Citation
A biosystematic review of the bloodsucking psychodid flies of Colombia (Diptera: Phlebotominae and Sycoracinae) /

Material Information

Title:
A biosystematic review of the bloodsucking psychodid flies of Colombia (Diptera: Phlebotominae and Sycoracinae) /
Creator:
Young, David Grier, 1940-
Publication Date:
Copyright Date:
1977
Language:
English
Physical Description:
xiii, 481 leaves : ill. ; 28 cm.

Subjects

Subjects / Keywords:
Female animals ( jstor )
Genitalia ( jstor )
Head ( jstor )
Holotypes ( jstor )
Male genitalia ( jstor )
Pharynx ( jstor )
Pumps ( jstor )
Species ( jstor )
Spermatozoa ( jstor )
Tree trunks ( jstor )
Dissertations, Academic -- Entomology and Nematology -- UF
Entomology and Nematology thesis Ph. D
Insects -- Colombia ( lcsh )
Psychodidae ( lcsh )
Sand flies -- Colombia
Genre:
bibliography ( marcgt )
non-fiction ( marcgt )

Notes

Thesis:
Thesis--University of Florida.
Bibliography:
Bibliography: leaves 451-480.
General Note:
Typescript.
General Note:
Vita.
Statement of Responsibility:
by David Grier Young.

Record Information

Source Institution:
University of Florida
Holding Location:
University of Florida
Rights Management:
Copyright David Grier Young. Permission granted to the University of Florida to digitize, archive and distribute this item for non-profit research and educational purposes. Any reuse of this item in excess of fair use or other copyright exemptions requires permission of the copyright holder.
Resource Identifier:
020620286 ( ALEPH )
03983178 ( OCLC )
AAB5221 ( NOTIS )

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Full Text












A BIOSYSTE MATiC REVIEW OF THE BLOODSUCKING PSYCHODID FLIES OF
COLOMBIA (DiPTERA: PHLi.EBOTOM'jN AE AND SYCORACTNAE)









By

DAVID GRI-R YOUNG

















A DISSERTATIEON PRESENTED 10 THE GRADUATE COUNCIL OF
THE UNIVERSITY OF FLORIDA
IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE
DEGREE OF DOCTOR OF PHILOSOPHY














INIVERSITY OF FLORIDA

1.77





































Dedicated to my father,

Howard G. Young













TABLE OF CONTENTS




LIS O FURES . . . . . . . . . . . . . . . . . . . . . . . viii

A ST AC . . . . . . . . . . . . . . . . . . . . . . . . . . . xii

INTRODUCTION . . . . . . . . . . . . . . . . . . . . . . . . 1

CLASSIFICATION AND TAXONOMIC TREATMENT . . . . . . . . . . . . . 4

Keys to the Subfamilies of New World Psychodidae. . . . . . 6

,GENEV.AL MOkd lOLOGY AND TERMINOLOGY . . . . . . . . . . . . . . . 8

TNEYHODS AND MATERIALS. . . . . . . . . . . . . . . . . . . . . 15

Field Collections . . . . . . . . . . . . . . . . ... .. . . 15
Laboratory Methods. . . . . . . . . . . . . . . . .... .. . 17

DlSTRIBUTION, COLLECTING LOCALITIES, AND CHARACTERISTICS OF
T':L FAJUNA. ................... ........ 19

Pacific Coast Region. . . . . . . . . . . . . . . . . .. . 20
Atlantic Coast Region . . . . . . . . . . . . . . . . ... . 23
Andean Region . . . . . . . . . . . . . . . . . . . .. . . . 24
Oriental Region . . . . . . . . . . . . . . . . . . ..... . 25

MEDICAL IKMORTANCE . . . . . . . . . . . . . . . . . . . . . . . 29

SUBFAMILY SYCORACAE . . . . . . . . . . . . . . . . ... ..... . 31

Keys to the [ycorax Species . . . . ... . . . . . . . . . 33

1. Sycorax andicola n. sp. . . . . . . . . . . . . . 35
2. S!corax colombiensis n. sp. . . . . . . . . . ... 36
3. Sycorax fairchildi n. sp. . . . . . . . . . . . . 38
4. Sy~orax tricpinoso n. sp. . . . . . . . . . . . . 39

SU GRAMILY ' iLE BOTOMINAE. . . . . . . . . . . . . . . .. . . . 50

Key to the New World Cener . . . . . . . . . . . . . . . . 50
Genur WVawily-f1c bertig, 1948 . . . . . . . . . . . . . . . . 52
Keys to Species . . . . . . . . . . . . . . . . ... . . . . 52

5. WaIieia (W.) nig osacculus. . . . . . . . .... 53
6. c.r:e'zueya (W.) rotu ndipennis. .. . . . . . . . . . 5/



iii







Pase

Genus Bri.mntomyia Franca & Parrot, 1921 . . . . . . . . . . . 60
Key to Species . . . . . . . . . . . . . . . . . . . . . . . 60

7. Brumptoia avllar . . . . . . . . . . . . . . . 61
8. BrE ptoryia beaperuyi. . . . . . . . . . . . . 62
9. Brrrptomyn.a galindoi. . . . . . . . . . . . . . ... 63
10. Brsipt ornyia haata. . . . . . . . . . . . . . . ... 65
i1. Brw ptormyia leopoldoi . . . . . . . . . . . . .. . 66

Genus Iutorrpjia Franca, 1924 . . . . . . . . . . . . ... . . 76
Keys to Subgenera, Species Groups, and Ungrouped Species. . . 76
Subgenus Lutzomyjia Franca, 1924.. . . . . . . . . . . . . . 91
Keys to Species . . . . . . . . . . . . . . . . . . . . . . . 91

12. Lutzomyia (L.) bifoliata. . . . . . . . . . . . ... 93
13. Lutsorr.ia (L.) Zichyi . . . . . . . . . . . . . . 94
14. Lutzomyia (L.) Zongipapis. . . . . . . . . . . ... 96
15. Lutzomyia (L.) gomc~i . . . . . . . . . . . . . .. .97
16. Lutzorrjia (L.) marinkel.ei n. sp. . . . . . . . ... 99

Migonei Group Theodor, 1965 . . . . . . . . . . . . . . . .. 112
Keys to Species . . . . . . . . . . . . . . . . . . . . . . . 112

17. Lutzomnyia migonei . . . . . . . . . . . . . ... .113
18. Lutzomnyia marajoensis . . . . . . . . . . . . . .. 114
19. Lutzomrfia sp. de Baduel . . . . . . . . . . . . .. 116
20. Lutzornyia walker . . . . . . . . . . . . . . . .. 116

Saulenuis Group Lewis et al., 1978. . . . . . . . . . . . . 125

21. Lutzomyia saulensis . . . . . . . . . . . . . . . 125

Verrucarum Group Theodor, 1965 . . . . . . . . . . . . . ... 130
Key to Species. . . . . . . . . . . . . . . . . . . . . . .. . 130

22. Lutzomyia odax . . . . . . . . . . . . . . . . .. 133
23. Lutomyia serrana . . . . . . . . . . . . . . . . . 134
24. Lutzomyia andina. . . . . . . . . . . . . ... .. 135
25. Lutzomria columbiana. .... . . . . . . . . . .136
26. Lutzomrnia disiuncta . . . . . . . . . . . . . . .137
27. Lutzonyia evansi. . ... . . . . . . . . . . . . . 137
28. Lutzomjia longiflocosa. . . . . . . . . . . .. . . 138
29. Lutzormnia moralesi n. sp. . . . . . . . . . . . ... 139
30. Luitsonjia nunetovari . . . . . . . . . . . . . .. .41
31. Lutzonq ia ovalcsi. . . . . . . . . . . . . . . .. . 143
32. Lutzomija quasitownsendi. . . . . . .. . . . . . . 145
33. Lutzomyia sauroida. . . . . . . . . . . . . . . . . 145
34. Li.tzonryia spinicrassa . . . . . . . . . . . . . .. 146

Vespertilionis Group Theodor, 1965. . . . . . . . . . . . .. . 161
Key to Species. . . . . . . . . . . . . . . . .. . . . . . . 161




Vj I







Page

35. Lutzomyia isovesportiion . . . . . . . . . . . . 161
36. Lut.omnria vespFeriZioic. . . . .. . . . . . . .. 162

Subgenus Darqpfornyicz Addis, 1945 . . . . . . . . . . . .. . . 167

37. Lutzomyia (D.) rosabi . . . . . . . . . . ... .. 167

Subgenus Pintormyia Costa Lima, 1932 . . . . . . . . . . . .. 171

38. Lutzsoryia (P.) spi:osa. . . . . . . . . . . . . ... 171

Subgenus Pr'ssatia Mangabeira, 1942 .. . . . . . . . . . .176
Keys to Species . . . . . . . . . .... . . . . . . . . . . . 176

39. Lutzorryia (Pr.) capo;i . . . . . . . . . . . . . . 177
40. Lutzsomrria (Pr.) dysponeta . . . . . . . . . ... . 178
41. Lutzoimyia (Pr.) trianntha. . . . . . . . . .. . . 180

Baityi Group Theodor, 1965. . . . . . . . . . . . . . . . .. .186

42. Lutomjyia bait yi. . . . . ... . . . . . . . . .i8

Subgenus Viannamyia Mangabeira, 1941. . . . . . . . . . . ... 190
Keys to Species . . . . . . . . . . . . . . . . . . . . . . . 190

43. Lutzomria (V.) caprit. . . . . . . . . . . . . . . 191
44. Lutzomyia (V.) fu.rcta . . . . . . . . . . . . ... 192
45. Lutzornyia (V.) tubeulata. . . . . . . . . . . ... 194

Shannoni Group Theodor, 1965. . . . . . . . . . . . . . . ... 200
Keys to Species . . .. . . . . . . . . . . . . . . . . . . . 200

46. Lutzormyia abonnenci . . . . . . . . . . . . . . . 202
47. Lutzomyia dasyera. . . . . . . . . . . . . . . ... 203
48. Lutzomyia dendrophyla . . . . . . . . . . . .... . 204
49. Lutzomjia puntigeniculata. . . . . . . . . . . ... 206
50. Lutzoryia shannoni . . . . . . . . . . . . . . . . 208
51. Lutzornia undulata . . . . . . . . . . . . . . .. . 210'

Longispina Group Theodor, 1965... . . . . . . . . . . . . . 224
Keys to Species . . . . . . . . . . . . . . . . . . . .. . . 224

52. Lutzovjyia longispina. . ... . . . . . . . . . . . 225
53. Lutzomyia trirvuZa . . . . . . . . . . . . . . .. 226

Aragaoi Group Theodor, 1965 . . . . . . . . . . . . ... . . 232
Keys to Species . . . . . . . . . . . . . . . . . . . . . . 232

54. Lutzornjia arag i . . . . . . . . . . . . . . . . 234
55. Lut;omyia boa'retboi ar-rrettoi . . . . . . . . . .. 236
56. Lutvi'rrm-ia barrettoi raf:,uscula n. ssp. . . . . . ... 237
57. L'utzo;yia carp ..en.t. .. .............. . .241
58. Lrutzomy'i runoidesK. . . . . . . . . . . . . . . . 243


V







Page

Dreisbachi Group Lewis et al., 1978 . . . . . . . .. . . 255
Key to Species . . . . . . . . . . . . . . . . . . . . . 255

59. Lutzoyia actydifera. . . . . . . . . . . . ... . . 255
60. Lutzomyia draisbaohi. . . . . . . . . . . . . . . .257

Subgenus Trichophorom'ia Barretto, 1962.. . . . . . . . . . 261
Key Lo Species. . . . . . . . . . . . . . . . ... .. . . . . 261

61. Lut :omria (T.) auaensi. . . . . . . . . . . . 262
62. Lutzormia (T.) cellulana n. sp. ... ....... 264
63. Lutzomjia (T.) houardi n. sp.. . . . . ... ..266
64. Lutzomyia (T.) reburra. . . . . . . . . . . . .. . 267
65. Lutzonyia (T.) saltzusa n. sp. . . . . . . . ... 269
66. Lutzomyia (T.) ubiquitalis. . . . . . . . . . . . 270

Subgenus Nyssormyia Barretto, 1962. . . . . . . . . . . . .. . 280
Keys to Species . . . . . . . . . . . . . . . . . . . . . . . 280

67. Lutzormyia (1.) antunesi . . . . . . . . . . . . .. 283
68. Lutzomyia (f.) flavia.uteo.i atat. . . . . . . . . . . 285
69. Lutomiyia (i'.) olmeca bicolor . . . . . . . . . . 287
70. Lutzomyia (11.) trapidoi . . . . . . . . . . . . .. 289
71. Lutzom'via (7.) Jmbratilis . . . . . . . . . . . .. 292
72. Lutzormyia (21.) yZephietor. . . . . . . . . . . ... 295
73. Lubzomyia (I.) yuilli . . . . . . . . . . . . . .. 297

Subgenus Psychodopygus Mangabeira, 1941 . . . . . . . . .. .311
Keys to Species . . . . . . . . . . . . . . . . . . . . . . . 312

74. Lutpomyia (P.) ber.zatei . . . . . . . . . . . . .. 317
75. Lutzonmia (P.) chagai. . . . . . . . . . . . . . . 317
76. Lutzotmyia (P.) fairtigi ...... . . . . . .. . 318
77. Lutzomyia (P.) bispinosa. . . . . . . . . . . . ... 319
78. Lutzomyia (P.) ayrozai . . . . . . . . . . . . . . 320
79. Lutzomyia (P.) carrerai cerrerai. . . . . . . . ... 323
80. Lutzomyia (P.) carrerai thula n. ssp. . . . . . . . 326
81. Lutzomyia (P.) daisi . . . . . . . . . . . . . . . 331
82. Lutzomjyia (P.) guyanensis . . . . . . . . . . . . 333
83. Lutzonmia (P.) hirsuta . . . . . . . . . . . . . . 335
84. Lutzomyia (P.) nocticola. . . . . . . . . . . . .. 338
85. L.utzornmia (P.) pacnenais . . . . . . . . . . . . .339
86. Lutsonria (P.) recrva. . . . . . . . . . . . . . . 341
87. Lubtzon ia (P.) sp. of Tres Esquinas . . . . . . . . 342

Vexator Group Theodor, 3965 .. .... ..... ............ . 366
Keys to Species . . . . . . . . . . . . . . . . ... . . . .. 367

88. Lutzomulia cirrita.. . . .. . .. . . . . . . . . . 369
89. Ltutzomyia hacrtma'.i i ..... . . . . . . . . . 370
90. Lutzomyia osonzoi . . . . . . . . . . . . . . . .. 371
91. LutzomU-ia sarguinariar . . . . . . . . . . . . . . 372



vi








Page

92. Luzo ia scorai . . . . . . . . . . . . . . . . 374
93. Lutsomyjia strictivi . . . . . . . . . . . . . 378
94. Lutsoryia sp. of Pichinde ............. 380

Cayennoensis Group Theodor, 1965 . . . . . . . . . . . . . . . 398
Keys to Species . . . . . . . . . . . . . . . . . . . . . . 398

95. Lutzomyia cayennensis. . . . . . . . . . . . . . 399
96. Lutzornyia micropyga . . . . . . . . . . . . . . . . 401
97. Lutzomyjia atroclavata . . . . . . . . . . . . . .. 405
98. Lutzomyia venezuelensis . . . . . . . . . . . . .. 407

OG0aldoi Group Theodor, 1965. . . . . . . . . . ... . ... 417
Keys to Species . . . . . . . . . . . . . . . . . . . . . . . 417

99. Lutzo ia pia . . . . . . . . . . . . . . . . . . . 418
100. Lutzormyia rorotaensis. .. . . . . . . . . . . .420
101. Lutzofyqia trinidadensis . . . . . . . . . . ... . 422

Pilosa GrcIuj 'J.L.hed, :196. .. ...... . . . . . . . . 432

102. Lutzomyia pilosa. . . . . . . . . . . . . . . . . . 432

Ungrouped Species . . . . . . . . . . . . . . . . . . . .. 437

103. Lutzoyia nordestina. . . . . . . . . . . . ... . . 437
10,. Lutzonjmia raneliana. . . . . . . . . . . . . . . . 439
105. Lutzomyia sp. of Anchicaya. . . . . . . . . . . ... 440

SYNOPSIS OF NEW TAXA, SYNONYMS, FEMALE DESCRIPTIONS, AND
NEW DISTRIBUTION RECORDS . . . . . . . . . . . . . . . .. . . . . 448

REFERENCES . . . . . . . . . . . . . . . . . . . . . . . . . . . . 451

BIOGRAPHICAL SKETCH. . . . . . . . . . . . . . . . . . . . . . . . 481






















vii













LIST OF FIGURES


Fiagureg

1. Eap of Colombia showing localities where phlebotomine
and sycoracine flies have been collected . . . . . . . .. 28

2. Syooax andicola. . . . . . . . . . . . . . . . . . . . .. . 43

3. Syc'. ra colombiensis. ... . . . . . . . . . . . . . . . 45

4. Syrax faichildi. . . . . . . . . . . . . . . . . . . ... 47

5, Sycorx trsispinosa. . . . . . . . . . . . . . . . . ... . 49

6. 57'qrZlya nign?,o alrIuZI. ..... ,... ........ 57

7. 1 rileya rotundipennis. . . . . . . . . . . . . . . . . . . 59

8. Brmrp,.tomyia avella-ri and Br; .ptonrmia beaupertuyi. . . . ... 69

9. Brumptortyjia galindoi. .. . . . . . . . . . . . . . . . . 71

10. Brumptomyia hamata . . . . . . . . . . . . . . . . . . .. . 73

13. Branptoryia Zeopoldoi and Brzcrptonria guimaraesi. . . . . . 75

12. Lutzornyia (P.) hirsuta nicaraguensis and Lutzonmia
tcwnsendi . . . . . . . . . . . . . . . . . . . . . . . . . 90

13. Lutzomyia (L.) bifoliata . . . . . . . ... . . . . . . . 103

14. Lutzsomyia (L.) ichyi . . . . . . . . . . . . . . . . . . . 105

15. Lutzorriyia (L.) Zongipapis. . . . . . . . . . . . . . . ... 107

16. Lizco:yzJa (L.) gomezi. ... . . . . . . . . . . . . . . .109

17. Lutscmyiaa (L.) r.rinkellei. . . . . . . . . . . . . . . ... 11

18. Lutbsom;yin migonei and Lutzom:ia sp. de Badnel .. . . . . . 120

19. Lut,'omyij a marajoensi . . . . . . . . . . . . . . . . . .. 122

20. Lut ia alkei . . . . . . . . . . . . . . . . . . . . . 124

21. a1 tsom ia saulensis . . . . . . . . . . . . . . . .. . . . 129



viii








Figur' Piage

22. Lutzor.ia serr na. . . . . . . . . . . . . . . . . . . . . 148

23. Lutzomftia andina, LutzomUyia disiuncta, arid LuitzoPrria
n'm eztov'.i . . . . . . . . . . . . . . . . . . . . . . .. . 150

24. Lutzsryia columbiana. .. . . . . . . . . . . . . . . . . . 152

25. Lutzomyia evansi . . . . . . . . . . . . . . . . . . . . . 154

26. Lutotrrria moraesi . . . . . . . . . . . ... .... .. . 156

27. Lutzomryia ovallesi . . . . . . . . . . . . . . . . .. . . . 158

28. Lutzornmia sauroida, Lutzormia quacitownsendi, LThtzormia
longiflocosa, Lutzomyia spinicrassa, and Lutzorgia odax. . . 160

29. Lutzorigia vespertilionic and Lutzomyia isovespr'tiiionis . 166

30. Lutztom ia (D.) rosbali. . . . . . . . . . . . . . . . . . . 170

31. Lutzornyza (Pi.) spinosa. . . . . . . . . . . . . . . . . . . 175

32. Lutzonmyia (Pr.) camposi . . . . . . . . . . . . . . . . ... 183

33. Tutbzomyia (Pr.) dysponeta. . . . . . . . . . . . . . . ... . 185

34. Lutzsomyia (Pr.) tricantha and Lutzoimyia baityi.. ...... . 189

35. zutzorm ia (V.) caprina and Lutzomyia (V.) furcata. .... . .197

36. Lutzomyia (V.) tubrculata . . . . . . . . . . . . . . . . . 199

37. Tutzoryia obonnenci . . . . . . . . . . . . . . . . . . . . 213

38. Lutzomyia dasymera . . . . . . . . . . . . . . . . .. . . . 215

39. Lutzornyia dendrophyla. . . . . . . . . . . . . . . . . ... . 217

40. Lutzomyia punctigeniculata . . . . . . . . . . . . . .... . 219

41. Lut o ia shannoni . . . . . . . . . . . . . . . . . . . . . 221

42. Lut'on la undulaba . . . . . . . . . . . . . . . . ... . . . 223

43. Lutornyia longipina . . . . . . . . . . . . . . . ... . . . 229

44. Lutzojia. triramula. . . . . . . . . . . . . . . . ... .. . . 231

45. Lutzom4ia aaao. ... . . . . . . . . . . . . . . . .. 246

46. Lutzorrjia barrcttoi majuscla. . . . . . . . . . . . . . . . 248




.x








'Figunre SPage

47. Lutzom .fa barrettoi majuscula, Lutzoryia barrettoi
barretboi, and Lutzomyia texcna . . . . . . . . .. . . 250

48. Lutzomqjia carpenteri . . . . . . . . . . . . . . . . . .. . 252

49. Latzomyio runoides. . . . . . . . . . . . . . . . . . . . .254

50. Lutzomyia aclydifera. . . . . . . . . . . . . . . . . ... . 260

51. ;utSzomyia dreisbachi, Lvtzoryia (T.) auraensis, and
Lutzonqmyia ubiquitalis . . . . . . . . . . . . . . . . . . . 273

52. Lutzormnia (T.) cellulana. . . . . . . . . . . . . . . ... . 275

53. Lutzomnia (T.) howardi and Lutzoimiia (T.) saltuosa. ... . 277

54. Lutzomnia (T.) reburra. . . . . . . . . . . . . . . . . . . 279

55. Lutzomyia (N.) antunesi and Lutsomyia (N.) flaviscutellactta. 300

56. Lfutzomyia (N.) olmeca bicolor . . . . . . . . . . . .... . 302

57. Lu.toryia (N.) trapidoi . . . . . . . . . . . . . . . . . . 304

58. Lutzonyia (N.) wnbratilis . . . . . . . . . . . . .... . 306

59. Lutzomyjia (N.) ylephiletor. . . . . . . . . . . . . . . ... 308

60. Lutzomyia (N.) yuilli . . . . . . . . . . . . . . . . . . . 310

61. Lutzormyia (P.) bernatei, Lutzomryia (P.) chagasi, and
Lutzonvjia sp. of Tres Esquinas. . . . . . . . . . . . . . 345

62. Lutzornyia (P.) fairtigi . . . . . . . . . . . . . . . . . . 347

63. Lutzomyia (P.) bispinosa. . . . . . . . . . . . . . . .. . 349

64. Lutzom pnia (P.) ayrozai . . . . . . . . . . . . . . . . . . 351

65. Lutzomrrica (P.) carrerai carrerai. .... . . ........ . 353

66. Lutzomyia (P.) carrerai thula . . . . . . . . . . . .... . 355

67. Luzonmyia (P.) davisi and LutzoVyia (P.) hirsuta hiru-sta. . 357

68. Tutsnomyra (P.) guyan.nsis . ................ . 359

69. Lutzonyia (P.) nocticola. . . . . . . . . . . . . . ... . 361

70. Lutsorjia (P.) panamensis. . . . . . . . . . . . . . . . 363

71. jutzomnia (P.) recurva. . . . . . . . . . . . . . . . . ... 365




A







FPgure Pago

72. Lut-om'ia cirrita. .. . . . . . . . . . . . . . . . . . . . 385

73. Lutz'oyia h manni. . . . . . . . . . . . . . . . . . . 387

74. Lutz yia o ornoi. . . . . . . . . . . . . . . . . . . 389

75. Lutzomyia sang iinaria. . . . . . . . . . . . . . . . . . . 391

76. Lutzonmyia strictivilla . . . . . . . . . . . . . . . ... . 393

77. Lutzomjia scorzai . ...... . . . . . . . . . . . . . . .395

78. Lutzomyia sp. of Pichinde.... . . . . . . . . . . . . . . 397

79. Lutszoria cayennensis. . . . . . . . . . . . . . . .. . . 410

80. Lutzomyia micropyga . . . . . . . . . . . . . . . . . . . .412

81. Lutzomyia atroclavata. . . . . . . . . . . . . . . . . ... . 414

82. Lutzo yia venezueLensis.. . . . . . . . . . . . . . . . . ... 416

83. Lutzo.rmia pia. . . . . . . . . . . . . . . . ... .. . . . 427

84. Lutzomyia rorotaenis.. . . . . . . . . . . . . . . . . . . 429

85. Lutzomyia trinidadensis. . . . . . . . . . . . . . . . .. . 431

86. Lut;ormnyia pilosa . . . . . . . . . . . .... . . . . . . . . 436

87. Lutzomiia nordestina . . . . . . . . . . . . . . . . . . . . 443

88. Lutaomyjia rangeliana . . . . . . . . . . . . . . . . . . . . 445

89. Lutzonmyia sp. of Anchicaya . . . . . . . . . . . . . . . . . 447



















r� i











Abstract of Dissertation Presented to the Graduate Council
of the University of Florida in Partial Fulfillment of the Requirements
for the Degree of Doctor of Philosophy


A BIOSYSTEMATIC REVIEW OF ITE BLOODSUCKING PSYC!{GDID FLIES OF
COLOMBIA (DIPTERA; FPHLFBOTOMINAIE AND SYCORACINAE)

By

David Grier Young

December, 1977

Chairman; Graham B. Fairchild
Major Department: Entomology and Nematology

The psychodid subfamily Sycoracinae, previously unknown in Colombia,

is represented by four new Sycorax species described and illustrated in

this ceview.

Phelbotominae, a much larger subfamily containing vectors of human

pathogens, is represented in the Republic by 101 species and subspecies --

a n ,umber equivalent to one-third of the described New World taxa. Ten of

these ir the genus Lutzormia are described as new and are given formal or

inform.i names. Tht previously unknown females of Lutzomyia pilosa, L.

(P.) ht�suta niccaraguensis ind L. bifoliota are described. Nineteen

phlebotomeine species are reported in Colombia for the first time. Other

new distributional records include those from Ecuador (22 spp.), Panama

(3 spp.), Honduras (2 spp.), Costa Rica, Cuba, Guatemala, and Paraguay

(1 sp. eanh). In order to clarify the status of some forms, an attempt

wa:s imade to study specimens from as many Colombian and non-Colombian

local tieu as possible. To this end, over 20,000 adults in both sub-

fa'itlies were examined from 1967 to the present.





x:ii








An extensive bibliography and synonomy is given for each previously

described taxon. Actual and/or probable misidentifications in the

literature are discussed. Identification keys, previously unavailable

for the Colombian fauna, are provided along with numerous illustrations,

most of which are original.

Although more information is needed, it is suggested that recent

speciation in the neotropical phlebotominae was due to climate changes

in the Pleistocene (refuge theory).














































xiiii














INTRODUCTION



Prior to the last decade, we knew very little about the phlebotomine

sand fly-; fa[un of Colombia. Osorno et al. (1967) in reviewing previous

studies by others and adding records of their own, reported less than 25

species in the Republic. From 1967 to 1972, the late Dr. E. Osorno-Mesa

andj his colleagues at Instituto National de Salud, Bugota added more

records, bringing the total to 77 species and subspecies (Osorno et al.,

1972a).

In this review 101 species and subspecies of Phlehotominae are re-

ported from Colombia. Thisrepresentsabout one-third of the described New

World species but I estimate that at least 40 additional species will be

discovered in the Republic, especially in the little-collected south-

eastern region.

The small subfamily Sycoracinae, whose members resemble those of

Phlebotominae in being able to take vertebrate blood, is treated here

for this reason.

In order to clarify the status of some taxa and to study intraspeci-

fic variation and distributional patterns, an effort was made to examine

specimens from as many Colombian and non-Colombian localities as possible.

To this end, T am very grateful to the following persons for providing

specimomns, inEormation or both: Dr. Jorge R. Arias, Instituto Nacional

de Peocuisas do Amazonia (INPA), Manaus; Drs. Stephen C. Ayala and Pablo


*� prefer to separate "sand" from "fly" in accordance with the suggestion
of borrer, Delong, and Triplehorn (1976).




.-.2--



Barretto, Universidad del Valle, Cali; Drs. Richard D. Uard and Habib

Fraiha, Instituto Evandro Chagas, Belem; Dr. David J. Lewis, British

Museum (Nat. Hist.); Dr. Cornelis J. Marinkelle, formerly Universidad

de Los Andes, Bogota; Dr. Alberto Morales-Alarcon and the late Dr. Ernesto

Osorno-Mesa, Instituto Nacional de Salud, Bogota; Prof. Almilcar Vianna

Martins, Universidade Federal de Minas Gerais; Dr. Nelson Papavero, Museu

de Zoologia, Universidade de Sao Paulo; Drs. Charles H. Porter and Thomas

M. Yuill, University of Wisconsin; Dr. Alan Stone, formerly Systematic

Entomology Laboratory (ARS); Mr. Richard C. Wilkerson, University of

Florida. Dr. Thomas H.G. Aitken, Yale University School of Medicine;

Dr. Elisha S. Tikasingh, Caribbean Epidemiology Center (CAREC), Trinidad;

Dr. Harold Trapido, Louisiana State University Medical Cencer; Dr. Jorge

E. Velasco, La Paz, Bolivia; Dr. Derek A. Duckhouse, University of

Adelaide, South Australia; Maj. John F. Reinert, U.S. Army Medical

Research and Development Command, Washington, D.C.

Others who have supported this study and to whom I am indebted

include members of the Atlantic-Pacific Interoceanic Canal Study Commis-

sion, Canal Zone from 1967-1968. Col. Bruce F. Eldridge of this organi-

zation was especially helpful in developing field programs and in

providing data on Choco collections.

Financial support from U.S. Army Medical Research and Development

Command under contract no. DADA 17-72-C-2139 is gratefully acknowledged.

Such support was provided also at various times by Drs. William G. Eden

and Hugh Popenoe, University of Florida.

Members of my graduate school committee at the University of Florida

have been helpful in many ways, too numerous to mention here. For their

help, I thank Drs. Franklin S. Blanton, Jerry F. Butler, Graham B.





-3-


Failchild, Dale H. Habeck, and Stephen G. Zam. Miss Stephanie Haney ably

slide mounted thousands of specimens from 1972 to 1977. For typing the

ranuscript, I wish to thank Mrs. Adele Koehler.

Dr. G.B. Fairchild deserves special recognition for his constant

encouragement, stimulating discussions and expert assistance throughout

the study. I ai deeply indebted to him. Particular appreciation is

expressed also to my wife Molly who was most understanding during the

preparation of this review.

Holotypas and allotypes of new taxa are to be deposited in the U.S.

National Museum (Nat. Hist.). Paratypes, when available, will be held by

the following institutions: Instituto Nacional de Salud, Bogota; Florida

State Collection of Arthropods, Gainesville; and British Museum (Nat.

Hist.), London. The remaining 20,000+ specimens, upon which most of this

review is based, are stored in the author's personal collection. A

synoptic collection will be given to the Instituto Nacional de Salud,

Bogota.














CLASSIFICATION AND TAXONOMIC TREATMENT



Of the 6 recognized subfamilies of Psychodidae (Duckhouse, 1972,

1973), all but one, Horaiellinae from the Oriental Region, are represented

in continental Colombia. The species of lorazieZla, Tr'icho�mJi, and

Sycorax and its allies are united by some in the subfamily Trichomyiinae

but this appears to be artificial and does not "express the wide discon-

tinuities and degrees of difference from other Psychodidae, especially in

Liet iis;ffiaurL, IN" (c ,12

The classification of Phlebotominae is interpreted differently

according to author(s). In a recent paper we reviewed most classifica-

tory schemes and proposed one which is used here for the New World taxa

(Lewis et al., 1978). It is essentially a modification of Theodor's 1965

classification. Three American genera are recognized -- Lutzonrjia Franga

(ra. 290 spp.), Bruwptomyia Franca and Parrot (ca. 23 spp.), Wlariteya

Hertig (5 spp.). The genus Hertigia Fairchild is considered to be a

junior synonym of WariZeya by Lewis et al. (1978).

The majority of the Lutzomyia spp. were described originally in the

genus Phleboto;Mr.s (= Flebotomus) Rondani. Some authors continue to place

them in this genus, ignoring the evidence to the contrary provided by

Theodor (1948, 1965) and Christensen et al. (1971).

The large genus Lutzomyia is divided into numerous suhgenera, species

groups, and series based upon adult morphology (Lewis at al., 1978). Tte

diversity of groups within the genus is attributed to "radiation or even

'explosive radiation' of some immigrating ancestral stocks in a newly




-4-





~5-


colonized area having a wealth of ecological niches" (Theodor, 1965).

This implies that these ancestral stocks originated in the Old World --

a hypothesis which may never be proven. It would appear that recent

speciation in some groups (e.g. subgenus Tricrhophoronjyia) may have been

due to climatic changes in the past, especially during the Pleistocene,

which served to isolate conspecific populations in moist refugia during

dry periods (Haffer, 1974).

Other remarks on the classification of New World phlebotomines are

given by Lewis et al. (1978) and in this review under each supraspecific

taxon.

An extensive bibliography and synnonomy is given for most taxa treated

here. I usually omitted references which contain species lists but some

of them including those by Fairchild (1955), Barretto (1955a, 1962),

Theodor (1965), and Martins & Morales (1972) are very important because

of discussions related to classification and distribution. For each

species, I cite the original description as well as most others published

since Barretto's 1947 catalog. In cases where a species was reviewed

subsequent to 1946 (e.g. Forattini, 1973), I refer to that publication

for full references.

Abbreviations of words used in the text which may not be familiar

to some readers are given below:

Insti utions

USNM-- United States National Museum (Nat. Hist.), Washington, D.C.

INPES -- Instituto Nacional para Programas Especiales de Salud, Bogota

INPA -- nstituto Nactional de Pesquisas de Amazonia, Manaus

UV -- Universidad del Valle, Cali





-6--



GML -- Crgas Memorial Laboratory, Panama

UF -- University of Florida


Collectors

C.H.P. -- Charles H. Porter

C.J.M. -- Cornelius J. Marinkelle

R.C.W. -- Richard C. Wilkerson

D.G.Y. -- David C. Young


Other Abbreviations

ca.-- about

cf.-- compared to

classif. -- classification

descr. -- description

dist. - distribution

morphol. -- morphology

pop. -- population



Key to the Subfamilies of Psychodidae of the New World


1. Wing venation with 4 branched radius, R2 fused into a single

vein (Fig. 2B). . . . . . . . . . . . . . . . . . . . . .... . 2


Wing venation with 5 branched radius (Fig. 12G) . . . . . . . . . . 3


2. Wing venation with short cubitus (cu). Mandibles

present . . . . . . . . . . . . . . . .... . . .Sycoracinae (Fig. 2)


Wing venation with long cubitus. Mandibles absent. . . .Trichomyiinae





-7--



3. Palpus of 4 segments, rarely 3. Eyes usually with eye bridge.

Antennal flagellomeres nodiform or barrel shaped.. . . . Psychodinae


Palpus of 5 segments. Eyes without eye bridge. Antennal flagel-

lomwres subcylindrical ................... .. .. 4


4. Females with 1 spermatheca. Mandibles absent. Male aedeagus

entire . . . . . . . . . . . . . . . . . . . . . . . .. Bruchomyiinae


Females with 2 spermathecae. Mandibles present. Males with bifid

aedeagus. . . . . . . . . . . . . . . . . . . Phlebotominae (Fig. 12F)














GENERAL MORPHOLOGY AND TERMINOLOGY



This brief discussion focuses on the structures commonly used in

taxonoitic studies of phlebotomine sand flies. General accounts of adult

anatomy include those by Fairchild and Hertig (1947a), Kirk & Lewis

(1951), Abonnenc & iinter (1965), Davis (1967), Abonnene (1972), Lewis

(1973), and Forattini (1973). The terminology varies according to author;

that used here applies also to the sycoracine flies except where otherwise

staLed.

Although little studied, the immature stages are probably no less

important in systematics; they are not considered here owing to lack

of material. For information on the anatomy of these stages see Barretto

(1941), Hanson (1968), Abonnenc (1972), Carnheiro & Sherlock (1964), Ward

(1977), and Zimmerman et al. (1977). Descriptions of immatures of those

species occurring in Colombia are cited in the appropriate species

bibliography.

The sand fly head and its appendages (Fig. 12A) are useful in

associating conspecific males and females and for grouping taxa below the

genus level. Head height, measured from the vertex to the tip of the

clypeus, in relation to its greatest width is of value in distinguishing

some of the Lubtomyia species.

As a probable adaptive feature related to host finding, the size of

the compound eyes varies according to species, especially those of

Lut:o';nrmia, but may be characteristic of subgenera and equivalent species




-8--








groups. Contrary to Davis (1967), ocelli are lacking in the Phlebuoomrines

and Sycoracines. Illustrations are by far the best way to indicate eye

size, the relative terms such as "snalL" or "large" being understood

easily with associated figures. The interocular distance or eye separa-

tion is the narrowest distance between the eyes and is measured directly

or may be expressed by the number of facet diameters (or fractions there-

of) needed to connect the eyes.

The interocular suture (= post frontal suture of mosquitoes,

Christophers, 1960) is complete in adults of Brurmp-tor.yia (Fig. 8B),

Warilea (Fig. 6A), and Syoorax (Fig. 2A) but is incomplete in those of

Lust;ompyia (Fig. 12A).

In Phlebotominae, the maxillary palpus (Fig. 12A) consists of 5

segments, the basal one (palp 1) always the smallest and least important

in taxonomy (Lewis, 1973). Being partially fused with the second, it is

difficult to accurately measure. The relative lengths of the segments

are expressed often by a palp formula or by ratios with palp 1 being unity

or 10 (Kirk & Lewis, 1951). A palp formula of 1-4-2-3-5 indicates that

palp 1 is the shortest, palp 4 the next shortest, etc. When two segments

are equal in length they are enclosed in brackets, e.g. 1-(4-2)-3-5. The

sycoracines have four, instead of five, palpal segments (Fig. 2E).

Species in both subfamilies have palpal sensilla (= Newstead's

scales) which are small clubbed sensory organs attached to one or more

segments in the phlebotomines and to palp 2 in the Colombian Sycorax spp.

The cylindrical antennal flagellomercs, unlike true segments, are

not indepedently musculated (Imms, 1938). For this reason I join the

increasing unmber of students who use the term "flagellomere" in the

place of "antennal segment." The antenna consists of a scape, pedicel,






-10-


and 1i flageilomtees, each of which is designated by a Roman numeral

(Fig. 12A). in SyNo0ax the terminal flagellciere is markedly reduced in

size (Fig. 2D). The length of flagellomere 1 (= antennal segment 3), the

most basal flagellomere, in relation to other flagellomeres, head height,

or labrum length is useful as a diagnostic feature at the species or

subspecies level.

The paired antcnnal ascoids (Fig. 12B), often difficult to observe,

are important owing to interspecific variation in their distribution,

shape, and length. Some specimens may have but a single ascoid on a

particular flagellcmere.

The mouthparts of New World phlebotomines in relation to feeding

habits and btaonomy were studied in detail by Lewis (1975a). Following

him, I use the term labrum for the labrum-epipharynx of various authors.

Its length is measured from the apex of clypeus to tip of proboscis.

The cibarium (= buccal cavity) lies within the clypeus and is armed

with a variab!e number of teeth in the Lutzormjia and Brurmptomyia females.

Those in the latter genus are characteristically arranged in 4 longi-

tudinal rows (Fig. 10HM). Cibarial teeth are absent in both sexes of

WariZeya and Sycorax and are poorly developed or absent in the Lutzonyia

and BrLumptomyia males.

Ih en viewed ventrally as in Fig. 12E, the horizontal teeth (= hind

teeth) of most Lutzomyia females indeed look like teeth, their tips

pointed, or not, depending on species. The vertical teeth (= fore teeth)

usuall appnar as dark dots distad uf the horizontal teeth at the same

angle of view (Fig. 12E). Lateral teeth may occur on both sides of

cibarium near the horizontal teeth (Fig. 30M). The number, position,

shape, and size of these cibarial teeth are very important in species

diagn sis and classification.





-11-


Two other features of the cibarium are noted in the descriptions

and keys. The r.chirinous arch (Fig. 12E), when complete, crosses the

ventral wall of the cibarium. Its development varies according to species

and may be complete or not. The pigment patch on the dorsum of cibarium

(Fig. 12E) varies in size, shape, and degree of infuscation (= pigmenta-

tion) in the Lulzomyia spp.

lhe pharynx (Fig. 12C), composed of 3 chitinous plates, is attached

to the cibarium. Many species of Old World phlebotomines (genus

Ser.gentom ria) have spines on the posterior part of the pharynx but they

are relatively rare in the Lutzomyia spp., occurring mostly in some

species in the cayenennts and os~aldoi groups. The majority of Lutzsoaia

females have transverse, unarmed ridges on the wider, posterior area of

the pharynx. The length of the pharynx is given in descriptions although

it is flexible and difficult to measure with accuracy (Lewis, 1967a).

I adopt the terminology of Davis (1957) as applied to the thorax.

The mesonotum. should correctly include the mesoscutum, mesoscutellum, and

postnotum and should not be restricted to the mesoscutum alone (Saether,

197!).

Tie degree and distribution of pigmentation of the thorax and other

body regions are important in associating sexes and for distinguishing

species of Lutzomyia. Coloration has not been given much importance in

systenacic works but I find it generally dependable as a taxonomic

character as applied to the New World phlebotomines.

PFeural setae on the anepisterum (upper episternal setae) and

katepisternum (lower episternal setae) are present in species of Lutzo3,yia,

BL'2i ptcmyia, and Stfcorax. The Warileya spp. and the majority of Old World

phlebotomines lack such setae (Abonnenc & Leger, 1975). Additional





-12-


pleural setae, not mentioned in other studies to my knowledge, occur

behiad the metathoracic spiracle in all species :f Phlebotominae and

Sycoraciliae examined, those in the latter subfamily being 4 in number

anm relatively stout EFig. 2H). The 4 postspiracular setae in the

iphlebotomine species are reduced in si2:e, barely visible (Fig. 12D).

The ving length is measured from the basal costal node (= rudiment

of tegula; Snodgrass, 1935) to its apex; its width at widest part (Fig.

12G). Certain wing vein sections (Fig. 12G) are given useful, easily

remembered names. Alpha (a) is the length of R2 from its junction with

R3 t t the costa. Beta (5) is the length or R from the junction of R to

the junction of R2 + R3. Garmna (y) is the section of R from the r - m

crossvein to the junction of R5. Delta (6) is that part of R1 distad of

the junction of R and R.. It is negative when R ends before this

junction.

The legs are measured as in Fig. 12H. A few phlebotoamine spp. have

a row of short spines on the hind femur (Fig. 31F). The length of the

femur in relation to that of the tibia and/or basitarsus is sometimes

used for distinguishing species of Lutzomyiza.

The setation of the abdominal tergites 2 and 6 is useful in the

classification of Old World phlebotomines but has been little studied

for the New World species. lewis (1975b), however, discussed and figured

the socket patterns for some species, noting that the patterns were of 5

types. Setae on the sidea of tergite 8 may be present or not in New

World females of Phlebotominae and Sycoracinae.

The nature of the abdominal sternites, especially sternites 2 and 3,

is useful in associating male and female sand flies (IHertig & Fairchild,

1.950) but the character state is variable among individuals of conspecific

populatiions (forattini, 1954).





13


The structures of the bilaterally symmetrical male genitalia,

labealld i- FiF. 2 a-ed 12, are extremely important ir systematics. After

eclosion, the genitalia rotates 1800 in the phlebotomine males but not in

those of Sycrioaz. Ihe style, attached to the apex of the coxite, bears

1 ro 6 major spines* and S to many additional sirall setae. The presence

of numerous strong spines is presumably a plesiomorphic feature (Theodor,

1965).

The coxite is the dorsal basal appendage in the phiebotomine males

and, of course, is the ventral basal one in the Sycorax males. It may or

may not bave a setal tuft or other nondeciduous setae on its inner sur-

Eace. These setae are those which remain on the coxite after maceration

and which usually differ in size or shape from other setae on the

appendage.

The fused, paired aedeagi, each of which is called the aedeagus is

subtriangular and well sclerotized in most phlebotomtne males. Some

species (e.g. L. walkeri) have aedeagi with dorsal projections. The

aedeagus of the Colombian Sycorax males is complex as in Fig. 2F. There

is a basal, laterally flattened sperm pump within the abdomen, distal

paired genital filaments and other structures mentioned on p. 31. The

genital pump of the phlebotomine males is probably homologous with the

sperm pump of the Sycorax spp. Leading from it are 2 genital filaments

which pass through each aedeagus nPd which vary in length, width, and

sclerotization according to species. The apices are simple, modified,

inflated or not.



*It is difficult to define major (= strong) spine, as it is a relative
tutrm referring here to size. One author's interpretation may differ from
that of another. In doubtful cases, the illustrations should be studied
to understand the meaning of statements in the text.





-14-


The phlebotomine parameres U r: between the coxites and ventral

lateral lobes when viewed laterally. Their shape and setation are im-

portant in differentiating species. Simple parameres, i.e., those without

arms or processes, are shown in Fig. 16F and 18A. Complex parameres are

illustrated in Fig. A41. The cercopods of the Sycorax males appear to

correspond to the parameres of Phlebotominae and are very complex.

The ventral, nonsegmented lateral lobes are more or less uniform in

the phlebotomine males, although their length, width, and setation may

offer good characters in some species. Corresponding structures are

wanting in the sycoracines.

The shape of the setose cerci may help associate conspecific sexes

but in general these structures are of limited use in species diagnosis.

The size and structure of the internal paired spermathecae and their

ducrs (Fig. 12F) are particularily important in the classification and

identification of female sand flies. An idea of the variety of sperna-

thecae can be gained by scanning the illustrations in this review. A well

sclerotized, "Y" shaped genital fork or furca in the phlebotomines is

associated with the spermathecae.














METHODS AND IMTERIALS



Field Collections


For sampling phlebotomine and sycoracine populations, several methods

should be used to determine species diversity and relative abundance. In

Choco Department and elsewhere to a lesser extent, we used the following

methods to capture these flies.

Batte 'r powered light traps (Sudia and Chamberlain, 1962) attract

many psychodids, the numbers varying according to weather conditions,

location, species composition, etc. We secured these traps to tree branches

2 meters above ground level but did not use carbon dioxide as an adjunct

attraction. At Curiche, where at least 1 trap per week was operated

throughout the night (1800 hrs.-0700 hrs.), we collected a total of 23

phlebotomine spp. (378.(l, 78999) from April to Dec., 1967 (39 trap nights).

We recorded 36 sand fly species from Curiche based on all collection

techi iquies.

Shannon traps (Shannon, 1939) made from muslin bedsheets do not trap

insects per se buc are collecting devices similar to those used by

lepidopterists in "sheeting" for moths at night. A gasoline lantern pro-

vides a light sourcc enabling collectors to readily aspirate the psychodids

which land oa the illuminated cloth. They are attracted to the light, to

the collectors, or to a combination of both. The species composition of

light and Shannon trap captures is generally similar.





-15-




-Ib-


a.iais'e traps (Townes, 196?) and flight traps (similar to the design

ol Gr(ressitt and Gressitt, 1962) were placed in forest clearings, across

trza:ls .-r othEt-: flyways at grountd level and in the forest canopy, Insects

which land on the trap baffles eventually die after making their way

upwards to the killing jar(s) charged with potassium cyanide. Ordinarily,

thene L tzaps do not capture large numbers of psychodids but they are

effective in sampling the species composition of a given area if operated

over extended periods of time. For example, in the Curiche Forest at

groLind level, one Malaise trap captured 24 Phlebotomine species (316'9,

4829 ) from 3 April 1967, to 30 Nov. 1967 (109 trap days ).

Whenever possible, we searched diurnal resting sites for sand flies.

1These included tree trunks, especially dark crevices between buttresses,

animal burrows, and litter on the forest floor. Tree trunks are favored

resting sites for several species, specimens of which are easily captured

with a simple aspirator.

During the survey in Choco Dept., especially at Curiche and Teresita,

we collected large numbers of phlebotomines on human bait. The majority

of collections were made by 2 men sitting on the forest floor between

1i30 and 2100 hours. Phlebotomines were captured with aspirators as they

attempted to feed, the results computed on a man-hour basis, i.e., the

total number captured in one hour divided by the number of collectors.

Specimens were preserved dry in cardboard pill boxes rather than in

alcohol to prevent hardening of muscle tissue.








*Oine trap day equals 24 hours.





J








Laboratory Methods


For routine. identification of phlebotomines and sycotacines, un-

dissected specimens are first macerated (i.e., cleared) in 10-20% NaOH.

The loss of pigmentation can be reduced by heating this solution (con-

taining the flies) to the boiling point rather than nacerating them at

room temperature for 12 hours or more. The flies should be left in the

hot NaOH for 5 minutes, then placed directly into a drop of 80-95% liquid

phenol (C6H OH) in a depression microslide. Within 5 minutes, the

important internal structures such as the spermathecae and ducts are

clearly visible and should be drawn if necessary. Once identified, the

flies can be discarded, preserved in vials of 70% alcohol, or processed

further for slide mounting. I follow the procedure of Fairchild aud

Hertig (1948c) except that Canada balsam or Euparal is substituted for

copal.

Other slide-mounting procedures are aiscussed uy Osorno et al. (1966),

Quate & Steffai (1966), and Lewis (1973) and others. The choice of which

to use is a matter of personal preference but in all cases the best pre-

parations are made from freshly killed flies.

Sometimes it is necessary to remount specimens because of undesirable

position and/or shrinkage. For those embedded in Canada balsam or other

xylene-soluble media, it is advisable to submerge the entire slide in

liquid phen<-! in a suitably closed container such as a petri dish. After

3 to 7 days, the specimens become soft and are easily handled. Xylene as

a solvent should be avoided as the specimens tend to become brittle and

easily damaged.





-18-


Specimens were drawn withl the aid of a camera lucida and Bausch and

Lomb microprojector. I rmeasured specimens with a calibrated ocular

SicroTneter, All such measurements are given in millimeters throughout

the text and figures.














