia (Fig. 36)

Base of paramere small and graduall bent, subapical

portion with globose process and tip splnose (Fig. 35-0)
Oecacta (Fig. 35)






61


Genus Austroconops Wirth and Lee

Austroconous Wirth and Lee, i958, Proc. Linn. Soc. N. S. Wales,

83:337 (Type-species: Austrocono2ps ncmillani Wirth and Lee,

mon.)

DIAGNOSIS. Eyes hairy, contiguous above; vertex sparsely haired. Antenna 15-segmented, flagella segments spherical basal, becoming oval distally. Clypeus small, with four long hairs; proboscis shorter than head; mouthparts well developed, mandible with eight large teeth. Palpi "four-segmented," the fourth and fifth segments fused; third segment swollen without sensory pit, sensoria covered almost half of the mid portion of the segment. Thorax robust, humeral pits conspicuous. Legs unarmed; hind tibia with plumose spur, tibial comb four spines. Claws small and equal; empodium. rudimentary. Wing milky, radial veins distinctly thickened, costa long, r-m crossvein very oblique, forming almost a straight line with base of media and posterior side of radial cells; radial cells long and broad; media with long petiole; anal cell large; macrotrichia absent. Abdomen short with blunt apex and short cerci. T-wo subequal spermathecae and a

rudimentary one. jale unknown.

Only one species known.

Distribution: W. Australia.

Adult habits and larval habitats: Female blood sucking, immature stages unknown.

This genus was erected in 1958 by Wirth and Lee for Austroconons mcmillani of Australia. It has not been discovered outside the Australian region.






62


The members of this genus have the characteristics of Cul-coilesh Leptoconos and Stilobezzia. It resembles Leotoconous by the following characters: palpi four-segmented, fourth and fifth segments fused into one segment. W.ing nmilky white, without macrotrichea; empodiu= absent; antennal segments similar in size and shape; blood sucking habit diurnal. Wing venation is similar to Stilobezzia, with long costa and well developed radial cells and petiolate-mdia. It has the characteristics of Culicoides in having the humeral pits on pronotun; small and equal claims, empodium vestigial; antennal segments not sculptured.

Illustration: Fig. 46 Austroconops mcrillani Wirth and Lee


Genus Camotooterohelea Wirth and Hubert

Camntooterohelea Wirth and Hubert, 1960, Fieldiana (Zool.), 42(7):89

(Type-species: Ca ptoterohelea hoogstraali Wdirth and 1Hbert, (mon.) DIAGNOSIS. Eyes bare, widely separated. Antenna 15-segmented, cylindrical. Palpi very short, only two segments present; the third and the fused segment of the fourth and the fifth. Proboscis vestigial, shorter than palpus, mandibles with very fine teeth. Thorax dark and stout; scutum with anterior median pair of oval hyaline pits and few hairs. Legs rather stout; front tibia with a lanceolate spur and a compact comb, hind tibia with more hairs on comb; each tarsomcre bearing a single long, stout, slightly curved claw. Wing broad on basal half and tapering distally; costa long; two radial cells present, the first one well developed but the second one small, the radial veins thickened; the rest of the wing venation includes the cubital stem, 1A and possibly Cu1 appears from posterior wing fold to margin of the wing.







63


Disc of wing modified with two folds transversely continuous from costal end to end of Cul proximally and about halfway from this line to wing tip distally, fold areas densely covered with microtrichia; along vein Cul with long, spinelike nicrotrichia as well as over radial cells and along margin of proximal fold and the patch in front of the distal fold; macrotrichia absent. Abdomen stout with single oval spermatheca. Male genitalia small; ninth tergite rounded; basistyle rather stout, dorsal root prominent; dististyle slender, tapering with hook-like tip; aedeagus arched -ith large body.

Only one species known.

Distribution: Philippines, Malaya.

This genus is readily separated from the other genera by having

two-segmented palps; proboscis rudimentary; all claws single; wing greatly modified w-ith broad basal half and tapering distally; wing vegnation with only radial vein, cubital stem and the first anal vein present; microtrichia spinelike. Illustration: Fig. 47 Carmotooterohelea hoogstraali Wf. & H.



Genus Ceratopogon Meigen

Ceratopogon Neigen, 1803, Illiger's Mag. Ins., 2:261; Kieffer, 1899,

Bull. Soc. Ent. France, p. 69; Johannsen, 1903, Bull. N. Y. State

Mus., 86:97; Kieffer, 1906, Gen. Insectorum, fasc. 42:48; 1919,

Ann. Mus. Nat. Hung., 17:1; Goetghebuer, 1920, Mem. Mus. Roy.

Hist. Nat. Belg. 8:19; Edwards, 1920, Ann. Mag. Nat. Hist., 6:129;

1926, Trans. Ent. Soc. London, 74:407; Johannsen, 1934, J. N.T. Ent.

Soc., 42:343; Tokunaga, 1940, Tenthredo, 3:151; Johannsen, 1943,






64


Ann. Ent. Soc. Amer., 36:780; Wirth, 1952, Univ. Calif. Publ.

Ent., 9:198; de Meillon, 1959, S. Afr. Anim. Life, 6:340;

Clastrier, 1961, Inst. Pasteur Alger., 39:401; Tokunaga, 1964,

Pacif. Ins. 6:292. (Type-species: Tipula barbicornis Fabricius,

mon. 7= Ceratooogon communis Meigen).

Helea Neigen, 1800, Nouv. Class. des mouches a deux ailes, p. 18;

Goetghebuer and Lenz, 1933, in Lindner, Flieg. Pal. Reg., 77:5,

48; Stone, 1941, Ann. Ent. Soc. Amer., 34:411 (Type-species:

Ceratopogon communis Meigen, desig. Coquillett, 1910).

Brachypogon Kieffer, 1899, Bull. Soc. Ent. France, p. 69 (Type-species:

Ceratoopogon vitiosus Winnertz, orig. desig.).

Psilohelea Kieffer, 1917, Ann. Mus. Nat. Hung., 15:294 (Type-species:

Ceratopogon candidatus Winnertz, mon.; = niveipennis Meigen). Isohelea Kieffer, 1917, Ann. Mus. Nat. Hung., 15:295 (Type-species:

Ceratopogon lacteipennis Winnertz, desig. Kieffer, 1919; =

sociabilis Goetghebuer).

Anakempia Kieffer, 1924, Bull. Soc. Hist. Nat. Moselle, 30:13 (Typekejia grandiforceps Kieffer, orig. desig.).

... a de 1illon, 1939, Ruwenzori Exped., 1934-35, 1(5):103

(Type-species: Ceratopogon (Fanthamia) adulator de Xeillon,

orig. desig.).

DIAGNOSIS. Body stout with few hairs. Eyes pubescent. Female antennal segments 3-10 rounded, the last five segments oval; male antennae plumose with last three segments longer than the preceding ones. Legs slender; empodium vestigial or absent; claws large in female, equal or almost equal. Wing without microtrichia and










macrotrichia or with few mcrotrichia near the wing tip; costa slightly longer than the middle of wing; usually with two radial cells, about the same size; intercalary fork indistinct; median fork usually with lower branch absent or interrupted. Male genitalia not inverted; ninth tergite small; basistyles swollen; aedeagus varies; parameres separate.

Approximately 219 species have been described. Distribution: World-wide.

Kettle and Lawson (1952) was the only paper published on the immature stages and the breeding places of Ceratopogon of British species.

LARVA. Head with posterior ventral suture long, well sclerotized and pigmented. Antennae more prominent (isohelea) or indistinct (Ceratooon). Dorsal comb with seven to nine teeth on each side. PUPA. Thorax dark, nodular. Operculum dark, covered with dark nodules; respiratory horn short and stout, very dark, except at the

tip. Metathorax not indentate. Caudal spine dark.

Breeding places always are associated with Culicoides. They were reported found in moorland, in marshes and often in large

numbers in Snharnum spp.
is
This genus is closely related to Culicoides but the wingA without microtrichia or pattern and the claws are very large.


Subgenus Ceratopogon s. str.

Vein M2 complete or narrowly interrupted; both radial cells distinct, longer than broad.

Illustration: Fig. 48 Cerato-ogon (Ceratoporon) culicoidithorax Hoffman.






66


Subgenus Isohelea Kieffer, 1917 Subgenus type: Ceratooogon lacteipennis Winnertz (orig. desig.) sociabilis Goet.

Vein widely interrupted at base; costa short not much beyond the middle of wing; radial cell small or absent. Illustration: Fig. 49 Ceratopoon (Isohelea) stigmalis Coq.


Subgenus Brachypogon Kieffer, 1899 Subgenus type: Ceratooogon vitiosus Winnertz (orig. desig.)

Wing with second branch of media absent.


Subgenus Nilohelea Kieffer, 1921 Subgenus type: Nilohelea albipennis Kieffer (mon.)

Wing with first and second radial cell obliterated.


Subgenus Fanthamia de ebillon, 1939 Subgenus type: Ceratopogon adulator de Meillon.

Wing with complete first radial cell, second one obliterated; male antenna as in female; hind claws of female greatly unequal; macrotrichia present on distal portion of wing.






67







" o : aCu... ;,h of C( ,one .....



T::;o co:,olo radial c ll.s o. .,t ..* *

One or both radil cells obliterated . . . . . .....



S o : 'b r .nch of :: dia p c . . . . . . .* . .* *. . * *





ba , co-ta




.-* . ... . c r O .................... . . . . . .. ......



. . . . ; ., .1




::-M le of 1 - * _ * * '






68


Genus Alluaudorria Kieffer

Alluaudomyia Kief., 1913, Voyage Ch. Alluaud et R. Jeannel en Afrique

Orientale, Dipt. 1:12; de Meillon, 1939, J. Ent. Soc. S. Africa 2:7; Okada, 1942, Trans. Nat. Hist. Soc. Formosa 32:315; Wirth,

1952, Ann. Ent. Soc. Amer. 45:423; Tokunaga and Marachi, 1959,

Ins. Micron. 12(3):352; Clastrier, 1961, Arch. Inst. Pasteur

Alger; 39:428; Tokunaga, 1963, Pacif. Ins. 5:215; Wirth and

Delfinado, 1964, Pacif. Ins. 6:602. (Type-species:

Alluaudoryia imoarunouis Kief. mon.).

Neoceratopogon Yalloch, 1915, Bull. Ill. State Lab. Nat. Hist.,

11:310 (Type-species: Ceratopogon bellus Coquillett).

Prionognathus Carter, Ingram and Yacfie, 1921, Ann. Trop. Med. &

Parasit., 16:309 (Type-species: Prionognathus marnoratus Carter,

Ingram and Macfie, orig. desig.).

Thysanognathus Ingram and Macfie, 1922, Ann. Trop. Med. & Parasit.,

16:244 (n. nom. for Prionognathus Carter, Ingram and cfie,

not Laferte-Senectere, 1851).

Isoecacta Garrett, 1925, Seventy New Diptera, p. 9. (Type-species:

Isoecacta poeyi Garrett, mon.).

DIAGNOSIS. Body slender, moderately hairy, mostly small size. Eyes bare or hairy, separated or contiguous. Female antennal segments 3-10 long, segments 11-15 elongated. Male antennae with well developed plumes. Palpi slender; third segment simple. Wing without microtrichia, macrotrichia appearing along the wing tip, wing usually with 1-15 or 20 small dark spots and sometimes with grayish streaks along the wing veins; male wing bare or hairy. One radial cell present (the first one






69


completely fused), media petiolate. Legs slender, in some species with long hairs. Male claws small and equal; female claws very unequal in hind legs, inner claw longer. Spermathecae one or two. MIale genitalia highly modified; ninth tergite always long, usually with well developed apicolateral processes; ninth steri;- short, concave mesally; basistyle and dististyle simple; aedeagus arched with large stem; parameres usually separated.

Approximately 122 species have been described.

Distribution: North America, Europe, Africa, Asia, and Pacific

Islands.

Immature stages and biology of Alluaudomyia of North American species have been reported in Thomsen (1937), illiams (1953) and \irth and Stone (1956).

EGG. Long and narrow, almost straight. Eggs of A. needh~ni placed side by side horizontally, forming a flat ribbon-like band, not jelly covered.

LARVA. Brown with pear-shaped head. The dorsal side of the body colored by pigments in the fat bodies, causing the larva to appear dark red. Last body segment with eight setae as long as or longer than last segment, with addition of four short setae. PUPA. Light brown with dark thorax. Respiratory trumpet funnel-form, covered entirely with scales; respiratory spiracles arranged in pairs. Cuticular projections on the body segments vary in shape and number. Last segments with slender and pointed projections. Larval habitats: larvae of Alluaudorlyia were found in algae and in mud in New York. Thomsen (1937) reported that larvae of A. splendida were found in algae, decaying leaves from swamps, and in the mud






70


near the edge of ponds in New York; A. needhami larvae were found in blanket algae in ponds. The larvae are predaceous. ,illia z (1953) sunnarized three breeding habitats of A. bella and A. needhani as follows: areas of permanent flowing water to areas of temporary -ater where the soil is firm enough to walk on; soil with dense growth of vegetation to areas with no vegetation and in sunny areas where the temperature reaches 900F as well as shady areas; pH of soil may range from 5.7 to 7.3.

This genus has distinctive characters and is readily recognized from the other genera in the follo-,wing points: wing usually adorned with one to twenty small black spots; rmicrotrichia absent; emDpodiun vestigial and claws of female unequal on hind legs. Illustrations: Fig. 50 Alluaudontia bella (Coq.) and Fig. 51 Alluaudoyia narva Wirth.



Genus Stilobezzia Kieffer

Stilobezzia Kieffer, 1911, Rec. Indian :s., 6:118; 1913, Rec. Indian

Mus., 9:184; 1917, Ann. Mus. Nat. Hungzarici, 15:307; 1919, Bull.

Soc. Ent. France 5:192; Goetghebuer, 1920, Mem. ZLus. Roy. Hist.

Nat. Belgique, 8:59; Carter, Ingram, and Macfie, 1921, Ann. Trop.

Med. Parasit., 15:324; Edwards, 1926, Trans. Ent. Soc. London,

74:411; 1929, Notulae Ent., 9:9; Johannsen, 1931, Arch. Hydrobiol.

Suppl., 9:430; Ingram and Macfie, 1931a, Dipt. Patagonia and S.

Chile, pt. 2, fasc. 4:191; Macfie, 1932, Ann. Trop. Xed. Parasi.t.,

26: 41; de Millon, 1938, Proc. Roy. Ent. Soc. London (B), 7:266; Tokunaga, 19140b, Tenthredo, 3:154; Johannsen, 1943, Ann. Ent.Soc.






71


Amer., 36:781; Lane, 1947, Rev. Ent., 18:197; Lee, 1948, Proc

Linn. Soc. N. S. 17., 72:345; 'Wirth, 1953, Proc. U. S. Nat. -us.,

103:57; Lane and Forattini, 1958, Rev. Bras. Ent., 8:203;

Tokunaga and NMrachi, 1959, Ins. TYcron. 12(3):363; Tokunaga, 1959, Pacif. Ins. 1:298; Lane and Forattini, 1961, Rev. Bras.
Ent., 10:83; Tokunaga, 1962, Pacif. Ins. 4:207; Clastrier, 1963,

Arch. Inst. Pasteur Algerie 41:41; Toku mrpan 1963, Pacif. Ins.

5:249; Das Gupta and Wirth, 1968, U. S. Nat. 1Ns. Bull., 283:16.

(Type-species: Stilobezzia festiva Kieffer, orig. desig.)

Hartomyia Malloch, 1915, Bull. Ill. State Lab. Nat. Iist., 10:339

(Type-species: Ceratopogon pictus Coquillett, orig. desig.).

Neostilobezzia Goetghebuer, 1934, in Lindner, Flieg. Pal. Reg., 78:53

(Type-species: genotype not designated).

DIAGNOSIS. Large size, slender with sparsely hairs. Eyes bare. Antennae of female with segments 3-10 oval to short cylindrical, segments 11-15 strongly cylindrical; male antennae plumose, hairs well developed on segments 3-12. Tesonotum with or without few bristles; humeral pits inconspicuous. Legs slender, without spines in all femora; fourth tarsal segment cordiform, fifth segment elongated and usually clavate; first tarsal segment of mid leg bearing subbasal spur; female claws very large and unequal, bifid; male claws small and equal; empodium absent. Wing long with few macrotrichia along wing tip; costa long; two radial cells present, second one well developed; median fork distinctly petiolate and long. Usually two spormathecae and a third small one. Male genitalia: very short ninth sternite; ninth termite rounded; basistyle and dististyle simple; aedeagus






72


reduced to pair of oblique lateral aclerites; parameres with modified, well developed submedian processes.

Approximately 248 species have been described. Distribution: World-wide.

The immature stages and biology are not well known. Few species have been described. The pupa of spirogyrae was described by Carter, Ingram and Macfie (1921). Ingram and Macfie (1922) gave the description of immature stages of poikiloptera from West Africa. In Europe, Thienemann (1928) gave the figures of the pupa of gracilis. Kettle and Lawson (1952) included two instars of ochracea and Pracilis in their paper. Mayer (1934) studied the Dutch East Indies species of perspicua, and the pupa of soror. Thomsen (1937) described and illustrated antennalis and bulla early stages from New York. Lane, Forattini, and Rabello (1955) described and illustrated pupa of glauca and wygodzinsky from Brazil. LARVA. Mouthparts reduced, directed anteriorly. Head capsule well developed; pharyngeal skeleton with several combs, principal comb of pharyngeal angulus undivided; larva without prolegs but with caudal bristles. Last abdominal segment without terminal hooks, with only short setae (Thomsen, 1937).

PUPA. As in Culicoides but differs in having trumpet constricted basally; no spines on operculum.

Adult habits and larval habitats: adult habits are poorly known. Edwards (1920b) indicated that the female of S. gracilis feeds on various small chironomid midges. Malloch (1915) and de Meillon (1936) reported that the Notata Group holding their wings when at






73


rest. Edwards (1926) and Malloch (1915) reported finding the same habit in the Subviridis Group. The larvae are commonly found in mud and algae along pond margins (Thomsen, 1937). In West Africa, Ingram and Macfie (1921) reported two species from Pistid in a swamp. Johannsen (1932) and Mayer (1934) reared the larvae of perspicua in hot springs and in a rice field in Sumatra.

This genus is readily recognized by having female claws large and very unequal or a single long claw with barb; fourth tarsomere cordiform; first tarsomere of mid legs with subbasal spur; wing with second radial cell much longer than first which may be entirely absent.


Subgenus Stilobezzia s. str.

Members of this subgenus are characterized by the following

characters: wing with two radial cells, without macrotrichia. Most of the species are described from the tropical region. Illustration: Fig. 52 Stilobezzia (Stilobezzia festiva K.


Subgenus Eukraiohelea Ingram and Macfie, 1921

Subgenus type: Eukraiohelea africana I. & M. (Macfie, 1940a)

Johannsen (1934) placed Eukraiohelea as a subgenus of Parabezzia Malloch (1915), but in comparing the male genitalia of Parabezzia petiolata Malloch 1915, with those of the four species of Eukraiohelea, the latter are more closely related to Stilobezsia.

This subgenus is characterized by having one radial cell on wing; vein R1 arising at or past r-m; legs with stout spines on fore femur midventrally or at base.

Illustration: Fig. 53 Stilobezzia (Eukraiohelea) elegantulata (Joh.)









Subgenus Neostilobezzia Goetghebuer, 1934

Subgenus types Ceratopogon ochraceus Winnertz, 1852 (Wirth 1953a)

Goetghebuer (1934) distinguished this subgenus from Stilobezzia by the presence of macrotrichia at the end of the wing. In addition, all the species known are yellowish or brown to black, and the male genitalia with aedeagus bent and humped in the middle and basistyle without inner lobe. Most of the species are described from the temperate region.

Illustration: Fig. 54 Stilobezzia (Neostilobezzia) lutea (Malloch).






75





Key for the Subgenera of Stilobezzia

1. Wing with one radial cell (Fig. 531E); legs with spines on
fore femora; hind femora slender and unarmed (Fig. 53G)
. . . . . . . . .. . . . . Eukraiohelea (Fig. 53)

Wing with two radial cells; femora unarmed . . . . . * * * * * 2

2. Wing entirely bare of macrotrichia (Fig. 52E)
S. .. * . . .. . . . Stilobezzia (Fig. 52)

Wing with macrotrichia on the surface or at tip
(Fig. 54E) o � � � � � . Neostilobezzia (Fig..54)






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Genus chinohelea Macfie

Echinohelea Maofie, 1940, Proc. R. Ent. Soc. Lend. (B) 9:187; Wirth,

1951, Proc. Ent. Soc. Wash. 53:319; de Meillon, 1959, Novos

Taxa Ent. 13:15; 1960, J. Ent. Soc. S. Afr. 23:409; Tokunaga, 1963, Pacif. Ins. 5:231. (Type-species: Echinohelea ornatipennis Macfie, orig. desig.).

DIAGNOSIS. Eyes bare. Antennae of female with segments 11-15 elongated, the last segment usually shorter than the preceding four segments. Male antennae without plumose hairs. Thorax without anterior tubercle or spine, but with few stout bristles. Wing without macrotrichia, costa long, not extending beyond RS, M, and M2 forked before or at r-m, two radial cells present; second one well developed. Femora of all legs not greatly swollen, almost all femora and tibia armed with long spines, scattered rather irregularly; fourth tarsal segment short, not distinctly bilobed or cordiform, fifth segment very long, slender, unarmed, claws equal or unequal, each with barb. Abdominal tergites of female without gland rods. Male genitalia: ninth sternite excavated mesally, basistyles large, dististyle small and slender, aedeagus arched, ending in two parts, each bearing hook; paramere fused mesally, expanded ends.

Approximately 11 species have been described.

Distributions British Guiana, Liberia, New Britain, New Guinea, Upper Volta, and U.S.A. (Florida and Virginia).

This genus was reported by Macfie (1940b) for three very similar species from British Guiana. They resemble in general appearance some species of Serronria but hind femora not greatly swollen. The distinctive characters of the genus are as follows: all femora and tibiae






77

armed strongly with well developed spines; last antennal segment always distinctly short. The male is readily recognized by having no plumes on antennae.

Illustration: Fig. 55 Echinohelea lanei Wirth


Genus Monohelea Kieffer

Monohelea Kieffer, 1917, Ann. Mus. Nat. Hung., 15:294; Goetghebuer,

1920, Meom. Mas. Roy. Hist. Nat. Belg., 8:63; Ingram and Macfie, 1921, Ann. Trop. Med. & Parasit., 15:344; Edwards, 1926, Trans.
Ent. Soc. London, 74:410; Tokunaga, 1940, Tenthredo, 3:156;

Johannsen, 1943, Ann. Ent. Soc. Amer., 36:781; Lane, 1948, Arg.

Fac. Hig. Saude Pub. Univ. Sao Paulo, 1:225; Wirth, 1953, Proc.

U. S. Nat. Mas., 103(3320):136; Tokunaga and Murachi, 1959,
Ins. Micron., 12(3):404; Clastrier, 1963, Arch. Inst. Pasteur

Algerie,41:55; Tokunaga, 1963, Pacif. Ins., 5:238; Lane and

Wirth, 1964, Studia Ent. (N.S.) 7:211; Wirth and Williams, 1964, Ann. Ent. Soc. Amer., 57:302. (Type-species: Monohelea hieroglyphica Kieffer, orig. desig.)

Schizohelea Kieffer, 1917, Ann. Mus. Nat. Hung., 15:295; 1918, Ann.

Mus. Nat. Hung., 16:57; 1919, Ann. Mus. Nat. Hung., 17:89;

Goetghebuer, 1920, Mem. Mus. Roy. Hist. Nat. Belg., 8:65;

Edwards, 1926, Trans. Ent. Soc. London, 74:411; Johannsen, 1943,

Ann. Ent. Soc. Amer., 361782 (Type-species: Ceratopogon leucopeza Meigen, as Ceratopogon copiosus Winnertz, mon.)

Allohelea Kieffer, 1917, Ann. Mus. Nat. Hung., 15:364 (Type-species:

Sphaeronvas pulchripennis Kieffer, orig. desig.).






78


DIAGNOSIS. Body short, moderately hairy. Eyes bare or with very fine pubescence. Humeral pits inconspicuous. Fore and mid legs simple, fourth tarsal segment short, cylindrical, claws large or small but equal in both sexes; hind legs with thickening of femora and tibiae, spines absent; ventral of basitarsi with a spine at base, with or without spine at tip; fifth tarsal segment of female with one long claw and with or without another short one; hind claws of male small and equal except in some species; empodium absent. Wing with few macrotrichia at the tip; costa long; two radial cells, second cell well developed; intercalary fork inconspicuous; media with short stem, 4~ sometimes interrupted near base. Two spermathecae present. Male genitalia with bilobed or truncated ninth tergite; ninth sternite convex mesally; aedeagus usually with a pair of lateral sclerites and an additional dorsomedian lobe; parameres fused or paired.

Approximately 93 species have been)described. Distribution: World-wide.

The habits of Monohelea are unknown. The larva of only one species, M. calcarata, has been found in moss in a spring habitat (Strenzke, 1950).

The males of this genus are easy to identify by characters of the male genitalia but the great variation among the females makes identification very difficult. Wing markings are not a reliable

character. Leg markings are more constant.

This genus resembles Stilobezzia but differs particularly in having the fourth tarsomere cylindrical, the female claws with the four anterior claws equal but the hind claw single and long.






79


Subgenaua Monohelea as. tr. This subgenus is distinguished by the following characters wing usually with pattern, first radial cell complete, base of M2 narrowly interrupted; scutum with pruinose pattern. Illustration: Fig. 56 Monohelea (Monohelea) hieroglyphica K.


Subgenus Schizohelea Kieffer, 1917 Subgenus types Ceratopogon copiosus Winnertz (mon.) = leucopeza (Meigen).

Members of the subgenus are characterized by: wing without

pattern, first radial cell complete, M broadly interrupted at base; scutum polished black.

Illustration: Fig. 57 Monohelea (Schizohelea) leucopeza (Mg.) Subgenus Isthmohelea I. & M., 1931 Subgenus types Isthmohelea disjuncta I. & M.

This subgenus is characterized by having two radial cells

separated for some distance by fusion of vein between them, first radial cell complete.


Subgenus Kiefferongia Mayer, 1937 Subgenus type: Kiefferonia gorana Mayer

This subgenus differs from the other subgenera by having first radial cell obliterated.






80




Key for the Subgenera of Nonohelea


1. First radial cell obliterated . .. . . . . . . .Kiefferoia

First radial cell complete . . . .. . . . . . . . . . . . . 2


2. Fourth tarsomere cordiform or bell-shaped . . . . . . Isthmohelea

Fourth tarsomere cylindrical or subcylindrical . . . . . . . . 3


3. Wing usually with pattern; i not broadly interrupted
at base (Fig. 56D) . . . . . . . . . . . Monohelea (Fig. 56)

Wing without pattern; 4 broadly interrupted at
base (Fig,. 57D) . . . . . . . . . Schizohelea (Fig. 57)






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Genus Seonw Meigen

Berronia (Megerle, M. S. in) Meigen, 1818, Syst. Beschr. Eur. Zefl.

Ins., 1:83; Johannsen, 1905, Bull. N. Y. State Mus., 86:108;

Kieffer, 1906, Gen. Insectorum, fasc. 42t64; Johannsen, 1908,

Bull. N. Y. State Mus., 124:269; Malloch, 1914, Bull, Ill. State Lab. Nat. Hist., 10:216; 1915, Bull. Ill. State Lab. Nat. Hist.,

10:331; Kieffer, 1917, Ann. Mus, Nat. Hung., 15:294; 1919,

Ann. Mus. Nat. Hung., 17:69; Goetghebuer, 1920, Mem. Mis. Roy.

Hist. Nat. Belg., 8:70; Edwards, 1926, Trans. Ent. Soc. London,

74:409; Tokunaga, 1940, Tenthredo, 3:156; Johannsen, 1943,
Annm. Ent. Soc. Amer., 36:781; Kettle and Lawson, 1952, Bull.

Ent. Res., 43:465; Clastrier, 1963, Arch. Inst. Pasteur Algerie,

41:60. (Type-species: Ceratopogon femoratus Meigen, mon.) Prionamyia Stephens, 1829, Cat. Brit. Inst., 21237 (Type-species:

Ceratopogon femoratus Meigen, desig. Westwood, 1840).

Ceratolophus (part) Kieffer (not Barboza de Bocage, 1873), 1899,

Bull. Soc. Ent. France, p. 69 (Type-species: Ceratopogon femoratus

Maigen, mon.).

Johannseniella Williston, 1907, Jour. N. Y. Ent. Soc., 15:1 (nom. nov.

for Ceratolophus K., not Barboza de Bocage).

DIAGNOSIS. Body slender and almost bare. Eyes bare. Female antennal segments 3-10 short, segments 11-15 cylindrical and elongated; male antennae plumose. Mesonotum covered with small bristles, humeral pits absent. First four legs slender, without spines; fourth segments of tarsi cordate, claws simple and almost equal; femora of hind legs distinctly swollen in both male and female, armed ventrally with






82


numerous spines, tibia slender And curved basally; fourth segments of tarsi cylindrical, female claws highly unequal, without empodium. Wings with microtrichiae, macrotrichiae present in some species; costa long; two subequal radial cells present; intercalary fork absent, media sessile to short petiolate. Two spermathecae. Male genitalia: ninth sternite small; ninth tergite cone-shaped, with large anal lobes; basistyles stout, dististyles long and narrow; aedeagus small with long arms; parameres small and simple.

Approximately 27 species have been described.

Distribution: North America, Europe, Africa, Malaysia and Caroline Island.

The larval and pupal stages of S. femorata were given in Kettle and Lawson (1952).

LARVA. Large eyes. Two combs on epipharyns and each armed with a slender lateral plate; seven teeth on each half of dorsal; and 11i teeth on ventral side. No pigment, only a pair of grey and clear latiral substances in mesothorax and metathorax. PUPA. Operculum thickly covered with nodules. Respiratory horn short, pale at base. Tip and distal area with eight or nine papillae. Metathorax concave. Arrangement of setae differs from Culicoides. Caudal spines dark at tips.

Larval habitats Strenzke (1950) reported finding the larvae of S. femorata in damp moss in Germay.

Female claws resemble Heteronia but the main point of difference is that the femur of hind leg is greatly swollen and armed, fore tibia arcuate. This character is found on fore leg of HeterogiM a. Illustration: Fig. 58 Serronyia femorata (Mg.)






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Genus Macrurohelea Ingram and Macfie

Macrurohelea Ingram and Macfie, 1931, Dipt. Patag. and S. Chile,

part II, fasc. 4:203; Lee, 1962, Proc. Linn. Soc. N.W., 87:

339; Wirth, 1965, Pan-Pacif. Ent., 41:46. (Type-species:

Macrurohelea caudata Ingram and Macfie, orig. desig.).

DIAGNOSIS. Eyes well separated, with fine pubescence. Palpi fivesegmented. Antennae with all segments short. Legs slender, without spines. Female claws small and equal. Wing without macrotrichia, microtrichia present; costa long; two radial cells present. Female abdomen with modified tenth segment, which curves forward and reaches the ventral side of the anterior segments. Spermathecae two or three. Male genitalia: with adeagus Y-shaped, parameres separated, long but larger at the distal portion, basal arms well developed.

Approximately five species have been described. Distribution: South America and Australia.

Ingram and Macfie, 1931, proposed this genus for two species, M. caudata I. & M. from southern Chile, and M. thoracica I. & M. from southern Argentina. The female of this genus is readily recognized by the uniquely modified abdomen, with the tenth segment long and bent forward and appressed to the venter of the preceding segment.

The wing venation and male genitalia are similar to Stilobezzia

but differ in having equal tarsal claws. This character also separates them from Monohelea. Macrulohelea can be separated from Echinohelea Macfie and Serronyia Meigen by the absence of femoral spines; and from Parabezzia Malloch by the presence of two radial cells. Illustration: Fig. 59 Macrurohelea setosa Wirth






84


Genus Parabezzia Malloch

Parabezzia Malloch, 1915, Bull. Ill. St. Lab. Nat. Hist., 10:358;

Johannsen, 1943, Ann. Ent. Soc. Amer. 36:782; Wirth, 1952,

Proc. Ent. Soc. Washington 54:23; 1965, Proc. Ent. Soc. Washington 67:215. (Type-species: Parabezzia petiolata Malloch,

orig. desig.)

Diaphanobezzia Ingram and Macfie, 1931, Dipt. Patagonia & S. Chile,

pt. 2, p. 223. (Type-species: Diaphanobezzia pellucida Ingram

and Macfie, by orig, desig.).

DIAGNOSIS. Body rather stout, almost bare. Eyes bare. Palpus foursegmented with one to six long sensillae, first and second segments fused. Female antennal segments 3-10 oval, segments 11-15 cylindrical and elongated; male antennae plumose. Mesonotum without ary sign of pits or spine or tubercle, but bristles and small pubescence present. Legs slender with some bristles; coxae of male densely covered with bristles; tarsi sometimes with ventral spines; fourth segments of tarsi cordiform; fifth segments swollen and flattened on the lateral side, sometimes with tubercles on ventral side; without empodium; male claws small and equal but long, curved and may be unequal in female; no basal tooth. Wing of male narrower than female, macrotrichia absent, microtrichia very fine; costa very long, extended beyond a single radial cell, costa shorter in male; median fork petiole. Spermathecae two. Male genitalia, aedeagus large, triangular, parameres small, fused to form triangular sclerite behind aedeagus.

Approximately 18 species have been described.






85

Distributions North America, S6uth America, Republic of Congo, Jamaica, and Puerto Rico.

PUPA. Pupa with very short respiratory horn, oval shape, about five pairs of spiracles; caudal spines short and stout, pointed laterally. Adult habits: reared from sand bar (Wirth, 1965c)

Malloch (1915a) proposed the genus Parabezzia for three Nearctic species: P. petiolata n. sp. (male), Ceratopogon inermis Coquillett, and Bezzia elegantulata Johannsen. He used their petiolate media of wing as a distinguishing character.

It resembles Neurohelea in having long costa, extending beyond R5 but differs in having one radial cell and four segmented palps. Illustration: Fig. 60 Parabezzia petiolata Mall.


Genus Clinohelea Kieffer

Clinohelea Kieffer, 1917, Ann. Mus. Nat. Hung., 15:295, 316; 1919,

Ann. Mus. Nat. Hung., 17:76; Goetghebuer, 1920, Mem. Ms. Roy.

Hist. Nat. Belg., 8:76; Edwards, 1926, Trans. Ent. Soc. London,

74:413; Johannsen, 1931, Arch f. Hydrobiol., 9:435; Macfie,

1934, Ruwenzori Exp., 351:91; 1939, Rev. Ent. Rio de J., 10:205; Johannsen, 1943, Ann. Ent. Soc. Amer., 36:783; Lane, 1944, Rev.

Ent. Rio. de J., 15:249; Goetghebuer, 1948, Explor. Parc. Nat.

Albert. Miss. de Witte fasc. 55:17; Lee, 1948, Proc. Linn. Soc.

N.S.W., 73:64; Wirth, 1951, Proc. Ent. Soc. Wash., 53:321; 1952, Univ. Calif. Publ. Ent., 9:209; Lane & Durat, 1954,

Dusenia, 5:247; de Meillon, 1959, Novos Taxa Ent. no. 13:20

(Type-species: Ceratopogon variegatus Winnertz, orig. desig.)






86


DIAGNOSIS. Eyes bare, widely separated. Palpi simple. Female antennal segments 3-10 long, segments 11-15 cylindrical. Humeral pits absent, anterior spine very short. Femora slender, without spines; fourth tarsal segment of at least four posterior legs strongly bilobed and each lobe terminates in a stout spine, fifth tarsal segment of fore legs greatly swollen in both male and female; claws of four posterior legs very unequal in female. Wing long, without macrotrichia; costa long; two radial cells well developed; intercalary fork absent; media widely sessile. Gland rods absent. Spermathecae two. Male genitalia: ninth sternite short; ninth tergite tapered; basistyles slender; dististyles very long and slender; aedeagus arched, median portion large; parameres slender, rounded tip and sometimes fused at particular point.

Approximately 26 species have been described. Distribution: World-wide.

For the immature stages of Clinohelea, Wirth (1951d) published on the pupa. One species was collected from the sandy edge of a stream at Falls Church, Virginia.

PUPA. Light brown; long and slender respiratory organ with 10 spiracles at apex; operculum as long as broad, with a pair of round tubercles each with a long seta, surface especially near distal areas .and between the tubercles with shagreen body surface appearance with crescent-shaped tubercles, sharp and setose at lateral margins; anal segment long with fine shagreening, the apicolateral processes well developed with sclerotized tip, subapical shagreening.






87


This genus differs from Palpoimia and Tetrabezzia in the following characters: fifth tarsomeres of fore legs swollen in both sexes; fourth tarsomeres of the four posterior legs of female highly bilobed, each with a stout spine; claws of four posterior legs of female very unequal.

The other characters which readily differentiate them from

Tetrabezzia are: hind leg not unusually long; claws unequal only on

four posterior legs; wing with two radial cells. Illustration: Fig. 61 Clinohelea bimaculata (Loew)



Genus Tetrabezzia Kieffer

Tetrabezzia Kieffer, 1917, AnD. Ms. Hung., 15:296; Ingram and Macfie,

1923, Bull. Ent. Res., 14:70; Macfie, 1940, Ann. Trop. Med.

Parasit., 34:24; de Meillon, 1961, Rev. Ent. Mocambique, 4:55;

Wirth, 1962, Ann. Ent. Soc. Amer., 55:274. (Type-species:

Tetrabezzia spinimera Kieffer, mon.)

DIAGNOSIS. Large size, body slender and almost bare. Eyes bare, separated. Female antennae very long and slender with very long hairs on segments 3-10; palpi slender, third segment simple. Mesonotum with anterior spine. Legs long, hind legs longest, hind femur enlarged near tip, fourth tarsomeres with bifid spinose lobes on fore and mid legs, but long and cylindrical on hind legs; bifid lobes of male fourth tarsomeres not as distinct as in female; claws very unequal on female, almost subequal on male; fore femur armed ventrally. Abdomen petiolate, two spermathecae, gonopore without hair tufts. Wing infuscate, macrotrichia absent, microtrichia fine, costa almost





88


reaches the wing tip. Male genitalia with ninth sternite short, ninth tergite large, basistyles and dististyles long and slender, aedeagus arch, triangular shape, parameres club-tipped.

Approximately four species have been described. Distribution: Africa, Malaya, ThAiland.

This genus resembles Clinohelea but the chief points of difference are as follows: female hind leg unusually long; fourth tarsomere of four anterior legs with bifid spinose lobes; all female claws very unequal; wing with one radial cell; abdomen petiolate. Illustration: Fig. 62 Tetrabezzia pictipennis (K.)


Genus Heteronyia Say

HeteroMwia Say, 1825, Amer. Ent., 2:79; Kieffer, 1906, Gen. Insectorum, faso,., 42:64; Edwards, 1926, Trans. Ent. Soc. London

:420; 1935, Konowia, 12:87; Macfie, 1940, Ann. Trop. Mod. Parasit.,

34:27; Johannsen, 1943, Ann. Ent. Soc. Amer., 36:774; Lane,

1946, Rev. Ent. Rio de J., 17:211; Lee, 1948, Proc. Linn. Soc.
N.S.W., 73:61; Duret and Lane, 1955, Dusenia, 6:35; Lane, 1961,

Rev. Bra. Biol., 21:41, Wirth, 1962, Ann. Ent. Soc. Amer., 55:274.

(Type-species: Heteronyia fasciata Say, mon.).