DISTRIBUTION, COLLECTING LOCALITIES, AND
CHARACTERISTICS OF THE FAUNA



The distribution of phlebotomines and sycoracines can be understood

by present ecological factors, both biotic and physical, and by knowledge

of past changes in climate, vegetation, and geology. Haffer (1967, 1974)

discussed the probable speciation and distribution of some neotropical

birds in relation to past changes, especially those occurring in the

Pleistocene and post Pleistocene periods. During times of droughc, popu-

lations of forest birds and other organisms were restricted to isolated

forest refugia which acted as core areas of speciation. As conditions

became wetter, these forests enlarged in area, often merging with others

to form zones of secondary contact. The presumed forest refugia in

Colombia and elsewhere were discussed and mapped by Haffer (1967, 1974)

end Brown (1975), the latter author studying speciation in forest butter-

flies in relation to these presumed refugia.

Although more data are needed, the refuge theory offers a plausible

explanation for understanding recent speciation in the Phlebotominae and

Sycoracinae. The majority of New World species inhabit forests, especially

those in the lowland tropics which receive 2000+ mm of rain per year.

For discussion purposes, it is convenient to divide Colombia into

natural regions. D'Allesandro & Barreto (1971) delineate 7 such areas,

their large "Oriental Region" consisting of both Amazonian forest and

eastern natural savannah (Ilanos). Within each region there are one or

more life zones, each defined by a combination of biotemperature, annual




-19-





-20-


it.recipi.tation, humidity, and vegetation (Holdridge, 1967). The life

:.-onr. in Colom::ia were studied by Espina. and Montenegro (1963). Their

'bcsque h'uriedo" translates to "tropical moist forest" (2000-4000 mmn of

rain per year) and "bosque muy humedo" corresponds to "tropical wet

forest" (4000-8000 T m of rain per year). The "bosque pluvial" (= "tropipal

rain forest"), the wettest lowland life zone, receives over 8000 ani of

rair. ?per year.

Figure 1 depicts where phlebotomines and sycoracines have been col-

lected in Colombia. Osorno et al. (1972a) provide specific data for most

localities; others from which collections were made by me or my colleagues

are discussed below in relation to faunal regions (D'Allesandro & Barreto,

1971) and distribution centers (Haffer, 1974).



Pacific Coast Region


Haffer (1967) attributes the "high concentration of endemic species

in the tropical lowland forests of western Colombia and Central America"

to a gradual accumulation of isolates in the trans-Andean forest region.*

The dominant refugium on the Colombian Pacific coast is the Choco refugium

extending from Lago Calima (Valle) to north of Quibdo (Choco) and across

the Rio Atrato and San Juan river systems (Brown, 1975).

An analysis of the phlebotomine species and subspecies inhabiting the

Pacific coast region indicates the following.

The vast majority of the 41 known taxa, probably originated in

forests east cf the Andes (cI-s-Andean region). Most of these reached the


'*The tr2ans-Andean region comprises "Middle America and the narrow Pacific
lowlands along the western base of the Andes" (Haffer, 1967). Cis-Andean
refers lo lowland forests east of the Andes.





-21-


Pacific coast forests by advancing around, not across, the northern Andes.

Thesu species which now occur on both sides of the mountains include:

Lz tzorqi; ayrozai, qL. gujanc-nsi-, L. bipiwnsa, L. comezi, L. ncrdestina,

L. senrvana, L. pilosa, L. spinosa, L. trinidadensis, L. shannoni, L.

tvi, h cuiata, L. triramulca, and others.

Nine of the 41 tpecies and subspecies do not occur in other regions

of the Republic. Of these, only Lutzormria sp. of Anchicaya is precinc-

tive, the others occurring also in the trans-Andean regions of Northwestern

Ecuador and/or Central America. These species include: Brurptomyia

hajrabaa, WariZeya nigrosacculus, L. reb rrac, L. recurva, L. isovesperti-

Zionis, L. yZephiletor, L. sanguinauria, and possibly L. odax. Two forms

wiich probably evolved in Pleistocene forest refugia within the trans-

Andean region include L. carrerai thula n.spp. and L. barrettoi majuscula

n.spp.,both of which now occur in trans-Andean regions of Ecuador, Colombia,

and Central America and east to the northern foothills of the Central

Cordillera of the Colombian Andes.

There is a slight possibility that some phlebotomines were

able to cross the Andes in southern Ecuador and northern Peru in times

past when wet forests occupied the now dry valleys. Some Amazonian birds

apparently colonized Pacific coast forests in such a manner (Haffer, 1967).

Phlebotomines, however, are rather weak fliers and it would have been

necessary for lowland forms to survive at elevations above 2000 meters

above sea level. The one possible example of successful colonization may

be that of L. rehurra -- a species whose closest allies occur entirely in

the nis--Andean region, especially in Amazonian forests.

A few species such as L. vespertilionis and its sister species,

L. iso'ecOertilionis probably invaded the Pacific coast region of Colombia




-22-



from Central Arerica, the former species also occurring east of the Andes,

at least as far south as Bolivar Department.



.Lepartment of Chocc


Three collecting sites within this Department were described by

Eldridge & Fairchild (1973) and Eldridge et a]. (1973). Maps accompany

their descriptions.

C'uriiche. On narrow coastal plain between western slopes of Serrania

del Baudo and Humboldt Bay on the Pacific coast; "tropical wet forest;"

elev..tion* less than 10 m. Sand flies were collected from April 1967, to

Dec. 1967, in discurbed and undisturbed evergreent forests and near a

mangrove swamp.

Alto C-urichf. About 3 km inland from Curiche on western ridge of

Sercania del Baudo: "tropical wet forest;" elevation 302 m. The sand

fly fauna is similar to that of nearby Curiche.

Teresita. At eastern foothills of Serrania de Baudo, near Rio

Truando; transitional life zone, "tropical moist/tropical wet forest";

elevation 35 m. Collections made in semi-disturbed forests on flat or

gently rolling terrain from March-Dec. 1967.



Department of Valle


Anehicaya Dcan. About 35 km inland from Pacific Ocean, on the Rio

Anchicaya; "tropical wet forest;" elevation ca. 560 m. This site is

similar to that near the Rio Anori (Antioquia Dept.) in terrain, elevation,




*Elevation is expressed here and elsewhere in meters above sea level.




U





-23-



and cl:r-:ate. Sycoracines were recorded fror. both localities and from one

other in C;auc Department.

25 ,.m E of Bu.enaventura. "Tropical rain forest" arcording to Espinal

(198) ; elevation ca. 50 m. Phlebotomines were captured in a patch of

disturbed f.rest surrounded by cultivated fields.



Atlantic Coast Region


This area encompasses not only the Carribean lowlands but also the

northern foothills of the western and central Cordilleras of the Andes.

The presumed Nechi refugium is located iere, a center "heavily influenced

by the Choco u-eter to the west" (Haffer, 1967). The present life zones

are nore diverse here than in the Pacific coast forests and except for the

area near the Rio Anori, the phlebotomine fauna has been little studied.

Intensive collections of phlebotomines made by Dr. C.H. Porter and

his colleagues near the Rio Anori (1970-1971) yielded 37 species and sub-

species. Thirty of these also occur in Pacific coast forests, a fact not

surprising based on the distribution of other organisms (HaEfer, 1967,

1974). Lutwzoiyia strictivilZa may be endemic. Lutzormyia yuilli and

L. nocticola are cis-Andean taxa. The former reaches its northernmost

li-mits in northarn Colombia; whereas L. nocticola invaded extreme western

Panaiaa but populations between there and northern Colombia have since

disappeared.



.Deoartmeit of Antioruia


Rio Anori (70 19'N; 750 04'W), about 24 km SW of Zaragoza; "tropical

wer forest;" elevations from 500-700 m. Located in the foothills of the





-24-


central Cordillera of the Andes, this site is mostly well forested with

steep. well drained slopes. A detailed description is given by Porter

& Defoliari: (MS).



Departmecit of tasdalena


Three sites near Santa Marta yielded sand flies from tree trunks or

under loose bark of standing trees in cultivated or semicleared areas

(August, 1973).

Rio Don Diego, E of Santa Marta within 1 km of Carribean Sea; "tropi-

cal wet forest;" elevation ca. 3 m.

Between Srn .ta Aarta and Mfnc-a; inear a small stream; probably a

transition zone between "tropical wet forest" and "tropical dry forest;"

elevation ca. 80 m.

Minca, SE of Santa Marta; life zone indeterminate but similar to

preceding site; elevation 200 m.



Andean Region


Nearly all records of phlebotomine species occurring above 1000 m are

given by Osorno et al. (1972a). Based on these and on my limited collec-

tions from the Western Cordillera, it is apparent that the Lutzornyia species

in the verrucarum and vexator groups dominate the rather depauperate

phlebotomine fauna.



Department of Valle


Three sites W or SW of Cali -- 1. Near Pichinde, 2. In forest at the

t;leavision towe, just E of Salidito, and 3. Near Rio Pance, SW of Call;





-25-


"lower montane very wet forest;" elevatiotns from 1570-1700 Im. Limited

collections from 31 July, 1973, to August. 1973.



DenartmenL of Cauca


fastern slope, near summit of Cerro Munchique, west of Popoyan;

"lower uontane rain forest;" elevation ca. 2500 m. Briefly surveyed on

5-6 August, .1973. One Sycorax sp. taken but no phlebotomines.



Oriental Region


Occupying a huge area east of the Andes, the Amazonian fcrest in

Colombia is apparently rich in phlebotomine species but few collections

have been made there or in the more northern Llanos which forms part of

the Oriental Region (Haffer, 1967, fig. 1). Many of the species known

from the Amazonian forests in Brazil, Peru, and Ecuador are expected to

occur also in Colombia.



Comisaria of Amazonas


About 17 km W of Leticia; "tropical moist forest;" elevation 84 m.

PhlebotoTmine flies were relatively rare in late July, 1973, the bulk of

the material being taken in semi-disturbed forests near the Amazon River

on tree trunks, less commonly in light traps. July and August are the

two driest months at this locality.





-26-



Intendencia of Caquota


Tyr-t Asqiu.inaj; transitional life zone, "tropical moist forest"/

"rrop:c-al weot forest;" elevation ca. 200 m. Using light traps, Dr. C.J.

Marinkelle collected numerous phleboto. mines in November, 1971.























Figure 1

Map of Colombia showing localities where phlebotomines and sycoracine
flies have been collected (records based on Osorno et al., 1972a and
this review). Shaded sections represent areas 1000 i+ above sea level.

Legend

S= Pllebotomine record

X = Sycoracine record

4= Phlebotomine and sycoracine record

I = Atlantico Dept.

2 = Magdalena Dept.

3 = Norte de Santander Dept.

4 = Caldas Dept.

5 = Risaralda Dept.

6 = Quindio Dept.

7 = CuDdinamarca Dept.












1h




--28-









Ccath'!an Seo a



z': "- " -*



ep
42
S"0 Venezuela


c %-.. ., . . - tn
S. =b .. m . ....
ccic

o " " -. - i'
c. .x V'. Ant,.o - ".. -" ."' A











".A "azooas
Pr-.- /


,." \ .. / .J . ,-a ../


_- ., i ...--"


"' -.." ."..'. ..G ai ia
../t . -- c ,, ^ v.- '^:.: : . ....... *:

., - , ' '.-S"-' :- ' .../ '* "". vutj� .t
ci ''4,: . ; "'--:f. '" --
,. s /.: c;-""- -- ' f-^': . Voupes 1

&< 'W.+.. 3--"S S.
S --" ,' / - " ---

-cuudo.t " '--" *" ... "





. Pru ./ /















MEDICAL IMPORTANCE



The phlebotomine sand flies, unlike the sycoracine flies, are

notorious vectors of leishmaniasis (Ward, 1977), bartonellosis (Schultz,

1968), and certain arboviruses (Tesh et al., 1974), diseases which

have caused a great deal of human suffering in the neotropics and

else-where.

Leishmaniasis, a collective term referring to several disease

entities caused by Leishmania spp., occurs an Colombia but the vectors

are poorly known. Reyes (1957) provided limited information on the dis-

tribution and nature of dermal (= cutaneous) leishmaniasis in the Republic,

noting that 206 out of 725 human infections involved the naso-pharyngeal

region (mucocutaneous leishmaniasis). Ward, citing figures from Reyes

(op. cit.) and Garnham (1962), stated that nearly 2000 human cases of

cutaneous leishmaniasis were reported in Colombia from 1948 to 1955. The

true incidence, distribution, and identity of leishmaniasis in the

Republic remains largely unknown, however.

Visceral leishmaniasis (kala-azar), a very serious disease, is

apparently rare in Colombia, the few human cases from Santander, Tolima,

and Cundinarmarca Departments having been discussed by Arjona et al.

(1971.).

An outbreak of Bartonellosis (oroya fever, Carrion's disease, etc.),

lasting from 1935 to the early 1940's, in southwestern Colombia (Cauca

and Narino), was reviewed by Jaramillo (1943). From 1940 to 1943, there




-29-





-30-


were 2,241 cases with a mortality rate of nearly 16% (Samaniego, 1944).

The disease, Rnso known to occur in Peru and Ecuador, was probably intro-

duced into Colombia "by returning soldiers or by 'colporteurs' who travel

from village to village" (Rozeboom, 1947b). Circumstantial evidence led

invastigators to believe that Lutzornria cozlmbiana was the responsible

vector.

Arboviruses ware recovered from wild caught phlebotomines in the

PacLfic lowlands of Colombia (Barreto, 1969). The strain, designated

Co Ar 3319, was 4iscussed by Theiler and Downs (1973) and Tesh et al.

(1974). Based upon the pioneering work of Dr. Tesh and his colleagues

in nearby Panama (1971-1975), it is safe to assume that other arboviruses

associated with phlebotomines exist in Colombia.

Sand flies also play a role in the transmission of nonhuman para-

sites, especially protozoans. References to studies related to them are

cited in the species bibilographies.














SUBFAMILY SYCORAC.LNAE



Following Duckhouse (1972) I presently recognize 3 extant genera in

the subfamily -- Sycorax Haliday (ca. 21 spp.), Parasycorax Duckhouse (2

spp.), and Apcsycorax Duckhouse (1 sp.). Until now, only 4 species have

been reported from the New World -- Sycorax assimilis Barretto, 1956 and

Parcwyacora satchelli (Barretto, 1956) from Sao Paulo State, Brazil,

Aposiyorax chiirensis (Tonnoir, 1929) from Chile, and an unnamed sycoracine,

the wing of which was figured by Falrchi.d (1955) from Paienque, Colon

Prov.. Panama.

The feeding habits of most species have not been studied but the

females of SycorCa and Aposycorax at least possess mouthparts adapted for

sucking blood. Sycorax silacea Curtis, the type species of the genus from

Europe, feeds on frogs and transmits a filarial worm to them (Desportes,

1942).

The four new Colombian species, described here in the genus Sycorax,

share several characters which set them apart from the other

sycoracine species. In addition to the paired genital filaments between

the parameres (= cercopods) of the male genitalia, there is a single median

process which seems to form part of the aedeagal complex. It lies above

the ducts and the proximal sperm pump (Fig. 2G). Aposycorax chilensis,

the only sycoracine with inverted male genitalia (Duckhouse, 1972), has

very long genital filaments but it lacks the elongate median process. The

stouL proximal spines (2 or 3) on each style of the Colombian and




-31-





-32-



Parc/:.crax species distinguish them from the extant species in the sub-

fau:i] y.

OTher features, not always repeated in the specific descriptions

which follow, are shared by the 4 Colombian Sycorax species. Coloration:

Dusky brown, mesonoturn but slightly darker than pleuron. Head: Broader

than long; frons putfed out behind antenna. Palpus of 4 segments, the

second with an inner group of 10-20 sensory rods. Antenna slender, of 16

"segments" (scape, pedicel, and 14 flagellomeres), the last reduced in

size and terminating in a cone shaped peg sensilla (two similar sensilla

on flageliomere XIV); outer base of scape with 2 short sensory hairs as

in A. chi.ensis; pedicel with 2-3 similar hairs; ascoids subequal in

size, often difficult to examine in available material. Female mouthparts

include toothed mandibles and 4 boot shaped sensilla at tip of labrum;

these features absent in males. Cibarium unarmed. Thorax: Pleura with

6+ setae mainly on the anepimeron but some may extend downwards to the

katepimeron; upper episternal setae present or not; both sexes with 4

straight bristles below base of haltere (Fig. 2H). Wing broadly rounded;

veins sparsely haired; radial fork distad of medial fork. Abdomen: Fe-

males without setae on tergites 8 and 10 or sternite 8. Male genitalia

not inverted. Style with a patch of 8-15 short pointed steae at base;

terminal spine slightly beveled at tip, 2 or 3 proximal spines present,

with or without a long subterminal hair; aedeagal complex as mentioned

above; the paired genital filaments turned upwards, sinuous or not;

parameres comewhat resembling those of A. chitensis but differing in the

nacure and position of the lobes, setae, and projections. Female geni-

talia: Each spermatheca and sperm duct forming a continuous tube with

faint transverse striations, terminating in a heavily sclerotized "button"





-33-


which pnpecars shiny black; a tenuous sac usually amorphic following

treatment in KOH, arises from the end of the button; an internal apodeme

or furca associated with paired sperma.thecae.

The combined length of the spermatheca and duct varies interspeci-

fically an: probably ccrrespontd� to the length of the male. genital

filaments. This feature and the distribution of the antennal setae and

distance between the compound eyes were also used in associating the

sexcs of three of the Colombian species but there remains the possibility

that the males and females were not correctly associated. The immature

stages of these and other New World species in the subfamily have not

been discovered.



Keys to the Sycoraxz Species


Mal es

1. Style of male genitalia with 3 strong spines and long subterminal

hair; median process of aedeagus parallel-sided in dorsal view; lower

distal projection of paramere pointed at tip. Antennal sensory setae,

excluding the paired ascoids, arranged in a whorl-like pattern on

flagellomeres. . . . . . . . . . . . . . . . . . . . . . . . . . . . 2


Style with 3-4 strong spines but without long subterminal hair;

median process of aedeagus with basal two-thirds

expanded and terminal one-third slender in dorsal view; lower distal

projection of paramere rounded at tip. Antennal sensory setae,

excluding the paired ascoids, not arranged in a whorl-like

pattern. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3





-34-


2. Aedecgal du-:ts r~elatively lotg, exceeding the length of the specrm

pump. A subtriangular plate, dorsoventrally flattened, with acute

biurutat_ tip present below the base of the median process of aedeagus.

Tip of median process of aedeagus rounded in lateral view

. . . . . . . . . . . . . . . . . . . . . . . . colombiensis (Fig. 3)


Aedeagal ductu shorter, less than length of sperm pump and not as

sinuous. Subtriangular plate absent. Tip of median process of

aedeagus angular in lateral view. . . . . . . ... fairchildi (Fig. 4)


3. Style oE male genitalia with 4 strong spines; median process of

aedeagus more slender (Fig. 2G). A larger species, wing length greater

than 2 mm . . . . . . . . . . . . . . . . ... . . . andicola (Fig. 2)


Style with 3 strong spines; median process of aedeagus relatively

stout (Fig. 51). Wing length less than 1.5 mm. . trispinosa (Fig. 5)



Fi-males

1. Flagellomeres I-VIII with paired ascoids, other antennal setae ran-

domly distributed. Eyes rather narrow, separated by distance = to

7 facet diameters . . . . . . . . . . . . . . . . trispinosa (Fig. 5)


Flagellomeres T-IV with paired ascoids, one group of antennal setae

arranged in whorl-like pattern. Eyes separated by 8 or more facet

diameters . . . . . . . . . .. . . . . . . . . . . . . . ... . . . . 2





-35



2. Firca spe e-shaped apically, its length at least � that of sperm duct

+ spenratheca. Flagellomero, 1, 0.20 mm, or longer . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . oo or fiens (Fig. 3)


Furea "v" shaped apically, its length but slightly less than that of

sperm duct + spermatheca. Flagellomere I less than 0.20 mm long

. . . . . . . . . . . . . . . . . . . . fa rchidi (Fig. 4)



1. Sycorax andicola n. sp.
(Fig. 2)


Male (holotype): Wing length 2.35; width 1.05. Head height 0.31;

,ridth, 0.37. Eyes scparated by 0.14 or by distance ro 8 facet dia-

meters. Antenna (including scape and pedicel) 1.61 long; flagellomere I

(0.26 long), about 1.8xlength of flagellomere 1; paired ascoids (Fig.

2C), subequal in size, on flagellomeres I-IV only; other antennal setae

mostly deciduous, not arranged in a whorl-like pattern. Palpal length

0.17; ratio of segments: 1-1.02-0.72-0.67; second segment with 20+ strap-

like sensory rods in a distinct patch. Pleura with 19-22 setae below

wing base, without episternal setae. Wing venation as figured. Length

of femora, tibiae, and basitarsi: Foreleg, 0.63, 0.73, 0.49; midleg, 0.63,

0.75, 0.47; hindleg 0.73, 0.82, 0.44. Visible sternites 2 and 3 with

paired circular openings laterally. Genitalia complex: Coxite 0.31 long.

Style 0.18 long with 1 terminil spine, slightly beveled at tip and with a

proximal row of 3 smaller spines. Parameie as shown with several setose

l.bes aind projections, the most distal finger-like with rounded, upturned

tips. Aedeagal complex of a laterally compressed sperri pump (0.20 long)

within the abdomen; a median distal process (0.165 long) compressed dorso-

ventrally, curved and slender in lateral view, upturned at tip, basal





-36--



two-thirds much wider than distal one-third when viewed dorsally; paired

sinuos, relatively long genital filaments. Cerci as figured.

Material examined: Colombia. d holotype (no. 537), Cerro Munchique

(Cauca), 2450 m above sea level, light trap in cloud forest, 6 Aug. 1973,

D.G.Y. and R.C.V. coll. c paratype (no. 538), same data except collected

by R.C.W., 8 Aug. 1975.

Di.scuss-ion: S. andicola, the largest Sycorca known from Colombia,

closely resembles S. trispinosa n. sp. in details of the male genitalia,

setation of the antenna and in other character states. The two species,

apparently allopatric, are separated by the characters given in the key.



2. SQcorax coZombiensis n. sp.
(Fig. 3)


Male (holotype): Wing length 1.32; width 0.47. Head height 0.21;

width 0.25; eyes separated by 0.126 mm or by distance = to 10 facet

diameters. Flagellomere I (0.19 mm long), about 2.2 xlength of flagello-

mere II. Paired ascoids visible only on flagellomeres I and II; other

sensory hairs in a whorl-like pattern on flagellomeres I-X, the remaining

flagellomeres missing. Palpal length 0.10, ratio of segments 1-0.91-0.80-

0.72. Pleura with 9-12 setae in one specimen, but lacking in the

holutype. Wing venation as figured. Legs missing. Nature of sternites

not determinable. Genitalia: Coxite (0.177 long); style (0.98 long)

with a terminal spine, a long subterminal bristle and 2 proximal spines.

Paramere as figured, the lower distal projection blade-like, pointed at

tip. Aedeagal complex: Sperm pump (0.10 long); median distal process

(0.09 long), slender and subequal in width in dorsal view, with a slender,

rounded tip in lateral view; a subtriangular plate, dorsoventrally





-37-


flattened, with acute bifurcate tip, pointing to the rear, present below

the .ase of the median process of aedeagus. Cerci as shown.

Female (allotype): Wing length 1.37; width 0.59. Head height, 0.24;

width, 0.29; eyes separated by 0.15 or by distance = to 10 facet diameters.

Antenna (1.15 long), flagellomere I (0.22 long), about 2.4 xlength of

flagellomere II; paired ascoids on flagellomeres I-IV; other sensory

hair- arranged in a whorl-like pattern on flagellomeres I-XV. Palpal

length, 0.113; ratio of segments 1-1-0.75-0.58. Pleura with 16-18 setae

below wing base and 0 or 1 upper episternal seta. Wing venation as shown.

Iegs partly or wholly missing in all specimens, length of femora, tibiae,

and basitarsi of foreleg: 0.43, 0.51, 0.27. Abdominal sternite 2 with

paired circular openings at sides; other sternites apparently lacking

these openings. Spernatheca and sperm duct length at least 0.15 from

base to tip of button; furca spade-shaped at end.

Material examined: Colombia. e holotype (no. 529), Anchicaya Dam

(Valle), elev. ca. 400 m above sea level, light trap, 28 Jan. 1975, R.C.W.

a allotype (no. 530), same data as holotype except collected 10 June

1975. Paratypes (nos. 531-538), all from type locality in light traps;

1 o, 11 Aug. 1973, D.G.Y. and R.C.W. 1 o', 2 oo, 28 Jan. 1975; 2 o9, 10

June 1975, R.C.W.

JDiscussion: The male of S. cooombiensis, the only Colombian species

with e flattened subtriangular plate below the base of the median distal

process of the aedeagus, was at first confused with S. Fairchildi but the

presence of the plate and the greater length of the aedeagal ducts

readily separate the 2 species. The female of S. cooombiensis share

several character states with S. fairchildi and S. trispinosa but may be

identified by the characters in the key.





-38-


3. Szycorax fairchi&di I:. sp.
(Fig. 4)


Male (holotype): Wing length, 1.13; width, 0.47. Head height,

0.23; width, 0.28; eyes separated by 0.13 or by distance = to 10 facet

diameters. Antenna about 0.79 long; flagellomere I (0.18 long), about

2.5 xlength of flagellomere II; paired ascoids visible only on flagello-

meres I-IIl, other sensory hairs in a whorl-like pattern on flageliomeres

I-XV, palpal length 0.10 mm; ratio of segments: 1-1.18-1-0.72. Pleura

with 5-11 setae below wing base and with 1-2 upper episternal setae (n = 5).

Wing venation as shown. Sternites apparently lacking clear circular

openings. Length of femora, tibiae, and basitarsi of slide 544: Foreleg,

0.42, 0.43, 0.24; midleg, 0.45, 0.40, 0.24; hindleg, 0.47, 0.37, 0.20.

Genitalia: Coxite, 0.16 long; style, 0.88 long, with a large terminal

spine, a long subterminal hair and 2 proximal spines. Lower distal end

of paramere terminating in a hook-like projection, acute at: tip. Aedeagal

complex: Sperm pump (0.11 long); median distal process (0.068 long),

angular at tip and relatively wide in lateral view, subequal in width

when viewed dorsally; aedeagal ducts short, each about 0.96 long. Cerci

as shown.

Female (allotype): Wing length, 1.47; width 0.61. Head height,

0.26; width, 0.30; eyes separated by 0.14 or by distance = to 9 6 facet

diameters. Flagellomere I (0.16 long) about 2.3 xlength of flagellomere

II; paired ascoids, subequal in size (but not conspicuous) on flagello-

meres 1-IV, absent from remaining flagellomeres, other nondeciduous

antennal setae as in male. Palpal length 0.116; ratio of segments 1-1-

0.92-0.78. Pleura with 10-11 setae below wing base and with 1-2 upper

episternal setae (n = 3). Wing venation as shown. Length of femora,





-39--


tibia., and basitarsi: Foreleg, 0.39, 0.40, 0.22; midleg, 0.42, 0.47,

0.24; hindlcg, 0.47, 0.50, 0.20. Sperilatheca + sperm duct length about

0.08 from base to end of subcircular button: furca somewhat "v" shaped

apically.

Mateelal examined: Colombia. o hclotype (no. 539), Anchicaya Dam

(Valle), elev. ca. 400 m above sea level, light trap in forest, 11 Aug.

1973, D.G.Y. and R.C.W. q allotype (no. 540), same data except collected

28 Jan. 1975, R.C.W. Paratypes (nos. 541-547) all collected in light

traps, 1 S, Rio Anori (Antioquia), Colombia, elev. ca. 410 m above sea

level, 22 Sept. 1970, D.G.Y. 1 (, Rio Anori, 23 Sept. 1970. 1 o,

Anchicaya Dam, 11 Aug. 1973, D.G.Y. and R.C.W. 3 W$, 1 o Anchicaya Dam,

28 Jan. 1975, R.C.W. 1 6', Anchicaya Dam, 5 March 1976, R.C.W.

Discussion: The short sperm ducts of S. fairchildi serve to dis-

tinguish this taxon from the other species of Synorax in Colombia.

I take pleasure in naming the species in honor of Dr. G.B. Fairchild

who has contributed so much to our knowledge of medically important

arthropods in the neotropics.



4. Sycorax trispinosa n. sp.
(Fig. 5)


Male (holotype): Wing length 1.22; width 0.56. Head height 0.23;

width 0.28; eyes separated by 0.08 or by distance = to 7 facet diameters.

Antenna, 0.88 long; flagellomere I (0.15), about 1.8 xlength of flagello-

mere II. Paired ascoids on flagellomeres I-IV, absent from remaining

flagellomeres; other antennal setae mostly deciduous, not arranged in a

whorl-like pattern. Palpal length, 0.12 am; ratio of segments: 1-1.15-

0.84-0.76, second segment with 15� rod sensilla. Pleura with 9-16 setae





-40-


below wing base ana 1-4 upper episternal setae (n = 10). Wing venation

as figured. Length of femora, tibiae, and basitarsi: Foreleg, 0.45,

0.43, 0.26; midleg, 0.49, 0.44, 0.26; hindleg, 0.53, 0.52, 0.24. Abdominal

sternites 2-5 with paired lateral openings. Genitalia: Coxite, 0.19 long.

Style, 0.09, with 2 proximal spines and a terminal spine. Paramere as

shown, similar to that of S. andicola, the most distal projection also

curved at tip, finger-like. Aedeagal complex of a laterally compressed

sperm pump (0.11 long), a median distal process (0.10 long), with the

basal two-thirds expanded, greater than twice the width of the terminal

third in dorsal view; aedeagal ducts and cerci as shown.

Female (allotype): Wing length 1.54; width 0.71. Head height, 0.31;

width, 0.27; eyes separated by distance of 0.1 or by distance = to 6.9

facet diameters. Antenna, 1.0 long; flagellomere I (0.16 long), about

1.7 x length of flagellomere II. Paired ascoids on flagellomeres I-VIII,

absent from remaining flagellomeres, other antennal setae mostly deciduous

not arranged in a whorl-like pattern. Palpal length 0.14 mm; ratio of

segments: 1-0.89-0.61-0.61, palpal sensilla as in male. Pleura with

10-21 setae below wing base and 1-5 upper episternal setae (n = 9); wing

venation as shown. Length of femora, tibiae, and basitarsi: Foreleg,

0.44, 0.42, 0.27; midleg, 0.49, 0.48, 0.27; hindleg, 0.53, 0.56, 0.27.

Abdominal sternites 2-5 with paired circular openings at sides, remaining

sternites indeterminate. Combined length of spermatheca and sperm duct,

from base to tip of sclerotized button, 0.15. Furca spade-shaped as

shown.

iMaterial examined: Colombia. S holotype (no. 548), Anchicaya Dam

(Valle), elev. ca. 400 m above sea level, light trap in forest, 5 March

1976, R.C.W. o allotype (no. 549), same data. Paratypes (nos. 550-573),





-41-


all collected from type locality in light or flight traps by D.G.Y.

and/or R.C.W. 2 q , 10 Aug. 1973; 1 o6, 1 o, 11 Aug. 1973; 13 dd', 4 o,

28 Jan. 1975; 1 6, 2 oo, 5 March 1976.

DisUcusion,: I associated the male and femaJe of S. trispinosa on

the basis of the following characteristics which. in combination, are not

shared by S. fairchiZdi or S. coorrbiensis. Flagellomere I relatively

short (0.14-0.19 mm, n = 27); compound eyes more narrowly separated than

chose of the other sympatric species; antennal setae (excluding the paired

ascoids) mostly deciduous, scattered; aedeagal ducts and spermathecae

generally corresponding in length.

The males of S. trispinosa and S. andicola are quite similar but

differ in size and by other characters given in the key. The female of

a.dicola, presently unknown, probably resembles trispinosa in details of

the spermathecae and palpi.

The specific name refers to the 3 strong spines on the style of the

male.
























Figure 2

SuCoras: oadicola male -- A. Head, B. Wing, C. Pedicel and flagellomeres
I and II showing ascoids, D. Terminal 4 flagellomeres, E. Palpus,
F. Genitalia, dorsal view, G. Sperm pump, genital filaments, and median
dorsal process of aedeagus, lateral view, H. Metathoracic spiracle and
postspiracular setae.

Male: Ccrro Munchique, Cauca Dept., Colombia




-43-










.0 . ,










/ ti
S'Ae--w / cs niotl rtr lamc1 ke ci


















*-*"f [-1! (^ "" . . r- . .. :
- I










1g. ,g. ' 7i,=-
.E o
Sp rm Vr-.p
1Me Da'I Pvac s
1 et . <0<;? yp

f A rd-ogut
il ISpern Pump et.. . .t. 2
'I N N -- -




001
























Figure 3

Sycor'ax c-oloimbt.nsi -- A. Male head, B. Female wing, C. Male wing,
D. Female head, E. Male genitalia, dorsal view, F. Male subtriangular
plate, G. Spermathecae, H. Sperm pump, genital filaments,and median
dorsal process of aedcagus, lateral view.

!]ale: Anchicaya Dam, Valle Dept., Colombia
Female: Same locality as male




-45-











*V? c~--c-'^


V D




i-Ser Po~mp
V/ 0




r y/ F: r1p
U i __ Se~mPerm hi


1..
r.O


I;' ~; :
~-�
E ; 1



H--S
KiAS
























Figure 4

Syco .r fairchildi -- A. Male head, B. Female wing, C. MHle wing,
D. Female head, E. Male genitalia, dorsal view, F. Sperm pump, genital
filaments,and median dorsal process of aedeagus, lateral view,
G. Spernathecae, H. Female pedicel and flagellomeres I and II showing
paired ascoids and other seLae.

Male; Anchicaya Dam, Valle Dept., Colombia
Female: Same locality as male




-47-












V N I



A V
C







UKCji. <,KN (

* * Ij/ I I It



- M- n
3 EI H



G
p-:)\ 'JV
























Figure 5

Sycorax triapinosa -- A. Male head, B. Female palpus, C. Female,
terminal 4 flagellomares, D. Female pedicel and flagellomeres I and
II showing ascoids, E. Female head, F. Female wing, G. Male wing,
H. 1ala genitalia, dorsal view, I. Sperm pump, genital filaments,and
median dorsal process of aedeagus, lateral view, J. Tip of furca,
dorsal view, K. Spermathecae,

Male: Anchicaya Dam, Valle Dept., Colombia
Female: Same locality as male




-49-














r Y. GI
B D






V Art G I


/'KS 'i (I




IlK (CX K
"-V ^ .. '













SUBFAMILY PHLEBOTOMINAE



Previous studies on this subfamily in Colombia were reviewed by

Osorno at al. (1972a) and will not be repeated here except for literature

citations in the species accounts. Nearly all of these studies involve

systematics, especially species descriptions and distribution records.

Owing to the paucity of bionomic studies on the Colombian phlebotomines,

I felt that it would be desirable to provide references to such studies

carried out in other countries. The results of these studies may not

always apply to conspecific populations in Colombia. Information on the

biting habits of some species in Choco Dept. is provided in the species

accounts.

With few exceptions I have not redescribed known taxa because of

adequate original or subsequent descriptions. Hopefully, the figures

and keys will serve to distinguish the taxa.



Key to the New World Genera


1. Wing broad, rounded at tip; R2 + R3 + R4 forks before, on same level

as, or slightly beyond r-m crossvein. Pleura without episternal

setae. Female cibarium unarmed. Male genitalia with style longer

than coxite . . . . . . . . . . . . . . . . ... . Warileya (Fig. 6-7)


Wing pointed at tip; R + R3 + R forks well beyond r-m crossvein.

Pleura with episternal setae. Female cibarium armed with teeth.

,ale genitalia with style shorter than coxite . . . . . . . . . . . 2



-50-





-51-



.. I terocular suture complete. Female cibarium with 4 longitudinal

rows of horizontal teeth. Male genitalia with 5 large spines, 2 of

which (usually basal pair) borne on a common tubercle . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . .Bumptomynia (Fig. 8-11)


Interocular suture incomplete. Female with 1 row of horizontal

teeth; vertical and lateral teeth present or not. Male genitalia with

style bearing 1-6 large spines, basal pair not borne on a common

tubercle in those species having 5 spines . . Lutzomyia (Fig. 12-89)





-52-


Genus Warileya Hertig, 1948


This genus, recently enlarged to accomodate W. (H.) hert-ii (Fchld.)

by Lewis et al. (1978), contains 5 species, 2 of which in the subgenus

Wayt',-eySa occur in Colombia. Little is known about the habits of any of

the species although W. (W.) rotundipennis, one of the Colombian species,

and W. (W.) phlebotomanica Hertig, the type species from Peru, are

anthropophilic.



Keys to Species


Males

1. Palpal segment 5 longer than combined length of segments 2 + 3.

Paired intraabdominal rods present. Genital filaments about 1.8X

length of pump. Style lacking a basal bristle. Wing length less

than 1.5 mm. . . . . . . . . . . . . . .... ..n-igrosacculus (Fig. 6)


Palpal segment 5 shorter than combined length of segments 2 + 3.

Intraabdominal rods absent. Genital filaments less than 1.3X length

of pump. Style with a basal bristle. Wing length over 1.8 mm . . .

. . . . . . . . . . . . . . . . . . . . . . . . rotundipennis (Fig. 7)



Females

1. Sperrathecae large and sac-like with smooth walls. Palpal segments

5 longer than combined length of segments 2 + 3. Wing length less

than 2.0 mm. . . . . . . . . . . . . . . . . . nigrosacculus (Fig. 6)


Spermathecae cylindrical, somewhat worm-like, with transverse stria-

tions. Palpal segment 5 shorter than combined length of segments

2 + 3. Wing length over 2.0 mm. . . . . . . . rotundipennis (Fig. 7)





-53-


5. Warileya (W.) nigrosrceulVus
(Fig. 6)


Waritcya nigrocaecclus Fairchild & Hertig, 1951b: 428 (o holotype.

Carro Campana, Panama Prov., Panama). Fairchild, 1953: 102 (cf. to

Hertigia hertigi). Barretto, 1955a: 188 (listed). Forattini, 1971a:

107 (listed). Tesh et al., 1971a: 153 (blood meals, Panama). Martins &

Morales, 1972: 366 (listed). Young & Chanictis, 1972: 97 (di, descr.).

Chaniotis et al., 1972: 95-96 (in tree hollows, Panama). Christensen,

1972a: 88 (listed). Forattini, 1973: 536-538 (9, fig. ). Velasco &

Trapido, 1974: (cf. to W. yungasi). Lewis, 1975a: 500 et seq. (mouth-

part morphol.). Lewis et al., 1978 (classif.).

Distribution: Colombia (Choco, Valle), Panama.

Material exanined: Colombia: 1 d , 1 , Curiche (Choco), Malaise

trap, 27 April 1977. 1 S, 1 �, Anchicaya Dam (Valle), light trap, 28 Jan.

3975, R.C.W. 1 d, same data but taken 16 July 1975. Panama: $ holotype,

Cerro CamTpana (Panama Prov.), hollow log, 7 Jan. 1947, M. Hertig. 2 & S,

5 T?, near Gamboa, Canal zone, tree hollow, 10 Sept. 1969, B. Chaniotis.

4 Sde, same data but taken 15 Jan. 1970.

Discussion: A little known species, W. nigrosacculus is easily

distinguished from W. rotundipennis by the characters given in the key.

Using a precipitin test for blood meal determinations, Tesh et al. (1971a)

found that blood from 4 recently engorged nigrosacculus females reacted

with mammalian and reptile-amphibian antisera. Four other blood neals

were nonreactive due to small volumes of blood and/or to the weakness of

the reptile-amphibian antisera.





-54-


6. Warileya (W.) ro tundipennis
(Fig. 7)


Wacuiieya rotundipennis Fairchild & Hertig, 1951b: 424 (( hol type,

o, Cerro Campana, Panama Prov., Panama). Fairchild, 1953: 102 (cf. to

Hertigia hertigi). Barretro, 1955a: 188 (listed). Fairchild & IIertig,

1959: 122 (Costa Rica). Johnson & Hertig, 1961: 765, 775 (rearing).

Ranson, 1968: 93 (larval fig.). Forattini, 1971a: 107 (listed). Young

& Chaniotis, 1972: 366 (listed). Christensen, 1972a: 88 (listed).

Forattini, 1973: 536-538 (c6, o fig.). Velasco & Trapido, 1974: 436 (cf.

to W. yungasi). Lewis, i975a: 500 et seq. (mouthpart morphol.). Lewis,

1975b: 366 (hair sockets, fig.). Lewis at al.: 1978 (classif.).

Distribution: Colombia (Antioquia, Choco, Valle), Costa Rica, Panama.

Material examined: Colombia. 2 99, Teresita (Choco), tree buttress,

15 June 1967. 27 �o, Rio Anori (Antioquia), light traps, Sept. 1970.

5 ~ , same data but 29 May 1970, C.H.P. 8 J', 2 oo, Anchicaya Dam (Valle),

light trap, 28 Jan. 1975, J.E. Browne. 6 d 6, 6 oo, same data but taken

10 June 1975, R.C.W. 5 6d', 4 q?, same data but taken 16 July 1975.

Panama. S holotype (no. 2335), o allotype (no. 2312), Cerro Canpana

(Panama), Shannon trap, 24 Aug. 1950, M. Hertig & G. Fairchild. 2 d',

Rio Changena (Bocas del Toro), Shannon trap, 8-11 Sept. 1961, R. Hartmann

& P. Galindo. Costa Rica. 2 'c', 1 ?, Turrialba, Shannon trap, 20 Aug.

1961, G. Fairchild & M. Hertig.

Discussion: WariZeya rotundipennis, the only W'arileya species lacking

intra-abdominal. rods (Lewis et al., 1978), has been reported feeding on man

in Panama (Fairchild & Ilertig, 1951b) but its role, if any, in disease

transmission is not known. Dr. C.H. Porter also has collected man-biting






-55-


femaies at the Rio Anori (Antioquia) locality, mostly in forest clearings

(pers. comm.) .

The records of W. rotundipennis and W'. nigrosacculus from Choco and

Valle Depts. indicate that both taxa probably occur in other areas of the

trans-Andean region perhaps as far south as Guayas Prov., Ecuador.



























r Pro., Ecadox
























Figure 6

Warilcya nifrosacculus -- A. Hale head, B. Male flagellomere II,
C. Female head, D. Female flagellomere II, E. Female cibarium and
pnarynx, F. Male genitalia, dorsal, G. Tips of genital filaments,
H. Female wing, I. Male wing, J. Spermathecae.

Male: Cutiche, Choco Dept., Colombia
Female: Same locality as male




-57-







N',


I I E1


, B









," ^ ^ ''-'^-- .--.

.. . ... " J.-
.-* - --_ _ _-^-----^ , ,










- __ , i
























Figure 7

Thariteya rotundipennis -- A. Male head, B. Male flagellomere II,
C. Female head, D. Female flagellomere II, E. Female cibarium and
pharynx, F. Male genitalia, G. Female wing, H. Male wing, I. Sperma-
thecae.

Nale: Cerro C:npana, Panama Prov., Panama
Female: Rio Anori, Antioquia Dept., Colombia












B D
II 9 ?4t K P / i/A

d ' ? �," j


S - iC E





K' -JN ^' _ 4

GO 4'

H N -*-'-!





-60-


Genus Blruzmwtomyia Franca and Farrot, 1921


Characterized by Theodor (1965), Lewis et a]. (1978), and others,

the genus Bruriptomyia now contains over 21 species, 5 of which occur in

Colonbia. Except for the female of B. hamnata which has very short sperm

ducts (Fig. 101), the females of the other species in Colombia are

virtually impossible to identify in the absence of males.

Brtputonrjia aveitari, one of the Colorrbian species, was reported

feeding on armadillos in Brazil (Mangabeira, J942b). This and other

Brunrptcrmia spp. often rest in burrows dug by these mammals. They do not

bite man and the preferred hosts of most species remain unknown.



Key to Species


Males

1. Genital filaments less than 4X length of pump. . .. hamata (Fig. 10)


Genital filaments greater than 4X length of pump . . . . . . . . . 2


2. Coxite tuft of mostly stout, spine-like setae . . . . . ... . . . 3


Coxite tuft of slender, hair-like setae. . . . . . . . . . . . . . 4


3. Coxite tuft of 20+ setae implanted on a distinct tubercle. Paramercs

more slender as shown. . . . . . . . . . . .... ..galindoi (Fig. 9)


Coxite tuft of fewer than 20 setae, usually about 12, not implanted

on a tubercle. Paramere broader as shown. . . . .avellari (Fig. 8)


4. Style with isolated spine well distad of proximal pair. Coxite tuft

implanted on a distinct raised tubercle. ... . . eaupertuzyi (Fig. 8)





-61-


Style with isolated spine at or near level of proximal spines. Coxite

tuft implanted on a slightly raised, raspberry-like (circular)

base. . . . . ... .. . . . . . . . . . .. . ... .eopoldoi (Fig. 11)



7. Brtmptomyia ave lani
(Fig. 8)


Fhtebotomus avellari Costa Lima, 1932: 48 (c, Lassance, Minas

Gerais, Brazil). Mangabeira, 1942b: 225 et seq. (immatures, adults,

descr., fig.). Barretto, 1947: 187-188 (full refs.). Fairchild &

Hertig, 1947a: 615-616 (cf. to galindoi). Barretto, 1951: 212 (dist.).

Floch & Abonnenc, 1952: 39, 45 (
(mention). Forattini, 1954: 214-217 (second sternite, fig.). Forattini

& dos Santos, 1955: 17 (Brazilian record). Ortiz, 1963: 320 (7, keyed).

Brumptor~ia avellari: Barretto, 1955a: 187 (listed). Martins et

al., 1961b: 309 (mention). Martins et al., 1962a: 380 (Goias, Brazil).

Sherlock, 1962: 332, 335 (mention). Carnheiro & Sherlock, 1964: 315

(pupa, keyed, fig.). Fraiha et al., 1970a: 468 (cf keyed). Christensen,

1972a: 88 (Panama). Osorno et al., 1972a: 14 (Boyaca, Colombia).

Forattini, 1973: 122 et seq. (gen. review, fig.). Llanos et al., 1976:

480 (Peru). Martin et al., 1976b: 496 (Peru). Ramirez et al., 1976:

599 (Venezuela).

Distr-ibution: Panamna Colombia (Boyaca), Peru, Venezuela, Brazil,

Paraguay.

Mat.erial examined: Colombia. 2 66', 1 , Puerto Boyaca (Boyaca),

light trap, 6 May 1973, C.J.M. Brazil. 1 6, Coqueiros (Sao Paulo),

armadillo burrow, Oct. 1953, Albertin. Panama. 1 ', Tocumen Airport

(Panama), light trap, 23 March 1953, F. Blanton. Paiaguay. 5 Vd, Aca-

Poi, San Pedro, burrow, 18 April 1950, M. Hertig.





-62-


Disc S_ ion: Brumptomyia avcllari and B. brt',ypti (Larousse) with

sympatric populations in Brazil and Paraguay, closely resemble one

ainolher, the males differing chiefly by the shape of the parameres. I

regard the Colombian specimens as being conspecific with the former

soecies.



8. Bruznptornyia beaupertuyi
(Fig. 8)


PhZebotomr.us beaupertziy Ortiz, 1954: 235 (dholotype, Duaca, Lara

State, Venezuela). Scorza & Ortiz, 1960: 434 et seq. (ecology). Pifano

et al., 1962: 383, 387 (r, q, keyed), 411-412 (C, ., descr., fig.).

Ortiz, 1963: 322 (Q descr.). Ortiz & Scorza, 1963: 350 (listed), 354

(C keyed). Ortiz, 1965a: 208 (mention). Scorza et al., 1967: 193,

195 (6, o keyed). Leon, 1969: 30 (listed).

Phlebot;oimus gaZindoi (not aalindoi Fairchild & Hertig, 1947): Pifano

& Ortiz, 1952: 138 (listed, Venezuela). Leon, 1969: 30 (listed).

Brwnptomyia beaupertuyi: Barretto, 1955a: 187 (listed). Sherlock,

1962: 321 et seq. (d, fig., Colombia). Fraiha et al., 1970a: 468 (6,

keyed). Forattini, 1971a: 98 (listed). Forattini, 1973: 522 et seq.

(gen. review, fig.).

Distribution: Colombia (Santander), Venezuela.

Material examined: Venezuela. 2 c6 (topotypes), Duaca (Lara), no

date, I. Ortiz.

Discussion: At present, this little-known species is represented in

Colombia by a single male, skillfully illustrated by Dr. 0. Mangabeira

(in Sherlock, 1962). Ortiz (1963) described the female based on Venezuelan

specimens.





-63-


9. Bru.Ttoz4yia ga -i*:doi
(Fig. 9)


Phlebotomus galindoi Fairchild & Hertig, 1947a: 615 (6 holotype,

near hJquete, Chiriqui Prov., Panama). Floch & Abonnenc, 1952: 40 (6

keynd). Ortiz, 1954: 238 (cf. to beaupertuyi). Rosabal, 1954: 30 et

seq. (S fig., Costa Rica). Fairchild & Hertig, 1959: 122 (geographic

records). Hanson, 1961: 320 et seq. (breeding sites, Panama). Johnson

& Hertig, 1961: 765 et seq. (rearing data). Diaz-Najera, 1963: 193

et seq. (9 desci., Mexico, fig.). Ortiz & Scorza, 1963: 354 (' keyed).

Biagi et al., 1966: 149 (Mexican records), 151 (d fig.). Strangways-

Dixon & Lainson, 1966: 193 (Belize). Hanson, 1968: 56-58 (larva, pupa,

descr., fig.).

Brumptomyia galindoi: Barretto, 1955a: 187 (listed). Osorno et al.,

1972a: 14-15 (Colombian records). Forattini, 1973: 122 et seq. (gen.

review, fig.). Lewis, 1975a: 500 et seq. (mouthpart morphol.).

Brurptomyia mesai Sherlock, 1962: 332 (6c, San Vicente de Chucuri,

Santander, Colombia). Williams, 1970: 331 (Belize). Fraiha et al.,

1970a: 468-469 (as synonym of galindoi). Forattini, 1971a: 98 (listed).

Williams, 1976a: 603 et seq. (in caves, Belize).

Distribution: Mexico, Belize, Honduras, Costa Rica, Panama, Colombia

(Caqueta, Choco, Bolivar, Boyaca), Ecuador, Paraguay.

zaterial examined: Colombia. 1 c', Puerto Boyaca (Boyaca), light

trap, 6 May 1973, C.J.M. 1 d', Tres Esquinas (Caqueta), light trap, 11

Nov. 1971, C.J.M. 27 6'd, 11 , Teresita (Choco), light & Malaise traps,

tree trunks, April-Aug. 1967, D.G.Y. Costa Rica. 1 d", San Carlos

(Alajuela) tree buttress, 23 June 1955, R. Rosabal. Ecuador. 5 dd',

4 po, Rio Napo at Limoncocha (Napo), light & flight traps, tree buttresses,





-64-


20-24 May 1976, D.G.Y. & T. Rogers. Honduras. 1 d, Tela, Lancetilla

Valley, tree buttress, 24 July 1953, W. Hils. 1 3, same data but light

trap, 13 Jan. 1954. Mexico. 2 65, 2 ?p, Ocoscoautia (Chiapas), tree

hollows, 8 April 1951, G. Fairchild & R. Hartmann. 2 '1, Paleuque

(Chiapas), tree buttresses, 30-31 March 1951, G. Fairchild & R. Hartmann.

Paraguay. 1 3, Sommerfeld, Yhu, tree cavity, 15 March 1950, V. Zelada.

1 (', same data but 18 March 1950, M. Hertig. Panama. 1 d (holotype no.

814), type locality, 17 Feb. 1947, P. Galindo. 1 o, same data but 26

March 1948. 1 d", 1 p, Mojinga Swamp neor Ft. Sherman, Canal Zone, light

trap, 19 Nov. 1951, F. Blanton.

Discussion: The holotype of B. galindoi is a large specimen, wing

length nearly 3.0 mm, from Chiriqui Province, Panama (ca. 1200 m above

sea level). Smaller specimens from lower elevations in Santander Depart-

ment, Colombia were later named B. mesai by Sherlock (1962) who separated

the males on size, number of distal setae on the coxites, shape of the

aedeagi and setation of the parameres. Fraiha et al. (1970a) treated the

taxa as conspecific, basing their decision on the original descriptions

and on a male (identified as B. galindoi by Dr. G.B. Fairchild) from a

lowland locality in Panama.

Aside from size and the number of distal coxite setae (7 in the

gazindoi holotype, 5-6 in males from the lowlands), I can detect no marked

differences among the specimens listed above. The number of distal setae

on the coxites probably varies according to the size of the insect and in

this case, apparently represents infraspecific variation. It remains to

be determined whether or not the size of galindoi individuals is correlated

with altitude, i.e., is there a continuous increase in size of specimens

from the lowlands to the highlands? Believing this to be the case but





-65-


'wthout evidenc- to support it, I tentatively consider B. mcsai and

B. galiridoi to be conspecific following Fraiha et al. (1970a).



10. Brunptomyia hcamcta
(Fig. 10)


Phlebotom4.us hcanatus Fairchild & Hertig, 1947a: 614 (d holotype,

Chilibrillo bat caves, near Chilibrc, Panama). Barretto, 1951: 217

(mention). Floch & Abonnenc, 1952: 39 (6 keyed). Rodriguez, 1953b: 53

(6, o measured, Ecuador), 55 (cf. to Zeopoldoi). Ortiz, 1954: 237

(listed). Lewis & Garnham, 1959: 83-84 (S, Belize, fig.). Garnham &

Lewis, 1959: 24 (Belize). Hanson, 1961: 320 et seq. (breeding sites,

Panama). Johnson & Hertig, 1961: 765 et seq. (rearing data). Diaz-

Najera, 1963: 193 et seq. (Mexico). Ortiz & Scorza, 1963: 353 (r',

keyed). Disney, 1966: 449 (66 in rodent-baited traps, Belize). Hanson,

1968- 58 (larvae identical to those of gali-.doi).

Brwmptomyia h~mata: Barretto, 1955a; 187 (listed). Sherlock, 1962:

332 (cf. to mesai). Williams, 1970: 331 (listed, Belize). Fraiha et

al., 1970a: 468 (cf keyed). Forattini, 1971a: 98 (listed). Rutledge &

Mosser, 1972: 300 et seq. (breeding sites, ecology, Panama). Christensen,

1972a: 88 (listed). Forattini, 1973: 139 et seq. (gen. review, fig.).

Williams, 1976a: 603 (in caves, Belize).

Distribution: Mexino, Belize, Panama, Colombia (Choco), Ecuador.

Material examined: Colombia. 1 e, Teresita (Choco), hollow tree,

25 March 1967, D.G.Y. 1 ? same data but Malaise trap, 17 June 1967.

Panama. I 6 (holotype no. 112), type locality, 4 Dec. 1943. 4 S5, 5 o,

lab reared from larvae collected in Canal Zone, 1957-1958, W. Hanson.





-66-


Diecussio': Although the adults of B. hanata are rarely encountered,

the larvae may be quite common in .oil at the base of trees. Hanson

(1961) and Rutledge & Mosser (1972) in Panama found that the larvae of

this species far outnumber those of other phlebotomine species in this

microhabitat. Like other Brz.'ptoria spp., the fourth instar larva has

only two caudal setae.



11. Bru3nptoy jia leopoldoi
(Fig. 11)


PhZebonomus leopoldoi Rodriguez, 1953b: 52 (T holotype, Quevedo,

Los Rios Prov., Ecuador; y, Naranjal, Guayas Prov., Ecuador). Ortiz,

1963: 320 (7 keyed).

Br-imptomyia leopoldoi: Barretto, 1955a: 187 (listed). Sherlock,

1962: 332 (cf. to mesai). Williams, 1970: 331 (Belize). Fraiha et al.,

1970a: 468-469 (d keyed, as possible synonym of guirmaraesi). Forattini,

1971a: 98 (listed). Christensen, 1972a: 88 (listed, Panama). Forattini,

1973: 530 (as synonym of guimaraesi). Williams, 1976a: 603 (in caves,

Belize).

Distribution: Belize, Panamna, Colombia (Antioquia, Choco, Valle),

Ecuador .

Material examined: Colombia. 3 So', Rio Anori (Antioquia), light

traps, May 1970, C.H.P. 1 d, Curiche (Choco), light trap, 26 May 1967,

D.G.Y. 2 od, 5 ??, ca. 10 km W of Call near Pichinde (Valle), rock

crevice, tree trunks and light trap, 31 July 1973-4,Aug. 1973, D.G.Y. &

R.C.W. Ecuador. I &T (paratype no. 4314), Quevedo (Los Rios), no date,

J. Rodriguez. 2 (d&, 2 oo, 17 km E of Santo Domingo de Los Colorados

(Pichincha), mammal burrow, 4 May 1976, D.G.Y. & T. Rogers. 1 o3, same





-67-


data but 27 May 1976. 1 ', same data but light trap, 27 May 1976.

Panama. 1 .. (paratype no. 4292), Almirante (Boras del Toro), animal

burrow, 22 Jan. 1953, W. Hils. 1 d6, same data but tree buttress, 27 Jan.

1956, R. Hartmanu.

Disczusion: B. leopoldoi, considered to be conspecific with

B. guimaraesi (Coutinho and Barretto) by Forattini (1973), is specifically

distinct based upon the following considerations:

1. The nature and size of the basal coxite tufts and the structure

and setation of the parameres differ markedly between the two

males (cf. Fig. 11C and 111). The original figure of leopoldoi

(Rodriguez, 1953b) shows a rather broad paramere but I believe

this is a distortion due to mounting technique, not.a reflection

of normal structure. Paratype no. 4314 is obviously flattened.

2. The genital pump of the guimaraesvi male is less flared and

notably shorter (0.12 mm-0.15 amm) than that of leopoldoi which

is 0.22-0.24 mm long (cf. Fig. 11G and 11H).

3. The isolated spine of the style of guimaraesi is inserted at

0.59-0.61 of the segment whereas that of Zeopoldoi is located

at 0.53.

In addition to the Zeopoldoi specimens listed above, I examined 4

guimaraesi males from southern Brazil: 2 d'd, Faz. Ribeirao de Baizo,

Patos, Minas Gerais; 1 d', Coqueiros, Sao Paulo; 1 d'reared from egg laid

by ? captured at the type locality (Itaporanga, Sao Paulo, Brazil). I

conclude that these allopatric species are distinct, B. gtzimaraesi

occurring only in southeastern Brazil as far as known.
























Figure 8

i3r.umpto-yia avellari male -- A. Genitalia.

Male: Puerto Boyaca, Boyaca Dept., Colombia



B.rwrptnoryia beaupertuyi male -- B. Head, C. Flagellomere II, D. Paramere,
E. Ning, F, Genital pump and filaments, G. Genitalia, H. Coxite tuft,
rame scale as Fig. 8A.

Male: Lara State, Venezuela




-69-





.. .....- :.. ^A . . ,,.
--. �'' . - " :\ ":-







� , 1 . .

\ V -1' 1-- J "-'',, ,
I: �i





i / l{ l '----" .. . -' ..... E ...."--

1 ^,t ' 5^ . . ---- 'S
sa \i F
0N

Vt
9


-: . '- ' " ' " '
iS




if




/-**^ 'T?^ ^-- p^
� �-- - .. -
.; -1 _-




/4 I1^ \ F .
C -







jN^^" ) V/*
61"
Hi: (At
























Figure 9

RrPirptomyia galindoi -- A. Male head, B. Male flagellomere II,
C. Female head, D. Female flagellomere II, E. Female cibarium and
pharynx, F. Genital pump, G. Genital pump and filaments, 11. Male
genitalia, 1. Paramere, J. Coxite tuft, K. Spermathecae, L. Body of
speinatheca, M. Female wing, N. Male wing, 0. Female cibarium.

Male: Teresita, Choco Dept., Colombia
Female: Same locality as male




-71-










U ' I iI
c%) r i i! ^




p n-4 B / rt .)

1/ c \ E f / :



4, (' / 1 - \
X I Zr- I I




\( 3 i \ i
N II






9i jl .-.L.


k/i H
iti nf�~ ~� \ ,I I
-F -ZZCi -So
I .-- V
NO' - N'
























Figure 10

Brumptomyia hamata -- A. Male head, B. Male flagellomere II, C. Female
head, D. Female flagellomere II, E. Female cibarium and pharynx,
F. Genital pump and filaments of male from Teresita, Choco Dept.,
Colombia, G. Aedeagus and Paramere, H. Male genitalia, I. Spermatheca,
the other not drawn, J. Body of spermatheca, K. Female wing, L. Male
wing; M. Female cibariurm.

Male: Barro Colorado Island, Panama Canal Zone (except Fig. 10F)
Female: Teresita. Choco Dept., Colombia




-73-







-, ,/ ,-, -_



:' I i
" i , 2
i :


I I, i, /i



r \;-. .Z 1;.:-;
s~ ~ i ffxL m ^f


Sic\





.".'-,. -, ..4'. -A,' \ '\ "\
a A


�i1




;iO


,.,I r -:, - .\.-- ....
A'
^'^^ */ / �� >^\-^^








rM





, .. . - _ ..L
s H /''' ^i r . v






,..\ C ^. f .- . .*. , - .: . ----: = . .- - _- :
L � " " - " ' '*

1"-^' >^"~ *
*'^v4 L--^ ^ f
























Figure 11

Brumptormiia leopoldoi male -- A. Head, B. Flagellomere II, C. Coxite
tuft, paramere and aedeagus of male from near Santo Domingo de Los
Colorados, Ecuador, D. Wing, E. Genital pump and filaments, F. Genitalia,
G. Genital pump of male from Fig. 11C locality.

Male: Rio Anori, Antioquia Dept., Colombia (except for Figs,
11C and G)

Brzuptomnyia guimaraesi male -- H. Genital pump, I. Coxite tuft, paramere
and aedeagus, same scale as 11C.

Male: Itaporanga, S5o Paulo State, Brazil




-75-













i si 'i rr- '' =



l IB Cr
44...,1 E Y - .
E "






(-, -._� '_.
F H ' .





"N 05c.:.
5�g -.c- CrQ5~ t to




-76-



Genus LutznroTia Franca, 3924


As mentioned earlier the classification of the Lutzomnyia sand flies

is based upon the scheme of Lewis et al. (1978). It should be emphasized

that this represents a flexible classification, subject to change as new

species, new character states, or unknown sexes are discovered.

It is hoped that the following keys and discussions will be helpful

to those interested in sand flies and disease in Colombia and elsewhere

in the neotropics.



Keys to Subgenera, Species Groups, and Ungrouped
Species of Lutzoyjia


In order to interpret some of the character states in these keys,

the user is urged to closely examine the illustrations corresponding to

the descriptive statements. Such relative terms as "strong," "small,"

"inflated," etc. are understood more easily with figures. This also holds

true for the male and female genitalia, both of which may be complex and

therefore difficult to describe by words alone.

To a certain extent, the Lutzoayia subgenera and equivalent species

groups are defined by the characters given here. It is important to

noce, however, that species in some groups, presently unknown in Colombia,

may not exactly "fit" into a subgenus or species group as characterized

by these keys. For example, L. nevesi (Damasceno and Arouck) belongs in

the verriuoarvm group although the male, unlike the others, lacks a coxite

setal tuft. This species is very common at Limoncocha, Napo Prov.,

Ecuador -- a locality not far from Putumayo Intendencia, Colombia.





-77-


As additicnal species are discovered in the Republic, these keys

obviously will have to be revised. For the present, they are intended

to be used with caution, applicable only to the Colombian fauna.



Males

1. Palp 5 very short, less than half length of third and less than twice

length of fourth. . . . . . subgenus Pccyhodopygus (Fig. 61-71) . . 2


Palp 5 subequal to or longer than half length of third and greater

than twice length of fourth. . . . . . . . . . . . . . .. . . . 4


2. (1) Style with 1 large terminal spine and 3 inconspicuous subapical

setae. Coxite with dorsal margin indeated near middle . . . . . .

. . . . . . . . . . . . . . . series squaniventris (Fig. 61)


Style with 2-6 major spines. Coxite not indented near middle . . . 3


3. (2) Style with 2 major spines, both terminal. Paramere simple. . .

. . . . . . . .........series arthuri, L. bispinosa (Fig. 63)


Style with 3-5 (sometimes 6) major spines. Paramere simple or

complex . . . . . . . . . . . . ... .series panamensis (Fig. 64-71)


4. (1) Lateral lobes markedly inflated . .vaspertilionis group (Fig. 29)


Lateral lobes not inflated. . . . . . . . . . . . . . . . ... ... 5


5. (4) Lateral lobes with spatulate setae at tips. . . . . . . . . . . .

. . . . . . . . . . . .. subgenus Evoxnromnia, series infraspinosa*



*Species of Evandirormia in this series have not been found as yet in
Colombia but it is probable that at least one species occurs in the
southeastern part of the Republic. The subgenus was reviewed by Young
and Arias (1977).




-78-



Lateral lobes without spatulate setae . . . . . . . . . . . . . 6


6. (5) Style deeply forked. Parameres with 2-4 modified apical setae as

shown . . . . . . . . . . . ... . subgenus Viannamnyia (Fig. 35-36)


Style simple, not forked. Parameres usually with simple setae but,

if modified, then different from above. . . . . . . . . . . . . . . 7


7. (6) Style with 2 major spines and 1 or 2 smaller accessory seta . . 8


Style with 3-6 major spines, smaller accessory setae present or not.9


8. (7) Paramere with a dorsal setiferous arm. Coxite without nondeciduous

Seae . . . . . . . . suibgenus Damcpfomyia, L. rosabati (Fig. 30)


Paramere simple, without a dorsal arm. Coxite with a basal tuft or

group of setae . ... . . .verrucarum group, series serrana (Fig. 22)


9. (7) Style with 3 major spines . . . . . . . . . . . . . . . . .. 10


Style with 4-6 major spines . . . . . . . . . . . . . . . . . .. 11


10. (9) Coxite with a basal tuft of simple and modified setae. Style with

a small subterminal and a small median seta in addition to the major

spines. . . . . . . . . . . . . ... . subgenus Pressatia (Fig. 32-34)


Coxite without a basal tuft but with a distal group of 15+ simple

setae. Style with only a small median seta and 3 major spines, no

subterminal seta. . . . . . .... .. pilosa group, L. pitosa (Fig. 86)


11. (9) Style with 4 major spines . . . . . . . . . . . . . . .... . 12


Style with 5, sometimes 6, major spines .. . . . . . . . ... . 32

*Males of L. rorotaensis usually have 5 major spines but those examined
from Valle Dept., have but 4 (see p. 421).





-79-


12. (11) Antennal asc(,ids with very long pointed or short blunt basal

spurs. Style without a subterminal seta . . . . . . . . . . . . 13


Antennal ascoids with or without basal spurs, but if present they

are very short and pointed. Style with or without a subterminal

seta . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . 14


13. (12) Coxite with 5-8 strong distal setae. Style with isolated basal

spine . . . . . . . . . . . . . . ... ..deisbachi group (Fig. 50-51)


Coxite without distal setae. Style with basal spines more or less

paired . . . . . . . . . . . . . . ... shannoni group (Fig. 37-42)


14. (12) Palp 5 shorter than palp 3. Style without a subterminal

seta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15


Palp 5 as long as or longer than palp 3. Style with or without a

sub terminal seta. . . . . . . . . . . . . . . . . .. . . . . . . 18


15. (14) Coxite with 2 or more basal and/or median setae. . . . . ... 16


Coxite without nondeciduous setae . . . . . . . . . . . . . . .. 17


16. (15) Style with basal spine isolated. Antennal ascoids with very

short pointed basal spurs. Flagellomere I shorter than flagellomeres

II + III. . . . . . . . ... .. subgenus Trichophoro,,yia (Fig. 51-54)


Style with paired basal spines. Antennal ascoids simple, no basal

spurs visible. Flagellomere I longer than II + III . . . . . . . .

. . . . . . . .subgenus yssomnia (in part), L. antunesi (Fig. 55)





-80--


17. (15) Antennal ascoids with short, pointed basal suprs. Paramere with

e dorsobasal hump. Flagellomere I shorter than II + III . . . . .

. . . . . . . . . . . . . . . . . . . . . a'~aao group (Fig. 45-49)


Antennal ascoids simple. Paramere lacking a dorsobasal hump. Flagel-

lomere I longer than II + I I . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . subgenus Pihssomyia (in part), (Fig. 55-60)


18. (,4) Coxite with 3+ basal and/or median setae in a tuft or group,

d-sscal setae may also be present. . . . . . . . . . . .. . . . .19


Coxi.te without a basal and/or redian group of setae but long ventral

setae may be present. . . . . . . . . . . . . . . . . . . . . . . Q2


19. (18) Style with 4 short stubby spines, 2 of which are terminal; sub-

terminal seta present. Paramere simple, undivided. Coxite with 4-8

strong distal setae . . . . . ... .baiti group, L. baityi (Fig. 34)


Style with longer spines; if 2 are terminal then parameres are

divided; subterminal seta present or absent. Coxite without distal

nondeciduous setae. . . . . . . . . . . . . .. . . . . . . ... .20


20. Hind femur with a row of 3-5 short spines . . . . . . . . . . . . .

. . . . . . . ....... subgenus Pintomyia, L. spinosa (Fig. 31)


Hind femur without spines . . . . . . . . . . . . . . . .... . . .21


21. (20) Paramere with 2 dorsobasal setae, much longer than others,

hooked or fan-shaped. . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . .subgenus Lutzc'.;Pjia, series long palpis (Fig. 13-15)


Paramere without such setae . . . . . . . ..... . . . . . . . 22





-8L-



22. (21) Style with a subterminal seta . ... .. . . . ..... .. . 23


Style without a subterminal seta. . . . . . . . . . . . . . . ... 26


23. (22) Genital filament tips enlarged, spoon-shaped, each with a dis-

tinct inner "tooth" . . . migonei group, series waZkeri (Fig. 19-20)


Genital filament tips enlarged or not but lacking an inner tooth. 24


24. (23) Coxite with setae of basal tuft shorter than width of coxite

. . .. . . . . . . . . . . .migonei group, L. migonei (Fig. 18)


Coxite with setae of basal tuft longer than width of coxite . . . 25


25. Mesonotum & pleura pale . . sautensis group,* L. sauZensis (Fig. 21)


Mesonotum distinctly darker than pleura . . . . . . . . . . . . . .

. . . . verruoarwu group, series verrucarrum (in part), (Fig. 23-24)


26. (22) Coxite with basal setae inserted on a raspberry-like, subcircular

base. . . ... . . subgenus Lutzomyia, series cruciata (Fig. 16-17)


Coxite with basal setae, if present, not implanted on suchabase. 27


27. (26) Style with paired basal spines. Coxite with basal tuft of

slender, subequal setae . . . . . . . . . . . . . . . .... ... ver-

rucaruml group, series verruCarum (in part), L. nuneztovari (Fig. 23)


Style with basal spine isolated. Coxite with either basal-median

setae, 1 at least larger than others, or with a patch of slender distal

setae . . . .. c.aycnnencis group, series atroclavata (Fig. 81-82)


*I am unable to separate the males on the group level using structural
characters.





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28. (18) Style uth a small subterminal seta . . . . . . . . . . . . 29


Style without a small subterminal seta. . . . . . . . . . . . . . 31


29. (28) Paramere divided. Coxite with long ventral setae . . . . . .

. . . . . . . . . . . . . ...... .on ispira group (Fig. 43-44)


Paramere simple, undivided. Coxite without long ventral setae. . 30


30. (29) Palp 5 shorter than segments 3 + 4. Eyes large. Paramere

slender, its width less than that of style . . . . . . . . . . . .

. . . . . . . . . . . . .. ungrouped, L. sp. of Anchicaya (Fig. 89)


Palp 5 longer than segments 3 + 4. Eyes small. Paramere broad, its

width greater than that of style. ungrouped, L. rangeliana (Fig. 88)


31. (28) Style with isolated basal spine. Flagellonere I longer than

head height; ascoids with very short basal spurs. Palp 5 shorter

than segments 3 + 4. . ...... . ungrouped, L. nordestina (Fig. 87)


Style with paired basal spines. Flagellomere I shorter than head

height; ascoids without visible basal spurs. Palp 5 longer than seg-

ments 3 + 4 . . cayennensis group, series cayennensis (Fig. 79-80)


32. (11) Style with a small subterminal seta . . . . . . . . . . . . .

. . . . . . . . . . . . . oswaldoi group (in part), L. pia (Fig. 83)


Style without a small subterminal seta. . . . . . . . . . . ... . 33


33. (32) Coxite with 2 or more setae at base; others may extend to middle

of structure forming a loose or compact tuft . . . . . . . . . . .

. . . . . . . . . . .. vxator group, series periuensis (Fig. 72-78)





-8--



Coxite without setae at base but with a few scattered setae near

middlt.. of coxite. . ..... . oaswudoi group (in part), (Fig. 84-85)



Females

1. Spermathecae associated with paired sclerotized structures, intracel-

lular ducts (= "hairs") covering most of spermathecae . . . . . . .

. . . . . . . . . . . . . . . . .. subgenus Vin(vacuryia (Fig. 35-36)


Spernathecae without associated sclerotized structures, intracellular

ducts confined to small area, usually but not always, to terminal

knob. .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2


2. (1)Pharynx with posterior spines . . . . .... .. . . . . . . .. 3


Pharynx without spines. . . . . . . . . ... .. . . . . . . . .. 6


3. (2) Cibarium with inner pair of horizontal teeth pointing inwards

. . . . . . .. oswadoi group (in part), L. trinidadensis (Fig. 85)


Cibarium with horizontal teeth pointing toward pharynx, not slanted

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4


4. (3) Cibarium with 10+ horizontal teeth in a comb-like row . . cayen-

nsis group, series caennensis (in part),L. cayennensis (Fig. 79)


Cibarium with 4 horizontal teeth not in a comb-like row ..... 5


5. (4) Foreleg with femur longer than tibia. Pharynx markedly enlarged

with prominent, subequally spaced transverse ridges and conspicuous

spines. Pleura pale . . . . . . . . . . . . . . . . . . . . . ...

....... . cayennen;i: group, series atroclavata (Fig. 81-82)





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Foreleg with femur shorter than tibia. kharynx nore slender, the

transverse ridges and spines less defined. Pleura dark . . . . . . .

cylenenosis group, series Cayenrnensis (in part), L. mic-opyga (Fig. 80)


6. (2) Spermathecae with bubble-like expansions. . . . . . . . . . . . 7


Spermathecae without bubble-like expansions . . . . . . . . . . . . 8


7. (6) Cibariumn with 4 broad and blunt horizontal teeth and conspicuous

lateral teeth. Mesonotum well pigmented, darker than pleura. Eyes

very small. . . . . . .. .subgenus Da..ipfomyia, L. rosabati (Fig. 30)


Cibarium with 4 slender, sharp horizontal teeth, without obvious

lateral teeth. Mesonotum and pleura pale. Eyes larger . . . . . .

. . . . . . . . . . . . . saulensis group, L. saulensis (Fig. 21)


8. (6) Antennal ascoids with long pointed or short blunt basal spurs . 9


Antennal ascoids simple or with short pointed basal spurs ... . .10


9. (8) Cibarium with 10+ horizontal teeth. Spermathecae distinctly

anndlated . . . . . . . . . . . . . . . .deisbachi group (Fig. 50-51)


Cibarium with 4-8 horizontal teeth. Spermathecae annulated or not

. . . . . . . . . . . . . . . . . . . .. . hannoni group (Fig. 37-42)


10. (8) Hind femur with row of 3-5 short spines . . . . . . . . . . . . .

. . . . . . . . . . . . . . .subgenus Pitomryia, L. spinosa (Fig. 31)


Ilind femur without spines . . . . . . . . . . . . . . . .... ... 11


11. (10) Cibarium with 6 or more horizontal teeth . . . . . .... . . .12





-85-



Cibarium with 4 horizontal teeth. . . . . . . . . . . .... . . 17


12. (1') Individual sperm ducts markedly convoluted, twisted . . . . .

. . . . . aagaoi group, series. brasiliensis, T. runoides (Fig. 49)


Individual sperm ducts not convoluted as above. . . . . . . ... . 13


13. (12) Palp 5 equal to or shorter than segments 3 + 4 . . . . . .. 14


Palp 5 longer than segments 3 + 4 . . . . . .... . . . . . . .. 16


14. (13) Spermathecae larger, sac-like, without annuli. Ascoids with

short pointed basal spurs. aragaoi group, series aragaoi (Fig. 45-48)


Spermathecae smaller, with incomplete or complete annuli. Ascoids

with or without short basal spurs . ... . . . . . . . . . . .. 15


15. (14) Flagellomere I longer than flagellomeres II + III. Ascoids with

short basal spurs . ..... .subgenus Trichophoromyia (Fig. 51-54)


Flagellomere I shorter than II + III. Ascoids without visible basal

spurs . . . . . . ... . ... .. ..subgenus Nyssomyia (Fig. 55-60)


16. (13) Cibarium with 6 horizontal teeth. Spermatheca with terminal

segment smaller than preceding segment. Common sperm duct longer

than individual ducts . .oswaldoi group (in part), L. pia* (Fig. 83)


Cibarium with 8+ horizontal teeth. Spermatheca with terminal segment

as large as others. Common sperm duct much shorter than individual





*L. pia is keyed twice because of intraspecific variation in the number
of horizontal teeth (see p. 4L9).





-85-


du ts . . . . . . . . . . . . . . . . . . . . . . . . . . subge-

nULsL-tzcomyia, series longipal-pis (in part), L. Longipalpis (Fig. 15)


17. (11) Palp 5 shorter than segments 3 + 4 . . . . . . . . . . 18


Palp 5 longer than segments 3 + 4 . . . . . . . . . . . . 21


18. (17) Spermathecae imbricated, annuli semi-telescoped. Individual

sperm ducts heavily sclerotized, wrinkled, or rugose in part or

whole . . . . . . . . . .... .subgenus Psyclhodopygua (Fig. 61-71)


Spermathecae not imbricated, annuli, if present, not semi-telescoped.

Individual sperm ducts non-rugose, smooth walled or with slight

thickenings . . . . . . . ..... . . . . . . . . . . . . . . . . 19


19. (18) Cibarium with inner pair of horizontal teeth pointing inwards.

Flagellomere I extremely long, subequal to or longer than combined

lengths of palpal segments. Ascoids with very short basal spurs.

. . . . . . . . . . . . . . . . ungrouped, L. nordestina (Fig. 87)


Cibarium with horizontal teeth pointing toward pharynx, not slanted

inwards. Flagellomere I shorter than palpus. Ascoids without visible

basal spurs . . . . . . . . . . . . . . . . . . . . . . . . . 20


20. (19) Spermathecae as shown, longer than individual sperm ducts.

Abdominal tergite 8 without lateral seta . . . . . . . . . . . . .

. . . . . . . .vespertilionis group, series vespertilionis (Fig. 29)


Spermathecae otherwise, shorter than individual sperm ducts. Ab-

dominal tergite 8 with lateral setae. . ........ .. . . 21





--87-


21. (20) Spermathecae less than twice as long as wide, terminal segment

smaller than preceding segment. Common sperm duct longer than in-

dividual ducts. .... . oswaltdoi group (in part), L. pia (Fig. 83)


Sperniathecae cylindrical, its length at least 3 times width, terminal

segment subequal to or smaller than others. Common sperm duct

shorter than individual ducts . . . . . . . . . . . . . . . . . . .

. . v exator group, seriesperuensis (in part), (Fig. 72-74; 76-78)


22. (17) Individual sperm ducts heavily pigmented in part or whole. . .

. . . . . . . . . . . . . . . . . . .subgenus Pressatia (Fig. 32-34)


Individual sperm ducts not pigmcnted. . . . . . . . . . . . . . . 23


23. (22) Cibarial arch absent or conspicuous only at sides. . ... . .24


Cibarial arch complete or nearly so . . . . . . . . . .. . .... . 26


24. (23) Cibarium with horizontal teeth like flattened plates viewed

almost edgewise. Common sperm duct absent . . . . . . . . . . . .

. . . . . . . . . . . . . . . .. pilosz group, L. pitosa (Fig. 86)


Cibarium with horizontal teeth otherwise. Common sperm duct

present . . . . . . . . . . . . . . . .... . . . . . . . . . ... 25


25. (24) Wing venation with beta less than half aZpha. Cibarium with

horizontal teeth pointing towards pharynx, not inwardly slanted .

. . . ..... vexator group, series peruensis (in part), (Fig. 75)


Wing venation with beta over half length of alpha. Cibarium with

horizontal teeth pointing inwards . . . . . . . . . . . . . . . . .

. . . . . . . . osaldoi group (in part), L. rorotaenais (Fig. 100)




Full Text

PAGE 1

A BIOSYSTEMATIC REVIEW GF THE BLOODSUCKING FSYCHODID FLIES OF COLOMBIA (DIPTERA: PHLEBOTOMIHAE AND SYCORACTNAE) By 'ID GRIER YOUNC 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 1077

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Dedicated to my father, Howard G. Young

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TABLE OF CONTENTS Page LIST OF FIGURES viii ABSTRACT xii INTRODUCTION 1 CLASSIFICATION AND TAXONOMIC TREATMENT A Keys to the Subfamilies of New World Psychodidae 6 GENERAL MORPHOLOGY AND TERMINOLOGY 8 METHODS AND MATERIALS 15 Field Collections 15 Laboratory Methods 17 DISTRIBUTION, COLLECTING LOCALITIES, AND CHARACTERISTICS OF THE FAUNA 19 Pacific Coast Region 20 Atlantic Coast Region 23 Andean Region 24 Oriental Region 25 MEDICAL IMPORTANCE 29 SUBFAMILY SYCORACINAE 31 Keys to the Sycorax Species 33 1. Sycorax coidieola n. sp 35 2. Sycorax colombiensis n. sp 36 3. Sycorax fairchildi n. sp 3rf 4. Sycorax trispinosa n. sp 30 SUBFAMILY PHLEBOTOMINAE 50 Key to the New World Genera 50 Genua Wariloya Hertig, 1948 52 Keys to Species 52 5. WariZeya (W.J nigrosacculus 53 6. Wan'.teya (W.) rotundipennis 5:

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Page Genus Brumptomyia Franga & Parrot, 1921 60 Key to Species 60 7. Brumptomyia av?.llavi 61 8. Brumptomyia beaupevtuyi 62 9. Brumptomyia galindoi 63 10. Brumptomyia hanata 65 11. Brumptomyia leopoldoi 66 Genus Lutzomyia Franca, 1924 76 Keys to Subgenera, Species Groups, and Ungrouped Species. . . 76 Subgenus Lutzomyia Franca, 1924 9.1 Keys to Species 91 12. Lutzomyia (L.) bifoliata 93 13. Lutzomyia (L.) lichyi 94 14. Lutzomyia (L.) longipalpis 96 15. Lutzomyia (L.) gomezi 97 16. Lutzomyia (L.) rnarinkellci n. sp 99 Migonei Group Theodor, l96i> Ii2 Keys to Species 112 17. Lutzomyia migonei 113 18. Lutzomyia marajoensis 114 19. Lutzomyia sp. de Baduel 116 20. Lutzomyia waVkevl 116 Saulsnvis Group Lewis et al., 1978 125 21. Lutzomyia saulensia 125 Vervucarum Group Theodor, 1965 130 Key to Species 130 22. Lutzomyia odax 133 23. Lutzomyia sevvona 134 24. Lutzomyia andina 135 25. Lutzomyia columbiana 136 26. Lutzomyia disiuncta 137 27. Lutzomyia evansi 137 28. Lutzorrpjia lonciflocosa 138 29. Lutzomyia movalesi n. sp 139 30. Lutzomyia nuneztovari 141 31. Lutzomyia ovaltesi 143 32. Lutzomyia quasitownsendi 14 5 33. Lutzomyia sauvoida 145 34. Lntzovvyia spinicrassa 146 Vbspertitionis Group Theodor, 1965 161 Key to Species 161

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Page 35. Lutzomyia isovesperti.li.onts 161 35. Lutzomyia vespertilionis 162 Subgenus Damp forty ia Addis, 1945 167 37. Lutzomyia (D.) rosdbali 167 Subgenus Pintomyia Costa Lima, 1932 171 38. Lutzomyia (P.) spinosa 171 Subgenus Pressatia Mangabeira, 1942 176 Keys to Species 176 39. Lutzomyia (Pp.) oamposi 177 40. Lutzomyia (Pr.) dysponeta 178 41. Lutzomyia (Pr.) triaoantha 180 Baityi Group Theodor, 1965 186 42. Lutzomyia baityi 180 Subgenus Viannamyia. Mangabeira, 1941 190 Keys to Species 190 43. Lutzomyia (V.) caprir/x 19.1 44. Lutzomyia (V.) furcata 192 45. Lutzomyia (V.) tuberculata 194 Shannoni Group Theodor, 1965 200 Keys to Species 200 46. Lutzomyia abonnenci 202 47. Lutzomyia dasymera 203 48. Lutzomyia dendrophyla 204 49. Lutzomyia punctigenioulata 206 50. Lutzomyia shannoni 208 51. Lutzomyia. undulata 210 Longispina Group Theodor, 1965 224 Keys to Species 224 52. Lutzomyia longispina 225 53. Lutzomyia triramula 226 Aragaoi Group Theodor, 1965 232 Keys to Species 232 54. Lutzomyia aragaoi 234 55. Lutzomyia barret toi b arret boi 236 56. Lutzomyia barrettoi najuscula n. ssp 237 57. Lutzomyia eavpenteri 241 58. Lutzomyia. runoides 243

PAGE 6

Page Drcisbachi Group Lewis et al . , 1978 255 Key to Species 25 5 5 C 2. Lutzomyia aolydifera 255 60. Lutzomyia drsisbaoki 257 Subgenus Triahopkororrryia Barretto, 1962 261 Key to Species 261 61. Lutzomyia (T.) auvaensis 262 62. Lutzomyia (T.) cellulana n. sp 264 63. Lutzomyia (T.) kouardi n. sp 266 64. Lutzomyia. (?.) rebur-va 267 65. Lutzomyia (T.) saltu^sa n. sp 269 66. IjUtzomyia (T.) ubiquitalis 270 Subgenus Nyssomyia Barretto, 1962 280 Keys to Species 280 67. Lutzomyia (II.) antunesi 283 68. Lutzomyia. ([.'. ) flavisscuLallata 285 69. Tjutzomyia (II.) olneoa bioolov 287 70. Lutzomyia (H.) brapidoi 289 71. Lutzomyia (!!.) urnbratilis 292 72. Lutzomyia (N.) ylephiletor 295 73. Lutzomyia (N.) yuilli 297 Subgenus Psycho dopy gus Mangabeira, 1941 311 Keys to Species 312 74. Lutzomyia (P.) bernalei 317 75. Lutzomyia (P.) chagasi 317 76. Lutzomyia (P.) fairtigi 318 77. Lutzomyia (P.) bispinosa 319 78. Lutzomyia (P.) ayrozzi 320 79. Lutzomyia. (P.) oar-rerai oewevai 323 80. Lutzomyia (P.) oarrerai thula n. ssp 326 81 . Lutzomyia. (P. ) davisi 331 82. Lutzomyia (P.) guyanensis 333 83. Lutzomyia (P.) hirsuta 335 84. Lutzomyia (P.) nooticola 338 85. Lutzomyia (P.) panarr.ensis 339 86. Lutzomyia (P.) vecuvva 341 87. Lutzomyia (P.) sp. of Tres Esquinas 342 VexatoT Group Theodor, 1965 366 Keys to Species 36 7 88. Lutzomyia cir-rita 369 89. Lutzomyia harbmar.ni 370 90. Lutzomyia. oso/noi 37.1 9.1. Lutzomyia sar.guinar-ia 372

PAGE 7

Page 92. Lutzomyia scorzai 374 93. Lutzomyia strictivilla 378 94. Lutzonyia sp. of Pichinde. 380 Cayennensis Group Theodor, 1965 398 Keys to Species 398 95. Lutzomyia cayennensis 399 96. Lutzomyia micropyga 401 9 7. Lutzomyia atroolavata 405 98. Lutzomyia venezuelensis 407 Oswaldci Group Theodor, 1965 417 Keys to Species 417 99. Lutzomyia via 418 100. Lutzorwyia vovotaensis 420 101. Lutzomyia trinidadensis 422 LUL>u;>a utuu[j jii£uuCl, irOJ h.iZ 102. Lutzomyia pilosa 432 Ungrouped Species 4 37 103. Lutzomyia nordestina 4 37 104. Lutzomyia vangeliana 4 39 105. Lutzomyia sp. of Anchicaya , 440 SYNOPSIS OF NEW TAXA, SYNONYMS, FEMALE DESCRIPTIONS, AND NEW DISTRIBUTION RECORDS 448 REFERENCES 451 BIOGRAPHICAL SKETCH 481

PAGE 8

LIST OF FIGURES Figure Page 1. Map of Colombia showing localities where phlebotomine and sycoracine flies have been collected 23 2. Syoo/'ox andicola 43 3. Syaorax colombiensis 45 4. Sycorax faivchildi 4 7 5. Syoorax trispinosa 49 6. V:av it °-ya nigvo^or-riylvs -,...,,<.-.,....,,., 57 7. Wavileya rotundipennis 59 8. Bmonptomyia avellai-i and Brumptomyia beaupertuyi 69 9. Brumptomyia galindoi 71 10. Erumptomyia hamata 73 11 . Bv.mptomyia leopoldoi and Brvrrotomyia guimavaesi 75 12. Lutzomyia (P.) hirsuta niearacuensis and Lutzomyia tounsendi 90 13. Lutzomyia (L.) bifoliaia 103 14. Lutzomyia (L.) liohyi 105 15. Lutzomyia (L.) longivalpis 107 16. Lutzomyia (L.) gomezi 109 17. Lutzomyia (L.) mavinkellei Ill 18. Lutzomyia migonei and Lutzomyia sp. de Baduel 120 19. Lutzomyia marajoensis 122 20. Lutzomyia walkeri 124 21. Lutzomyia saulensis 129

PAGE 9

Fi gur e Page 22. Lutzomyia serrana 148 23. Lutzomyia andina, Lutzomyia disiuneta, and Lutzomyia nuneztovavi 150 24. Lutzomyia aolurribiana 152 25. Lutzomyia evansi 154 26. Lutzomyia moralesi 156 27. Lutzomyia ova'llesi 158 28. Lutzomyia sauvoida, Lutzomyia quasi towns endi > Lutzomyia. tongiflocosa } Lutzomyia spiniopassa, and Lutzonyia odax. . . 160 29. Lutzomyia vesper til ionis and Lutzomyia isovespsrtilionis . . 166 30. Lutzopvyia (D.i rosaboli 170 31. Lutzomy%a (Pi.) spinosa 175 32. Lutzomyia (Pp.) eamposi 183 33. Lutzomyia (Pp.) dysponeta 185 34. Lutzomyia (Pp.) tpicantha and Lutzomyia ha.ityi 189 35. Lutzomyia (V.) cappina and Lutzomyia (V. ) furaata 197 36. Lutzomyia (V.) tubepoulata 199 37. Lutzomyia abonnenei 213 38. Lutzomyia dasymeva 215 39. Lutzomyia dendpophyla 217 40. Lutzomyia punetigeniculata 219 41. Lutzomyia shannoni 221 42. Lutzomyia undulata 223 43. Lutzomyia longispina 229 44. Lutzomyia triramula 231 45. Lutzomyia avagaoi 246 46. Lutzomyia barvettoi majusoula 248

PAGE 10

Figure Page. 47. Lutzormjia barrettoi maQusoula 3 Lutzormjia barret toi barrettoi, and Lutzorrryia texana 250 43. Lutzormjia cavpentevi 252 49. Lutzormjia vunoides 254 50. Lutzormjia aelydifeva 260 51. Lutzormjia dreisbachi, Lutzormjia (T.) auraensisj and Lutzormjia ubiquitaiis 273 52. Lutzormjia (T.) cetlulana 275 53. Lutzormjia (T.) howardi and Lutzormjia (T.) sdltuosa 277 54. Lutzormjia (T.) reburra 279 55. Lutzormjia (f-1.) antunesi and Lutzormjia (N.) flaviscutellata. 300 56. Lutzormjia (N.) olmeca bioolor 302 57. Lutzormjia (N.) trapidoi 304 58. Lutzorrpjia (11.) umbratitis 306 59. Lutzormjia (N.) ylephiletor 308 60. Lutzormjia (N.) yuilli 310 61. Lutzonrjia (P.) bevnalei, Lutzormjia (P.) ch.aga.si, and Lutzormjia sp. of Tres Esquinas 345 62. Lutzormjia (P.) fairtigi 347 63. Lutzomjia (P.) bispinosa 349 64. Lutzormyia (P.) ayvozai 351 65. Lutzormjia, (P.) cawevai carrerai 353 66. Lutzormjia (P.) cavrerai thula 355 67. Lutzormjia (P.) davisi and Lutzormjia (P.) hirsuta hirsute/.. . 357 68. lutzormjia (P.) guyanensis 359 69. Lutzormjia (P.) nooticola 361 70. Lutzorrpjia (P.) panamensis 363 71. Lutzormjia (P.) vecurva 365

PAGE 11

Figure Page 72. Lutzomyia air-vita 385 73. Lutzomyia kca>tmanni 387 74. Lutzomyia osornoi 389 75. Lutzomyia sanguinaria 391 76. Lutzomyia strictivilla 393 77. Lutzomyia soorzai 395 78. Lutzomyia sp . of Pichinde 39 7 79. Lutzomyia cayerm&nsis 410 80. Lutzomyia micropyga 412 81. Lutzomyia atroolavata 414 82. Lutzomyia venezuetensis 416 83. Lutzomyia pia 427 84. Lutzomyia r-orotaensis 429 85. Lutzomyia tvinidadensis 431 86. Lutzomyia, pilosa 4 36 87. Lutzomyia nordestina 443 88. Lutzomyia rangeliana 445 89. Lutzomyia sp . of Anchicaya 447

PAGE 12

Abstract of Dissertation Presented to the Graduate Council if cite University of Florida in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy A BIOSYSTFi'tATIC REVIEW OF THE BLOODSUCKING PSYCHCDID FLIES OF COLOMBIA (DJ.PTERA: FHLFBOTOMINAE AND SYCORACINAR) By David Crier Young December, 1.977 Chairman: Graham B. Fairchild Major Departmenc: Entomology and Nernatology The psychodid subfamily Sycoracinae, previously unknown in Colombia, is represented by four new Sycovcx species described and illustrated in this review. Phlebotorainaa, a much larger subfamily containing vectors of human pathogens, is represented in the Republic by 101 species and subspecies — a number equivalent to one-third of the described New World taxa. Ten of these in the genus Lutzomyia. are described as new and are given formal or informal names. The previously urknown females of Lutzomyia pilosa, L. (P.) hi-vsuta ni ccwaguensis ?nd L. bifoliata are described. Nineteen phlebotcmin.e species are reported in Colombia for the first time. Other new distributional records include those from Ecuador (22 spp.), Panama (3 spp.), Honduras (2 spp.), Costa Rica, Cuba, Guatemala, and Paraguay (1 sp. each). In order to clarify the status of some forms, an attempt .-/as made to study specimens from as many Colombian and non-Colombian localities as possible. To this end, over 20,000 adults in both subfamilies were examined from 1967 to the present.

PAGE 13

An extensive bibliography and synonomy is given for each previously described taxon. Actual and/or probable misidentif ications in the literature are discussed. identification keys, previously unavailable for the Colombian fauna, are provided along with numerous illustrations, most of ivhich are original. Although more information is needed, it is suggested that recent speciation in the neotropical phlebotominae was due to climate changes in the Pleistocene (refuge theory) .

PAGE 14

INTRODUCTION Prior to the last decade, we knew very little about the phlebotomine sand fly* fauna of Colombia. Osorno et al . (1967) in reviewing previous studies by others and adding records of their own, reported less than 25 species in the Republic. From 1967 to 1972, the late Dr. E. Osorno-Mesa and his colleagues at Instituto Nacional de Salud, Bogota added more records, bringing the total to 77 species and subspecies (Osorno et al., In this review 101 species and subspecies of Phlebotominae are reported from Colombia. This represents about one-third of the described New World species but I estimate that at least 40 additional species will be discovered in the Republic, especially in the little-collected southeastern region. The small subfamily Sycoracinae, whose members resemble those of Phlebotominae in being able to take vertebrate blood, is treated here for this reason. In order to clarify the status of some taxa and to study intraspecific variation and distributional patterns, an effort was made to examine specimens from as many Colombian and non-Colombian localities as possible. To this end, I am vary grateful to the following persons for providing specimens, information or both: Dr. Jorge R. Arias, Instituto Nacional de Pesquisas da Amazonia (INPA) , Manaus ; Drs . Stephen C. Ayala and Pabln *I prefer to separate "sand" from "fly" in accordance with the suggestion of Borrer, Delong, and Triplehorn (1976) .

PAGE 15

Barretto, Universidad del Valle, Cali' Drs. Richard D. Ward and Habib Fraiha, Institute Evandro Chagas , Belem; Dr. David J. Lewis, British Museum (Nat. Hist.); Dr. Cornells J. Marinkelle, formerly Universidad de Los Andes, Bogota; Dr. AlberLo Morales-Alarcon and Che lace Dr. Ernesto Osorno-Mesa, Instituto Nacional de Salud, Bogota; Prof. Almilcar Vianna Martins, Universidade Federal de Minas Gerais; Dr. Nelson Papavero, Museu de Zoologia, Universidade de Sao Paulo; Drs. Charles H. Porter and Thomas M. Yuill, University of Wisconsin; Dr. Alan Stone, formerly Systematic Entomology Laboratory (ARS) ; Mr. Richard C. Wilkerson, University of Florida. Dr. Thomas H.G. Aitken, Yale University School of Medicine; Dr. Elisha S. Tikasingh, Caribbean Epidemiology Center (CAREC) , Trinidad; Dr. Harold Trapido, Louisiana State University Medical Center; Dr. Jorge E. Velasco, La Paz, Bolivia; Dr. Derek A. Duckhouse, University of Adelaide, South Australia; Maj . John F. Reinert, U.S. Army Medical Research and Development Command, Washington, D.C. Others who have supported this study and to whom I am indebted include members of the Atlantic-Pacific Interoceanic Canal Study Commission, Canal Zone from 1967-1968. Col. Bruce F. Eldridge of this organizaCion was especially helpful in developing field programs and in providing data on Choco collections. Financial support from U.S. Army Medical Research and Development Command under contract no. DADA 17-72-C-2139 is gratefully acknowledged . Such support was provided also at various times by Drs. William G. Eden and Hugh Popenoe, University of Florida. Members of my graduate school committee at the University of Florida have been helpful in many ways, too numerous to mention here. For their help, T thank Drs. Franklin S. Blanton, Jerry F. Butler, Graham B.

PAGE 16

Fairchild, Dale H. Habeck, and Stephen G. Zam. Miss Stephanie Haney ably slide mounted thousands of specimens from 1972 to 1977. For typing the manuscript, I wish to thank Mrs. Adele Koehler. Dr. G.B. Fairchild deserves special recognition for his constant encouragement, stimulating discussions and expert assistance throughout the study, I ai., deeply indebted to him. Particular appreciation is expressed also to my wife Molly who was most understanding during the preparation of this review. Holotypes and allotypes of new taxa are to be deposited in the U.S. National Museum (Nat. Hist.). Paratypes, when available, will be held by the following institutions: Instituto Nacional de Salud, Bogota; Florida State Collection of Arthropods, Gainesville; and British Museum (Nat. Hist.), iondon. The remaining 20,000+ specimens, upon which most of this review is based, are stored in the author's personal collection. A synoptic collection will be given to the Instituto Nacional de Salud, Bogota.