Pacyleptus Walker, 1856, Insecta Saundersiana, Diptera. London :426. DIAGNOSIS. Large size, body slender with few hairs. Eyes bare, broadly separated. Female antennal segments 3-10 long, segments 11-15 very long and dark; male antennae with inconspicuous plumes. Palpi long and slender, third segment simple. Mesonotum dark; with small anterior spine. All legs different, femora of fore legs greatly swollen and armed ventrally with spines, fore tibiae arched along






89


the femora; mid legs long; hind*legs of female very long, tip of hind femora club-shaped; fourth tarsal segments on all legs of male and on the anterior four of female cordate, but long and slender on hind legs of female. Hind claws of female single, very long and barbed basally; the others including all male claws equal. Wing fasciate, with one or two radial cells; costa long. Male genitalia: ninth sternite short, ninth tergite tapering with well developed apicolateral process; basistyles and dististyles simple; aedeagus arched with large body; parameres club-tipped and lateral arms distinct.

Approximately 20 species have been described.

Distribution: North America, South America, Asia, and Australia.

The study of the immature stages was made by Thomsen (1937).

One pupa was found from wet algae on the cliffs near Ithaca, New.York. PUPA. Respiratory trumpet flat and short, constricted at base, apex bearing 11 spiracles; operculum with two setae and also with small hooks scattered on the surface; abdominal segments with three mid dorsal sclerotized areas, the surface beside these areas covered with small round hooks; last segments with two projections.

This genus is closely related to Palpowyia and some authors have misplaced them in Palponyia. It is differentiated from Palporgia by the following characters fore femur greatly swollen and armed ventrally with spines; fore tibia arcuate to the shape of the femur; hind leg of female very long with a single barbed claw; fourth tarsal segment on all legs of male and anterior four of female cordate, with equal claws; wing fasciate.

In Serromyia, hind femur is greatly swollen and armed and tibia also arcuate as in fore leg of Heteronvia.




Full Text

PAGE 1

AN ILLUSTRATED KEY FOR THE GENERA OF CERATOPOGONIDAE (DIPTERA) OF THE WORLD By NIPHAN CHANTHAWANICH RATANAWORABHAN A DISSERTATION PRESENTED TO THE GRADUATE COUNCIL OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY UNIVERSITY OF FLORIDA 1969

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ACKNOWLEDGMENTS The author expresses her deep gratitude to Dr. F. S. KLanton, chairman of her supervisory committee, for his help and encouragement in all phases of her graduate study. She wishes to express her most sincere appreciation to Dr. W. W. Wirth for all assistance, criticism and for providing the available specimens from the U. S. National Museum for this taxonomic study. For their criticism of the manuscript, appreciation is also extended to committee members: Dr. M. J. Westfall, Dr. L. C. Kuitert and Dr. D. H. Habeck. The author also expresses her thanks to the U. S, Army (project number D A-*f9-193-MD-2177) for sponsoring this program and finally, to her husband for his help, encouragement, cooperation and understanding during her graduate study.

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TABLE OF CONTENTS Page AC KNOWLEDGMENTS ii LIST OF FIGURES ^ INTRODUCTION 1 KEY FOR THE GENERA OF CERATOFO GONIDAE DESCRIPTIONS 1** Genus Leptoconops Skuse . . . . . Ik Key for the Subgenera of Leptoconops 19 Genus Atrichoporcon Kieffer 20 Key for the Subgenera of Atrichoporcon .... 25 Genus ForcinoTnyla Keigen 26 Key for the Subgenera of Forcipomyia ..... 37 Genus Dasyhelea Kieffer . 40 Key for the Subgenera of Dasyhelea h$ Genus Paradasyhelea Macfie 46 Genus Culicoides LatreiUe 47 Key for the Subgenera of Culicoides 58 Genus Austroconops Wirth and Lee ... 61 Genus C anotopter ohelea Wirth and Hubert 62 Genus Ceratopogon 1-eigen 63 Key for the Subgenera of Ceratopogon ..... 67 Genus Alluaudonyia Kieffer 68

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Monohelea . Genus Stilobezzia Kief f er Key for the Subgenera of Stilobezzia Genus Echinohelea Macfie Genus Monohelea Kieffer e Key for the Subgenera of Genus SerroTnvi,a Meigon . Genus Macrurohelea I. & M, Genus Parabezzia Malloch Genus Clinohelea Keiffer Genus Tetrabezzia Kieffer Genus Keteromyia Say Genus Poll's ici d prr/ia Macfie Genus Keurohelea Kieffer . Genus Johanns enotnyia Malloch Genus Xenohelea Kieffer . Genus Jenkinshelea Macfie Genus Probezzia Kieffer • « • • • • • • Genus Calyptoporcon Kieffer Genus Sphaerotnias Curtis Key for the Subgenera of Sphaeromias Genus Mall oc hohele a Wirth Genus Nilobezzia Kieffer Key for the Subgenera of Nilobezzia Genus Stenoxenus Coquillett .... Genus Paryphoconus Enderlein .... 70 75 76 77 80 81 83 84 85 87 88 90 91 93 95 97 98 100 102 105 106 107 110 111 112 iv

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I I Genus Pachyhelea *-JIrth Genus Pslponyla Meigen Key for the Subgenera of Palpowyia Genus Bezzia Keif f er Key for the Subgenera of Bezzia . . V> Genus Phaenobezzia Haeselbarth . . . . Genus A • Genus B Genus C Genus D Genus E Genus F Genus G Genus H Gems J Genus K Genus L Genus M Genus N Genus 0 Genus P Genus Q Genus R Genus S Genus T 113 115 118! 119 122 123 12k 125 126 126 12? 128 128 129 129 130 130 131 132 132 132 133 134 134 135

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Gems Dolichoheloa Edwards , . . Genus Acnnthohelea Kieffer ... Genus Luciaryia de Meillon . . . Genus Ceratobezzia Kief fer ... ALPHABETICAL LIST OF GENERA MID SUBGENERA LITERATURE CITED BIOGRAPHICAL SKETCH vi

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LIST OF FIGURES Fipore P a 2 e 1. Culicoidos furens (Poey) 140 2. Leptoconops ( Leptoconops ) stygius Skuse. ...... 141 3. Leptoconops ( Holoconops ) kerteszi K 142 4. Leptoconops ( Styloconops. ) albiventris (Meijere). . . 143 5. Leptoconops ( Styloconops ) splnosifrorts Carter. . . . 144 6. Atrichopogon (Atrichopogon) levis (Coq.) 145 7. Atrichopogon ( Psilokempia ) arcticus (Coq.) 146 8. Atrichopogon ( Heloehelea ) meloesugans K 147 9. Forcipomyia ( Forcipomyia ) bipunctata (L.) 148 10. Forcipomyia ( Prof orcipomyia ) -wirthi Saunders .... 149 11. Forcipomyia ( Warmkea ) aeria Saunders 150 1 1 12. Forcipomyia ( Warmkea ) tuberculata Saunders 15 1 13. Forcipomyia ( Calof orcipomyia ) glauca Edwards .... 152 14. Forcipomyia ( Metaf orcipomyia ) pluvialis Malloch . . 153 15. Forcipomyia ( Thyridomyia ) palustris Saunders .... 154 16. Forcipomyia ( Synthyridomyia ) colemani Wirth .... 155 17. Forcipomyia ( Trichohelea ) fijiensis (Macfie) .... 156 18. Forcipomyia ( Lasiohelea ) fairfaxensis Wirth .... 157 19. Forcipomyia ( Keof orcipomyia ) egues (Joh.) 158 20. Forcipomyia ( Dacnof orcipomyia ) anabaenae Chan & Saunders 159 21. Forcipomyia ( Phasmidohelea ) fulip:inosa (Mg.) .... 160 vii

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Page F;i£ure 22. Forcipomyia ( Phasmidohelea ) sp 23. Forcipomyia ( Lepidohelea ) annulatipes Macfie 162 24. > Forcipomyia ( Pterobosca ) fusicornis (Coq.) 163 25. Dasyhelea ( Dasyhelea ) grisea (Coq.) ^ 26. Dasyhelea ( Pseudoculicoides ) mutabilis (Coq.) .... 165 27. Dasyhelea ( Prokempia ) cincta (Coq.) 28. Paradasyhelea roinuta Wirth & Lee • ••'»•••_••• 29. Cvaicoides ( Cvaicoides ) yvtkonensis Hoffman iw 30. Cvaicoides ( Trithecoides ) flaviscutatus W. & H. . . . 169 31. Cvaicoides ( Haemophoructas ) gemellus Macfie 170 32. Cvaicoides ( Meijerehelea ) guttifer (Meijire) 171 33. Cvaicoides ( Hoff mania ) insignis Lutz 17 2 34. Cvaicoides ( Avaritia ) obsoletus (Mg.) 173 35. Cvaicoides ( Oecacta ) furens (Poey) *74 36. Cvaicoides ( Drymodesmyia ) copiosus R. & H. . . . • • 175 37. Cvaicoides ( Diphaeomyia ) baueri Hoffman 176 38. Cvaicoides ( Anilomyia) covagarciai Ortiz 177 39. Cvaicoides ( Beltranmyia ) crepuscvaaris Malloch. . • • 178 40. Cvaicoides ( Glaphiromyia ) scopus R. & H. 179 41. Culicoides ( Haematomyidivim ) paraensis (Goeldi). ... 180 42. Cvaicoides ( Macfiella ) phlebotomus (Will.) 181 43. Cvaicoides ( Hataemyia ) mojingaensis W. & B 182 2j4. CvOicoides ( Monocvaicoides ) variipennis (Coq.). ... 183 45. CvOicoides ( Selfia ) hieroglyphicus Malloch 184 46. Austroconops mcmillani Wirth & Lee 185 viii

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Fipnre 186 ( -.T ,_4. 0 ^ 0(Ton (Cer?to"DO' ,, on) culicoidi thorax Hoffrnan. . 187 188 50. 189 51. 190 52. 191 53. 192 rIt 193 55. 19^ 56. 195 57. 196 58. 197 59. 198 60. x ciTTcioczzicx pe (-fj_o_Lci uj. jl -lci-'— l * •••••• 199 01. 200 62. 201 63. • 202 1, 64. 203 65. 204 66. 205 67. 206 68. Jonkinshelea magniponnis (joh.)* ••«••••••• 207 69. 2 OS 70. 209 71. Sphaeromias (Snhaoromias) lonripenrds (Loew) .... 210 ix

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72. Sphaeroreias ( Honohelea ) barkudensis Edwards. .... 211 73. Hallochoheloa albibasis (Hall.) 212 i 7^. Nilobezzia schwarzii (Cog.) 213 75. Stenoreenus insigninervis Macfie 214 76. Paryphoconus anf^ustipennis Enderlein 215 77. Pachyhelea pachynera (Will.) 2l6 78. Palponyia plebeia (Loew) 217 79. Bezzia (Bezzia ) setulo-sa (Loev) 218 80. Bezzia ( Pseudobezzia ) flavitarsis (Kail.) 219 81. Phaenobezzia pistiae (I. & M.) 220 82. Genus A 221 83. Genus B 84. Genus C 223 85. Genus D 224 86. Genus D sp. 2 225 87. Genus E 226 88. Genus F sp. 1 227 89. Genus F.sp. 2 228 90. Genus G 229 91. Genus H 230 92. Genus J 231 93. Genus K 232 94. Genus L .' 233 95. Genus :M 234 96. Genus N 235 X i

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Figure 97. Genus 98. Genus p 99. Genus 100. Genus 101. Genus 102. Genus &3££ 236 237 238 239 2MO 24-1 3d

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INTRODUCTION Ceratopogonidae is a large family of minute flies, which was brought to the attention of taxonomists in 1852 by Winnertz in his monograph of the European species of the genus Ceratopogon . The contributions following Winnertz were those of Coquillett who published on American species from 1900 to 1905. He included all the species in Ceratopoeon. Kieffer (1899-1926) described several species and erected many genera. Ceratopogonidae was recognized as a subfamily of Chironomidae until Malloch (1917) finally elevated them as' belonging to a distinct family. Later Edwards ( 1926) published the diagnostic characters of many genera which are now being used as the basis for modern classification. Since 1925 Macfie has been monographing groups on a worldwide basis. The early publications on the genera of American ceratopogonids were published by Malloch (1915), Thomson (1935, 1937), Hoffman (1924-1939), Root and Hoffman (1937) and Johannsen (19^3). Recent ones are those of Wirth (1952, 1962a) and Wirth and Stone (1956). The important papers on European Ceratopogonidae are those of Kieffer (1925), Edwards (1926), Ooetghebuer (1920), Lindner (193 2 *) and Clastrier (1961). Elsewhere, Lutz (1912-1914) and Lane (19^1956) worked on the Neotropical genera, Carter, Ingram and Macfie (1920-1922),

PAGE 13

on the West African, de Meillon (1931) on the South African, Tokunaga (1939) on the Japanese and the Ficronesian (Tokunaga and Karachi, 1959), and Lee (19^8) and Skuse (1889) on the Australasian species. A key to the genera of Ceratopogonidae was published in 1926 by Xieffer who separated the family into 59 genera. Other keys have been made by many authors who mainly limited their keys to the genera and the species in a certain region. Edwards (1926) published the diagnostic characters of the genera occuring in Great Britain. Johannsen (1932) constructed a key based on the collection from Malayan Subregion with the additional genera which were recorded from the entire Oriental region. Goetghebuer (1933) published a key for the genera of the Palearctic region. Xacfie's key (19^0a) was considered to be the complete key to the known genera 0 f that time. It was a concise key which contained 63 genera but he failed to give the generic definitions, which made the key ineffective to taxonomists. The best key to North and South American genera, Johannsen (19^3), deluded the illustrations of many imoortant generic characters. Lee (19^8) published a key to the genera known to occur in the Australasian region. He divided the faraly into 10 groups, using Macfie's groupings (19^0a). He also presented a comparison of closely related genera. A key to some American genera was published in 1952 by Hirth, based on his studies of California ceratopogonids . A key to the North American genera vas also presented by Wirth and Stone (1956). Recent keys to the subfamilies, genera and subgenera of the Ceratopogonidae of the Vfo&A by Wirth (1959c, unpublished) and the genera of the tribe Sphaeromiinae

PAGE 14

(1962) are probably the most comprehensive ones. The latter provided some illustrations of the important characters and generic diagnoses were added. There has been a great deal of discrepancy among the current keys because of the confusion -which is almost always present with the key and the generic diagnosis. Many previously established genera were synonymized or sometimes reduced to subgeneric rank. It is very difficult to use available keys with this added confusion. Therefore a complete and comprehensive key is needed to facilitate the classification of ceratopogonids. An attempt is made in these studies to provide the diagnostic characters of each genus with detailed illustrations of its representative species. Whenever possible the type species was selected for study. Keys to the subgenera are also presented and the important references are provided. The following genera: Dolichohelea , Acanthohelea, Luciamyia and Ceratobezzia are not included in the key for the genera of Ceratopogon idae. The specimens of these genera are not represented in the collection. The information obtained is only that published by the authors and the fragmentary information was summarized in this report. The information on the immature stages, biology and habitats of any genus is included whenever it is available. The specimens for this study were obtained from the U. S. National Museum.

PAGE 15

KEY FOR THE GENERA OF CERATOPOGONIDAE 1. Female antenna 13-1^-segmented 2 Female antenna 15-segmented • 3 2. The r-m crossvein absent Leptoconops (Fig. 2) The r-m crossvein present Genus P (Fig. 98) 3. Female empodium large • ** Female empodium small or vestigial . . . 5 4. Radial cells not well developed, first small or obliterated, second small, wing with abundant flattened macrotrichia; body densely covered with hairs; male genitalia: ninth tergum short and tapered, setose apicolateral process present Forcipomyia (Fig. 9) Radial cells well developed, second one large, wing with moderate stiff macrotrichia; body hairs sparse; male genitalia: ninth tergum rounded, apicolateral process absent Atrichopoffon (Fig. 6) 5. Antenna! segments sculptured Dasyhelea (Fig. 25) Antenna! segments not sculptured . 6 6. Radial cells both absent 7 Radial cells present, at least one 10

PAGE 16

7. Palpi five-segmented Genus M (Fig. 95) Palpi two-four-segmented ..8 8. Palpi four-segmented; wing not broad; male antenna 14-segmented Paradasyhelea (Fig. 28) Palpi two-segmented; wing broad; male antenna eight-segmented ...9 9. Posterior branch of media narrowly interrupted at base, cell Mj. large, costa extended about half of wing length Genus Q (Fig. 99) Posterior branch of media broadly interrupted, cell M^ small, costa short extended about onethird of wing length Genus S (Fig. 101) 10. Media petiolate , usually forking behind r-m crossvein ... .11 Media sessile, forking at or before r-m crossvein ...... 24 11. Claws small, equal and simple in both sexes; one or two radial cells almost equal in size 12 Claws of female usually large; one or two radial cells, second one always much longer than the first 14 12. Palpi five-segmented Culicoides (Fig. 29) Palpi three-four-segmented • 13 13. Palpi three-segmented; one radial cell, lower branch of media absent; one spermatheca Genus N (Fig. 96) Palpi four-segmented; two medial cells, long and broad, lower branch of media present; two spermathecae Austroconops (Fig. 46)

PAGE 17

14. Second radial cell not or little longer than the first, one or both cells may be obliterated (in Alluaudomyia first one usually obliterated) costa about half of wing length 15 Second radial cell much longer than the first, costa extended to wing tip or two-thirds of wing length .... 17 15. Wing with one radial cell, usually with 1-20 small dark spots and sometimes with grayish streaks along the wing veins; two claws on each leg . Alluaudomyia (Fig. 50>51) Wing usually with two radial cells, usually without dark spots; one or two claws on each leg 16 16. All claws single, stout and slightly curved; palpi twosegmented; wing veins with only cubital stem and 1A present, Cu^ indistinct, microtrichia spinelike * Camotopterohelea (Fig. 47) Two claws on each leg, equal or subequal; palpi fivesegmented; wing with lower branch of media often absent or interrupted, without microtrichia . . Ceratopofton (Fig. 48) 17. Costa extending beyond vein R,-, one or two radial cells; claws usually equal 18 Costa not extending beyond vein R^* two radial cells present, claws equal or unequal 20 18. Two radial cells, first one small; palpi threesegmented Genus 0 (Fig. 97) One radial cell, palpi four-segmented 19

PAGE 18

7 19. First tarsomere of mid leg much longer than that of the other legs and bearing spines; Coxae of male with few hairs; male genitalia: aedeagus arched, paramere absent Genus R (Fig. 100 ) First tarsomere of mid leg simple; coxae of male denselycovered with bristles; male genitalia: aedeagus large, paramere small and fused to form a triangular sclerite behind aedeagus Parabezzia (Fig. 60) 20. Hind femur simple, armed or unarmed, tibia not curved .... 21 Hind femur swollen in both sexes, armed ventrally with numerous spines, tibia curved basally . . Serromyia (Fig. 58) 21. Female abdomen simple 22 Female abdomen highly modified with tenth segment curved forward and reaching the ventral side of anterior segments .. Macrurohelea (Fig. 59) 22. Male antenna plumose; femora and tibiae unarmed; last antennal segment simple 23 Male antenna without plumes; almost alt , femora and tibia armed with long spines; last antennal segment usually shorter than the prece ding four segments Echinohelea (Fig. 55) 23. Female claws all unequal; first tarsomere of mid leg bearing subbasal spur; male genitalia: ninth tergite rounded, aedeagus reduced to a pair of oblique lateral sclerites, pararaeres with modified, well developed submedian process Stilobezzia (Fig. 52)

PAGE 19

8 Female anterior four claws equal, hind claw with one long and with or without another short one; male genitalia: bilobed or truncate ninth tergum, aedeagus usually with a pair of lateral sclerites and additional dorsomedian lobe, parameres fused or paired Monohelea (Fig. 56) 24. Female fifth tarsomere with numerous ventral blunt spines or batonnets 25 Female fifth tarsomere with or without sharp batonnets ... 37 25. Female claws highly unequal; wing with two radial cells ... 26 All female claws equal with internal or external basal tooth; one or two radial cells 27 26. All female claws unequal; batonnets appear on basal half of fifth tarsomere Xenohelea (Fig. 67) Claws highly unequal only on four posterior legs, each with external basal tooth; fifth tarsomere with numerous batonnets along the segment .... J ohannse nomyia (Fig. 66) 27. Female abdomen with ventral sclerotized plate on seventh segment ......... 28 Female abdomen without sclerotized plate on seventh segment . . 29 28. Wing with two radial cells; anterior part of thorax cone-shaped Genus H (Fig. 91) Wing with one radial cell; anterior part of thorax rounded Genus K (Fig. 93)

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9 29. Hind tarsus about three times longer than mid tarsus Calyptopogon (Fig. 70) Hind tarsus simple 30 30. Wing of female with usually broad anal lobe . Jenkins helea (Fig. 68) Wing with anal lobe narrow and simple 31 31. Female claws each with internal basal tooth (Genus J: claws with internal tooth in one and external tooth in the other claw); ventral hair tufts absent 3 2 Female claws each with external basal tooth; abdomen with ventral hair tufts 3^ 32. One spermatheca; anterior tubercle well developed Genus A (Fig. 82) Two spermathecae; anterior tubercle poorly developed or absent .33 33. One radial cell; claws each with internal tooth in one and external tooth in the other; leg unarmed . • Genus J (Fig. 92) Two radial cells; claws each with internal basal tooth; legs with numerous spines on ventral side of femora and bristles on dorsal side of tibiae ... Sphaeromias (Fig. 71) 3^. Legs armed with ventral spines on femora and dorsal spines on tibiae; male genitalia: highly modified, with basistyle poorly developed and fused to form a cone-shaped process Nilobezzia (Fig. 7*0 Leg with femora and tibiae unarmed 35

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10 35. Wing with two radial cells Mallochohelea (Fig. 73) Wing viith one radial cell 36 36. Costa very long reaching the wing tip; fourth tarsomere bilobed; spermatheca small; mandible vithout serrate margin; male genitalia: parameres fused mesally, tip slender and separated Frobezzia (Fig. 69) Costa short, not reaching the wing tip; fourth tarsomere oval; spermathecae large; mandible with serrate margin; male genitalia: parameres separated then fused again at the tip Genus T (Fig. 102) 37. Fifth tarsomere at least on one leg with one to five ventral sharp spines, or abdomen cone-shaped 38 Fifth tarsomere without ventral sharp spines . . 44 38. Abdomen cone-shaped 39 Abdomen simple, not cone-shaped ... 40 39. Fifth tarsomere with about three ventral spines on distal half; gland rods on seventh abdominal segment only Pachyhelea (Fig. 77) Fifth tarsomere without ventral sharp spines; gland rods present on segments five, six, and seven .Genus D (Fig. 85,86) 4-0. Female abdomen with gland rods • 41 Female abdomen without gland rods 43

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11 41. Femora armed with ventral spines; first hind tarsonere longest ^nus B ( Fi S' 8 3) Femora -unarmed; first hind tarsomere simple ^2 42. One radial cell; tarsal segments tiro to five rather short; all femora rather swollen Genus L (Fig. 94) Two radial cells; tarsal segments two to five simple; four anterior femora slender, hind femora enlarged Genus E (Fig. 8?) 43. One spermatheca, large, pear-shaped; claws small; hind femora slender; fifth tarsomere stout with one to two sharp spines; two radial cells; mandible with five large teeth Genus G (Fig. 90) Two spermathecae; claws large especially of hind legs, each with inner basal tooth; hind femora enlarged and curved; fifth tarsomere slender, sometimes with spines; mandible with about 11 teeth Genus F (Fig. 88,89) 44. Hind tarsus of female about three or more times longer than midtarsus; claws highly unequal, single long hind claws with barb 45 Hind tarsus of female simple; all claws equal, if unequal without any single claws 47 45. Femora of fore legs greatly swollen and armed ventrally with spines; fore tibiae arched along femora . Heteromyia (Fig. 63) Femora of fore legs simple; fore tibiae not arched 46

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12 46. Fourth tarsoneres with bifid spinose lobes on fore and mid legs but long and cylindrical on hind legs; fore femora armed ventrally; abdomen petiolate; wing fasciate; fifth tarsomere of fore leg simple . . . Tetrabezzia (Fig. 62) Fourth tarsomeres cordiform to transverse on fore and mid legs but elongated on hind legs; fore femora unarmed; abdomen slightly curved downward; wing not fasciate; fifth tarsomere of fore leg swollen Pellucidomyia (Fig. 64) 47. Fourth tarsomeres of at least four posterior legs strongly bilobed and each lobe terminates in a stout spine Clinohelea (Fig. 61) Fourth tarsomeres cordate or cylindrical 48 48. Antennal segments with stiff hairs; radial cell narrow ... 49 Antennal segments without stiff hairs; radial cell simple . . 50 49. Vein M£ elbowed in female; female antennal segments three to ten with stiff hairs; palpi four-segmented; fourth tarsomere spoon-shaped to subcylindrical; last three tarsomeres simple; male genitalia: parameres fused with broad rounded tip; two spermathecae . . . Stenoxenus (Fig. 75) Vein not elbowed; female antennal segments 1115 with stiff hairs; palpi five-segmented; fourth tarsomere cordate to subcylindrical; last three tarsomeres relatively short; male genitalia: parameres separated; one spermatheca Paryphoconus (Fig. 76)

PAGE 24

13 50. Costa extended beyond R^; two radial cells; femora unarmed Neurohelea (Fig. 65) Costa not extended beyond R^; one or two radial cells; femora usually armed, at least on one leg 51 51. One radial cell 52 Two radial cell 53 52. Male genitalia: basistyle and dististyle reduced but distinctly separated Phaenobezzia (Fig. 81) Male genitalia: basistyle and dististyle well developed Bezzia (Fig. 79) 53. Hind claws longer than the claws on the first four legs; fourth tarsomere oval to enlongated; gland rods absent Genus C (Fig. 8*0 All Slaws equal; fourth tarsomere bilobed or cordiform; gland rods always present Palpomyia (Fig. 78)

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DESCRIPTIONS Genus Leptoconops Skuse Leptoconops Skuse, 1890, Proc. Linn. Soc. N. S. Wales, 4:288; Johannsen, 1905, Bull. N. T. State Mas., 86:94; Kieffer, 1906, Gen. Insectorum, fasc. 42:48; 1908, Ann. Mas. Nat. Hung., 6:577; 1921, Arch. Inst. Pasteur Afr. Nord, 1:107; Carter, 1921, Bull. Ent. Res., 12:1; Johannsen, 1943, Ann. Ent. Soc. Amer., 3^:777; Lee, 1948, Proc. Linn. Soc, N. W. W. 72:332-8; Wirth, 1952, Uhiv. Calif. Pubis. Ent., 9:109; Smee, I966, Aust. J. Zool., 14:993; Chantha-wanich and Delfinado, I967, J. Med. Ent., 4:294 (Typespecies: Leptoconops stylus Skuse, Mon). Tersesthes Towns end, 1893, Psyche, 6:370 (Type-species: Tersesthes torrens Townsend, mon.). Centrotypus Grassi, 1901, Studi di Zool. sulla Mai. Roma, p. 118 (nomen nudum). Mycterotypus Noe, 1905, Rend, atti Accad. Lincei, 14:114 (Type-species: Mycterotypus bezzii Noe« desig. Carter, 1921). Mycteronyia Lutz, 1912 (not Philippi, I865), Mem. Inst. Oswaldo Cruz, 4:24 (lapsus for Mycterotypus Noe). Holoconops Kieffer, 1918, Ann. Mus. Nat. Rung., 16:135 (Type-species: Leptoconops kerteszi Kieffer, orig. desig. ) . 14

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15 Schizoconops Kleffer, 1918, Ann. Mis. Nat. Hung., 16:135 (Type-species: Schizoconops indicus Kieffer, orig. desig.). Styloconops Kleffer, 1921, Arch. Inst. Pasteur Afr. Nord, 1:107 (Type-species: Leptoconops albiventris de Meijere, orig. desig.). Protersesthes Kieffer, 1921, Arch. Inst. Pasteur Afr. Nord, 1:107 (Type-species: Tersesthes brasiliensis Lutz, orig. desig.). Microconops Kieffer, 1921, Arch. Inst. Pasteur Afr. Nord, 1:108 (Type species: Microconops vexans Kieffer, orig. desig.). Acanthoconops Carter, 1921, Bull. Ent. Res., 12:24 (Type-species: Acanthoconops spinosifrons Carter, orig. desig.). DIAGNOSIS. Eyes bare, widely separated in both sexes; vertex with bristles; frons and clypeus bare or hairy. Mandible about 16 teeth. Palpi "f our-segmented" ; distal two segments fused; male palpi longer than in female. Female antennae 13or i^-segmented, the flagella segments subspherical or oval, each with few hairs and transparent sensory spines, the terminal segment elongate. Male antennae 15segmented, plumose. Thorax arched anteriorly; humeral pits present. Wings usually milky white; R separate at base, fused with costa about mid wing forming a stigma; crossvein absent; intercalary vein above vein M^ conspicuous. Legs slender, hind legs longest; femora unarmed, tibiae each with stout apical spur; hind tibial comb five to seven; basitarsus with prominent ventral spines. Tarsal claws with small bristle or toothed; empodium small and branched. Abdomen with ninth segment bearing long or short lamellae or cerci, two well developed spermathecae. Male genitalia conspicuous; basistyles

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16 •with subbasal ventromesal lobe; dististyles short, small and tapering; aedeagus simple; parameres usually stout, heavily sclerotized. Approximately 62 species have been described. Distribition: World-vide. Keys to the larval and pupal stages of North American species are presented by Wirth and Stone (1956) and Thomsen (1937). The immature stages of the Bodega black gnat were described in detail by Smith and Lowe (19^). EGG. The egg is unsculptured, usually banana-shaped, blunt at one end and narrower at the other, slightly curved and circular in cross section. LARVA. The head is partly sclerotized, inside the head and first thoracic region is a system of heavily chitinized rods connected with the mandibular processes. Labrum scoop-like. Antennae at the extreme tip of the head are protrusible. Abdominal segments each with an intercalary segment, giving the appearance of 23 segments in larva. PUPA. Respiratory tube two-segmented, bearing 15 spiracles apically. The head is rugose. Abdominal segments covered with sharp recurved spines, located on large tubercles. Last segment is forked, without spines or tubercles. Adult habits and larval habitats: adult feeds on vertebrate blood, female may feed only once or as many as four times, males do not feed. Carter (1921) states that the adults are most active during the hottest parts of the day. Larvae are found in damp soil and along the seacoast in sand with organic matter. The genus Leptoconops was erected by Skuse in 1889 for Leptoconops stygius Skuse. In 1893» Townsend set up the new genus Tersesthes and Noe (1905) proposed the genus Ifycterotypus , which are now placed as

PAGE 28

17 synonyms of Leptoconops . In 1918 and 1921 the genus was divided into five new genera by Kieffer, these were: Eoloconops , Schizoconops, Styloconops, Proteresethes and Microconops . Carter (1921) retained only the genus Leptoconops, with two subgenera, Leptoconops s.s. and Holoconops , and also erecting a new genus, Acanthoconops . Acanthoconops was synonymized under Styloconops Kieffer by Kieffer (1921a) and it was reduced to a subgenus by Wirth (1952). Three subgenera of Leptoconops are retained: Leptoconops , Holoconops and Styloconops . This genus is different from most of the ceratopogonids and readily recognized by the following characters: Palpus four-segmented. Antennae of female with 13 or 1Usegments. Wings milky white; crossvein r-m absent; intercalary vein above vein M, prominent. Legs, basi tarsi usually armed with ventral spines. Lamellae or cerci of female usually prominent. Male genitalia, basistyle with subbasal lobe; dististyle with terminal appendage. Subgenus Leptoconops s. str. Subgenus type: Leptoconops stygius Skuse, mon. Distinguished from the others by having toothed tarsal claws; sensory area of the third palpal segment large occupying almost the entire length. Female antennae l^segmented. The lamellae of female long slender. Illustration: Fig. 2 Leptoconops (Leptoconops) stygius Skuse.

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Subgenus Holoconops Kieffer, 1918 Subgenus type: Leptoconops kerteszi Kieffer, orig. des. This subgenus is characterized by having simple tarsal claws and subapical pit on third palpal segment. Female antennae 13-segmented. Female genital i a with moderately long, broad lamellae. Kale genitalia with large, sclerotized parameres having lateral projections; dististyle bifid. Illustration: Fig. 3 Leptoconops ( Holoconops ) kerteszi K. Subgenus Styloconops Kieffer, 1921 Subgenus type: Leptoconops albiventris de Meijere, orig. des. This subgenus is distinguished from the others by the following characters: Head with numerous spines or short bristles on frons and vertex; female antennae ^segmented; palpus with sensory pit centrally or subbasally located. Tarsal claws with small basal tooth. Female genitalia with very short lamellae; parameres of male varied; the aedeagus fairly large. Illustrations: Fig, 4 Leptoconops ( Styloconops ) albiventris (Keijere) and Fig. 5 Leptoconops ( Styloconops) spinosifrons Carter.

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19 Key for the Subgenera of Leptoconops 1. Female antenna with 13 segments Holoconops (Fig. 3) Female antenna with 14 segments ......... 2 2. Female genitalia with lamella very long and slender; head with frons bare, few bristles on vertex; palpal sensory area occupying almost entire length of the third segment (Fig. 2B) Leptoconops (Fig. 2) Female genitalia with lamella very short (Fig. 5H); frons and vertex with numerous spines or short bristles (Fig. 4C, 5C); palpus with sensory pit subbasally or centrally located (Fig. 4A, 5 A ) • • Styloconops (Fig. 4,5)

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Genus Atrichopogon Kief f er Atrichopogon Kieffer, 1906, Ann. Soc. Sci. Brux., 30: ?; 1906, Gen. Insectorum, fasc. 42:53; Coquillett, 1910, Proc. U. S. Nat. Mas., 37:512; Kieffer, 1913, Pec. Ind. Mas., 9:1?4; 1919, Bull. Soc. Ent. France, p. 193; 1919, Ann. Mus. Nat. Hung., 17:20; Goetghebuer, 1920, Mam. Mus. Roy. Hist. Nat. Belg., 8:29; Carter, Ingram, and Macfie, 1921, Ann. Trop Mad. & Parasit., 15:321; Kieffer, 1924, Arch. Inst. Pasteur d* Algerie, 2:39^; 1924, Bull. Soc. R. Ent. Egypto, p248; 1925, Faune de France, fasc. 11, p. 51; Edwards, 1926, Trans. Ent. Soc. London, 74:399; Johannsen, 1931, Arch. f. Hydrobiol., 9:4l4; Ingram and Macfie, 1931, Dipt. Pat. & S. Chile, pt. 2, fasc. 4, p. 172; Macfie, 1932, Ann. Trop. Med. Parasit., 26:31; 1934, Ann. Trop. Mad. & Parasit., 28:184; 1939, Rev. Ent., 10:173; Tokunaga, 1940, Tenthredo, 3:109; Macfie, 1940, Ann. Trop. Med. & Parasit., 34:15; Zilahi-Sebess, 1940, Folia. Ent. Hungar. 5:34; Johannsen, 1943, Ann. Ent. Soc. Amer., 36:775; Macfie, 1947, Proc. R. Ent. Soc. Lon. (B), 16:72; Wirth, 1956, Proc. Ent. See. Wash.,, 58:16; Ewen and Saunders, 1958, Canad. J. Zool., 36:719; Tokunaga, 1959, Pacif. Ins., 1:179; Tokunaga and Murachi, 1959, Ins. Micron., 12(3) :ll6; Reran, 1959, Ent. Obozr., 38:683; Reran, 1961, Ent. Obozr., 40:925; Tokunaga, 1962, Pacif. Ins., 4:154. (Type-species: Ceratopogon levls Coquillett as exiles Coquillett, by desig. CoquiUett, 1910). Didymophleps Weyenbergh, I863, Stett, Ent. Zeit., 44:108 (Typespecies : Didymophleps hortorum Weyenbergh, mon. ) .

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21 Kempia Kieffer, 1913, Rec. Ind. Mas., 9:1^2 (As subgenus of Dasyhelea ) (Type-species: Dasyhelea ( Kempia ) calcuttensis Kieffer, by ? desig. ) . Gymnohelea Kieffer, 1921, Arch. Inst. Pasteur Afr. Nord, 1:115 ( Type-species : ? ) . Lophomyidium Cordero, 1929, An. Mus. Hist. Nat. Montevideo, 3:9^ (Type-species: Lophomyidium uruguayense Cordero, mon.). Psilokempia E nderlein, Tierwelt Mitteleur., 6(X7I):^ (Type-species: Kempia appendiculata Goetghebuer , mon. ) . DIAGNOSIS. Body stout, moderately hairy. Eyes pubescent or bare. Antennae 15-segmented, segments 4-10 globular or transverse, last five segments elongate, usually last segment terminates with nipple or style. Mesonotum "bare or -with few hairs. Legs slender; fore femora setose anteriorly. Wings moderately covered with macrotrichia, finer than in Forcipomyia and always lesser in male; costa long, about two-thirds of wing length, second radial cell well developed; intercalary fork distinct; median fork with short stem. Female abdomen usually bearing abdominal armature on sixth or seventh steral processes absent; aedeagus large, usually trilobed; paratneres small, usually hook-shaped, or absent. Approximately 296 species have been described. Distribution: World-wide. The biology and immature stages of Atrichopogon were studied by Johannsen (1952) and Ewen and Saunders (1958). The life history and

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22 all stages of Atrichopopion levis -were described in Boesel and Snyder ( 1944) . Keys -were provided by Thomson ( 1937) and Wirth and Stone (1956) for North American species. EGG. Atrichopogon levis egg is smooth, subshining, black; elongate and slightly curved. Most of the species lay eggs in masses, in aquatic situations. LARVA. Body usually appears flattened as the sides of the segments often protrude into lateral processes. Head hypognathous , antennae spur-like. Head capsule commonly bears eleven pairs of setae; abdominal segments one to seven with six pairs of setae which is similar to metathorax. The mesothorax has two lateral setae and three on the prothorax. Prothorax with bilobed, pseudopod bearing backward-curved booklets arranged in two transverse rows. The ninth abdominal segment is cylindrical, covered dors ally by an unpaired sclerite. Posterior segment with a pair of protrusible anal papillae and the anal pseudopod. PUPA. Body broad anteriorly and tapering posteriorly. Respiratory horn variable in shape, appearing boot-shaped, flattened, or cylindrical. Thorax with six pairs of cuticular processes on the dorsal side. Posterior median point of mesothorax never overlaps the first abdominal segment. Abdomen with branched or setaceous projections on first five segments. Segment nine terminates with a pair of diverging and tapering processes and covered with small stout spinules directed forward. Male genital sheaths are ventral and protrusible. Adult habits and larval habitats: Adult habits of Atrichopor;on levis are given by Boesel and Snyder (1944). Males and females have been found flying in broad daylight above moist mud covered with algae.

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23 Some were collected along streams and stagnant water. The species is abundant in the short grasses of parks, lawns or roadsides. Males •usually s-warm during sunset or other times of equivalent light. Larvae occur chiefly in shaded localities where mold, bacteria and algae are present in areas along the streams covering with algae, or wood and soil covering -with moss. Most species are aquatic. The adults of this genus are difficult to identify or even to separate distinctly into subgenera. As pointed out by Edwards (1926), Macfie (1939c), Nielsen (195D and Wirth (1952, 1956), the adults offer no good, clear-cut character or combination of characters. The male genitalia might be very useful if there were no variation in appearance caused by protrusion or retraction. The form of the ventral armature of the female abdomen and the spermathecae might make a more satisfactory key, if an intensive study was made of the above characters. Even and Saunders (1958) based their taxonomy mostly on the immature stages, -which show excellent and accurate separations. Atrichopogon is a well defined genus. It resembles Forcipomyia by having a large empodium, but can be distinguished from Forcipomyia adults by the following characters: second radial cell long, macrotrichia few or absent, microtrichia conspicuous and the nature of the fringe on the posterior margin of wing. Subgenus Atrichopogon s. str. Proboscis normal, straight. Eyes separated. Female antenna, segments 3-10 oval, segments 1115 cylindrical and elongated. Spormatheca one. Illustration: Fig. 6 Atrichopogon ( Atrichopogon ) levis (Coq.)