PAGE 17

CLASSIFICATION AND TAXONOMIC TREATMENT Of the 6 recognized subfamilies of Psychodidae (Duckhouse, 1972, 1973), all but one, Horaiellinae from the Oriental Region, are represented in continental Colombia. The species of IloraieVla, Trtahomyia, and Suaorax and its allies are united by some in the subfamily Trichomyiinae but this appears to be artificial and does not "express the wide discontinuities and degrees of difference from other Psychodidae, especially in LIUi'»»^^.v. .,._^. t ,v,., v^^C OUS>C, A.JI*.;. The classification of Phlebotominae is interpreted differently according to author(s) . In a recent paper we reviewed most classificatory schemes and proposed one which is used here for the New World taxa (Lev/is et al . , 1978). It is essentially a modification of Theodor's 1965 classification. Three American genera are recognized — LutzonryLa Franca (ca. 290 spp.), Brumptomyia Franca and Parrot (ca. 23 spp.), Wariteya Hertig (5 spp.). The genus Hex'tigia Fairchild is considered to be a junior synonym of Waviteya by Lewis et al. (1978). The majority of the Lutzomyia spp. were described originally in the genus Phlebotomus (= Flebotomus) Rondani. Some authors continue to place them in this genus, ignoring the evidence to the contrary provided by Theodor (1948, 1965) and Christensen et al. (1971). The large genus Lutzomyia is divided into numerous subgenera, species groups, and series based upon adult morphology (Lewis et al., 197S) . The diversity of groups within the genus is attributed to "radiation or even 'explosive radiation' of some immigrating ancestral stocks in a newly

PAGE 18

colonized area having a wealth of ecological niches" (Theodor, 1965). This implies that these ancestral stocks originated in the Old World -a hypothesis which may never be proven. It would appear that recent speciation in some groups (e.g. subgenus TlH-chophovornyia) may have been due to climatic changes in the past, especially during the Pleistocene, which served to isolate conspecific populations in moist refugia during dry periods (Haffer, 1974). Other remarks on the classification of New World phlebotomines are given by Lewis et al. (1978) and in this review under each supraspecif ic taxon. An extensive bibliography and synonomy is given for most taxa treated here. I usually omitted references which contain species lists but some of them including those by Fairchild (1955), Barretto (1955a, 1962), Theodor (1965) , and Martins & Morales (1972) are very important because of discussions related to classification and distribution. For each species, I cite the original description as well as most others published since Barretto' s 1947 catalog. In cases where a species was reviewed subsequent to 1946 (e.g. Forattini, 1973), I refer to that publication for full references. Abbreviations of words used in the text which may not be familiar to some readers are given below: Inst i tutions USNM — United States National Museum (Nat. Hist.), Washington, D.C. INPES — Instituto Nacional para Programas Especiales de Salud, Bogota INPA — Instituto Nactional de Pesquisas de Amazonia, Manaus UV -Univcrsidad del Valle, Call

PAGE 19

GML -•Corgas Memorial Laboratory, Panama UF — University of Florida Collectors C.H.P. •— Charles H. Porter C.J.M. — Cornelius J. Marinkelle R.C.lv. — Richard C. Wilkerson D.G.Y. — David G. Young Ot her Abbreviations ca. — about cf. — compared to classif. -classification dsscr. — description (list. -distribution morphol. — morphology pop. — population Key to the Subfamilies of Psychodidae of the New World 1. Wing venation with 4 branched radius, P ? ,o fused into a single vein (Fig. 2B) 2 Wing venation with 5 branched radius (Fig. 12G) 3 2. Wing venation with short cubitus (cu) . Mandibles present Sycoracinae (Fig. 2) Wing venation with long cubitus. Mandibles absent. . . .Trichomyiinae

PAGE 20

-/3. Palpus of 4 segments, rarely 3. Eyes usually with eye bridge. Antennal f lagellomeres nodiform or barrel shaped Psychodinae Palpus of 5 segments. Eyes without eye bridge. Antennal f lagellomeres subcylindrical 4 4. Females with 1 spermatheca. Mandibles absent. Male aedeagus entire Bruchorayiinae Females with 2 spermathecae. Mandibles present. Males with bifid aedeagus. . . Phlebotominae (Fig. 12F)

PAGE 21

GENERAL MORPHOLOGY AND TERMINOLOGY This brief discussion focuses on the structures commonly used in taxonomic studies of phlebotomine sand flies. General accounts of adult anatomy include those by Fairchild and Hertig (1947a), Kirk & Lewis (1951), Abonnenc & Minter (1965), Davis (1967), Abonnenc (1972), Lewis (1973), and Forattini (1973). The terminology varies according to author; that used here applies also to the sycoracine flies except where otherwise stated. Although little studied, the immature stages are probably no less important in systematics; they are not considered here owing to lack of material. For information on the anatomy of these stages see Barretto (1941), Hanson (1968), Abonnenc (1972), Carnheiro & Sherlock (1964), Ward (1977), and Zimmerman et al. (1977). Descriptions of immatures of those species occurring in Colombia are cited in the appropriate species bibliography. The sand fly head and its appendages (Fig. 12A) are useful in associating conspecific males and females and for grouping taxa below the genus level. Head height, measured from the vertex to the tip of the clypeus, in relation to its greatest width is of value in distinguishing some of the Lutzomyia species. As a probable adaptive feature related to host finding, the size of the compound eyes varies according to species, especially those of Lutzomyia, but may be characteristic of subgenera and equivalent species

PAGE 22

groups. Contrary to Divis (1967), ocelli are lacking in the Phlebotomines and Sycoracines. Illustrations are by far the best way to indicate eye size, the relative terms such as "snail" or "large" being understood easily with associated figures. The interocular distance or eye separation is the narrowest distance between the eyes and is measured directly or may be expressed by the number of facet diameters (or fractions thereof) needed to connect the eyes. The interocular suture (= post frontal suture of mosquitoes, Christophers, 1960) is complete in adults of BrumptonyLa (Fig. 8B) , Wavileya (Fig. 6A) , and Syaovax (Fig. 2A) but is incomplete in those of Lutzomyia (Fig. 12A) . In Phlebotominae, the maxillary palpus (Fig. 12A) consists of 5 segments, the basal one (palp 1) always the smallest and least important in taxonomy (Lev/is, 1973). Being partially fused with the second, it is difficult to accurately measure. The relative lengths of the segments are expressed often by a palp formula or by ratios with palp 1 being unity or 10 (Kirk &. Lewis, 1951). A palp formula of 1-4-2-3-5 indicates that palp 1 is the shortest, palp 4 the next shortest, etc. When two segments are equal in length they are enclosed in brackets, e.g. 1(4-2) -3-5. The sycoracines have four, instead of five, palpal segments (Fig. 2E) . Species in both subfamilies have palpal sensilla (= Newstead's scales) which are small clubbed sensory organs attached to one or more segments in the phlebotomines and to palp 2 in the Colombian Syoovox spp. The cylindrical antennal f lagellomeres, unlike true segments, are not independently musculated (Imms, 1938). For this reason I join the increasing number of students who use the term "flagellomere" in the place of "antennal segment." The antenna consists of a scape, pedicel,

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-10and 14 flagellomeres, each of which is designated by a Roman numeral (Fig. 12,\;. In Sycorax the terminal flagellomere is markedly reduced in size (Fig. 2D). The length of flagellomere 1 (= antennal segment 3), the most basal flagellomere, in relation to other flagellomeres, head height, or labrum length is useful as a diagnostic feature at the species or subspecies level. The paired antennal ascoids (fig. 12B) , often difficult to observe, are important owing to interspecific variation in their distribution, shape, and length. Some specimens may have but a single ascoid on a particular f lageilcmere. The raouthparts of New World phlebotomines in relation to feeding habits and taxonomy were studied in detail by Lewis (1975a). Following him, I use the term labrum for the labrum-epipharynx of various authors. Its length is measured from the apex of clypeus to tip of proboscis. The cibarium (= buccal cavity) lies within the clypeus and is armed with a variable number of teeth in the Lutzomyia and Brumptorrqiia females. Those in the latter genus are characteristically arranged in 4 longitudinal rows (Fig. 10M). Cibarial teeth are absent in both sexes of Warileya and Sycorax and are poorly developed or absent in the Lutzomyia and Bvurrp terry ia ma 1 e s . When viewed ventrally as in Fig. 12E, the horizontal teeth (= hind teeth) of most Lutzomyia females indeed look like teeth, their tips pointed, or not, depending on species. The vertical teeth (= fore teeth) usually appear as dark dots distad of the horizontal teeth at the same angle of view (Fig. 12E) . Lateral teeth may occur on both sides of cibarium near the horizontal teeth (Fig. 30M) . The number, position, shape, and size of these cibarial teeth are very important in species diagnosis and classification.

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-11Two other features of the clbarium are noted in the descriptions and keys. The nhitinous arch (Fig. 12E), when complete, crosses the ventral wall of the cibarium. Its development varies according to species ana may be complete or not. The. pigment patch on the dorsum of cibarium (Fig. 12E) varies in size, shape, and degree of infuscation (= pigmentation) in the Lutzomyia spp. The pharynx (Fig. 12C) , composed of 3 chitinous plates, is attached to the cibarium. Many species of Old World phlebotomines (genus Sevgentomyid) have spines on the posterior part of the pharynx but they arc relatively rare in the Lutzomyia spp., occurring mostly in some species in the eayannensis and oswatdoi groups. The majority of Lutzomyia females have transverse, unarmed ridges on the wider, posterior area of the pharynx. The length of the pharynx is given in descriptions although it is flexible and difficult to measure with accuracy (Lewis, 1967a). I adopt th.p terminology of Davis (195 7) as applied to the thorax. The mesonotum should correctly include the mesoscutum, mesoscutellum, and postnotum and should not be restricted to the. mesoscutum alone (Saether, 1971). Trie degree and distribution of pigmentation of the thorax and other body regions are important in associating sexes and for distinguishing species of Lutzomyia. Coloration has not been given much importance in systematic works but 1 find it generally dependable as a taxonomic character as applied to the New World phlebotomines . Pleural setae on the anepisterum (upper episternal setae) and katepisternum (lower episternal setae) are present in species of Lutzomyia. Brumptomyia, and Sycovax. The WariZeya spp. and the majority of Old World phlebotomines lack such setae (Abonnenc & Leger, 1975). Additional

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-12pleural setae, not: mentioned in other studies to my knowledge, occur behind the metathoracic spiracle in all species of Phlebotominae and Sycoracinae examined, those in the latter subfamily being 4 in number and relatively stout ^Fig. 2ii) . The 4 postspiracular setae in the phlebotomine species are reduced in sii'.e, barely visible (Fig. 12D) . The wing length is measured from the basal costal node (= rudiment of tegula; Snoigrass, 1935) to its apex; its width at widest part (Fig. 12G.i . Certain wing vein sections (Fig. 12G) are given useful, easily remembered names. Alpha (a) is the length of R from its junction with R to the costa. Beta (3) is the length or R from the junction of R to the junction of R„ + R, . Gamma (y) is the section of R from the r m crossvein to the junction of R . Delia (5) is that part of R distad of the junction of R and R . It is negative when R ends before this junction . The legs are measured as in Fig. 12H. A few phlebotomine spp. have a row of short spines on the hind femur (Fig. 31F) . The length of the femur in relation to that of the tibia and/or basitarsus is sometimes used for distinguishing species of Lutzomyta. The setation of the abdominal tergites 2 and 6 is useful in the classification cf Old World phlebotomines but has been little studied for the New World species. lewis (1975b), however, discussed and figured the socket patterns for some species, noting that the patterns were of 5 types. Setae on the sides of tergite 8 may be present or not in New World females of Phlebotominae and Sycoracinae, The nature of the abdominal sternites, especially sternites 2 ^nd 3, is useful in associating male and female sand flies (Hertig & Fairchild, 1950) but the character state is variable among individuals of conspecific I ipulations (Forattini, 1954).

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The structures of the bilaterally symmetrical male genitalia, labelled in Fig. 2 and 12, sr-i extremely important ir systematics. After eclosion, the genitalia rotates 180° in the phlebotomine males but not in those of Sycor-ax. The style, attached to the apex of the coxite, bears i to 6 major spines* and to many additional small setae. The presence of numerous strong spines is presumably a plesiomorphic feature (Tbeodor, 1965). The coxite is the dorsal basal appendage in the phlebotomine males and, of course, is the ventral basal one in the Sycovax males. It may or may not have a setal tuft or other nondeciduous setae on its inner surface. These setae are those which remain on the coxite after maceration and which usually differ in size or shape from other setae on the appendage. The fused, paired aedeagi, each of which is called the aedeagus is sub triangular and well sclerotized in most phlebotomine males. Some species (e.g. L. walkeri) have aedeagi with dorsal projections. The aedeagus of the Colombian Sycorax males is complex as in Fig. 2F. There is a basal, laterally flattened sperm pump within the abdomen, distal paired genital filaments and other structures mentioned on p. 31. The genital pump of the phlebotomine males is probably homologous with the sperm pump of the Sycovax spp. Leading from it are 2 genital filaments which pas.? through each aedeagus and which vary in length, width, and sclerotization according to species. The apices are simple, modified, inflated or not. *It is difficult to define major (= strong) spine, as it is a relative term referring here to size. One author's interpretation may differ from that of another. In doubtful cases, the illustrations should be studied to understand the meaning of statements in the text.

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-14The phi ebo torn! ne parameres lie between the coxites and ventral lateral lobes when viewed laterally. Their shape and setation are important in differentiating species. Simple parameres , i.e., I hose without arras or processes, are shown in Fig. 16F and ISA. Complex parameres are illustrated in Fig. 44H. The cercopods of the Syaorax males appear to correspond to the parameres of Phlebotominae and are very complex. The ventral, nonsegmented lateral lobes are more or less uniform in the phlebotomine males, although their length, width, and setation may offer good characters in some species. Corresponding structures are wanting in the eycoracines . The shape of the setose cerci may help associate conspecific sexes but in general these structures are of limited use in species diagnosis. The size and structure of the internal paired spermathecae and their ducts (Fig. 12F) are particularily important in the classification and identification of female sand flies. An idea of the variety of spermathecae can be gained by scanning the illustrations in this review. A well sclerotized, "Y" shaped genital fork or furca in the phlebotomines is associated with the spermathecae.

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METHODS AND MATERIALS Field Collections For sampling phlebotomine and sycoiacine populations, several methods should be used to determine species diversity and relative abundance. In Choc.o Department and elsewhere to a lesser extent, we used the following methods to capture these flies. Battery powered light traps (Sudia and Chamberlain, 1962) attract many psychodids, the numbers varying according to weather conditions, location, species composition, etc. We secured these, traps to tree branches 2 meters above ground level but did not use carbon dioxide as an adjunct attraction. At Curiche, where at least 1 trap per week was operated throughout the night (1800 hrs.-0700 hrs.), we collected a total of 23 phlebotomine spp. (3786*6*. 789oo) from April to Dec, 1967 (39 trap nights). We recorded 36 sand fly species from Curiche based on all collection techniques . Shannon traps (Shannon, 1939) made from muslin bedshaets do not trap insects per se but are collecting devices similar to those used by lepi'dopterists in :, sheeting" for moths at night. A gasoline lantern provider a light source enabling collectors to readily aspirate the psychodids which land on the illuminated cloth. They are attracted to the light, to the collectors, or to a combination of both. The species composition of light and Shannon trap captures is generally similar. -15-

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-16Malalse ti'aps (Townes, 196?) and flight traps (similar to the design oi Gressitt and Oressitt, 1962) were placed in forest clearings, across trails or othei flyways at ground level and in the forest canopy. Insects which land on the trap baffles eventually die after making their way upwards to the killing jar(s) charged with potassium cyanide. Ordinarily, there trapp do not capture large numbers of psychodids but they are effective in sampling the species composition of a given area if operated over extended periods of time. For example, in the Curiche Forest at groend level, one Malaise trap captured 24 Phlebotomine species (3166*.:?, 482^o) from 3 April 1967, to 30 Nov. 1967 (109 trap days"). Whenever possible, we searched diurnal resting sites for sand flies. These included tree trunks, especially dark crevices between buttresses, animal burrows, and litter on the forest floor. Tree trunks are favored resting sites for several species, specimens of which are easily captured with a simple aspirator. During the survey in Choco Dept., especially at Curiche and Teresita, we collected large: numbers of phlebotomines on human bait. The majority of collections were made by 2 men sitting on the forest floor between 1830 and 2100 hours. Phlebotomines were captured with aspirators as they attempted to feed, the results computed on a man-hour basis, i.e., the total number captured in one hour divided by the number of collectors. Specimens were preserved dry in cardboard pill boxes rather than in alcohol to prevent hardening of muscle tissue. *0ne trap day equals 24 hours.

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-17Laboratory Methods For routine. identification of phlebotomines and sycoracines, undissected specimens are first macerated (i.e., cleared) in 10-20% NaOH. The loss of pigmentation can be reduced by heating this solution (containing the files) to the boiling point rather than macerating them at room temperature for 12 hours or more. The flies should be left in the hot NaOH for 5 minutes, then placed directly into a drop of 80-95% liquid phenol (C,H OH) in a depression microslide. Within 5 minutes, the important internal structures such as the spermathecae and ducts are clearly visible and should be drawn if necessary, Once identified, the flies can be discarded, preserved in vials of 70% alcohol, or processed further for slide mounting. 1 follow the procedure of Fairchild and Hertig (1948c) except that Canada balsam or Euparal is substituted cor copaJ . Other slide-mounting procedures are discussed ny Osorno et al. (1966) Quate & Steffan (1966), and Lewis (1973) and others. The choice of which to use is a matter of personal preference but in all cases the best preparations are made from freshly killed flies. Sometimes it is necessary to remount specimens because of undesirable position and/or shrinkage. For those embedded in Canada balsam or other xylene—soluble media, it is advisable to submerge the entire slide in liquid phenol in a suitably closed container such as a petri dish. After i to 7 days, the specimens become soft and are easily handled. Xylene as a solvent should be avoided as the specimens tend to become brittle and easily damaged.

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-18Specimens were drawn with the aid of a camera lucida and Bausch and Lomb microprojector. I measured specimens with a calibrated ocular micrometer. All such measurements are given in millimeters throughout the text and figures.

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DISTRIBUTION, COLLECTING LOCALITIES, AND CHARACTERISTICS OF THE FAUNA The distribution of phlebotomines and sycoracines can be understood by present ecological factors, both biotic and physical, and by knowledge of past changes in climate, vegetation, and geology. Haffer (1967, 1974) discussed the probable speciation and distribution of some neotropical birds in relation to past changes, especially those occurring in the Pleistocene and post Pleistocene periods. During times of drought, populations of forest birds and other organisms were restricted to isolated forest refugia which acted as core areas of speciation. As conditions became wetter, these forests enlarged in area, often merging with others to form zones of secondary contact. The presumed forest refugia in Colombia and elsewhere were discussed and mapped by Haffer (1967, 1974) snd Brown. (1975), the latter author studying speciation in forest butterflies in relation to these presumed refugia. Although more data are needed, the refuge theory offers a plausible explanation for understanding recent speciation in the Phlebotominae and Sycoraciiiae. The majority of New World species inhabit forests, especially those in the lowland tropics which receive 2000+ mm of rain per year. For discussion purposes, it is convenient to divide Colombia into natural regions. D'Aliesandro & Barreto (19 71) delineate 7 such areas, their large "Oriental Region" consisting of both Amazonian forest and eastern natural savannah (llanos) . Within each region there are one or more life zones, each defined by a combination of biotemperature, annual -19-

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20prec.Ipitation, humidity, and vegetation (Holdridge, i 96 /) . The life zones in Colombia were studied by Espinal and Montenegro (1963) . Their ''bosque humedo" translates to "tropical raoist forest" (2000-4000 mm of rain per year) and "bosque muy humedo" corresponds to "tropical wet forest" (4000-8000 mm of rain per year) . The "bosque pluvial" (= "tropipal rain forest"), tne wettest lowJand life zone, receives over 8000 runi of rain per year. Figure 1 depicts where phlebotomines and sycoracines have been collected in Colombia. Osorno et al. (1972a) provide specific data for most localities; others from which collections were made by me or my colleagues are discussed below in relation to faunal regions (D'Allesandro ?* Barreto, 1971) and distribution centers (Haffer, 1974). Pacif ic Coast Reg ion Haffer (1967) attributes the "high concentration oi endemic species in the tropical lowland forests of western Colombia and Central America" tc a gradual accumulation of isolates in the trans-Andean forest region.* Tao dominant refugium on the Colombian Pacific coast is the Choco refugium extending from Lago Calima (Valle) to north of Quibdo (Choco) and across the Rio Atrato and San Juan river systems (Brown, 1975) . An analysis of the phlebotomine species and subspecies inhabiting the Pacific coast region indicates the following. The vast majority of the 41 known taxa, probably originated in forests east cf the Andes (cis-Audean region) . Most of these reached the *The tiVms-Andean region comprises "Middle America and the narrow Pacific lowlands along the western base of the Andes" (Haffer, 1967). Cis-Andean refers to lowland forests east of the Andes.

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-21Pacific coast forests by advancing around, net across, the northern Andes. These species which now occur on both sides of the mountains include: Lutzomyta ayvozai, L. guyanensis, L. bispinosa> L. gomezi., L. ncrdestina, L. sewcena, L. pilosa, L. spinosa, L. trinidadensis , L. shannonij L. tvbeYculata.1 L. trn.rarr.ulCj and others. Nine of the 41 species and subspecies do not occur in other regions of the Republic. Of these, only Lutzomyia sp. of Anchicaya is precinctive, the others occurring also in the trans-Andean regions of Northwestern Ecuador and/or Central America. These species include: Brumptomyia hcj-.ata, Warileya nigrosaoaulus ., L. rebur-rc.., ."'. veauvva, L. isovespertilionis, L. ylephiletov , L. sangu.ina.via, and possibly L. od.ax. Two forms which probably evolved in Pleistocene forest refugia within the transAndean region include L. cavvevai thula n.spp. and L. barrettoi rnajusouta n.spp. , both of which now occur in trans-Andean regions of Ecuador, Colombia, and Central America and east to the northern foothills of the Central Cordillera of the Colombian Andes. There is a slight possibility that some phlebotomines were able to cross the Andes in southern Zcuador and northern Peru in times past when wet forests occupied the now dry valleys. Some Amazonian birds apparently colonized Pacific coast forests in such a manner (Haffer, 1967) . Phlebotomines, however, are rather weak fliers and it would have been necessary for lowland forms to survive at elevations above 2000 meters above sea level. The one possible example of successful colonization may be that of L. vehuri'a — a species whose closest allies occur entirely in the (•?•?'&— Andean region, especially in Amazonian forests. A few species such as L. vespertitionis and its sister species, L. isovespertitionis probably invaded the Pacific coast region of Colombia

PAGE 35

from Central America, the former species also occurring east of the Andes, at least as far south as Bolivar Department. Depart mant of Cho cc Three collecting sites within this Department were described by Eidridge & Fairchild (1973) and Eldridge et a] . (19 73) . Maps accompany their descriptions. Cuptche. On narrow coastal plain between western slopes of Serrania del Baudo and Humboldt Bay on the Pacific coast; "tropical wet forest;'' elevation* less than 10 m. Sand flies were collected from April 1967, to :tec. 1967, in disturbed and undisturbed evergreen forests end near a mangrove swamp . Alto Cupi-dhe. About 3 km inland from Curiche on western ridge of Serrania del Baudo; "tropical wet forest;" elevation 302 m. The sand fly fauna is similar to that of nearby Curiche. Teresita. At eastern foothills of Serrania de Baudo, near Rio Truando; transitional life zone, "tropical moist/tropical wet forest"; elevation 35 m. Collections made in semi-disturbed forests on flat or gently rolling terrain from March-Dec. 1967. Depar tment of Va lle Ancth.iccvja Dam. About 35 km inland from Pacific Ocean, on the Rio Anchicaya; "tropical wet forest;" elevation ca . 560 m. This site is similar to that near the Rio Anori (Antioquia Dept.) in terrain, elevation, ^Elevation is expressed here and elsewhere in meters above sea level

PAGE 36

-23and climate. Sycoracines were recorded from both localities and from one other in Cauca Department. 25 km E of Buenaventura. "Tropical rain forest" according to Espinal (1968); elevation ca. 50 m. Phlebotomines were captured in a patch of disturbed forest surrounded by cultivated fields. A tlantic Coast Region This area encompasses not only the Carribean lowlands but also the northern foothills of the western and central Cordilleras of the Andes. The presumed Nechi refugiura is located here, a center "heavily influenced by the Chor.o center to the west" (Haffer, 196 7 ) . The present life zones are more diverse here than in the Pacific coast forests and except for the area near the Pvio Anori, the phlebotomine fauna has been little studied. Intensive collections of phlebotomines made by Dr. C.H. Porter and his colleagues near the Rio Anori (19 70-1971) yielded 37 species and subspecies. Thirty of these also occur in Pacific coast forests, a fact not surprising based on the distribution of other organisms (Haffer, 1967, 1974). Lutzorrvjia. stvictivitta may be endemic. Lutzomyia yuil-ti and L. noeticola are eis-Ardean taxa. The former reaches its northernmost limits in northern Colombia; whereas L. noeticola invaded extreme western Panama but populations between there and northern Colombia have since disappeared. Depart ment of Antioq uia Rio Anori (7° 19'N; 75° 04'W), about 24 km SW of Zaragoza; "tropical wet forest;" elevations from 500-700 m. Located in the foothills of the

PAGE 37

centra] Cordillera of the Andes, this site is mostly well forested with steep, well drained slopes. A detailed description is given by Porter & Defoliart (MS) . Pep art meat of Hagdalena Three sites near Santa Marta yielded sand flies from tree trunks or under loose bark of standing trees in cultivated or semicleared areas (August, 1973). Rio Don Diego, E of Santa Marta within 1 km of Carribean Sea; "tropical wet forest;" elevation ca . 3m. Fetuccn Santa McOfta and Mvnco.-, near a small stream; probably a transition zone between "tropical wet forest" and "tropical dry forest;" elevation ca. 80 m. Minca, SE of Santa Marta; life zone indeterminate but similar to preceding site; elevation 200 m. Andean Region Nearlv all records of phlebotoraine species occurring above 1000 m are given by Osorno et al. (1972a). Based on these and on my limited collections from the Western Cordillera, it is apparent that the Lutzornyia species ill the Vewusaxtm and vexatov groups dominate the rather depauperate phiebotomine fauna. De partroent of Vail e Three sites W or SW of Cali — 1. Near Piohinde , 2. In forest at the television tower just E of Salidito, and 3. Near Rio Ranee, SW of Cali;

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-25"lower montane very wet forest;" elevations from 3 570-1700 m. Limited collections from 31 July, 1973, to August, 1973. Departm ent; ot Ca uca iTasterr slope, near summit of Cerro Munchique, west of Popoyan; "lower montane rain forest;" elevation ca. 2500 in. Briefly surveyed on 5-6 August, 1973. One Si/eorax sp . taken but nc phlebotomines. Oriental Region Occupying a huge area east of the Andes, the Amazonian forest in Colombia is apparently rich in phlebotomine species but few collections have been made there or in the more northern Llanos which forms part of the Oriental Region (Haffer, 1967, fig. 1). Many of the species known from the Amazonian forests in Brazil, Peru, and Ecuador are expected to occur also in Colombia. C ore isana of Amazonas About 17 km W of Leticia; "tropical moist forest;" elevation 84 m. PhlebotoTtine flies were relatively rare in late July, 1973, the bulk of the material being taken in semi-disturbed forests near the Amazon River on tree trunks., less commonly in light traps. July and August are the two driest months at this locality.

PAGE 39

[n c eudcncia of Caq uota Tres Esquinaj; transitional life zone, "tropical moist forest"/ 'tropical wet forest;" elevation ca. ?00 m. Using light traps, Dr. C.J larinkelle collected numerous phleboto.mines in November, .1971.

PAGE 40

Figure 1 Map of Colombia showing localities where phlebotomises and sycoracine flies have been collected (records based on Osorno et al., 1972a and this review). Shaded sections represent areas 1000 m+ above sea level, Legend » = Fhlebotomine record X = Sycoracine record 4 Fhlebotomine and sycoracine record 1 = Atlantico Dept. 2 = Magdalena Dept. 3 = Norte de Santander Dept. 4 = Caldas Dept. b = Risaralda Dep 1 ". 6 Quindio Dept. 7 Cundinaniarcs Dept.

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-12. V M V Choco ^f't^soM.K',, 3oyac< S
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MEDICAL IMPORTANCE The phlebotomlne sand flies, unlike the sycoracine flies, are notorious vectors of leishmaniasis (Ward, 1977), bartonellosis (Schuitz, 1968), and certain arboviruses (Tesh et al., 1974) , diseases which have caused a great deal of human suffering in the neotropics and elsewhere. Leishmaniasis, a collective term referring to several disease entities caused by Leishmanial spp . , occurs j.n Colombia but the vectors are poorly known. Reyes (1957) provided limited information on the distribution and nature of dermal (= cutaneous) leishmaniasis in the Republic, noting that 206 out of 725 human infections involved the naso-pharyngeal region (mucocutaneous leishmaniasis) . Ward, citing figures from Reyes (op. cit.) and Garnham (1962), stated that nearly 2000 human cases of cutaneous leishmaniasis were reported in Colombia from 1948 to 1955. The true incidence, distribution, and identity of leishmaniasis in the Republic remains largely unknown, however. Visceral leishmaniasis (kala-a:'.ar) , a very serious disease, is apparently rare in Colombia, the few human cases from Santander, Tolima, and Cur;dinamarca Departments having been discussed by Arjona et al . (1971). An outbreak of Bartonellosis (oroya fever, Carrion's disease, etc.), lasting from 1925 to the early 1940' s, in southwestern Colombia (Cauca and Narino) , was reviewed by Jaramillo (1943). From 1940 to 1943, there •29-

PAGE 43

were 2 , 24 L cases with a mortality rate of nearly 16% (Samaniego, 1944). The. disease, also known to occur in Peru and Ecuador, was probably introduced into Colombia "by returning soldiers or by 'colporteurs' who travel from village to village" (Rozeboom, 1947b) . Circumstantial evidence led investigators to believe that LutzoryCa Columbiana was the responsible vector. Arboviruses were recovered from wild caught phlebotomines in the Pacific lowlands of Colombia (Barreto, 1969). The strain, designated Co Ar 3519, was discussed by Theiler and Downs (1973) and Tesh et al . (1974). Based upon the pioneering x;ork of Dr. Tesh and his colleagues in nearby Panama (1971-1975), it is safe to assume that ether arboviruses associated with phlebotomines exist in Colombia. Sand flies also play a role in the transmission of nonhuman parasites, especially protozoans. References to studies related to them are cited in the species bibliographies.

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SUBFAMILY SYCORACINAE Following Duckhouse (1972) I presently recognize 3 extant genera in the subfamily — Sycovax Haliday (ea. 21 sop.), Pavasyoovax Duckhouse (2 spp.), and Aposyoovax Duckhouse (1 sp.). Until now, only 4 species have been reported from the New World — Sycovax assimilis Barretto, 1956 and Pavasyoovax satohelli (Barretto, 1956) from Sao Paulc State, Brazil, Aposyoovax chi'lensis (Tonnoir, 1929) from Chile, and an unnamed sycoracine, the wing of which was figure;' by Fairchild (1955) from Paienque, Colon Prov. , Panama. The feeding habits of most species have not been studied but the females of Sycorax and Aposyoovax at least possess mouthparts adapted fcr sucking blood. Sycovax silacea Curtis, the type species of the genus from Europe, feeds on frogs and transmits a filarial worm to them (Desportes, 1942). The four new Colombian species, described here in the genus Sycovax, share several characters which set them apart from the other sycoracine species. In addition to the paired genital filaments between the parameres (= cercopods) of the male genitalia, there is a single median process which seems to form part of the aedeagal complex. It lies above the ducts and the proximal sperm pump (Fig. 2G) . Aposyoovax chilensis, the only sycoracine with inverted male genitalia (Duckhouse, 1972), has very long genital filaments but it lacks the elongate median process. The stout proximal spines (2 or 3) on each style of the Colombian and 31-

PAGE 45

Parasysoz'OX species distinguish then from the extant species in the subfamily. Other features, not always repeated in the specific descriptions which follow, are shared by the 4 Colombian Sycorax species. Coloration: Dusky brown, mesonotum but slightly darker than pleuron. Head: Broader than long; frons putted out behind antenna. Palpus of 4 segments, the: second with an inner group of 10-20 sensory rods. Antenna slender, of 16 "segments" (scape, pedicel, and 14 f lagellomeres) , the last reduced in size and terminating in a cone shaped peg sensilla (two similar sensilla on flagelloraere XIV) ; outer base of scape with 2 short sensory hairs as in A. chilensis; pedicel with 2-3 similar hairs; ascoids subequal in Size, often difficult to examine in available material. Female mouthparts include toothed mandibles and 4 boot shaped sensilla at tip of labrum; these features absent in males. Cibarium unarmed. Thorax'Pleura with 6+ setae mainly on the anepimeron but some may extend downwards to the katepimaron; upper episternal setae present or not; both sexes with 4 straight bristles below base of haltere (Fig. 2H) . Wing broadly rounded; veins sparsely haired; radial fork distad of medial fork. Abdomsn: Females without setae on tcrgites 8 and 10 or sternite 8. Male genitalia not inverted. Style with a patch of 8-15 short pointed steae at base; terminal spine slightly beveled at tip, 2 or 3 proxi.mal spines present, with or without a long subterminal hair; aedeagal complex as mentioned above; the paired genital filaments turned upwards, sinuous or not; parameres comewhat resembling those of A. chilensis but differing in the nature and position of the lobes, setae, and projections. Female genitalia: Each spermatheca and sperm duct forming a continuous tube with faint transverse striations, terminating in a heavily sclerotized "button"

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-33which appears shiny black; a tenuous sac usually amorphic following treatment in KOH, arises from the end of the button; an internal apcderae or furca associated with paired spermathecae. The combined length of the spermatheca and duct varies interspecifi rally am; probably corresponds to the length of the male genital filaments. This feature and the distribution of the antenna! setae and distance between the compound eyes were also used in associating the sexes o': three of Lite Colombian species but there remains the possibility that the males and females were not correctly associated. The immature stages of these and other New World species in the subfamily have not been discovered. Keys to the Syeorax Spe ci es Hales 1. Style of male genitalia with 3 strong spines and long subterminal hair; median process of aedeagus parallel-sided in dorsal view; lower distal projection of paramere pointed at tip. Antannal sensory setae, excluding the paired ascoids, arranged in a whorl-like pattern on flagellomeres 2 Style with 3-4 strong spines but without long subterminal hair; median process of aedeagus with basal two-thirds expanded and terminal one-third slender in dorsal view; lower distal projection of paramere rounded at tip. Antennal sensory setae, excluding the paired ascoids, not arranged in a whorl-like pattern 3

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2. Aedeagal ducts relatively long, exceeding the length of the. sperm pump. A. subtrlangular plate, dorsoventrally flattened, with acute bifurcate tip present below the base of the median process of aedeagus . Tip of median process of aedeagus rounded in lateral view colornbiensis (Fig. 3) Aedeagal ducts shorter, less than length of sperm purap and not as sinuous. Subtrlangular plate absent. Tip of median process of aedeagus angular in lateral view faivchildi (Fig. 4) 3. Style of male genitalia with A strong spines; median process of aedeagus more slender (Fig. 2G) . A larger species, wing length greatei than 2 mm andioola (Fig. 2) Style with 3 strong spines, median process of aedeagus relatively stout (Fig. 51). Wing length less than 1.5 mm. . trispinosa (Fig. 5) Ffiinalos 1 . Flagellomeres I-VIII with paired ascoids, other antennal setae randomly distributed. Eyes rather narrow, separated by distance = to 7 facet diameters trispinosa (Fig. 5) Flagellomeres T -IV with paired ascoids, one group of antennal setae arranged in whorl-like pattern. Eyes separated by 8 or more facet diameters 2

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2. Furca spade-shaped apically, its length at least % that of sperm duct + spennatheca. Flagellomere, 1, 0.20 mm, or longer , . ootombvensis (Fig. 3) Furca "v" shaped apically, its length but slightly less than that of sperm duct + spermatheca. Flagellomere 1 less than 0.20 mm long ........... fairchildi (Fig. 4) 1. Sycorax andioola n. sp. (Fig2) Male (holotype): Wing length 2.35; width 1.05. Head height 0.31; W.H1I.I1, u . .,/ » l_,^O.o ciCpaidCeu u_y U . J.T >->-. "; v_.j_._i _.<_.-.._ t Lu O j-o.i_c.i_ uj.u meters. Antenna (including scape and pedicel) 1.61 long; flagellomere I (0.26 long), about 1.8xlength of flagellomere II; paired ascoids (Fig. 2C) , subequal in size, on f lagellomeres I-IV only; other antennal setae mostly deciduous, not arranged in a whorl-like pattern. Palpal length 0.17; ratio of segments: .1-1.02-0.72-0.67; second segment with 20+ straplike sensory rods in a distinct patch. Pleura with 19-22 setae below wing base, without episternal setae. Wing venation as figured. Length of femora, tibiae, and basitarsi: Foreleg, 0.63, 0.73, 0.49; midleg, 0.63, 0.75, 0.47; hindieg 0.73, 0.82, 0.44. Visible sternites 2 and 3 with paired circular openings laterally. Genitalia complex: Coxite 0.31 long. Style 0.18 long wich 1 terminal spine, slightly beveled at tip and with a proximal row of 3 smaller spines. Paramere as shown with several setose lobes and projections, the most distal finger-like with rounded, upturned tips. Aede.agal complex of a laterally compressed sperm pump (0.20 long) within the abdomen; a median distal process (0.165 long) compressed dorsoventrally, curved and slender in lateral view, upturned at tip, basal

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-36two-thirds much wider than distal one-third when viewed dorsally; paired. sinuos, relatively long genital filaments. Cerci as figured. Material examirted: Colombia, c? holotype (no. 537), Cerro Munchique (Cauca) , 2450 m above sea level, light trap in cloud forest, 6 Aug. 1973, D.G.Y. and R.C.W. coll. c? paratype (no. 538), same data except collected by R.C.W., 8 Aug. 1975. Discussion: S. andicola, the largest Syoorax known from Colombia, closely resembles S. tvispinosa n. sp. in details of the male genitalia, setation of the antenna and in other character states. The two species, apparently allopatric, are separated by the characters given in the key. 2. Sycorax colombiensis n. sp, (Fig. 3) Male (holotype): Wing length 1.3 2; width 0.47. Head height 0.21; width 0.25; eyes separated by 0.126 mm or by distance = to 10 facet diameters. Flagellomere I (0.19 mm long), about 2.2xlength of flagellomere II. Paired ascoids visible only on f lagellomeres I and II; other sensory hairs in a whorl-like pattern on f lagellomeres I-X, the remaining f lagellomeres missing. Palpal length 0.10, ratio of segments 1-0.91-0.800.72. Pleura with 9-12 setae in one specimen, but lacking in the holotype. Wing venation as figured. Legs missing. Nature of sternites not determinable. Genitalia: Coxite (0.177 long); style (0.98 long) with a terminal spine, a long subterminal bristle and 2 proximal spines. Parainere as figured, the lower distal projection blade-like, pointed at tip. Aedeagal complex: Sperm pump (0.10 long); median distal process (0.09 long), slender and subequal in width in dorsal view, with a slender, rounded tip in lateral view; a subtriangular plate, dorsoventrally

PAGE 50

flattened, with acute bifurcate tip, pointing to the rear, present below the base of the median process of aedeagus. Cerci as shown. Female (allotype): Wing length 1.37; width 0.59. Head height, 0.24; width, 0.29; eyes separated by 0.15 or by distance = to 10 facet diameters, Antenna (1.15 long), flagellomere I (0.22 long), about 2.4xlength of flagellomere II; paired ascoids on flagellomeres IIV; other sensory hairarranged in a whorl-like pattern on flagellomeres I-XV. Palpal length, 0.113; ratio of segments 1-1-0.75-0.58. Pleura with 16-18 setae below wing base and or 1 upper episternal seta. Wing venation as shown. legs partly or wholly missing in all specimens, length of femora, tibiae, and basitarsi of foreleg: 0.43, 0.51, 0.27. Abdominal sternite 2 with paired circular openings at sides; other sternites apparently lacking these openings. Spernatheca and sperm duct length at least 0.15 from base to tip of button; furca spade-shaped at end. Material examined: Colombia, o* holotype (no. 529), Anchicaya Dam (Valle) , elev. ca. 400 m above sea level, light trap, 28 Jan. 1975, R.C.W. Q allotype (no. 530), same data as holotype except collected 10 June 1975. Paratypes (nos. 531-538), all from type locality in light traps; 1 c., 11 Aug. 1973, D.G.Y. and R.C.W. 1
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-383. Sijcovax faivrfriildi •.-.. sp. (Fig. 4) Male (holotype) : Wing length, 1.13; width, 0.47. Head height, 0.23; width, 0.28; eyes separated by 0.13 or by distance = to 10 facet diameters. Antenna about 0.79 long; flagellomere I (0.18 long), about 2.5 xlength of flagellomere IT; paired ascoids visible only on flagellomeres I-III, other sensory hairs in a whorl-like pattern on flageliomeres I-XV, palpal length 0.10 mm; ratio of segments: 1-1.18-1-0.72. Pleura with 5-11 setae below wing base and with 1-2 upper episternal setae (n = 5), Wing venation as shown. Sternites apparently lacking clear circular openings. Length of femora, tibiae, and basitarsi of slide 544: Foreleg, 0.42, 0.43, 0.24; raidleg, 0.45, 0.40, 0.24; hindleg, 0.47, 0.37, 0.20. Genitalia: Coxite, 0.16 long; style, 0.88 long, with a large terminal spine, a long subterminal hair and 2 proximal spines. Lower distal end of parainere terminating in a hook-like projection, acute at tip. Aedeagal complex: Sperm pump (0.11 long); median distal process (0.068 long), angular at tip and relatively wide in lateral view, subequal in width when viewed dorsally; aedeagal ducts short, each about 0.96 long. Cerci as shown . Female (allotype): Wing length, 1.47; width 0.61. Head height, 0.26; width, 0.30; eyes separated by 0.14 or by distance = to 96 facet diameters. Flagellomere I (0.16 long) about 2.3 xlength of flagellomere IT; paired ascoids, subequal in size (but not conspicuous) on flageliomeres IIV, absent from remaining flageliomeres, other nondeciduous antennal setae as in male. Palpal length 0.116; ratio of segments 1-10.92--0.78. Pleura with 10-11 setae below wing base and with 1-2 upper episternal setae (n 3). Wing venation as shown. Length of femora,

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39tibiae, and basitarsi: Foreleg, 0.39, 0,40, 0.22; midleg, 0.42, 0.47, 0.24; hindleg, 0.47, 0.50, 0,20. Spermatheca + sperm duct length about 0.08 from base to end oi subcircular button; furca somewhat "v" shaped apieally . Material examined: Colombia. 6* hclotype (no. 539), Anchicaya Dam (Valle) , elev. ca. 400 m above sea level, light trap in forest, 11 Aug. 1973, D.G.Y. and R.C.W. o allotype (no. 540), same data except, collected 23 Jan. 1975, R.C.W. Paratypes (nos. 541-547) all collected in light traps, 1 c?, Rio Anori (Antioquia) , Colombia, elev. ca. 410 m above sea level, 22 Sept. 1970, D.G.Y. 1 6*, Rio Anori, 23 Sept. 1970. 1 g, Anchicaya Dam, 11 Aug. 1973, D.G.Y. and R.C.W. 3 6*6* , 1 o. Anchicaya Dam, 28 Jan. 1975, R.C.W. 1 cf, Anchicaya Dam, 5 March 1976, R.C.W. Discussion: The short sperm ducts of S. faivchildi serve to distinguish this taxon from the other species of Sycovax in Colombia. I take pleasure in naming the species in honor of Dr. G.B. Fairchild who has contributed so much to our knowledge of medically important arthropods in the neotropics. 4. Syeorax trispinosa n. sp. (Fig. 5) Male (holotype): Wing length 1.22; width 0.56. Head height 0.23; width 0.28; eyes separated by 0.08 or by distance = to 7 facet diameters. Antenna, 0.88 long; flagellomere I (0.15), about 1.8xlength of flagelloniere II. Paired ascoids on f lagellomeres IIV, absent from remaining flagellomeres; other antennal setae mostly deciduous, not arranged in a whorl-like pattern. Palpal length, 0.12 mm; ratio of segments: 1-1.150.84-0.76, second segment with 15"-t rod sensilla. Pleura with 9-16 setae

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-40below wing base ana 1-4 upper epistemal setae (n = 10). Wing venation as figured. Length of femora, tibiae, and basitarsi: Foreleg; 0.45, 0.43, 0.26; midleg, 0.49, 0.44, 0.26; hindleg, 0.53, 0.52, 0.24. Abdominal sternites 2-5 with paired lateral openings. Genitalia: Coxite, 0.19 long. Style, 0.09, with 2 proximal spines and a terminal spina. Paramere as shown, similar to that of 5'. andicola, the most distal projection also curved at tip, finger-like. Aedeagal complex of a laterally compressed sperm pump (0.11 long), a median distal process (0.10 long), with the basal twc-thirds expanded, greater than twice the width of the terminal third in dorsal view; aedeagal ducts and cerci as shown. Female (allotype): Wing length 1.54; width 0.71. Head height, 0.31; width, 0.27; eyes separated by distance of 0.1 or by distance = to 6.9 facet diameters. Antenna, 1.0 long; flagellomere 1 (0.16 long), about 1.7 x length of flagellomere II. Paired ascoids on f lagellomeres I-VIII, absent from remaining flagellomeres, other antennal setae mostly deciduous not arranged in a whorl-like pattern. Palpal length 0.14 mm; ratio of segments: 1-0.89-0 .61-0. 61, palpal sensilla as in male. Pleura with 10-21 setae below wing base and 1-5 upper episternal setae (n 9) ; wing venation as shown. Length of femora, tibiae, and basitarsi: Foreleg, 0.44, 0.42, 0.27; midleg, 0.49, 0.48, 0.2.7; hindleg, 0.53, 0.56, 0.27. Abdominal sternites 2-5 with paired circular openings at sides, remaining sternites indeterminate. Combined length of spermatheca and sperm duct, from base to tip of sclerotized button, 0.15. Furca spade-shaped as shown . Mater-ial examined: Colombia, o holotype (no. 548), Anchicaya Dam (Valle) , elev. c.a. 400 m above sea level, light trap in forest, 5 March 1976, R.C.W. o allotype (no. 549), same data. Paratypes (nos. 550-573),

PAGE 54

all collected from type locality in light or flight traps by D.G.Y. and/or R.C.W. 2 oo, 10 Aug. 1973; 1
PAGE 55

Figure 2 Syoorax andioola male — A. Head, B. Wing, C. Pedicel and f lagellomeres 1 and II showing ascoids, D. Terminal 4 f lagellomeres, E. Palpus, F, Genitalia, dorsal view, G. Sperm pump, genital filaments, and median dorsal process of aedeagus, lateral view, H. Metathoracic spiracle and postspiracular setae. Male: Cerro Munchique, Cauca Dept., Colombia

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43j£\ v " FW1 J t for* K !: a MM Ns tl ^ I "'^v^^^fe^*/' G— i'-.i Filc^-nt

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Figure 3 SyaGPax eolorribiensis — A. Male head, B. Female wing, C. Male wing, D. Female head, E. Male genitalia, dorsal view, F. Male subtriangular plate, G. Spermathecae, H. Sperm pump, genital filaments, and median dorsal process of aedeagus, lateral view. Male: Anchicaya Dam, Valle Dept., Colombia Female: Same locality as male

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c / a V o U ^ i__ >u ^V v^ -*/< V* /

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Figure 4 SycOTPax fatvohi'ldi — A. Male head, B. Female wing, C. Male wing, D, Female head, E. Male genitalia, dorsal view, F. Sperm pump, genital filaments, and median dorsal process of aedeagus, lateral view, G. Spermathecae, H. Female pedicel and flageilomeres I and II showing paired ascoids and other setae. Male: Anchicaya Dam, Valle Dept., Colombia Female: Same locality as male

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X'.: ^:Pmj j >' s^~ i I J H Ml •" ^ F ^^; c_^

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Figure 5 Sycovax trispinosa — A. Male head, B. Female palpus, C. Female, terminal 4 f lagellomeres , D. Female pedicel and f lagellomeres I end TI showing ascoids, E. Female head, F. Female wing, G. Male wing, H. Male genitalia, dorsal view, I. Sperm pump, genital filaments, and median dorsal process of aedeagus, lateral view, J. Tip of furca, dorsal viev.'j KSpenhathecae. Male: Anchicaya Dam, Valle Dept., Colombia Female: Same locality as male

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-49/ H . s r; 1 \ .1 B g ill j * 9 £> / If ml

PAGE 63

SUBFAMILY PHLEBOTOMINAE Previous studies on this subfamily in Colombia were reviewed by Osorno et al. (1972a) and will not be repeated here except for litez-ature citations in the species accounts. Nearly all of these studies involve systematica, especially species descriptions and distribution records. Owing to the paucity of bionomic studies on the Colombian phlebotomines , I felt that it would be desirable to provide references to such studies carried out in other countries. The results of these studies may not always apply to conspecific populations in Colombia. Information on the biting habits of some species in Choco Dept. is provided in the species accounts . With few exceptions I have not redescribed known taxa because of adequate original or subsequent descriptions. Hopefully, the figures and keys will serve to distinguish the taxa. K ey to the New World Genera 1. Wing broad, rounded at tip; R„ + R„ + R, forks before, on same level as, or slightly beyond r-ni crossvein. Pleura without episternal setae. Female cibarium unarmed. Male genitalia with style longer than coxite Wcwileya (Fig. 6-7) Wing pointed at tip; R 4R + R forks well beyond r-m crossvein. Pleura with episternal setae. Female cibarium armed with teeth. Male genitalia with style shorter than coxite 2 •50-

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-51Interocular suture complete. Female cibarium with 4 longitudinal rows of horizontal teeth. Male genitalia with 5 large spines, 2 of which (usually basal pair) borne on a common tubercle Brumptonrjia (Fig. 8-11) Interocular suture incomplete. Female with 1 row of horizontal teeth; vertical and lateral teeth present or not. Male genitalia with style bearing 1-6 large spines, basal pair not borne on a common tubercle in those species having 5 spines . . .Lutzornyia (Fig. 12-89)

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-52Genus Warn, ley a Hert ig, 1948 This genus, recently enlarged to accomodate W. (H.) hertigi (Fchld.) by Lewis et al. (1978), contains 5 species, 2 of which in the subgenus Waviteya occur in Colombia. Little is known about the habits of any of the species although W. (W.) rotundipennis , one of the Colombian species, and W. (W.) phlebotomanica Hertig, the type species from Peru, are anthropophilic . Keys to Species Males 1. Palpal segment 5 longer than combined length of segments 2 + 3. Paired intraabdominal rods present. Genital filaments about 1.8X length of pump. Style lacking a basal bristle. Wing length less than 1.5 mm nigrosaccuhMS (Fig. 6) Palpal segment 5 shorter than combined length of segments 2+3. Intraabdominal rods absent. Genital filaments less than 1 . 3X length of pump. Style with a basal bristle. Wing length over 1.8 mm . . . rotundipennis (Fig. 7) Females 1. Spermathecae large and sac-like with smooth walls. Palpal segments 5 longer than combined length of segments 2+3. Wing length less than 2.0 mm nigrosaoculus (Fig. 6) Spermathecae cylindrical, somewhat worm-like, with transverse striations. Palpal segment 5 shorter than combined length of segments 2 + 3. Wing length over 2.0 mm rotundipervnis (Fig. 7)

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-535. Warileya (W. ) nigrosacaulus (Fig. 6) Warileya nigrosacaulus Fairchild & Hertig, 1951b: 428 (o. holotype, Cerro Campana, Panama Prov., Panama). Fairchild, 1953: 102 (cf. to Her tig ia hertig i) . Barretto, 1955a: 183 (listed). Forattini, 1971a: 107 (listed). Tesh et al., 1971a: 153 (blood meals, Panama). Martins & Morales, 1972: 366 (listed). Young & Chaniotis, 1972: 97 (cf, descr.). Chaniotis et al., 1972: 95-96 (in tree hollows, Panama). Christensen, 1972a: 88 (listed). Forattini, 1973: 536-533 (o, fig. ). Velasco & Trapido, 1974: (cf. to W. yungasi) . Lewis, 1975a: 500 et seq. (mouthpart morphol.). Lewis et al., 1978 (classif.). Distribution: Colombia (Choco, Valle) , Panama. Material examined: Colombia: 1 o", 1 o, Curiche (Choco), Malaise trap, 27 April 1977. 1 holotype, Cerro Campana (Panama Prov.), hollow log, 7 Jan. 1947, M. Hertig. 2 6cf, 5 op, near Gamboa, Canal zone, tree hollow, 10 Sept. 1969, B. Chaniotis. 4 6V, same data but taken 15 Jan. 1970. Discussion: A little known species, W. nigrosacaulus is easily distinguished from W. rotundipennis by the characters given in the key. Using a precipitin test for blood meal determinations, Tesh et al. (1971a) found that blood from 4 recently engorged nigrosacaulus females reacted with mammalian and reptile-amphibian antisera. Four other blood meals were nonreactive due to small volumes of blood and/or to the weakness of the reptile-amphibian antisera.

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6. Warileya (W.) rotundipennis (Fig7) Warileya votundipennis Fairchild & Hertig, 1951b: 424 (c? hole type, o, Cerro Campana, Panama Prov. , Panama). Fairchild, 1953: 102 (cf. to Eevtigia hertigi) . Barretto, 1955a: 188 (listed). Fairchild & Hertig, 1959: 122 (Cosira Rica). Johnson & Hertig, 1961: 765, 775 (rearing). Hanson, 1968: 93 (larval fig.). Forattini, 1971a: 107 (listed). Young & Chaniotis, 1972: 366 (listed). Christensen, 1972a: 88 (listed). Forattini, 1973: 536-538 (cf, o. fig.). Velasco & Trapido, 1974: 436 (cf. to W. yungasi) . lewis, 1975a: 500 et seq. (mouthpart morphol.). Lewis, 1975b: 366 (hair sockets, fig.). Lewis et al. : 1978 (classlf.). Distribution: .Colombia (Antioquia, Choco, Valle), Costa Rica, Panama. Material examined: Colombia. 2 no, Teresita (Choco), tree buttress, 15 June 1967. 27 oo. Rio Anori (Antioquia), light traps, Sept. 1970. 5 w, same data but 29 May 1970, C.H.P. 8 6V, 2 no, Anchicaya Dam (Valle), light trap, 28 Jan. 1975, J.E. Browne. 6 def, 6 oo, same data but taken 10 June 1975, R.C.W. 5 <5b", 4 ot) , same data but taken 16 July 1975. Panama. 6* holotype (no. 2335), o allotype (no. 2312), Cerro Campana (Panama), Shannon trap, 24 Aug. 1950, M. Hertig & G. Fairchild. 2 c?6*, Rio Changena (Bocas del Toro) , Shannon trap. 8-11 Sept. 1961, R. Hartmann 6 P. Galindo. Costa Rica. 2 &<$, 1 o, Turrialba, Shannon trap, 20 Aug. 1961, G. Fairchild & M. Hertig. Discussion: Warileya rotundipennis , the only Warileya species lacking intra-abdominal rods (Lewis et al . , 1978), has been reported feeding on man in Panama (Fairchild & Hertig, 1951b) but its role, if any, in disease transmission is not known. Ur. C.H. Porter also has collected man-biting

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-55feraales at the Rio Anori (Antioquia) locality, mostly in forest clearings (pers. comm. ) . The records of W. rotundipennis and W. nigvosaeovXus from Choco and Valle Depts. indicate that both taxa probably occur in other areas of the trans-Andean region perhaps as far south as Guayas Prov. , Ecuador.

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Figure 6 Wariloya. nigvosaocutus — A. Hale head, B. Male flageilomere II, C. Female head, D. Female flageilomere II, E. Female cibarium and pharynx, F. Male genitalia, dorsal, G. Tips of genital filaments, H. Female wing, I. Male v.dng, J. Spermathecae. Male: Curiche, Choco Dept., Colombia Female: Same locality as male

PAGE 70

-57fife,, CM) Steffi V <3 A

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Figure 7 Warileyo. vctund.ipenn-Ls — A. Male head, B. Male f lagellomere II, C. Female head, D. Female f lagellomere II, E. Female cibarium and pharynx, F. Male genitalia, G. Female wing, H. Male wing, I. Spermachecaa. Male: Cerro Campana, Panama Prov. , Panama Female: Rio Anori, Antioquia Dept., Colombia

PAGE 72

f / f ^ i l °% \\\ r ( / " \ \i/ I l( l )/l Hi/A r7 V v

PAGE 73

-60Genus Brurnptornijia Franca a nd Parro t, 1921 Characterized by Tkecdor (1965), Lewis et al. (1978), and others, the genus Bvu^tomyi-a now contains over 21 species, 5 of which occur in Colombia. Except for the female of B. hatnata which has very short sperm ducts (Fig. 101) , the females of the other species in Colombia are virtually impossible to identify in the absence of males. Brwnptomyia avelXarn.-, one of the Colombian species, was reported feeding on armadillos in Brazil (Mangabeira, J 942b). This and other Brumptomyia spp. often rest in burrows dug by these mammals. They do not bite man and the preferred hosts of most species remain unknown. Key to Species Males 1. Genital filaments less than 4X length of pump . . . . hamata (Fig. 10) Genital filaments greater than 4X length of pump 2 2. Coxite tuft of mostly stout, spine-like setae 3 Coxite tuft of slender, hair-like setae 4 3. Coxite tuft of 20+ setae implanted on a distinct tubercle. Paramercs more slender as shown gdlindoi (Fig. 9) Coxite tuft of fewer than 20 setae, usually about 12, not implanted on a tubercle. Paramere broader as shown avell-ari(Fig. 8) 4. Style with isolated spine well distad of proximal pair. Coxite tuft implanted on a distinct raised tubercle beaupertuyi (Fig. 8)

PAGE 74

-61Style with isolated spine at or near level of proximal spines. Coxite tuft implanted on a slightly raised, raspberry-like (circular) base leopoldoi (Fig. 11) 7. Brumptomyia avellari (Fig." 8) Fhlebotomus avellari Costa Lima, 1932: 48 ((?, Lassance, Minas Gerais, Brazil). Mangabeita, 1942b: 225 et seq. (iir.matures , adults, descr., fig.). Barretto, 1947: 187-188 (full refs.). Fairchild & Hertig, 1947a: 615-616 (cf. to galindoi) . Barretto, 1951: 212 (dist.) . Floch & Abonnenc, 1952: 39, 45 (6*, o, keyed). Rodriguez, 1953b: 55 (mention). Forattini, 1954: 214-217 (second sternice, fig.). Forattini & dos Santos, 1955: 17 (Brazilian record). Ortiz, 1963: 320 (o, keyed). Brumptomyia avellari: Barretto, 1955a: 187 (listed). Martins et al., 1961b: 309 (mention). Martins et al., 1962a: 380 (Goias, Brazil). Sherlock, 1962: 332, 335 (mention). Carnheiro & Sherlock, 1964: 315 (pupa, keyed, fig.). Fraiha et al . , 1970a: 468 (6* keyed) . Christensen, 1972a: 88 (Panama). Osorno et al., 1972a: 14 (Boyaca, Colombia). Forattini, 1973: 122 et seq. (gen. review, fig.). Llanos et al., 1976: 480 (Peru). Martin et al . , 1976b: 496 (Peru). Ramirez et al . , 1976: 599 (Venezuela) . Distribution: Panama, Colombia (Boyaca), Peru, Venezuela, Brazil, Paraguay . Material examined: Colombia. 2 6b*, 1 o, Puerto Boyaca (Boyaca). light trap, 6 May 1973, C.J.M. Brazil. 1 6\ Coqueiros (Sao Paulo), armadillo burrow, Oct. 1953, Albertin. Panama. 1 o", Tocumen Airport (Panama), light trap, 23 March 1953, F. Blanton. Paraguay. 5 do\ AcaPoi, San Pedro, burrow, 18 April 1950, M. Hertig.

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-62Discussion: Brumptomyia avellari and E. brurnpti (Larousse) with sympatric populations in Brazil and Paraguay, closely resemble one another, the males differing chiefly by the shape of the parameres. I regard the Colombian specimens as being conspecific with the former species. 8. Brumptomyia beaupertuui (Fig. 8) Fhlebotomus beaupertuyi Ortiz, 1954: 235 (o* holotype, Duaca, Lara State, Venezuela). Scorza & Ortiz, 1960: 434 et seq. (ecology). Pifano et al., 1962: 383, 337 (o*, q, keyed), 411-412 (cf, o, descr., fig.). Ortiz, 1963: 322 (^ descr.). Ortiz & Scorza, 1963: 350 (listed), 354 {&> keyed) . Ortiz, 1965a: 208 (mention). Scorza et al., 1967: 193, 195 (d\ o keyed) . Leon, 1969: 30 (listed). Phlobotoinus galindoi (not galindoi Fairchild h Hertig, 1947) : Pifano & Ortiz, 1952: 138 (listed, Venezuela). Leon, 1969: 30 (listed). Brumptomyia beo-upertuyi: Barretto, 1955a: 187 (listed). Sherlock, 1962: 321 et seq. (6*. fig., Colombia). Fraiha et al., 1970a: 468 (6*, keyed). Forattini, 1971a: 98 (listed). Forattini, 1973: 522 et seq. (.gen. review, fig.). Distribution: Colorrbia (Santander), Venezuela. Material examined: Venezuela. 2 6$ (topotypes), Duaca (Lara), no date, T. Ortiz. DiscussionAt present, this little-known species is represented in Colombia by a single male, skillfully illustrated by Dr. 0. Mangabeira (in Sherlock, 1962). Ortiz (1963) described the female based on Venezuelan specimens .

PAGE 76

•639. Brumptomyia galindoi (Fig". 9) Phlebotomus galindoi Fairchild & Hertig, 1947a: 615 (cfholotype, near Roquete, Chiriqui Prov., Panama). Floch & Abonnenc, 1952: 40 (6* keyed). Ortiz, 1954: 238 (cf. to beaicpertuyi) . Rosabal, 1954: 30 et seq. (
PAGE 77

20-24 May 1976, B.G.Y. & T. Rogers. Honduras: 1 6*, Tela, Lancetilla Valley, tree buttress. 24 July 1953, W. Hils. i 6", same data but light trap, 13 Jar-. 1954. Mexico. 2 cfd*, 2 og, Ocoscoautla (Chiapas), tree hollows, 8 April 1951, G. Fairchild & R. Hartnann. 2 cfo", Palenque (Chiapas), tree buttresses, 30-31 March 1951, G. Fairchild & R. Hartmarm. Paraguay. 1 cT, Sommerfeld, Yhu, tree cavity, 15 March 1950, V. Zelada. 1 6\ same data but 18 March 1950, M. Hertig. Panama. 1 6* (holotype no. 814), type locality, 17 Feb. .1947, P. Galindo. 1 £, same data but 26 March 1943. 1 cf, 1 o, Mojinga Swamp near Ft. Sherman, Canal Zone, light trap, 19 Nov. 1951, F. Blanton. Discussion: The holotype of B. gatindoi is a large specimen, wing length nearly 3.0 mm, from Chiriqui Province, Panama (ca. 1200 m above sea level). Smaller specimens from lower elevations in Santander Department, Colombia were later named B. mesai by Sherlock (1962) who separated the males on size, number of distal setae on the coxites, shape of the aedeagi and setation cf the parameres. Fraiha et al . (1970a) treated the taxa as conspecific, basing their decision on the original descriptions and on a male (identified as B. gatindoi by Dr. G.B. Fairchild) from a lowland locality in Panama. Aside from size and the number of distal coxite setae (7 in the gatindoi holotype, 5-6 in males from the lowlands) , I can detect no marked differences among the specimens listed above. The number of distal setae on the coxites probably varies according to the size of the insect and in this case, apparently represents inf raspecif ic variation. It remains to be determined whether or not the size of gatindoi individuals is correlated with altitude, i.e., is there a continuous increase in size of specimens from the lowlands to the highlands? Believing this to be the case but

PAGE 78

-65without evidence Lo support it, I tentatively consider B. mesai and B, galindoi to be conspecific following Fraiha et al. (1970a) . 10. Brumptomyia hamate (Fig. 10) Phtebotomus hamatus Fairchild & Hertig, 1947a: 614 (c? holotype, Chilibrillo bat caves, near Chilibre, Panama). Barretto, 1951: 217 (mention). Floch & Abonnenc, 1952: 39 (t? keyed) . Rodriguez, 1953b: 53 (t? s p measured, Ecuador), 55 (cf. to leopoldoi) . Ortiz, 1954: 237 (listed). Lewis & Garnham, 1959: 83-84 (6*, Belize, fig.). Garnham & Lewis, 1959: 24 (Belize). Hanson, 1961: 320 e'e seq. (breeding sites, Panama). Johnson & Hertig, 1961: 765 et seq. (rearing data). DiazNajera, 1963: 193 et seq. (Mexico). Ortiz & Scorza, 1963: 353 (
PAGE 79

Dioeussion: Although the adults of B. hamaia are rarely encountered, the larvae may bo quite common in soil at. the base of trees. Hanson (1961) and Rutledge & Mosser (19 72) in Panama found that the larvae of this species far outnumber those of other phlebotomine species in this microhabitat. Like other Brumptomyia spp., the fourth instar larva has only two caudal setae. 11. Brumptomyia leopoldoi (Fig. 11) Phlebobomus leopoldoi Rodriguez, 1953b: 52 (6* nolo type, Quevedo, Los Rios Prov., Ecuador; ri, Naranjai, Guayas Prov., Ecuador). Ortiz, 1963: 320 (^ keyed). Br-wnptomyia leopoldoi : Barretto, 1955a: 187 (listed). Sherlock, 1962: 332 (cf. to mesai) . Williams, 1970: 331 (Belize). Fraiha et al . , 1970a: 468-469 (6" keyed, as possible synonym of guimaraesi) . Forattini, L97ia: 98 (listed). Christensen, 1972a: 88 (listed, Panama). Forattini, 1973: 530 (as synonym of guimaraesi) . Williams, 1976a: 603 (in caves, Belize) . Distribution: Belize, Panama, Colombia (Antioquia, Choco, Valle) , Ecuador. Material examined: Colombia. 3 6*0*, Rio Anori (Antioquia), light traps, May 1970, C.H.P. 1 6*, Curiche (Choco), light trap, 26 May 1967, D.G.Y. 2 6*0*, 5 05, ca. 10 km W of Cali near Pichinde (Valle), rock crevice, tree trunks and light trap, 31 July 1973-4, Aug. 1973, D.G.Y. & R.C.W. Ecuador-. 1 o 1 (paratype no. 4314), Quevedo (Los Rios), no date, J. Rodriguez. 2 def, 2 go, 17 km K of Santo Domingo de Los Colorados (Pichincha) , mammal burrow, 4 May 1976, D.G.Y. & T. Rogers. 1 6", same

PAGE 80

-67data but 27 May 1976. 1 6*, same data but light trap., 27 May 1976. Panama. 1 c? (paratype no. 4292), Almirante (Bocas del Toro) , animal burrow, 22 dan. 1953, W. Hils. 1 o 1 , same data but tree buttress, 27 Jan. 1956, R. Eartmann. Discussion'. 3. leopotdoi, considered to be conspecific with B. gui'P.araesi (Coutinho and Barretto) by Forattini (1973) , is specifically distinct based upon the following considerations: 1. The nature and size of the basal coxite tufts and the structure and setation of the parameres differ markedly between the txvo males (cf. Fig. 11C and 111). The original figure of leopoldoi (Rodriguez, 1953b) shows a rather broad paramere but I believe this is a distortion due to mounting technique, not a reflection of normal structure. Paratype no. 4314 is obviously flattened. 2. The genital pump cf the gui.rnavaesi rale is less flared and notably shorter (0.12 mm-0.15 mm) than that of leopotdoi which is 0.22-0.24 mm long (cf. Fig. 11G and 11H) . 3. The isolated spine of the style of guimor>aesi is inserted at 0.59-0.61 of the segment whereas that of leopoldoi is located at 0.53. In addition to the leopoldoi specimens listed above, I examined 4 guimavciPsi males from southern Brazil: 2 <3a, Faz. Ribeirao de Baizo, Patos, Minas Gerais ; 1
PAGE 81

Figure 8 Br-wnp terry ia o.vellar-i male — A. Genitalia. Male: Puerto Boyaca, Boyaca Dept . , Colombia Bvumptomyia beauper-tuyi. male — B. Head, C. Flagellomere II, D. Paramere, E. Wing, F, Genital pump and filaments, G. Genitalia, H. Coxite tuft, same scale as Fig. 8A. Male: Lara State, Venezuela

PAGE 82

X k3 \ V; Yi /

PAGE 83

Figure 9 Brumptomyia ga'Lindoi — A. Male head, B. Male f lagellomere II, C, Female hsad, D. Female flagellomere II, E. Female cibarium and pharynx., F. Genital pump, G. Genital pump and filaments, H. Male genitalia, 1. Paramere, J. Coxite tuft, K. Spermathecae, L. Body of spermatheca, M. Female wing, N. Male wing, 0. Female cibarium. Male: Teresita, Choco Dept., Colombia Female: Same locality as male

PAGE 84

71<:•. -y-./.-ys \ to ,438 L m I Ph 6 I : I / f\\ ft A // if I is / ^A 1/ A\ V v

PAGE 85

Figure 10 Brvjnptomyi-a hconata — A. Male head, B. Male flagellomere II, C. Female head, D. Female flagellomere II, E. Female cibarium and pharynx, F. Genital pump and filaments of male from Teresita, Choco Dept., Colombia, G. Aedeagus and Faramere, H. Male genitalia, I. Spermatheca, the other not drawn, J. Body of spermatheca, K. Female wing, L. Male wing, M, Female cibarium. Male: Barro Colorado Island, Panama Canal Zone (except Fig. 10F) Female: Teresita, Choco Dept., Colombia

PAGE 86

/ 3MS /v; BfcT 9 ,1\ CA> ! C r D / cos ^H /' V /* 8 . /> I v -" \ \ *& c ^ ^ Ml ^

PAGE 87

Figure 11 Bx'imptomyia leopotdoi male — A. Head, B. Flagellomere II, C. Coxite tuft, paramere and aedeagus of male from near Santo Domingo de Los Colorados, Ecuador, D. Wing, E. Genital pump and filaments, F. Genitalia, G. Genital pump of male from Fig. 11C locality. MaJe: Rio Anori, Antioquia DeptS Colombia (except for Figs, 11C and G) Bmenptomyia guimaraesi male — H. Genital pump, I. Coxite tuft, paramere and aedeagus, same scale as 11C. Male: Itaporanga, Sao Paulo State, Brazil

PAGE 89

•76Germs _£ u tzorrcj ia Franga, 3 924 As mentioned earlier the classification of the Lutzomyia sand flies is based upon the scheme, of Lewis et al. (1978). It should be emphasized that this represents a flexible classification, subject to change as new species, new character states, or unknown sexes are discovered. It is hoped that the following keys and discussions will be helpful to those interested in sand flies and disease in Colombia and elsewhere in the neotrooics. Keys to Subgenera, Species Gr oups, and Un g ro upejd Species of Lutzonrjia In order to interpret some of the character states in these keys, the user is urged to closely examine the illustrations corresponding to the descriptive statements. Such relative terms as "strong," "small," "inflated," etc. are understood more easily with figures. This also holds true for the male and female genitalia, both of which may be complex and therefore difficult to describe by words alone. To a certain extent, the Lutzornyia subgenera and equivalent species groups are defined by the characters given here. It is important to note, however, that species in some groups, presently unknox^n in Colombia, may not exactly "fit" into a subgenus or species group as characterized by these keys. For example, L. nevesi (Damasceno and Arouck) belongs in the verpueavwri group although the male, unlike the others, lacks a coxite setal tuft. This species is very common at Limoncocha, Napo Prov. , Ecuador — a locality not far from Putumayo Intendencia, Colombia.

PAGE 90

As additional species are discovered in the Republic, these keys obviously will have to be revised. For the present, they are intended to be used with caution, applicable only to the Colombian fauna. Males 1. Palp 5 very short, less than half length of third and less than twice length of fourth subgenus Psyahodopygus (Fig. 61-71). . 2 Palp 5 subequal to or longer than half length of third and greater than twice length of fourth 4 2. (1) Style with 1 large terminal spine and 3 inconspicuous subapical setae. Coxite with dorsal margin indented near middle series sqi'.amiventr-is (Fig. 61) Style with 2-6 major spines. Coxite not indented near middle ... 3 3. (2) Style with 2 major spines, both terminal. Paramere simple. . . series ar-ihirei, L. bisp-inosa (Fig. 63) Style with 3-5 (sometimes 6) major spines. Paramere simple or complex series panamensis (Fig. 64-71) 4. (1) Lateral lobes markedly inflated . . vsspertilionis group (Fig. 29) Lateral lobes not inflated. .... 5 5. (4) Lateral lobes with spatulate setae at tips . . . subgenus Evandromyia, series infraspincsa* "Species of Evandvorrvji.a in this series have not been found as yet in Colombia but it is probable that at least one species occurs in the southeastern part of the Republic. The subgenus was reviewed by Young and Arias (19 7 7) .

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•78Lateral lobes without spatulate setae 6. (5) Style deeply forked. Parameres with 2-4 modified apical setae as shown subgenus Viannamyia. (Fig. 35-35) Style simple, not forked. Parameres usually with simple setae but, if modified, then different from above 7 7. (6) Style with 2 major spines and 1 or 2 smaller accessory seta . . 8 Style with 3-6 major spines, smaller accessory setae present or not. 9 8. (7) Paramere with a dorsal setiferous arm. Coxite without nondeciduous setae •••.•J*.!., subgenus DoiTiV^oiwia L, vosciliciZi (Fig. 30) Paramere simple, without a dorsal arm. Coxite with a basal tuft or group of setae vevpv.cavwn group, series sewana (Fig. 22) 9. (7) Style with 3 major spines 10 Style with 4-6 major spines 11 10. (9) Coxite with a basal tuft of simple and modified setae. Style with a small subterminal and a small median seta in addition to the major spines subgenus Pressatia (Fig. 32-34) Coxite without a basal tuft but with a distal group of 154simple setae. Style with only a small median seta and 3 major spines, no subterminal seta pilosa group, L. pilosa (Fig. 86) 11. (9) Style with 4 major spines 12 Style with 5, sometimes 6, major spines 32 *Males of L. vovotaensis usually have 5 major spines but those examined from ValJe Dept., have but 4 (see p. 421).

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-79L2. (11) Antenna! ascoids with very long pointed or short bluut basal spurs. Style without a subterminal seta 13 Antermal ascoids with or without basal spurs, but if present they are very short and pointed. Style with or without a subterminal seta 14 13. (12) Coxite with 5-8 strong distal setae. Style with isolated basal spine clreisbacki group (Fig. 50-51) Coxite without distal setae. Style with basal spines more or less paired shannoni group (Fig. 37-42) 14. (12) Palp 5 shorter than palp 3. Style without a subterminal seta 15 Palp 5 as long as or longer than palp 3. Style with or without a subterminal seta 18 15. (14) Coxite with 2 or more basal and/or median setae 16 Coxite without nondeciduous setae 17 16. (15) Style with basal spine isolated. Antennal ascoids with very short pointed basal spurs. Flagellomere I shorter than f lagellomeres II + III subgenus Tviohophovorryia (Fig. 51-54) Style with paired basal spines. Antennal ascoids simple, no basal spurs visible. Flagellomere I longer than II + III . .subgenus NyssomyCa (in part), L. cmtunesi (Fig. 55)

PAGE 93

-8017. (15) Antennal ascoids with short, pointed basal suprs. Paramere with ? dorsobasal hump. Flagellomere I shorter than II + III avagaoi group (Fig. 45-49) Antennal ascoids simple. Paramere lacking a dorsobasal hump. Flagellomere I longer than II + III subgenus Nyssomyia (in part), (Fig. 55-60) 18. (14) Ccxite with 3+ basal and/or median setae in a tuft or group, d-istal setae may also be present 19 Coxite without a basal and/or median group of setae but long ventral ?et_ci£ llldy Dtr jJj.toCn>_ «.o 19. (18) Style with 4 short stubby spines, 2 of which are terminal; subterminal seta present. Paramere simple, undivided. Coxite with 4-8 strong distal setae baityi group, L. haityi (Fig. 34) Style with longer spines; if 2 are terminal then parameres are divided; subterminal seta present or absent. Coxite without distal nondeci.dtious setae 20 20. Hind femur with a row of 3-5 short spines subgenus Fintomyia, L. spinosa (Fig. 31) Hind femur without spines 21 21. (20) Paramere with 2 dorsobasal setae, much longer than others, hooked or fan-shaped subgenus Lutzomyia, series longipalpis (Fig. 13-15) Paramere without such setae 22

PAGE 94

22. (21) Style with a subterminal seta 23 Style without a subterminal seta 26 23. (22) Genital filament tips enlarged, spoon-shaped, each with a distinct inner "tooth" . . . migonei group, series walkevi (Fig. 19-20) Genital filament tips enlarged or not but lacking an inner tooth. 24 24. (23) Coxite with setae of basal tuft shorter than width of coxite migonei group, L. migonei (Fig. IS) Coxite with setae of basal tuft longer than width of coxite ... 25 25. Mesonotum & pleura pale . . saulensis group,* L. saulensis (Fig. 21) Mesouotum distinctly darker than pleura .... verrucarum group, series verrucar-ion (in part), (Fig. 23-24) 26. (22) Coxite with basal setae inserted on a raspberry-like, subcircular base subgenus Lutzomyia, series oruaiata (Fig. 16-17) Coxite with basal setae, if present, not implanted on suchabase. 27 27. (26) Style with paired basal spines. Coxite with basal tuft of slender, subequal setae . verrucarum group, series verrucavum (in part), L. nuneztovaTi (Fig. 23) Style with basal spine isolated. Coxite with either basal-median setae, 1 at least larger than others, or with a patch of slender distal sotae cayennencis group, series atvoclavata (Fig. 81-82) V T am unable to separate the males on the group level using structural characters .

PAGE 95

28. (18) Style with a small sub terminal seta 29 Style without a small sub terminal seta 31 29. (28) Paramere divided. Coxite with long ventral setae loagi spina group (Fig. 43-44) Paramere simple, undivided. Coxite without long ventral setae. . 30 30. (29) Palp 5 shorter than segments 3+4. Eyes large. Paramere slender, its width less than that of style ungrouped, L. sp . of Anchicaya (Fig. 89) Palp 5 longer than segments 3+4. Eyes small. Paramere broad, its width greater than that of style, ungrouped, L. ronyeliana (Fig. 88) 31. (28) Style with isolated, basal spine. Flagellomere I longer than head height; ascoids with very short basal spurs. Palp 5 shorter than segments 3 + 4 ungrouped, L. -aovdestina (Fig. 87) Style with paired basal spines. Flagellomere I shorter than head height; ascoids without visible basal spurs. Palp 5 longer than segments 3 + 4 . . • cayennensis group, series ccrjennsnsi-s (Fig. 79-80) 32. (11) Style with a small subterminal seta oswaldoi group (in part) , L. pia (Fig. 83) Style without a small subterminal seta 33 33. (32) Coxite with 2 or more setae at base; others may extend to middle of structure forming a loose or compact tuft vexatov group, series per-uens'is (Fig. 72-78)

PAGE 96

Coxite without setae at base but with a few scattered setae near middle of coxite oswa.Zdoi group (in part), (Fig. 84-85) Females 1. Spermathecae associated with paired sclerotized stiuctures, intracellular ducts (= "hairs") covering most of spermathecae subgenus Viannar.tyia (Fig. 35-36) Spermathecae without associated sclerotized structures, intracellular ducts confined to small area, usually but not always, to terminal knob 2 2. (J) Pharynx with posterior spines 3 Pharynx without spines 6 3. (2) Cibariuir. with inner pair of horizontal teeth pointing inwards OSWdldoi group (in part), L. trinidadsnsis (Fig. 85) Cibarium with horizontal teeth pointing toward pharynx, not slanted 4 4. (3) Cibarium with 10+ horizontal teeth in a comb-like row . . cayennsis group, series cayennensis (in part),/,, cayennensis (Fig. 79) Cibarium with 4 horizontal teeth not in a comb-like row 5 5. (4) Foreleg with femur longer than tibia. Pharynx markedly enlarged with prominent, subequally spaced transverse ridges and conspicuous spines. Pleura pale cayennensic group, series atroolavata (Fig. 81-82)

PAGE 97

Foreleg with femur shorter than tibia. Pharynx nore slender, the transverse ridges and spines less defined. Pleura dark oayettnensi-s group, series oayennensis (in part), L. micvopyga (Fig. 80) 6. (2) Spermathecae with bubble-like expansions 7 Spermathecae without bubble-like expansions 8 7. (6) Cibariuni with 4 broad and blunt horizontal teeth and conspicuous lateral teeth. Mesonotum well pigmented, darker than pleura. Eyes very small subgenus Darrtpfomyia f L. vosdbali (Fig. 30) Cibarium with 4 slender, sharp horizontal teeth, without obvious lateral teeth. Mesonotum and pleura pale. Eyes larger . . . . saulensis group, I. sav.lensis (Fig. 21) 8. (6) Antennal ascoids with long pointed or short blunt basal spurs . 9 Antennal ascoids simple or with short pointed basal spurs 10 9. (8) Cibarium with 10+ horizontal teeth. Spermathecae distinctly aimulated dreisbadki group (Fig. 50-51) Cibarium with 4-8 horizontal teeth. Spermathecae annulated or not . . . . shannonigroup (Fig. 37-42) 10. (8) Hind femur with row of 3-5 short spines .... .subgenus Pintomyia, L. spinosa (Fig. 31) Hind femur without spines 11 11. (10) Cibarium with 6 or more horizontal teeth 12

PAGE 98

-85Cibarium with 4 horizontal teeth 17 12. (13) Individual sperm ducts markedly convoluted, twisted . , . . . .avagaoi group, series, bras Hi ens is, L. runoides (Fig. 49) Individual sperm ducts not convoluted as above 13 13. (12) Palp 5 equal to or shorter than segments 3 + 4 14 Palp 5 longer than segments 3 + 4 16 14. (13) Spermathecae larger, sac-like, without annuli. Ascoids with short pointed basal spurs, avagaoi group, series aragaoi (Fig. 45-48) Spermathecae smaller, with incomplete or complete annuli. Ascoids with or without short basal spurs 15 15. (14) Flagellomere I longer than f lagellomeres II + III. Ascoids with short basal spurs subgenus THohophoromyia (Fig. 51-54) Flagellomere I shorter than II + III. Ascoids without visible basal spurs subgenus Nyssomyia (Fig. 55-60) 16. (13) Cibarium with 6 horizontal teeth. Spermatheca with terminal segment smaller than preceding segment. Common sperm duct longer than individual ducts . .OSWaldoi group (in part), L. pia* (Fig. 83) Cibarium with 8+ horizontal teeth. Spermatheca with terminal segment as large as others. Common sperm duct much shorter than individual k L. via is keyed twice because of intraspecif ic variation in the number of horizontal teeth (see p. 419).

PAGE 99

-86ducts subgenus Lutsomyia, series long ip alp is (in part), L. longipalpis (Fig. 15) 17. (11) Palp 5 ahorter than segments 3 + 4 18 Palp 5 longer than segments 3 + 4 21 18. (17) Spermathecae imbricated, annuli semi-telescoped. Individual sperm ducts heavily sclerotized, wrinkled, or rugose in part or whole subgenus Psychodopygus (Fig. 61-71) Spermathecae not imbricated, annuli, if present, not semitelescoped. Individual sperm ducts non-rugose, smooth walled or with slight thickenings 19 19. (18) Cibarium with inner pair of horizontal teeth pointing inwards. Flagellomere I extremely long, subequal to or longer than combined lengths of palpal segments. Ascoids with very short, basal spurs. . ungrouped, L. nord.es tin 'a (Fig. 87) Cibarium with horizontal teeth pointing toward pharynx, not slanted inwards. Flagellomere I shorter than palpus. Ascoids without visible basal spurs -0 20. (19) Spermathecae as shown, longer than individual sperm ducts. Abdominal tergite 8 without lateral setae .... vespevtilionis group, series vespertilionis (Fig. 29) Spermathecae otherwise, shorter than individual sperm ducts. Abdominal tergite 8 with lateral setae 21

PAGE 100

-8721. (20) Spermathecae less than twice as long as wide, terminal segment smaller than preceding segment. Common sperm duct longer than individual ducts oswaldoi group (in part), L. pia (Fig. S3) Spermathecae cylindrical, its length at least 3 times width, terminal segment subequal to or smaller than others. Common sperm duct shorter: than individual ducts . . . vexatoP group, series perv.ensis (in part), (Fig. 72-74; 76-78) 22. (17) Individual sperm ducts heavily pigmented in part or whole. . . subgenus Prsssatia (Fig. 32-34) 23. (22) Cibarial arch absent or conspicuous only at sides 24 Cibarial arch complete or nearly so 26 24. (23) Cibarium with horizontal teeth like flattened plates viewed almost edgewise. Common sperm duct absent pilosx group, L. pilose. (Fig. 86) Cibarium with horizontal teeth otherwise. Common sperm duct present ?.j 25. (24) Wing venation with beta less than half alpha. Cibarium with horizontal teeth pointing tox^ards pharynx, not inwardly slanted . . vexator group, series per-uensis (in part), (Fig. 75) Wing venation with beta over half length of alpha. Cibarium with horizontal teeth pointing inwards oswaldoi group (in part), L. rofotaensis (Fig. 100)

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-8826. (23) Spermathecae completely smooth-walled 27 Spei.raathecae wrinkled, with transverse striations or indentations forming complete annul! or not 28 21, (26) Spermathecae tubular, much longer than wide and only slightly wider than individual ducts. Common sperm duct short, not exceeding onethird length of individual ducts migonei group, series migonei, L. migonei (Fig. 18) Spermathecae capsular, about as long as wide, distinctly wider than individual ducts where they enter spermathecae. Common duct over onethird length of individual ducts .... migonei group, series walkeri (Fig. 19-20) 28. (26) Spermathecae as shown, with some clearly defined annuli, terminal one hemispherical and larger than others, .subgenus Lutzomyia, series longipalpis (in part), and series cruciata (Fig. 13-14; 16-17) Spermathecae otherwise, with transverse striations or wrinkles, terminal segment, if present, not hemispherical 29 29. (28) Spermathecae as shown, pear-shaped with fine transverse striations, complete or not longispina group (Fig. 43-44) Spermathecae otherwise, saclike and wrinkled ..... 30 30. (29) Individual sperm ducts nearly absent, spermathecae seemingly joxned directly to common duct. Eyes very small. . . ungrouped, L. r-angetiana (Fig. 8S) Individual sperm ducts at least as long as half length of spermathecae. Eyes large vewucavion group (Fig. 24-27)

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Figure 12 Lutzomyia (P.) hirsuta nioaraguensis female — A. Head with palpal segments marked in Arabic numerals and flage Homer es designated by Roman numerals, B. Flagellomere II showing ascoids, C. Pharynx attached to cibarium, D. Metathoracic spiracle and postspiracular setae, E. Cibarium, F. Spe.rmathecae and associated structures, G. Wing, H. Femur, tibia, and basitarsus of front leg. Female: Rio Changena, Eocas del Toro Prov. , Panama Lutzomyia townsendi male — I. Male genitalia; small subterminal seta on sryle is present, J. Genital pump and filaments, sane scale as Fig. 121. Male: Rancho Grande, Aragua State, Venezuela

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-90S : II H Jim .-« i \ , fl--J/-w its ?::;' A Kv7A\. lnd;>:duol Sp-rm Ouct ss: '/7 >// .1 H U \\\ C-r..'al FilniKn!

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•91 Subgenus Lutzomyia _F r anca. , 1 924 This subgenus, divided into 3 series (Lewis et al., 1978), contains 21 species including L. mafinkcllei n. sp. described in this review. Sever, of these are anthropophilic and others probably share the £ ime habit. Lutzomyia longipalpis , the principal vector of visceral leishmaniasis in the New World (Lewis, 1974), ranges from southern Mexico to central Argentina where it is locally common in dry, nonforested areas. Lutzomyia gomezi, also reported in Colombia, has been implicated as a vector of dermal leishmaniasis in Panama (Christensen et al., 1969). In Colombia the series longipalpis is represented by 3 species — L. longipalpis, L. liahyi, and L. bifoliata. Two species, L. gomezi and L. marinkellei are included in the series aruoiata. Except for longipalpis, the females of these species are remarkably similar and are separated only by minor nonsexual characteristics. Keys to S pecies Males 1. Paramere with long dorsobasal arm bearing apical setae. Coxite with 2 very broad fan-like setae at inner bass 2 Paramere without dorsobasal arm. Coxite with 4 or more simple setae at inner base. . 3 2. Paramere with dorsobasal arm bearing 2 slender, recurved setae. Coxite with patch of long setae distad of fan-like setae bifoliata (Fig. 13)

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Pararaere with dorsobasal arm bearing 2 fan-like setae. Coxite without a distal patch of long setae lichyi (Fig. 14) 3. Paramere with 2 strongly developed, dorsobasal setae. Coxite tuft of 4 setae. Style with a subterrninal seta. . . longipalpis (Fig. 15) Paramere without, such setae. Coxite tuft of 10+ setae. Style without a subterminal seta 4 4. Pleura entirely pale. Style over half length of lateral lobe. Paramere with dorsal setae on distal half of structure, .gomezi (Fig. 16) Pleura mostly dark. Style shorter than half length of lateral lobe. Paramere with short dorsal setae confined to distal third of structure marinkellei (Fig. 17) Females 1. Cibarium with at least 8 horizontal teeth. Spermathecae as shown, terminal annulation not markedly larger than others longipalpis (Fig. 15) Cibarium with 4 horizontal teeth. Spermathecae otherwise, terminal annulation spherical much larger than others 2 2. Pleura pale, contrasting with dark mesonotum. Flagellomere I shorter than or equal to length of labrura gomezi (Fig. 16) Pleura mostly dark. Flagellomere I longer than labrum 3 3. Flagellomere I less than 0.32 mm long marinkellei (Fig. 17) Flagellomere 1 greater than 0.32 mm long 4

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4. Terminal flagellomere longer than preceding f lagellomere (XV) . . . lichyi (Fig. 14) Terminal flagellomere shorter than preceding flagellomere (XV) . . . bi.foZia.ta (Fig. 13) Series longipalpis 12. Lutzomyia (L.) bifoliata (Fig. 13) Lutzomyia bifoliata Osorno, Morales, Osorno, & Hoyos, 1970: 8 (c?, El Terminal, Municip. Puerto Boyaca, Boyaca, Colombia). Osorno et al., 1972a: 15 (listed). Forattini, 1973: 248 (gen. review, 6" fig.). Distribution: Colombia (Antioquia, Boyaca) . Material examined: Colombia. 13 <$d, 19 o, Rio Anori (Antioquia), tree buttresses, Sept. 1970, D.G.Y. 3 6*0", 1 cp , light traps, same locality May 1970, C.H.P. Discussion'. Lutzomyia bifoliata males from Rio Anori generally agree with the description and figures given by Osorno et al. (1970) but the strong erect seta on the dorsum of the paramere is lacking in our material. The female, described below, closely resembles L. lichyi but the terminal flagellomere is shorter, not longer than the preceding flagellomere. The longer labrum of bifoliata females (0.37-0.42 mm as opposed to 0.25-0.36 mm, n = 17, in lichyi) also may be useful in distinguishing the species if it proves to be a consistent, nonoverlapping feature . The following description is based on 10 females captured near the Rio Anori .

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-94Female: Wing length 2,35-2.57; width 0.73. Head, mesonotum, and abdominal tergites strongly pigmented: Rest of insect, including most of pleura, paler. Head height 0.46; width 0.44. Eyes separated by 0.14 or distance = to ca . 7.8 facet diameters. Flagellomere I (0.40-0.50 long), 1.2 x length of II + III; ascoids longer than those of 6", but not reaching ends of flagellcmeres , with almost indiscernible posterior spurs, present on all but last flagellomere. Length of palpal segments: 1 (0.06) 2 (0.15-0.18), 3 (0.19-0.21), 4 (0.09-0.12), 5 (0.30-0.37); palpal sensi 11a on segments 2 and 3. Labrum length = 0.37-0.42. Cibarium with 4 nearly straight, sharp horizontal teeth and 20-30 vertical teeth distributed as shown; chitinous arch complete, well defined; pigment patch sub triangular, darker, and broader posteriorally . Pharynx (0.2C long) unarmed. Pleura with 14-22 upper and 3-5 lower episternal setae. Length of wing vein sections: Alpha. (0.64-0.71), beta (0.27-0.34), delta (0.13-0.19), gamma (0.27-0.34). Length of femora, tibiae, and basitarsi of slide 618: Foreleg, 0.95, 1.22, 0.74; midleg, 0.91, 1.47, 0.81; hindleg, 0.98, 1.67, 1.27. Abdominal sternite 2 horseshoe-shaped, hollow in middle. Tergite 8 without setae. Spermathecae and sperm ducts as figured. 13. Lutzonyia (L.) lidhvi (Fig. 14) Phlebotomus lichyi Floch & Abonnenc, 1950a: 1 (o holotype, Rio Borburata, Carabobo, Venezuela). Morales et al., 1969a: 381 (Meta, Colombia) . Phlebotomus VexillaHus Fairchild & Hertig, 1952: 514 (6*, o, Panama) Plfano et al., 1960: 65 (Miranda State, Venezuela). Pifano et al.,

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-951962: 385 (keyed). McConnell & Correa, 1964: 527 (infected with fungi/. Flcch & Kramer, 1965: 1 (as synonym of lichyi) . Hanson, 1968: 90 (larva & pupa). Scorza et al., 1968: 35 (bionomics). Calderon, 1973: 87 (Merida State, Venezuela). PhZebotomus foliatus Mirsa & Ortiz, 1952: 249 (
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-96Disoussion: Man biting records for this species include those from Panama (Fairchild and Hertig, 1952), Venezuela (Scorza et al., 1968), and Colombia (Osorno et al . , 1972a). During the interoceanic canal survey in Choco Dept. at Alto Curiche, we collected only 4 females on human bait during a 5 month period. Each was a daytime catch although routine biting collections were carried out at night as well. Dr. Pablo Barreto (pers. com.) has collected biting females within the city limits of Gali, Colombia. In Panama, McConnell (1963) found non-mammalian (nucleated) red blood cells in the gut of a resting female and Thytcher and Hertig (1966) captured females feeding on Didelphis and Potos . According to Hanson (1968) , the larvae of L. lichyi (as veorillarius) and L. gomezi share certain characters not observed in other species. This, along with adult morphology, supports the placement of L. gomezi in the subgenus Lutzomyia. 14. Lutzomyia (I.) longipalpis (Fig. 15) PhUbotomus longipalpis Lutz & Neiva, 1912: 90 (
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Material examined: Colombia. 1 6*, 2 op, Baraya (Huila) , rock T T crevices, March 1945, M. Hertig. Venezuela. 7 do*, 13 oo, near Calabozo (Guarico), outside wall of house, 17 July 1965, D.G.Y. Bolivia., Brazil, Costa Rica, El Salvador, Mexico, it Pavagvwj . Numerous specimens from various localities within these countries. Discusc-ion: Lutzormjia longipalpis, first reported in Colombia by Osorno et al. (1969), has received much attention because of its role as a vector of kala-azar in the New World. It would be redundant to cite all important references to this species, the type species of the genus, because Forattini (1973) has recently reviewed most of them. Fhlebotomus otarnae Nunez-Tovar, 1924, and Fhlebotomus almazani Galliard, 1934, are well established junior synonyms of L. longipalpis. Series cvuoiata 15. Lutzomyia (L.) gom&zi (Fig. 16) Fhlebotomus gomezi Nitzulescu, 1931: 247 (^, San Cristobal, Tachira State, Venezuela). Barretto, 1947: 202 (full references). Fairchild & Hertig, 1948a: 252 (taxonomy). Fairchild & Hertig, 1953b: 382 (6\ o, redescr., refs., figs.). Hanson. 1968: 60 (larva & pupa). Barreto, 1969: 464 (Meta & Valle, Colombia). Morales et al . , 1969a: 378 (Meta, Colombia) . Fhlebotomus suis Rozeboom, 1940: 8 (6*, o, Panama). Barretto, 1946a: 1 (as synonym of gomezi). Fairchild & Hertig, 1948a: 252 (as synonym of gomezi) . Fhlebotomus japigmji Floch & Abonnenc, 1944b: 2 (6*, o, French Guiana). Fairchild & Hertig, 1948a: 252 (as syonoym of gomezi) .

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-98Lutzomyia gomezi: Martins et al., 1962b: 90 (cf. to L. gasparviannai) . Osorno et al., 1972a: 19 (Colombian records). Forattiru , 1973: 240 (gen. review). Lewis, 1975a: 500 (mouthpart morphol.). Interiflill & Muller, 1976: 543 (Honduras). Miles et al. , 1976: 531532 (mating behavior). Ward & Ready, 1975: 128 (egg, figs.). Zimmerman et al., 1977: 575 (egg, figs.). Distribution: El Salvador to southern Brazil. Colombia (Amazonas, Antioquia, Bolivar, Boyaca, Caldas, Caqueta, Cesar, Guajira, Huila, Magdalena, Meta, Norte do Santander, Santander, Tolima, Vichada) . Material examined: Colombia. 19 33, 14 oo, Curiche (Choco) , Malaise trap, May-Nov. 1967, D.G.Y. 23 c^, same data but biting man. 2 33, same data but on tree trunk, 4 April 196 7. 1 o, Sautata, Rio Atrato (Choco), Malaise trap, 20 Jan. 1968, D.G.Y. 8 33, 3 05, Rio Anori (Antioquia), light traps. May 19 70, C.H.P. 1
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dermal leishmaniasis in that country (Schneider and Hertig, 1966; Christensen and Herrer, 1973) . Forattinl (1973) provides additional references and information on the biology and disease relationships of L. gomezi. 16, Lutzomyia (L.) marinkellei n. sp. (Fig. 17) Male: Wing length 1.7; width 0.42. Head, mesonotum, abdominal tergites, and gentialia strongly pigmented; rest of insect, including most of pleura, moderately pigmented. Head height 0.34; width 0.33. Eyes large, separated by 0.09 or by distance = to 5.3 facet diameters. Flagellomere I (0.33 long), 1.1 x length of II + III; ascoids with very shore posterior spurs, their distal tips ending before apical third of flagellomere 11, on all flagellomeres except last 2. Length of palpal segments: 1 (0.04), 2 (0.12), 3 (0.15), 4 (0.12), 5 (0.32); palpal sensilla, fewer than 10, on middle third of segment 3. Labrum length 0.195. Cibarium unarmed except for vestiges of vertical teeth; chitinous arch complete, diffuse in middle; pigment patch subtriangular, more narrow th3n p. Pharynx length 0.146. Pleura with 19 upper and 5 lower episternal setae. Length of wing vein sections: Alpha (0.30), beta (0.21), delta (0.07), gamma (0.30). Legs missing but length of femora, tibiae, and basitarsi of paratype no. 631: Foreleg, 0.71, 0.70, 0.39; midleg, 0.71, 0.93, 0.A6; hindleg, 0.74, 0.98, 0.56. Abdominal sternite 2 horseshoe-shaped, the median opening not enclosed. Style short (0.13 long) with 4 strong spines as shown, no sub terminal bristle. Coxite stout (0.25 long x 0.10 wide at greatest width), basal tuft of 12-13 long setae on individual tubercles which form a raspberry-like base. Paramere

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-100 as shown, upturned apically, dorsal setae few and rather short, these and lateral setae restricted to apical third of structure, ventral setae on distal half. Aedeagus (ca. 0.11 long) slender apically. Lateral lobe (0.28 long), wider distally, perhaps due to mounting. Genital pump (0.126 long): each filament ca. 0.43 long or 3.4 x length of pump, rip simple, not significantly enlarged. Cercus as shown. Female: Wing length 1.85; width 0.57. Coloration same as 6". Head height 0.38. width 0.37. Eyes large, separated by 0.98 or distance = to 5.5 facet diameters. Flagellomere I (0.28 long), 1.2 x length of II + III; ascoids with short posterior spurs, longer than those of o" but not reaching end of flagellomere II, last 7 flagellomeres missing. Length of palpal segments: 1 (0.04), 2 (0.13), 3 (0.16), 4 (0.10), 5 (missing); palpal sensilla as in 6*. Labrum length 0.23. Cibarium with 4 sharp horizontal teeth, 11-15 small vertical teeth in a single row; chitinous arch complete, diffuse in middle; pigment patch as shown, darker and wider pcsterio rally . Pharynx length 0.17. Pleura with 19 upper and 7 lower episternal setae. Length of wing vein sections: Alpha (0.44), beta (0.25), delta (0.11), gamma (0.30). Length of femora, tibiae, and basitarsi of slide 632: Foreleg, 0.78, 0.71, 0.39; midleg, 0.75, 0.90, 0.46; hindleg, 0.83, 1.1, 0.56. Abdominal sternite 2 apparently open in middle as in o" but hardly noticeable. Tergii.e 8 lacking setae. Tergite 9 without dorsolateral papillate lobes. Spermathecae, sperm ducts and cerci as shown . Material examined: Colombia. 6* holotype (no. 628), Tres Esquinas (Caqueta) , light trap, 10 Nov. 1971, C.J. Marinkelle. o allotype (no. 629), same data. 1 c?, 1 o paratypes (no. 630, 633) same data. Brazil. 1 d", 1 o paratypes (nos. 631-632), near Rio Aripuana (Mato Grosso) , J . Arias .