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2h Subgenus Psilokempia Enderlein, 1936 Subgenus type: Kempia appendiculata End, (orig. desig.) Enderlein (1936) erected genus Psilokempia for Kempia appendiculata End. It was reduced to subgenus by Remm (1959). This subgenus is distinguished from the others by the f ollowing points: proboscis not curved, antennae with last five segments together about as long as segments 3-10 combined, palpi stout; macrotrichia of wing few or absent; one spermatheca; does not parasitize insects. Illustration: Fig. 7 Atrichopogon ( Psilokempia ) arcticus (Co<*) Subgenus Meloehelea Wirth, 195 6 Subgenus type: Atrichopogon meloesugans Kieffer (orig. desig.) Wirth (1956c) erected this subgenus based on specimens that attack the bettles of family Meloidae. This group is characterized by having an unusually upcurved proboscis, palpi slender, antennae segments 11-15 together about twice as long as segments 3-10 combined; two spermathecae ; macro trichia abundant on wing; hind basi tarsus about 2.5 times as long as the second segment. Illustration: Fig. 8 Atrichopogon ( Msloehelea ) roeloesugans K.

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Key for the Subgenera of Atrichooogon Proboscis upcurved; two spermathecae (Fig. 8L) . Meloehelea (Fig. 8) Proboscis straight; one spermatheca 2 Antennal segments 3-10 rounded, 11-15 subspherical (Fig. 7A); female palpi with short segments (Fig. 7C); eyes contiguous (Fig. 7F) Psilokempia (Fig. 7) Antennal segments 3-10 oval, 11-15 cylindrical and elongated (Fig. 6A); female palpi with slender segments (Fig. 6C); eyes separated (Fig. 6F) Atrichopogon (Fig. 6)

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26 Genus Forcipomyia Meigen Forcipomyia (Megerle MS) Meigen, 1818, Syst. Beschr. Eur. Zweifl. Ins., 1:59; Kieffer, 1906, Gen. Insectorum, fasc, 42:52; 1913, Rec, Ind, Mus., 9:166; Malloch, 1915, Bull. 111. State Lab. Nat. Hist., 10:311; Ingram and Macfie, 1924, Ann. Trop. Med. & Parasit. 18:541; Edwards, 1926, Trans. Ent. Soc. London, 74:393; Johannsen, 1931, Arch. f. Hydrobiol., 9:407; Ingrain and Macfie, 1931, Dipt. Pat. & S. Chile, pt. 2, fasc. 4, pp. 156,177; Macfie, 1932, Tijdschr, Ent., 75:279; 1934, Ann. Trop. Med. & Parasit., 28:178; Goetghebuer, 1935, Rev. Zool. Bot. Afr., 27:145; Macfie, 1939, Rev. Ent., 10:142; 1940, Ann. Trop. Med. & Parasit., 34: 16; Tokunaga, 1940, Tenthredo, 3:6l; Zilahi-Sebess, 1940, Folia. Ent. Hungar. 5*21; . Johannsen, 1943, Ann. Ent. Soc. Amer. , 36:777; Harant and Huttel •. & N. , 1951, Ann* Parasit. Hum. Comp. , 26:471; Saunders, 1956, Canad. J. Zool., 34:657; Tokunaga and Murachi, 1959, Ins. Micron., 12 (3): 144; Tokunaga, 1959, Pacif. Ins., 1:256; Dessart, 196l, Bull. Ann. Soc. R. Ent. Belg., 97: 315; Remm, 1962, Loodusuur. Seltsi Aastar., 54:168; Tokunaga, 1962, Pacif. Ins., 4:169; Dessart, I963, Mem. Inst. Sci. Nat. Belg. (2) fasc. 71: 13* (Type-species: Ceratopogon ambiguus Meigen = Tipula bipunctata Linnaeus by desig. Coquillett, 1910) Tipula (part) Linnaeus, I767, Syst. Nat., ed. 12, 2:978. Ceratopogon (part) Meigen, 1804, KLass . , 1:31. Labidomyia Stephens, 1829, Cat. Brit. Ins., 2:239 (Type -species: Tipula bipunctata Linnaeus, by desig. Westwood, 1840).

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Tetraphora Philippi, 1865, Verh. Zool.-Bot. Ges. Wien, 15:630 (Typespecies: Tetraphora fusca Philippi, mon.). Prohelea Kieffer, 1912, Spolia Zeylanica, 8:1 (Type-species: ?). Euforcipomyia Malloch, 1915, Bull. 111. State Lab. Nat. Hist., 11:312 (Type-species: Euforcipomyia hirtipennis Malloch, by orig. desig.) Lepidohelea Kieffer, 1917, Ann. Mis. Nat. Hung., 15:364 (Type-species: Ceratopogon chrysolophus Kieffer, mon. ) . Microhelea Kieffer, 1917, Ann. Mas. Nat. Hung., 15:364 (Type-species: Atrichopogon tropicus Kieffer, mon.). Apelma Kieffer, 19 19 (not Billberg, 1820), Ann. Mas. Nat. Hung., 17:64 (Type-species: Apelma auronitens Kieffer, ? desig. by Macfie, 1940). Trichohelea Goetghebuer, 1920, Mem. Mas. Roy. Hist. Nat. Belg., 8:18 (Type-species: Trichohelea tonnoiri Goetghebuer, mon.). Thyridomyia Saunders, 1925, Parasit. , 17:268 (Type-species: Thyridomyia palustris Saunders, orig. desig.). Phasmidohelea Mayer, 1937, Arb. Morph. Tax. Ent. Berlin-Dahlem, 4:233 (Type-species: Phasmidohelea crudelis Mayer, orig. desig.). DIAGNOSIS; Body stout, densely covered -with hairs. Eyes bare. Antennae 15-segmented, segments 3-10 pear-shaped to vase-shaped in female,, with dense verticils, segments 1115 usually cylindrical to elongate, last segment often ending in a nipple; in male, segments 12-14 swollen at base, segment 12 very long, and segments 12-15 each with a single verticil. Humeral pits absent, mesonotum densely covered with hair and a few bristles. All legs and abdomen, especially

PAGE 39

in male, vith long bristles; femora large, empodium well developed, usually as long as claws (absent in male of Apelma = Trlchohelea) , Wings with flattened macrotrichia abundant; costa ending about the middle of wing; first radial cell small or almost obliterated, second cell small but distinct; intercalary fork present; median fork with short stem, usually faint. Male genitalia not inverted; aedeagus large, shield-shaped; parameres usually present, slender and tapering, often fused at base or connected by transverse rod or plate. Approximately 507 species have been described. Distribution: World-wide. Biology and early stages of Forcipomyia have been studied in detail by quite a number of taxonomists. In 1923, Frew described the larval and pupal stage of F. piceust Saunders (1924, I925) published the most extensive papers on the immature stages of Apelma and Thyridomyia ; de Meillon (1931) on one species from Transvaal; Mayer (1933) on F. corals; Tokunaga (193*0 on F. crinume ; Lane (I94?a) on F. argenteola and F. inornatipennis . Saunders (1956) had done the most extensive study in distinguishing the subgenera of Forcipomyia using characters of all stages. Recent papers on the immature stages and biology of Forcipomyia were published by Saunders ( 1959) on some species of Proforcipomyia and Thyridomyia. Saunders (1963) on subgenus Laslohelea t Chan and Saunders (1965) on F. ( Dacnof orcipomyia ) anabaenae ; and Chan and Le Roux (I965) on F. ( Neof orcipomyia ) eques and F. saundersl . Keys to early stages of North American species were presented by Thomson (1937) and Wirth and Stone (I956).

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EGO. Mayer (193*0 stated that eggs of F. plcea are found in irregular masses and those of F. pallida in garlands, LARVA. The larvae of this genus exhibit a considerable range of variation of structure throughout the subgenera. Most of them have elongate body, distinctly segmented, each segment as long as wide and bearing either dome-shaped projections ending in hooks, or spearshaped, or fringed setae. Head hypognathous ; they have eversible pro thoracic and anal pseudopods. PUPA. The pupa are less variable and may be confused with Atrichopogon. The posterior three segments remain tightly attached to the larval exuviae. Respiratory trumpets short, knob-like, bearing spiracles only at apex. Adult habits and larval habitats: Saunders (1959) found many species of the subgenera Proforcipomyia and Thyridomyia as pollinators of cacao. Chan and Saunders (1965) stated that F. (D.) anabaenae females are diurnal man-biters, prefering the shaded parts of the body. The females of Pterobosca group have specialized habit as ectoparasite of dragonflies. Females of eques (Johannsen) and related species suck the blood from the wing veins of lacewing flies, butterflies and other insects (Wirth and Stone, 1956). Larvae are mostly terrestrial, the gregarious larvae breeding under tree bark, in manure or moist decaying vegetable matter, in ant nests, and the like. F. calcarata (Coquillett) larvae were found along the rocky margin of a warm mineral spring at Alum Rocky Park in Santa Clara County, California (Wirth, 1952).

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30 This genus is very large and shows a vide range of variation among the species. Many attempts have been made to subdivide the genus into subgenera. Saunders' system seems to be the most acceptable. He set up eight subgenera based on larval and pupal characters in addition to adult characters (Saunders, 1956). Tokunaga and Kurachi (1959) mentioned using the following main characters to separate the subgenera: structure of male hypopygium; number of spermathecae; development of female antennae; value of hind tarsal ratio; segmentation of maxillary palp; wing venation, and development of scales and colored markings. Members of this genus are readily recognized by having abundant macrotrichia on wing, radial cells very small and empodium well developed as long as claws. Subgenus Forcipomyia , s. str. It is characterized by having two spermathecae; palpi usually five-segmented, third segmented with distinct sensory pit. Female antennal segments 3-10 flask-shaped, not compressed. Wing with dense macrotrichia, costa short, first radial cell small or absent, second one short. Male genitalia with parameres separated, pointed tip; aedeagus shield-shape, rather variable. Illustration: Fig. 9 Forcipomyia ( Forcipomyia ) bipunctata (L. ) Subgenus Proforcipomyia Saunders, 195& Subgenus type: Forcipomyia wirthi Saunders (orig. desig.) Saunders (1956) proposed the new subgenus Proforcipomyia with wirthi Saunders as type, to substitute for the uncertain identification

PAGE 42

31 of hirtipennis of subgenus Buforcipomyia Malloch 1915. Since Malloch* s subgenus had been placed in miscellaneous species by Johannsen, also Virth misplaced the species calcarata (Coquillett) in this subgenus. Members of this subgenus can be differentiated from the others by the following characters: -wing densely covered -with slender macrotrichia; costa about half of wing length; palpi five-segmented, sometimes the last two segments incompletely fused; first and second radial cells small or absent; spermathecae two. Kale genitalia: paraneres TJ„ or V-shaped, arched, arising from basistylar apodemes; aedeagus shield-shaped. Illustration: Fig. 10 Forcipomyia ( Fr of orcipomyia ) wirthi Saunders. Subgenus Warmkea Saunders, I956 Subgenus type: Forcipomyia bicolor Saunders (orig. desig.) Saunders (1956) believed this new subgenus to be a "young" offshoot from the parent stock of the subgenus Forcipomyia and that it had not had time to evolve any great variety of species. Larval habitat is very uniform, all species but one have been found breeding in the leaf axils of water -holding plants in the Caribbean area. The larvae live above the water line and moved further up to pupate. • The pupa has the best specific characters. The pupa cements itself to the leaf surface by the penultimate segment. The larval exuviae can usually be found stretched out on the leaf a few millimeters behind the pupa. The distinctive characters of this subgenus are: female antenna long, segments 3-10 with hyaline sensory hairs, segment 11 rather short; wing with thin microtrichia, second radial cell long. One spermatheca.

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32 Male genitalia} parameres with' hooked tips. Illustrations i Fig. 11 Forclpomyia ( Warmkea) aeria Saunders and Fig. 12 Forclpomyia ( Warmkea ) tuberculata Saunders. Subgenus Caloforcipomyia Saunders, 1956 Subgenus type: Forclpomyia caerulea Saunders (orig. desig.) This subgenus is represented by two species from Brazil (Saunders, 1956). The larvae have subcutaneous green or blue pigment granules in the fat body. Female antenna with hyaline sensory hairs. Segment 11 of the male antenna longest. Wing with elongated second radical cell. Male genitalia: parameres V-shaped; ninth sternite transverse. Illustration: Fig. 13 Forclpomyia ( Caloforcipomyia ) glauca Edwards Subgenus Metaf ore lpomyia Saunders , 195 6 Subgenus type: Forclpomyia cerlf era Saunders (orig. desig.) A Brazilian species, cerifera Saunders was designated as a type of the subgenus (Saunders, 1956). The characteristics which differentiate it from other subgenera are: female antennae with basal segments flask-shaped, each with a hyaline hair; two sperraathecae; male genitalia: parameres fused, basistylar apodemes curved and joined; larvae bearing wax-covered globules of liquid. Illustration: Fig. 14 Forclpomyia ( Mataf orcipomyia ) pluvlalis Malloch.

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33 Subgenus Thyridomyia Saunders, 1925 Subgenus type: Thyridomyia palustris Saunders (orig. desig.) It is characterized "by having the basal antennal segments compressed in female; macrotrichia of wing not flattened, second radial cell subtriangular in female; one spermatheca; male genitalia with complicated aedeagus, lateral sclerites chitinized; parameres forming a pair of subtriangular internal plates reduced to small slender sclerites of well developed and connected caudal external lobes absent. Illustration: Fig. 15 Forcipomyia ( Thyridomyia ) palustris Saunders. Subgenus Synthyri domyia Saunders, 195 6 Subgenus type: Lasiohelea acidicola Tokunaga (orig. desig.) Members of this subgenus share the common characteristic of male genitalia with club-shaped parameres extending backward from variously shaped basal roots. Larva with well developed antennal flagellum. Body usually without scales. Spermatheca one. Illustration: Fig. 16 Forcipomyia ( Synthyridomyia ) colemani Subgenus Trichohelea Goetghebuer, 1921 Trichohelea was adopted to replace Kieffer's genus Apelma, which was occupied previously by a genus of beetles. The important characteristics of this group are: empodium of male absent or vestigial. Wing with long second radial cell, thinly covered with macrotrichia. Male genitalia: parameres H-shaped with cross bar between bases of coxites and paired forward apodemal projections and baclacard external caudal lobes arising from bases of coxites; aedeagus large and shieldshaped. Illustration: Fig. 17 Forcipomyia ( Trichohelea ) fijiensis (Macfie)

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3^ Subgenus Lasiohelea Kieffer, 1921 Subgenus type: Atrichopogon pilosipennis Kieffer (orig. desig.) = velox (Winnertz) Tokunaga (1959) reduced Lasiohelea to a subgenus of Forcipomyia . The f ollowing characteristics differentiate this subgenus from the others: wing with distinct bare areas adjoining veins, second radial cell long and very narrow; male genitalia with aedeagus subdivided into a pair of recurved sclerites, each bearing apical hook-like process; paramere simple, arch-like. Illustration: Fig. 18 Forcipomyia ( Lasiohelea ) fairfaxensis Wirth Subgenus Neoforcipomyia Tokunaga & Murachi, 1959 Subgenus type: Forcipomyia pectinunguis de Meijere (Tokunaga & Murachi, 1959:200-202) This subgenus was erected by Tokunaga (1959) for species which are closely related to Lasiohelea but the male genitalia have arch-like parameres and spoon-shaped caudal lobes arising from coxite joints, coxites at lateral ends of paramere arms. The other characters are: basal antennal segment subspherical, distal segments elongated; wing densely covered with slender macrotrichia; two spermathecae. Illustration: Fig. 19 Forcipomyia ( Neoforcipomyia ) eques (Joh.) Subgenus Dacnoforcipomyia Chan and Saunders, 1965 Subgenus type: Forcipomyia anabaenae C. & S. (Orig. desig.) Chan and Saunders (I965) erected this subgenus based on the larval stages and the male. Members of this subgenus are characterized by the following points: larval head with one pair of small,

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35 sharp, forward-pointed prominences on vertex; antennae short, stout; prothoracic pseudopod long, armed dis tally -with two stout black hooks; anal pseudopod -with only six stout black hooks; cauda short, rounded; F'emale antenna with basal five segments compressed, each with a pair of hyaline setae. One spermatheca. Kale genitalia: basistyles slender; dististyles straight, uptapered, equal in length to basistyles, basistylar apodemes joined; parameres absent; aedeagus with two parts. Illustration: Fig. 20 Forcipomyia ( Dacnof orcipomyia ) anabaenae Chan and Saunders Subgenus Phasmidohelea Mayer, 1937 Subgenus type: Phasmidohelea crudelis Mayer (orig. desig.) Mayer (1937) proposed the genus Phasmidohelea for the three species which attack phasmids. Wirth ( 1956c) reduced to subgenus of Forcipomyia based on the following characters which resembles Forcipomyia : palpi with third segment swollen to apex; basitarsus short with strong spines; empodium large; two spermathecae. They are differentiated from the other subgenera by having a tick-like abdomen in female and mouthparts short, stout and highly sclerotized. Illustrations: Fig. 21 Forcipomyia ( Phasmidohelea ) fuliginosa Mg. and Fig. 22 Forcipomyia ( Phasmidohelea ) sp. Subgenus Lepidohelea KLeffer, 19 17 Subgenus type: Ceratoppgon chrysolophus K. (orig. desig.) Members of this subgenus are characterized by having foursegmented palpi; two spermathecae; male genitalia with parameres

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36 separated; hind tibial coirib present. Illustration: Fig. 23 Forcipomyia ( Lepidohelea ) annulatipes Macfie Subgenus Pterobosca Macfie, 1932 Subgenus type: Ceratopogon aeschnosuga Meijere (Macfie 19^0 a: 16) It differs from subgenus Forcipomyia by having six elongated terminal antennal segments; empodium of female large and broad, adapted for clinging; wing with moderate macrotrichia but -without scale-like hairs; spermatheca one or two. Illustration: Fig. 2k Forcipomyia ( Pterobosca ) fusicomis (Coq.)

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Key for the Subgenera of Forcipomyia 1. Antennae with last six segments elongated (Fig. 2^A); legs with empodium very broad, modified for clinging (Fig. 2^H) (female mainly ectoparasite on Odonata or rarely on lacewing flies) Pterobosca (Fig. 24) Antennae with last five segments elongated; empodium large but normal 2 2. Two spermathecae 3 One spermatheca 9 3. Last five segments of antenna combined not longer than the preceding eight segments combined kLast five segments of antenna combined much longer than the preceding eight segments combined 6 4. Sensory pit deep (Fig. 9C); antenna with segments almost similar in shape and size (Fig. 9A) . . . Forcipomyia (Fig. 9) Sensoria scattering, without distinct pit; antenna with last five segments long, each segment more than three timos longer than wide 5 5. First tarsomere short or about the same size as the second segment (Fig. IkG); palpi four-segmented (Fig. IkC); male parameres fused midline, truncate anteriorly (Fig. 14K) Metaforcipomyia (Fig. Ik) First tarsomere longer than the second segment (Fig. 13 j); palpi five-segmented (Fig. 13c, D); male parameres V-shaped, fused in the middle (Fig. I3M) . . . Caloforcipomyia (Fig. 13)

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38 6. Basitarsus very short about half the size of the second segment (Fig. 2 IK, 22G) Phasmidohelea (Fig. 21,22) Basitarsus simple, not shorter than the second segment .... 7 7. Antennal segments 3-10 oval to flask-shape (Fig. 10A) Proforcipomyia (Fig. 10) Antennal segments 3-10 depressed to transverse 8 8. Antennal segments three to seven depressed, 8-10 slightly rounded (Fig. 17A); parameres H-shaped with cross bar and paired forward apodemal projections and backward external caudal lobes (Fig. 17*0) Trichohelea (Fig. 17) Antennal segments 3-10 slightly transverse to oval (Fig. 19A); parameres fused arch-like, arising from joint of basistyles (Fig. 19*0) Neoforcipomyia (Fig. 19) 9. Palpi four-segmented 10 Palpi five-segmented 11 10. Parameres separated (Fig. 23*0); antennal segments similar in sise and flask-shaped (Fig. 23A) . . Lepidohelea (Fig. 23) Parameres fused basally (Fig. 12D) ; antenna with last five segments very long about four times as long as broad, segments 3-10 oval (Fig. 11A). Warmkea (Fig. 11,12) 11. Last five antennal segments combined not longer than all the preceding eight segments combined 12 Last five antennal segments combined much longer than the preceding eight segments combined 13

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39 12. Male genitalia: parameres with basal apodeme forming a pair of subtriangular internal plates (Fig. 15K) Thyridomyia (Fig. 15) Male genitalia: parameres club-shaped, arise from basal apodemes (Fig. 16K) Synthyridomyia (Fig. 16) 13. Third palpal segment broad without distal neck; sensorypit large (Fig. 20B) ; antenna with first five segments compressed, bearing a pair of hyaline setae (Fig. 20A); parameres absent, aedeagus separated, each with median rounded lobe, anterior arms straight, posterior tip blunt (Fig. 201) Dacnoforcipomyia (Fig. 20) Third palpal segment slender or slightly swollen (Fig. 18C); antennal segment 3-10 oval, 11-15 elongated (Fig. 18A); parameres slender joined mesally, arch-like (Fig. 18N), aedeagus subdivided into pair of recurved sclerites, each with apical hook-like process (Fig. 18*0) . Lasiohelea (Fig. 18)

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Genus Dasyhelea Kief f er Dasyhelea Kieffer, 1911, Bull. Soc. Hist. Nat. Metz, 27:5; 1913, Rec. Ind. Mus., 9*179, 1913, Ann. Mus . Nat. Hung., 17:48; 1919, Bull. Soc. Ent. France, :192; Goetghebuer, 1920, Mam. Mas. Roy. Hist. Nat. Belg., 8:40; Carter, Digram, and Macfie, 1921, Ann. Trop. Mad. & Parasit., 16:178; Kieffer, 1924, Bull. Soc. R. Ent. Egypte :253; Edwards, 1926, Trans. Ent. Soc. London, 74:401; Thienemann, 1926, Mitt. Geogr. Ges. Lubeck, 31:104; Ingram and Macfie, 1931, Dipt. Pat. & S. Chile, pt. 2, fasc. 4, :177; Sebess V. Zilahi, 1931, Arch. Hydrobiol. 23:310; Goetghebuer, 1935, Rev. Zool. Bot. Afr., 27:164; Tokunaga, 1940, Tenthredo, 3:116; Zilahi-Sebess, 1940, Folia Ent. Hungar., 5:44; Johannsen, 1948, Ann. Ent. Soc. Amer., 36:778; Tokunaga & Murachi, 1959, Inst. Micron. 12(3) :238; Remm, 1962, Tartu Riik. Ulik. Toim, 120:108; Tokunaga, 1962, Pacif. Ins. 4:189. (Type -species : Dasyhelea halophila Kieffer (orig. desig.) Prokempia Kieffer, 1913, Rec. Ind. Mas., 9:163, 179 (Type-species: not desig., two orig. incl. spp., Dasyhelea ( Prokempia ) ornaticornis Kieffer and D. ( Prokempia ) longicomls Kieffer). Pseudoculicoldes Malloch, 1915, Bull. m. State Lab. Nat. Hist., 10:309 (Type-species: Ceratopogon mutabilis Cooulllett. orig. desig.). Tetrahelea Kieffer, 1925, Arch. Inst. Pasteur d' Algeria, 3:422 (Type-species: Cullcoides insignicornis Kieffer. orig. desig.). Cryptoscena Enderlein, 1936, Tierwelt Mitteleur., 6:51 (Type-species: Ceratopogon palustris Meigen, mon.).

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mcrvotoscona Bndorlein, 1936, Tiarwelt Mltteleur., 6:51 (Typespecies: Daryholea inclnra Kieffer, won.) Totrnphora of authors (not Philippi, 1865). DIAGNOSIS. Body stout and hairy but not as dense as in Forc-i joongla . Eyes contiguous with short pubescence. Female antennal segments similar in shape, usually with verticils; male antennae with distal four segments elongate. Maxillary palp slender, first segment indistinctly separated, only four segments visible. Humeral pits absent, mesonotum covered with hairs. Legs slender; tarsal segments simple; claws small and equal; empodium small. Wings with macrotrichia over all or part of surface; costa ending about half of wing length; first radial cell obliterated, second radial cell small with square end; median fork sessile or short petiolate. Abdomen stout; one to three spermathecae. Male genitalia with long and tapering ninth tergite, apicolateral processes well developed; basis tyles short and broad; dististyies slender to stout, with or without basal branch; aedeagus large, often with paired apodemes ; parameres usually asymmetrical, with single median sclerite and pairs of anterior arms. Approximately 315 species have been described. Distribution: World-wide . Very few studies have been made on the biology and early stages of Dasyhelca . Carter, Ingram and Macfie (1921) and Goetghebuer (1925) reported on the early stages of D. bilineata . A key to larvae and papae was presented by Thienemann (1926) and the life history of Hungarian species, D. geleiana has been worked out by Zilahi-Sebess (1931). Vattier (1965) described the morphology and biology of D. adar-i from Congo.

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The following descriptions are taken from Thomsen (1937)» Mirth and Stone (1956) and Wirth (1965): EGG. Dasyhelea eggs are horseshoe-shaped, the masses are small, detected by microscope. LARVA. Head oval. Mouthparts hypognathous . Eyes crescent-shaped with heavily pigmented. Mandibles usually three-toothed. Hypopharynx heavily sclerotized, composed of dorsal and ventral parts, with posterior portions bearing teeth. Body segments short and thick, colored by pigmented fat-bodies beneath the hypodermis. Protrusible proleg on last segment bearing hooks. Retractile filamentous anal gills posterior to prolegs. PUPA. Anal segment -with pair of setigerous protuberances in addition to apicolateral processes. Opercula, lacking long setae, with or •without hooks in some species. Adult habits and larval habitats: adults visit flowers, feeding on nectar. The immature stages are semiaquatic. Larvae are found in the blanket algae of ponds or slow streams, algae on dripping banks, and the fermenting sap of tree ulcers. The larvae cannot swim but travel by crawling over the substrate. Most members of the genus spin tubular cases in the last instar. This genus was erected by Kieffer in 1911 for the Indian species, Dasyhelea halophila . It has characteristics intermediate between Forcipomyia and Culicoides . The wing venation resembles Forciponyia but tarsi lacks empodium as in Culicoides , however, it can be distinguished from the above genera by the following points: eyes pubescent; palpi four-segmented; wing vein R^ and Rg fused forming

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43 a single vein; humeral pits absent; male genitalia: parameres often asymmetrical with paired basal arms and single median lobe. Dasyhelea is rather a difficult genus. The males are comparatively easier to identify than females. Tokunaga and Murachi (1959) suggested the use of the female subgenital plate for identification. Subgenus Dasyhelea , s. str. Wing -with one radial cell usually slightly longer than broad, rarely square or missing. Scutum pruinose grayish or bluish. Female antenna with flagella segments basally striped, segment 15 with long apical style. Spermatheca one. Legs usually banded. Male genitalia: ninth sternite not modified, tergite with long apicolateral processes; hook or lobes on inner side of basis tyles. Illustration: Fig. 25 Dasyhelea ( Dasyhelea) grisea (Coq.) Subgenus Dicryptoscena End. , 193& Wing with two radial cells, both longer than broad, more or less equal in length. Palpal segments slender. Female antenna with flagella segments not basally striped, segment 15 with apical style. Spermatheca one, subspherical, with short duct. Male genitalia: ninth tergite with apicolateral processes very short. Subgenus Pseudoculic oides Malloch, 19 15 Subgenus type: Ceratopogon mutabilis Coq. Wing with second radial cell square or slightly longer than broad. Female antenna with flagella segments not striped, without apical style. Scutum usually dull, black, with or without pruinose gray pattern and rarely shining. Spermatheca one. Male genitalia:

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ninth sternite produced caudad in the middle, joined with aedeagus; parameres symmetrical or asymmetrical. Illustration: Fig. 26 Dasyhelea ( Pseudoculic oides ) mutabilis Coq. Subgenus Prokempia Kieffer, 1913 Wing with square second radial cell, macrotrichia scanty. Female antenna with flagella segments not striped, no apical style. Scutum usually shining, without pruinose pattern. Spermatheca one. Itele genitalia: aedeagus irregular, H-shaped, separated from caudal border of ninth stemite; latter with two long processes; parameres asymmetrical. Illustration: Fig. 27 Dasyhelea ( Prokempia ) cincta (Coq.)

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45 Key for the Subgenera of Dasyhelea 1. Two radial cells present on wing Dicryptoscena One radial cell on wing 2 2. Female antenna with flagella segments basally striped, last segment with long apical stylo (Fig. 25A). Second radial cell rarely square (Fig. 25C,D) Dasyhelea (Fig. 25) Female antenna with flagella segments not striped, without apical style (Fig. 26A, 27A). Second radial cell square (Fig. 26E, 27D) 3 3. Scutum usually dull, black, with or without pruinose gray pattern Pseudoculicoides (Fig. 26) Scutum usually shining, without pruinose pattern' . Prokempia (Fig. 27)

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46 Genus Paradasyhelea Macfie Paradasyhelea Macfie, 1940, Ann. Trop. Med. Parasit, 34:17; Wirth and Lee, 1959, Bull. Brooklyn Ent. Soc., 54:114; Wirth and Blanton (in press). (Type-species; Dasyhelea brevipalpis Ingram and Macfie orig. desig.). DIAGNOSIS. Eyes widely separated, covered densely -with fine hairs. Clypeus hairy. Proboscis short about half as long as the length of the head, vestigial mouth parts. Palpi four-segmented, the largest segment bearing long modified sensory hairs on distal portion, the shallow, small sensory pit indistinct. Antenna 15-segmented. First segment with a row of hairs, third to fifteenth segment uniform in shape. Male antenna plumose. Humeral pits present. Legs slender, tibial comb with four spines, claws small, equal, with bifid tip in male; vestigial hair-like empodium. Wing with short costa, radial cells obliterated, r-m cross vein oblique almost in line with a single radial vein, intercalary fork well developed; anal angle obtuse; macro trichia abundant. One spermatheca. Male genitalia with ninth sternum excavated posteriorly; ninth tergum with well developed apicolateral processes; basis tyles and dististyles simple; aedeagus arched, median part small; parameres appear only as a sclerotized rod connecting the two ventral roots, middle portion broken. Approximately three species have been described. Distribution: South America and Australia. PUPA. Elongate conical as in Culicoides and Dasyhelea. respiratory trumpet stalked, flattened, mid portion wrinkled transversely and bearing papillae around apex. Tubercles of cephalothorax with spines,

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47 abdominal tubercles each with fine hair; apicolateral processes ending in a blade-like spine pointed almost perpendicular to the body. Macf ie ( 1940a) proposed this genus for Dasyhelea brevipalpis Ingram and Macfie, which he differentiated from Dasyhelea and Forcipomyia, as discussed by Ingram and Macfie (1931). The important characteristic is the unsculptured antennal segments; not binodose in segments 12-14 of male. Paradasyhelea is closely related to Forcipomyia , Dasyhelea , and Culicoides . It resembles Forcinomyia in having abundant macrotrichea on wing; scutellum hairy; eyes well separated; antennae with sensory pegs scattered. It is similar to Dasyhelea by the following points: palpi four-segmented; eyes hairy; proboscis and mouthparts poorly developed; and the shape of apicolateral processes of ninth tergite in male. Like Culicoides , it has prominent humeral pits; antennal sensory pits and aedeagus Y-shaped. Illustration: Fig. 28 Paradasyhelea minuta Wirth and Lee Genus Culicoides Latreille Culicoides Latreille, 1808, Gen. Crust, et Ins., 4:251; Johannsen, 1905, Bull. N. Y. State Mas., 86:101; KLeffer, 1906, Gen. Insectorum, fasc. 42:53; Johannsen, 1908, Bull. N. Y. State Mas., 124:267; Kieffer, 1913, Rec. Ind. Mas., 9:162; 1919, Ann. Kus. Nat. Hung., 17:31; Goetghebuer, 1920, Mem. Mus. Roy. Hist. Nat. Belg., 8:48; Carter, Ingram and Macfie, 1920, Ann. Trop. Med. Parasit., 14:212; Hoffman, 1925, Amer. Hyg., 5:274; Edwards, 1926, Trans. Ent. Soc. London, 74:403; Johannsen, 1931, Arch, f. Hydrobiol., Suppl., 9:428; Maker ji, 1931, Ind. J. Med. Res.,

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48 18:105**, 1056; 1931, Nature, 127:339; Pomerantzen, 1932, Mag. Parasit. Leningrad, 3:183; Vircraer, 1932, Shorn. Ent. Odd. Narod. Mas. Praze 10:132; Goetghebuer, 1933, Flieg. Pal. Reg. 13a:37; 1935, Rev. Zool. Bot. Afr., 27:170; Tokunaga, 1937, Tenthredo, 1:273; Costa Lima, 1937, Mem. Inst. Oswaldo Cruz, 32:411; Root and Hoffman, 1937, Amer. J. Hyg., 25:156; Macfie, 1937, Proc. Roy. Ent. Soc. London (B), 6:112; Causey, 1938, Amer. J. Kyg., 27:399; Kono and Takahasi, 1940, Ins. Mats. 14:69; Tokunaga, 1940, Tenthredo, 3:1*1-1; Takahasi, 1941, Ins. Matsum. 15:80; Zilahi-Sebess, 1941, Folia Ent. Hung., 6:32; Fox, 1942, Puerto Rico J. Pub. Health and Trop. Med., 17:412; James, 1943, Pan-Pacif. Ent., 19:148; Johannsen, 1943, Ann. Ent. Soc. Amer., 36:779; Colaco, 1946, Ann. Inst. Med. Trop. Lisbon, 3:222; Fox, 1946, Ann. Ent. Soc. Amer., 39:2*48; Barbosa, 1947, Amu Soc. Biol. Pernambuco, 7:3; Hill, 1947, Ann. Trop. Med. Parasit., 41:55; Macfie, 1948, Ann. Trop. Med. Parasit., 42:67; Fiedler, 1951, Onderstepoort J. Vet. Res., 25:10; "Mirth, 1951, Ann. Ent. Soc. Amer. 44:76; Downes and Kettle, 1952, Proc. R. Ent. Soc. Lond. (B), 21:77; Goetghebuer, 1952, Biol. Jaarb. , 19:186; Gutsevich, 1952, Parazit. Shorn. Zool. Inst. Akad. Nauk SSSR, 14:79; Kettle and Lawson, 1952, Bull. Ent. Res., 43:428; Lee and Reye, 1953, Proc. Linn. Soc. N.S.W., 77:373; Ortiz, 1953, Rev. Sanid. Asist. Soc. 18:?; Foote and Pratt, 1954, Publ. Hlth. Monogr., Washington No. 18:10; Fox, 1955, J. Agr. Univ. P. R., 39:219; Kettle and Lawsen, 1955, Proc. R. Ent. Soc. Lond. (B) 24:46; Ortiz and Leon, 1955,

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Bol. Inf., Cient. Nac. 7, No. 67:566; Arnaud; 1956, Microentomology, 21:134; Jones, 1956, ProC. Ent. Soc. Wash., 58: 2 5; Lewis, 1956, Canad. Ent., 88:551; Forattini, 1957, Agr. Fac. Hig. Univ. S. Paulo, 11:194; Khalaf, 1957, Amer. Midi. Nat. 58:187; Wirth and Blanton, 1956, Proc. Ent. Soc. Wash., 58:308; Wirth, 1958, J. Kansas Ent. Soc, 31:85; De Prada and Gil Collado, 1959, Med. Trop. Madrid, 34:4-23; Ortiz, 1959, Rev. Sanid., Asist. Soc, 24:355; Sen and Das Gupta, 1959, Ann. Ent. Soc. Amer., 52:629; Tokunaga, 1959, Pacif. Ins., 1:204; Tokunaga and Murachi, 1959, Ins. Micron., 12, No. 3:322; Wirth and Blanton, 1959, Proc TJ. S. Nat. Mus., 109:262; Wirth and Hubert, 1959, Pacif. Ins., 1:5; Campbell and Pelhan-Clinton, i960, Proc Roy. Soc Edirib. (B) 67:196; Gutsevich, i960, Opred. Faune SSSR, 72:44; Wirth and Hubert, i960, Ann. Ent. Soc. Amer., 53:643; Callot and Kremer, 1961, Ann. Parasit. Hum. Comp. 36:686; Delfinado, I96I, Fieldiana (Zool.), 33:636; Hubert and Wirth, 1961, Proc. Ent. Soc Wash., 63:236; Jones, 1961, Ann. Ent. Soc Amer., 54:741; Khalaf, 1961, Beitr. Ent. 11:450; Nagaty and Morsey, 1961, J. Arab. Vet. Med. Ass., 21:364; Pais Caeiro, I96I, Estud. Ens. Docum. Jta. Inv. Ultramar., No. 86:46; Wirth and Hubert, 1961, Pacif. Ins. 3:12; Galukhova, 1962, Trudy. Zool. Inst. Akad. Nauk SSSR, 31:218; Tokunaga, 1962, Pacif. Ins. 4:203, 458; Wirth and Hubert, I962, Ann. Ent. Soc. Amer., 55:184; Jamnback and Wirth, I963, Ann. Ent. Soc Amer., 56:186; Tokunaga, 1963, Plant Prot. Bull. 5:120; Wirth and Blanton, I967, Fla. Ent. 50(3) :209. (Type-species: Culex pulicaris Linnaeus, as Culicoides punctata Latreille, orig. desig.)

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50 Oecacta Poey, 1851, Mem. Hist. Nat. Isla Cuba, 1:236. (Type-species: Oecacta furens Poey, mon.). Psychophaena Philippi, 1865, Verh. Zool.-Bot. Ges. Wien, 15:628 (Type-species: Psychophaena pictipennis Philippi, mon.). Haematomyidium Goeldi, 1905, Mem. Mas. Goeldi, 4:137 (Type-species: Haematomyidiun paraense Goeldi, orig. desig.). Cotocripus Brethes, 1912, Ann. Mas. Nac. Buenos Aires, 22:451 (Type-species: Cotocripus caridei Brethes, mon.). Haemophoructus Macfie, 1925, Bull. Ent. Res., 15:349 (Type-species: Haemophoructus maculipennis Macfie, mon.). Synhelea Kieffer, 1925, Arch. Inst. Pasteur d'Algerie, 3:423 ( Type-species : ? ) . Prosapelma Kieffer, 1925, Arch. Inst. Pasteur d*Algerie, 3:417 (Type-species: Prosapelma cinerea Kieffer, orig. desig.). Hoff mania Fox, 1948, Proc. Biol. Soc. Wash., 6l:21 (Type-species: Culicoides inamollae Fox and Hoffman, orig. desig.). DIAGNOSIS. Body slender, with moderate hairs. Eyes usually bare. Female antennae with segments 3-10 oval or rounded and usually much shorter than the last five segments which are usually cylindrical in shape. Male antennae plumose, three apical segments long. Mesonotum usually with pattern, covered with short hairs and usually with few bristles; humeral pits well developed. Legs and tarsi simple; claws of both sexes tiny but equal; empodium vestigial. Wings densely covered with microtrichia; macro trichia always present near wing tip; wings often with spotted pattern; costa usually to two-thirds wing

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51 length; two radial cells, almost equal; media petiolate with parallel branches. Kale genitalia with ninth sternite short, with mesal excavation; ninth tergite usually with apicolateral processes present; basistyle often with both dorsal and ventral root developed; dististyle long and curved; aedeagus variable, usually cone-shaped with anterior lateral arms; parameres usually separate with sharp point or irregular shape. Approximately 1000 species have been described. Distribution: World-wide. The immature stages of Culicoides have been studied by several authors. The American species were described by the following authors: Painter (1926), Dove, Hall and Hull (1932), Thomsen (1937), Fox (19^2), Smith and Lowe (19^8), Williams (1951), Wirth (1952), Jamnback, Wall and Collins (1958), Jones (196l). On the European species, the studies had been done by: Rieth (1922), Thienemann (1928), Lenz (193*0, Mayer (193*0 and Jobling (1953); descriptions of the British species have been given by: Hill (19^7), Dowries (1950), Laws on (1951) and Kettle and Laws on (195 2 ). Culicoides immature stages were reported; from Africa by Goetghebuer (1933), Carter, Ingram and Macfie (1920), and Ingram and Macfie (1921); from Brazil by Lutz (1912), Forattini, Rabello and Pattoli (1956, I960); from India by Patton (19 13) and Patel (1921); from French Guiana by Forattini and Rabello (1956); and from Japan by Tokunaga (1937). Particularly detailed descriptions have been given by Carter, Ingram and Macfie (1920), Hill (19^7), Kettle and Lawson (1952) and Jones (I96I).