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Discussion: LutsomyLa marinkeZlei is closely allied with L. evueiata (Coq.;, L. evangelistai Martins and Fraiha, L. gomezi, and L. shevZoaki Martins, Silva, and Falcao, all of v.'hich form the series oruoiata. From, these, the male, of ravinksllei is separated by the following combination of characters: Both rnesonotum and pleura infuscated. Style short, its length less than or equal to half the length of lateral lobe. Paramere with few dorsal setae, these and lateral setae confined to apical third of structure. Coxite tuft of fewer than 20 setae. The dark pleura differentiates the female from evangelistai, gornezi, and shevlocki — species with pale pleura which are (or may be sympatric with mavirikeZlei , The status of L. evueiata and the form described as Pklehotonrus diaboticus Hall is still uncertain but the latter taxon is probably a subspecies of evueiata as treated by Lewis (1975a) . The females of moj'inkellei and c. diabolica, unlike e. evueiata, lack papillate lobes of tergite 9 and have well pigmented pleura. It would be difficult to separate these females in the absence of males were it not for the fact that e. diabetica occurs well outside the range of marinkeZlei in Mexico and Texas . Tiie length of the first f lagellomere, the character used in the key to separate mavinkellei from females of lickyi and bifoZiata, may prove to be variable as a diagnostic character. With these taxa specific identification should be based on associated males. I am pleased to name this species after Dr. C.J. Marinkelle who was most helpful in providing material for this study.

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Figure 13 Lutzontyia (L.) hifoliata — A. Male head, B. Male flagelloraere II. C. Female head, D. Female flagellomere II, E. Female cibarium and pharynx, F. Dorsal arm of paramere, G. Genital pump and filaments, H. Male genitalia, I. Spermathecae, J. Female wing, K. Male wing, l>. Female cibarium. Male: Rio Anori, Antioquia Dept., Colombia Female: Same locality as male

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103-

PAGE 117

Figure 14 Lutzonvjia (L.) liohyi — A. Male head, B. Male flagellomere II, C. Female head, D. Female flagellomere II, E. Female cibarium and pharynx, F. Spermachecae, G. Female cibarium, H. Female wing, I. Male v/ing, J. Male genitalia. Male: Alto Curiche, Choco Dept., Colombia Female: Curiche, Choco Dept., Colombia

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-105-

PAGE 119

Figure 15 Lutzormjia (L.) longipalyis — A, Male head, B. Male flagellomere II, C. Feraale head, D. Female flagellomere II, E. Female cibarium and pharynx, F. Female wing, G. Male wing, H. Genital pump and filaments, I. Male genitalia, J. Body of spermatheca, K. Spermathecae, L. Genita] pump, same scale as Fig. 151, M. Paramere of male from Huila Dent., Colombia, same scale as Fig. 151, N. Female cibarium. Male: Calabozo, Guarico State, Venezuela Female: Same locality as male (except for Fig. 15M)

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-107-

PAGE 121

Figure 16 Lutzorwjia (L.) gomezi — A. Male head, B. Male flagellomere II, C. Female head, D. Female f lagellomere II, E. Female cibarium and pharynx, F. Male genitalia, G. Spermathecae, H. Female wing, I. Male wing, J. Female cibarium. Male : Trinidad Female: Trinidad

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Figure 17 Lutzomyia (L.) mapinkellei — A. Spermathecae, B. Female wing, C. Male. wing, D. Female flagellomere IT, E. Base of ascoid, F. Female head, G. Female cibarium, H. Male head, I. Male genitalia. Kale: Tres Esquinas, Caqueta Dept., Colombia Female; Rio At"ipuana, Mate Grosso -State, Brazil

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-111V

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-112Migonei Group Th eodor, 1965 This group is divided into 3 series following Lewis et al. (1978). The series rrrCgonei , walkeri, and cGstalimai (= subgenus Boj.'rettomyia Martins and Silva, 1965) . Species in the latter series occur in Brazil but some of those in the other series are more widespread with 4 species occurring in Colombia. iSarretto (1562) and Forattini (1971a, 1973) include some of the mi-gonei group taxa in the subgenus Coromyia Barretto, 1962. Based on present knowledge, the only man-biting species in this group is L. migonei, here reported in Colombia for the first time. Keys to Species Males 1. Genital filaments over 3 x length of pump, with simple tips. Coxite with basal tuft of 4-6 short setae migonei (Fig. 18) Genital filaments less than 3 x length of pump, their tips spherically enlarged with an inner "tooth." Coxite with basal tuft of 10+ longer setae and a median ventral group of 6+ setae 2 2. Aedeagus simple, without a dorsal projection. Genital filaments 0.30 mm or longer mava^oensis (Fig. 19) Aedeagus complex, with a dorsal cylindrical projection. Genital filaments less than 0.30 mm loalkeri (Fig. 20)

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113Fema las 1. Spernathecae narrow and tubular, at least 3 x longer than wide. . . migonei (Fly. 18) Spermathecae capsule-shaped, less than 2 x as long as wide 2 2. Individual sperm ducts over 3 x length of common duct, wider and with fine transverse striations basally sp.de Baduel (Fxg. 18) Individual sperm ducts about 1/2 as wide as common duct r.arajoensis (Fig. 19) Individual sperm ducts more slender, about 1/3 as wide as common duct walkevi (Fig. 20) Series miaonei 1 7 . Lutzomyia migonei (Fig. 18) Phlebotomy^ migonei Franca, 1920: 230 (c? holotype, Assuncion, Paraguay). Barretto, 1947: 211-213 (full refs., synonyms). Forattini, 1954: 214 et seq. (sternites, figs.). Forattini, ]960: 479 (Amapa, Brazil). Pifano et al., 1962: 387, 388 (6*, o, keyed). Calderon, 1973: 87 (Merida State, Venezuela) . Lutzomyia migonei: Theodor, 1965: 182 (cf, o, figs.). Forattini, 1973: 122 et seq. (gen. review, refs., immatures, figs.). Lewis, 1975a: 500 et seq. (mouthpart morphol . ) . Distribution: Colorrbia (Magdalena) , Venezuela, Trinidad, Brazil, Paraguay, Argentina.

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-114Material examined: Colombia. 2 SJ', Rio Don Diego, E of Santa Marta (Magdalena), tree trunks, 15 Aug. 1973, D.G.Y. & R.C.W. Brazil. 2 3$, Laphina (Ninas Gerais) , flight trap near cave, 28 Aug. 1974, D.G.Y. & P. Williams. Trinidad. 2 6*6', Bush Bush Forest, Nariva Swamp, chickenbaited trap, 17-18 May 1961, T.H.G. Aitken. 1 o_, same data but 22-23 Kay. Discussion: Ranging from Argentina to northern Colombia, L. nrigonei has been found naturally infected with flagellates of uncertain identity in Venezuela and Brazil (Johnson et al . , 1963; Forattini, 1973). Other information dealing with the biology of this anthropophilic species was reviewed by Forattini (1973) and need not be repeated here. Series walkeri 18. Lutzomyia mavajoensis (Fig. 19) Pklebotomus marajoensis Damasceno & Causey, 1944: 339 (6* holotype, Ilha do Marajo, Brazil). Fairchild & Hertig, 1961b: 250-254 (cf, o, redescr., figs., refs., distrib.). Pifano et al., 1962: 386 et seq. (
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-115Pklebotomus dubitans Sherlock, 1962: 324 (6* hoiotype, San Vicente de Chucuri, Santander, Colombia). Osorno eU al., 1967: 28 (mention). Lutzormfia dubitans: Theodor, 1965: 182 (listed). Lewis, 1967b: 131 (mention). Forattini, 1971a: 103 (listed). Osorno et al . , 1972a: 23, 78 (mention). Martins & Morales, 1972: 366 (listed). Forattini, 1973: 292 (as synonym of walkeri and marajoensis) . Distribution: Panama, Colombia (Boyaca, Huila, Santander), Brazil, Venezuela, Trinidad. Material examined: Colombia . 1 o", Soledad, San Vicente de Chucuri (Santander), chicken coop, 16 Aug. 1944, A. Cast, E. Osorno, & 0. Mangabeira. Fanama, Venezuela, and Trinidad, those specimens listed by Fairchild & Hertig (1961b) . Discussion: A L. marajoensis male, the one listed by Fairchild and Hertig (1961b) from San Vicente de Chucuri, Santander, Colombia, was given to Corgas Memorial Laboratory by Dr. 0. Mangabeira in the 1940' s (Dr. G.B. Fairchild, pers . comm.). With some reservations, Sherlock (1962) later described L. dubitans based on other males taken at the same time and place as this male. Forattini (1973) treated both dubitans and marajoensis as junior synonyms of L. walkeri but only the first two are conspecific, the latter species being easily separated from marajoensis by the characteristics given in the key and by the shape of the parameres (cf . Figs. 19G and 20C) . Although L. walkeri appears to be more widely distributed than marajoensis , both species occur together in some localities in Colombia and Trinidad.

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-11619. Lutzorrpjic. sp. de Baduel (Fig. 18) Phlebotomies sp. de Baduel Floch & Abonnenc, 1945b: 1 (o, Baduel, French Guiana). Floch & Abonnenc, 1945c: 12 (o keyed). Floch & Abonnenc. 1952: 21 et acq. (o, keyed, redescr., figs.). Forattini, 1960: 480 (Amapa, Brazil) . Lutzotrvjia sp. de Baduel: Theodor, 1965: 196 (listed). Martins et al., 3 965: 4 (Rondonia & Maranhao, Brazil). Osorno et al . , 1972a: 66 (Vichada, Colombia) . Vistvibution: Colombia (Vichada), Brazil, French Guiana. Material examined: Colombia. 1 g (no. 652 1NPES) , Cumariana (Vichada), light trap, 15 May 1967, C.J.M. Discussion. The male of this informally named taxon remains unknown but it is certainly possible that it may be conspecific with one of the described males in the series, possibly L. sericea (Floch and Abonnenc) or L. williamsi (Damasceno, Causey, and Arouck) . The single known female from Colombia, seen by me, compares favorably with the description and figures given by Floch and Abonnenc (1945b, 1952). I have no reservations about placing it in the series walkeri of the migonei group based on spermathecae, cibarial structure, and palpi. 20 . Lutzomyia wdlkevi (Fig. 20) Phlebovomus walkevi Newstead, 1914: 188 (6*, o, Rio Abuna, BoliviaBrazil boundary). Barrctto, 1947: 230 (full refs.). Fairchild & Hertig, 1961b: 250, 254 (cf. to marajoensis) .

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•117Lutzomyia Walkeri: Barretto, 1962: 98 (listed). Martins et al., 1965: 4 (Rondonia, Brazil). Lewis, 1967b: 132 et seq. (cf, p, redescr., figs.). Forattini, 1971a: 101 (listed). Osorno et al . , 1972a: 23 (Colombian records) . Llanos, 1973: 31 (cf, g, redescr., figs.). Forattini, 1973: 274 et seq. (in part, gen. review, figs.). Lewis, 1975a: 500 et seq. (mouthpart morphol.). Llanos et al., 1975b: 671 (Peru). Phlebotomus gasti Sherlock, 1962: 326 (cf, San Vicente de Chucuri, Santander, Colombia). Osorno et al . , ]967: 28 (mention). NEW S 'YNON "I'M. Lutzomyia gasti: Theodor, 1965: 132 (listed). Lewis, 1967b: 131 (listed). Forattini, 1971: 103 (listed). Christensen, 1972a: 88-89 (Panama) Osorno et al . , 1972a: 23 (Colombian records). Forattini, 1973: 348 et seq. (gen. review, fig.). Distribution: Panama, Colombia (Antioquia, Caqueta, Santander), Ecuador, Peru, Brazil, Tri,nidad. Material examined: Colombia. 1 cf (no. 3924 INPES) , La Nevera , Solano (Caqueta), tree hole, 25 June 1969, A. Gast £< J. Ochoa. 3 oo , Tres Esquinas (Caqueta), light trap, 10 Nov. 1971, C.J.M. 1 cf, labelled P. gasti Mang. MS, no. 1120, Colombia (probably from the type locality of gasti). Brazil. 1 cf from type series, Abuna River, Bolivia-Brazil boundary, Dr. F.D. Walker, the Elders, Brixham, S. Devon. 2 6*d\ Labrea (Amazonas), tree trunk, 9 Oct. 1972. D.G.Y. 1 cf, 1 9, same data but light trap, 10 Oct. 1972. Ecuador. 1 o, Rio Napo at Limoncocha (Napo), flight trap, 22 May 1976, D.G.Y. & T. Rogers. Trinidad. 150+ specimens of both sexes from various localities to be discussed in a forthcoming paper. Discussion: Both Lewis (1967b) and Llanos (1973) carefully redescribed the male and female of L. walkeri, the former author studying

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-118tnater.ial from the type series; the latter basing her description on Peruvian specimens. I remounted a male from the type series, the one mentioned by Lewis (op. cit.), as being in the collection at Gorgas Memorial Laboratory but which is now at the University of Florida. I also examined a male from Santander Dept., Colombia which Mangabeira collected and which bears the name P. gasti MS. These two males are identical in all aspects including the presence of the dorsal projection of the aedeagus as figured by Sherlock (1962) for L. gasti. and by Llanos (op. cit.) for L. walkeri . I am convinced therefore that L. gasti is a junior synonym of L. walkevi . I should point out that the dorsal projection of the aedeagus is lightly sclerotized, less so than Sherlock (1962) depicts, and this probably is the reason why Lewis (op. cit.) did not observe this character state in the specimens at the British Museum (Nat. Hist.) . I consider the Ecuadorian female to be conspecific with walker-i based on morphology and distribution.

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Figure 18 Lutzorrtyia mtgonei — A. Male genitalia, B. Female cibarium, C. Spermathecae . Male: Trinidad Female: Trinidad 'jutzomyia sp. de Baduel female — D. Cibarium, E. Spermathecae , Female: Cumariana, Vichada Comisaria, Colombia

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A .^ S£*22«j> cM >*-^, ~\\ \ [\ i l 7/ R|

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Figure 19 Lutzowyia marajoensis — A. Male head, B. Male f lagellomere II, C. Female head, D. Female f lagellomere II, E. Female cibarium and pharynx, F. Body of spermatheca, G. Male genitalia, H. Genital pump, I. Genital pump and filaments, J. Tips of genital filaments, X. Spermathecae, L. Female wing, M. Male wing, N. Female cibarium. Male: San Jose, Los Santos Prov. , Panama Female: Puerto Mensabe, Los Santos Prov., Panama

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Figure 20 Lutzomyia walkevi — A. Spermathecae, B. Female cibarium, C. Male genitalia, D. Tip of genital filament, greatly enlarged. Male: Trinidad Female r Trinidad

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-124( J \ \ 1® \ w i 4 i n A vs IV'X /v;;.m

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-125So-'.lcnsis Group Lewis et. al. , 1978 This group was created to accomodate two closely related species -L. saulensis and L. wilsoni (Damascene) and Causey). Forattini (1971a, 19/3) places the former species in the subgenus Dampfomyia Addis but the spermathecae are structurally different (Theodor, 1965), the male parameres lack a dorsal arm, and the style has no subterminal bristle. The females of saulensis and wilsoni are nearly identical (Vianna Martins, pers. comm.), their populations probably overlapping in northern Brazil. 21 . Lutzomyia saulensis (Fig. 21) Phlebotomus saulensis Floch & Abonnenc, 1944a: 11 (6* holotype, Saul, Haute Mana, French Guiana). Barretto, 1947: 222 (listed). Damasceno et al., 194'?: 831 (Brazilian records). Floch & Abonnenc, 1952: 35, 112114 (6* keyed, redescr., figs.). Fairchild & Hertig, 1958b: 204-205 (refs., taxonomic discussion). Fairchild & Hertig, 1959: 122 (distrib.). Sherlock, 1962: 321, 330 (Santander, Colombia). Pifano et al., 1962: 386, 389 (o*, o, keyed, Venezuela). Osorno et al . , 1967: 28 (mention). Phlebotomus pinealis Floch & Abonnenc, 1944a: 11 (cj> holotype, Crique Anguille, French Guiana). Fairchild & Hertig, 1958b: 204-205 (refs. as synonym of saulensis). Johnson &. Hertig, 1961: 765, 774 (rearing attempt). Hanson, 1968: 69 (1st instar larva, descr., fig.). Lutzonniia saulensis: Martins et al., 1963: 335 (Roraima, Brazil). Martins et al . , 1965: 3 (Rondonia, Brazil). Theodor, 1965: 196 (o*, g, figs.). Barreto, 1969: 466-467 (Valle, Colombia). Lewis et al., 1970: 215 (parous study). Fraiha et al . , 1970b: 215 (mention).

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-126Christensen & Fairchild, 1971: 301 (Panama), Chaniotls et al., 1971a: 344 et seq. (pop. dynamics, Panama). Osorno et al., 1972a: 66 (Colombian records). Christensen, 1972a: 88 (listed). Chaniotls et al., 1972: 95 et seq. (resting sites). Shaw & Lainson, 1972: 710 et seq. (infected with nonlelshmanial flagellates, collecting data, Brazil). Llanos, 1973: 34 (6*, o, redescr., figs., Peru). Forattini, 1973: 327 et seq. (gen. review, figs.). Christensen & Herrer, 1973: 579 (listed). Ward et al., 1973: 178 (attracted to rodents and man, Brazil). Lainson et al . , 1973: 190 (negative for Leishmanial . Lewis, 1975a: 504 et seq. (mouthpart morphol.). Llanos et al., 1975b: 6 71 (Peru). Llanos et al., 19 76: 480 (Peru). Martins et al., 1976b: 496 (Peru). Distribution: Costa Rica, Panama, Colombia (Antioquia, Caqueta , Choco, Santander, Valle) , Peru, Brazil, French Guiana. Material examined: Colombia. 2 op, Rio Anori (Antioquia), light traps, 3 May 1970, C.H.P. 8 d'o", 11 oo, same data but some in tree hollows, Sept. 1970. D.G.Y. 14 6*d\ 55 ^o, Curiche (Choco), light, Shannon & Malaise traps, tree trunks, May-Dec. 1967, D.G.Y. 3 og, Alto Curiche (Choco), Malaise & light traps, Aug., Spet., Nov. .1967, D.G.Y. 1 g, Teresita (Choco), Malaise trap, 22 April 1967, D.G.Y. 4 ^o, Rio Atrato at Sautata (Choco), Malaise traps, Nov. -Dec. 1967, Jan. 1968, D.G.Y. 2 cfo", Anchicaya Dam (Valle), tree trunks, 9 Aug. 1973, D.G.Y. & R.C.W. 1 6", same data but light trap, 10 Aug. 1973. 2 o^ , 25 km E of Buenaventura (Valle), flight trap, 12 Aug. 1973, D.G.Y. & R.C.W. Brazil. 1 6", Belem (Para), 1 Feb. 1944, R. Damasceno & 0. Causey. Discussion: Although widely distributed in lowland forests, L. saulensis is a little known species. Ward et al . (1973) collected

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-127specimens on rodent and human bait in Para, Brazil, but this sper-.ies can not be regarded as being anthropophilic. Both sexes are readily identifiable.

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Figure 21 Lutzomyia saulensis — A. Male head, B. Male flagellomere II, C. Female head, D. Female flagellomere II, E. Female cibarium and pharynx, F. Male genitalia, G. Genital pump and filaments, H. Spermathecae, I, Female wing, J. Male wing, K. Female cibarium. Male: Curiche, Choco DepL., Colombia Female: Same locality as male

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-130V erruoarum Group Theodor, 196 5 This group of 22 species is divided into 2 series based on male characteristics (Theodor, 1965) -the series verruaarvm (14 spp.) and the series ssrrana (7 spp.). At present, L. buvsifortnis (Floch & Abonnenc) from French Guiana cannot be placed because the male remains undiscovered. Some authors include these species in the. subgenus Lutzomyia Franca (Forattini, 1971a, 1973) or in the subgenus Cororraia Barretto, 1962. Two species in the series sewana and 11 in the series vevruearum occur within or very near the borders of Colombia. The majority of species in the latter series are Andean, although L. ovallesi and L. evansi are widespread in lowland areas. The Andean taxa are morphologically similar and for the most part have limited, allopatric distributions. Unfortunately, the females of most species are difficult, if not impossible, to identify without associated males. For this reason I am omitting keys to the females but will point out, when possible, the distinguishing characteristics for some of the species. It may be appropriate to add that the presence of L. verrucarum (Townsend) in Venezuela (Anduze et al . , 1947; Floch and Abonnenc, 1950a; Forattini, 1973) can be confirmed only when the male is discovered in that country. K ey to Sp ecies Males 1. Style of male genitalia with 2 or 3 strong spines (series s err ana) . 2 Style with 4 strong spines (series verruc-ai^uni) 3

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-1312. Coxite tuft of 5-6 setae, upturned but not strongly recurved at tips, implanted in a straight row sen-ana (Fig. 22) Coxite tuft of 6-7 setae, markedly recurved apical ly, not implanted in a straight row odax (Fig. 28) 3. Basal spine of style isolated Basal spines of style paired, inserted more or less on the same level 7 4. Terminal spine of style very thick and crooked. Paramere as in Fig. 28£ spinier as sa (Fig. 28) Terminal spine of style relatively slender, not crooked. Paramere otherwise 5 5. Middle third of paramere with dorsal and ventral setae sauroida (Fig. 28) Middle third of paramere without dorsal or ventral setae 6 6. Paramere club-like, angular at tip and with dorsal subspical row of 13-16 short spine-like setae longiflocosa (Fig. 28) Paramere more or less rounded at end, less club-like, dorsal subapical setae longer, not spine-like quasitounsendi (Fig. 28) 7. Coxite with subapical group of long hairs. Basal spines of style inserted on well marked rather long tubercles 8 Coxite without subapical group of long hairs. Basal spines of style not implanted on tubercles 9

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•1328. Setae of basal coxite tuft thicker. Paramere as shown, with distal lobe on the ventral margin and with dorsal setae confined to extreme tip. Style with simple basal spines, no subterminal bristle. . . . moralesi n. sp. (Fig. 26) Setae of basal coxite tuft slender. Paramere more clubbed at end, without ventral lobe but with numerous dorsal setae in a subapical patch. Style with shorter basal spine modified as shown, subterminal bristle present andina (Fig. 23) 9. Dorsal setae of paramere restricted to distal third of structure. .10 Dorsal setae of paramere covering at least distal half of structure 11 10. Paramere strongly clubbed. Coxite tuft of 10+ setae. Flagellomere I at least 20% longer than lab rum ........ columbiana (Fig. 24) Paramere not clubbed. Coxite tuft of fewer than 7 setae. Flagellomere 1 short, subequal in length to labrum OVallesi (Fig. 27) 11. Basal coxite tuft of 18+ setae. Distal third of paramere wider than middle third disinnata (Fig. 23) Basal coxite tuft of 10 or fewer setae. Distal third of paramere not enlarged 12 12. Genital filaments over 4 x length of pump. Style with a subterminal seta evansi. (Fig. 25) Genital filaments less than 4 x length of pump. Style without a subterminal seta nuneztovari (Fig. 23)

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-133U"ie3 8Cl'r\:-l:7 22. Lutzomyia odax (Fig. 28) Phlebotomus odax Faircbild & Hertig, 1961b: 239 (6* holotype, o, Almirante, Bocas del Toro, Panama). Lutzomyia odax: Barretto, 1962, 97 (listed). Martins et al . , 1965: 7 (cf. to dubia) . Christensen & Fairchild, 1971: 302 (Darien, Panama). Christensen, 1972a: 88 (listed). Christensen et al., 1972: 57 (Panama). Forattini, 1973: 213 et seq. (gen. review, figs.). Distribution: Guatemala, Honduras , Nicaragua, Panama. Materialexamined'Panama, cf holotype (no, 4168), Almrirante (Bocas del Toro), light trap, 15 Oct. 1952, A. Quinones. o allotype (no. 3327), same data, but human bait collection, 20 June 1951, A. Vivanco. 1 cf, same data but 17 June 1950, R. Hartmann. discussion: There is little doubt that this species occurs in Colombia, Christensen and Fairchild (1971) having reported its presence within 4 km of the Colombian border in Darien Province, Panama. Females closely resemble those of L. serrana but differ in the lengths of the individual sperm ducts (Fairchild & Hertig, 1961b) . Both species occasionally bite man but are more common in light trap and resting collections. Forattini (1973) may be correct in treating L. Cybia Martins, Falcao, & Silva as a junior synonym of L. odax but more evidence is needed to support this view. Martins et al . (1965) point out differences between the taxa, especially those of the male genitalia.

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•134 23. Lutzorrgia serrana (Fig. "22) Vhlebotomus serranus Damasceno & Arouck, 1949: 843 (6* holotype, Serra da Piriabas, Para, Brazil). Forattini, 1960: 479 (Amapa, Brazil). Blancas, 1959-1960: 125 (Peru). Fairchild & Hertig, 1961b: 237 (refs., descr., figs.). Fianson, 1961: 320 (breeding sites). Johnson & Hertig, 196.1: 765 (rearing data) . Sherlock, 1962: 330 (Santander, Colombia). Rosabal, 1966: 1 (Costa Rica). Hanson, 1968: 76 (larva, pupa, descr.). Phlebotomus guayasi Rodriguez, 1956: 76 (cf, o, Guayas, Ecuador). Fairchild & Hertig, 1961b: 237 (as synonym of serranus) . Lutzomyia serrana: Barretto, 1962: 97 (listed). Martins et al . , 1965: 3 (Rondonia, Brazil), 6 (cf. to dvbid) . Lewis, 1967a: 74 (listed) Forattini, 1971a: 100 (listed). Tesh et al . , 1971a: 152 (blood meals). Chaniotis et al., 1971a: 345 (pop. dynamics, Panama). Osorno et al., 1972a: 30 (Colombian records). Tesh et al., 1972: 90 (blood meals). Chaniotis et al., 1972: 95 (listed). Christensen, 1972a: 88 (listed). Forattini, 1973: 123 et seq. (gen. review, figs.). Llanos, 1973: 32 (Peru, figs.). Velasco, 1973: 77 (Bolivia). Llanos et al . , 1975b: 668 (unusual spermathecae, fig.), 67]. (Peru). Herrer & Christensen, 1976a: 62 (Panama). Llanos et al . , 1976: 480 (Peru). Martins et al . , 1976a: 488 (Peru) . Martins et al., 1976b: 496 (Peru). Distribution: Mexico, Honduras , Nicaragua, Costa Rico,, Panama, Colombia (Antioquia, Bolivar, Boyaca, Caldas, Chocc, Cundinamarca, Santander), Ecuador, Peru, Bolivia, Brazil. Material examined: Colombia. 1 c?, Alto Curiche (Choco) , tree, buttress, 21 May 1967, D.C,.^. 1 cf, Rio Anori (Antioquia), tree buttress, 6 Feb. 1970, C.H.P. 1 0, same data but at light, 21 Sept. 1970, D.G.Y.

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-135Panama. 1 ] <2, Rio Platanar (Panama), hollow tree, 20 July 1950, R. Hartmaan. Discussion: In Panama, Tesh et al .. (1971a, 1972) identified the blood meals of 14 females using the precipitin test. Ten had fed upon rodents, 2 on edentates and 2 on marsupials. Dr. C.H. Porter (in litt.) collected less than 10 specimens on human bait over a one year period at Ric Anori (Antioquia) , Colombia. Series verrucarum 24 . Lutzomyia andina (Fig. 23) Lutzomyia andina Osorno, Osorno, & Morales, 1972b: 2 (c? holotype, p, El Chariquito, Cundinamarca, Colombia). Osorno et al., 1972a: 24 (Colombian records) . Fh'lebotomus verruaaricn: Osorno et al., 1967: 29 (not verrueanm Townsend) . Distribution: Colombia (Cundinamarca). Material examined: 1 6", paratype, type locality, 2550 m above sea level, Aug. 1966. 1 o, paratype, same data but Sept. 1966. Discussion: Specimens of this recently described species, known only from the type locality, were taken on human bait and in tree holes. Females bite man during the day and night. The shorter of the paired basal spines of the male sytle is curiously modified, the apical part widened and with a flange (Fig. 2 3A) . This characteristic, if shared by other andina males, will serve to separate it from other males in the verrucar-jm group.

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-13625 . Lu tzomyia co I vjrb i an a (Fig. 24) Phlebotomus columbianus Ristorcel]i & Van Ty, 194.1: 263 (o holotype, Valle He Capuli, Narino, Colombia). Rozeboom, 1947b: 705 (6*, figs.). Sherlock, 1962: 322 (cf, o, figs.). Osorno et a.l . , 1967: 28 (mention). Phlebotomus montieolus var. incarwn Ristorcelli & Van Ty, 1941: 266 (o, Valle de Capuli, Narj.no, Colombia). Rozeboom, 1947b: 705 (as synonym of colurrbiand) . Lutzomyia colvmbiana: Rarretto, 1962: 97 (listed). Osorno et al., 1972a: 24-26 (Colombian records). Forattini, 1973: 206 et seq . (gen. review, figs . ) . Distribution: Colorrbia (Cauca, Guajira, Narino, Valle). Material examined: Colombia. 5 6*6", 3 oo, Bolivar (Cauca), Jan. 1944, J. Burbano. 1 (J 1 , 3 oo, San Pedro (Narino), Jan. 1944, J. Burbano . 2 6*6*, ca. 30 km NW of Cali, near El Carmen (Valle), tree trunks, 12 Aug. 1973, D.G.Y. & R.C.W. Discussion: This precinctive Colombian species was the suspected vector of bartonellosis during the outbreak in southeastern Colombia over 40 years ago (Rozeboom, 1947b) . The record of colvmbiana. from Montes de Oca and Serra Kacuira in Guajira (Osorno et al . , 19 72a) is very interesting, these localities being widely separated from southeastern Colombia where colurrbiana, was first discovered. The Andes, especially the eastern chain, may have acted as a corridor for the dispersal of this species. 1 have seen other specimens of colunfoiana in the collections of the California Academy of Sciences and University of Florida. All were captured in southeastern Colombia and were identified by 0. Mangabeira and P. Galindo.

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13726. Lutzorr^ia disiunata (Fig. 23) Lutzomjia disiunata Morales, Osorno, & Osorno, 1974: 446 (cfholotype, o, Hacienda La Conejera, Municip. de Suba, Bogota, Cundinamarca, Colombia) . Distribution: Col-orrhia (Cesar, Cundinamarca). Material examined: Colombia. 8 3d 1 , 21 oo, Municip. Agustin Cadazzi (Cesar), Sept. 1969 (INPES) . 3 do" (including holotype) , 9 o^, type locality, Aug. -Sept. 1969 (INPES). Discussion: This is another Andean species which is anthropophilic , specimens having been taken on human bait during the day and night (Morales et al., 1974). Other specimens were captured in tree holes in Cesar Department. 27. Lutzcyia evansi (Fig.' 25) Phlebotomus evansi: Nunez-Tovar, 1924: 44 (cf, Mariara, Carabobo, Venezuela). Barretto, 1947: 198 (full refs.). Mirsa, 1953: (rearing data). Fairchild & Kertig, 1959: 122 (distrib.). Pifano et al . , 1962: 412 (refs., redescr., figs.). Rosabal & Trejos, 1964: 167 (El Salvador) Rosabal & Trejos, 1965: 222 (El Salvador). Osorno et al . , 1967: 28 (Narino, Colombia) . Lutzomyia evansi: Theodor, 1955: 183 (listed). Morales et al., 1969b: 385 (mention). Osorno et al . , 1972a: 27 (Colombian records). Llanos, 1973: 34 (d*, o. redescr., figs., Peru). Forattini, 1973: 122 et seq. (gen. review, figs.). Llanos et al . , 19 75b: 670 (Peru).

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-138Distribution: Guatemala,, El Salvador, Honduras, Nicaragua, Costa Rica, Colombia (Bolivar, Guajira, MagJalena, Narino, Santander) , Peru, Venezuela. Material examined: Colombia. 1 q, Arjona (Bolivar), 27 May 1940. 1 o, 10 km SE of Santa Marta (Magdalena) , tree trunk, 17 Aug. 1973, D.G.Y, & K.C.W. Honduras: 30+ d*cf, 100+ oo, Chumbagua, Santa Barbara, light traps, July 1966, J. Matta. Other specimens from Central America and Venezuela in collection at UF. Discussion: The female of evansi has very long sperm ducts and relatively small spermathecae (Fig. 25J) , features which separate it from other verrucavum group females in Colombia. This species has been reported biting man in several countries including Colombia (Osorno et al., 1972a). It and L. longipalpis often, but not always, occur together in Central America, Colombia, and Venezuela. The apparent absence of this species from Panama is most peculiar. 28. Lutzomyia longifloeosa (Fig. 28) Lutzomyia longifloeosa Osorno, Morales, Osorno, & Hoyos, 1970: 1 (c? holotype, Municip. de Tello , Huila, Colombia. Osorno et al., 1972a: 28 (Colombian records). Osorno et al . , 1972b: 18 (cf. to andina and quasitownsendi) . Distribution: Colombia (Huila) . Material examined: Colombia. 1 d" (paratype no. 1641), type locality, 9 Feb. 3 968, E. Osorno et al. Discussion: Other than resting in tree holes at the. type locality, nothing is known about the habits of this species. The female is unknown.

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•13929. LutzomyLa moralesi n. sp. (Fig. 26) Male: Wing length 1.96; width 0.58. Head, mesonotum, and genitalia strongly pigmented, rest of insect pale. Head height 0.35; width 0.32. Eyes separated by 0.11 or by distance = to 5.9 facet diameters. Flagellomere I (0.33 long), 1.2 x length of II + III; ascoids simple, those on flagellomere IT ending near middle of structure, on all f lagellomeres except last. Length of palpal segments: 1 (0.04), 2 (0.15), 3 (0.16), 4 (0.1 1), 5 (0.32); palpal sensilla (ca. 10) grouped loosely at middle of segment 3. Labrum length 0.21. Cibarium with 30+ vestigial, dot-like teeth; chitinous arch complete, diffuse in middle; pigment patch invisible. Pharynx 0.15 long, unarmed. Pleura with 8-9 upper and 4 lower episternal setae. Length of wing vein sections: Alpha (0.53), beta (0.18) delta (0.17), garmr.a (0.32). Length of femora, tibiae, and basitarsi: Foreleg, 0.78, 0.90, 0.54; midleg, 0.74, 1.07, 0.61; hindleg 0.83, 1.22, 0.69. Abdominal sternites hardly visible some with lateral, circular openings. Genitalia: Style 0.13 long, shaped as shown, the proximal 2 spines inserted at the same level, the longer on a long process, no subterminal seta. Coxite (0.32 long x 0.12 wide) with a distal patch of 30+ straight or nearly straight setae, a dense tuft of ca . 30 thick sinuous setae at base. Paramere as shown but variable according to angle of view; the apical part with an incipient lateral lobe and a ventral heel. Aedeagus subtriangular, unmodified. Lateral lobe relatively long (0.42). Genital pump (0.155 long), each filament 0.63 long or 4 x length of pump, tip enlarged. Cercus as shown. Female: Wing length 2.45; width 0.76. Coloration as in d 1 . Head height 0.43; width 0.39. Eyes separated by 0.13 or distance = to ca. 6.6

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-140facet diameters. Flagellomere I (0.31 long) nearly 1.3 x length of II + III; .ascoids longer than those of o but not reaching end of flagellomere II; on all flagellomeres except last. Length of palpal segments: 1 (0.014), 2 (0.20), 3 (0.19), 4 (0.12), 5 (0.39); palp 2 with ca. 5 apical sent, ilia, palp 3 with 25+ sensilla on apical half. Lab rum 0.36 long, Cibarium with 4 sharp horizontal teeth evenly spaced, 2 transverse rows of subequal vertical teeth, ca . 13 teeth in each row; chitinous arch complete; pigment patch subtriangular , well defined. Pharynx 0.20 long, unarmed but with distinct ridges. Pleura with 9-10 upper and 2 lower episternal setae. Length of wing vein sections: Alpha (0.67), beta (0.20), delta. (0.26), gemma. (0.41). Length of femora, tibiae, and basitarsi: Foreleg, 0.88, 0.99, 0.59; midleg, 0.83, 1.18, 0.66; hindleg, 0.87, 1.34, 0.78. Abdominal sternite 3 with paired circular openings laterally. Tergite 8 with 1 or 2 setae on each side. Spermathecae saclike, as figured; common duct ca. 3 x length of individual duct. Cerci unremarkable . Distribution: Colombia. (Valle) . Material examined: Colombia. 6* holotype (no. 512), ca. 15 km SW of Cali (1600 m above sea level) near Rio Pance (Valle), on tree trunk, 8 Aug. 1973, D.G.Y. & S. Ayala. ^ allotype (no. 513), ca . 10 km W of Cali (1800 m above sea level), near Pichinde (Valle), rock crevice, 2 Aug. 1973, D.C.Y. & R.C.W. Paratypes (nos. 514-526), 1
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-141The male of this species resembles that of L. and,ina in details of the male genitalia but the setae of the basal coxi te tuft are thicker, the paramere and its setation are different, the style lacks a small subterminal seta, and the shorter of the two proximal spines is unmodified. The male of L. vei'vucavum (Townsend) , an allopatric Peruvian species, is similar to these species, but the proximal spines of the style are not inserted on long or well marked tubercles and the parameres are different. Without males, I am unable to separate the females of L. morale si from those of most other verruaarien group species. Identifications based on this sex alone require confirmation. I am pleased to name this species after Dr. Alberto Morales-Alarcon, not only for his valuable assistance in this study, but to acknowledge his significant contribution to our knowledge of Colombian phlebotomines . 30. Lutzomyia nuneztovari (Fig. 23) Phlebotomus nunez-tovari Ortiz, 1954: 232 (6" holotype, Duaca, Lara, Venezuela). Scorza & Ortiz, 1960: 434 et seq. (ecology). Pifano et al., 1962: 387 (d* keyed) . Ortiz & Scorza, 1963: 344 (Venezuela). Scorza et al., 1967: 194 (6* keyed) , 195 (o keyed), 196 (collecting data). Lutzomyia nuneztovari'. Theodor, 1965: 183 (mention), Martins & Silva, 1965: 275 (mention). Martins et al., 1965: 9 (mention). Morales et al., 1969b: 385 (mention). Forattini, 1971a: 103 (listed). Martins & Fraiha. 1971: 364 (cf. to evangelistai) . Young, 1972a: 312 (mention). Osorno et al., 1972b: 18 (mention). Forattini, 1973: 265 (as synonym of ovallesi) . Distribution: Venezuela, Colombia (Huila) .

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-142Materia'L examined: Colombia. 1 6*, Timana (Huila) , Shannon trap, 21 July 1971, C. Aguila. 1 6", same data but 1-2 Aug. 1971. 1 p, same data buc light trap, 2-3 Aug. 1971. 1 o, same data but Shannon trap, 12-13 Aug. 1971. Discussion: Unlike Forattini (1973), I consider L. nv.neztovo.ri and L. OVdltesi to be distinct species, the male of the latter having a subterminal on the style (lacking in nuneztovari) and with different parameres. The female of ovaVlesi has elongate spermathecae, slightly constricted in the middle (Fig. 27J) unlike those of nuneztovari which are stout and non-constricted (Scorza et al., 1957, in key and Fig. 23E this paper) . Fairchild (pers. coram.) examined the holotype of nuneztovari in 1963, and noted the following: Tips of genital filaments slightly expanded. Subterminal seta of style lacking. Parameres long and slender, very slightly clubbed, not: pointed as shown by Ortiz (1954) but blunt, slightly shorter than lateral lobes. Alpha of wing venation very long, at least equal to beta + gamma, not as shown by Ortiz (1954) . The males from Huila, Colombia, agree with the description of the holotype by Ortiz (1954) (as slightly emended by Fairchild) and I therefore have no doubt that they are conspecific. The Colombian females of nuneztovari are in poor condition with partially missing palpi and antennae, but other structures are intact and are described as follows. Female (n = 2): Wing length 2.25; width 0.74. Head, mesonotum, and abdominal tergites strongly pigmented, rest of insect, including pleura, pale or dusky but the degree of pigmentation difficult to determine due to overstaining. Head height 0.42-0.43. Eyes separated by 0.11-0.12 or by distance = to ca. 6 facet diameters. Length of palpal segment 1 (0.05),

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-1432 (0.19). others missing; 2 palpal sensilla on apical part of segment 2. Labrum 0.32 long. Cibarium with 4 sharp equidistant horizontal teeth and 12-20 small vertical teeth in a row; chitinous arch complete; pigment patch sub triangular, the posterior (wider) half more heavily pigmented. Pharynx 0.20 long, unarmed. Pleura with 12 upper and 4 lower episternal setae. Length of wing vein sections: Alpha (0.71), beta (0.22), delta (0.32), gamma (0.31). Length of femora, tibiae, and basitarsi: Foreleg, 0.78, 0.86, 0.54: midleg, 0.83, 1.05, 0.64; hindleg, 0.88, 1.27, 0.71. Abdominal sternite 2 with a median clear area not enclosed posteriorally . Tergite 8 with 5-7 setae on each side. Spermatheca wrinkled and sac-like, terminal knob deeply recessed; common sperm duct and most of individual ducts invisible. Cerci unremarkable. 31. Lutzorirjia ovallesi (Fig". 27) Phlebotomus ovallesi. Ortiz, 1952: 155 (6* holotype, Duaca, Lara, Venezuela, not San Felipe, Yaracuy, Venezuela). Ortiz, 1954: 239 (6*, redescr., figs.). Lewis & Garnham, 1959: 87 (o descr., Belize). Fairchild & Hertig, 1959: 122 (Central American records). Scorza & Ortiz, 1960: 434 (ecology). Hanson, 1961: 317 (breeding sites). Johnson & Hertig, 1961: 765 (rearing data). Pifano et al . , 1962: 370 et seq. (d*, o keyed, redescr., figs.). Strangways-Dixon & Lainson, 1962: 297 (with flagellates, Belize). Johnson et al . , 1963: 110 (negative for flagellates). McConnell & Correa, 1964: 527 (infected with fungus). Williams et al., 1965: 65, 70 (Belize). Rosabal, 1966: 1 (Costa Rica). BiagL et al., 1966: 368 (Mexico, o figs.). Thatcher & Hertig, 1966: 50 (biting man and Potos , Panama). Scorza et al., 1967: 191 (distrib.).

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-144Hanson, 1968: 64 (larva, pupa, descr.). Osorno et al., 1967: 29 (Colombia) . Sergentomyia ovallesi: Barretto, 1955: 185 (listed). Barret to et al., 1956: 51 (cf. to firmatoi) . Lutzomyia ovallesi: Barretto, 1962: 98 (listed). Aitken et al., 1968: 264 (Trinidad, on man). Williams, 1970: 332 et seq. (summary of studies, refs., Belize). Tesh et al., 1971a: 153 (blood meals). Chaniotis et al., 1971a: 100 (listed). Osorno et al., 1972a: 28 (Colombian records). Shaw et al., 1972: 719 (mention). Chaniotis et al . , 1972: 95 (collecting data). Christensen, 1972a: 88 (listed). Christensen et al., 1972: 57 (light trap catch). Rutledge & Mosser, 1972: 366 (listed). Forati:ini, 1973: 122 et seq. (gen. review, refs., figs.). Lewis, 1975a: 500 et seq. (mouthpart morphol.). Rutledge et al., 1975: 181 (ecology). Rutledge & Ellenwood, 1975a: 73 (ecology, refs.), 1975c: 87 (mention). Williams, 1976a: 603 (in caves, Belize); 1976b: 618 (mention). Bruinptonryia ovallesi: Lewis, 1965: 378 et seq. (internal morphol.). Distribution: Mexico, Belize, Honduras, Nicaragua, Costa. Rica, Panama, Colombia (Antioquia, Boyaca, Caldas, Choco, Cundinamarca, Magdalena, Santander) , Venezuela, Trinidad. Material examined: Colombia. 34 do", 19 go, Teresita (Choco), in tree buttresses, March-Nov. 1967, D.G.Y. 1 6", near Rio Don Diego, E of Santa Marta (Magdalena) , 15 Aug. 1973, D.G.Y. & R.C.W. Panama. 2 oo, Almirante (Bocas del Toro) , Shannon trap, 13 July 1951, A. Quinones. Trinidad. Both sexes from various localities, to be discussed in a forthcoming paper. Discussion: The. female of ovallesi, a man biter in several countries including Colombia (Osorno et al . , 1972a), is readily identified by the

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-145elongate constricted spermathecae (Fig. 27J) . Flagellates, apparently not Leishmania, have been recovered from wild caught specimens in Panama and Belize (Williams, 1970). Based on evidence gathered by Thatcher and Hertig (1966) and Tesh et al . (1971a), this species feeds primarily, if not exclusively, on mammals. In Panama, immatures were found in soil at the base of trees (Hanson, 1961), less commonly on the open forest floor (Rutledge and Ellenwood, 19 75a) . 32. Lutzomuio. quasi towns endi (Fig. 28) Lutzorajia quasitounsendi Osorno, Osorno, & Morales, 1972b: 10 (
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46Material examined'Colombia. 1 d* (paratype no. 2690), 1 o (paratype no. 2685, both in collection at INPES) , type locality, 29 Oct. 1968. Discussion: This species and L. townsendi (Ortiz) (Fig. 12) from Venezuela are very similar, the males of sauroida having slightly stronger setae on the dorsum of the parameres but are otherwise indistinguishable. I can find no differences between the females of the. two species but hesitate to treat sauroida as a junior synonym of townsendi until more material of the former species becomes available. 34 . Lutzomyia spinicrassa (Fig. 28) Lutzomyia spinicrassa. Morales, Osorno, Osorno, & Hoyos , 1969b: 383 (c? holotype, ve.reda Umbabita, Municip. Almeida, Boyaca, Colombia; o, vereda ChLtavita, Municip. Guayata, Boyaca, Colombia). Osorno et al., 1972a: 30 (Colombian records). Forattini, 1973: 213 et seq. (o fig-, keyed) . Distribution: Colombia (Boyaca) . Material examined: Colombia. 1 d" (paratype no. 2654), 1 o (paratype no. 2652), Guateque (Boyaca), 22 Oct. 1968, A. Morales & J. Ochoa. Eoth specimens in collection at UF. D-iscussion: The male of spinicrassa differs from other males in the vcrrucavvm group by the size and shape of the terminal spine of the style (Fig. 28U) . This species, known only from Boyaca Dept. at elevations ranging from 1430 to 1640 m above sea level, was found resting in tree holes and on walls inside of houses.

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Figure 22 Lutzomyia serrana — A. Male head, B. Rile flagellomere II, C. Female head, D. Female flagellomere II, E. Female cibarium and pharynx, F. Male genitalia, G. Paramere, H. Spermathecae, I. Genital pump and filaments, J. Female wing, K. Male wing, L. Female cibarium. Male: Alto Curiche, Choco Dept., Colombia Female: Rio Anori, Antioquia Dept., Colombia

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•148-

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Figure 23 Lutzomyia andina male — A. Male genitalia. Male: El Charquito , Cundinamarca Dept., Colombia Lutzomyia disiuneta male — B. Male genitalia. Male: Bogota, Municip . de Suba, Cundinamarca Dept., Colombia Lutzomyia nuneztovari -C. Female cibarium, D. Male genitalia, E. Spermatheca; the common duct and part of individual duct not visible. Male: Timana, Huila Dept., Colombia Female: Same locality as male

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-150-

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Figure 24 Lubzomyia oolwrbiana — A. Genital pump and filaments, B. Male head, C. Mais flagellomere II, D. Female head, E. Female flagellomere II, F. Femals cibarium and pharynx, G. Male genitalia, H. Female cibarium, 1. Sperraathecae, J. Body of spermatheca, K. Female wing, L. Male wing, M. Spermathecae. Male: San Pedro, Narino Dept., Colombia Female: Same locality as male

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—1 52—

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Figure 25 Lutzomyia evansi — A. Male head, B. Male flagellomere II, C. Female head, D. Female flagellomere II, E. Female cibarium and pharynx, F. Genital pump and filaments, G. Genital pump, same scale as 25H, H. Male genitalia, I. Paramere, outer aspact, J. Spermathecae, K. Body of spermatheca, L. Female wing, M. Male wing, N. Female cibarium. Male: Guapilonar, Carazo, Nicaragua Female: Same locality as male

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M cf

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Figure 26 Lutzorwjia movalasi— A. Female head, B. Female wing, C. Male wing, D. Male head, E. Male flagellomere II, F. Female flagellomere II, G. Female cibarium, H. Genital pump, same scale as Fig. 26 J, I. Tip of genital filament, same scale as Fig 26G, J. Male genitalia, K. Spermathecae. Male: Near Rio Pance, Valle Dept., Colombia Female: Near Pichinde, Valle Dept., Colombia

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-156-

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Figure 27 Lutzomyia ovaltesi — A. Male head, B. Male flagellomere II, C. Female head, D. Female flagellomere II, E. Female cibariun and pharynx, F. Genital pump and filaments, same scale as Fig 27G, G. Male genitalia, H. Female wing, I. Male wing, J. Spermathecae, K. Female cibarium. Male: Rio Sabana, Darien Prov., Panama Female: Same locality as male

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153-

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Figure 28 Lutzomyia sauvoida -A. Male genitalia, B. Female cibarium. Hale: Municip. de Santa Ana, Boyaca Dept., Colombia Female: Same locality as male Lutzomyia quasitcwnsendi — C. Male genitalia. Male: Barbosa, Santander Dept,, Colombia Lutzomyia longiflocosa — D. Male genitalia. Male: Municip. de Tello, Huila Dept., Colombia Lutzomyia spiniorassa — E. Male genitalia. Male: Guateque, Boyaca Dept., Colombia Lutzomyia odax — F. Coxite tuft and paramere. Male: Almirante, Bocas del Toro Prov., Panama

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-160A ( \!Sk*m ,J'; c I ^K \ -'> v ^> :'';>:': _j,\\ *5bLJ>

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-161Vespert ilionis Group Theodor, 1965 With the center of distribution in Central America, the vespertili.onis group species are divided into 2 series following Theodor (1965) -the series vespertitinois (5 spp.) and the series deleoni (3 spp.). Barretto (1962) created the subgenus CoTomyia to accomodate them and other species, mostly those in the verrucamm group. Forattini (1971a> 1973) divides the vesper tiliorvis group species by placing 1 in the subgenus Lutzomyia Franga, 2 in the subgenus Dampfomyia Addis, while leaving the others in Coromyia. In my opinion, this arrangement is unnecessary when the species are classified using a combination of character states. Two sibling species in the series vespertili-onis are known to occur in Colombia. The females feed on bats and are morphologically indistinguishable . Key to Species Males 1. Paramere with dorsal setae on distal half of structure L. vespertilionis (Fig. 29) Paramere with dorsal setae restricted to distal third of structure L. isovespertilionis (Fig. 29) 35. Lutzomyla isovespei-tili-on-ls (Fig. 29) Phlebotomies isovespertilioni-s Fairchild & Hertig, 1958a: 516 (6* holotype, p, reared from eggs laid by p from tributary of Rio Cocoli, Canal Zone). Johnson & Hertig, 1961: 765 et seq. (rearing data).