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52 Breeding places of Culicoides were reported by Lutz (1912), Rieth (1922), Thorpe (1927), Jobling (1929), Mayer (193^), Buckley (1938), Virth, 0. M. , W. W. Wirth and F. S. Blanton (1968). EGGS. Eggs are elongated and slightly curved. They are deposited in masses. LARVA. Larvae are eel-like form, without prolegs. The body segments are short and thick cylinders, each segment carrying a pair of lateral setae. The last segment bears three pairs of anal setae, short and light in color. The head is well sclerotized with prognathous mouthparts. Mandibles are curved. Pharyngeal skeleton has combs. PUPA. The pupa is well sclerotized. It has dark brown color and is free from larval exuvium. Respiratory organ is elongated, tubular, and constricted at the base. The operculum is either with or without spines. The anal segment has one pair of apicolateral spines. Adult habits and larval habitats: The adults are generally most active at dawn and dusk (Kettle, I962). Their flight and other activities are strongly influenced by temperature and wind and other conditions. The females require a blood meal, but males feed only on nectar or plant juices. Each species has a range of vertebrate hosts on which it will feed. Feeding may occur anytime, but some species are crepuscular. The resting places are usually in damp cool situations, such as algae, the base of trees, moss, grass or dead leaves (Kettle, I962). Larvae may be found in a wide variety of moist environments, such as ponds, creeks, lakes, tree holes, slime-covered bark, rotting forest litter, rotting banana stalks. In the recent paper, Wirth, 0. M. , W. W. Wirth and F. S. Blanton (1968) reported finding two

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53 habitats of Panama Culicoides . These habitats ware rotting spadics of the Panama hat palm, Carludovica palraata Ruiz and Pavon and older decaying in florescences of the "beefsteak heliconia" Eelicor/i a mariae Eooker.f. The majority of the species of this genus may be recognized by having spotted -wings which is absent in some species. Malloch (1915) was the first to show the sensory pits as a distinct generic character. These pits do not appear in Dasyhelea , the nearest genus to Culicoides . They are present in Ceratopogon . Honohelea and some Stilobezzia . -which can be differentiated from Culicoides since the females have large claws or modified legs. This genus comprises a large number of species widely distributed over the world. Many attempts have been made to divide Culicoides into subgenera. The important references are Vargas (i960) who separated the subgenera based on wing patterns arid male genitalia and Fox (1955) who emphasized only the male genitalia. Subgenus Culicoides s. str. Cell with a marginal dark spot shaped like an hour-glass; cubital fork in dark area; second radial cell in light spot. Male genitalia: basistyles short and strong pilosity at the inner margin; parameres with short hairs at tip; aedeagus V-shaped. Illustration: Fig. 29 Culicoides ( Culicoides ) yukonensis Hoffman Subgenus Trithecoides Wirth and Hubert, 1959 Subgenus type: Culicoides flaviscutatus W. & H., 1959 (orig. desig.) Characters as follow: three well developed spermathecae; second

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5 h radial cell long; wing with two pale areas, over r-m and over apex of second radial cell; large pale area on the basal part of wing across extreme wing tip. Male genitalia with ventral root reduced; aedeagus blunt tip. Illustration: Fig. 30 Culicoides ( Trithecoides ) flaviscutatus W. & H. Subgenus Haemophoructus Mac fie, 1925 Subgenus type: Haemophoructus maculipennis Macfie (orig. desig.) Female wing with a single radial cell. Illustration: Fig. Jl Culicoides ( Haemophoructus ) gemellus Macfie Subgenus Mei.jerehelea Wirth and Hubert, 1961 Subgenus type: Ceratopogon guttifer Meijere (orig. desig.) One sperrnatheca; wing with costal pale spot in cell Rtj immediately distal to second radial cell and a round pale spot at extreme apex of cell. Male genitalia: aedeagus usually flaring at tip; parameres with large basal knob directed laterad. Illustration: Fig. 32 Culicoides ( Mei.jerehelea ) guttifer (Heijere) Subgenus Hoffmania Fox, 19^7 Subgenus type: Culicoides inamollae Fox and Hoffman (orig. desig. )= insignis Lutz, 1913. Pale spot on vein M 2 ; cubital fork in light area; macrotrichia abundant; r-m crossvein dark with large light spot. Male genitalia: aedeagus bottle-shaped, parameres fused in some species. Illustration: Fig. 33 Culicoides ( Hoffmania ) insignis Lutz

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55 Subgenus Avarltia Fox, 1955 Subgenus type: Ceratopogon obsoletus Meigen (orig. desig.) Wing marking faint; macrotrichia scanty. Male genitalia: ninth tergite without apicolateral processes; ventral root of basistyle very long. Illustration: Fig. 34 Culicoides ( Avaritia ) obsoletus (Mg.) Subgenus Oecacta Poey, 185 1 Subgenus type: Oecacta furens Poey (mon.) Second radial cell in dark spot; clear spot on vein absent; three pale spots on Mj_. Male genitalia: ventral root with retrorse tooth; parameres with spinose tip; aedeagus triangular. Illustration: Fig. 35 Culicoides ( Oecacta ) furens (Poey) Subgenus Drymodesmyia Vargas, i960 Subgenus type: Culicoides copiosus Root and Hoffman, 1937 (orig. desig.) Wing hairy; second radial cell within dark area; cell Re with two pale spots, one is about the middle of cell. Male genitalia: basistyles with ventral hook, dististyles hook-shaped, paramere with sharp tip. Illustration: Fig. 36 Culicoides ( Drymodesmyia ) copiosus R. & H. Subgenus Diphaeomyia Vargas , i960 Subgenus type: CxTlicoides baueri Hoffman, I925 (orig. desig.) Male genitalia with ventral root long, curved, rounded apically with a small notch at base; parameres with many short spines. Illustration: Fig. 37 Culicoides ( Diphaeomyia ) baueri Hoffman

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56 Subgenus Anilomyia Vargas, i960 Subgenus type: Culicoides covagarciai Ortiz, 1950 Wing with darkest spot occurring basal to second radial cell. Male genitalia with inner margin of basis tyle spinose; aedeagus bottle-shaped; parameres fused or closely appressed with a terminal brush. Illustration: Fig. 38 Culicoides ( Anilomyia ) covagarciai Ortiz Subgenus Beltranmyia Vargas , 1953 Subgenus type: Culicoides crepuscularis Malloch, 1915 (orig. desig.) Wing dark with pale markings and many long hairs. Male genitalia: aedaegus Vor Y-shaped, truncate, broad tip. Illustration: Fig. 39 Culicoides ( Beltranmyia ) crepuscularis Malloch Subgenus Glaphiromyia Vargas, i960 Subgenus type: Culicoides scopus Root and Hoffman, 1937 (orig. desig.) Wing with pale spot on vein Mj_ and Mg; r-m crossvein dark in some species. Male genitalia: ventral root long and sharp; parameres with terminal hairs; aedeagus arch-shaped. Illustration: Fig. 40 Culicoides ( Glaphiromyia ) scopus R. & H. Subgenus Haematomyidium Goeldi, 1905 Subgenus type: Culicoides paraensis (Goeldi, 1905) Crossvein r-m usually dark but clear basally. Male genitalia: ventral root well developed with retrorse tooth; parameres with spinose tips; aedeagus rounded apically. Illustration: Fig. 41 Culicoides ( Haematomyidium ) paraensis (Goeldi)

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57 Subgenus Kacfiella Fox, 1955 Subgenus type: Ceratopogon phlebotomus Willis ton Second radial cell in a dark spot. Male genitalia with ninth tergite rounded apically; basal root long; dorsal root somewhat boot-shaped. Illustration: Fig. 42 Culicoides ( Hacfiella ) phlebotomus (Will.) i Subgenus Kataemyia Vargas, 19 60 Subgenus type: Culicoides mo.jingaensis Wirth and Blanton, 1953 (orig. desig.) Vein usually without pale spot; Mj, and ML with long light bands. Ventral root of male genitalia prominent; parameres sharp pointed. Illustration: Fig. 43 Culicoides ( Hataemyia ) mojingaensis W. & B. Subgenus Konoculic oides Khalaf , 195^ Subgenus type: Ceratopogon nubeculosus Meigen (orig. desig.) Wing with second radial cell in a dark spot. One spermatheca. Male genitalia with paramere fused forming a plate and usually bifid distally; aedeagus deeply bifid; dorsal root often boot-shaped. Illustration: Fig. 44 Culicoides ( Konoculic oides ) variipennis (Coq} Subgenus Selfia Khalaf , 1954 Subgenus type: Culicoides hieroglyphicus Malloch (Orig. desig.) Wing dark without pale marking. Spermathecae not apparent. Kale genitalia with parameres fused. Aedeagus with two sclerotised pieces often fused apically. Illustration: Fig. 45 Culicoides ( Selfia ) hiero^lyphicus Malloch

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58 Key for the Subgenera of Culicoides 1. Spermathecae unsclerotized, not apparent; wing dark without pale marking Selfia (Fig. 45) Spermathecae sclerotized; wing usually -with distinct pale spots 2 2. One spermatheca 3 Two to three spermathecae 5 3. Parameres separated 4 Parameres fused to form a plate usually bifid distally vTith two apical points (Fig. 44«0) . . Monoculicoides (Fig. 44) 4. Paramere with large basal knob directed laterad (Fig. 32P), stem short, rather stout simple tip without distal fringing spines Meijerehelea (Fig. 32) Paramere with small base, tapering, directed basally, and pointed tip. (Fig. 39*0) Beltranmyia (Fig. 39) 5. Three well developed spermathecae (Fig. 30H) . Trithecoides (Fig. 30) Two well developed spermathecae 6 6. Female wing with a single radial cell (Fig. 31E) Haemophoructus (Fig. 31) Female wing with two radial cells, almost equal in size ... 7 7. Ventral root absent or poorly developed 8 Ventral root large and well developed 10

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59 8. Base of paramere sharply bent and close together, tip with short hairs (Fig. 29L); -wing without pale spot on vein Culicoides (Fig. 29) Base of paramere gradually bent; -wing -with pale spot on vein M> 9. Wing with cubital fork in a dark area (Fig. 38S) Anilomyia (Fig. 38) Wing with cubital fork in a light area (Fig. 332) Koffreania (Fig. 33) 10. Second radial cell included in a pale area (Fig. 3^F) Avaritia (Fig. 34) Second radial cell included in a dark area 11 11. Middle of vein M^ not straddled by pale spot 12 Middle of vein straddled by pale spot 13 12. Vein M 1 and M2 -within long dark bands (Fig. 4 IE); ventral root with retrorse tooth (Fig. 4lN). . Haeiaat oroyidium (Fig. 4l) Vein M ± and Mg within long pale bands (Fig. 43D); ventral root prominent Mataemyia (Fig. 43) 13. Vein and M 2 within dark bands 14 Vein M^ and Mj> within pale bands 15 14. Third palpal segment with sensory pit scattering, without a true pit (Fig. 42C); aedeagus without a prominent arch, apical portion long, truncate distally, basally with a long, curved hook-like pointed processes (Fig. 42N) Macfiolla (Fig. 42)

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60 Third palpal segment with a true and large sensory pit (Fig. 40C); aedeagus arch with long stem (Fig. 40N) Glaphiromyia (Fig. 40) 15. Aedeagus with small, tapering tip, with short process on arch (Fig. 37N) Dinhaeomyia (Fig. 37) Aedeagus with broad or truncate tip, without process on arch *6 16. Base of paramere bent perpendicularly, sharp tip, not spinose (Fig. 36*0) Drymodesmyia (Fig. 36) Base of paramere small and gradually bent, subapical portion with globose process and tip spinose (Fig. 35*°) Oecacta (Fig. 35)

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61 Genus Austroconops Wirth and Lee Austroconops •irth and Lee, 1958, Proc. Linn. Soc. N. S. Wales, 83:337 (Type-species: Austroconops mcmillani Wirth and Lee, mon.) DIAGNOSIS. Eyes hairy, contiguous above; vertex sparsely haired. Antenna 15-segmented, flagella segments spherical basal, becoming oval distally. Clypeus small, with four long hairs; proboscis shorter than head; mouthparts well developed, mandible with eight large teeth. Palpi "four-segmented," the fourth and fifth segments fused; third segment swollen without sensory pit, sensoria covered almost half of the mid portion of the segment. Thorax robust, humeral pits conspicuous. Legs unarmed; hind tibia with plumose spur, tibial comb four spines. Claws small and equal; empodium rudimentary. Wing milky, radial veins distinctly thickened, costa long, r-m crossvein very oblique, forming almost a straight line with base of media and posterior side of radial cells; radial cells long and broad; media with long petiole; anal cell large; nacrotrichia absent. Abdomen short with blunt apex and short cerci. Tito subequal spermathecae and a rudimentary one. Male unknown. Only one species known. Distribution: W. Australia. Adult habits and larval habitats: Female blood sucking, immature stages unknown. This genus was erected in 1958 by Wirth and Lee for Austroconops mcmillani of Australia. It has not been discovered outside the Australian region.

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62 The members of this genus have the characteristics of Culicoides, Leptoconoos and Stilobezzia . It resembles Leptoconops, by the f ollowing characters: palpi four-segmented, fourth and fifth segments fused into one segment. "Wing milky white, without macrotrichea; empodium absent; antennal segments similar in size and shape; blood sucking habit diurnal. Wing venation is similar to Stilobezzia , with long costa and -well developed radial cells and petiolate-media. It has the characteristics of Culicoides in having the humeral pits on pronotum; small and equal claws, empodium vestigial; antennal segments not sculptured. Illustration: Fig. ^+6 Austroconops mcmillani "Mirth and Lee Genus Camotopterohelea Wirth and Hubert Camotooterohelea Wirth and Hubert, i960, Fieldiana (Zool.), ^2(7) :89 (Type-species: Camptopterohelea hoogstraali Wirth and Hubert, (men. DIAGNOSIS. S!yes bare, widely separated. Antenna 15-segmented, cylindrical. Palpi very short, only two segments present; the third and the fused segment of the fourth and the fifth. Proboscis vestigial, shorter than palpus, mandibles with very fine teeth. Thorax dark and stout; scutum with anterior median pair of oval hyaline pits and few hairs. Legs rather stout; front tibia with a lanceolate spur and a compact comb, hind tibia with more hairs on comb; each tarsomere bearing a single long, stout, slightly curved claw. Wing broad on basal half and tapering dis tally; costa long; two radial cells present, the first one well developed but the second one small, the radial veins thickened; the rest of the wing venation includes the cubital stem, 1A and possibly Cuji appears from posterior wing fold to margin of the wing.

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63 Disc of wing modified with two folds transversely continuous from costal end to end of Cv. ± proximally and about halfway from this line to wing tip distally, fold areas densely covered with microtrichia; along vein Cu 1 with long, spinelike microtrichia as well as over radial cells and along margin of proximal fold and the patch in front • of the distal fold; macrotrichia absent. Abdomen stout with single oval spermatheca. Male genitalia small; ninth tergite rounded; basistyle rather stout, dorsal root prominent; dististyle slender, tapering with hook-like tip; aedeagus arched with large body. Only one species known. Distribution: Philippines, Malaya. This genus is readily separated from the other genera by having two-segmented palps; proboscis rudimentary; all claws single; wing greatly modified with broad basal half and tapering distally; wing vernation with only radial vein, cubital stem and the first anal vein present; microtrichia spinelike. Illustration: Fig. k7 Camptopterohelea hoogstraali W. & H. Genus Ceratopogon Meigen Ceratopogon Meigen, 1803, THiger's Mag. Ins., 2:261; Kieffer, 1899, Bull. Soc. Ent. France, p. 69; Johannsen, 1903, Bull. N. Y. State Mas., 86:97; Kieffer, 1906, Gen. Insectorum, fasc. 42:^+8; 1919, Ann. Mus. Nat. Hung., 17:1; C-oetghebuer, 1920, Mem. Mas. Roy. Hist. Nat. Belg. 8:19; Edwards, 1920, Ann. Mag. Nat. Hist., 6:129; 1926, Trans. Ent. Soc. London, 7^07; Johannsen, 193^, J« K.J. Ent Soc, ^-2:343; Tokunaga, 19^0, Tenthredo, 3:151; Johannsen, 19^3,

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m Ann. Eat. Soc. Amer., 36:780; Wirth, 1952, Univ. Calif. Publ. Ent., 9:198; de Meillon, 1959, S. Afr. Anim. Life, 6:340; Clastrier, 1961, Inst. Pasteur Alger., 39:401; Tokunaga, 1964, Pacif. Ins. 6:292. (Type-species: Tipula barbie ornis Fabricius, mon. ?= Ceratopogon communis Meigen). Helea Meigen, 1800, Kouv. Class, des mouches a deux ailes, p. 18; Goetghebuer and Lenz, 1933, in Lindner, Flieg. Pal. Reg., 77:5, 48; Stone, 1941, Ann. Ent. Soc. Amer., 34:411 (Type-species: Ceratopogon comminis Meigen, desig. Coquillett, 1910). Brachypogon Kieffer, 1899, Bull. Soc. Ent. France, p. 69 (Type -species: Ceratopogon vitiosus Winnertz, orig. desig.). Psilohelea Kieffer, 1917, Ann. Mus. Nat. Hung., 15:294 (Type-species: Ceratopogon Candida tus Winnertz , mon.; = niveipennis Meigen). Isohelea Kieffer, 1917, Ann. Mus. Nat. Hung., 15:295 (Type-species: Ceratopogon lacteipennis Winnertz, desig. Kieffer, 1919; = sociabilis Goetghebuer) . Anakempia Kieffer, 1924, Bull. Soc. Hist. Nat. Moselle, 30:13 (TypeAnakempia grandiforceps Kieffer, orig. desig. ) . Far.-. ...-a de Meillon, 1939, Ruwenzori Exped., 1934-35, 1(5): 103 (Type-species: Ceratopogon ( Fanthamia ) adulator de Meillon, orig. desig.). DIAGNOSIS. Body stout -with few hairs. Eyes pubescent. Female antennal segments 3-10 rounded, the last five segments oval; male antennae plumose with last three segments longer than the preceding ones. Legs slender; empodiutn vestigial or absent; claws large in female, equal or almost equal. Wing without microtrichia and

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65 macrotrichia or with few macrotrichia near the wing tip; costa slightly longer than the middle of wing; usually with two radial cells, about the same size; intercalary fork indistinct; median fork usually with lower branch absent or interrupted. Male genitalia not inverted; ninth tergite small; basistyles swollen; aedeagus varies; parameres separate. Approximately 219 species have been described. Distribution: World-wide. Kettle and Laws on (1952) vas the only paper published on the immature stages and the breeding places of Ceratopogon of British species. LARVA. Head with posterior ventral suture long, well sclerotized and pigmented. Antennae more prominent (isohelea) or indistinct ( Ceratopogon) . Dorsal comb with seven to nine teeth on each side. PUPA. Thorax dark, nodular. Operculum dark, covered with dark nodules; respiratory horn short and stout, very dark, except at the tip. Metathorax not indentate. Caudal spine dark. Breeding places always are associated with Culicoides. They were reported found in moorland, in marshes and often in large numbers in Snhatmum spp. is This genus is closely related to Culicoides but the wing A without microtrichia or pattern and the claws are very large. Subgenus Ceratopogon s. str. Vein 2^ complete or narrowly interrupted; both radial cells distinct, longer than broad. Illustration: Fig. Ceratopogon ( Ceratopogon ) culicoidithorax Hoffman.

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66 Subgenus Isohelea Kieffer, 19 17 Subgenus type: Ceratopogion lacteipennis Winnertz (orig. desig.) sociabilis Goet. Vein -widely interrupted at base; costa short not much beyond the middle of -wing; radial cell small or absent. Illustration: Fig. ^9 Ceratopogon ( Isohelea ) stipmalis Coq. Subgenus Brachypogon Kieffer, 1899 Subgenus type: Ceratooo^on vitiosus Winnertz (orig. desig.) Wing -with second branch of media absent. Subgenus Nilohelea Kieffer, 1921 Subgenus type: Nilohelea albipennis Kieffer (mon.) Wing -with first and second radial cell obliterated. Subgenus Fanthamia de Meillon, 1939 Subgenus typei Ceratopogon adulator de Meillon. Wing -with complete first radial cell, second one obliterated male antenna as in female; hind claws of female greatly unequal; macrotrichia present on distal portion of -wing.

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67 Key for the Subgenera of Ceratopofon 2 1. Two complote radial cells present One or both radial cells obliterated ..... 2. Second branch of media present ' Second branch of media absent ->v — 3. YSo complete or narrowly interrupted at base, costa ending about two-thirds of wing length (Fig. **8E) Ceratopogon (Fig. *t8) M 2 widely interrupted at bat
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68 Genus Alluaudomyia Kief f er Alluaudomyia Kief., 1913, Voyage Ch. Alluaud et R. Jeannel en Afrique Orientale, Dipt. 1:12; de MeilLon, 1939, J. Ent. Soc. S. Africa 2:7; Okada, 1942, Trans. Nat. Hist. Soc. Formosa 32:315; Wixth, 1952, Ann. Ent. Soc. Amer. 45:423; Tokunaga and Murachi, 1959, Ins. Micron. 12(3): 352; Clastrier, 1961, Arch. Inst. Pasteur Alger; 39:4-28; Tokunaga, 1963, Pacif. Ins. 5:215; Wirth and Delfinado, 1964, Pacif. Ins. 6:602. (Type-species: Alluatriomyia imoarunjniis Kief. inon.). Neoceratopofcon Malloch, 1915, Bull. 111. State Lab. Nat. Hist., 11:310 (Type-species: Ceratopogon bellus Coquillett). Prionognathus Carter, Ingram and Macfie, 1921, Ann. Trop. Med. & Parasit., 16:309 (Type-species: Prionognathus marmoratus Carter, Ingram and Macfie, orig. desig.). Thysanognathus Ingram and Macfie, 1922, Ann. Trop. Med. & Parasit., 16:244 (n. nom. for Prionognathus Carter, Ingram and Macfie, not Laferte-Senectere, I85I). Isoecacta Garrett, 1925, Seventy New Diptera, p. 9. (Type-species: Isoecacta poeyi Garrett, mon. ). DIAGNOSIS. Body slender, moderately hairy, mostly small size. Eyes bare or hairy, separated or contiguous. Female antennal segments 3-10 long, segments 11-15 elongated. Male antennae with well developed plumes. Palpi slender; third segment simple. Wing without microtrichia, macrotrichia appearing along the wing tip, wing usually with 1-15 or 20 small dark spots and sometimes with grayish streaks along the wing veins; male wing bare or hairy. One radial cell present (the first one

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69 completely fused), media petiolate. Legs slender, in some species -with long hairs. Kale claws small and equal; female claws very unequal in hind legs, inner claw longer. Spermathecae one or two. Kale genitalia highly modified; ninth tergite always long, usually with well developed apicolateral processes; ninth sternife short, concave mesally; basistyle and dististyle simple; aedeagus arched with large stem; parameres usually separated. Approximately 122 species have been described. Distribution: North America, Europe, Africa, Asia, and Pacific Islands . Immature stages and biology of Alluaudomyia of North American species have been reported in Thomsen ( 1937) , Williams ( 1953) and Wirth and Stone (1956). EGG. Long and narrow, almost straight. Eggs of A. needhami placed side by side horizontally, forming a flat ribbon-like band, not jelly covered. LARVA. Brown with pear-shaped head. The dorsal side of the body colored by pigments in the fat bodies, causing the larva to appear dark red. Last body segment with eight setae as long as or longer than last segment, with addition of four short setae. PUPA. Light brown with dark thorax. Respiratory trumpet funnel-form, covered entirely with scales; respiratory spiracles arranged in pairs. Cuticular projections on the body segments vary in shape and number. Last segments with slender and pointed projections. Larval habitats: larvae of Alluaudomyia were found in algae and in mud in New Tork. Thomsen (1937) reported that larvae of A. splendida were found in algae, decaying leaves from swamps, and in the mud

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70 near the edge of ponds in New York,' A. needhami larvae were found in blanket algae in ponds. The larvae are predaceous. Williams (1953) summarized three breeding habitats of A. bella and A. needhami as follows: areas of permanent flowing water to areas of temporary water where the soil is firm enough to walk on; soil with dense growth of vegetation to areas with no vegetation and in sunny areas where the temperature reaches 90°F as well as shady areas; pH of soil may range from 5.7 to 7.3. This genus has distinctive characters and is readily recognized from the other genera in the following points: wing usually adorned with one to twenty small black spots; microtrichia absent; empodium vestigial and claws of female unequal on hind legs. Illustrations: Fig. 50 Alluaudomyia bella (Coq.) and Fig. 51 Alluaudomyia parva Wirth. Genus Stilobezzia Kief f er Stilobezzia Kieffer, 1911, Rec. Indian Mas., 6:118; 1913, Rec. Indian Mus., 9:184; 1917, Ann. Mas. Nat. Hungarici, 15:307; 1919, Bull. Soc. Ent. France 34:192; Goetghebuer, 1920, Mam. Mas. Hoy. Hist. Nat. Belgique, 3:59; Carter, Ingram, and Macfie, 1921, Ann. Trop. Med. Parasit., 15: 324; Edwards, 1926, Trans. Ent. Soc. London, 74:411; 1929, Notulae Ent., 9:9; Johannsen, 1931, Arch, ^ydrobiol Suppl. , 9:430; Ingram and Macfie, 1931a, Dipt. Patagonia and S. Chile, pt. 2, fasc. 4:191; Macfie, 1932, Ann. Trop. Med. Parasit. 26: 4*1 ; de Meillon, 1938, Proc. Roy. Ent. Soc. London (B), 7:266; Tckunaga, 1940b, Tenthredo, 3:154; Johannsen, 1943, Ann. Ent. Soc.

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71 Amer., 36:781; Lane, 19^7, Rev. Ent., 18:197; Lee, 19^8, Proc Linn. Soc. N. S. W., 72:3^5; T ^th, 1953, Proc. U. S. Hat. Mas., 103:57; Lane and Forattini, 1958, Rev. Bras. Ent., 8:203; Tokunaga and Karachi, 1959, Ins. Micron. 12(3) 13*3; Tokunaga, 1959, Pacif. Ins. 1:298; Lane and Forattini, 1961, Rev. Bras. Ent., 10:83; Tokunaga, 1962, Pacif. Ins. 4:207; Clastrier, 1963, Arch. Inst. Pasteur Algerie Tokunaga, 1963, Pacif. Ins. 5:2^9; Das Gupta and Wirth, I968, U. S. Nat. Mus. Bull., 283:16. (Type-species: Stilobezzia f estiva Kieffer, orig. desig.) Hartomyia Malloch, 1915, Bull. HI. State Lab. Nat. Hist., 10:339 (Type-species: Ceratopogon pictus Coquillett, orig. desig.). Neostilobezzia Goetghebuer, 193^, in Lindner, Flieg. Pal. Reg., 78:53 (Type-species: genotype not designated). DIAGNOSIS. Large size, slender with sparsely hairs. Eyes bare. Antennae of female with segments 3-10 oval to short cylindrical, segments 11-15 strongly cylindrical; male antennae plumose, hairs well developed on segments 3-12. Mesonotum with or without few bristles; humeral pits inconspicuous. Legs slender, without spines in all femora; fourth tarsal segment cordiform, fifth segment elongated and usually clavate; first tarsal segment of mid leg bearing subbasal spur; female claws very large and unequal, bifid; male claws small and equal; empodium absent. Wing long with few macrotrichia along wing tip; costa long; two radial cells present, second one well developed; median fork distinctly petiolate and long. Usually two spermathecae and a third small one. Male genitalia: very short ninth sternite; ninth tergite rounded; basistylo and dististyle simple; acdeagus

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72 ,/ reduced to pair of oblique lateral aclerites; parameres with modified, well developed submedian processes. Approximately 248 species have been described. Distribution: World-wide. The immature stages and biology are not well known. Few species have been described. The pupa of splrogyrae was described by Carter, Ingram and Macfie (1921). Ingram and Macfie (1922) gave the description of immature stages of poikiloptera from West Africa. In Europe, Thienemann (1928) gave the figures of the pupa of gracilis . Kettle and Laws on (1952) included two ins tars of ochracea and gracilis in their paper. Mayer (193^) studied the Dutch East Indies species of perspicua , and the pupa of soror . Thomson (1937) described and illustrated antennalis and bulla early stages from New York. Lane, Forattini, and Rabtjllo (1955) described and illustrated pupa of glauca and wygodzinsky from Brazil. LARVA. Mouthparts reduced, directed anteriorly. Head capsule well developed; pharyngeal skeleton with several combs, principal comb of pharyngeal angulus undivided; larva without prologs but with caudal bristles. Last abdominal segment without terminal hooks, with only short setae (Thomsen, 1937). PUPA. As in Culicoides but differs in having trumpet constricted basally; no spines on operculum. Adult habits and larval habitats: adult habits are poorly known. Edwards ( 1920b) indicated that the female of S. gracilis feeds on various small chironomid midges. Malloch (1915) and de Meillon (1936) reported that the Notata Group holding their wings when at

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73 rest. Edwards (1926) and Malloch (1915) reported finding the same habit in the Subviridis Group. The larvae are commonly found in mud and algae along pond margins (Thomson, 1937). In West Africa, Ingram and Macfie (1921) reported two species from Pistia in a swamp. Johannsen ( 1932) and Mayer ( 193*0 reared the larvae of perspicua in hot springs and in a rice field in Sumatra. j This genus is readily recognized by having female claws large and very unequal or a single long claw with barb; fourth tarsomere cordiform; first tarsomere of mid legs with subbasal spur; wing with second radial cell much longer than first which may be entirely absent. Subgenus Stilobezzia s. str. Members of this subgenus are characterized by the following characters: wing with two radial cells, without macro trie hia. Most of the species are described from the tropical region. Illustration: Fig. 52 Stilobezzia ( Stilobezzia) f estiva K. Subgenus Eukraiohelea Ingram and Macfie, 1921 Subgenus type: Eukraiohelea africana I. & M. (Macfie, 19*40a) Johannsen (1934) placed Eukraiohelea as a subgenus of Parabezzia Malloch (1915), but in comparing the male genitalia of Parabezzia petiolata Malloch 1915, with those of the four species of Eukraiohelea . the latter are more closely related to Stjlobezzia . This subgenus is characterized by having one radial cell on wing; vein Ri arising at or past r-m; legs with stout spines on fore femur midventrally or at base. Illustration: Fig. 53 Stilobezzia ( Eukraiohelea ) elegantulata (Joh.)

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Subgenus Neostllobezzla Goetghebuer, 1934 Subgenus type: Ceratopogon ochraceus Winnertz, 1852 (Wlrth 1953a) Goetghebuer ( 1934) distinguished this subgenus from Stllobezzia by the presence of macrotrichia at the end of the wing. In addition, all the species known are yellowish or brown to black, and the male genitalia with aedeagus bent and humped in the middle and basistyle without inner lobe. Most of the species are described from the temperate region. Illustration: Fig. 54 Stllobezzia ( Neostilobezzia ) lutea (Malloch).

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75 Key for the Subgenera of Stilobezzia 1. Wing with one radial cell (Fig. 53E); legs wi th spines on fore femora; hind femora slender and unarmed (Fig. 53G) Eukraiohelea (Fig. 53) Wing with two radial cells; femora unarmed 2. Wing entirely bare of macrotrichia (Fig. 5 2E ) Stilobezzia (Fig. 52) Wing with macrotrichia on the surface or at tip (Fig. 5*IE) Neostllobezzia (Fig. 5*0

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Genus Echlnohelea Macf ie Echlnohelea Macfle. 19**0, Proc. R. Ent. Soc. Lend. (B) 9*187; Wirth, 1951, Proc. Ent. Soc. Wash. 53019; de Meillon, 1959, Novos Taxa Ent. 13t 15; i960, J. Ent. Soc. S. Afr. 23:409; Tokunaga, 1963, Pacif. Ins. 5:231. (Type-species: Echlnohelea ornatipennis Macfie, orlg. desig. ). DIAGNOSIS. Eyes bare. Antennae of female with segments 11-15 elongated, the last segment usually shorter than the preceding four segments. Male antennae without plumose hairs. Thorax without anterior tubercle or spine, but with few stout bristles. Wing without macrotrichia, costa long, not extending beyond R^, M^ and forked before or at r-m, two radial cells present; second one well developed. Femora of all legs not greatly swollen, almost all femora and tibia armed with long spines, scattered rather irregularly; fourth tarsal segment short, not distinctly bilobed or cordiform, fifth segment very long, slender, unarmed, claws equal or unequal, each with barb. Abdominal tergites of female without gland rods. Male genitalia: ninth sternite excavated mesally, basistyles large, dististyle small and slender, aedeagus arched, ending in two parts, each bearing hook; paramere fused mesally, expanded ends. Approximately 11 species have been described. Distribution: British Guiana, Liberia, New Britain, New Guinea, Upper Volta, and U.S.A. (Florida and Virginia). This genus was reported by Macfie (I9*40b) for three very similar species from British Guiana. They resemble in general appearance some species of Serromyia but hind femora not greatly swollen. The distinctive characters of the genus are as follows: all femora and tibiae

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armed strongly with well developed spines; last antennal segment always distinctly short. The male is readily recognized by having no plumes on antennae. Illustration: Fig. 55 Echinohelea lanei Wirth Genus Mbnohelea Kieff er Monohelea Kleffer, 1917, Ann. Mas. Nat. Hung., 15:29**; Goetghebuer, 1920, Mem. Mus. Roy. Hist. Nat. Belg., 8:63; Ingram and Macfie, 1921, Ann. Trop. Med. & Parasit., 15:344; Edwards, 1926, Trans. Ent. Soc. London, 74:410; Tokunaga, 1940, Tenthredo, 3:156; Johannsen, 1943, Ann. Ent. Soc. Amer., 36:781; Lane, 1948, Arg. Fac. Hig. Saude Pub. Oniv. Sao Paulo, 1:225; Wirth, 1953, Proc. U. S. Nat. Mus., 103(3320): 136; Tokunaga and Murachi, 1959, Ins. Micron., 12(3) :4o4; Clastrier, 1963, Arch. Inst. Pasteur Algerie, 41:55; Tokunaga, 1963, Pacif. Ins., 5:238; Lane and Wirth, 1964, Studia Ent. (N.S.) 7:211; Wirth and Williams, 1964, Ann. Ent. Soc. Amer., 57:302. (Type-species: Monohelea hieroglyphic a Kleffer, orig. desig.) Schizohelea Kieffer, 1917, Ann. Mus. Nat. Hung., 15:295; 1918, Ann. Mus. Nat. Hung., 16:57; 1919, Ann. Mus. Nat. Hung., 17:89; Goetghebuer, 1920, Mem. Mus. Roy. Hist. Nat. Belg., 8:65; Edwards, 1926, Trans. Ent. Soc. London, 74:411; Johannsen, 1943, Ann. Ent. Soc. Amer., 36:782 (Type-species: Ceratopogon leucopeza Meigen, as Ceratopogon copiosus Winnertz, mon. ) Allohelea Kieffer, 1917, Ann. Mus. Nat. Hung., 15:364 ( Type-species 1 Sphaeromyas pulchrlpennis Kieffer, orig. desig.).

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DIAGNOSIS. Body short, moderately hairy. Byes bare or with very fine pubescence. Humeral pits inconspicuous. Fore and mid legs simple, fourth tarsal segment short, cylindrical, claws large or small but equal in both sexes; hind legs with thickening of femora and tibiae, spines absent; ventral of basitarsi with a spine at base, with or without spine at tip; fifth tarsal segment of female with one long claw and with or without another short one; hind claws of male small and equal except in some species; empodium absent. Wing with few macrotrichia at the tip; costa long; two radial cells, second cell well developed; intercalary fork inconspicuous; media with short stem, M> sometimes interrupted near base. Two spermathecae present. Male genitalia with bilobed or truncated ninth tergite; ninth sternite convex mesally; aedeagus usually with a pair of lateral sclerites and an additional dorsomedian lobe; parameres fused or paired. Approximately 93 species have been described. Distribution: World-wide. The habits of Monohelea are unknown. The larva of only one species, M. calcarata, has been found in moss in a spring habitat (Strenzke, 1950). The males of this genus are easy to identify by characters of the male genitalia but the great variation among the females makes identification very difficult. Wing markings are not a reliable character. Leg markings are more constant. This genus resembles Stilobezzia but differs particularly in having the fourth tarsomere cylindrical, the female claws with the four anterior claws equal but the hind claw single and long.

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Subgenus' Mbnohelea s. str. This subgenus is distinguished by the following characters : wing usually with pattern, first radial cell complete, base of Mg narrowly interrupted; scutum with pruinose pattern. Illustration: Fig. 56 Monohelea ( Monohelea ) hieroglyphlca K. Subgenus Schizohelea RLeffer, 1917 Subgenus type: Ceratopogon copiosus Wlnnertz (mon.) = leucopeza (Meigen). Members of the subgenus are characterized by: wing without pattern, first radial cell complete, Mg broadly interrupted at base scutum polished black. Illustration: Fig. 57 Monohelea ( Schizohelea ) leucopeza (Mg.) Subgenus Isthmohelea I. & M., 1931 Subgenus type: Isthmohelea dls.tancta I. & M. This subgenus is characterized by having two radial cells separated for some distance by fusion of vein between them, first radial cell complete. Subgenus Kiefferomyia Mayer, 1937 Subgenus type: Kiefferomyia gorana Mayer This subgenus differs from the other subgenera by having first radial cell obliterated.