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-162Thatcher & Hertig, 1966: 46 (mention). Hanson, 1968: 88-90 (larva, pupa, descr.) . Phleho tonus vespevtitionis : Fairchilci b. Hertig, 1947b: 617 et seq. (in part, descr., fig.). Luizcmuia isovespevtilionis: Barretto, 1962: 97 (listed). Chaniotis et al . , 1971a: 344 et seq. (in part, pop. dynamics, Panama). Chaniotis et al., 1972: 94 et seq. (resting sites, Panama). Christensen et al . , 1972: 57 et seq. (collecting data, infected with trypanosomes , Panama). Christensen, 1972a: 88-89 (mention). Forattini, 1973: 122 et seq. (gen. review, fig.). Christensen & Herrer, 1973: 579 et seq. (collecting data, Panama). Rutledge & Mosser, 1975: 411 (mention). Herrer & Christensen, 1969: 62 (collecting data, Panama) . Pistx'ibution: Panama, Colombia (Choco). Material examined'. Colorrbia. 1 o", Curiche (Choco), tree buttress, 5 July 1967, D.G.Y. 4 dcf, Teresita (Choco), tree buttress, 18 May 1967, D.G.Y. Panama. 1 6", Cruces trail, Madden Forest, Canal Zone, hollow tree, 7 Sept. 1949, M. Hertig. 3 6*cf, Cerro Campana (Panama Prov.), tree hollows and buttresses, 2 Oct. 1949, M. Hertig & H. Trapido. Discussion: See L. vespertilicnis 36. Lutzomyia vespertilioni (Fig. 29) Phlebotomies vesperhitionis Fairchild & Hertig, 1947b: 617 (6" holotype, Cerro Campana, Panama Prov., Panama). Hertig & Fairchild, 1950: 91 (abdor.iin.-il steraites). Barretto, 1951: 225 (distrib.). Vargas & Difiz-Najera, 1951b: 101 (cf. to beltrani) . Floch & Abonnenc, 1952: 30-48 (d\ o, keyed). Rosabal, 1954: 10, 28-29 (measurements, figs.,

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Costa Rica, sp. no. 12 = Vesper tilionis) . Rodriguez, 1956; 76 et seq. (Ecuador). Fairchild £< Hertig, 1958a: 510 (keyed). 514 (o, o, redescr., figs.). Fairchild & Hertig, 1959: 122 (Central American records). Hanson, 1961: 320 (breeding site, Panama). Johnson & Hertig, 1961: 765 et seq. (rearing). McCormell & Correa, 1964: 523 et seq. (infected with trypanosomes & fungi, Panama). Thatcher & Hertig, 1966: 46 et seq. (mention). Hanson, 1968: 88-90 (larva, pupa, descr., figs.). Lutzomyia vespertilionis: Barretto, 1962: 97 (listed). Theodor, 1965: 184 (o*, o, figs.). Barreto, 1969: 468 (Valle, Colombia). ZeleT don & Rosabai, 1969: 221-227 (as vector of bat trypanosome) . Tesh et ai., 1971a: 152-153 (in part, blood meals). Chaniotis et al . , 1971a: 344 et seq. (in part, pop. dynamics, Panama). Christensen et al., 1971: 116 et seq. (pyloric armature, figs.). Tesh et al . , 1972: 90 et seq. (in part, blood meals). Chaniotis et al., 1972: 94 et seq. (resting sites, Panama). Christensen, 1972a: 88-89 (mention). Christensen, 1972b: 683 et seq. (rearing). Osorno et al . , 1972a: 32-33 (Colombian records). Rutledge & Mosser, 1972: 307 (mention). Christensen & Herrer, 1973: 579 et seq. (collecting data, infected with trypanosomes, Panama). Forattini, 1973: 123 et seq. (gen. review, figs.). Rutledge & Mosser, 1975: 411 (mention). Distribution: Costa Rica, Panama, Colombia (Antioquia, Bolivar, Boyaca, Choco, Valle), Ecuador. Material examined: Colombia. 3 66, Rio Anori (Antioquia), tree trunks, 6 Feb. 1970, C.H.P. 2 oV, 1 o, same data but Sept. 1970, D.G.Y. 4 dc/, 2 co, Anchicava Dam (Valle), tree trunks, 9 Aug. 1974, D.G.Y. & t i R.C.W. 7 dV, 14 oo, same data but light trap at entrance of tunnel,

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-16410-11 Aug. 1974. 1 6*, 1 o, 25 km E of Buenaventura (Valle) , tree trunks, 11 Aug. 1974, D.G.Y. & R.C.W. Ecuador. 15 c?tf, 1 o, 17 kn E of Santo Domingo do los Colorados (Pichincha) , tree trunks, 4-6 May 1976, D.G.Y. & 1. Rogers. Panama. 1 o* (holotype no. 761), Cerro Campana (Panama), shallow cave, 17 Jan. 1947. Discussion: In structure and habits, L. vesper til-ionis and L. isovespertilionis are remarkably similar, both species often being found in tree hollows associated with bats. At least one species of Trypanosoma is transmitted to these mammals by one or both of these flies (Zeledon and Rosabal, 1969) . The preferred breeding site(s) of vespertilionis and -isovespertilionii has not been determined (Rutledge and Mosser, 1972), the only wiJd caught larvae being reported by Hanson (1961, 1968) who recovered this stage on two occasions in soil and bat guano inside of hollow trees. Apparently the larvae and pupae of these species are indistinguishable (Hanson, 1968). Lubzomyia vespertiZionis ranges as far south as Guayas Province, Ecuador. Luty.omijia isovespertilionis probably has a more limited distribution as noted by Fairchild and Hertig (1958a) , its southernmost locality at present being Choco Department, Colombia. These are the only vespertilionis group species which occur in South America. Fairchild and Hertig (1958a) provide further information on collecting data, distribution, and taxonomy.

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Figure 29 Lutzonyia Vespertilonrs — A. Male head, R. Male flagellomere II, C. Female head, D. Female flagellomere II, E. Female ciharium and pharynx, G. Paramere, H. Paramere of male from Panama, I. Male genitalia, J. Spermathecae, K. Female wing, L. Male wing, M. Female cibarium. Male: Rio Anori, Antioquia Dept. , Colombia (except Fig. 29H) Female: Same locality as male Lutzomyia isovespeptilionis — F. Paramere. Male: Panama Canal Zone

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-166Cf :<:?f\ D ~^^2rf> H t\it-'^7 ^ x^ V \. \ M w Ml. ; % /e =Z7 w \ ! \

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-167S ubg enus Dampfomyia Add is, 1 945 The 5 species in this subgenus are characterized by several features (Fairchild and Hertig, 1956; Theodor, 1965), some of which are shared by L. aquilonia (Fairchild and Harwood) of the vespsrti.li.onis group and members of the delpozoi group Lewis et al. (1973). The spermathecae of the Dampfomyia species are large with bubble-like expansions and could have been derived from the d&leoni-type spermathecae of the vespertilionis group. Like species in these two groups, ths Dampfomyia spp. do not ordinarily bite man and there are more species in Mexico and Central America than elsewhere. The preferred host(s) of L. vosabali, the only Dampfomyia sp . in Colombia or South America, is unknown. Related species such as L. anthophora (Addis) and L. permiva (Fairchild and Hertig) feed on small mammals, the former species probably transmitting Rio Grande virus to Neotoma woodrats in Texas (Calisher et al., 1977). 37. Lutzomyia (D.) rosabali (Fig. 30) Phlebotomus vosabali Fairchild & Hertig, 1956: 310 (6* holotype, Puerto Armuelles, Chiriqui Prov. , Panama; o, Guarare, Los Santos Prov., Panama). Rosabal, 1954: 10 et seq. (as sp . no. 9, Costa Rica). Fairchild & Hertig, 1959: 122 (Central American records). Rosabal & Trejos, 1964: 168 (cf . to dodgei) . Lutzomyia rosabali: Barretto, 1962: 95 (listed). Theodor, 1965: 193 (listed, spermatheca fig.). Forattini, 1971a: 100 (listed). Osorno et al., 1972a: 61 (Narino, Colombia). Christensen, 1972a: 88 (listed). Forattini, 1973: 208 (spermatheca fig.), 329 (as synonym of dodgei).

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•168Distribution: Costa Rica, Panama, Colombia (Cauca, Narino). Material examined'. Colombia. 6 c5c?, 8 oo, Capuli (Narino), rock crevices, March 1945, M. Hertig. 3 go, Bolivar (Cauca), Jan. -Feb. 1944, J. Burbano. Panama. 1 o" (holotype no. 3209), type locality, tree buttress, 9 June 1951, M. Hertig. 1 o (allotype), Guarare (Lcs Santos), light trap, 23 Oct 1952, F. Blanton. Discussion: Forattini (1973) treats L. rosabali as a junior synonym of L. dodgei (Vargas and Najera), believing, perhaps correctly, that the differences between the males are due to intraspecif ic geographic variation. Without a larger sample of specimens from various localities, however, I prefer to consider them a distinct based on the. discussion and figures of Fairchild and Hertig (1956) . The Colombian males (Fig. 30) agree more closely with L. rosabali than with L. dodgei in the shape and setation of the dorsal arm of the paramere but the basal spine of the style is longer and more slender. The small bristle near the basal spine is absent or: one style of the rosabali hoiotype and from the styles of the Colombian males.

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Figure 30 Lutzomyia (D.j rosabali — A. Male head, B. Male flagellomere II, C. Female head, D. Female flagellomere II, E. Female cibarium and pharynx, Panamanian specimen, F. Male genitalia, G. Dorsal arm of puramere, H. Different views of paramere, I. Genital pump and filaments, J. Spermathecae, K. Female wing, L. Male wing, M. Female cibarium, Panamanian specimen. Male: Capuli, Narino Dept., Colombia Female: Same locality as male (except Fig. 30E and 30M)

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170-

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-171Sub genus Pintomyia Cost a Lima, 1932 The 5 described species in this subgenus are characterized by a row of short spines on the hind femur. Forattlni (1971a, 197 3) treats Pintomyia as a genus, placing, I believe, undue importance on these spines which, although not exactly the same, are shared by 2 species of Serge atomy ia Franca and 1 of Bnanptomyia. The Pintomyia females, like those of Pressatia, have well sclerotized individual sperm ducts but differ from them in other aspects (Theodor, 1955). At present, only L. spinosa is known to occur in Colombia. Lutzomyia damaseetioi (Mar.g.), a clos^ ally, was repoi'ted in Meta Department by Morales et al. (3 969a) but was excluded in a later work on the Colombian fauna (Osorno et al., 1972a) probably because of the uncertain identity of the specimens (2 oo) . Except for L. pessoai (Coutinho and Barretto) and L. fisokeri (Pinto) . the Pinto.nyia spp . have not been reported feeding on man. Lutzomyia gihsoni (Pifano and Ortiz), recently described from Venezuela from a single female, definitely belongs in the subgenus but its identify remains uncertain based upon the original description. 38. Lutzomyia (Pi.) spinosa (Fig. 31) Pklebntomv3 spinosus Floch & Abonnenc, 1942b: 1 (o* holotype, Gallion, near Cayenne, French Guiana). Barretto, 1947: 224 (full refs.). Hertig L Fairchild, 1950: 92, 93 (abdominal sternites 1 & 2, figs., Panama). Barretto, 1951: 224 (aistrib.). Floch & Abonneuc, 1952: 20 et seq . (a*, o, keyed, redescr. , figs.).

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-172Blancas, 1959-1960: 129-130 (cf. to gorbitzi) . Johnson & Hertig, 1961: 765 et seq. (rearing data). Hanson, 1968: 80 (larva, pupa, descr., figs.). Morales et al . , 1969a: 381 (Meta, Colombia). Pifano & Ortiz, 1972: 31 (cf. to gibsoni) . Pifano et al., 1973: 153-154 (cf. to gibsoni) . Lutzonvjia spinosa: Barretto, 1962: 93 (listed). Martins et al., 1962a: 381 (Ooias, Brazil). Martins et al . , 1963: 335 (Brazilian records). Christensen & Fairchild, 1971: 302 (Darien Prov., Panama). Christensen, 1972a: 88 (listed). Osorno et al., 1972a: 60-61 (Colombian records). Ward et al., 1973: 178-181 (collecting data, Para, Brazil). Christensen & Herrer, 1973: 579 (Collecting data, Panama). Lewis, 1975a: 503 et seq. (mouthpart morphol.). Pintomyia spinosa: Forattini, 1971a: 104 (listed). Forattini, 1973: 123 et seq. (gen. review, figs.). Distribution: Panama, Colombia (Amazonas, Antioquia, Boyaca, Caqueta, Choco, Meta), French Guiana, Brazil. Material examined: Colombia. 70 6*0*, 12 09, 17 km W of Leticia (Amazonas), tree trunks, 24-26 July 1973, D.G.Y. & R.C.W. 1 6", Rio Anori (Antioquia) , light trap, 14 May 1970, C.H.P. 38 $3, 20 ^, same data but tree trunks, Sept. 1970, D.G.Y. 1 o, Curiche (Choco), Malaise trap, 7 July 1967, D.C.Y. Brazil. 11 6*0*, 1 o, Labrea (Amazonas), tree trunks, 9 Oct. 1972, D.G.Y. 2 dV, near Altamira (Para), tree trunks, 1 Oct. 1972, D.G.Y. 1 ^, Itiatuba (Para), tree trunk, 4 Oct. 1972, D.G.Y. Panama. 2 33 , 1 o, Cerro Campana (Panama) , tree buttresses. Discussion: The male parameres of spinosa are rather slender and the dorsal setae are distributed over the distal half of the structure. The coxite tuft consists of 6-8 setae. In L. domascenoi the parameres

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-173are broader, slightly clubbed apically, and the dorsal setae are less numerous, being restricted to the distal third of the structure. The coxite tuft consists of 12+ setae. The females of the two species are very similar, but differ in the relative lengths of the sperm ducts and in the distribution of their sclerotization. The common sperm duct of spinosa is longer than 2 x the length of the individual ducts and the basal one-fourth of the latter is nonsclerotized, unlike damascenoi in which the individual ducts are almost completely sclerotized and the common duct is shorter, less than 2 x the length of the individual ducts. The female cibarium of L. gihsoni has well defined, long horizontal teeth as shown by Pifano and Ortiz (1972) . Those of spinosa and damascenoi are short and irregularly spaced. Adults of L. spinosa are commonly collected on tree trunks. Near Leticia (24-26 July 1973), this species was by far the most abundant sand fly taken in such resting sites.

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Figure 31 LutzomyLa spinosa — A. Male head, B. Male f lagellomere II, C. Female head, D. Female flagellomere II, E. Female cibarium and pharynx, F. Female hind femur showing spines, G. Male genitalia, H. Sperraathecae. I. Female wing, J. Male wing, K. Female cibarium. Male : Panama Canal Zone Female: Same locality as male

PAGE 187

-175X n o / A f to I VO*Ot 1! W c \ , : ;-.i C / w iV>/7 u 'I fe,y?4$0Z~\ ^V^l. xx: 7 4 / ~vk / VI / A n o w. ! s

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•17 71. Individual sperm duct as long as or longer tnan common duct; basal 1/2 thin vailed, not well sclerotized camposi (Fig. 32) Individual sperm duct much shorter than common duct, thick walled, well sclerotized throughout most of its length . .dysponeta (Fig. 33) triacantha 39. Lutzorryia (Pp.) camposi (Fig. 32) Vh.lebotorn.us camposiRodriguez, 1952 (1950): 7 (o* holotype, Quevedo, Los Rios Prov., Ecuador). Fairchild & Hertig, 1958b: 205 (refs.). Fairchild & Hertig, 1959: 121 (Central American records). Arzube, 1960: 157 (Ecuadorian records). Hanson, 1961: 320 et seq. (breeding sites, Panama). Johnson & Hertig, 1961: 765, 773 (rearing data). Sherlock, 1962: 322 (Santander, Colombia). McConnell & Correa, 1964: 526 (infected with gregarines & fungi). Hanson, 1968: 52 (larva, descr., figs.) Phlebotomies acanthobasis Fairchild & Hertig, 1952: 508 (cf, o, Panama, figs.). Fairchild & Hertig, 1958b: 205 (as syonoym of camposi). Lutzomyia. camposi: Barretto, 1962: 95 (listed). Barreto, 1969: 463 (Valle, Colombia). Chaniotis et al . , 1971a: 345 et seq. (pop. dynamics, Panama). Osorno et al . , 1972a: 59 (Colombian records). Chaniotis et al., 1972: 95 (resting sites). Christensen, 1972a: 88 (listed). Christensen et al . , 1972: 57 et seq. (collecting data, Panama). Christensen & Herrer, 1973: 579 et seq. (collecting data). Rutledge & Mosser, 1975: 410 (biting man, Panama). Lewis, 1975a: 502 et seq. (mouthpart morphol.). Herrer & Christensen, 1976a: 62 (collecting data).

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-178Pvessatia oamposi: Forattini. 1971a: 106 (listed). Forattlni, 1973: 123 et sec. (gen. review, figs.). Gomes, 1975: 8 (listed). Distribution: Costa Rica, Panama, Colombia. (Antioquia, Boyaca , Caldas, Choco, Santander, Valle), Ecuador. Material examined: Colombia. 26 (?
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-179 figs.). Rodriguez, 1956: 79 (Ecuadorian records). Fairchild & Hertig, i959: 122 (Central American records). Arzube, 1960: 157 (Ecuadorian records). Hanson, 1961: 320 et seq. (breeding sites). Johnson & Hertig, 1961: 765 (mention). Leon, 1969: 31-33 (, same locality, Shannon trap, 20 Aug. 1967, D.G.Y. 6 do*, 17 o<^, Teresita (Choco), light, Malaise, Shannon traps, tree trunks, March-Dec. 1967, D.G.Y. Panama. 6* holotype (no. 2012), type locality, 13 Dec. 1949, light trap, M. Hertig. p allotype (no. 2012), type locality, 15 Dec. 1949, M. Hertig.

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-180Disaussion: In Panama, Thatcher (1968b) recovered dysponeta larvae from dead leaves and detritus 6+ meters above the ground in trees. Rutledge and Ellenwood (1975a), using soil emergence traps, collected 1 adult which had developed on the open forest floor. The immature stages have not been described. Observations on hosts include pigs (Fairchild and Hertig, 1952) , rodents (Christensen and Hertig, 1973), and horses (Rutledge and Mosser, 1975) . 41. Lutzomyia (Pi'.) triaoantha (Fig. 34) PklebotonvAS triacanthus Mangabeira, 1942a: 119 (
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-181Material examined: Colombia. 1 o (INPES no. 8605), Puerto Lieras, Mapiripan (Meta) , 13 May 1973, A. Morales & Vidales. Brazil. 2 , Rio Napo at Limoncocha (Napo) 21-24 May 1976, light and flight traps, D.G.Y. & T. Rogers. Discussion: The record of L. triaoantha from Venezuela is based on Pifano et al. (1962) who included the male in a key to the phlebotomines of chat country. No additional data were given. Without males, I am unable to separate the females of triacantka from those of dysponeta, although the species probably do not occur together in Colombia or elsewhere.

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Figure 32 Lutzomyia (Pr.) carrtposi — A. Male head, R. Male f lagellonere II, C. Female head, D. Female flagellomere II, E. Female cibarium and pharynx, F. Male genitalia, G. Genital pump and filaments, H. Spermathecae, slightly collpased, normal shape similar to those of Fig. 33H, I. Female wing, J. Male wing, K. Female cibarium. Male: Curiche, Choco Dept., Colombia Female: Same locality as male

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-183vfely fl

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Figure 33 Lutzornyia (Pr. ) dysponeta — A. Male head, B. Male f lagellomere II, C. Female head, D. Female f lagellomere II, E. Female cibarium and pharynx, F. Male genitalia, G. Genital pump and filaments, H. Spermathecae, I. Female wing, J. Male wing, K. Female cihariurn. Male: Teresita, Choco Dept., Colombia Female: Same locality as male

PAGE 197

-185-

PAGE 198

•18( Bai tyi Gr oup Theoclor, 196 5 The two species comprising this group do not" show obvious affinities to other Lutzorrryia species and I prefer to separate them, unlike Forattini (1973) who places them in the subgenus Cororyia Barretto. The female of L. baityi remains unknown but it probably resembles that of L. gorbitzi (Blancas) figured by Blancas (1959-1960) , Fairchild & Hertig (1961b, as hansoni) , and Theodor (1965). At present, only L. baityi is known to occur in Colombia but populations of gorbitzi, discovered in Darien Prov., Panama, may extend into Choco Department. 42. Lutzorrryia baityi (Fig. 34) Phlebotomies baityi Damasceno, Causy, & Arouck, 1945: 22 (o^holotype, Cuiteua, Irituia, Para, Brazil). Barretto, 1947: 188-189 (listed). Damasceno et al . , 1949: 819 (Brazilian records). Floch & Abonnenc , 1952: 34 (o* keyed) . Fairchild & Hertig, 1961b: 2^6 (cf. to hansoni). Pifano et al . , 1962: 386 (d* in key to Venezuelan spp.). Leon, 1968: 31-32 (6" redescr., figs., Venezuela). Lutzomyia baityi: Martins et al., 1963: 334 (Roraima, Brazil). Theodor, 1965: 194 (listed, fig.). Forattini, 1971a: 101 (listed). Forattini, 1973: 274 et seq. (gen. review, fig.). Distributional Colombia (Boyaca) , Ecuador, Venezuela, Brazil. Material examined: Colombia . 1 cf [UV], San Pablo de Borbur (Boyaca), tree hole, 10 March 1945, A. Cast. Brazil. 1 cf, Boa Vista (Roraima), 16 Dec. 1960, J. Silva & L. Ferreira. Ecuador. 3 $$ \ Rio Napo at Limoncocha (Napo) , tree trunks, 18-22 May 1976, D.G.Y. & T. Rogers .

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-187Disoussion: The single Colombian male of baityi, examined by me through the kindness of Dr. Pablo Earreto, is housed in the collection at the University of Valle, Faculty of Medicine, Cali, Colombia. Future search will probably reveal its presence in other localities within the Republic, especially those in Amazonas.

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Figure 34 Lutzormjia (~Pr.) triacantha — A. Male genitalia, B. Spermathecae, C. Female cibarium. Male: Limoncocha, Napo Prov., Ecuador Female: Puerto Lleras, Meta Dept., Colombia Lutzomyia baityi male — D. Genitalia. Male: Limoncocha, Napo Prov., Ecuador

PAGE 201

189

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-190Subgenus V iar.no /rpj ig H angab eir a , 1941 This small distinctive group of four species is treated as a genus by Fcrattini (1971a, 1973). Three species occur in Colombia; the fourth, L. faviasi (Damasceno, Causey, and Arouck) , is known from Brazil and French Guiana. The preferred hosts of these species remain unknown. Specimens are observed usually on tree trunks, sometimes in burrows and at light. Keys to S p ecies Males 1. Style with both proximal spines inserted on long arm or process. Parameres relatively slender 2 Style with proximal spines inserted on separate processes or tubercles. Parameres broader tvberculata (Fig. 36) 2. Parameres slender and downwardly turned apically. . fuvoata (Fig. 35) Parameres enlarged distally, shaped like a goat's head caprina (Fig. 35) Females 1. Spermathecae cylindrical, associated sclerotized processes funnelshaped tuberoutata (Fig. 36) Spermathecae oval, sclerotized processes subequal in width throughout eaprina (Fig. 35) furcata

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-19143. Lutzomyic, (V.) oapvina (Fig. 35) LutsomyCa oapvina Osorno, Morales, & Osornn, 1972c: 437 (6* holotype, veroda El Llano, Municip. Victoria, Caldas, Colombia; p, El Terminal, Municip. Puerto Boyaca, Boyaca, Colombia). Osorno et al . , 1972a: 58 (Colombian records). Gomes, 1975: 8 (listed). Martins et al., 1976b: 496 (Peru) . Phlebotomus furcatus (not furcatus Mangabeira, 1941) : Fairchild, 1943: 571 (Panama records) . Phlebotomus arborealis (not arborealis Floch & Abonnenc, 1944) : Fairchild & Hertig, 1959: 121 (in part, Honduras & Panama records only). Johnson & Hertig, 1961: 765 et seq. (rearing data). Hanson, 1968: 4850 (larva, pupa, descr., figs.). Thatcher, 1968a: 296 (6*, in Disney trap) . Lutzomyia furcata (not furcatus Mangabeira, 1941): Osorno et al., 1970: 11 (Boyaca, Colombia). Christensen 5. Fairchild, 1971: 302 (Darien, Panama). Christensen, 1972a: 88 (listed). Christensen & Herrer, 1973: 579 (in light traps, Panama). Lutzomyia arborealis (not arborealis Floch & Abonnenc, 1944): Chaniotiset al., 1971a: 344 et seq. (pop. dynamics, Panama). Chaniotis et al., 19/2: 95 (resting sites). Distribution: Honduras _, Panama, Colombia (Antioquia, Boyaca, Caldas, Choco) , Peru. Material examined: Colombia. 1 o, Rio Anori (Antioquia), tree buttress, 6 Feb. 1970, C.H.P. 1 o, 1 g, same data but Sept. 1970, D.G.Y. 1 g, Curiche (Choco), 27 July 1967, D.G.Y. 1 g, Alto Curiche (Choco), tree buttress, 31 May 1967, D.G.Y. 1 o. same data but light trap 27 Aug.

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192 1967. 3 op, Teresita (Choco) , light trap & tree buttresses, May & Sept. 1967, D.G.Y. Honduras. 1 <$ , 4 oo, Tela, Lancetilla Valley, light traps 6 tree buttresses, Dec. 1953 & Jan. -Feb. 1954, V.'. Hils. Panama. 10 do', 20 go from various localities in the Canal Zone, Bocas del Toro Prov. , Color) Prov., Darien Prov., & Panama Prov., tree buttresses & light or Shannon traps. Discussion: See L. furcata. 44. Lutzomyia (V.) furcata (Fig. 35) Phlebotomus furcatus Mangabeira, 1941b: 256 (cf holotype, Aura, Belem, Para, Brazil). Barretto, 1947: 201 (refs.) . Barretto, 1950a: 104 (keyed). Barretto, 1951: 217 (Brazilian records). Floch & Abonnenc, 1952: 31 (keyed), 78-80 (c?, redescr., figs., French Guiana). Fauran, 1960: 8, 9 (mention. Forattini, 1960: 479 (Amapa, Brazil). Pifano et al., 1962: 38 3 (c? keyed) . Phlebotomus arborealis Floch & Abonnenc, 1944a: 8 (o, Baduel, French Guiana). Barretto, 1947: 186 (refs.). Floch & Abonnenc, 1952: 43 (keyed), 179-181 (redescr.. figs.). Fairchild, 1955: 190 (as probable o of furcatus). Fauran, 1960: 9 (mention). Lutzomyia fur cata: Barretto, 1962: 94 (listed). Martins et al., 1963: 334 (Roraima, Brazil). Martins et al., 1965: 3 (Rondonia, Brazil). Wijers & Huisenga, 1967: 397 (cf. to tuberculata) . Osorno et al., 1972a: 58-59 (Colombian records). Osorno et al . , 19 72c: 441 (cf. to caprina) . Shaw & Lainson, 1972: 713 (mention). Lewis, 1975a: 502 et seq . (mouthpart morphol.). Llanos et al., 1976: 480 (Peru). Lainson et al., 1977 (Mato Grosso, Brazil).

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-193butzomyia arbovealis: Barretto, 1962: 94 (as synonym of furcata) . Viannamyia furcata: Forattini, 1971a: 106 (listed). Forattini, 19 73: 123 et seq. (in part, refs. based on material from Panama not furcata Mang., gen. review, figs.). Distribution: Brazil, French Guiana, Venezuela, Peru, Colombia (Caqueta, Santander) . Material examined: Colorrbia. 1 6* (INPES no. 3825), Vereda La Nevera, Solano (Caqueta), tree hole, 25 Aug. 1969. Brazil. 1 d 1 , 1 o, Serra do Navio (Araapa), tree trunks, 21 Sept. 1972, D.G.Y. & H. Fraiha. 1 if, 2 oo., Labrea (Amazonas) , tree trunk, 9 Oct. 1972, D.G.Y. 2 o'o", 2 cj>o , Rio Aripuana at Humboldt (Mato Crosso) , tree trunks, 17-20 Aug. 1974, D.G.Y., J. Shaw, & II. Fraiha. 1 6*, Belem (Para), 3 Aug. 1964, F. Guimaraes. 1 o, same locality, light trap, 18 Aug. 1972, D.G.Y. 2 o^, same locality, flight traps, 29 July-6 Aug. 1974, D.G.Y. 1 ^, Itiatuba (Para), light trap, 4 Oct. 1972, D.G.Y. 1 o. , 27 km E of Maraba (Para), light trap, 27 Setp. 1972, D.G.Y. & H. Fraiha. 1 d", 1 £, W of Altamira, km 164 (Para), tree trunk, 9 Nov. 1974, J. Reinert. Discussion: The distal part of the parameres of L. furcata (description of Mangabeira, 1941b, and males from Brazil, French Guiana, and Caqueta, Colombia) is long and slender, downwardly curved with a short, stout apical spine (Fig. 35G) . In contrast, the parameres of L. caprina (description of Osorno et al . , 1972c, and male from Antioquia, Colombia) are broader apically and lack a stout terminal spine (Fig. 35H) . The occurrence of caprina in Loreto Prov. , Peru (Martins et al . , 1976b) suggests that it and furcata are distinct taxa. The females, however, from these and all other localities are morphologically inseparable, the record of fui'cata (o only) in Antioquia Dept . (Osorno et al., 1972a) probably representing not that species but L. caprina.

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-194Without additional material from Central America, the identity of the specimens from Panama and Honduras remains uncertain. Panamanian males generally resemble capr-ina in the shape of the parameres but each terminates in an apical spine, longer than that of fuTCata. The male from Honduras, on the other hand, more closely resembles furcata in paramere shape but again the apical spine is much longer. It is difficult to determine whether or not these character states reflect inter or intraspecific variation but for the present I arbitrarily treat specimens from Panama and Honduras as variants of caprina, primarily on the basis of distribution. Lutzomyia furcata was included in a key to the phiebotomines of Venezuela (Pifano et al . , 1962) but no additional information was given. 45. Lutzomyia (V.) tuberoulata (Fig. 36) P'hlebotomus tuberculo.tus Mangabeira, 1941b: 251 (6* holotype, Aura, Belem, Para, Brazil). Barretto, 1947: 227 (refs.). Damasceno et al., 1948: 699 (cf. to f arias i) . Damasceno et al . , 1949: 833 (Brazilian records). Barretto, 1950a: 104 (keyed). Barretto, 1951: 225 (distrib.) Floch & Abonnenc, 1952: 31 (keyed). Fauran, 1960: 8-9 (6", redescr., figs.). Phlebotomus sp. x Floch & Abonnenc, 1944a: 10-12 (o, French Guiana). Floch & Abonnenc, 1952: 43 (keyed), 181 (redescr., figs.). Barretto, 1962: 94 (as female of tubex'cu'lata) . Lutzomyia. tuberoulata: Barretto, 1962: 94 (listed). Martins et al., 1963: 335 (Roraima, Brazil). Theodor, 1965: 190 (6*, £, figs.). Barreto, 1969: 467 (Valle, Colombia). Lewis et al . , 1970: 215 (ovaries

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-195& accessory glands). Christensen 1972a: 38 (Panama). Osorno et al., 1972a: 59 (Colombian records). Lewis, 1975a: 502 et seq . (mouthpart morphol.). Lainson et al., 1977 (Mato Grosso, Brazil). Lutzomyia munangai Wijers & Huisenga, 1967: 395 (d 1 holotype, near Moengo, Surinam), Lewis, 1975a: 511 (as synonym of tubsroulata) . Viannamyia munangai: Forattini, 1971a: 106 (listed). Forattini, 1973: 516 (cf. to tuberculata.) . Gomes, 1975: 8 (listed). Viannamyia tuberculata: Forattini, 1971a: 106 (listed). Forattini, 1973: 507 et seq. (gen. review, figs.). Gomes, 1975: 8 (listed). Distribution:. Panama, Colombia (Antioquia, Caqueta, Choco, Valle), French Guiana, Surinam, Brazil. Material examined: Colombia. 1 o, Rio Anori (Antioquia), tree trunk, 4 March 1971, C.II.P. 1 £, Curiche (Choco), Malaise trap, 25 July 1967, D.G.Y. 2 00, Alto Curiche (Choco), 7 July 1967, D.G.Y. 1 o, Anchicaya + T "f Dam (Valle), tree trunk, 9 Aug. 1973, D.G.Y. & R.C.W. 6 JcT, 8 00, 25 km E of Buenaventura (Valle), tree trunks & light trap, 11-12 Aug. 1973, D.G.Y. & R.C.W. Brazil. 2 op, Serra do Navio (Amapa) , tree trunks, 21 Sept. 1972, D.G.Y.& H. Fraiha. 1 0, W of Altamira (Para), light trap, 1 Oct. 1972, D.G.Y. 2 00, Belem (Para), light trap, 18 Oct. 1972, D.G.Y. 2 06 1 . 5 00, Belem (Para), tree trunks, flight trap, 29-31 July, 1974, D.G.Y. 3 0*0*, 2 00, Rio Aripuana at Humboldt (Mato Grosso), tree trunks, 16-19 Aug. 1974, D.G.Y., J. Shaw, & H. Fraiha. Panama. 2 00, Almirante (Bocas del Toro) , Aug. -Sept. 1951, A. Quinones. Discussion: Lutzomyia tuberculata, a little known species, is not considered to be anthropophilic although Barreto (1969) reported the capture of a single female on man in Valle Dept., Colombia. Both sexes are easily recognized.

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Figure 35 Lutzomyia (V.) oaprina — A. Male head, B. Male flagellomere II, C. Female head, D. Female flagellomere II, E. Female cibarium and pharynx, F. Male genitalia, H. Tip of paramere of male from Rio Anori, Antioquia Dept., Colombia, I. Genital pump and filaments, J. Spermathecae, K. Female wing, L. Male wing, M. Female cibarium. Male: Panama Canal Zone (except Fig. 35H) Female: Curiche, Choco Dept., Colombia Lutzomyia (V. ) furcata — G. Tip of paramere. Male: Para State, Brazil

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•197 "X

PAGE 210

Figure 36 Lutsonyia (V.) tubevoulata — A. Male head, B. Male flagellomere II, C. Female head, D. Female flagellomere II, E. Female cibarium and pharynx, F. Male genitalia, G. Spermathecae, H. Female wing, I. Male wing, J. Female cibarium. Male: Valle Dept., Colombia Female: Curiche, Choco Dept., Colombia

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-199-

PAGE 212

-200Shanno ni Group The odor, 1965 Barret to (1962) created the subgenus Psathyromyia to accomodate L. shanncmi and rits allies, some of which (the aragaoi group species) were separated later by Theodor (1965) . Rased on both sexual and nonsexual characters, the shannon!, and aragaoi species are very similar as noted by Fairchild (1955) and Barretto (1962). Forattini (1971a, 1973), for reasons not fully understood, places the species in the aragaoi group in the genus Psychodopygus but retains L. channoni and its relatives in the genus Lutzorryia in the expanded subgenus Trickopygomyia Barretto. This seems to be unnecessarily artificial ana I prefer to follow the arrangement of Theodor (1965) and Lewis et al. (1978). Six of the 14 described shannoni group species occur in Colombia but others such as L. lutziana (Costa Lima), L. campballi (Damascene, Causey & Arouck) , and possibly I. soaffi (Damascene & Arouck) may eventually be discovered in the Amazonian region of the Republic. Keys to Species Males 1. Paramere with a basal tuft of long setae implanted on a short dorsal process dasyrrera (Fig. 33) Paramere simple, lacking a setiferous dorsal process 2 2. Antennal ascoids with short, blunt posterior spurs punetigenieulata (Fig. 40) Antennal ascoids with long pointed posterior spurs 3

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-2013, Paramere with sinuous dorsal setae undulata (Fig. 42) Paramere without such dorsal setae 4 4. Paramere with dorsal setae restricted to apical third of structure . , abonnenci (Fig. 37) Paramere with dorsal setae covering at least the apical half of structi i 5. Paramere us in Fig. 39A. The apical dorsal setae slightly foliaceous and recurved unlike the shorter proximal setae, dendrophyla (Fig. 39) Paramere otherwise (Fig. 41G) , the dorsal setae unmodified, more or less erect shannoni (Fig. 41) F emales 1. Cibarium with 6+ horizontal teeth . 2 Cibarium with 4 horizontal teeth* shannoni (Fig. 41) dendrophyla (Fig. 39) abonnenci 2. Spermathecae annulated undulata (Fig. 42) Spermathecae smooth-walled, without annulations 3 *Morales et al. (1974) noted that the common sperm duct of dendrophyla is shorter than that of shannoni (0.049-0.099 mm as opposed to 0.1450.198 mm). This may be helpful in separating the taxa but without associated males I nesitate to identify females to species.

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-2023. Antennal ascoid wich short blunt posterior spur. Individual sperm duct, about half length of common duct and less than 2 >: length of spermatheca punctigenioulata (Fig. 40) Antennal ascoid with long pointed posterior spur. Individual sperm duct much longer than common duct and over 3 x length of spermatheca dasymsra (Fig. 38) 46. Lutzomyia abonn&noi (Fig. 37) Fhlebotomus abonnenoi Floch & Chassignet, 1947c: 1 (
PAGE 215

203Material examined: Colombia. 1
PAGE 216

-204Distribution: Mexico, Nicaragua, Cost'.. Rica, Panama, Colombia (Antioquia, Boyaca, Choco, Valle) , Venezuela. Material examined: Colombia. 1 o, Curiche (Choco), tree buttress, 5 July 1957, D.G.Y. 2 <$<$, same data but 22 Sept. 1967. 1
PAGE 217

-205(d"', keyed, redescr.). Barretto & Duret, 1953: 344 (cf. to microcephalus) . Vargas & Dlaz-Najera, 1959: 146 (mention). Forattini, 1960: 476 (Amapa, Brazil). Pifano et al . , 1962: 385 (6* keyed) . Ortiz, 1965c: 416 (mention). Ortiz, 1972a: 23, 24 (listed). Pifano et al., 197 3: 158 (mentioo) . Lutzornyia dendvophila. (or dendx-ophyla) : Barretto, 1962: 99 (listed). Martins & Silva, 1964: 128 (Acre, Brazil). Martins et al . , 1965: 2 (Rcndonia, Brazil). Lewis et al . , .1970: 215 (parous rates). Forattini, 1971a: 102 (listed). Osorno et al., 1972a: 51 (Colombian records). Martins & Morales, 1972: 36/ (listed). Llanos, 1973: 30 (Peruvian record). Forattini, 1973: 293 et seq. (gen. review, figs.). Velasco, 1973: 85 (Bolivia). Morales et al., 1974: 443 (^ descr., figs., Colombia). Ward & Killick-Kendrick, 1974: 216 (Para, Brazil). Lewis, 1975a: 502 et seq. (mouthpart morphol.). Llanos et al . , 1975b: 670 (Peru). Uard & Ready, 1975: 128 et seq. (egg structure). Llanos et al . , 1976: 480 (Peru). Martins et al . , 1976b: 496 (Peru). Lainson et al., 1977 (Mato Grosso, Brazil). Distribution: Colombia (Amazonas, Caqueta, Meta), Ecuador , Peru, Bolivia, Venezuela, French Guiana, Brazil. Material examined: Colombia. 1 63, 9 ocj>, 17 km W of Leticia (Amazonas), tree trunks, 24-26 July 1973, D.C.Y. & R.C.W. Bolivia. 1 6\ 1 o, 5 km N of Tajlewi (Larecaja) , tree trunks, 13 Aug. 1971, J. Velasco. 1 d", 1 o, Misquirnayo (Sud Yungas) , chicken coop, 1700 m above sea level, 3 Sept. 1971, J. Velasco. Brazil. (all on tree trunks). 5 c?cf, Serra do Kavio (Amapa), 21 Sept. 1972, D.G.Y. & H. Fraiha. 12 6"d\ Labrea (Amazonas), 9-10 Oct. 1972, D.C.Y. 72 o'c?, Rio Aripuana at Humboldt (Mato Grosso), 13-20 Aug. 1974, D.G.Y. , J. Shaw, & H. Fraiha. 3 S3, 27 km

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-206SE of Maraba (Para), 26-28 Sept. 1972, D.G.Y. & H. Fraiha. 1 6*, W of Altamira (Para), 1 Oct. 1972, D.G.Y. 5 6*0*, Belern (Para), 29 July-1 Aug. 1974, D.G.Y. Ecuador. 90 cf6*, Rio Napo at Limoncocha (Napo) , tree trunks, 19-24 Kay 1976, D.G.Y. & T. Rogers. French Guiana. 1 6*, Baduel, Dec. 1947, no other data. Peru. 2 6*6*, Cachicote (Huanuco) , 12 Sept. 1965, J. Hitchcock. Discussion: Lutzormjia dendrophyla is an Amazonian species which occurs often but not always with L. shannoni where their ranges overlap. We collected only dendrophyla at Leticia, the spermathecae (Fig. 39B) drawn from a female taken from there. The females of both species occasionally bite man (Lainson et al . , 1977; Forattini, 1973) and are very difficult to separate in the absence of associated males. 49. Lutzomyia puna tig eniculata (Fig. 40) Phlebotomus punctigeniculatus Floch & Abonnenc, 1944b: 5 (6* holotype, near Cayenne, French Guiana). Barretto, 1947: 220 (refs.). Floch & Abonnenc, 1948b: 1, 12, 24 (Venezuela, 0*, figs.). Damasceno et al . , 1949: 831 (Brazilian records). Fairchild & Hertig, 1950: 524 et seq. (measurements, taxonomy, figs.). Barretto, 1951: 222 (distrib.). Pifano & Ortiz, 1952: 144, 146 (keyed). Floch & Abonnenc, 1952: 37, 46 (keyed), 153 (6*, o, redescr.). Barretto & Duret, 1953: 344 (mention) Pifano et al., 1962: 385, 388 (6", o, keyed), 397 (6*, o, redescr., refs., figs., Venezuela). Ortiz, 1965c: 416 (mention). Leon, 1968: 30 (listed). Ortiz, 1972a: 23 (mention). Pifano et al., 1973: 158 (mention) .

PAGE 219

207Phlebotomus chrisiophersoni Damasceno & Causey, 1944: 347 (
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-208The male resembles L. saaffi (Damasceno & Arouck) , known from Brazil grid Peru, in the shape of the genitalia but the ascoids of that species have long, pointed posterior spurs. The female of saaffi, discovered but unde scribed (unpublished data) resembles that of puna tig eniculata in spermathecae and color (pleura moderately infuscated) but there are only 4 horizontal teeth in the cibarium. 50 . Lutzonyia sharmoni (Fig. 41) Phleboiornus sharmoni Dyar, 1929: 121 (d\ Cano Saddle, Gatun, Canal Zone). Barretto, 1947: 222-223 (full refs., synonyms). Fairchild & Hertig, 1950: 523 et seq. (collecting data, review of habits, c?, o, figs.). Forattini, 1954: 214 et seq. (sternites, figs.). Fairchild a Hertig, 1959: 122-124 (distrib.). Johnson h Hertig, 1961: 765 et seq. (rearing data) . Hanson, 1968: 78 (larva, pupa, descr., figs.). Morales et al., 1969a: 380 (Meta, Colombia). Lutzomyia shannoni: Martins et al., 1961b: 311 (Brazil). Theodor, 1965: 189 (listed, 6", o, figs.). Osorno et al., 1972a: 52-57 (Colombian records). Forattini, 1973: 84 et seq. (gen. review, refs., figs.). Zeledon & Alfaro, 1973: 416 (infected with promastigotes) . Ward et al., 1973: 178 (biting man, Para, Brazil). Morales et al . , 1974: 445-446 (o cf. to dendr'ophyla) . Williams, 1976a: 604 (in caves, Belize). Miles et al., 1976: 532 (mating aggregation, Panama). Distribution: SE United States to Paraguay, Coloraoia (Antioquia, Bolivar, Boyaca, Caldas , Caqueta, Choco, Cundinamarca, Guijira, Magdalena, Norte de Santander, Santander, Tolima, Valle) . Material examined: Colorrbia. 2 6V, Rio Anorl (Antioquia), light traps, May 1970, C.H.P. 14 6"cf, 22 oo., same data but tree trunks, Sept.

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-2091970, D.G.Y. 238 d"d\ 164 po, Curicha (Choco) , light Malaise, & Shannon traps, biting man & tree trunks, April-Dec. 1967, D.G.Y. 2 S3, 6 oo. , Alto Curiche (Choco), tree trunks & Shannon trap, May, July-Aug. 1967, D.G.Y. 268 <$£, 167 no, Teresita (Choco), light, Malaise, & Shannon traps, tree trunks, March-Dec. 1967, D.G.Y. 1
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-210preference studies in Panama (Tesh et al . , 1972) revealed that 78.5% of 312 blood meals reacted with rodent antiserum. At Curiche, we collected only one shannoni female biting man from April to November 1967 (141 man hours). As expected, most specimens (232 6*6*, 155 yy) were captured on tree trunks where only L. trinidadensis was taken in greater numbers (333 6*o*, 301 on) during the same time period. 51. Lutzomvia undulata (Fig". 42) Phlebotomus undulatus Fairchild & Hertig, 1950: 527 (6* holotype, near Esquintla, Guatemala). Vargas & Diaz-Najera, 1953b: 313 (Mexico). Fairchild & Hertig, 1959: 122 (Central American records). Fairchild & Hertig, 1961b: 244 (cf. to cvatifer) . Rosabal & Trejos, 1965: 222 (listed). Thatcher, 1968a: 296 (eft? in Disney trap). Phlebotomus humboldit Vargas & Diaz-Najera, 1959: 143 (6\ o_, Oaxaca. Mexico). Rosabal & Trejos, 1964: 169 (as synonym of undulatus). Lutzomyia undulata: Barretto, 1962: 99 (listed, as synonym of humboldti) . Williams, 1970: 333 et seq. (Belize, negative for flagellates). Chaniotis et al., 1971a: 344 et seq. (pop. dynamics, Panama). Forattini, 1971a: 102 (listed). Tesh et al., 1971a: 152 (blood meals). Tesh et al., 1972: 90 (blood meals). Martins & Morales, 1972: 355 (listed). Chaniotis et al . , 19 72: 95 (resting on or in trees). Christensen, 1972a: 88 (listed). Christensen et al . , 1972: 57 (collecting data). Forattini, 1973: 294 et seq. (gen. review, figs.). Christensen & Herrer, 1973: 579 (in light trap). Lutzomyia cvatifeva: Osorno et al . , 1972a: 61 (not cvatifer Fairchild & Hertig, 1961b, Boyaca, Colombia).

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-211Distribution: Mexico, Guatemala, Belize, El Salvador, Panama, Colombia (Koyaca, Choco, Narino), Ecuador. Material examined: Colombia. 1 o, Curiche (Choco), tree buttress, 15 Aug. 1967, D.G.Y. 1 cf, 2 op, Rio Aguaclara, Tumaco (Narino), light trap, 21 July 1969, P. Barreto. Ecuador. 1 o, 17 km E of Santo Domingo de los Colorados (Pichincha) , tree trunk, 27 May 1976, D.G.Y. Guatemala. 3 eft? (paratypes & holotype) , near Esquintla, tree buttresses, May, June, 1945. Panama. 1 b*, Rio Tuira at mouth of Rio Paya (Darien) , Shannon trap, 25 Feb. 1958, P. Galindo. Discussion: Lutzomyia undulata is generally rare throughout its range, the relatively few specimens having been captured in or on trees and in light traps. Using the precipitin test, Tesh et al. (1971a, 1972) in Panama found that blood meals from recently engorged females reacted with antisera from rodents, marsupials, and carnivores. The females of undulata and L. cratifer (Fairchild & Hertig) are morphologically inseparable, the record of the latter species in Colombia (Osorno et al., 1972a) probably referring to undulata, the males of which have been found in Colombia and Panama. Males of cratifer are known only from Mexico and Honduras .