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80 Key for the Subgenera of Monohelea U First radial cell obliterated Kiefferomyia First radial cell complete 2 2. Fourth tarsomere cordiform or bell-shaped Istbaohelea Fourth tarsomere cylindrical or subcylindrical 3 3. Wing usually with pattern; ^ not broadly interrupted at base (Fig. 56D) Monohelea (Fig. 56) Wing without pattern; ^ broadly interrupted at base (Fig. 57D) Schizohelea (Fig. 57)

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Qenas Serromvia Meigen Serromyia (Megerle, M. S. in) MBigen, 1818, Syst. Beschr. Eur. Zweifl. Ins., 1:83; Johannsen, 1905, Bull. N. Y. State Mas., 86:108; Kieffer, 1906, Gen. Insectorum, fasc. 42:64; Johannsen, 1908, Bull. N. Y. State Mas., 124:269; Malloch, 1914, Bull, HI. State Lab. Nat. Hist., 10:216; 1915, Bull. 111. State Lab. Nat. Hist., 10:331; Kieffer, 1917, Ann. Mas. Nat. Hung., 15:294; 1919, Ann. Mas. Nat. Hung., 17:69; Goetghebuer, 1920, Mem. Mus. Roy. Hist. Nat. Belg., 8:70; Edwards, 1926, Trans. Ent. Soc. London, 74:409; Tokunaga, 1940, Tenthredo, 3:156; Johannsen, 1943, Ann. Ent. Soc. Amer., 36:781; Kettle and Lawson, 1952, Bull. Ent. Res., 43:465; Clastrier, 1963, Arch. Inst. Pasteur Algerie, 4l:6o. (Type-species: Ceratopogon femoratus Meigen, mon.) Prionomyia Stephens, 1829, Cat. Brit. Inst., 2j237 (Type-species: Ceratopogon femoratus Meigen, desig. Westwood, 1840). Ceratolophus (part) Kieffer (not Barboza de Bocage, 1873), 1899, Bull. Soc. Ent. France, p. 69 (Type-species: Ceratopogon femoratus Meigen, mon.). Johannseniella V fnn 1907, Jour. N. Y. Ent. Soc., 15:1 (nom. nov. for Ceratolophus K., not Barboza de Bocage). DIAGNOSIS. Body slender and almost bare. Eyes bare. Female antennal segments 3-10 short, segments 11-15 cylindrical and elongated; male antennae plumose. Mesonotum covered with small bristles, humeral pits absent. First four legs slender, without spines; fourth segments of tarsi cordate, claws simple and almost equal; femora of hind legs distinctly swollen in both male and female, armed ventral 1 y with

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numerous spines, tibia slender and curved basally; fourth segments of tarsi cylindrical, female claws highly unequal, without empodium. Wings with microtrichiae, macrotrichiae present in some species; costa long; two subequal radial cells present; intercalary fork absent, madia sessile to short peti elate. Two spermathecae. Male genitalia: ninth sternite small; ninth tergite cone-shaped, with large anal lobes; basistyles stout, dististyles long and narrow; aedeagus small with long arms; parameres small and simple. Approximately 27 species have been described. Distribution: North America, Europe, Africa, Malaysia and Caroline Island. The larval and pupal stages of S. femorata were given in Kettle and Lawson (1952). LARVA. Large eyes. Two combs on epipharynx and each armed with a slender lateral plate; seven teeth on each half of dorsal; and ii teeth on ventral side. No pigment, only a pair of grey and clear latiral substances in meso thorax and metathorax. PUPA. Operculum thickly covered with nodules. Respiratory horn short, pale at base. Tip and distal area with eight or nine papillae. Metathorax concave. Arrangement of setae differs from Culicoides . Caudal spines dark at tips. Larval habitat; Strenzke (1950) reported finding the larvae of £. femorata in damp moss in Germany. Female claws resemble Heteromyia but the main point of difference is that the femur of hind leg is greatly swollen and armed, fore tibia arcuate. This character is found on fore leg of Heteromyia . Illustration: Fig. 58 Serromyia femorata (Mg.)

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83 Genus Macrurohelea Ingram and Macfie Macrurohelea Ingram and Macfie, 193 1> Dipt. Patag. and S. Chile, part H, fasc. 4:203; Lee, I962, Proc. Linn. Soc. N.W.W., 87: 339; Wirth, 1965, Pan-Pacif. Ent., kUk6, (Type-species: Macrurohelea caudata Ingram and Macfie, orig. desig.). DIAGNOSIS. Eyes -well separated, with fine pubescence. Palpi fivesegmented. Antennae with all segments short. Legs slender, without spines. Female claws small and equal. Wing without macrotrichia, microtrichia present; costa long; two radial cells present. Female abdomen with modified tenth segment, which curves forward and reaches the ventral side of the anterior segments. Spermathecae two or three. Male genitalia: with adeagus Y-shaped, parameres separated, long but larger at the distal portion, basal arms well developed. Approximately five species have been described. Distribution: South America and Australia. Ingram and Macfie, 1931, proposed this genus for two species, M. cauda ta I. & M. from southern Chile, and M. thoracica I. & M. from southern Argentina. The female of this genus is readily recognized by the uniquely modified abdomen, with the tenth segment long and bent forward and appressed to the venter of the preceding segment. The wing venation and male genitalia are similar to Stjlobezzia but differ in having equal tarsal claws. This character also separates them from Monohelea. Macrulohelea can be separated from Echinohelea Macfie and Serromyia Meigen by the absence of femoral spines; and from Parabezzia Malloch by the presence of two radial cells. Illustration: Fig. 59 Macrurohelea setosa Wirth

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Genus Parabezzia Mai loch Parabezzia Malloch. 1915, Bull. 111. St. Lab. Nat. Hist., 10:358; Johannsen, 19^3, Ann. Ent. Soc. Amer. 36:782; Wirth, 1952, Proc. Ent. Soc. Washington 5^:23; 1965, Proc. Ent. Soc. Washington 67:215. (Type-species: Parabezzia petiolata Malloch, orig. desig.) Diaphanobezzia Ingram and Macfie, 1931» Dipt. Patagonia & S. Chile, pt. 2, p. 223. (Type-species: Diaphanobezzia pellucida Ingram and Macfie, by orig. desig.). DIAGNOSIS. Body rather stout, almost bare. Eyes bare. Palpus foursegmented with one to six long sensillae, first and second segments fused. Female antennal segments 3-10 oval, segments 11-15 cylindrical and elongated; male antennae plumose. Mesonotum without any sign of pits or spine or tubercle, but bristles and small pubescence present. Legs slender with some bristles; coxae of male densely covered with bristles; tarsi sometimes with ventral spines; fourth segments of tarsi cordiform; fifth segments swollen and flattened on the lateral side, sometimes with tubercles on ventral side; without empodium; male claws small and equal but long, curved and may be unequal in female; no basal tooth. Wing of male narrower than female, macrotrie hia absent, micro trichia very fine; costa very long, extended beyond a single radial cell, costa shorter in male; median fork petiole. Spermathecae two. Male genitalia, aedeagus large, triangular, parameres small, fused to form triangular sclerite behind aedeagus. Approximately 18 species have been described.

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Distribution: North America, South America, Republic of Congo, Jamaica, and Puerto Rico. PUPA. Pupa with very short respiratory horn, oval shape, about five pairs of spiracles; caudal spines short and stout, pointed laterally. Adult habits: reared from sand bar (Wirth, V)6$c) Malloch (1915a) proposed the genus Parabezzia for three Nearctic species: P. petlolata n. sp. (male), Ceratopogon lnermis Coquillett, and Bezzia elegantulata Johannsen. He used their petiolate media of wing as a distinguishing character. It resembles Neurohelea in having long costa, extending beyond but differs in having one radial cell and four segmented palps. Illustration: Fig. 60 Parabezzia petlolata Mall. Genus Clinohelea Kieff er Cllnohelea Kieffer, 1917, Ann. Mus. Nat. Hung., 15:295, 316; 1919, Ann. Mus. Nat. Hung., 17:76; Goetghebuer, 1920, Mem. Mus. Roy. Hist. Nat. Belg., 8:76; Edwards, 1926, Trans. Ent. Soc. London, 74:413; Johannsen, 1931, Arch f. Hydrobiol., 9:435; Macfie, '.1934, Ruwenzori Exp., 351:91; 1939, Rev. Ent. Rio de J., 10:205; Johannsen, 1943, Ann. Ent. Soc. Amer., 36:783; Lane, 1944, Rev. Ent. Rio. de J., 15:249; Goetghebuer, 1948, Explor. Pare. Nat. Albert. Miss, de Witte fasc. 55:17; Lee, 1948, Proc. Linn. Soc. N.S.W., 73:64; Wirth, 1951, Proc. Ent. Soc. Wash., 53:321; 1952, Univ. Calif. Publ. Ent., 9:209; Lane & Durat, 195 2 *, Dusenia, 5:247; de Meillon, 1959, Novos Taxa Ent. no. 13:20 (Type-species: Ceratopogon variegatu3 Winnertz, orig. desig.)

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DIAGNOSIS. Eyes bare, widely separated. Palpi simple. Female antennal segments 3-10 long, segments 11-15 cylindrical. Humeral pits absent, anterior spine very. short. Femora slender, without spines; fourth tarsal segment of at least four posterior legs strongly bilobed and each lobe terminates in a stout spine, fifth tarsal segment of fore legs greatly swollen in both male and female; claws of four posterior legs very unequal in female. Wing long, without macrotrichia costa long; two radial cells well developed; intercalary fork absent; media widely sessile. Gland rods absent. Spermathecae two. Male genitalia: ninth sternite short; ninth tergite tapered; basistyles slender; dististyles very long and slender; aedeagus arched, median portion large; parameres slender, rounded tip and sometimes fused at particular point. Approximately 26 species have been described. Distribution: World-wide. For the immature stages of Cllnohelea f Wirth (195 Id) published on the pupa. One species was collected from the sandy edge of a stream at Falls Church, Virginia. PUPA. Light brown; long and slender respiratory organ with 10 spiracles at apex; operculum as long as broad, with a pair of round tubercles each with a long seta, surface especially near distal areas .and between the tubercles with shagreen body surface appearance with crescent-shaped tubercles, sharp and setose at lateral margins; anal segment long with fine shagreening, the apicolateral processes well developed with sclerotized tip, subapical shagreening.

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This genus differs from Palpomyia and Tetrabezzia in the following characters: fifth tarsomeres of fore legs swollen in both sexes; fourth tarsomeres of the four posterior legs of female highly bilobed, each with a stout spine; claws of four posterior legs of female very unequal. The other characters which readily differentiate them from Tetrabezzia are: hind leg not unusually long; claws unequal only on four posterior legs; wing with two radial cells. Illustration: Fig. 6l Clinohelea bimaculata (Loew) Genus Tetrabezzia Kieffer Tetrabezzia Kieffer, 1917, Ann. His. Hung., 15:296; Ingram and Macfie, 1923, Bull. Ent. Res., 14:70; Macfie, 1940, Ann. Trop. Med. Parasit., 34:24; de Meillon, I96I, Rev. Ent. Mozambique, 4:55; Wirth, 1962, Ann. Ent. Soc. Amer., 55:274. (Type-species: Tetrabezzia spinigera Kieffer, mon. ) DIAGNOSIS. Large size, body slender and almost bare. Eyes bare, separated. Female antennae very long and slender with very long hairs on segments 3-10; palpi slender, third segment simple. Mesonotum with anterior spine. Legs long, hind legs longest, hind femur enlarged near tip, fourth tarsomeres with bifid spinose lobes on fore and mid legs, but long and cylindrical on hind legs; bifid lobes of male fourth tarsomeres not as distinct as in female; claws very unequal on female, almost subequal on male; fore femur armed ventral 1 y. Abdomen petiolate, two spermathecae, gonopore without hair tufts. Wing infuscate, macrotrichia absent, microtrichia fine, costa almost

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reaches the wing tip. Male genitalia with ninth sternite short, ninth tergite large, basis tyles and dististyles long and slender, aedeagus arch, triangular shape, parameres clubtipped. Approximately four species have been described. Distribution: Africa, Malaya, Thailand. This genus resembles Clinohelea but the chief points of difference are as follows: female hind leg unusually long; fourth tarsomere of four anterior legs with bifid spinose lobes; all female claws very unequal; wing with one radial cell; abdomen petiolate. Illustration: Fig. 62 Tetrabezzia pictipennis (K.) Genus Heteromyia Say Heteromyia Say, 1825, Amer. Ent., 2:79; RLeffer, 1906, Gen. Ihsectorum, fasc., 42:64; Edwards, 1926, Trans. Ent. Soc. London :420; 1935, Konowia, 12:8?; Macfie, 1940, Ann. Trop. Med. Parasit. 34:27; Johannsen, 1943, Ann. Ent. Soc. Amer., 36:774; Lane, 1946, Rev. Ent. Rio de J., 17:211; Lee, 1948, Proc. Linn. Soc. N.S.W., 73:61; Duret and Lane, 1955, Dusenia, 6:35; Lane, 1961, Rev. Bra. Biol., 21:4l, Wirth, I962, Ann. Ent. Soc. Amer., 55:274. (Type-species: Heteromyia fasclata Say, mon.). Pachyleptus Walker. I856, Insecta Saundersiana, Diptera. London :426. DIAGNOSIS. Large size, body slender with few hairs. Eyes bare, broadly separated. Female antennal segments 3-10 long, segments 11-15 very long and dark; male antennae with inconspicuous plumes. Palpi long and slender, third segment simple. Mesonotum dark; with snail anterior spine. All legs different, femora of fore legs greatly swollen and armed ventrally with spines, fore tibiae arched along

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the femora; ndd legs long; hind legs of female very long, tip of hind femora club-shaped; fourth tarsal segments on all legs of male and on the anterior four of female cordate, but long and slender on hind legs of female. Hind claws of female single, very long and barbed basally; the others including all male claws equal. Wing fasciate, with one or two radial cells; costa long. Male genitalia: ninth sternite short, ninth tergite tapering with well developed apicolateral process; basistyles and dististyles simple; aedeagus arched with large body; parameres club-tipped and lateral arms distinct. Approximately 20 species have been described. Distribution: North America, South America, Asia, and Australia. The study of the immature stages was made by Thomson (1937)« One pupa was found from wet algae on the cliffs near Ithaca, New. York. PUPA. Respiratory trumpet flat and short, constricted at base, apex bearing 11 spiracles; operculum with two setae and also with small hooks scattered on the surface; abdominal segments with three mid dorsal sclerotized areas, the surface beside these areas covered with small round hooks; last segments with two projections. This genus is closely related to Palpomyia and some authors have misplaced them in Palpomyia . It is differentiated from Palpomyia by the following characters: fore femur greatly swollen and armed ventrally with spines; fore tibia arcuate to the shape of the femur; hind leg of female very long with a single barbed claw; fourth tarsal segment on all legs of male and anterior four of female cordate, with equal claws; wing fasciate. In Serromyia , hind femur is greatly swollen and armed and tibia also arcuate as in fore leg of Heteromvia .

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Illustration: Fig. 63 Heteromyia fasclata Say, Heteromyia sp. Genus Pellucidomyia Macfie Pellucidomyia Macfie, 1939, Ruwenzori Exped. , | 193^-5, 1(5) :99; Wirth, i960, Bull. Brooklyn Ent. Soc, 55:1 (Type-species: Pellucidomyia ugandae Macfie, mon.) Macfiehelea Lane, 19^6, Rev. de Ent. , 17:208 (Type-species: Macfiehelea oliveirai Lane, Mon.); Lane, 1956, Rev. Brasil. Biol. 16:435 (key to 3 Neotropical spp.) I DIAGNOSIS. Dorsal side of the body eoveped thickly pollinose with dark and white areas. Head with anterior and posterior portions flattened; eyes widely separated; antennae usually elongated; palpi five-segmented, all segments simple, third segment with sensory pit absent. Mesonotum without spine or tubercle but with anterior cone. Legs slender, without spines in all femora, distal part almost clublike; hind legs longest; fourth tarsal segment cylindrical and elongate, but cordiform to transverse on the first fore legs; fifth segment unarmed, simple on mid and hind legs, enlarged on fore legs; claws on fore and mid legs short and equal, hind leg with single and long claw. Wing without macrotrichia or microtrichia; one radial cell present; costa long. Abdomen with two tufts of long hairs on ventral side of the eighth segment; spermathecae two; abdomen slightly curved downward. Male with all claws equal, hind leg simple not as long as in female; wing long and narrow; antennae with few plumes; male genitalia with ninth sternite small, ninth tergito tapered, basistyles simple, dististyles rather stout with curved, pointed tips, aodoagus archod sides almost parallel, parameres separated and reduced.

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91 Approximately five species have been described. Distribution: Panama, Brazil, Uganda, Australia, Colombia, and Mexico. The character of the unarmed fifth tarsomere and unusually long hind leg closely relate this genus to the genera Clinohelea Kieffer, Tetrabezzia Kieffer, and Heteromyia Say. It is readily separated from Heteromyia and Tetrabezzia by having fore femora unarmed. It differs from Clinohelea particularly in having a cordif orm to cylindri cal fourth tarsomere not strongly bilobed as in Clinohelea . The wing venation is similar to Neurohelea and Parabezzia with the long costa extending well beyond while Bezzia has similar venation without extension of costa, all these genera have one radial cell, fivesegmented palpi. Illustration: Fig. 64 Pellucidomyia sp. Trans. Ent. Soc. London, ?4:413; Ingram and Macfie, 1931, Dipt. Pat. & S. Chile, pt. 2, fasc. 4:212; Johannsen, 1943, Ann. Ent. Soc. Amer., 36:783; Wirth, 1952, Univ. Calif. Publ. Ent., 9:208. (Type-species: Ceratopogon luteitarsls Meigen, mon.) DIAGNOSIS. Moderate size, body almost bare. Eyes bare and separated. Female antennal segments 3-10 oval with sparse verticils, segments 11-15 cylindrical and elongated. Male antennae with few plumes; palpi slender, third segment not swollen. Mesonotum without anterior tubercle. All femora slender and unarmed; fifth tarsal segment of Genus Neurohelea Kieffer Neurohelea Kieffer, 1925, Faune de France, 11:112; Edwards, 1926,

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fore tarsi swollen in both sexes. Wings moderately broad; macrotrichia absent; costa extending much beyond R^; two well developed radial cells; intercalary fork absent; median fork barely sessile or with very short stem. Female abdomen without eversible glands or gland rods. Two spermathecae. Male genitalia: ninth stemite small, transverse, with continuous spicuate membrane from ventral face of aedeagus to apex; ninth tergite narrow, tapering with a pair of well developed setose lobes; basistyles and dististyles stout; aedeagus a sclerotized arch with rounded apex and parallel sides; parameres with a pair of flattened plates distally. Approximately five species have been described. Distribution: North America and South America. Characters are as in Palpomyia with these exceptions: anterior tubercle absent; all femora slender and unarmed; last segment of fore tarsi swollen in both sexes; costa extended beyond R^; median fork barely sessile or with very short stem; female abdomen without eversible glands. In Parabezzia wing with costa extended beyond R^ as in Neurohelea but only one radial cell present; fifth tarsomere swollen. Neurohelea with two radial cells on wing. i Illustration: Fig. 65 Neurohela nigra Wirth I i

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Genus Johannsenomvla Malloch Johannsenomyia Malloch, 1915, Bull. HI. State Lab. Nat. Hist., 10:332; Qoetghebuer, 1920, Mem. Mus. Roy. Hist. Nat. Belg., 8:73; 1921, Mem. Mus. Belg. 8 fasc. 4:182; Edwards, 1926, Trans. Ent. Soc. London, 74:413; Ingram and Macfie, 1931, Dipt. Patag. & S. Chile, pt. 2:231; Johannsen, 1931, Arch. f. Hydrobiol., Suppl., 9:^35; Goetghebuer, 1933, Rev. Zool. Bot. Afr. 24:149; 1938, Bull. Arm. Soc. Ent. Belg., 78:380; Macfie, 1940, Proc. Roy. Ent. Soc. London, (B) 9:75; Johannsen, 1943, Ann. Ent. Soc. Amer., 36:784; Wirth, 1952, Univ. Calif. Publ. Ent., 9:210; Clastrier, I962, Arch. Inst. Pasteur Alger., 40:282; Wirth, 1962, Ann. Ent. Soc. Amer., 55:276; 1965, Proc. Ent. Soc. Wash., 67:4. (Type-species: Johannsenomyia halteralis Malloch, designated in Wirth, 1952, Univ. Calif. Publ. Ent., 9:211 = argentata (Loew). Dicrohelea KLeffer, 1917, Ann. Mus. Natl. Hung., 15:363. (Type-species: Palpomyia filicomis Kieffer (des. Macfie, 1940, Ann. Trop. Med. Parasitol., 34:26). DIAGNOSIS. Body slender, almost bare. Eyes bare and contiguous below. Female antennae elongated, male antennae plumose. Palpi slender. Mesonotum with anterior tubercle; with microtubercles bearing short setae over the surface. Legs long and quite slender; femora especially of hind legs enlarged near apexes; no spines on femora or tibia; fourth tarsal segment simple; fifth segment with very blunt batonnets on the ventral side, numerous in female but only two to three pairs in male. Female claws highly unequal on four posterior legs, fore legs with equal claws; all claws each with external basal tooth; male claws simple, inconspicuously bifid at tip.

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9 l> Wings with short costa, two radial cells. Abdomen narrower near base, hair tufts present near gonopore; spermathecae two. Male genitalia rotated; ninth tergite rounded with strong bristles near posterior margin; aedeagus arched, body large, with caplike tip; parameres paired with large platelike tip. Approximately 18 species have been described. Distribution: North America, South America, Europe, Asia, and Africa. Information on the immature stages is presented in Malloch (1914), Thomson (1937), Johannsen (1952), Wirth (1952), Wirth and Stone (1956) and Wirth (1962a). j EGG. Thomson (1937) indicated that eggs of two unknown species and J. argentata (Lw.) were arranged in more or less spiral bands, eggs of the latter were found at the edge of a pond in New York. LARVA. The first ins tar larva has a long, conical head and anal segment with four long hairs and four slightly shorter hairs. PUPA. The respiratory horn is very short; abdominal tubercles are small, not conical or angulate; grandular discs on abdominal sterna are absent (Wirth, 1962a). The pupa of J. sybleae was described by Wirth (1952); it was taken from a cattail-choked pond near the beach. Malloch (191^a) also described the pupa of J. flavidula taken from the Illinois River. Williston (1907) proposed the new name J ohanns enlella to replace the former genus Ceratolophus Kieffer (1899) which was preoccupied by Ceratolophus Barbosa de Bocage (I873). Malldch (1915a) noticed that the genotype, Ceratopofton femoratus (Fabricius) was also being used as the genotype of Serromyia Meigen (1819). In this new revision,

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he divided the species into two groups under two new genera. Johanns enoiwi a was proposed for those with media sessile. The type species was not designated until Wirth (1952) utilized the characters of the allotype female and designated Malloch's last species J. halteralis (male) as genotype. After careful study, Wirth (1962a) suggested that this genus must have a new name since Malloch (1915) had misplaced the female of the type species halteralis in a different genus. Johannsenomyia is related to Jenkins helea. Mallochohelea . Nilobezzia. Sphaeromias. Xenohelea. Probezzia. and CalyptopoKon in having batonnets on fifth tarsomere but the following distinctive characters of the genus can be differentiated from the others: hind leg enlarged especially near knee; femora unarmed; claws highly unequal on four posterior legs, each claw with external basal tooth; ventral hair tufts present on eight abdominal segment and wing simple with two radial cells. Illustration: Fig. 66 Johannsenomyia argentata (Loew) Genus Xenohelea Kief f er Xenohelea Kieffer, 1917, Ann. Mus. Nat. Hong., 15:295; Macfie, 1934, Tijdschr. Ent., 77:226; Macfie, 19*K), Ann. Trop. Med. Parasit., 3*1:28; de Meillon, 19^2, Estac. anti-Malar. Lourenco Marques :23; Lee, 1948, Proc. Linn. Soc. N. S. W., 73:^5; Mayer, 1957, Dtsch. Ent. Z. (N. F.) f 4:31. (Type-species: Xenohelea pruinosa ______ Kieffer, orig. desig.) Mixohelea Kieffer (1917).

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DIAGNOSIS. Body slender, almost bare. Eyes bare, nearly contiguous. Palpi slender, all segments simple. Female antennal segments 3-10 oval, segments 11-15 very long. Mesonotum sometimes with a spine at mid anterior portion. Wing without macrotrichia, costa long, two well developed radial cells present. Legs slender, hind legs slightly longer than the preceding legs, femora sometimes with a few spines; fourth tarsal segments cylindrical; fifth segments armed with very stout, distally bent batonnets. Female with single long claw with basal barb on all legs. Male unknown. Approximately six species have been described. Distribution: Africa, Asia, and Australia. PUPA. Mayer ( 1957) described the pupa of Xenohelea galatea de Meillon: prothoracic horn with about nine spiracles; operculum absent; two long setae on prothoracic horn; three to seven abdominal segments with most tubercles surrounded by a round chitin edge; exuviae dark. This genus has most of the characters of Palpomyia but all claws are unequal and simple and the fifth tarsomere is armed with batonnets. It is particularly differentiated from Jenkinshelea . Johannsenomyia. Mallochohelea . Nilobezzia and Sphaeromias. those genera bearing batonnets on fifth tarsomere but the batonnets of Xenohelea are located only on the anterior half of the segment. Illustration* Fig. 67 Xenohelea tonnoiri Lee

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97 Genus Jenkinshelea Macf ie Jenkinshelea Macfie, 193^, Ann. Trop. Med. Parasit., 28:177; de Meillon, 1936, Pub. S. Afr. Inst. Med. Res., 38:189; 1937, Ann. S. Afr. Mus. 32:261; Johannsen, 19^2, Ent. News, 53:76; de Meillon, 19^2, Proc. Rhod. Sci. Ass., 39: 116; 1943, J. Ent. Soc. Sthn. Afr., 6:109; Johannsen, 19*4-3, Ann. Ent. Soc. Amer., 3^:774; Mayer, 1952, Beitr. Ent., 2:584; Wirth, 1962, Bull. Brooklyn Ent. Soc, 57:2 (Typespecies: Jenkinshelea setosipennis (Kieffer). DIAGNOSIS. Body slender. Eyes broadly separated. Male antennae with few plumes. Mesonotum with fine whitish gray pubescence. Anterior portion of thorax somewhat jutting over the head but not pointed. Legs slender, femora unarmed; fourth tarsal segment wider than long, cordate or bell-shaped; fifth tarsomere with strong ventral batonnets; tarsal claws equal on all legs, each with external basal tooth. Wing with unusually broad anal lobe, one or two radial cells, costa extended mostly to wing tip, medial fork broadly sessile, pos terior branch of media not elbowed in female; microtrichia present. Abdomen of female, eighth segment with sclerotization and pair of ventral hair tufts. Male genitalia, aedeagus stout, paramere very Approximately six species have been described. Distribution: North America, South Africa, and India. PUPA. Large, respiratory trumpet very small and oval-shaped with with about 60 degree apart from each other; abdomen with glandular discs present. slender, separated at tip hooklike. about 10 pairs of spiracles, caudal spines long, pointed posteriorly

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Kieffer (1913a) erected the new genus Jenklnsia for a species of midge taken in Calcutta. Since this name had been previously I occupied by a genus of fishes, Macfie (193**) suggested the name Jenkins helea to substitute for it. This genus resembles Johannsenomyia but can be readily separated from it in the characters of thorax and wing. The thorax of JenkLnshelea is somewhat projecting over the head and wing with enlarged anal lobe. Illustration: Fig. 68 Jenkins helea magnipennls (Joh.) Genus Probezzia Kieffer Probezzia Kieffer, 1906, Gen. Insectorum, fasc. b2i$7\ Coquillett, 1910, Proc. U. S. Nat. Mus., 37:59^; Malloch (part), 191^, Proc. Biol. Soc. Wash., 27:137; Malloch (part), 1915, Bull. 111. St. Lab. Nat. Hist., 10:352; de Meillon, 19^+2, Estac. anti-Malar. Lourenco Marques J319; Wirth, 1951, Proc.jEnt. Soc. Wash., 53:27? (Type-species: Ceratopogon venustus Meigen, desig. Coquillett, 1910). Dlcrobezzia Kieffer, 1919, Ann. Mus. Nat. Hungariei, 17:127; Ingram & Macfie, 1921, Ann. Trop. Med. and Parasitol., 15071; Edwards, 1926, Trans. Ent. Soc. London, 7^1^; Johannsen, 19^3, Ann. Ent. Soc. Amer., 36:785; Lane, 19^7, Arq. Fac. Hig. S. Pub. U. Sao Paulo, 1:232. (Type-species: Ceratopogon venustus Meigen, orig. desig.). I DIAGNOSIS. Body slender and almost bare. Eyes bare and separated. Female antenna! segments 3-10 short, globose, segments 11-15 long;

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99 i , male antennae with few pluses • Mesonotum without anterior spine or tubercle; short bristles scattered on the surface. Legs slender, femora unarmed; fourth tarsomere broad and short; female fifth tarsomere with ventral batonnets; claws with blunt outer basal tooth. Wing usually milky white with obscure broad infuscated band; single radial cell well developed, vein R2 missing; costa long; media widely sessile. Female abdomen with a pair of ventral hair tufts on eighth segment. Spermathecae two. Male genitalia: ninth stemite narrow and highly concave; ninth tergite long and tapering distally, lateral lobes well developed; basis tyles elongate but dististyles small; aedeagus arched, tapered, ending in caplike tip; parameres fused with bifid tip. Approximately 36 species have been described. Distribution: World-wide. Little information concerning the immature stages of Probezzia have been reported. The following statements are taken from Thomson (1937). EGG. Eggs of Probezzia glabra (CoqO are long, narrow and arranged almost vertical in the mass. LARVA. Larvae are similar to those of Bezzia , the head is oblong, three to four times as long as wide; anal hairs are varied in length. Flap-like structure inside of cast skin is absent, PUPA. Pupae of Bezzia and Probezzia can be separated by the number of setae on operculum. They are also similar to those of Palpomyia and Heteromyia . Differences are in numbers of setae on operculum and numbers of respiratory spiracles on the trumpet. According to

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100 Mayer ( 1937) » the pupae of this' genus cannot be separated from those of Johannsenomyia, This genus resembles Bezzia by having slender unarmed femora; fifth tarsomere of female with ventral batonnets. Males differ from Bezzia males by the great prolongation of the radial cell. The presence of a pair of ventral hair tufts on the eighth abdominal segment is similar to Nilobezzia , Jenkins helea, Johannsenomyia and Mallochohelea . The presence of only one radial cell and simple anal lobe readily separates the genus from Jenkins helea, Johannsenomyia and Mall ochohelea . The male genitalia -with well developed claspers differentiates Probezzia from Nilobezzia . Illustration: Fig. 69 Probezzia pallida Mall och Genus Calyptopogon Kieffer Calvptopogon Kieffer, 1910, Mem. Ind. Mus., 2:209; Johannsen, 1927, Ent. Mitt., 16:423; Seguy, 1931, Bull. Soc. Ent. France, :208 1934, Mem. Acad. Cienc. Zaragoza, 3*10; de Meillon, 1936, Pub. S. Afr. Inst. Med. Res., 38:204; 1937, Pub. S. Afr. Inst. Med. Res., 40:351; 1938, Proc. R. Ent. Soc. Lond., (B) 7:270; Macfie, 1939, Trans. R. Ent. Soc. Lond., 89:2; de Meillon, 19^, Trans. R. Ent. Soc. Lond., 90:463; 1942, Proc. Rhod. Sci. Ass., 39:116; Mayer, 1952, Beitr. Ent., 2:584; Lane, 1961, Rev. Bras. Biol., 21 37. (Type-species: Calyptopogon albitarsis Kieffer, mon.). Haaslella Kieffer (1913), Macroptilum Becker (1903), Macropeza Meigen (1818).

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DIAGNOSIS. Body large, very slender. Eyes bare, videly separated. Palpi short, rather pale; mandible teeth large. Female antennae greatly elongated. Male antennae with few plumes. Thorax conical shaped in front. Legs slender; first femora sometimes club-shaped distally; no spines on femora or tibiae; hind legs longest, hind femora and tibiae very long and slender; fourth and fifth tarsal segments of fore legs swollen; tarsomeres of mid and hind legs slender, fifth segment with three pairs of ventral batonnets; two pairs on fore legs; male fifth tarsomere without batonnets, claws small and equal, each with external basal tooth; male claws simple with cleft tip. Wing with one narrow radial cell, costa very long extended beyond R c , crossvein r-m very long and causing cell R5 to appear broad. 5 Abdomen elongated; eighth abdominal segment bearing two ventral hair tufts; two spermatheca. Male genitalia: basistyles slender, dististyles tapering; ninth sternite narrow and highly concave; ninth tergite tapering, lateral lobes well developed; aedeagus arched; parameres fused at base, tips elongated and slender. Approximately 20 species have been described. Distribution: South America, Eruope, Asia, and Africa. This genus is related to Stenoxenus but differs in the following points: hind leg very long; fifth tarsomere bearing batonnets; r-m crossvein long; vein M 2 not elbowed, thorax conical in front and ventral hair tufts present. It resembles Tetrabezzia , Heteromyia , Pellucidomyia , and Clinohelea in having the unusually long hind leg, but differs in having batonnets on fifth tarsomeres.

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The wing with eosta prolonged beyond Rj is similar to those of Neurobezzia and Parabezzia but the radial cell of Stenoxenus is very narrow and cell R«j is wide. Illustration: Fig. 70 Calyptopogon gibbosus (Wied) Genus Sphaeromias Curtis Sphaeromlas (Stephens Ms, sine descr.) Curtis, 1829, Brit, Ent., 6:285; Kieffer, 1906, Gen. Insectorum, fasc. 42:62; Edwards, 1926, Trans. Ent. Soc. London, 74:415; Johannsen, 1931* Arch. f. Hydrobiol., Suppl., 9:436; 1943, Ann. Ent. Soc. Amer., 36:784; Wlrth, 1952, Univ. Calif. Publ. Ent., 9:214; Freeman, 195^, Arch. Hydrobiol. 48:445; de Meillon, I96I, Rev. Ent. Mozambique, 4:54. (Type-species: Sphaeromias albomarginatus Curtis (mon.) = fasciatus (Meigen). Xylocrypta Kieffer, 1899, Bull. Ent. Soc. France, p. 69; 1917» Ann. Mus. Nat. Hung., 15:295 (Type-species: Ceratopogon fasciatus Meigen, mon.). Schizodactylus Ingram and Mac fie, 1921, Ann. Trop. Med. & Parasit. , 15:353 (Type-species: Schlz odac tylus telmatoscopus Ingram and Macfie, mon.) Ankistrodactylus Ingram and Macfie, 1922, Ann. Trop. Med. & Parasit., 16:272 (new name for Schizodactylus Ingram and Macfie, not Schlzodactyla Claus, 1876, Coelenterata.)

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DIAGNOSIS. Body large, very stout, almost bare. Eyes bare and very narrowly separated. Antenna long, almost bare, male antenna with few plumes. Palpi slender. Mesonotum densely pollinose, anterior tubercle poorly developed. Legs slender with numerous spines on ventral side of femora and dorsal side of tibiae. Fourth tarsal segment simple; fifth segment with numerous ventral blunt dark spines or batonnets; claws large, equal, each with long inner basal tooth. Wing with long costa, radial cells two. Spermathecae two. Male genitalia: basistyles and dististyles elongate; aedeagus arched, broad tip; par aire res fused, lateral arms developed, distal part club-shaped. Approximately 40 species have been described. Distribution: North America, South Americaj Europe, Africa, and Asia. PUPA. Respiratory trumpet long, apex wide and rounded, tapering basally; abdominal tubercle conspicuous, conical and pointed; abdomen with ventral glandular discs absent; caudal spines slender. Members of this genus have the characters of Palpomyia , with the following exceptions: eyes very narrowly separated; antennae very short; anterior tubercle poorly developed or absent; femoral spines more numerous but smaller and inconspicuous; fourth tarsomere short, cylindrical; fifth tarsomere bearing batonnets, claws large and male antennae with very few plumes. Subgenus Sphaeromias s. str. Fifth tarsomere with three or more pairs of batonnets along the length of the segment. Illustration: Fig. 71 Sphaeromias ( Sphaeromias ) lonrf.pennis (Loew)

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10^ Subgenus Homohelea Kieffer, 1917 Subgenus type: Homohelea abjure ta (Kieffer)* Fore femora slightly swollen; femora armed; tibiae unarmed, mid tibia sometimes with an apical ventral spine; fourth tarsomere slender oval; fifth segment with one or two pairs of basal batonnets. Ulustrationj Fig. 72 Sphaeromias ( Homohelea ) barkudensis (Edwards) <

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105 Key for the Subgenera of Sphaeromlaa 1. Fifth tarsomere with few batonnets located on the basal half of segment (Fig. 72Q) Homohelea (Fig. 72) . Fifth tarsomere with numerous batonnets along the segment (Fig. 71K) Sphaeromias (Fig. 71)

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106 Genus Mallochohelea Wirth Mallochohelea Wirth. 1962, Ann. Ent. Soc. Amer., 55: 278. Johannsenomyia Malloch, ndsident.; Johannsen, 19^3, Ann. Ent. Soc. Amer., 36:784; Wirth, 1952, Univ. Calif. Publ. Ent., 9:210; Johannsen, 1952, Connecticut State Geol. Nat. Hist. Surv. Bull., 80:164. (type-species: Johannsenomyia albibasis Malloch, orig. desig.). \ 1 1 DIAGNOSIS. Body slender or fairly stout, almost bare, integument shining. Eyes bare, broadly separated. Palpi slender; mandible teeth large. Female antennae rather long. Male antennae with few plumes. Anterior spine, and tubercle on mesonotum absent. Legs slender or enlarged; femora sometimes with few ventral spines; tibiae without spines; fourth tarsal segment simple; fifth segment with five to eight pairs of ventral batonnets; claws long and equal, each with blunt external basal tooth. Male claws simple with tip slightly cleft. Wing with two radial cells. Abdomen tapering and elongated posteriorly; eighth segment bearing two ventral hair tufts; spermathecae two. Male genitalia: basis tyles elongated, dististyles slender and short, aedeagus arched, stout body, distal part tapering ending in caplike tip; parameres fused at base, separated anteriorly into a pair of long processes. Approximately 21 species have been described. Distribution: World-wide. PUPA. Respiratory trumpet very short, about twice as long as broad; abdominal segments with conspicuous spinose tubercles, project posteriorly; sixth and seventh sternal segments with membranous glandular discs. (Wirth, 1962)

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Wirth (1962a) erected this genua for the majority of Johannsenomyia which were mlsidentified by Malloch (1915). Mallochohelea is closely related to Johannsenomyia and Jenkinshelea by the following characters: claws with external basal tooth; abdomen with a pair of ventral hair tufts on eighth segment, and wing with two radial cells. It is distinguished from Johannsenomyia by the equal tarsal claws in the female, absence of batonnets on the male fifth tars one re, and absence of anterior tubercle. The wing with simple anal lobe can be readily separated from the wing of JenkinThis genus also resembles Nilobezzla but the male genitalia with well developed claspers distinctly separates them. Illustration: Fig. 73 Mallochohelea albibasis (MallO Genus Nilobezzia Kieffer Nilobezzla Kieffer, 1921, Ann. Soc. Ent. France, 90:24; 1922, Ann. Soc. Linn. Lyon, 68:161; Edwards, 1929, Notul. Ent., 9:12; Macfie, 1939, Ruwenzori Exp., 1934-35, 1:101; de Meillon, 1943, J. Ent. Soc. Sthn. Afr., 6:111; Lee, 1948, Proc. Linn. Soc. N.S.W., 72:343; de Meillon and Hardy, 1954, J. Ent. Soc. S. Afr., 17*70; Clastrier, 1958, Arch. Inst. Pasteur Alger., 36:498; Sen & Das Gupta, 1958, Bull. Calcutta Sen. Trop. Med., 6:69; Lane, I96I, Stud. Ent. (N.S.), 4:449; Clastrier and Wirth, 1961, Arch. Inst. Pasteur Alger., 39:212; Clastrier, 1962, Arch. Inst. Pasteur Alger,, 40:121; Tokunaga, 1962, Pacif. Ins., 4:216; Wirth, 1962, Ann. Ent. Soc. Amer., 55:284; Vattier and Adam, 1966, Ann.

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Speleol., 21: 7//, (type-species: Nllobezzia armata Kieffer f mon. ) • DIAGNOSIS. Body very stout, almost bare. Eyes bare, very narrowly separated. Antennae not elongated; male antennae with few plumes. Palpi slender; mandible with large teeth. Thorax densely pollinose, anterior tubercle absent. Legs slender; femora fairly stout, with stout bristles scattered on ventral side, tibia with bristles on dorsal side. Fourth tarsal segments simple; fifth segment with numerous ventral batonnets, claws large and equal, each with blunt external basal tooth. Wing with one or two radial cells; costa short. Abdomen with a pair of ventral hair tufts on eighth segment; spermathecae two. Male genitalia: basistyle and dististyle barely developed and fused forming a cone-like process; aedeagus with slender arch and caplike distal portion; parameres fused mesally, tip slender. Approximately 26 species have been described. Distribution: North America, South America, Africa, and Asia. PUPA. Respiratory trumpet short, about three times longer than the distal portion; abdominal tubercles distinctly large, conical, pointed and directed caudad; glandular discs present on abdominal sterna five, six and seven in schwarzll (Coq.), but absent in minor (Wirth). (Wirth, 1962) This genus is similar to Jenkinshelea . Mallochohelea and Johannsenomyia in having an external basal tooth on claws; eighth abdominal segment bearing a pair of ventral hair tufts. But it is readily separated from those three genera by the character of armed femora (subgenus Nllobezzia) which resembles Bezzla . However, the fourth t arsons re in all Nllobezzia is subcylindrical and the spines

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109 I on femora are not as well developed as in Bezzia. The male genitalia with reduced claspers is the unique character of Nilobezzia. subgenus Nilobezzia s. str. Femora armed ventrally with black spines. Illustration: Fig. 7k Nilobezzia ( Nilobezzia) schwarzii (Coq.) Subgenus Parrotia KLeffer, 1924 Subgenus type: Parrotia flaviventris Kieffer Femora unarmed.

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110 Key for the Subgenera of Nilobezzia 1. Femora armed ventrally with black spines (Fig. 7^ F) Nilobezzia (Fig. 7*0 Femora unarmed EaECB&i

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Ill Genus Stenoxenus CoquHlett . Stenoxenus Coquillett, 1899, Ent. News Philad., 10:61; Johannsen, — — — — i, 1927, Ent. News, 38:70; Seguy, 1931, Bull. Soc. Ent. Fr., p209; Macfie, 193^-, Ann. Trop. Med. Parasit., 28:290; 1939, Rev. Ent. Rio de J., 10:138; 19^, Ann. Trop. Med. Parasit., 3^:28; Johannsen, 19^+3, Ann. Ent. Soc. Amer., 36:77^; Lane, 1956, Rev. Bras. Biol., 16:305; de Meillon, 1959* S. Afr. Anim. Life, 6:355; Lane, 1961, Stud. Ent. (N.S.), 4:^52; Vattier and Adam, 1966, Ann. Speleol. 21:7//. (Type-species: Stenoxenus iohnsoni Coquillett, mon.). DIAGNOSIS. Moderate size, slender, dorsoventrally flattened. Eyes bare, widely separated. Female antennal segments 3-10 with stout setose hairs, segments 1^-15 elongated with few hairs; male antennae with very few plumes. Palpi four-segmented, third segment slender. Mesonotum broad and round anteriorly without anterior tubercle. Femora unarmed, hind femur and tibia long and slender, vestiture with very fine erect hairs; first and second tarsomeres of hind leg with ventral tubercles; fourth tarsal segment spoon-shaped or subcylindrical; fifth segment unarmed; claws small and equal. Wing with r-m crossvein very short, causing the cell between vein R and M to appear narrow; costa long extended to the tip of wing; on radial cell media sessile, in male forking at level of crossvein; elbowed in female. Female abdomen with pairs of eversible glands and gland rods Male genitalia: ninth stemite small, ninth tergite short with well developed lateral lobes; basistyles slender, dististyles elongated; aedeagus with slender arch, pointed tip; parameres fused with broad rounded tip.

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Approximately 12 species have been described. Distribution: North America, South America, Africa, Malaysia, and Tonkin. Related to Calyptopogon with the following exceptions: hind leg simple, not very long; fifth tarsomere without batonnets, wing with vein elbowed, r-m crossvein short; thorax not conical in front. It also resembles Bezzia in the presence of gland rods on the female abdomen and the male parameres fused in one long lobe but it can be separated from Bezzia by the above mentioned characters. The female of this genus is readily recognized by the wing with vein Mg elbowed, this character is not found in other ceratopogonids.. Illustration: Fig. 75 Stenoxenus insigninervis Macfie Genus Paryphoconus Enderlein Paryphoconus Enderlein, 1912, Stett. Ent. Zeitg., 73*57; Kieffer, 1917 Ann. Mas. Nat. Hang., 15:332; Macfie, 1939» Trans. Roy. Ent. Soc. London, 89(1) :1; 1939, Rev. Ent., 10:142; 1940, Ann. Trop. Med. Parasit., 34:24; Johannsen, 1943, Ann. Ent. Soc. Amer., 36:773; Lane, 1946, Rev. Ent. Rio. de J., 17:203; Mayer, 1952, Beitr. Ent., 2:583; Lane, 1956, Rev. Bras. Biol., 16:299; Wirth, 1959, Dtsch. Ent. Z. (N.F.), 6:236; Mayer, 1959, Dtsch. Ent. Z. (N.F.), 6:232; Lane, 1961, Stud. Ent. (N.S.), 4:453. (Type-species: Paryphoconus angustipennis Enderlein . orig. desig.). DIAGNOSIS. Body slender, dorsoventrally flattened. Eyes bare, widely separated. Female antennae much elongated, segments 1115 with dense setose hairs; male antennae with few plumes, segments 12-14 with setose hairs. Palpi five-segmented, third segment simple.

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Thorax narrowed in front, more or less conical. Legs long and slender; femora unarmed; femora and tibial vestiture with very fine erect hairs; last three tarsal segments relatively short; fourth segment cordate to subcylindrical; female claws equal, each with a small internal basal barb. Wing long and narrow, often with color pattern; r-m crossvein distinct but short; costa long extended almost to wing tip; one radial cell; posterior branch of radius closely approaching costa; median fork broadly sessile. Female abdomen slender with pairs of eversible glands and gland rods. Male genitalia: ninth sternite short, ninth tergite tapering with well developed lateral lobes; basistyles slender, dististyles elongated; aedeagus with slender arch and pointed tip; parameres separated. Approximately 24 species have been described. Distribution: South America. This genus resembles Stenoxenus . Bezzia and Calyptopogon with the following exceptions: thorax not conical in front; parameres separated and antennae with setose hairs, these three characters are different from Bezzia and Calvptoppgon . The lack of elbow of wing vein and the male genitalia separated separates this genus from Stenoxenus. Illustration: Fig. 76 Paryphoconus angustipennis Enderlein Genus Pachyhelea Wirth Pachyhelea Wirth, 1959, Bull. Brooklyn Ent. Soc. (N.S.), 54:50. (Type -species: Ceratopogon magnus Coquillet, mon.) DIAGNOSIS. Body stout and nearly bare. Eyes bare, well separated. Palpus small, all segments simple. Antennal segments 310 oval, segments 11-15 cylindrical and elongated. Scutum very robust large,

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ilk with an anterior spine; humeral pits conspicuous. Abdomen stout, cone shaped. Legs large, hind coxa greatly enlarged; hind femur much enlarged and curved; hind tibia rather thick; no spines on femora and tibia. First tarsal segments of hind legs very long, fourth segment slightly cordiform, fifth segment slender with three pairs of ventral spines on distal half. Claws equal, small on the first fore legs, longer on hind legs; each with small mesal tooth. Wing broad without macrotrichia, microtrichia distinct; costa long; two radial cells; media widely sessile. Male genitalia: ninth sternite small, rounded; ninth tergite tapering with well developed lateral lobes; basistyle large; dististyle short with hook tip; aedeagus triangular shape, truncate end; parameres separated,. Only one species known. Distribution: Mexico, Canal Zone, and Brazil. J Wirth (1959b) proposed this genus and selected Ceratopogon magnus Coq. as the type, as it is not related to any other genera of Ceratopogonidae . PachybLea resembles Palpomyia in general, but differs in having unarmed femora and very short radial crossvein. Hind femora swollen as in Serromyia but unarmed and of different structure, closer to Sphaeromias . The la-tter genus is distinctly separated from Pachyhelea in having typical ventral blunt spines (batonnets) on the fifth tarsomere. Illustration: Fig. 77 Pachyhelea pachymera (Will.)

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Qenus Palpomyia Meigen Palpomvia (Megerle, M. S., in) Meigen, 1818, Syst. Beschr. Eur. Zweift. Ins., 1:65; Stephens, 1829, Cat. Brit. Ins., 2:238; Johannsen, 1905, Bull. N. Y. State Mas., 86:105; Kieffer, 1906, Gen. Insectorum, fasc. 42:61, 1913, Rec *nd. Mus., 9:187; Malloch, 1914, Bull. HI. State Lab. Nat. Hist., 10:216; 1915, Bull, HI. State Lab. Nat. Hist., 10:319; Kieffer, 1917, Ann. Mus. Nat. Hung., 15:295-318; Goetghebuer, 1920, Mem, Mas. Roy. Hist. Nat. Belg., 8:77; Edwards, 1926, Trans. Ent. Soc. London, 74:416; Ingram and Macfie, 1931, Dipt. Pat. & S. Chile, pt. 2, fasc. 4, p. 215; Johannsen, 1931, Arch. f. Hydrobiol., Suppl., 9:437; Goetghebuer, 1932, Bull. Ann. Soc. Ent. Belg., :212; Macfie, 1939, Ruwenzori Exped., 1934-35, 1:93; 1939, Rev. Ent., 10:207; Tokunaga, 1939, Tenthredo, 2:295; 1940, Tenthredo, 3*157; Zilahi-Sebess, 1940, Folia Ent. Hungar., 5:85; Johannsen, 1943, Ann. Ent. Soc. Amer., 36:734; Lee, 1948, Proc. Linn. Soc. N.S.W., 73:57; Wirth, 1952, Univ. Calif. Pub. Ent., 9:217; Lane, i960, Rev. Bras. Biol., 20:382. (Typespecies: Ceratopogon flavipes Meigen, rnon.). Apogon Rondani, I856, Dipt. Ital. Prod., 1:175. (Type-species: Ceratopogon hortulanus Meigen, orig. desig. ) . Alasion Rondani, 1857, Dipt. Ital. Prod., 2:14 (new name for Apogon Rondani, not Lacepede. 1802, Pisces). Heteromyia Malloch (not Say), 1915, Bull. HI. State Lab. Nat. Hist., 10:324; Kieffer, 1917, Ann. Mus. Nat. Hung., 15:324.

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DIAGNOSIS. Moderate to large size, slender and almost bare body. Eyes bare and separated. Female antenna! segments 3-10 oval with sparse verticils, segments 11-15 cylindrical and elongated; male antennae with few plumes; palpi slender, third segment simple. Mesonotum usually bearing a small tubercle or spine; surface clothed with fine hairs. Fore femora with stout ventral spines, the mid and hind ones usually without; fore femora often enlarged; the other legs usually slender; fourth tarsal segment bilobed beneath or cordiform fifth tarsal segment bare rarely with a few sharp ventral spines; claws equal, moderate size, sometimes with an inner basal tooth. Wings long and narrow, without macrotrichia; costa long; two radial cells, the second one well developed; intercalary fork absent; media widely sessile. Female abdomen with paired eversible glands opening between terga, these marked internally by slender sclerotized rodlike projections pointed to the base of abdomen. Male genitalia inverted; ninth tergite small, apicolateral processes large; basistyles and dististyles simple; aedeagus triangular to conical, apex caplike and clear; parameres fused or separated. Approximately 241 species have been described. Distribution: World-wide. Thorns en (1937) described the larvae and pupae of Palpomyia tibialis and P. longioennis and pointed out the character which differentiated Palpomyia from the allied genera. Lee ( 1948) studied the larvae and pupae of the Palpomyia Group which included Palpomyia,, Cllnohelea. Heteromyia . Xenphelea, Johannsenomyia. and Dicrohelea.

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EGO. Palpomyia flavlpes eggs are placed nearly vertical in the mass; nearly straight, long and narrow shape. LARVA. Snake-like, without anterior or posterior prologs. Largest size compared to other genera of Ceratopogonidae. Anal hairs usually shorter than last segment. Larvae can be separated from those of Bezzla and Probezzia by the flap-like structures attached to the conjunctiva on the inside of cast skin. PUPA. Largest pupae of Ceratopogonidae. Operculum with one pair of setae. Respiratory trumpet with 10-23 spiracles. Adult habits and larval habitats: larbae are aquatic, mostly occur in blanket algae or in the mucky or sandy margins or bottoms of lakes, ponds and streams. Some species breed in the profundal zone of deep lakes. The adults are predacious with strong, spinose legs. (Wirth and Stone, 1956) Abdomen with gland rods present resembles Stenoxenus , Bezzia , Phaenobezzia and Paryphoconus, but the presence of two radical cells on wing and simple wing veins separates Palpomyia from the above. The fore femora is always enlarged as in Heteromyia but the hind leg is not unusually long as in Heteromyia . Subgenus Palpomyia s. str. At least fore femur armed ventral! y with black stout spines; claws moderately large. Illustration: Fig. ?8 Palpomyia plebela (Loew) Subgenus Iftpldhelea Kief for, 1925 Subgenus type: Diplohelea parvula K. All femora unarmed; claws very small.

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Key for the Subgenera of Palpomyia i. At least fore femur armed ventrally -with black stout spines (Fig. ?8G) . . . Palpomyia (Fig. 78) All femora unarmed Diplohelea

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Genus Bezzia Kieffer Bezzla Kieffer, 1899, Bull. Soc. Ent. Prance, p. 69; Johannsen, 1905> Bull. N. Y. State Mus., 86:102; Kieffer, 1906, Gen. Insectorum, fasc. 42:57; Johannsen, 1908, Bull. N. Y. State Mis., 124:267; Kieffer, 1913, Rec. Ind. Mas., 9:192; Malloch, 1914, J. N. Y. Ent. Soc, 20:281; 1915, Bull. HI. State Lab. Nat. Hist., 10: 345; Kieffer, 1919, Ann. Mas. Nat. Hung., 17:112; Goetghebuer, 1920, Mam. Mus. Roy. Hist. Nat. Belg., 8:96; Ingram and Macfie, 1921, Ann. Trop. Mad. & Parasit., 15*361; Edwards, 1926, Trans. Ent. Soc. London, 74:422; Johannsen, 1931» Arch. f. Hydrobiol., 9:439; Macfie, 1932, Ann. Mus. Nat. Hist., ser. 10, 9:496; Sebess V. Zilah, 1938, Mat. Term. Ert., Budapest, 57:1092; Tokunaga, 1939, Tenthredo, 2:273; 1940, Tenthredo, 3:160; Zihali-Sebess, 1940, Folia Ent. Hungar., 5:98; de Meillon, 1942, Estac. anti-Malar. Lourenco Marques :19; Johannsen, 1943, Ann. Ent. Soc. Amer., 36:785; Lee, 1947, Proc. Linn. Soc. N.S.W., 72:339; Wirth, 1952, Univ. Calif. Pub. Ent., 9:226; Lane, 1958, Rev. Bras. Ent., 8:26; Tokunaga & Murachi, 1950, Ins. Micron., 12(3) :4l7. (Type-species: Ceratopogon ornatus Meigen, orig. desig. ) . Probezzia Malloch (in part, not Kieffer), 1914, Proc. Biol. Soc. Wash., 27:137; 1915, Bull. HI. State Lab. Nat. Hist., 10:352 (part); Macfie, 1940, Ann. Trop. Med. & Parasit., 34:29; Johannsen, 1943, Ann. Ent. Soc. Amer., 36:785, and authors. Peeudobezzia Malloch. 1915, Bull. HI. State Lab. Nat. Hist., 10:351; Kieffer, 1919, Bull. Soc. Ent. France, p. 193; Johannsen, 1934, J. N.Y. Ent. Soc., 42:344; 1943, Ann. Ent. Soc. Amer., 36:762.

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120 (Type-species: Ceratopogoh expolitus Coquillett, orig. desig.). Allobezzia Kieffer, 1917, Ann. His. Nat. Hung., 15:296 ( -type-species: CeratopoEon expolitus Coquillett, orig. desig.). Lasiobezzia Kieffer, 1925, Ann. Soc. Ent. France, 9^:5^ (Type-species: Bezzia pilipennis Lundstrom, orig. desig.). DIAGNOSIS. Body slender to stout. Eyes bare, widely separated. Female antennal segments 3-10 short, oval, segments 11-15 cylindrical and elongated; male antennae plumose. Mesonotum with spar— bristles, without anterior spine or tubercle. Fore femora with stout ventral spines, sometimes also on mid and hind pairs; fore femora often somewhat swollen; fourth tarsal segment cordate, fifth segment cylindrical; claws simple and equal, basal internal tooth sometimes present. Wing narrow, without macrotrichia; one radial cell; R 1 and R g widely separated; costa long not reaching wing tip; intercalary fork absent; media sessile ( Bezzia ) or short petiolate ( Pseudobezzia ). Female abdomen with a pair of eversible glands as in Palpomyia . Male genitalia inverted; ninth stemite large, ninth tergite small with well developed apicolateral processes; basistyle and dististyle short; aedeagus triangular to conical; parameres fused, appearing as one long lobe. Approximately 23^species have been described. Distribution: World-wide. The immature stages of this genus are very similar to those of Palpomyia . No distinct characters have been found to separate these two genera. Some descriptions of immature stages of Bezzia are presented in Thomson (1937) who also gave the descriptions of Bezzia varicolor and B. flavitarsis .

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EGG. Eggs are visually straight,' long and narrow arranged almost vertical in the mass. Eggs of Bezzia varicolor are black, about twelve times longer than wide, they are surrounded by jelly. LARVA. Larvae are snake-like without anterior or posterior prologs. Cast skin has no flap-like structure. PUPA. Pupa has three setae on operculum. Respiratory trumpet is short and dark. Caudal spines are stout with sharp black tips. Glandular discs are absent. Larval habitat: the habitat of Bezzia larvae has been reported in Thomson (1937). Bezzia varicolor larvae are found in algae in ponds. This genus resembles Palpomyia but differs in lacking an anterior tubercle; femoral spines present or absent; fourth tarsomere short and cordate; vein R 2 + 3 absent, only one radial cell on wing; median fork narrowly sessile. Subgenus Bezzia s. str. Median fork sessile; femora armed at least on fore legs; costa short; r-m crossvein ends slightly beyond forking point of Mj and Mg. Illustration: Fig. 79 Bezzia ( Bezzia) setulosa (Loew) Subgenus Pseudobezzia Malloch, 1915 Subgenus type: Ceratopogon expolitus Malloch, 1915 (orig. desig.) Median fork barely petiolate or very short petiole in male; costa short; legs very spiny; male genitalia large with extreme contraposition of the aedeagus and parameres. Illustration: Fig. 80 Bezzia ( Pseudobezzia) flavitarsis (Mall.)

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122 Key for the Subgenera of Bezzia Medial fork just petiolate (Fig. 8QE) or with very short petiole in male wing; legs very spiny (Fig. 80F); , Pseudobezzia (Fig. 80) Jfedial fork sessile (Fig. 79E); legs armed at least on fore legs (Fig. 79H) 562215 ( Fi S« ^

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123 Genus' Phaenobezzia Phaenobezzia Haeselbarth, 1965* S» Afr. Inst. Med. Res., Johannesburg, 5^:297. (Type-species: Probezzia pistiae Ingram and Macfie, orig. desig.). DIAGNOSIS. Body slender with fine hairs. Eyes bare and separated. Palpi slender; mandible with seven large teeth. Female antennal segments 3-10 oval, segments 11-15 elongated; male antennae with few plumes. Legs pale; hind femora rather stout; femora and tibiae unarmed; fourth tarsal segment simple; fifth segment with numerous, sharp, ventral batonnets; no batonnets on male tarsomere; female claws equal and small with internal barb; male claws simple, tip bifid. Wing with one radial cell, costa long. Abdomen hairy, elongated seventh abdominal segment bearing a pair of gland rods; spermathecae two. Male genitalia: ninth steraite large, ninth tergite tapering with well developed lateral lobes; basistyles and dististyles much reduced but distinctly separated; aedeagus slender with clear tip; parameres fused into one long lobe. Approximately 12 species have been described. Distribution: Africa, Sahara, and North America. The Immature stages were studied by Ingram and Macfie (1921) and Haeselbarth (1965). EGG. Elongated with one end rather pointed. LARVA. Pale or nearly white colored. Antenna and palpi small. Mental plate with a strong, pointed central tooth, and two more delicate teeth on each side. Hypopharynx bearing about 12 pointed teeth. Mandible large. Anal segment with 14 strong hairs dis tally.

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124 PUPA. Respiratory trumpets visually bent posteriorly, rather short and broad, distal end infuscated. Spiracles about 18. Abdomen with spiculated integument and pigmented areas. Anal segment with acutely pointed, darktipped processes. Several pupal specimens were collected in stagnant or flowing water, often among vegetation. In 1965, Haeselbarth proposed a new genus Phaenobezzia to enclose species formerly placed in the genera Bezzia, Palpomyia and Nilobezzia . It is well defined by the hypopygium of the male. This genus is related to Bezzia. Probezzia and Nilobezzia . It has the following similar characters of Bezzia ; gland rods present on abdomen; wing with one radial cell, and male parameres fused in one long lobe. It differs in having unarmed femora and reduced claspers of male genitalia. Male genitalia with basistyles and dististyles distinctly separated, not fused to form a thumblike process which separates it from Nilobezzia . Batonnets present only on fifth tar some re of female similar to Probezzia . but without ventral hair tufts in Phaenobezzia and the character of male genitalia separates these two genera. Illustration: Fig. 81 Phaenobezzia pistiae (I. & M.) Genus A to This genus is closely related, Sphaeromi as by the following characters: fifth tarsal segments armed with numerous batonnets; anterior tubercle present; all claws equal, each with inner barb; wing with two radial cells; legs with bristles on femora and tibiae; fifth tarsomeres of male with few batonnets.

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125 It differs from Sphaeromias by the following characters: claw with inner tooth very large, subequal to the claw; legs with spines only on fore femora; eighth abdominal segment without ventral hair tufts; fore femora swollen, fore tibiae arched as in Heteromyia; spermathoca one, eyes separated distinctly; anterior strong erect spine well developed; male genitalia with parameres separated. Four species: species 1. Thailand Fig. 82 species 2. Ceylon species 3. Viet Nam species Sumatra (? Palpomyia caesia Macfie 193*0 i Genus B This genus resembles Pachyelea by the following characteristics: scutum with a median anterior spine; female abdomen with gland rods on seventh segment; hind femur enlarged and curved; hind tibia with rough margin; first tarsal segment of hind legs very long; fifth tarsomere with ventral spines; claws small and equal on first four legs, longer on hind legs, each claw with mesal tooth; spermathecae two; eyes widely separated. The f ollowing characters differentiate this genus from Pachyhelea : all femora with ventral spines; mid tibia with an apical spine; wing with one radial cell; ventral spines on fifth tarsal segments approximately four pairs, scattered along the entire segment, while in Pachyhelea they appear only on distal half; abdomen slender and elongated, female abdomen with sclerotized plate; male genitalia large

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126 I with spinelike lobe on basistyle. One species: Rio Raposo, Colombia Fig. 83 Genus C This genus has the following characters of Palpomyia ; mesonotum with small tubercle; femora armed with ventral spines; two radial cells present on wing, eyes separated; spermathecae two. It also resembles Homohelea by having mandible with apical teeth large; all femora armed with ventral spines; mid tibia with an apical ventral spine. It can be differentiated from Homohelea particularly in having hind femora swollen and fifth tarsomere without batonnets. The following additional characters separate it from Palpomyia and Homohelea : claws of hind legs longer than those of the first four legs; fore femora slender not enlarged; hind first tarsomere very long; fourth tarsomere elongated, simple; fifth tarsomere of mid leg short; mandible spiny; no gland rods on abdomen. One species, female only: Chiapas, Mexico Fig. 8^ Genus D This genus is closely related to Pac hyhelea by the following characters: eyes well separated; anterior tubercle present on mesoj notum; abdomen cone-shaped with gland rods; hind femora enlarged and curved, hind tibiae with toothed apex; femora and tibiae unarmed; fifth tarsomeres sometimes with spines; spermathecae two; wing with

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two radial cells; fifth tarsomere of hind leg long. The following characters differentiate the genus from Pachyhelea: hind tarsal claws longest; spines (if present) on fifth tarsomere about two to four pairs; gland rods appear on abdominal segments five, six and seven; wing with costa prolonged beyond nearly to wing tip. Two species: species 1. Tex., Panama, Colombia; legs and wing dark; Fig. 85. species 2. Panama, Colombia; legs pale bands, wing whitish; Fig. 86. Genus E This genus has the following characteristics of Mallochohelea : all claws equal; wing with two radial cells; mandible teeth large; antennae elongated; eyes broadly separated; hind femur enlarged; batonnets present on fifth tarsomere. It resembles Pachyhelea but differs from Mallochohelea by the following points: anterior spine present; hind legs enlarged, femora unarmed; two pairs of batonnets on fifth tarsomere; claws equal, each with inner basal tooth; no ventral hair tufts on eighth abdominal segment; abdomen with gland rods but widely separated from each other; wing with two radial cells. The following characteristics differentiate this genus from Pachyhelea : abdomen not cone-shaped; hind femora not curved but swollen in the middle; tibia without toothed margin; first tarsomere of hind leg simple not long; ventral spines of fifth tarsomere not

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located only on distal half of the segment; spermathecae one; mandible with few large teeth. One species, female only: Colombia, Fig. 87. Genus F This genus resembles Sphaeromias by the following characters: legs usually with ventral spines or bristles on femora and dorsal side of tibiae; fifth tarsomere with batonnets; claws large with inner basal tooth; spermathecae two. No gland rods. It differs from Sphaeromias by the following characters: eyes widely separated; fourth tarsomere cordate to oval; hind claws longest; wing with one or two radial cells; costa extends beyond nearly to wing tip; male genitalia: basistyles with additional processes, dististyle large; ninth tergite small, parameres separated. Two species: species 1. Panama, Colombia; one radial cell; Fig. 88. species 2. Panama, Costa Rica; two radial cells; Fig. 89. Qenus G This genus is characterized by the following characters: anterior tubercle present; femora unarmed below; tibia with bristles; fourth tarsomere cordate; fifth tarsomere stout, no batonnets; claws equal, short, sharp with internal tooth; abdomen slender, with pattern on ventral side; one large spermatheca, pear-shaped; wing with two radial cells, costa short, r-m crossvein short, media sessile, microtrichia well developed; proboscis short with large mandible; palpi

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129 with third segment swollen; eyes separated; antennal segments three to nine oval, 10-15 elongated. One species: Colombia, Fig. 90. Genus H This genus resembles Mallochohelea by the following points: eyes broadly separated; mandible with large teeth; female antennae long, fifth tarsomere bearing about five pairs of batonnets; claws long and equal, wing with two radial cells; spermathecae two. It is differentiated from Mai ^hph^e^ bv the following characters anterior part of thorax somewhat cone-shaped; anterior tubercle poorly developed; legs long and slender, subequal size; tibiae with dorsal bristles, mid tibia with an apical spine; second tarsomere of mid leg distinctly shorter than that of the other legs, claws with internal basal tooth as in Sphaeromias; seventh abdominal segment with a pair of ventral sclerotized plates and hairy. One species: Colombia, Fig. 91. Genus J This genus is closely related to Sphaeromias by the following characters: eyes contiguous; fifth tarsomere bearing batonnets; claws large and equal; two spermathecae. It differs from Sphaeromias by having short antennae with segments 3-10 oval; anterior tubercle absent; legs without spines on femora or tibiae; tibiae of second leg short; claws with internal tooth

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in one and external tooth in the other; wing with one radial cell. One species t Brazil Amazonas, Rio Paru; Fig. 92. Genus K It resembles genus H by the following characters i female seventh abdominal segment with ventral, hairy, sclerotized plates; two spermathecae, legs with bristles on femora and tibiae second tarsomere of mid leg shorter than that of other legs; mid tibiae with distal spine-, legs long and slender; fifth tarsomere with batonnets;; claws with internal tooth, anterior tubercle poorly developed; eyes broadly separated, mandible with large teeth, antennae long. It is differentiated from genus H by having one radial cell on wing. This genus is also related to Bezzia by having equal claws ; wing with one radial cell; leg sometimes with spine on femur and tibia. One species: Colombia, Fig. 93. Genua L This genus is similar to genus E in the following points I claws equal, mandible with few large teeth; antennae long; eyes broadly separated; femora unarmed; hind femur enlarged; fifth tarsomere with few batonnets; abdomen with gland rods and widely separated; one spermatheca.

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It resembles Bezzia by having one radial cell on the -wing; fore femora rather swollen, fourth tarsomere cordate, claws equal, abdomen with a pair of eversible glands. The genus is differentiated from genus E and Bezzia by having tarsal segments two to five distinctly short. One species: Colombia, Fig. 9^. Genus M This genus resembles Parabezzla by the following characters: female antennal segments 3-10 oval, segments 11-15 cylindrical to elongated; male antenna plumose; tarsi with ventral spines; fifth tarsomere swollen, claws long, unequal in female, small and equal in male; wing with media petiolate; male genitalia, aedeagus arched, parameres reduced to form triangular sclerite behind aedeagus. The character of unequal claws on hind legs resembles AHu'audonod.a . It differs from Parabezzia and Alluaudomyia by having palpi four segmented; coxae of male with few bristles; fourth tarsomere cordiform to oval; costa short, wing with radial cells obliterated; one large spermatheca; male genitalia with narrow ninth tergite, basistyles large, dististyles slender with claw-like tips; claws with both external and internal barbs; eyes narrowly separated. The following points are particularly different from Alluaudomyia : third palpal segment swollen with deep sensory pit; antennal segments 3-10 oval, 11-15 cylindrical to elongated; wing with microtrichia present. One species: Brazil, Fig. 95.

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132 i Genus N This genus is similar to Brachypogon in the absence of the lower branch of media on -wing; legs slender and unarmed. It also resembles Austroconops in the following characters: hind tibia with plumose spur; four tibial comb; claws small and equal. It differs from Brachypogon and Austroconops by the following points: eyes bare; antennae with segments 3-10 oval, 11-15 long; one spermatheca; palpi three-segmented; mandible with five teeth; one triangular radial cell. It particularly differs from Brachypogon in having small, equal claws and from Austroconops by the absence of the lower branch of media. One species, female only: Colombia, Fig. $6. Genus 0 It resembles Parabezzia by the following points: wing with costa extended beyond and media petiolate. The genus is differentiated from Parabezzia by having pubescent eyes; claw single with barb; wing with two radial cells and palpi three-segmented. One species: Colombia, Fig. 97. Genus P This genus is related to C ampt opter ohelea by the following characteristics: eyes widely separated; proboscis short; thorax

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dark; legs rather stout, hind tibia with more hairs on comb; wing broad, two radial cells, radial vein thickened, cubital stem and 1A distinct. It differs from Camptopterohelea by having pubescent eyes; antenna 14-segmented, all segments similar in shape; palpi four-segmented, third segment robust; proboscis short but large; mandible teeth very small and blunt; two claws present; wing with first" radial cell not well developed, one median vein present, microtrichia simple; spermathecae two. One species: Florida, California; Fig. 98. Genus Q It is related to Camptopterohelea by the following points: eyes widely separated; palpi two-segmented; proboscis vestigial; thorax dark; legs rather stout; wing broad; one spermatheca. It is differentiated from Camptopterohelea by having pubescent eyes; two claws, wing with radial cells absent, median fork present with petiolated media, microtrichia simple; antennal segments 3-10 round, 11-15 oval to cylindrical; male antennae with eight segments, without plume; male genitalia with ninth tergite large and truncated end, aedeagus arched, pararaere absent. One species: Ecuador, Fig. 99.

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13^ Genus R This genus resembles Parabezzia by the following characters: eyes widely separated; palpi four-segmented, first and second segments indistinctly separated; antennal segments 3-10 oval, 11-15 cylindrical to elongated; male antennae with well developed plumes, tarsi with ventral spines, fourth tarsomere cordate, fifth segment swollen, claws long and equal; wing with one radial cell, costa prolonged beyond R,-, media petiolate; spermathecae two. It differs from Parabezzia by having pubescent eyes; first tarsomere of mid leg longest and with spines; coxae of male with few hairs; claws with inner barb, tibia of fore leg swollen centrally; wing broad; male genitalia with basistyles large, bearing basal spine-like processes, dististyles with hook-like tips, aedeagus arched, paramere absent. One species: Amazonas, Brazil; Pig. 100. Genus S The male of this genus is characterized as follows: eyes pubescent and widely separated; proboscis vestigial; palpi two-segmented; antennae with eight segments, segments three and four oval, segments five to eight elongated, without plumes; legs slender; tarsal segments two to five similar in shape and subequal; fourth tarsal segment cylindrical; claws small and equal; wing without radial cells, posterior branch of media short, + i± short; male genitalia with basistyles and dististyles small, ninth sternite small, ninth tergite large with truncated margin, aedeagus

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135 arched with arras parallel, parameres absent. One species, male only: Ecuador, Fig. 101. Genus T This G enus has the following characteristics of Mallochohelea : eyes widely separated; mandible teeth large; female antennae long; no anterior spine or tubercle; legs slender; fifth tarsomere with about si* pairs of batonnets; claws long and equal with external basal tooth; male claws small, slightly cleft at tip; spermathocae two; ventral hair tufts on eighth abdominal segment present. It differ from f|f^ **»t ** **** nandible Witt serrate margin; all femora curved and enlarged distally, spermathecae large; male genitalia with parameres fused basaTLy, separated and fused again' distally; abdomen slender. One species: Mallochohelea amnicola Macfie; Brazil, Colombia; Fig. 102. The following four genera are not represented in our collections. Since specimens were not available for study and since the original descriptions were inadequate they were not included in the keys. All available information is included here. These can be worked into the keys when specimens are available for study.

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136 Genus Dolichohclca Edwards, 1929 (Type-species: Dolichoholea polita Edwards , mon.) Thorax produced anteriorly, mesonotum with an anterior tubercle; ,dn, narrow, second radial cell very long extending to more than fourfifths of wing length, median fork with short stem, anal vein straight* proxi.nl antennal segments of female more elongate, one to five times as long as broad. Genus Acanthohelea Kief for, 1917 (Type-species: Acanthohelea pruinosa Kieffer, mon.) Fourth tarsomeres bilobed on all legs, all femora and tibiae spincse; male antenna with last three segments elongated. Genus Luciamyia de Kbillon, 1937 (Type-species: Luciaryia biloba de Msillon, mon.) Wing densely hairy, *i?S venation greatly modified, vein 1% absent, second radial cell greatly prolonged and broad dis tally. The costa, vein R^, and vein ^ meet in a point at the wing tip where there is a distinct indentation at the ra.ng margin and the fringe is modified into lanceolate scales. Genus Ceratobezzia Kieffer, 1917 (Type-species: Ceratobeasia fallax K.) Fourth tarsomeres bilobed each armed with too stout spines, fifth tarsomeres greatly swollen on fore legs; claws unequal on mid and hind legs but equal on fore legs; one spormatheca; one radial cell,

PAGE 148

ALPHABETICAL LIST OF GENEFA AND STBCSIERA (Subgenera underlined) . _ f o r S 14-3 £ A ec mm S Caloforciuonyia . .... 32 . 56 Camptopterohelea . .... 62 . 56 .... 136 . 5^ .... 63 . 5^ .... 65 .... 85 .... ^7 . 66 Dacnoforcipomyia . .... 3^ Isthnohelea ...... I' 137

PAGE 149

138 Page Jenkinshelea 97 Johannsenonyia 93 Kiefferomyia 79 Lasiphelea . . . 3^ Lepidohelea 35 Leptoconops ^ LeptocoRO'os ....... 17 Luciamyia 136 Kacfiella 57 Kacrurohelea . . . .... 83 Mallochohelea 106 Mataemyia 57 Mei.ierehelea . 54 Me loehelea . . . 24 Ketaforciporayla 32 Kbnocrulic oides ...... 57 Konohelea 77 Tfonohelea 79 Neof orci'Domyia ...... 3^ Neo stilobezzla . . . . . . 74 Ifeurohelea 91 Nilobezzia 107 Kilobozzia 109 Kilohelea 66 Oecacta 55 i ?a?o. . . . 115 . . . 1 17 84 , 46 , 109 112 JrGJ^uCXCLOIujJ-cl • • • . . . 90 . . . 123 35 . . . 98 ... 44 ... 121 A b w UUU^ Uiwv w • • ... 43 . . <• • • ... 24 . . 36 Sohn £ohG~J GcL m * • • • • • ... 57 . . . 102 Stilobozzia . . . . ... 73

PAGE 150

139 Page Stvloconops lo Synthyridorsyia 33 Tetrabezzia 87 Thyridorwla 33 Trichohelea 33 Trithecoides 53 Warmkea 31 Xenohelea 95 Genus A 12^ Genus B 125 Genus C 126 Genus D 126 Genus E 127 Genus F 128 Genus G 128 Genus H 129 Genus J 129 Genus K 130 Genus L 130 Genus K 131 Genus N 132 Genus 0 132 Genus P 132 Genus Q 133 Genus R 13^ Genus S 13^ Page Genus T 135

PAGE 151

Figure i, — Culicoides furens (Poey) : lateral view of female, left wing and right leg removed, with parts labeled.

PAGE 152

Figure 2 . — Lept oc onops ( Leptoconops ) stygius Skuse; A, female antenna; B, female palpus; C, female wing; D, female head; E, mandible; F, female legs; G, enlarged hind tibial comb; H, female tarsi; I, female fifth tarsomeres and claws; J, female spermathecae.

PAGE 153

Figure 3 , — Leptoc onopa ( Holoconops ) kerteszi K.: A, female antenna; B, male antenna; C, female palpus; D, male palpus; E, female wing; F, female head; G, female legs; H, enlarged hind tibial comb; I, female tarsi; J, female fifth tars ome res and claws; K, female spermathecae; L, female lamella; M, male genitalia.

PAGE 154

ik3 Figure 4. — Leptoconops ( Styloconops ) alblventrls (Meljere): A, female palpus; B, female wing; C, female head; D, male head; E, mandible; F, enlarged hind tibial comb; G, female front tarsus; H, female spermathecae; I, male genitalia.

PAGE 155

Figure 5 1 -Leptoconopg (Styloconope) spinosifrona Carter) A, female palpus; B, female wing; C, female head; D, mandible; E, enlarged hind tibial comb; F, female front tarsus; G, female spermathecae; H, female lamella; I, male genitalia.

PAGE 156

COOCCQOO Figure 6. — Atrichopogon (Atrlchopogon) levls (Coq^: A, female antenna; B, male antenna; C, female palpus; D, female wing; E, male wing; F, female head; G, female legs; H, enlarged hind tibial comb; I, female tarsi; J, female fifth tars ome res and claws; K, female spermatheca; L, female abdomen; M, male genitalia.

PAGE 157

Figure 7»— Atrlchopogon ( Psilokempia ) arcticua (Coq.): A, female antenna; B, male antenna; C, female palpus; D, male palpus; E, female wing; F, female head; G, mandible; H, female legs; I, enlarged hind tibial comb; J, female tarsi; K, female fifth tarsomeres and claws; L, female spermatheca; M, dorsal view of female abdomen; N, ventral view of female abdomen; 0, male genitalia.

PAGE 158

1*7 Figure 8.-Atrichopogon ( Maloehelea ) meloesugans K. : A, female antenna; B, male antenna; C, female palpus; D, male palpus; E, female wing; F, male wing; G, mandible; H, female legs; I, enlarged hind tibial conib; J, female tarsi; K, female fifth tars ome res and claws; L, female spermathecae; M, male genitalia.

PAGE 159

Figure 9. — Forcipomyia ( Forcipomyia ) bipunctata (L. ): A, female antenna; B, male antenna; C, female palpus; D, male palpus; E, female wing; F, female legs; Q, enlarged hind tibial comb; H, female tarsi; I, female fifth tarsomeres and claws; J, female spermathecae; K, female abdomen; L, male genitalia; M, male parameres.

PAGE 160

D Figure 10 . — Forcipomyia ( Prof orcipomyia ) wirthl Saunders: A* female antenna; B, male antenna; C, female palpus; D, male palpus; E, female wing; F, female head; G, mandible; H, female legs; I, enlarged hind tibial comb; J, female tarsi; K, female fifth tars ome res and claws; L, female spermathecae; M, female abdomen; N, male genitalia.

PAGE 161

150 Figure 11. Forclpotnyia ( Warmkea) aerla Saunders: A, female antenna; B, female palpus; C, female wing; D, female head; E, mandible; F, female legs; G, enlarged hind tibial comb; H, female tarsi; I, female fifth tarsomeres and claws; J, female spermatheca.

PAGE 162

Figure 12.~ Forcipomyia ( Warmkea ) tuberculata Saunders: A, male palpus; B, male wing; C, male genitalia; D, male parameres.

PAGE 163

Figure 13. — Forcipomyia ( Calof orcipomyia) glauca Edwards: A, female antenna; B, male antenna; C, female palpus; D, male palpus; E, female wing; F, male wing; G, female head; H, female legs; I, enlarged hind tibial comb; J, female tarsi; K, female fifth tar scire res and claws; L, female spermathecae ; M, male genitalia.

PAGE 164

Figure 14. — Forcipomyia ( Metaforcipomyia ) pluvialis Malloch: A, female antenna; B, male antenna; C, female palpus; D, female wing; E, female legs; F, enlarged hind tibial comb; G, female tarsi; H, female fifth tarsomeres and claws; I, female spermathecae; J, female abdomen; K, male genitalia.

PAGE 165

154 Figure 15 . — Forciponyia ( Thyrldonyia ) palustrls Saunders: A, female antenna; B, male antenna; C, female palpus; D, male palpus; E, female legs; F, female tarsi; 0, female fifth tarsomeres and claws; H, female spermatheca; I, female abdomen; J, male genitalia; K, male parameres .

PAGE 166

155 Figure l6.--Forcipomyia ( Synthyridomyia ) colemanl Wirth: A, female antenna; B, male antenna; C, female palpus; D, male palpus; E, female legs; F, enlarged hind tibial comb; G, female tarsi; H, female fifth tarsomeres and claws; I, female spermatheca; J, male genitalia; K, male parameres.

PAGE 167

Figure 17. — Forclpomyla ( Trlchohelea ) fijiensis (Macfie): A, female antenna; B, male antenna; C, female palpus; D, male palpus; E, female wing; F, female head;,G, mandible; H, female legs; I, enlarged hind tibial comb; J, female tarsi; K, female fifth tarsomere and claws; L, female spermathecae; M, female abdomen; N, male genitalia; 0, male parameres .

PAGE 168

Figure 18 . ~ Forcipomyia ( Lasiohelea ) fairfaxensls Wirth: A, female antenna; B, male antenna; C, female palpus; D, male palpus; E, female wing; F, male wing; G, female head; H, female legs; I, enlarged hind tibial comb; J, female tarsi; K, female fifth tarsomares and claws; L, male fifth tarsomere and claws; M, female sperma theca; N, male genitalia; 0, male parameres.

PAGE 169

158 Figure 19. — Forclpomyla ( Neof orclpomyla ) eques (Joh. ): A, female antenna; B, male antenna; C, female palpus; D, male palpus; E, female wing; F, male wing; G, female head; H, mandible; I, female legs; J, enlarged hind tibial comb; K, female tarsi; L, female fifth tarsomeres and claws; M, male fifth tarsomere and claws; N, female spermathecae; 0, male genitalia. 4*

PAGE 170

I Figure 20,-Forclpomyia ( Dacnoforcipomyia ) anabaenae Chan & Saunders: A, female antenna; B, female palpus; C, female head; D, mandible; E, female legs; F, enlarged hind tibial comb; G, female tarsi; H, female fifth tarsomeres and claws; I, male genitalia.

PAGE 171

160 Figure 21. — Forcipomyia ( Phasmldohelea ) fuliglnosa (Mg.): A, female antenna; B, male antenna; C, female palpus; D, male palpus; E, female wing; F, male -wing; Q, female head; H, mandible; I, female legs; J, enlarged hind tibial comb; K, female tarsi; L, female fifth tarsomeres and claws; M, female spermathecae; N, male genitalia; 0, male parameres.

PAGE 172

G Figure 22 . — Forcipomyia ( Phasitddohelea) sp. t A, female antenna; B, female palpus; C, female wing, D, mandible; E, female legs; F, enlarged hind tibial comb; G, female tarsi; H, female fifth tarsomeres and claws; I, female spermathecae; J, female abdomen.

PAGE 173

162 Figure 23 . — Forclpomyia ( Lepldohelea ) annulatipes Macfio: A, female antenna; B, male antenna; C, female palpus; D, male palpus; E, female wing; F, male wing; G, female head; H, female legs; I, enlarged hind tibial comb; J, female tarsi; K, female fifth tarsomere and claws; L, female spermatheca; M, female abdomen; N, male genitalia; 0, male parameres.

PAGE 174

Figure 2k, — Forcipomyia ( Pterobosca ) fusicornia (Coq.)t A, female antenna; B, female palpus; C, female wing; D, mandible; E, female legs; F, enlarged hind tibial comb; G, female tarsi; H, female fifth tarsomeres and claws; I, female spermathecae.

PAGE 175

Figure 25 . — Dasyhelea ( Dasyhelea ) grisea (Coq.): A, female antenna; B, female palpus; C, female wing; D, male wing; E, female head; F, female legs; G, enlarged hind tibial comb; H, female tarsi; I, female fifth tarsomeres and claws; J, female spermatneca; K, female abdomen; L, male genitalia; M, male paramere.

PAGE 176

Figure 26. — Dasyhelea ( Pseudoculicoide3 ) mutabilis (Coq.) : A, female antenna; 8, male antenna; C, female palpus; D, male palpus; E, female wing; F, male wing; G, female head; H, female legs; I, enlarged hind tibial comb; J, female tarsi; K, female fifth tars ome res and claws; L, female spermatheca; M, female abdomen; N, male genitalia; 0, male paramere.

PAGE 177

Figure 27»— Dasyhelea (Prokempla) cincta (Coq.): A, female antenna; B, male antenna; C, female palpus; D, female wing; E, female head; F, female legs; G, enlarged hind tibial comb; H, female tarsi; I, female fifth tarsomeres and claws; J, female spermathecae; K, female abdomen; L, male genitalia.

PAGE 178

167 Figure 28 , — Paradasyhelea a&rata Wirth & Lee: A, female antenna; B, male antenna; C, female palpus; D, male palpus; E, female wing; F, female head; G, female legs; H, enlarged hind tibial comb; I, female tarsi; J, female fifth tarsomeres and claws; K, female spermatheca; L, female abdomen; M, male genitalia.

PAGE 179

Figure 29.-Culicoides ( Culicoides) yukonensls Hoffman: A, female antenna; B, male antenna; C, female palpus; D, female wing; E, male wing; F, female legs; G, enlarged hind tibial comb; H, female tarsi; I, female fifth tarsomeres and claws; J, female spermathecae; K, male genitalia; L, male parameres.

PAGE 180

Figure 30. — Cullcoides ( Trithecoides ) flaviscutatus W. & H. : A, female antenna; B, male antenna; C, female palpus; D, male palpus; E, female wing; F, male wing; G, female head; H, mandible; I, female legs; J, enlarged hind tibial comb; K, female tarsi; L, female fifth tarsomeres and claws; M, female spennathecae; N, female abdomen; 0, male genitalia; P, male parameres.

PAGE 181

Figure 31«-Culicoides ( Haemophoructus ) gemellus Macfie: A, female antenna; B, male antenna; C, female palpus; D, male palpus; E, female wing; F, male wing; G, female head; H, mandible; I, female legs; J, enlarged hind tibial comb; K, female tarsi; L, female fifth tars ome res and claws; M, female spermathecae; N, female abdomen; 0, male genitalia; P, male parameres.

PAGE 182

171 Figure 32.-Culicoides ( Meijerehelea ) guttlfer ( Mel j Ire ) : A, female antenna; B, male antenna; C, female palpus; D, male palpus; E, female wing; F, male wing; G, female head; H, mandible; I, female legs; J, enlarged hind tibial comb; K, female tarsi; L, female fifth tarsomeres and claws; M, female spermatheca; N, female abdomen; 0, male genitalia; P, male parameres.

PAGE 183

Figure 33»~ Culicoldes ( Hoff mania ) insignls Lutz: A, female antenna; B, male antenna; C, female palpus; D, male palpus; E, female wing; F, male wing; G, female head; H, mandible; I, female legs J, enlarged hind tibial comb; K, female tarsi; L, female fifth tars ome res and claws; M, female spermathecae; N, female abdomen; 0, male genitalia; P, male pararaeres.

PAGE 184

Figure 34.-Culicoides ( Avarltia ) obsoletus (Mg.): A, female antenna; B, male antenna; C, female palpus; D, male palpus; E, female wing; F, male wing; G, female head; H, mandible; I, female legs; J, enlarged hind tibial comb; K, female tarsi; L, female fifth tarsomeres and claws; M, female spermathecae ; N, male genitalia; 0, male parameres.

PAGE 185

Figure 35 »~ Cullcoides ( Oecacta ) furens (Poey): A, female antenna; B, male antenna; C, female palpus; D, male palpus; E, female ving; F, male wing; G, female head; H, mandible; I, female legs; J, enlarged hind tibial comb; K, female tarsi; L, female fifth tarsomeres and claws; M, female spermathecae; N, male genitalia; 0, male parameres ,

PAGE 186

Figure 36 . -Cull c oldes ( Drymodesmyla ) coplosus R«& H. : A, female antenna; B, male antenna; C, female palpus; D, male palpus; E, female wing; F, male wing; G, female head; H, mandible; I, female legs; J, enlarged hind tibial comb; K, female tarsi; L, female fifth tarsomeres and claws; M, female spermathecae; N, male genitalia; 0, male parameres .

PAGE 187

176 Figure 37* — Culicoldes ( Diphaeomyia ) baueri Hoffman: A, female antenna; B, male antenna; C, female palpus; male palpus; E, female wing; F, male wing; G, female head; H, mandible; I, female legs; J, enlarged hind tibial comb; K, female tarsi; L, female fifth tarsomeres and claws; M, female spermathecae; N, male genitalia; 0, male parameres; P, pupal respiratory organ.

PAGE 188

Figure 38.-Cullcoides ( Anilomyia ) covagarciai Ortiz.: A, female antenna; B, male antenna; C, female palpus; D, male palpus; E, female wing; F, male wing; G, female head; H, mandible; I, female legs; J, enlarged hind tibial comb; K, female tarsi; L, female fifth tarsomeres and claws; M, female spermathecae; N, male genitalia; 0, male parameres.

PAGE 189

Figure 39. — Cullcoides ( Beltranmyla ) crepuscularis Malloch: A, female antenna; B, male antenna; C, female palpus; D, male palpus; E, female wing; F, male wing; 0, female head; H, mandible; I, female legs; J, enlarged hind tibial comb; K, female tarsi; L, female fifth tarsomeres and claws; M, female spermatheca; N, male genitalia; 0, male parameres.

PAGE 190

Figure 40 . — -Culicoides ( Qlaphiromyla ) scopus R. & H. i A, female antenna; B, male antenna; C, female palpus; D, male palpus; E, female wing; F, male wing; G, female head; H, mandible, I, female legs; J, enlarged hind tibial comb; K, female tarsi; L, female fifth tarsomeres and claws; M, female spermathocae; N, male genitalia; 0, male parameres .

PAGE 191

180 * V IMf**:j«^ r . :A • ••-.•;'-.},.'< r •t5*' '"""'•"HVj :s;.„,..i : A..:< • Figure 41 . — Cullcoldes ( Haematomyidium ) paraensls (Goeldi) : A, female antenna; B, male antenna; C, female palpus; D, male palpus; E, female wing; F, male wing; 0, female head; H, mandible; I, female legs; J, enlarged hind tibial comb; K, female tarsi; L, female fifth tars ome res and claws; M, female spermathecae; N, male genitalia; 0, male parameres.

PAGE 192

Figure kZ. --Culicoides (J&cfiella) phlebotomus (Will.)* A, female antenna; B, male antenna; C, female palpus; D, male palpus; E, female wing; F, male wing; G, female head; H, mandible; I, female legs; J, enlarged hind tibial comb; K, female tarsi; L, female fifth tarsomeres and claws; M, female spermathecae; N, male genitalia; 0, male parameres; P, pupal respiratory organ.

PAGE 193

Figure *»3. ~ Culicoldes ( Mataemyia ) mo.llngaensis W. & B.» A t female antenna; B, female palpus; C, male palpus; D, female wing; E, mandible; F, female legs; G, enlarged hind tibial comb; H, female tarsi; I, female spermathecae.

PAGE 194

183 Figure Cullcoides ( Monoculicoldes ) varlipennls (Coq.): A, female antenna; B, male antenna; C, female palpus; D, male palpus; E, female wing; F, male wing; G, female head; H, mandible; I, female legs; J, enlarged hind tibial comb; K, female tarsi; L, female fifth tarsomeres and claws; M, female spermatheca; N, male genitalia; 0, male paramere; P, pupal respiratory organ.

PAGE 195

184 : :. • N 1 '•"(v'»rr rr mttrrff i f ,>: »'.-i.. Figure 45.-Culicoides ( Self ia ) hleroglyphlcus Malloch: A, female antenna; B, female palpus; C, male palpus; D, female wing; E, male wing; F, female head; G, mandible; H, female legs; I, enlarged hind tibial comb; J, female tarsi; K, female fifth tarsomeres and claws; L, male genitalia.

PAGE 196

185 Figure 46. — Austroconops mcmillani Wirth & Lee: A, female antenna; B, female palpus; C, female wing; D, mandible; E, enlarged hind tibial comb; F, female tarsi; G, female fifth tars ome res and claws.

PAGE 197

J Figure 47 . — C amp topter ohele a hoogstraali W. & H, : A. female antenna; B, male antenna; C, female palpus; D, female wing; E, female head; F, mandible; 0, female legs; H, enlarged hind tibial comb; I, female tarsi; J, female fifth tarsomeres and claws; K, female spermatheca; L, female abdomen; M, male genitalia.

PAGE 198

187 Figure 48.-Ceratopogon ( Ceratopogon ) culicoidithorax Hoffman: A, female antenna; B, male antenna; C, female palpus; D, male palpus; E, female wing; F, mandible; G, female legs; H, enlarged hind tibial comb; I, female tarsi; J, female fifth tarsomeres and claws; K, female spermathecae; L, male genitalia.

PAGE 199

188 Figure 49.— . Ceratopogon ( Isohelea ) stigmalis Co(j.: A, female antenna; B, female palpus; C, female -wing; D, female legs; E, female tarsi; F, female fifth tarsomeres and claws; G, female spermatheca; H, male genitalia.

PAGE 200

189 Figure 50. — Alluaudomyia be 11a (Coq.): A, female antenna; B, male antenna; C, female palpus; D, male palpus; E, female wing; F, male wing; G, female head; H, female legs; I, enlarged hind tibial comb; J, female tarsi; K, male tarsi; L, female fifth tarsomeres and claws; M, male fifth tarsomeres; N, female spermatheca; 0, male genitalia; P, male parameres.

PAGE 201

Figure 51. — Alluaudoiryla parva Wirth: A, female antenna; B, male antenna; C, female palpus; D, male palpus; E, female wing; F, male wing; G, female head; H, female legs; I, enlarged hind tibial comb; J, female tarsi; K, female fifth tarsomeres and claws; L, male fifth tarsomeres; M, female spermathecae ; N, male genitalia; 0, male parameres.

PAGE 202

191 Figure 52,— Stilobezzia (Stilobezzla) f estiva K.j A, female antenna; B, male antenna; C, female palpus; D, male palpus; E, female wing; F, male wing; 0, female head; H, female legs; I, enlarged hind tibial comb; J, female tarsi; K, female fifth tarsomeres and claws; L, male fifth tarsomeres; M, female spermathecae; N, female abdomen; 0, male genitalia; P, male parameres.

PAGE 203

Figure 53«--Stilobezzia ( Eukrai ohelea ) elegantula (Joh.): A. female antenna; B, male antenna; C, female palpus; D, male palpus; E, female wing; F, female eye separation; G, female legs; H, enlarged hind tibial comb; I, female tarsi; J, female fifth tarsomeres and claws; K, male fifth tarsomeres; L, female spermathecae; M, female abdomen; N, male genitalia; 0, male parameres.

PAGE 204

193 Figure 5^. — Stilobezzia ( Neostilobezzia ) lutea (Malloch): A, female antenna; B, male antenna; C, female palpus; D, male palpus; E, female wing; F, female eye separation; G, female legs; H, enlarged hind tibial comb; I, female tarsi; J, female fifth tarsomeres and claws; K, male fifth tarsomeres, L, female spermathecae; M, male genitalia; N, male parameres.

PAGE 205

19^ Figure 55 • — Echlnohelea lanel VHLrth: A, female antenna; B, male antenna; C, female palpus; D, male palpus; E, female wing; F, female head; 0, female legs; H, enlarged hind tibial comb; I, female tarsi; J, female fifth tars ome res and claws; K, female spermatheca; L, female abdomen; M, male genitalia; N, male parameres.

PAGE 206

195 I Figure 56. — Monohelea ( Monohelea ) hleroglyphlca K. : A, female antenna; B, male antenna; C, female palpus; D, female wing; E, female head; F, female legs; G, enlarged hind tibial comb; H, female tarsi; I, female fifth tarsomeres and claws; J, male fifth tarsomeres; K, female spormathocae; L, male genitalia; M, male paramoro3.

PAGE 207

196 i Figure 57. — Monohelea ( Schlzohelea ) leucopeza (Mg.): A, female antenna; B, female palpus; C, male palpus; D, female wing; E, female eye separation; F, female legs; G, enlarged hind tibial comb; H, female tarsi; I, female fifth tarsomeres and claws; J, male fifth tarsomeres; K, female spermathecae; L, male genitalia; M, male parameres.

PAGE 208

197 Figure 58. — Serromyia femora ta (Mg.): A, male antenna; B, male palpus; C, female wing; D, male wing; E, female head; F, mandible; G, female legs; H, enlarged hind tibial comb; I, female tarsi; J, female fifth tars ame res and claws; K, male fifth tarsomeres; L, female spermathecae; M, male genitalia; N, male parameres.

PAGE 209

Figure 59* — Macrurohelea setosa Wirth: A, female antenna; B, female palpus; C, male palpus; D, female wing; E, female head; F, female legs; G, enlarged hind tibial comb; H, female tarsi; I, female fifth tarsomeres and claws; J, female spermathecae; K, female abdomen L, male genitalia; M, male parameres.

PAGE 210

199 L Figure 60. — Parabezzia petiolata Mall.: A, female antenna; B, male antenna; C, female palpus; D, male palpus; E, female wing; F, female head; G, male coxae; H, female legs; I, enlarged hind tibial comb; J, female tarsi; K, female fifth tarsomeres and claws; L, male fifth tarsomeres; M, female spermathecae; N, male genitalia; 0, female pupal respiratory organ; P, male pupal respiratory organ; Q, female anal segment; R, male anal segment.

PAGE 211

200 Figure 6l. — Clinohelea bimaculata (Loew): A, female antenna; B, male antenna; C, female palpus; D, female wing; E, female head; F, mandible; G, female legs; H, enlarged hind tibial comb; I, female tarsi; J, female fifth tars one res and claws; K, female spermathecae; L, ventral view of female abdomen; M, male genitalia; N, male parameres.

PAGE 212

201 Figure 62. — Tetrabezzla pictlpennis (K. ): A, female antenna; B, female palpus; C, male palpus; D, female wing; E, female head; F, mandible; G, female legs; H, enlarged hind tibial comb; I, female tarsi; J, male tarsi; K, female fifth tarsomeres and claws; L, male fifth tarsomeres; M, female spermathecae ; N, female abdomen; 0, male genitalia; P, male parameres.

PAGE 213

202 Figure 63. — Heteromyia fasciata Say: A, female antenna; C, female palpus; E, female wing; G, female head; H, mandible; I, female legs; J, enlarged hind tibial comb; K, female tarsi; M, female fifth tarsomeres and claws. Heteromyia sp. (male): B, male antenna; D, male palpus; F, male wing; L, male tarsi; N, male fifth tarsomeres; 0, male genitalia; P, male parameres.

PAGE 214

203 Figure 6k. — Pellucidomyia sp. : A, female antenna; B, proximal segments of male antenna; C, female palpus; D, male palpus; E, female wing; F, female head; 0, female legs; H, enlarged hind tibial comb; I, female tarsi; J, male tarsi; K, female fifth tars one res and claws; L, male fifth tarsomeres; M, female spermathecae; N, female abdomen; 0, male genitalia; P, male parameres.

PAGE 215

Figure 65. — Neurohelea nigra Wirth: A, female antenna; B, male antenna; C, female palpus; D, male palpus; E, female wing; F, female head; G, mandible; H, female legs; I, enlarged hind tibial comb; J, female tarsi; K, female fifth tarsomeres and claws; L, female spermathecae; M, male genitalia; N, male parameres.

PAGE 216

205 Figure 66. — Johannse nomyia argentata (Loew)t A, female antenna; B, male antenna; C, female palpus; D, male palpus; E, female wing; F, female head; 0, anterior mesonotum; H, female legs; I, enlarged hind tibial comb; J, female tarsi; K, female fifth tarsomeres and claws; L, male fifth tarsomeres; M, female spermathecae; N, female abdomen; 0, male genitalia; P, male parameres.

PAGE 217

206 H Figure 67. — Xenohelea tonnoiri Lee J A, female antenna; B, female palpus; C, female wing; D, female head; E, female legs; F, enlarged hind tibial comb; G, female tarsi; H, female fifth tarsomeres and clavs; I, female spermathecae; J, ventral view of female abdomen.

PAGE 218

207 Figure 68. — Jenkins helea magnipennis (Joh. ): A, female antenna; B, male antenna; C, female palpus; D, female wing; E, female head; F, female legs; G, enlarged hind tibial comb; H, female tarsi; I, female fifth tarsomeres and claws; J, male fifth tarsomeres; K, female spermathecae; L, female abdomen; M, male genitalia; N, male parameres; 0, female pupal respiratory organ; P, male pupal respiratory organ; Q, female anal segment; R, male anal segment.

PAGE 219

208 A Figure 69.— Probezzla pallida Malloch: A, female antenna; B, male antenna; C, female palpus; D, male palpus; E, female wing; F, female head; 0, female legs; H, enlarged hind tibial comb; I, female tarsi; J, female fifth tarsomeres and claws; K, male fifth tarsomeres; L, female spermathecae; M, female abdomen; N, male genitalia; 0, male paramere; P, pupal respiratory organ; Q, male anal segment.

PAGE 220

209 Figure 70. — Calyptopogon glbbosus (WLed.)j A, female antenna; B, male antenna; C, female palpus; D, male palpus; E, female wing; F, female head; G, anterior mesonotum; H, female legs; I, enlarged hind tibial comb; J, female tarsi; K, male tarsi; L, female fifth tarsomeres and claws; M, male fifth tarsomeres; N, female spermathecae; 0, female abdomen; P, male genitalia; Q, male parameres.

PAGE 221

210 Figure 71 » — Sphaeromlas ( Sphaeromias ) longipennls (Loew) : A, female antenna; B, male antenna; C, female palpus; D, male palpus; E, female wing; F, female head; G, anterior mesonotum; H, female legs; I, enlarged hind tibial comb; J, female tarsi; K, female fifth tarsomeres and claws; L, male fifth tars ome res; M, female spermathecae; N, ventral view of female abdomen; 0, male genitalia; P, male paramere; Q, pupal respiratory organ; R, pupal abdominal segment; S, female anal segment; T, male anal segment.

PAGE 222

Figure 72. — Sphaerondaa ( Homohelea ) barkudensis Edwards: A, female antenna; B, female palpus; C, female wing; D, female head; E, female legs; F, enlarged hind tibial comb; 0, female tarsi; H, female fifth tarsomeres and claws; I, female spermathecae; J, ventral view of female abdomen.

PAGE 223

212 -:r~--x.--.->.. A Figure 73 . — Malloc hohelea albibasis (Mall.): A, female antenna; B, male antenna; C, female palpus; D, male palpus; E, female wing; F, female head; G, female legs; H, enlarged hind tibial comb; I, female tarsi; J, female fifth tarsomeres and claws; K, male fifth tarsomeres; L, female spermathecae; M, female abdomen; N, male genitalia; 0, male paramere.

PAGE 224

213 Figure 7^»— Nilobezzia schwarzil (Coq.): A, female antenna; B, male antenna; C, female palpus; D, female wing; E, female head; F, female legs; G, enlarged hind tibial comb; H, female tarsi; I, female fifth tarsomeres and claws; J, male fifth tarsomeres; K, female spermathecae; L, female abdomen; M, male genitalia; N, pupal respiratory organ; 0, pupal seventh abdominal segment and glandular disc; P, pupal anal segment.

PAGE 225

Figure 75 . — Stenoxenus inslgninervis Macfie: A, female antenna; B, male antenna; C, female palpus; D, male palpus; E, female wing; F, male wing; G, female head; H, mandible; I, female legs; J, enlarged hind tibial comb; K, female tarsi; L, female fourth and fifth tarsomeres and claws; M, female spermathecae; N, female abdomen; 0, male genitalia; P, male paramere.

PAGE 226

215 Figure % . — Paryphoconus angustipennis Enderlein: A, female antenna; B, male antenna; C, female palpus; D, male palpus; E, female wing; F, male wing; G, anterior mesonotum; H, female legs; I, enlarged hind tibial comb; J, female tarsi; K, female fifth tarsomeres and claws; L, female spermatheca; M, female abdomen; N, male genitalia; 0, male parameres.

PAGE 227

Figure 77. — Pachyhelea pachymera (Will.): A, female antenna; B, female palpus; C, female -wing; D, female eye separation; E, mandible; F, front view of mesonotum; G, female legs; H, enlarged hind tibial comb; I, female tarsi; J, female fifth tarsomeres and claws; K, female spermathecae; L, female abdomen; M, male genitalia. *

PAGE 228

Figure 78. — Palpomyia plebela (Loew): A, female antenna; B, male antenna; C, female palpus; D, male palpus; E, female wing; F, female head; G, female legs; H, enlarged hind tibial comb; I, female tarsi; J, female fifth tarsomeres and claws; K, female apermathecae; L, female abdomen; M, male genitalia; N, male paramere.

PAGE 229

Figure 79.— Bezzia ( Bezzia ) setulosa (Loew): A, female antenna; B, male antenna; C, female palpus; D, male palpus; E, female wing; F, female head; G, mandible; H, female legs; I, enlarged hind tibial comb; J, female tarsi; K, female fifth tarsomeres and claws; L, female spermathecae; M, female abdomen; N, male genitalia; 0, male paramere; P, pupal respiratory organ; Q, pupal anal segment.

PAGE 230

219 c D Figure 80. — Bezzla (Pseudobezzia) flavitarsis (Mall.): A, female antenna; B, male antenna; C, female palpus; D, male palpus; E, female ving; F, female legs; G, enlarged hind tibial comb; H, female tarsi; I, female fifth tarsomeres and clavs; J, female spermathecae; K, male genitalia; L, male paramere.

PAGE 231

Figure 81.— Phaenobezzia plstiae (I. & M, ): A, female antenna; B, male antenna; C, female palpus; D, male palpus; E, female wing; F, female head; G, female legs; H, enlarged hind tibial comb; I, female tarsi; J, female fifth tarsomeres and claws; K, male fifth tarsomeres; L, female spermathecae; M, female abdomen; N, male genitalia; 0, male paramere.

PAGE 232

221 Figure 82. — Genus A: A, female antenna; B, female palpus; C, female wing; D, female head; E, mandible; F, anterior mesonotum; G, female legs; H, enlarged hind tibial comb; I, female tarsi; J, female fifth tarsomeres and claws; K, male fifth tarsomeres; L, female spermatheca; M, male genitalia.

PAGE 233

222 Figure 83. --Genus B: A, female antenna; B, male antenna; C, female palpus; D, male palpus; E, female wing; F, female eye separation; G, mandible; H, anterior mesonotum; I, female legs; J, enlarged hind tibial comb; K, female tarsi; L, female fifth tarsomeres and claws; M, male fifth tarsomeres; N, female spermathecae; 0, female abdomen; P, male genitalia.

PAGE 234

223 Figure 84. — Genus C: A, female antenna; B, female palpus; C, female wing; D, female head; E, mandible; F, anterior mesonotum 0, female legs; H, enlarged hind tibial comb; I, female tarsi; J, female fifth tars ome res and claws; K, female spermathecae.

PAGE 235

224 Figure 85. — Genus D sp. 1: A, female antenna; B, male antenna; C, female palpus; D, male palpus; E, female wing; F, female head; G, mandible; H, female legs; I, enlarged hind tibial comb; J, female tarsi; K, male tarsi; L, female fifth tars ome res and claws; M, male fifth tarsomeres; N, female spermathecae; 0, female abdomen; P, male genitalia.

PAGE 236

Figure 86. — Genus D sp. 2: A, female antenna; B, female palpus; C, female wing; D, female head; E, mandible; F, anterior mesonotum; G, female legs; H, enlarged hind tibial comb; I, female tarsi; J, female fifth tarsomeres and claws; K, male fifth tarsomeres; L, female spermathecae; M, female abdomen; N, male genitalia.

PAGE 237

226 Figure 87. — Genus E: A, female antenna; B, female palpus; C, female wing; D, female head; E, mandible; F, anterior mesonotum; G, female legs; H, enlarged hind tibial comb; I, female tarsi; J, female fifth tarsomeres and claws; K, female spermatheca; L, female abdomen.

PAGE 238

227 Figure 88. — Genus F sp. 1: A, female antenna; B, male antenna; C, female palpus; D, male palpus; E, female wing; F, female head; G, mandible; H, anterior mesonotum; I, female legs; J, enlarged hind tibial comb; K, female tarsi; L, female fifth tarsomeres and claws; K, male fifth tarsomeres; N, female spermathecae; 0, male genitalia; P, pupal respiratory organ; Q, pupal anal segment.

PAGE 239

Figure 89. — Genus F sp. 2: A, female antenna; B, male antenna; C, female palpus; D, male palpus; E, female wing; F, female eye separation; G, mandible; H, female legs; I, enlarged hind tibial comb; J, female tarsi; K, female fifth tarsomeres and claws; L, male fifth tarsomeres; M, female spermathecae; N, male genitalia; 0, pupal respiratory organ; P, pupal abdominal segment; Q, pupal anal segment.

PAGE 240

229 Figure 90.— Genus G: A, female antenna; B, female palpus; C, female wing; D, mandible; E, anterior mesonotum; F, female legs; G, enlarged hind tibial comb; H, female tarsi; I, female fifth tarsomeres and claws; J, female spermatheca; K, female abdomen.

PAGE 241

230 Figure 91. — Genus H: A, female antenna; B, female palpus; C, female wing; D, female head; E, mandible; F, anterior mesonotum G, female legs; H, enlarged hind tibial comb; I, female tarsi; J, female fifth tarsomeres and claws; K, female spermathecae; L, female abdomen.

PAGE 242

231 Figure 92. --Genus J: A, female antenna; B, female palpus; C, female wing; D, female head; E, mandible; F, female legs; G, enlarged hind tibial comb; H, female tarsi; I, female fifth tar some res and claws; J, female spermathecae.

PAGE 243

232 Figure 93. — Genus K: A, female antenna; B, female palpus; C, female wing; D, female head; E, mandible; F, anterior mesonotum; G, female legs; H, enlarged hind tibial comb; I, female tarsi; J, female fifth tarsomeres and claws; K, female spermathecae; L, female abdomen.

PAGE 244

233 Figure 9^. — Genus L: A, female antenna, B, female palpus; C, female wing; D, female head; E, mandible; F, female legs; G, enlarged hind tibial comb; H, female tarsi; I, female fifth tarsomeres and claws; J, female spermatheca; K, female abdomen.

PAGE 245

Figure 95.— Genus Ms A, female antenna; B, male antenna; C, female palpus; D, male palpus; E, female wing; F, female head; G, mandible; H, female legs; I, enlarged hind tibial comb; J, female tarsi; K, female fifth tarsomeres and claws; L, male fifth tarsomeres M, female spermatheca; N, male genitalia.

PAGE 246

Figure 96. — Genus N: A, female antenna; B, female palpus; C, female wing; D, female legs; E, enlarged hind tibial comb; F, female tarsi; G, female fifth tarsoraeres and claws; H, female spermatheca.

PAGE 247

236 Figure 97. — Genus 0: A, female antenna; B, female palpus; C, female wing; D, mandible; E, female legs; F, enlarged hind tibial comb, G, female tarsi; H, female fifth tarsomeres and claws; I, female spermathecae.

PAGE 248

237 G Figure 98.— Genus P: A, female antenna; B, female palpus; C, female wing; D, female head; E, mandible; F, female legs; G, enlarged hind tibial comb; H, female tarsi; I, female fifth tarsomeres and claws; J, female spermathecae.

PAGE 249

238 A Figure 99. — Genus Q: A, female antenna; B, male antenna; C, female palpus; D, female wing; E, female head; F, male head; G, female legs; H~ enlarged hind tibial comb; I, female tarsi; J, female fifth tarsomeres and claws; K, female spermatheca; L, male genitalia.

PAGE 250

239 Figure 100. — Genus R: A, female antenna; B, male antenna; C, female palpus; D, male palpus; E, female wing; F, female head; G, mandible; H, female legs; I, enlarged hind tibial comb; J, female tarsi; K, female fifth tarsomeres and claws; L, female spermathecae; M, male genitalia

PAGE 251

Figure 101. — Genus S: A, male antenna; B, male palpus; C, male ving; D, male legs; E, male tarsi; F, male fifth tarsomsres and claws G, male genitalia.

PAGE 252

241 Figure 102. — Genus T: A, female antenna; B, male antenna; C, female palpus; D, male palpus; E, female wing; F, female head; G, mandible; H, female legs; I, enlarged hind tibial comb; J, female tarsi; K, female fifth tarsomeres and claws; L, male fifth tarsomeres; M, female spermathecae; N, female abdomen; 0, male genitalia; P, male paramere.

PAGE 253

LITERATURE CITED Arnaud, P. 1956. The heleid genus Cullcoides in Japan, Korea And Ryukyu Islands. Microentomology 21j85-207. Barbosa, F. A. S. 19^7. Cullcoides (Diptera: Heleidae) da Regiso Neotropica. Ann. Soc. Biol. Pernambueo (Brazil) 7:3-30. Boesel, W. M., and E. G. Snyder. 19^. Observations on the earlystages and life history of the grasspunky, Atrlchopoeon levis (Coquillett). Ann. Ent. Soc. Amer. 37:37/+ 6« Buckley, J. J. C. 1938. On Culicoldes as a vector of Onchocerca gibsoni (Cleland and Johnston, 1910). J. Helminthol. 16:121158. Campbell, J. A., and E. C. Pelham-Clinton. 1959-1960. A taxonondc review of the British species of Culicoldes Latreille (Dipteraj Ceratopogonidae). Royal. Soc. Edinburgh Proc. Sect. B., Biol. 67:181-302. Carter, H. F. 1921. A revision of the genus Leptoconops Skuse. Bull. Ent. Res. 12:1-28 (Diptera). Carter, H. F., A. Ingram, and J. W. S. Macfie. 1920-1922. Observations on the Ceratopogonine midges of the Gold Coast with descriptions of new species. I-IV. Ann. Trop. Med. & Parasit. lkt 187— 27^, 309-331 5 15:177-212. 5 pis. Causey, 0..R.. 1938. Culicoides of Siam with descriptions of new species. Amer. J. Hyg. 23:399-^16. 8 pis. (Dipt.). Chan, K. L., and E. J. LeRoux. 1965. Description of Forcipomvia ( Neoforcipomyia ) eques (Johannsen) (Diptera: Ceratopogonidae), with an account of the digestive and reproductive systems. Phytoprotection 46(2) :7*J-104. Illus. (French summary). Chan, K. L., and L. G. Saunders. I965. Forcipomyia ( Dacnoforcipomyia ) anabaenae . a new blood-sucking midge from Singapore, described in all stages (Diptera: Ceratopogonidae). Can. J. Zool. ^3:5275^0. Illus. Chanthawanich, N., and M. Delfinado. 1967. Some species of Leptoconops of the Oriental and Pacific regions (Dipteral Ceratopogonidae). J. Med. Ent. 4j29^-303. ' 2^2

PAGE 254

243 Clastrler, J. 1958a. Notes sur les Ceratopogonides. IH. Culicoides semimaculatus n. sp. d' Algeria. Archs. Inst. Pasteur Alger., Algiers. 36:55-60. ; i . 1958b. Notes sur les Ceratopogonides IV, V. Ceratopogonides d'Afrique Occidentale Francaise (1,2). Ibid. 36:192-258, 52 figs; 487-505. 15 figs. . 1959a. Notes sur les Ceratopogonides. VI. Ceratopogonides d^rique Occidentale Francaise (3). Ibid. 37:167-197. . 1959b. Notes sur les Ceratopogonides. VH. Ceratopogonides de 1' Afrique Occodentale Francaise (4). Ibid. 37:340-383. . 1959c Notes sur les Ceratopogonides. VUI. Ceratopogonides de 1« He de la Reunion. Ibid. 37:412-446. . 196I. Notes sur les Ceratopogonides XV. Ceratopogon et Alluaudomyia de la region palearctique. (Notes on the Ceratopogonidae XV Ceratopogon and Alluaudomyia of the palearctic region.) Ibid. 39:401-438. Illus. . 1962a. Notes sur les Ceratopogonides. XVI. Especes du genre Bezzia Kieffer ou apparentees de la region, palearchique. Ibid. 40:53-125. 32 figs. . 1962b. Notes sur les Certopogonides XVH. Nouveaux Palpomyia Meig. et Johannsenomyia Mall, de la region palearctique. (Notes on the ceratopogonids XVII. New species of Palpomyia Meig. and J ohannsenomyia Mall, of the palearctic region.) Ibid. 40:225-288. Illus. . 1963. Notes sur les Ceratopogonides XVITI. Expeces du genre Stilobezzia Kieffer ou apparentees de la region palearctique. Ibid. 4:41-68. Clastrier, J., and W. W. Wirth. 1961. Notes sur les Ceratopogonides XHI & XIV. Ceratopogonides de la region ethiopienne. Ibid. 39:190-240. 32 figs.; 302-337. 24 figs. Coquillett, D. W. 1900. Papers from the Harriman Alaska Expedition. IX. Entomological results (3):Diptera. Proc. Wash. Acad. Sci. 2:389-464. . 1901. New Diptera in the U. S. National Museum. Proc. U. S. Nat. Mus. 23:593-618. . 1902a. Three new species of nematocerous Diptera. Ent. News 13:84.

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Coquillett, D. W. 1902b. New Diptera from North America. Proc. U. S. Nat. Mus. 25:83-126. . 1905. New nematocerous Diptera from North America. J. N. Y. Ent. Soc. 13:56-69. Costa Lima, A. da. 1937a. Chave das especies de Culicoides da reglao Neotropica. Mem. Inst. Oswaldo. Cruz. 32:411-22. . 1937b. Primeira especie americana do genero Pterobosca (Diptera: Ceratopogonidae) . Mem. Inst. Oswaldo Cruz. 32:615616. 2 pis. Das Gupta, K. S., and W. W. Wirth. 1968. Revision of the Oriental Species of Stilobezzia Kieffer (Diptera: Ceratopogonidae). U. S. Nat. Mus. Bull. 283. 143 pp. Delfinado, M. D. I96I. The Philippines biting midges of the genus Culicoides (Diptera: Ceratopogonidae). Fieldiana. Zool. 33: 629-675. Dove, W. E., D. Q. Hall, and L. B. Hull. 1932. The salt marsh sand fly problem ( Culicoides ). Ann. Ent. Soc. Amer. 15:505-527. 3 pis. 2 figs. (Dipt.) .Downes, J. A. 1950. Habits and life-cycle of Culicoides nubeculosus . Nature 166:510-511 (Dipt.) Downes, J. A., and D. S. Kettle. 1952. Descriptions of three species of Culicoides Latreille (Diptera: Ceratopogonidae) new to science together with notes on a revised key to the British species of the pulicaris and obsoletus groups. Proc. Roy. Ent. Soc. London 21:61-78. Edwards, F. W. 1920a. On the use of the generic name Ceratopofcon Meigen (Diptera: Chironomidae) . Ann. Mag. Nat. Hist. 6:127-130. . 1920b. Some records of predaceous Ceratopogonidae (Diptera). Ent. Mo.. Mag. 6:203-205. . 1926. On the British biting midges (Diptera: Ceratopogonidae). Trans. Ent. Soc. London 74:389-426. 2 pis. 3 figs. . 1929. Philippine Nematocerous Diptera H, m. Notul. Ent. Helsingfors. 9:1-14, 70-78. 2 fig. I . 1933. Die Ausbeute der deutschen Chaco Expedition 1925-26 Diptera. XXXiX-XLi. Cecidomyiidae, Chironomidae, Ceratopogonidae. Konowia., Vienna. 12:86-88. Enderlein, G. 1936. ZweiflUgler: Diptera. Tierwelt Mittelour. 6, HI Tell, 2 Lief. XVi. 259 PP. 317 figs.l

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245 Ewen, A. B., and L. 0. Saunders. 1958. Contributions toward a ' revision of the genus Atrichopogon based on characters of all stages (Diptera: Heleidae). Can. J. Zool. 36:671-724. 21 figs. Fiedler, 0. G. H. 1951. The South African biting midges of the genus Culicoides (Ceratopogonidae) (Diptera). Onderstepoort J. Vet, Res. 25:1-33. Foote, R. H., and H. D. Pratt. 1954. The Culicoides of the eastern United States (Diptera, Heleidae). Publ. Health Monogr. Wash., D. C. 18:1-53. 126 figs. Forattini, 0. P. 1957. Culicoides da Regiao Neotropical (Diptera; Ceratopogonidae). Arg. Fao. Hig. Univ. Sao Paulo. 11: 161-526. 132 figs. Forattini, 0. P., and E. X. Rabello. 1956. As formas imaturas de J Culicoides guyanensjs Floch. e Abonnene, 1942 e de algemas especies de Stilobezzia . Rev. Brasil. Ent. 6:43-50. Forattini, 0. P., E. X. RabeLty and D. Pattoli. 1956. Nota sobre a larva e pupa de Culicoides ins ignis Lutz, 1913 (Diptera: Ceratopogonidae). Ibid. 4:195-198. 7 figs. . i960. Sobre as formas imaturas de Culicoides recticulatus Lutz e de Stilobezzia panamensis Lane e Forattini (Diptera: Ceratopogonidae). Ibid. 9:1-4. 3 figs. I Fox, I. 1942. The respiratory trumpet and anal segment of the pupae of some species of Culicoides (Diptera: Ceratopogonidae). Puerto Rico J. Pub. Health & Trop. Med. 17:412-425. 24 figs. . 1946. A review of the species of biting midges or Culicoides from the Caribbean Region (Diptera: Ceratopogonidae). Ann. Ent, Soc. Amer. 39:248-258. . 1955. A catalogue of the bloodsucking midges of the Americas ( Culicoides , Leptoconops and Lasiohelea ) with keys to the subgenera, index and bibliography. J. Agric. Univ. Puerto Rico. 39:214-285. Freeman, P. 1954. East African Chironomidae and Ceratopogonidae (Dipt.). (Ergebnisse der Deutschen Zoologischen Ostafrika Expedition 1951-52, Gruppe Lindner-Stuttgart, Nr. 9). Arch. Hydrobiol. 48:441-446. 1 fig. ^*rew, J. G. H. J. G. H. 1923. On the larval and pupal stages of Forcipomyia piceus Winn. Ann. Appl. Biol. 10:^09-441.

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246 Galukhova, V. M. 1962. Biting midges ( Diptera j Heleidae) of Karelia. (In Russina). Trudy Zool. Inst. Adad. Nauk SSSR, Leningrad. 31:197-249. 21 figs. Goetghebuer, M. 1920. Ceratopogoninae de Belgique. Mem. Mus. Roy. Hist. Nat. Belg. 8:1-116. . 1921. Chironomides de Belgiques. Brvocelles. 8 fasc. 4:1-208. 233 figs. Mem. Mus, Belgique . 1925. Notes biologiques et morphologiques sur Dasyhelea bilineata Goetgh. Encyc. Ent. Ser. B. H. Dipt. 1:121-124. 1933* Ceratopogonidae et Chironomidae du congo Beige. Rev. Zool. Bot. Afr., Brussels. 24:129-151. 42 figs. . 1934. Ceratopogonides de Chironomides du Congo Beige. 2 e Note. Rev. Zool. Bot. Africaines 25:191-205. 1938. Note sur quelques Ceratopogonides de Belgique. Bull. Ann. Soc. Ent. Belg. 78:375-380. 9 figs. 1948. Ceratopogonidae 1 (Diptera: Nematocera). Explor. Pare. Nat. Albert. Miss, de Witte (1933-35). Brussels fasc. 55:21 pp. 7 figs. Goetghebuer, M., and F. Lenz. 1933» 1934. Heleidae (Ceratopogonidae). In Under, Flieg. Pal. Reg., Fan. 13, 77:1-48; 78:49-133. 12 pis. OutseviVh, A. V. 1952. Contribution to the fauna of sandflies of the genus Cailicoides of the forest zone (Diptera: Heleidae). ' Parasit. Symp. Zool. Inst. Parasit. Sec. 14:75-94. . i960. Bloodsucking midges (Diptera: Heleidae) of the fauna of the USSR. Akad. Nauk. of the USSR Detector. No. 72: 3-130. Haeselbarth, E. I965. Phaenobezzia . a new genus of biting midges. S. Afr. Inst. Med. Res. 55:297-324. Hill, M. A. 1947. The life-cycle and habits of Cullcoldes impunctatus Goetghebuer and Culicoides obsoletus Meigen, together with some observations on the life-cycle of Culicoides obldilis Austen, Culicoides pallirticornis Kieffer, Culicoidqs eubi talis Edwards, and Culicoides chioptcrus Moigon. Ann. Trop. Mod. & Parasit. 41:55-115. 12 figs. 6 graphs (Dipt.). Hoffman, W. A. 1924. The presence of an eversible gland in a midge. Proc. Ent. Soc. Wash. 26:144. . 1925. A review of the species of Culicoides of North and Central America and the West Indies. Amer. J. Hyg. 5:274-301.

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22*7 Hoffman, W. A, 1926a. Two new species of American Leptoconops (Diptera: Chironomidae). Bull. Ent. Res. 17:133-136. 1926b. Notes on Ceratopogoninae (Diptera). Proc. Ent. Soc. Wash. 28:156-159. . 1939. Culicoides filariferus. new J. Pub. Health & Trop. Med. 15:172-174 species. Puerto Rico Hubert, A. A., and W. W. Wirth. I96I. Key to the Culicoides of Okinawa and description of two new species (Diptera: Ceratopogonidae). Proc. Ent. Soc. Wash. 63:235-239. Ingram, A., and J. W. S. Macfie. 1921. West African Ceratopogoninae. Ann. Trop. Med. & Parasit. 15:313-37^. 1 pi. 23 figs. . 1922. West African Ceratopogonidae. Part II. Ibid. 16:243-282. I923. Notes on some African Ceratopogonidae. Bull. Ent. Res. 14:41-74. 1931. Diptera of Patagenia and South Chile based mainly on material in the British Museum (Natural History). Part H. Fasc. 4. Ceratopogonidae. London Br. Mus. (Nat. Hist.): 155-232. 21 figs. James, M. T. 1943. The genus Culicoides in northern Colorado (Diptera: Ceratopogonidae). Pan. Pac. Ent. 19:148-153. Jamnback, H., and W. W. Wirth. I963. The species of Culicoides related to obseletus in eastern North America. (Diptera: Ceratopogonidae). Ann. Ent. Soc. Amer. 56:185-198. Jamnback, H., W. J. Wall, and D. L. Collins. 1958. Control of Culicoides melleus (Coq.) (Diptera: Heleidae) in small plots, with brief descriptions of the larvae and pupae of two coastal Culicoides . Mosquito News 18:64-70. 11 figs. Jobling, B. 1929. Some blood-sucking midges of the genus Culicoides from North London. Tran. R. Soc. Trop. Med. Hyg. 22:304-305. . 1953* On the blood-sucking midge Culicoides vexans Stager, including the description of its eggs and the first-stage larvae. Parasitology 43:148-159. 4 figs. (Dipt.). Johannsen, 0. A. 1905. Aquatic nematocerous Diptera. U. Bull. N. Y. State Mus. 86:76-327. . 1908. New North American Chironomidae. Ibid. 124: 264-285.

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248 „ n a io2?a Kacrooeaa and its allies (Chironomidae: Johannsen, 0. A. i^/a. ii^at Diptera). Ent. Mitt. Berlin. 16:423-425. 1927b. The genus Stcnoxenus. (Chironomidae: Diptera). Ent. News 33:70-72. . 1932. Ceratopogoninae from the Malayan Region of the S&h East Indies. Arch. Ilydrobiol. Suppl. 9^3-^5 rfL* (Dipt.). I93L. New soecies of North American Ceratopogonidae and "Chironomidae. J. & *. Ent. Soc. 42:343-3531942. Caratonogon albarius, Coquillett and related species "(Diptera: Ceratopogonidae). Ent. News 53:76-77. iota a ^Bttric svnoosis of the Ceratopogonidae (Heleidae) of the Americas, a ux^y^ v , species. Ann. Ent. Soc Aner. 36:763-79-. 19^2 Guide to the insects of Connecticut. Part VI. The ^7+CiT , a nT . +r. U e flies of Connecticut. Fxxtn lascicie. -'-^ 149-175. 3 ?ls. Jones, R. H. 1956. New species of C^Acoides from Wisconsin. Free. 4 Ent. Soc. Wash. 58:25-331961a. Observations on the larval habitats of some North African species of Culicoidgs (Diptera: Ceratopogonidae). Ann. .weriudn ouevj-^j ~— — , — _ Ent. Soc. Aner. 5^:702-710. i . 1961b. Descriptions of pupae of thirteen Nor^ ^erican ~ecies of Culicoides (Diptera: Ceratopogonidae). Ibid. 729-745. Rattle D S. 1962. The bionomics and control of Culicoides and L^cLps (Diptera: Ceratopogonidae, Heleidae). Ann. Rev. .nt. 7:401-418. Kfettte D S and J. VI. K. Lawson. 1952. The early stages of British allied genera. Bull. Ent. Res. 43:421-467. 6 pis. 123 fig?. Khalaf , X. T. 1957. Light-trap survey of ^M^^^^^ (Diptera: Heleidae). Amer. Midi. Nat. Notre Dame. 58.182-221 22 figs. (Dipt.) IHeffer J J. 1899. Description d'un nouveau genre et tableau des Snres euro P iens de la famille des Chironomides. Bull. Soc. Ent, France, p. 66-70.

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2^9 Kicffer, J. J. 1906. Diptera, Fam. Chironomidae. In •ytsman. Gen. Insectorum. fasc. 42, 73 pp. . 190 3. Description d'une espece nouvelle de CMxonoaWee d'Egypte. Ann. Mus. Hat. fiang. 6:576-577. . 1911, Description de nouvcaux Chironomides de l* Indian feemde Calcutta. Rec. Ind. Mas, 6:113-177. 2 pi.. 1013a. Nouvelle etude sur les Chironomides de 1» Indian KeUm de Calcutta. Rec. Ind. No*. 9*119-197. 2^. .UUVU.ll V»w — ' 1913b. Chironomides et Cecidomyidae. Voyage Ch. Alluaud — R. Jeannel en Afriquc Oriental* (1911-1912). Res. Sex. Ins. Dipt! 1:1-43. . 1917. Chironomides d'Amerique conserves au M^es Katibnal l£n"grois de Budapest. Aim. Mas. Nat. Hung. i5:292-3^. ; 1918. Chironomides d'Afrique et d'Asie. Ann. Mqs. Nat. Hum. 16:31-136. 1019a. Chironomides d'Europe conserves au T-^es National l^rois ^ Budapest. Ann. Mas, Nat. Hungarici. 17:l-l60. . 1919b. Observations sur les ^ronomides (DipO decrits J. R. Kalloch. Bull. Soc. Ent. France 32. x91-19*. V 1921a. Etude sur les Chironomides de Formose. Ann. Soc. Linn. Lyon. 68:149-163. Urribe des Ceratopogoninae. Arch. Inst, easxavr **» H. la lrioe uea aw^ b — -Algiers and Tunis. 1:262-268. 1921c. Sur quelques Dipteres piqueurs de la f 1 ^? 3 . -^^ , 1 p 2< ,+p> :: ^ de 1'Ffrique du Nora. Arcn. Ceratopogoninae. Inst. Pasteur ae ± 1:107-115. 6 fi 2 3 « 1921d. Chironomides de 1'Afrique equatoriale l re partie, T3a£ Soc. Ent. France 90:1-56 pis. 192? Nouvcaux Chironomides piqueurs habitant d'Algerie. Irch. Inst. Pasteur Afr. Nord. 1:494-518. ; 192ft, Chironomides d'Egypte (Dipt.) Bull Soc. R. Ent. Egypte, Cairo. 69 pp. 24 figs. 1925; Dipteres (nematoceres piqueurs): Chironomidae Ceratopogoninae*. Faune de France. 11:1-138. 192 6. Ceratopogonidae:cle de determination des genres. Arch. Inst. Pasteur Alger. 4:96.

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Lane, J. 1944. As especies neotropicais do ienero Cllnohelea Kleffer, 1917 (Diptera: Ceratopogonidae]). Rev. Ent. Rio de Janeiro 15:249-261. 8 figs. . 1946. New Neotropical Ceratopogonidae (Heleidae). (Diptera: Nematocera). Rev. de Entomologia 17. fasc. 1-2: 202-215. . 1947a. A biologia e taxononda de algumas especies dos grupos Forcipomyia , Culicoides . Arguiv. Fac. Hig. Saude Public a Univ. Sao Paulo I:l6l-l62. j . 1947b. Edpecies Brasileiras de Stilobezzia (Dipt. Ceratopogonidae) e Zygoneura stonei , nov. nom. (Dipt. Ifycetophilidae). Rev. Ent. 18:197-214. . 1956. On " Paryphoconus" and ' Stenoxenus" (Diptera: Ceratopogonidae). Rev. Bras. Biol., Rio de Janeiro. 16:299-308. 2 figs. . 196la. Further notes on Neotropical Ceratopogonidae (Diptera). Ibid. 21:37-44. . 1961b. Insecta Amapaensia. Diptera: Ceratopogonidae and Anisopodidae. Studig Entomologica 4:449-452. 1 fig. . 1961c. The genera Stenoxenus and Parypaoconus in the Nootropics (Diptera: Ceratopogonidae). Ibid. 4:452-458. 3 figs. Lane, J., and J. P. Duret. 1954. (71955). Clinohelea da Argentina e do Brasil. (Diptera; Ceratopogonidae). Dusenia Curitiba 5:247-254. Lane, J., and A. P. Forattini. 1958. Neotropical Stilobezzia H. Fourteen new species, chiefly from Panama (Diptera: Ceratopogonidae). Rev. Brasil Ent. 8:203-224. . 196I. Neotropical Stilobezzia Keiff^r, 1911. HI. Key the adults of this genus and description of one new species (Diptera: Ceratopogonidae). Ibid. 10:83-94. Lane, J., and W. W. Wirth. 1964. The biting midge genus Monohelea Kieffer in the Neotropical region (Diptera: Ceratopogonidae). Studia Entomologica 7:209-236. Lane, J., A. P. Forattini, and E. X. Rabello. 1955. Biologia e especies novas de Palpomyia e Stilobezzia (Diptera: Ceratopogonidae). Dusenia 6:81-88. 1 Laws on, J. W. H. 1951. The anatomy and morphology of the early stages v of Culicoides nubeculosus Meigen (Diptera: Ceratopogonldae-Heleidae) Trans. R. Ent. Soc. London 102:511-570. 1 pi. 46 figs. I

PAGE 262

251 Lee D J. 19*8. Australasian Ceratopogonidae (mptera: Ke^tocera) . Part I-W. Proe. Linn. Soc. N. S. V. 72:313-356. 2 pis. . 19*9. Australasian Ceratopo e onidae (Diptera: lle^tocera) . p^Tt V. Tho Palpoinyia group of genera. Ibid. WCf"* J. UJ. " w — | — 1962. Australian Ceratopogonidae (Diptera: Kenatocera). p~t The genus >*acrurohelea . Ibid. 87:339-3*0. rv T ~i v t 1Q<53. Australasian Coratopoconidao Lee, D. J., and h. J. R*y*. W> « spe cies of Cullcoides . (Diptera: Nematocera) . Ft. o. Ausx.raj.ict" Ibid. 77:369-39*. Lindner, E. g& Die Fliegen der Palearlctischen Region. Stuttgart. (Heleidae: Ceratopogonidae) 3'!pp. o jus. LerF R. 193*-. t&* Metamorphose der Heleidae. In Lindner. E. Die Fliegen der Palaearktischen Region. 13a. lief. 7-9^-123. Tutz J 1912. Contribuicao para o estudo das "Ceratopogoninas" 'h^atofagas encontradas no Brasil (parte geral). Hera. Inst. Oswaldo Cruz. *-:l-33. . m3 , Contribuicao para o estudo das ^ratopogoninas" ^tofagas do Brasil (parte sistematica) . Ibxd. 5**5-73? her 3 pis. i9 lft. Contribuicao para o conhecimento d^erat^lnas" — brasil (Aditamento terceiro e descricao des especxes qu nao do crasxx ^*cqL-wuwuvv ^t r rf ^ sugam sangue). Ibid. 6:81-99. 2 pis. ^acfie, J. W. W. 1925. A new blood sucking midge from. Singapore. Bull. Bat. Res. 15:3*9-351. iQ 3 ? a . 'ceratopogonidae from the wrings of dragonflies. Tijdschr. Ent. 75:265-285. 7 figs. j 93 2b. New Zealand biting midges (Diptera: Ceratopogonidae). "E. Trop. Med. & Parasit. 26:23-5310 ^xgs. 193*a. Fauna sumatrensis. Bijdrage no. 75Ceratopogonidae IHptera). Tijdschr. Ent. 77:202-231. 6 figs. 193*-b. Reoort on a collection of Ceratopogonidae from ISTayal Ann. Trop. Med; * Parasit. 28:177-195; 279-293. 9 figs. (Dipt.). # 19 37. Notes on Ceratopogonidae (Diptera). Proc. R. Ent. Soc*. London (3) 6:111-118. 1939a. A key to the species of Ceratopogonidae akin to Sropeza Mg. (Diptera). Trans. R. Ent. Soc. London 39:1-12. 1 pi.* 3 figs.

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252 Macfie, J. W. S. 1939b. Ceratopogonidae. (With an appendix by B. de Meillon) Ruvenzori Exped. 193^35 London, Brit. Mus. (Nat. Hist.) 1:81-107. 2pls. 12 figs. (Dipt.) , 1939c. A report on a collection of Brazilian Ceratopogonidae. Rev. Ent. 10:137-219. . 1940a. The Genera of Ceratopogonidae. Ann. Trop. Med. & Parasit. 3^:13-30. . 1940b. Ceratopogonidae (Diptera) from British Guiana and Trinidad. Parts 1 and 2. With an appendix by J. Smart, Proc. R. Ent. Soc. London 9:179-186, 187-195. 5 figs. . 19^8. Some species of Culicoides (Diptera: Ceratopogonidae) from the State of Chiapas, Mexico. Ann. Trop. Med. & Parasit. 42:67-87. Malloch, J. R. 1914a. Notes on North American Diptera. Bull. 111. State Lab. Nat. Hist. 10:213-243. 3 pis. . 1914b. Synopsis of the genus Probezzia , with description of a new species (Diptera). Biol. Soc. Wash. Proc. 27:137-139. . 1914c. Synopsis of North American species of the genus Bezzla . J. N. Y. Ent. Soc. 22:281-285. . 1915a. The Chirononddae or midges of Illinois. Bull. m. State Lab. Nat. Hist. 10:275-5^3. 23 pis. , 1915b. Some additional records of Chironomidae for Illinois Bull. 111. State Lab. Nat. Hist. 11:303-363. / . 1917. A preliminary classification of Diptera, exclusive V of" Pupipara, based on larval and pupal characters, with keys to imagines in certain families. Part I. Bull. HI. State Lab. Nat. Hist. 12:161-409. 29 pis. 1 Mayer, K. 1933. Die Metamorphose von Forclpomyia ( Apelma ) comis Johannsen. Arch. Hydrobiol. Suppl. XH. Tropische Binnengewassen. IV: 224-238 , 1934. Die Metamorphose der Ceratopogonidae (Dipt.) Ein Beitrag zur Morphologie, Systematik, Oekologie und Biologie der Jugendstadien deisor Dipterenfamilie. Arch. Naturgesch., Leipzig. (N.F.) 3:205-288. 93 figs. 1937. Beobachtung liber blutsaugende Ceratopogoniden. Arb. Morph. Tax. Ent. Berlin-Dahlem 4:231-234. . 1952. Die Macropeza. Gruppe der Heleiden (Diptera: Heleidae). Beitr. Ent. 2:582-5. I

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253 Mayer, K. 1957. Die Beschreibung der Puppen von Dlcrohelea albidorsata de M. und Xenohelea galatea de M. miteinem Uberblick | uber das System der Heleinae (Dipt.: Heleidae). Deut. Ent. Z., Berlin (N.E.) 4:30-33. . 1959. Die Puppen brasilionischer Heleiden (Diptera)., Ibid. 6:230-233. 5 figs. Meillon, B. de 1931. A new species of Forcinomyia (Diptera: Ceratopogonidae) from the Transvaal, with a description of its early stages. Trans. Ent. Soc. London 79:335-3^. . 1936a. South African Ceratopogonidae. Part II. Some new and unrecorded species. Publ. Afr. Inst. Med. Res. 7j141-207. . 1936b. Entomological Studies. Studies on insects of medical importance in South Africa (Part HI). Ibid. 7(38): 129-215. 32 pis. 2 figs. 1 map. . 1937. Studies on insects of medical importance from Southern Africa and adjacent territories (Part 4). Ibid. 7(40):305-*U1. 31 Pis. , 1938. Notes on African Ceratopogonidae (Diptera). Proc. Roy. Ent. Soc. London (B) 7:266-270. , 1939. Notes on Ceratopogonidae (Diptera: Nematocera) from Southern Africa. H. J. Ent. Soc. S. Africa 2:7-17. . 1940. Ceratopogonidae (Diptera: Nematocera) from Southern Africa. Trans. R. Ent. Soc. London 90:455-466. 7 figs. , , 1942a. Ceratopogonidae (Dipt. Nematocera) from Southern Rhodesia. Proc. Rhod. Sci. Ass. 39:113-119. 3 figs. . 1942b. Simuliidae e Ceratopogonidae (Dipt. Nematocera) da Colonia de Mocambique. Lourenco Marques, Estac. Anti. Malar. 27 p. 4 pis. . 1943. New records, and new species of Nematocera (Diptera) from the Ethiopian region. J. Ent. Soc. Afr. 6:90-113. 3 figs. . 1959a. New Ceratopogonidae (Diptera: Nematocera) from Africa. Novos Taxa Ent., Lourenco Marques, no. 13:1-24. . 1959b. Diptera (Nematocera); Ceratopogonidae. In Hanstrom, Brinck and Rudebeck, South African Animal Life VI: 325-355. 17 figs. 1

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25# Meillon, B. do i960. New Ceratopogonidae Kerratooera) from the Subsahara region. J. Ent. Soc. S. Air. 2^.W3 *iu. 9 figs. . 1961. The Madagascar* Ccratopogonidae. Rev. Ent. Itocarribique. 4(1): 37-64. XLlus. Dillon, B. de, and F. Hardy. 195*.. New records and species of biting insects from the Ethiopian region. V. J. BEit. t>oc. d. 17:62-85. 6 figs. Mokerji, S. 1931a. Morphology of the pharynx of female Culicoides, and its taxonomic importance. Nature 127:339-3^. 1 ixg. (Dipt.) 1931b On a new st>ecies of Culicoides (Culicoides claviSSiS tt. nov.) with notes on the morphology of the mouthparts Sa£te tfermlnalta of an Indian Oulicoidos,. Indian J. Bad. Res. 18:1051-1053. 1 pi. (Dipt.) Nagaty, E. F., and T. A. Horsey. l 9 6l. Taxonomic on Culicoides in Egypt. Part i, 3, JArab. Vet. Kas. Ass. 21:185-19^, 345-369. 3 pis. ^elsen A. 1951. Contributions to the metamorphosis and biology of IZ j&Fjj&tem *^ er (Diptera: Ceratopogonxdae) wxth remarks on the evolution and taxonomy of the genus. Bxdr. Skrift., Copenhagen 6:95. 2 pis. 33 figs* Noe G. 1905. rm nueve genera appartenente all famiglia Chironomidae. ' Atti. della Reale Accad. del Lincei, Set. 5, Rendxcontx XW. 114-120. Okada, T. 19^2. Alluadomria (Diptera: Heleidae) from Formosa with a revised key. Trans. Kat. Hist, Soc. Formosa 32.315-320. Ortiz, I. 1953. Sobre una nueva especie del Genero Culicoidcg (Diptera: Ceratopogonidae) proxima de C. horticola Lutz 1913Rev. Sanid. Asist. Soc. 18:807-12. Ortiz, I., and L. A. Leon. 1955Los C ulicoides (Diptera: Ceratooo^onidae) de la Republica del Ecuador. Bol. Inf. Cxentxi. I Nac. n. 67:5^4-90. OrtiI. 1959. Contribucion al estudio de los Ceratopogonidos (Jeiires) hematofages de Venezaela (Leotocoriops Skuso, 1890, Lasiohelea ISeffer, 1921 y Culicoides Latreille, 1809) (Diptera: Ceratopogonidae) Rev. Sanid. Asist. Soc. 24:34-9-369. 5 pis.

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255 3 Painter, R. H. 1926. The biology, immaoure stages and control of V the sandflies (biting Ccratopogoninae) at Puerto Castxlia, Honduras. Ann. Re?. Med. Dept. United Fruit Co., Boston, Mass. 15:245-262. 9 figs(Dipt.) Patel P. C-. 1921. Koto on the life-history of Culicoides, custom * iith some remarks on the early stages of Ceratorxyon. Proc. 4th Ent. Mtg. Pusa. 272 p. Patton, W. S. 1913. Culicoidos. &g££fl& »• s P! a now Indian blood " sucking nidge. Indian J. ifed. Res. 1:33c. Pomerantzen, B. E. 1932. Beitrage sur Iforphologie und Anatomi der Genitalein von Cu licoides (Diptera: Hematocera) (In Russian if*b Gorman stuttnaxy) Hag. Parasit. 3:183-214. 26 figs. Reran. H. 1959. Estonian species of the genus Atricho^o^on Kieffer (Dipt.: Keleidae). I. Subgenus Psilokempia End. Ent. Ogozr., 38:682-692. 36 figs. 1962. The genus Dasyhelea Kieffer in Estonia (Diptera: Heleidae). Tartu. Riik. Ulik. Toim. 120:108. rj . J.V, t m vnn 1022. Die Metamorphose der Culicoidinen (CeratoRexth, J. 1. von. iy<^~» , ^ _ ., o.^nn hh.'i 93 fiss. pogoninen). Arch. f. ^drobiol., Suppl. 3d. 2.377-^3» ^S s Root P. VL , and*. A. Hoffman. 1937. The North American species of \f * Cv-licoides . Amer. J. Pyg. 25:150-176. Saunders L. G. 1924. On the life history and the anatomy of the v early stages of Forciconyia (Diptera: Kemat., Ceratopogonxnae) . Parasitology. 16:154-213. 3 pis. 192^. Or the life history, morphology and systematic ~1 p*ositioii of Aoelma Kieff.', and Thyridomyia n. g. (Diptera: Kemat. Ceratopogoninae) . Ibid. 17:252-277. 1 p. . 1956. Revision of the genus Forci^orvia based on characters 4 of all stages. (Diptera: Ceratopogonidae) . Can. J. Zool. 34:657-705. . 1959. i-bthod for studying Forcipomyia midges -with special references to cacao pollinating species (Diptera: Ceratopogonidae) Ibid. 37:33-57. . 1964. New species of Forcipomyia in the Lasiohelea complex described in all stages (Diptera: Ceratopogonidae). Ibid. 42:463-482. HIus.

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256 Seguy, E. 1931. Sur les affinites des genres Stenoxenus et Macroptilum et description d'une espece nouvelle. (Dipt.: Ceratopogonidae). Bull. Soc. Ent. France 208-211. 3 figs. 193*1.. Bipteres d*Espagne. Etude systematique basie principalement les collections, formees par le R. P. Longin Navas, S. J. Mwu Acad. Cienc. Zaragoza. 315^ P. 7 rigs. Sen, P., and S. K. Das Gupta. 1958. "Dyar's law" in the determination of larval instars in mosquitoes. Bull. Calcutta Sch. Trop. Med. 6:69-70. Skuse, F. A. A. I889. Diptera of Australia, VI. The Chironoraidae. Proc. Linn. Soc. N.S.W. 4:215-3H. Smee, L. I966. A revision of the subfamily Leptoconopinse Noe (Diptera: Ceratopogonidae) in Australia. Aust. J. Zool. 14:993-1025. Smith, L. M„ and H. Lowe. 1948. The black gnats of California. Hilgardia. 18:157-183. Strenzke, K. 1950. Systematik, Morphologie und Ikilogie der terres1 trischen Chironomiden. Arch, f . Ifydrobiol. Suppl. Ed. 18:207-414. Thienemann, A. 1926. Dipteren aus den Salzgewassern von Oldesloe. (Mit einer Bestimmungstabelli der Larven und Puppen der Gattung Dasvhelea .) MLtt. Geogr. Ges. Lubeck (2). 31:102-126. , 1928. Chironomiden ffetamorphosen 1. Arch. IJydrobiol. 19:585-623. 31 figs. (Dipt.) Thomsen, L. 1935. New species of New York State Ceratopogonidae. J. N. Y. Ent. Soc. 43:283-296. . 1937. Aquatic Diptera. Part V. Ceratopogonidae. Cornell Univ. Agr. Exp. Sta. Mom. 210:57-80. 9 pis. Thorpe, W. H. 1927. Culicoides nubeciilosus breeding in the sea. Ent. Mon. Mag. 63:138. Tokunaga, M. 1932. A new biting midge from Japan (Diptera: Ceratopogonidae), -with anatomical notes on the larval head capsule and mouthparts. Trans. Kansai Ent. Soc. (Osaka) 3:1-12. . 1934. Chironomidae from Japan (Diptera). m. The early stages of a seashore biting modge, Forcipomyia crinume (Tokunaga). Philipp. J. Sci. 53:^9-487. . 1937. Sandflies (Ceratopogonidae, Diptera) from Japan. Tenthredo1:233-338. 6 pis.

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25? * bkunaca m. 1939. Japanese biting of Bozzia and Pa ^pory i a roicunaga, n. v 2^73-313. "15 figs* Ceratopogonidae, Diptera,. Ibid. a . 19>40. Biting midges from Japan and ^hborins lauding Hicronesian Islands *** ^ golia (Diptera: Ceratopogonidae) . Ibid. ^5° 10:3 1959. New Guinea biting midges (Diptera: Ceratopogonidae) &f* Insects 1:177-35-3. W fig?. . 1962a. Biting ftddges of the RyuXyu Islands (Diptera: Ceratopogonidae). Ibid. kil&ftfl 1962b. Biting nidges of the genus Culicoidcs from Hew lulr.ea (Diptera : Ceratopogonidae) Ibid. ^7-561. Hlus. 1963. Jfew Guinsa biding midges (Diptera: Ccratcpogcnidae) . Ibid. 5:211-2791Q64. Biting midges of the genus Ceratopogon from New G^ea (Diptera: Ceratopogonidae) . Ibid. 6:2 9 2-299. Hit*. w w j i? re Mrt-achi. 1959. Insects of I'dcronesia. 108-^. 98 figs. ^ r " T m 1893. An interesting blood-sucking gnat of the Townsend, C. H. ±. a.q » l-torphology and Biology. Bull. Soc. Pathol. Bxongue. 1159-1171 (English oummarj-). w p arv 4 j p Adam. 1966. Les Ceratopogonidae (Diptera) ^Ss g-rotSs t ll tSm* ** ^ngo (Brazzaville). Ann. Speleol. 2l(3):7H-773. Viirr-r £ 1932. Keca specie aus der Unterfamiiien Ceratopogoninaa. SbomEnt. Odd. Narod. Mas. Praze. 10:1^0-^. Lilians, R. * 1951. T^e ior^a^e eb^^os or C^ol^oxdea ^g^xg^us / Hoffman (Diptera: Heleidae). -ton. Ent. Soc. Aner. ^30-^0. 2 pis. fcW&. R. tf. 1953. Notes on the bionomics of the m>WZ^ of "" m -Bler County/Borgia. I. Observations on breeding habixats of balla and needhami . Proc. Ent. Soc. Wash. 55 klliston, W. W. 1907. Dipterological notes. J. H. X. Ent. Soc. 15:1.

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258 Winnertz, J. 1852. Beitrage zur Kenntris der Gattung Ceratopogon . Linn. Ent. 6:1-80. Wirth, 0. M., W. W. Wirth, and F. S. Blanton. I968. Plant materials as breeding places of Panama Culicoides . Proc. Ent. Soc. Wash. 70:132. Wirth, W. W. 1951a. The genus Probezzia in North America (Diptera: Heleidae). Proc. Ent. Soc. Wash. 53:25-34. 1 fig. . 1951b. The genus Culicoides in Alaska (Diptera: Keleidae) Ann. Ent. Soc. Amer. 44:75-86. 2 pis. . 1951c. A new biting midge of the genus Leptoconops from Florida, -with new records of other American species (Diptera: Heleidae). Proc. Ent. Soc. Wash. 53:281-28'+. 7 figs. . 195 Id. New species and records of Virginia Heleidae. Ibid. 53:313-326. . 1952. The Heleidae of California. Univ. Calif. Publ. Ent. 9:95-266. 33 figs. . 1953a. Biting midges of the heleid genus Stilobezzia in North America. Proc. U. S. Nat. Mas. 103:47-85. . 1953b. American biting midges of the heleid genus Nonohelea. Ibid. 103:135-5^. . 1956a. The biting midges ectoparasitic in blister beetles. (Diptera: Heleidae). Proc. Ent. Soc. Wash. 58:15-23. 5 figs. . 1956b. The Heleid midges involved in the pollination of rubber trees in America (Diptera: Heleidae). Ibid. 58:241-250. 2 figs. . 1956c. New species and records of biting midges ectoparasitic on insects (Diptera: Heleidae). Ann. Ent. Soc. Amer. 49:356-364. . 1959a. New species and records of Heleidae from Brazil (Diptera). Dtsch. Ent. Z., Berlin (N.F.) 6:234-237. 3 figs. . 1959b. Pashyhelea, a new genus of American Ceratopogonidae related to Palpomyla (Diptera). Bull. Brooklyn Ent. Soc. 54:50-52. . 1959c Systematic Arrangement of Ceratopogonidae (Heleidae). U. S. D. A. (mimeograph, unpublished) 20 p. . i960. The genus Pellucidomyia Macfie (Diptera: Ceratopogonidae). Bull. Brooklyn Ent. Soc. 55:1*

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1 259 i Wirth, W. W. 1962a. A Reclassification of the Palpomyia Eezzia Macropeza Groups, and a Revision of the North American Sphaeromiini (Diptera: Ceratopogonidae). Ann. Ent. Soc. Amer. 55:272-287. . 1962b. The North American species of the biting midge genus Jenkinshelea Macfie (Diptera: Ceratopogonidae). Bull. Brooklyn Ent. Soc. 57: 14. . 1965a. Two new species of T-kcrurohelea from Chile. PanPacific Ent. huhS-so. I . 1965b. A new J ohanns en o myia from Brazil (Diptera: Ceratopogonidae). Proc. Ent. Soc. Wash. 67:^. . 1965c. A revision of the genus Parabezzia Hal loch (Diptera: Ceratopogonidae). Ibid. 67:215-230. Ilius. Wirth, W. W., and F. S. Blanton. 1956. Studies in Panama Colic oldes . VIII. The neotropical species of the Cuttatus group of the subgenus Eoffmania (Diptera: Heleidae). Ent. Soc. Wash. 58: 305-32l£ 10 figs . . 1959. Biting midges of the genus Culicoides from Panama (Diptera: Heleidae). Proc. U. S. Nat. Mas. 109:237-^2. 91 figs. . 1967. The North American Culicoides of the Guttipennis group (Diptera: Ceratopogonidae). Fla. Ent. 50:207-232. HIus. Wirth, W. W., and M. D. Delfinado. 196^. Revision of the Oriental species of Alluaudomvia T&effer (Diptera: Ceratopogonidae). Pacif. Insects 6:599-6^8. Wirth, W. W., and A. A. Hubert. 1959. Trithecoides , a new subgenus of Culicoides (Diptera: Ceratopogonidae). Pacif. Insects 1:1-38 38 figs. . 1960a. C amptopter ohelea a new genus of Ceratopogonidae from the Philippines (Diptera). Fieldiana. Zool. 47(7):89-91. Ulus. . 1960b. Ceratopogonidae (Diptera) reared from cacti, with a review of the Copiosus group of Ctilicoides . Ann. Ent. Soc. Amer. 53:639-658. . I96I. New species and records of Taiwan Culicoides (Diptera: Ceratopogonidae). Pacif. Insects 3:11-26. Ulus. . 1962. The species of Culicoides related to piliferus Root and Hoffman in eastern North America (Diptera: Ceratopogonidae). Ann. Ent. Soc. Amer. 55:182-195. 18 figs.

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260 Wirth W W. and D. J. Log. 1958. Australasian Ccratoposonidae ^i^.fc-loo.M) Part VIH, A new genu, fro* Australia attacking man. Proc. Linn. Soc. H.S.W. 8^7 5 figs. 1969 The genus Paradasyhelea Macfie, with descriptions _ ; wo new snecies from Eastern Australia (Diptera: Ceratopogonxdae). of two new s"cecies . Bull. Brooklyn Ent. Soc. &:Uh-12L 2 pis. Wirth, W. W., and A. Stone. 1956. Awuatio Diptera, In Usinger, Aquatic insects of California 60 pp. 64 figs. Wirth w W and R. W. Williams. 1964. New species and records of Ceratopogonidae). Ann. Ent. Soc. Amer. 57:302-310. Illus. Zilahi-Sebess, G. 1931. Anabiotische Dipteren. Arch. Kydrobiol. 23:310-329. 1940. Heleiden aus Ungarn. (In Magyar with German summary). Folia Ent. Hungar., Budapest 5:10-133. 3 pis. (Ih.?tJ

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BIOGRAPHICAL SKETCH Niphan Chanthawanich Ratanaworabhan vas bom on April 3» 1939* in Chainat, Thailand. She attended primary and secondary classes at the Assumption Convent School and finished her high school at Chulalongkorn University High School in Bangkok. She enrolled at Chulalongkorn University, where she received the degree of Bachelor of Science in Zoology in 1962. Upon her graduation she was employed as an assistant instructor at that institution until 1964-. She was awarded a fellowship from the Center for Cultural and Technical Interchange (Easi>tfest Center) to continue her graduate study at the University of Hawaii in June, 1964. She was granted the degree of Master of Science in 1966. She continued her graduate study at the Department of Entomology and Hematology, University of Florida in September, 1966, where she completed graduate work leading to the degree of Doctor of Philosophy in March, 1969. Niphan married Sawart Ratanaworabhan in August, 1967. 261

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1 This dissertation was prepared under the direction of the chairman of the candidate's supervisory committee and has been approved by all members of that committee. It was submitted to the Dean of the College of Agriculture and to the Graduate Council, and was approved as partial fulfillment of the requirements for the degree of Doctor of Philosophy. March, 1969 ^^Dean, College of Agriculture Dean, Graduate School Supervisory Committee:


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