PAGE 224

Figure 37 Lutzomyia abormenoi male — A. Head, B. Flagellomere II, C. Wing, D. Genitalia, E. Genital pump and filaments. Male: Teresita, Choco Dept . , Colombia

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Figure 38 Lutzomyia (lasymeva — A. Male head, B. Male flagellomere II, C. Female head, D. Female flagellomere II, E. Female clbarium and pharynx, F. Genital pump and filaments, G. Male genitalia, H. Body of spermatheca, I. Spermathecae, J. Female wing, K. Male wing, L. Female c ibarium. Male: Rio Anori, Antioquia Dept., Colombia Female: Panama Canal Zone

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•215-

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Figure 39 Lutzomyla dendrophyla — A. Male genitalia, B. Spermatbecae, C. Female cibarium. Male: Leticia, Amazonas Comisaria, Colombia Female: Same locality as male

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-217-

PAGE 230

Figure 40 Lutzomyia punctigeniculata — A. Male head, B. Male flagellomere II. C. Female head, D. Female flagellomere II, E. Female cibarium and pharynx, F. Paramere, G. Male genitalia, II. Body of spermatheca, I. Spermathecae, J. Female wing, K. Male wing, L. Female cibarium. Male: Curiche, Choco Uept., Colombia Female: Panama Canal Zone

PAGE 231

-219.4 1? |/(J W o" / WS-;Z^ ML b / / "/ / I / / 1 / I I L

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Figure 41 Lutzomyia shannoni — A. Male head, B. Male flagellomere II, C. Female head, D. Female flagellomere II, E. Female cibarium and pharynx, F. Spermathecae, G. Male genitalia, H. Body of spermatheca before maceration in NaOH, Alachua Co., Florida, U.S.A., I. Female wing, J. Male wing, K. Female cibarium. Male: Teresita, Choco Dept., Colombia Female: Same locality as male (except Fig. 41H)

PAGE 233

-221i W 3 //ill < X I ° / f\ ;,1X3 C W :. W 9 /i ^ (j ,/ // i7 J \

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Figure 42 Lutzomyia undulata — A. Female head, B. Female flagellomere II, C. Ascold on female flagellomere III, D. Female cibarium, E. Spermathecae, F. Female wing. Male: Chiapas State, Mexico Female: Curiehe, Choco Bept., Colombia

PAGE 235

-223-

PAGE 236

224Longispina Group Theodo r, 3965 This group corresponds to the subgenus Trichopygomyia Barretto, 1962, later enlarged by Porattini (1971a, 1973) to accomodate 30+ additional species. Seven of the 8 longispina group species occur in or near the Amazon Basin. Lv.tzomyia triramula, a rowans-Andean species, ranges as fat north as Belize and is especially common in lowland foiests in Panama and northwestern Colombia. Most species seem to be readily attracted to light, often inhabit mammal burrows but do not attack humans. Two longispina group species, apparently allopatric, have been reported in Colombia (Osorao et al., 1972a). Keys to Sp ecies Males 1. Parameres as shown, trifurcate tvivaxnula (Fig. 44) Parameres otherwise, bifurcate longispina (Fig. 43) Females 1. Length of common sperm duct less than 2 x width of spermatheca. Cerci broad apically longispina (Fig. 43) Length of common sperm duct greater than 2 x width of spermatheca. Cerci slender or constricted apically tiri.ramula (Fig. 44)

PAGE 237

>2552. Lutzomyia longispina (Fig. 43) Phlebotomus longispinus Mangabeira, 1942a: 186 (c? holotype, Aura, Belem, Para, Brazil). Mangabeira, 1942d: 251 (iramatures descr.). Barretto, 1947: 209 (catalog). Causey & Damasceno, 1948: 628 et seq . (cf. to wagleyi, 6* fig.). Damasceno et al . , 1949: 827 (Brazil). Vargas & Diaz-Najera, 1951a: 22 (cf. to pratti) . Pifano & Ortiz, 1952: 142 (Bolivar, Venezuela). Floch & Abonnenc, 1952: 33, 48 (cf, y keyed). Fairchild & Her tig, 1952: 518 (cf. to triramulus) . Pifano et al . , 1962: 385, 389 (cf, o keyed). Sherlock & Pessoa, 1964: 333 (Bahia, Brazil). Hanson, 1968: 88 (larva, cf. to triranulus) . Lutzomyia longispina: Barretto, 1962: 98 (listed). Theodor, 1965: 189 (d", figs.). Martins et al . , 1965: 14 (cf. to rondoniensis) . Osorno et al . , 1972a: 57 (Caqueta, Colombia). Forattini, 1973: 172 et seq. (gen. review, figs.). Lewis, 1975a: 502 et seq. (mouthpart morphol.). Ramirez et al., 1976: 602 (mention). Martins et al . , 1976a: 491 (cf. to elegans) . Distribution: Brazil, Venezuela, Colombia (Caqueta) . Material examined'. Colombia. 1 o (INPES no. 7068), San Miguel, Municip. Florencia (Caqueta), tree hole, 2j June 1971, E. Osorno & A. Morales. Brazil. 1 6*, Cacheira (Bahia), armadillo burrow, 7 July 1959, Afonso. 1
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-226individual sperm ducts are wrinkled in part, unlike those of L. triramula, but this could be a result of mounting technique and not a reflection of normal structure. The spermathecae (Fig. 43A) are larger than those of triramula and the cerci are broader apically (Fig. 43C) . 53. Lutzomyia tvivcxula (Fig" 44) Phlebotomus trivaniulus Fairchild & Hertig, 1952: 517 (d* holotype, p, Rio del Medio, near Rio Gatun, Colon. Prov., Panama). Johnson & Hertig, 1961: 765, 775 (rearing data, biting man in laboratory). Hanson, 1968: 86 (larva, pupa, descr.). Sherlock, 1962: 321 (Santander, Colombia). Barreto, 1969: 467 (Valle, Colombia). Lutzomyia triratnula: Barretto, 1962: 98 (listed). Martins et al., 1965: 14 (cf. to rondoniensis) . Williams, 1970: 333 et seq. (summary of collecting data, Belize). Chaniotis et al., 1971a: 344 (pop. dynamics, Panama). Chaniotis et al., 19 72: 95 (in burrows). Osorno et al., 1972a: 57 (Colombian records). Christensen, 1972a: 88 (listed). Forattini, 1973: 123 et seq. (gen. review, figs.). Chaniotis & Correa, 1974: 115 (in light traps). Lewis, 1975a: 503 et seq. (mouthpart morphol.). Rutledge et al., 19 75: 180 (in light traps). Martins et al., 1976a: 491 (cf. to elegans) . Herrer b. Christensen, 1976a: 62 (collecting data, Panama). Ramirez et al., 1976: 602 (mention). Distribution: Belize, Panama, Colombia (Autioquia, Choco, Santander, Valle). Material examined: Colombia. 78 Stf, 40 op, Rio Anori (Antioquia) , light & flight traps, April-June 1970, CM. P. 30 &d, 54 oo, same locality light traps, Sept. 1970, D.G.Y. Ill 6*c?, 277 oq_, Curiche (Choco), light,

PAGE 239

-227Malaise, & Shannon traps, April-Nov. 1967, D.G.Y. 4 6<3 , 19 o^, Alto Curiche (Choco), July-No v. , 1967, U.G.Y. 1 d*, Teresita (Choco) , tree trunk, 23 Jane 1967, D.G.Y. 1 cf, same locality, light trap, 8 Oct. 1967, D.G.Y, 1 6", 2 oq, 25 km E of Buenaventura (Valle) , tree trunks, 11 Aug. 1973, D.G.Y. & R.C.W. 5 0*0*, 13 o^, same locality, light & flight traps, ]2 Aug. 1973, D.G.Y. & R.C.W. Panama. 1 6* (holotype no. 1754), type locality, tree buttress, 14 Oct. 1949, R. Hartmann. 1 y (allotype no. 2376), La Victoria, Cerro Jefe (Panama Prov.), Shannon trap, 20 Aug. 1950, M. Hertig & P. Galindo. Numerous other specimens, most of which were iisted by Fairchild & Hertig (1952). Discussion: Females of triramv.la from a population E of Buenaventura vary from those in Antioquia and Choco Dapts., Colombia, and Panama in the shape of the cerci. They are relatively slender (Fig. 44C) ; more so than those of longispina (Fig. 43C) and lack the nipple-like end which is characteristic of females from the more northern localities. The males from near Buenaventura are generally similar to those from other localities but the main lobe of the paramere is more recurved and slender apically. I attribute these differences to geographic variation and for the present, at least, regard them all as conspecific.

PAGE 240

Figure 43 Lutzomyia longispina — A. Spermathecae, B. Female cibarium, C. Female cercus, I). Male genitalia. Male: Bahia State, Brazil Female: Florencia, Caqueta Intendencia, Colombia

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>29-

PAGE 242

Figure 44 Lutzomyia trtvamula — A. Male head, B. Male flagellomere II, C. Female head, D. Female flagellomere II, E. Female cibarium and pharynx, F, Female cercus, G. Genital pump and filaments, same scale as Fig. 44H, H. Male genitalia, I. Spermathecae, J. Female wing, K. Male wing, L. Female cercus, specimen from Valle Dept., Colombia, M. Femalei cibarium. Male: Curiche, Choco Dept., Colombia Female: Same locality as male (except Fig. 44L)

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-23J V \ XSA

PAGE 244

-232Avagaoi Croup Theo do r, 1965 Theodor (1965) divides these species into 2 categories, the series avagaoi (7 spp.) and series bvasitiensis (4 spp.), based upon the spermathecae which are sac-like in the series avagzoi females, tubular in those of the bvasiliensis series. Fairchild (1955), Barretto (1962), Ortiz (1972a) , and others combine the avagaoi group species with those in the sharmoni group on the basis of several characteristics, the latter two authors placing them in the subgenus Psatkyvonyla Barretto, 1962. The status of L. pifa.noi (Ortiz), a Venezuelan species is not yet clear. It certainly belongs in the aragaoi group but is known from a single male which somewhat resembles L. vv.noid.es . Lutzomyia abunaensis Martins, Falcao and da Silva (1965), from Brazil and Ecuador, also described from the male, is similar to that of L. bvasiZiensis (Costa Lima, 1932) . I examined a male and several females of abunaensis , to be described in another paper, from Limoncocha, Napo Prov., Ecuador. The females differ from those of bvasiliensis in having 8+ horizontal cibarial teeth instead of 4 and by the shorter sperm ducts. The spermathecae of both females are similar in shape and size. The presence of one or both of these species in Colombia would not be surprising. Ke ys to Specie s M ales 1. Genital filament tips hooked or twisted 2 Genital filament tips simple, pointed, or spear shaped .3

PAGE 245

-2332. Style with isolated basal spine. Genital filament tips shaped like a burton hook. Aedeagus rao.re slender. Paramere as shown . . carpenteri (Fig. 48) Style with paired or nearly paired basal spines. Genital filament tips twisted like a corkscrew. Paramere variable but never as above aragaoi (Fig. 45) 3. Genital filament tips markedly inflated, spear-shaped. Coxite without scattered setae on inner surface runoides (Fig. 49) Genital filament tips simple, not inflated. Coxite with scattered setae on inner surface 4 4. Lateral lobes greater than 0.56 mm. Parameres as shown, the ventral "heel" better developed barrettoi majuscula (Fig. 4 6) Lateral lobes less than 0.50 mm. Parameres otherwise, more slender, lacking a ventral heel bavrettoi barrettoi (Fig. 47) females 1. Spermathecae tubular with convoluted individual ducts runoides (Fig. 49) Spermathecae sac-like, much wider than nonconvoluted ducts, .... 2 2. Spermathecae pear shaped with an expanded neck, individual ducts with fine transverse striations carpenteri (Fig. 48) Spermathecae otherwise, ovoid or spherical, individual ducts saiooth walled 3

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-2343. Spermathecae smaller, subspherical with apical "hairs" not arising from a differentiated area. . . , avagaoi (Fig. 45) Spermathecae larger, ovoid with apical "hairs" arising from a raised circular area 4 4. Spermathecae larger, their ducts expanded where they enter spermathecae bawettoi bavvettoi (Fig. 47) Spermathecae smaller, their ducts more slender, subequal in width throughout barrettoi majuscula (Fig. 46, 4 7) Series avagaoi 54. Lutzomgia avagaoi, (Fig". 45) Phlebotomus avagaoi Costa Lima, 1932: 48 (6" Lassance, Minas Gerais, Brazil). Barretto, 194 7: 185-186 (full refs.). Floch & Abonnenc, 1952: 131-134 (in part, c? only) . Fairchild & Hertig, 1953a: 21 et seq. (o", o, redescr., figs., keyed, refs., Panama). Vargas & Diaz-Najera, 1959: 146 (cf. to humboldti). Fairchild & Hertig, 1961b: 250 (o. cf. to botella) Lucena & Almeida, 1964: 187 et seq. (cf, o redescr., figs.). Sherlock & Pessoa, 1964: 333 (resting sites, Brazil). Lucena, 1967: 271 (mention). Ortiz, 1972a: 23-25 (listed, cf. to pifanoi) . Phlebotomus keckenvothi Floch u AbonneiiCj 1942b: 8 (o, French Guiana) Barretto, 1946c: 430 (listed). Floch & Abonnenc, 1952: 47, 176 (c. keyed, redescr., figs.). Fairchild & Hertig, 1953a: 21 et seq. (o keyed, cf. to avagaoi, refs.). Ortiz, 1972a: 23 (listed). Pifano et al . , 1973: 158 (listed). Forattini, 1973: 456 (as synonym of avagaoi)-

PAGE 247

-235Lutzomyia aragaoi: Barretto, 1962: 99 (listed). Theodor, 1965: 186 ((?, o, figs). Martins et al., 1965: 2 (Rondonia, Brazil). Barreto, 1969: 462-463 (Valle, Colombia). Lev/is et al., 1970: 215 (parous rates). Chanictis et al., 1971a: 344 et seq. (collecting data, Panama). Christensen, 1572a: 88 (listed). Osorno et al., 1972a: 37 (Vichada, Colombia). Chcmiotis et al., 1972: 95 (resting sites). Lewis, 1975a: 501 (inouthpart morphol., cibarium fig.). Martins et al . , 1976a: 488 (Peru) . Lutzomyia heckenrothi: Barretto, 1962: 99 (listed) . Martins & Morales, 1972: 367 (listed). Psychodopygus aragaoi: Forattini, 1971a: 105 (listed). Forattini, 1973: 416 et seq. (gen. review, figs.). Distribution: Panama, Colombia (Amazonas, Antioquia, Choco, Valle, Vichada), Peru, Brazil, Paraguay, French Guiana, Trinidad. Material examined: Colombia. 1 6", 17 km W of Leticia (Amazonas) , light trap, 26 July 1973, D.G.Y. & R.C.W. 2 66, 1 g, Rio Anori (Antioquia) 2-5 May 1970, C.H.P. 7 ^o., same data but 20-22 Sept. 1970, D.G.Y. 1 6, same data but 14 April 1971, C.H.P. Id 1 , Tres Esquinas (Caqueta) , light trap, 10 Nov. 1971, C.J.M. 52 do', 70 ^>§, Curiche (Choco), light h Malaise traps, April-Nov. 1967, D.G.Y. 2 oo., Alto Curiche (Choco), light trap, 2 July 1967, D.G.Y. 2 do", 2 ^o, Anchicaya Dam (Valle), light traps, 10-11 Aug. 1973, D.G.Y. & R.C.W. 1 o., 25 km E of Buenaventura (Valle), light trap, 12 Aug. 1973, D.G.Y. & R.C.W. Brazil, French Guiana, Panama, and Paraguay. Those specimens listed by Fairchild & Hertig (1953a). Trinidad. 50+ 66, 50+ oo, from various localities to be treated in a forthcoming paper.

PAGE 248

•236Discussion: According to locality, the Tiales of L. aragaoi vary in the shape of the parameres. Those seen by me or figured by "•(hers from Brazil (Costa Lima, 1932), Colombia (Rio Anori, near Leticia, and Anchicaya Dam), Panama (Fairchild & Hertig, 1953a), and Peru (Lucena & Almeida, 1964) have relatively slender parameres compared to those from French Guiana (Floch & Abonnenc, 1952), Trinidad, and Colombia (Choco Dept. and Tres Esquinas in Caqueta Dept.)Whether clinal or not, I believe that this variation is intraspecif ic, the other character states of both sexes being remarkably similar. Like Forattini (1973), I consider L. heskenrothi (Floch & Abonnenc, to be the female of L. aragaoi. Floch & Abonnenc' s subsequent figure (1945c) shows long sperm ducts with warty protuberances, as noted by Fairchild & Hertig (1953a) , but this probably represents an abnormal specimen, a poor slide mount or, less ]ikely, another species. 55. Lutzomyia bavrettoi bavvettoi New Status (Fig. 47) Phlebotorrrus bavrettoi Mangabeira, 1942a: 148 (o* holotype, Belem, Aura, Para, Brazil). Barretto, 1947: 189 (refs.). Floch & Abonnenc, 1952: 38 (c? keyed), 144 (6* redescr . , figs.). Fairchild & Hertig, 1953a: 26 (refs.). Fairchild & Hertig, 1959: 121 (in part, Brazil & French Guiana). Vargas & Diaz-Najera, 1959: 146 (cf. to humboldti) . Sherlock & Carnheiro, 1964: 206, 208 (internal morphol., (7, fig.). Rosabal, 1966: 1-2 (in part, distrib.). Ortiz, 1972a: 23-24 (listed). Phtebotomus sp. de Haripa Floch & Abonnenc, 1946: 3 (o, French Guiana). Floch & Abonnenc, 1952: 46, 191 (o keyed, redescr., figs.).

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-237Lutzomyia barrettoi: Barretto, 1962: 99 (listed). Martins et al., 1976c-: 483 (Peru). Psychodopygus barrettoi: Forattini, 1971a: 105 (listed). Forattini, 1973: 416 et seq. (in part). Distribution: Colombia (Caqueta), Peru, Brazil, French Guiana, Trinidad. Material examined: Colombia. 2 , Tres Esquinas (Caqueta), light trap. 10-11 Nov. 1971, C.J.M. Brazil. 1 d", , Mangabeira, no. 1075, no other data. Trinidad. 1 o, Macqueripe U.S. Naval Station, light trap, 19 Nov. 1956. T.H.G. Aitken. 1 6", Tucker Valley U.S. Naval Station, light trap, 14 Nov. 1957, T.H.G. Aitken. 1 tf, San Jose Point U.S. Naval Station, light trap, 18 Nov. 1957, T.H.G. Aitken. Discussion: Specimens of L. barrettoi from east of the Andes in Colombia (Caqueta), Brazil, French Guiana, Trinidad, and probably Peru agree closely with the descriptions of Mangabeira (1942a) and Floch & Abonnenc (1946, 1952, as sp. de Maripa) . I am convinced that Lutzomyia sp. de Maripa is the female of L. b. barrettoi based upon morphology and collecting data in Trinidad and Caqueta Dept., Colombia. The spermathecae are large and oval, their ducts subequal in width throughout (Fig. 47B) . Other features of diagnostic importance are discussed under L. barrettoi majuscula n. ssp. 56. Lutzomyia barrettoi majuscula. n. ssp. (Fig. 46, 47) Phlebotorrus barrettoi: Fairchild 6 Hertig, 1953a: 21 et seq. (d\ p, descr., keyed, figs.). Rodriguez, 1956: 79 (Ecuador). Fairchild & Hertig, 1959: 121 et seq. (in part, Central American records). Fairchild

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-238& Hertig, 1961b: 250 (cf. to botella) . Johnson & Hertig, 1961: 765 (rearing data). Rosahal & Trejos, 1964: 167 (El Salvador). Rosabal, 1966: 1--2 (Costa Rica). Hanson, 1968: 51 (first instar larva, descr., tig.). Fklebotomus sp. no. 6 Rosabal, 1954: 6, 46 (Costa Rica). Lutzomyia barrettoi: Barreto, 1969: 463 (Valle, Colombia). Chaniotis et si., 1971a: 344 et seq. (collecting data, Panama). Christensen, 1972a: 88 (listed). Christensen et al., 1972: 57 (in animal burrow, Panama). Chaniotis et al., 1972: 95 (resting sites). Osorno et al., 3 972a: 38 (mention). Psyehodopygus barrettoi Forattini, 1973: 416 et seq. (in part). Distribution El Salvador, Nicaragua, Costa Rica, Panama, Colombia (Antioquia, Choco, Valle) , Ecuador. Material examined: Panama. 1 6* (holotype no. 6320), Cruces Trail, Madden Forest Reserve, Canal Zone, animal burrow, 6 Jan. 1957, M. Hertig. 1 c (allotype no. 4732), Camp Pina, Canal Zone, light trap, 24 May 1954, F. Blanton. 1 6" (paratype no. 4200), San Lorenzo, Canal Zone, light trap, 15 Aug. 195.2, F. Blanton. 1 o (paratype no. 3914), Mojinga Swamp, Canal Zone, light trap, 31 Jan. 1952, F. Blanton. 1 o (paratype no. 4735), Canal Zone, light trap, 29 June 1954, G. Field. 3 $S , 4 jo. (paratypes nos. 270-276), Cerro Azul (Panama), Burrow, 24 March 1968, D.G.Y. Colombia. 1 o (paratype no. 277), Rio Anori (Antioquia), light trap, 22 Sept. 1970, D.G.Y. 1 cf (paratype no. 278) Alto Curiche (Choco), light trap, 7 July 1967, D.G.Y. 1 o (paratype no. 279), Curiche (Choco), Malaise trap, 29 June 1967, D.G.Y. Costa Rica. 1 6* (paratype no. 3447), Suerre de Guapiles, tree buttress, 29 April 1951, R. Rosabal. Ecuador. 1 6* (paratype no. 280), 17 km E of Santo Domingo de Los Colorados

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-239(Pichincha), light trap, 27 May 1976, D.G.Y., T. Rogers, & G. Fairchild. 1 o (paratype no. 281), same data but May 1976. Nicaragua. 1 cf (paratype no. 4457), Guapilonar, Carrazo, light trap, 21 Oct. 1953, A. Adames. Other material examined: Colombia. 1 d", 5 09, Curiche (Choco) , light & Malaise traps, 5 July-28 Aug. 1967, D.G.Y. 1 o., 25 km E of Buenaventura (Valle), light trap, 12 Aug. 1973, D.G.Y. & R.C.W. Discussion: Male: As described and illustrated by Fairchild & Hertig (1953a, as L. baivettoi) with the following additions based upon the holotype of L. b. majuscula n. ssp. Wing length 2.47; width 0.78. Head height 0.38; width 0.44. Eyes separated by 0.10 or by distance = to 4.6 facet diameters. Flagellomere I (0.30 long); combined length of f lagellomeres II + III = 0.38. Length of palpal segments: 1 (0.03), 2 (0.08), 3 (0.12), 4 (0.07), 5 (0.16). Labrum 0.20 long. Cibarial arch nearly complete; pigment patch distinct, subtriangular. Pharynx 0.20 long. Pleura with 13 upper and 4-5 lower episternal setae. Length of wing vein sections: Alpha (0.80), beta (0.22), delta (0.42), gamma (0.19). Length of femora, tibiae, and basitarsi: Foreleg, 1.06, 1.56, 0.95; midleg, 0.98, 1.66, 0.98. Hindleg missing. Genitalia: Length of style 0.28; coxite 0.49; lateral lobe 0.60; genital pump 0.21, filaments 0.78 (the latter 3.7X length of pump) . Female: As described and illustrated by Fairchild & Hertig (1953) with the following additions based upon the allotype of L. b. majuscula n. ssp. Wing length 2.60; width 0.88. Head height 0.42; width 0.46. Eyes separated by 0.45 or by distance = to 5 facet diameters. Flagellomere I 90.29 long); combined length of f lagellomeres II + III = 0.37. Length of palpal segments: 1 (0.03), 2 (0.09), 3 (0.14), 4 (0.07), 5

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-240(0.17). Cibarium with 15 horizontal teeth, about 20 vertical teeth in 2-3 transverse rows, the 7 closest to horizontal teeth markedly enlarged; cibarial arch nearly complete, pigment patch prominent as shown by Fairchild & Hertig (1953a, fig. 21). Labrum 0.19 long. Pharynx 0.20 long. Pleura with 15-16 upper and 4-5 lower episternal setae. Length of wing vein sections: Alpha (0.88), beta (0.22), delta (0.51), gamma (0.19). Legs missing. Spermathecae and ducts as figured. Although larger than L. b. barvettoi , the female of L. b. ma jus aula has smaller spermathecae and the ducts are noticeably swollen where they enter the spermathecae (Fig. 47A) . The tip of the genital fork stem, sha.llowly bifid in majusQula, is variable in b. bavvettoi (Fig. 47C-E). Without females or locality data, the males are difficult to distinguish but the parameres of majusaula are slightly broader with a better developed ventral "heel." The lateral lobes of this taxon range between 0.57-0.61 mm in length; those of b. bavrettoi are shorter, not exceeding 0.50 mm. The wing length of majuscula 8<3 ranges from 2.3 to 2.7 mm; go from 2.5 to 2.8 mm. The wing length of b. bavrettoi 6o ranges from 1.8 to 2.2; go from 2.1 to 2.4. The subspecific name refers to the larger size of the majuseula specimens. This subspecies like L. eavvevai thula n. ssp., occurs west of the Andes in Ecuador and Colombia, then northward into Central America to El Salvador. Sherlock & Carnheiro (1964) suggest that L. texana (Dampf) from Texas (U.S.A.), Mexico, and Honduras (unpub . data) and L. barvettoi may be conspeeific, basing their interpretation on the description of texana by Dampf (1938) . While examining numerous specimens of the former species (Young, 1972b), 1 noted that the males of the two species can not

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-241be separated by the key given by Fairchild & Hertig (1953a)., the styles of both species with the isolated spine set upon a well marked tubercle doner to the basal spines than to the terminal spine. They differ, however, in the relative lengths of the genital filaments (less than 2.6 r. length of the pump for texana.; 2.8 x or greater for the barrettoi subspecies) . The scattered hairs on the inner surface of the coxites are long in texana but rather short in barrettoi. The females are more easily distinguished, as was done by Fairchild & Hertig (op. cit., in key) and by the nearly spherical, smaller spermathecae of texana. (Fig. 471). 57. Lutzomyia earpenteri (Fig. 43) Phlebotomus earpenteri Fairchild & Hertig, 1953a: 28 (6* holotype, Chiva Chiva, Canal Zone; o, near Pacora, Panama Prov. , Panama). Fairchild & Hertig, 1959: 121-122 (Central American records). Fairchild & Hertig, 1961b: 2.50 (cf. to botella) . Johnson & Hertig, .1961: 765, 773 (rearing data). Biagi et al . , 1966: 141 et seq. (d", p, figs., Mexico). Hanson, 1968: 53 (first instar larva, descr., fig.). Ortiz, 1972a: 23-24 (listed) . Lutzomyia aarpenteri.: Barretto, 1962: 99 (listed). Williams, 1970: 332 et seq. (summary of collecting data, Belize). Tesh et al., 1971a: 153 (blood meal from bird). Chaniotis et al., 1971a: 344 et seq. (collecting data, Panama) . Christensen, 1972a: 88 (listed) . Chaniotis et al., 1972: 95 et seq. (resting sites). Shaw & Lainson, 1972: 709 (mention). Christensen et al . , 1972: 57 et seq. (collecting data, negative for flagellates, Panama). Christensen & Herrer, 1973: 579 et seq. (collecting data, negative for trypanosomatids) . Chaniotis & Correa,

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-2A21974: 115 (light trap data). Lewis, 1975a: 501 et seq. (mouthpart morphol.)Rutledge & Ellenwood, l?75a: 73 (breeding site, open forest floor, Panama). Rutledge et al . , 1975: 179 et seq. (light trap data). Williams, 1976a: 604 (in caves, Belize). Psyehodopygus carpentevi: Forattini, 1971a: 105 (listed). Forattini, 1973: 390 et seq. (gen. review, figs.). Distribution: Mexico, Belize, Costa. Rica, Panama, Colombia (Antioquia, Choco) . Material examined'. Colombia. 72 6*cf, 10 on, Rio Anori (Antioquia), light traps, May 1970, C.H.P. 12 cfc?, 25 o^, same data but Sept. 1970, D.C.Y. 29 eft?, 33 go, Curiche (Choco), light, Shannon, & Malaise traps, April-Dec. 1967, D.G.Y. 1 o, Teresdta (Choco), Malaise trap, 22 April 1967, D.G.Y. 1 o", same data but light trap, 29 Sept. 1967. 1 o, same data but 28 Oct. 1967. Panama. 1 6* (holotype no. 1345), Chiva Chiva, Canal Zone, light trap, Oct. 1948, S. Carpenter. 1 c(allotype no. 2301) near Pacora (Panama Prov.), burrow, 13 Aug. 1950, M. Hertig. 9 <$
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-243Se?. ies bras i Itensii 58. Lutzomyia runoida (Fig. 49) Phlebotomus runoides Fairchild & Hertig, 1953a: 30 (d 1 holotype, Almirante, Bocas del Toro Prov. , Panama; o, La Victoria, Cerro Jefe, Panama Prov., Panama). Fairchild & Hertig, 1959: 122 (Central American record;;)Vargas &. Diaz-Najera, 1959: 146 (cf. to humboldti) . Johnson &. Hertig, 1961: 765 et seq. (rearing data) . Mangabeira & Sherlock, 1962: 318 (cf. to bras Hi ens i s) . Thatcher & Hertig, 1966: 52 (in mammal burrows). Hanson, 1968: 71-71 (fourth instar larva, descr., figs.). Ortiz, 1972a: 23-24 (listed). Lutzontyia runoides: Barretto, 1962: 99 (listed). Martins et al., 1965: 3, 7, 19 (Rondonia, Brazil, cf. to abunaensis) . Barreto, 1969: 466 (Valle, Colombia). Chaniotis et al., 1971a: 344 et seq. (collecting data, Panama). Christensen, 1972a: 88 (listed). Osorno et al., 1972a: 38 (Caqueta, Colombia). Chaniotis et al., 1972: 95 (resting sites). Christensen & Herrer. 1973: 579 (collecting data, Panama). Psyahodopygus runoides: Forattini, 1971a: 105 (listed). Forattini, 1973: 469 (as synonym of in flat a) . Distribution: Costa Rica, Panama, Colorrbia (Amazonas, Antioquia, Caqueta, Valle), Brazil. Material examined: Colombia. 2 po, 17 km W of Leticia (Amazonas), flight trap, 26 July 1973, D.G.Y. & R.C.W. 16 d?6*, Rio Anori (Antioquia), light traps, May 1970, C.H. P. 7 oV, 2 05, same data but Sept. 1970, D.G.Y. Panama. 1 6" (holotype no. 3593), type locality, tree buttress, 25-27 July 1951, A. Quinones. 20 6V, 13 go from various localities in

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-244the Canal Zone (Mojinga Swamp, Barro Colorado Island, Juan Mina) , and in the Republic of Panama (Almirante, Cerro Jefe, Cerro Carapana) . Discussion: This species and L. inflata (Floch & Abonnenc) from French Guiana are very close and may indeed by conspecific as Forattini (197J) proposes. Without material from French Guiana, however, and in view of the distinguishing characteristics outlined by Fairchild & Hertij (1953a), I prefer to treat them as distinct taxa, realizing that the status of L. vunoides may have to be reconsidered in the future.

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Figure 45 Lutzorryia aragaoi — A. Male head, B. Male flagellomere II, C. Female head, D. Female rlagellomere II, E. Female cibarium and pharynx, F. Spermathecae, G. Male genitalia, H. Body of spermatheca, I. Female ving, J. Mile wing, K. Female cibarium. Male: Curiche, Choco Uept., Colombia Female: Same locality as male

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) E j

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Figure 46 Lutzomyia barrettoi ma jus aula — A. Male head, B. Male flagellomere II. C. Female head, D. Female flagellomere II, E. Female cibarium and pharynx, F. Male genitalia, G. Genital pump and filaments, same scale as Fig. 47F, H. Spermathecae, I. Female wing, J. Male wing, K. Female cibarium. Male: Cerro Azu] , Panama Prov. , Panama Female: Same locality as male

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>48i l§ "V =^-" ft \ i

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Figure 4 7 Lutzomyia, barpettoi majusaula — A. Sparmatheca. Female: Cerro kzu.1, Panama Prov. , Panama Lutzomyia bajrrettoi barvettoi — B. Spermathecae, same scale as Fig. 46A, C. Tip of genital fork stem, D. Tip of genital fork stem, E. Tip of genital fork stem of female from Caqueta, Colombia, F. Female cibarium, G. Female f lagellomere II, H. Male genitalia. Kale: Trinidad Female: Trinidad (except Fig. 46E) Lutzomyia texana — I. Spermatheca, same scale as Fig. 46A. Female: Sinton, Texas, U.S.A.

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-250-

PAGE 263

Figure 48 Lutzonyia oavpentevi — A. Male head, B. Male flagellomere II, C. Female head, D. Female flagellomere II, E. Female cibarium and pharynx, F. Body of spermatheca, G. Genital pump and filaments, same scale as Fig. 48H, H. Mai e genitalia, I. Tips of genital fiimanets, J. Spermathecae, K. Female wing, L. Male wing, M. Female cibarium. Male: Curiche, Choco Dept. , Colombia Female: Same locality as male

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Figure 49 Lutzomyia vunoides — A. Male head, B. Male flagellomere II, C. Female head, D. Female flagellomere II, E. Female cibarium and Pharynx, F. Genital pump and filaments, G. Male genitalia, H. Spermathecae, I. Female wing, J. Male wing, K. Female cibarium. Male: Rio Anori, Antioquia Dept., Colombia Female: Same locality as male

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-254-

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-255Drei s bac hi. Gro up Lewis et a 1. , 1978 Ry virtue of the annulated spermathecae and cibarial armature, the females of this group resemble those in the subgenera Nyssomyia Barretto and Tviakopkovomyia Barretto but the ascoids have long posterior spurs and the males have 2 or more strong distal setae on each coxite. Four species comprise the dveisbaahi group, two of which occur allopatrically in Colombia. The females ordinarily do not attack man and are very similar, if not indistinguishable, in structural details. Without associated males or knowledge of locality, I am unable to identify them beyond the group level. Key to Species Males 1. Paramere simple but with modified dorsal seta at apex and tuft of apically recurved setae near middle. Coxite with 2 large modified setae at ventral median surface aalydifeva (Fig. 50) Paramere bifurcate; it & coxite without modified setae dveisbaohi (Fig. 51) 59 . Lutzomyia aalydifeva (Fig". 50) Phlebotomus aclydifevus Fairchild & Hertig, 1952: 511 (c? holotype, Mcjinga Swamp, near Catun, Canal Zone; (j>, Juan Mina, Canal Zone) . Fairchild & Hertig, 1959: 121 (Mexican & Central American records). Johnson & Hertig, 1961: 765, 773 (rearing data). Hanson, 1968: 44 (larva, pupa. descr. , figs.) .

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-256Lutzomyia aclydifera: Theodor, 1965: 193, 194 (6*, n figs., listed). Barreto, 1969: 46?. (Valle, Colombia). Sherlock & Guitton, 1970: 140 (classif.). Martins et al . , 1970a: 550 (cf. to 1 ienna.nl en bi) . Chaniotis et al . , 1971a: 344 et seq. (pop. dynamics, Panama). Young, 1972a: 312 (mention). Martins & Morales, 1972: 365 (listed). Osorno et al . , 1972a: 61 (mention). Chaniotis et al., 1972: 95 (resting sites, mostly burrows) Christcnsen, 1972a: 88 (listed). Christensen et al . , 1972: 57 et. seq. (collecting data). Chaniotis & Correa, 1974: 115 (light trap data). Rutledge et al., 1975: 181 (light trap data). Rutledge & Ellenwood, 1975a: 72 (breeding on open forest floor). Lewis , 1975a: 502 et seq. (rnouthpaxt morphol.). Martins et al . , 1976a: 493 (cf. to ruparupa) . Herrer & Christensen, 1976a: 62 (collecting data). Psychodopygus aclydifevus: Forattini, 1971a: 105 (listed), Forattini, 1973: 189 et seq. (gen. review, figs.). Gomes, 1975: 9 (listed). Distribution: Mexico, Honduras, Nicaragua., Costa. Rica, Panama, Colombia (Antioquia, Choco, Valle), Ecuador. Material examined: Colombia. 58 dcT 1 , 16 oo, Rio Anori (Antioquia), light traps, May 1970, C.H.P. 6 6*o*, 6 (ft, same data but Sept. 1970, D.G.Y. 190 6*6*, 296 o^, Curiche (Choco), light, Shannon & Malaise traps, tree trunks, April-Dec. 1967, D.G.Y. 26 do*, 28 oo, Alto Curiche (Choco), light & Shannon traps, Aug. -Nov. 1967, D.G.Y. 28 do 1 , 59 o^, Teresita (Choco), light & Shannon traps, June-Nov. 1967, D.G.Y. 1 o, Anchicaya Dam (Valle), light trap, 11 Aug. 1973, D.G.Y. & R.C.W. Ecuador. 15 dcf, 10 op , 17 km E of Santa Domingo de los Colorados (Pichincha) , light & Malaise traps, May 1976, D.G.Y., T. Rogers, & G. Fairchild. Panama. d'holotype, o allotype and 50+ specimens from various localities, most of which were listed by Fairchild & Hertig (1952) .

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Discussion: At Curiche, L. aolydifeva was the most abundant phlebototoine species taken in light traps from April to Dec, 1967 (39 trap nights). Of a total catch of 1167 specimens (23 species), accounted for 30% or 350 specimens (155 6*o", 195 oo) . The feeding habits of this species remain unknown. 60 . Lutzomyia dreisbachi (Fig. 51) Phlebotomus dreisbachi Causey & Damascene, 1945: 649 (6* holotype, Coarij Rio Solimoes, Amazonas, Brazil). Barretto, 1947: 197 (refs.). Damasceno et al., 1949: 824 (Brazilian records). Barretto, 1950a: 106 (c? keyed). Barretto, 1951: 216 (distrib.). Floch h Abonnenc, 1952: 20 et seq. (cf, o, keyed, redescr., figs., French Guiana). Vargas h DiazNajera, 1953a: 49 (listed). Lutzomyia dreisbachi: Theodor, 1965: 185 (listed). Martins et al., 1965: 2 (Rondonia, Brazil). Sherlock & Guitton, 1970: 140 (classif.) . Martins et al., 1970: 550 (cf. to hermanlenti) . Young, 1972a: 312 (mention). Martins & Morales, 1972: 367 (listed). Martins et al., 1976a: 493 (cf. to ruparupa) . Martins et al . , 1976b: 498 (mention). Psychodopygus dreisbachi: Forattini, 1971a: 105 (listed) . Forattini, 1973, 390 et seq. (gen. review, figs.). Gomes, 1975: 9 (listed). Distribution: Brazil, French Guiana, Colonbia (Amazonas) . Material examined: Colombia. 1 d*, 17 km W of Leticia (Amazonas), light trap, 26 July 1973, D.G.Y. & R.C.W. Discussion: A little known Amazonian species, L. dreisbachi was discovered first in an armadillo burrow in Amazonas, Brazil, where over 2,000 males were captured (Causey and Damasceno, 1945). Future collections

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-258should reveal its presence in other localities within the Amazon Basin. I have not examined females of this species but a description and good figures are provided by Floch and Abonnenc (1952) .

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Figure 50 Lutzomyia aclydifevo. — A. Male head, B. Male flagellomere II, C. Female head, D. Female flagellomere II, E. Female cibarium and pharynx, F. Genital pump and filaments, G. Spermathecae, H. Male genitalia, I. Body of spermatheca, J. Female wing, K. Male wing, L. Female cibarium. Male: Curiehe, Choco Dept., Colombia Female: Same locality as male

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-260-

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-261Subg enus Tr-ichopk o vonvjio. Ba rretto, 196 2 This is a distinct group as treated by Barretto (1962) and Sherlock and Guitton (1970), the species of which occur in or near the Amazon Basin except for L. reburra. The 20+ Triahophoromjia species, especially females, resemble those in the subgenus Nyssomyia Barretto, 1962, as noted by several authors. Forattini (197] a, 1973) in fact enlarges TvLchophor-omyia to accomodate these and other species, some of which such as L. har>trnanni (Fairchild and Hertig), L. sangv.-inaria (Fairchild and Hertig) , L. fluvCatilis (Floch and Abonnenc) , andL. nordest-ina (Mang.) do not appear to be closely related to the others based on a combination of characters. Sherlock and Guitton (19 70) provide a useful key to the Trichophoromyia males. The few described females, being similar in structure, are difficult to identify in the absence of males. At present, 6 Tvichophovonvjia species are known to occur in Colombia but others certainly exist in the Amazonian region. The species are attracted to lights but their preferred resting sites and hosts remain virtually unknown. None of the species, unlike those of Nyssomyia, is known to be anthropophilic. Key to Species Males 1. Genital filaments less than 4 times length of pump 2 Genital filaments greater than 4 times length of pump

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•2622. Paramere broader, less clubbed, most of dorsal setae sinuous. Eyes large, separated by less than 4 facet diameters . . vphurra (Fig. 54) Paremere otherwise, dorsal setae straight or nearly so. Eyes smaller. separated by more than 6 facet diameters. . . . vbiquitdlis (Fig. 51) 3. Coxite with only a distal group of 4-6 straight setae. Paramere as shown saltuosa (Fig. 53) Coxite with basal and/or median tufts, the total number of setae greater than 10. Paramere otherwise 4 4. Coxite with 3 groups of setae, a basal patch of about 7 short setae, a row of 4 longer hairs on median ventral surface & a distal patch of about 8 slender setae. Paramere as shown. . . eelluZana (Fig. 52) Coxite wir.h 1 or 2 groups of setae. Paramere otherwise 5 5 Coxite tuft circular and compact, the tips of nearly all setae downwardly curved howardi (Fig. 53) Coxite tuft otherwise, the setae loosely arranged, the tips of most not downwardly curved avJ'O.ensis (Fig. 51) 61. Lutzomyia (T.) aw?o.ensis (Fig. 51) Phlehotomu.s auvaensis Mangabeira, 1942a: 161 (d 1 holotype, Aura, Belem, Para, Brazil). Barretto & Coutinho, 1943: 188 (mention). Damasceno et al . , 1949: 819 (Brazilian records). Barretto, 1950a: 108 (o keyed). Vargas & Diaz-Najera, 1951a: 22 (mention). Barretto, 1951: 212 (Brazilian records). Floch & Abonnenc, 1952: 36 (c? keyed) . Llanos. 1964: 373-375 (V auvaensis, o^redescr., figs., Peru).

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-263Lutzomyia auraensis: Barrettc, 1962: 97 (listed). Martins et al., 1965: 2 (Rondonia, Brazil). Theodor, 1965: 185 (listed, 6*, o. figs.). Sherlock & Guitton, 1970: 143 et seq. (
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-26^ 62. Lutzomyia (T.) oellulana n. pp. (Fig. 52) Male: Wing length 1.91; width 0.52. Whole insect dusky, mesonotura darker than pleura. Head height 0.34; width 0.33. Eyes separated by 0.11 or by distance = to 6.6 facet diameters. Flagellomere I (0.20 long), combined length of flagellomeres I and II = 0.24; ascoids with short posterior spurs, the distal tips of those on flagellomere II reaching beyond end of f 3 agellomere, on flagellomeres 2-8, remainder missing. Length of palpa3 segments: 1 (0.03), 2 (0.08), 3 (0.13), 4 (0.05), 5 missing; palpal sensilla at end of segment 2 and on distal two-thirds of the third. Labrum 0.20 long. Cibarium with vestiges of teeth, mostly dot-like but 3 slender ones present; cibarial arch and pigment patch invisible. Pharynx 0.14 long. Pleura with 11-12 upper and 4 lower episternal setae. Lengtli of wing vein sections: Alpha (0.51), beta (0.27), delta (0.31), gamma (0.20). Legs missing. Abdominal stermites 2-6 entire, lacking clear or open areas. Genitalia: Style (0.21 long) with 4 major spines inserted at different levels, no subterminal seta. Coxite (0.35 long x 0.15 wide), with 2 median groups of long setae and a group of 7 very small setae at base as shown. Paramere sub triangular apically. Aedeagus stout. Lateral lobe 0.44 long. Genital pump (0.18 long), each filament ca. 0.92 long or 5.1 x length of pump, filament tips expanded. Cerci as shown. Female: Wing length 2.45; width 0.69. Coloration as in d*. Head Height 0.40; width 0.37. Eyes separated by 0.14 or by distance = to 7.9 facet diameters. Flagellomere I (0.20 long), combined length of flagellomeres II and III = 0.27; ascoids as in (?, on all but last flagellomere. Length of palpal segments: 1 (0.04), 2 (0.12), 3 (0.15), 4 (0.05),

PAGE 277

5 (0.11); palpal sensilla as in 6*. Labrum 0.29 Jong. Cibarium with 18-20 horizontal teeth, those at extreme sides difficult to observe, with about 30 vertical teeth arranged as shown; clbarial arch complete though hardly visible in middle; pigment patch well infuscated, subtriangular. Pleura with 26 upper and 5 lower episternal setae. Length of wing vein sections: Alpha (0.70), beta (0.29), delta (0.46), gamma (0.25). Length of femora, tibiae, and basitarsi: Foreleg, 0.86, 1.12, 0.74; midleg, 0,84, 1.37, 0.83; hin.dleg, 0.93, 1.62, 0.90. Abdominal sternites not completely visible but apparently lacking clear or open areas. Tergite 8 with 10 setae en each side. Spermathecae as shown, cylindrical with complete annulations, the terminal one button-like, larger than others; sperm ducts as shown. Cerci rather broad and short. Distribution: Colombia (Caqueta) . Material exarrrined: Colombia. 1 6* (holotype no. 612), Tres Esquinas (Caqueta), light trap, 10 Nov. 1971, C.J.M. 1 g (allotype no. 613), same data as male. Discussion: The male of L. cellulana differs from the other Trichophopomyia males in several characteristics including the shape of the parameres, the presence of a cluster of 7 or so short setae at the base of the coxite, and the number and nature of coxite setae in the other two groups. There is a strong possibility that the female described above is not the female of cellulana. Further collections and/or rearing studies will be necessary to confirm the association. I am describing the specimen, however, because it differs from other known T^'ichophororm/ia females in the spermathecae and because it and the cellulana male were captured together, L. auvaensis being the only other Tr-ichophopomyia species taken in the light trap at Tres Esquinas.

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-266The specific name refers to "hermit" or "recluse" owing to the possibility that the single known male may not be conspecific with the female . 63. Lutzomyia (T.) hoioardi n. sp. (Fig. 53) Male: Wing length 2.03; width 0.61. Whole insect dusky, mesonotum darker than pleura. Head height 0.36; width 0.35. Eyes separated by 0.13 or by distance = to 7 facet diameters. Flagellomere I (0.22 long), combined length of f lagellomeres II and III = 0.26; ascoids with short posterior spurs, the tips of ascoids on flagellomere II reaching beyond distal end, on all f lagellomeres except last 2. Length of palpal segments: 1 (0.04), 2 (0.09), 3 (0.14), 4 (0.06), 5 (0.15); palpal sensilla at end of segment 2 and on distal two-thirds of the third. Labrum 0.21 long. Cibarium with ca. 15 dot-like vestiges of teeth; chitinous arch and pigment patch invisible. Pharynx (ca. 0.15 long but awkwardly turned due to mounting) . Pleura with 13-15 upper and 5 lower episternal setae. Length of wing vein sections: Alpha (0.54), beta (0.27), gamma (0.21), delta (0.36). Length of femur, tibia, and basitarsus of hind leg, 0.91, 1.52, and 0.86 respectively; other legs missing. Abdominal ste^nite 2 entire, without clear or open areas. Genitalia: Style (0.21 long), with 4 major spines at different levels, no subterminal seta. Coxite (0.41 long x 0.13 wide) with a dense median tuft of 30+ setae, most downwardly curved apically, 7 or so straight setae below those of the tuft. Paramere as shown, with a setose lobe on dorsal surface. Aedeagus stout with pointed tips. Lateral lobe (0.44 long). Genital pump (0.16 long), each filament ca. 0.93 long or 5.8 x length of pump;

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-267filament tip pointed, slightly expanded subapically. Cerci as shown . Distribution: Colombia (Amazonas) . Material examined: Colombia. 1 6* (holotype no. 614), 17 km W of Leticia (Amazonas), light trap, 26 July 1973, D.G.Y. & R.C.W. Discussion: The male of howardi differs from the other Trickophororrnjia males in the structure of the genitalia, especially the shape of the parame.res and the nature and position of the coxite tuft. The parameres of L. brachipy ga (Mang.) and L. viannamartinsi Sherlock and Guitton are somev/hat similar but they are more upturned apically and the setation is different. The coxites of these two Brazilian species are notably broader and there are 2 or more very strong setae in the coxite tuft. The female of howardi, presently unknown, probably has long sperm duci.s which correspond to the long ducts of the male. This species is named after ray father in appreciation for his constant encouragement and assistance throughout this study. 64. Lutzomyia (T.) reburra (Fig. 54) Phlebotomus reburms Fairchild & Hertig, 1961b: 246 (d* holotype, La Zumbadora, Cerro Azul, Panama Prov. , Panama; o, Rio Mandinga, Colon Prov. , Panama) . Lutzonvjia reburra: Barretto, 1962: 97 (listed). Lewis, 1967b: 136 (mention). Barreto, 1969: 466 (Valle, Colombia). Sherlock & Guitton, 1970: 141 et seq. (tf keyed, fig.). Christensen & Fairchild, 1971: 301 (mention). Christensen, 1972a: 88 (listed). Osorno et al . , 1972a: 37 (mention) .

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-268Psyahodopygns reburrus : Forattini, 1971a: 105 (listed). Forattini, 1973: 417 et seq. (gen. review, figs.). Gomes, 1975: 9 (listed). Distribution: Panama, Colombia (Choco, Valle), Ecuador. Material examined: Colombia. 178 33, 231 po, Curiche (Choco) , Malaise, Shannon, & light traps, April-Nov., 1967, D.G.Y. 18 33, 11 oo, Alto Curiche (Choco), Malaise, Shannon, & Light traps, JuneNo v. , 1967, D.G.Y. 38 S3, 55 oo, Anchicaya Dam (Valle), light & flight traps, tree trunks, 9-11 Aug. 1973, D.G.Y. & R.C.W. 57 6*d*, 126 oc. , 25 km E of Buenaventura (Valle), flight trap, 12 Aug. 1973, D.G.Y. & R.C.W. 129 33, 110 op, same data but light trap. Ecuador. 1 p, Puerto Quito (Pichincha) light trap, 30 Aug. 1976, J. Cohen. Panama. 1 3 (holotype no. 6427), type locality, Shannon trap, 12 Sept. 1957. 1 g (allotype no. 6414), Rio Mandinga (Colon), Shannon trap, 10 May 1957, P. Galindo & A. Adames. Discussion: The reburra female is recognized by the refringent excrescences on the individual sperm ducts, the elongate terminal knobs of the spermathecae, and the dark mesonotum and pleura. The female of L. (Nyssomyia) yuilli Young and Porter also has excrescences on the sperm ducts but it is a pale species occurring east and slightly north of the Andes unlike reburra which is restricted to Pacific coast forests in Ecuador and Colombia. It was first discovered in Panama but is much less common there. At Curiche, we collected 798 specimens of 24 phlebotomine species in a Malaise trap operated continuously at ground level from April to December, 1967. Lutzomyia reburra was the dominant species taken, accounting for 137 33, 145 op (31%) of the total catch. This species is readily attracted to light, one notable light trap catch (listed above) near Buenaventura yielding many specimens, some of which were observed

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-269in copula. We did not observe females attacking man at Curiche or else where nor did we discover the preferred resting sites. 65. Lutzomyia (T.) sdltuosa n. sp. (Fig. 53) Mate: Wing length 2.05; width 0.61. Whole insect dusky, mesonotum darker than pleura. Head height 0.38; width 0.34. Eyes separated by 0.12 or by distance = to 7 facet diameters. Flagellomere I (0.23 long), combined length of flagellomeres IT. and III = 0.25; ascoids with short posterior spurs present, on all flagellomeres except last 2; tips of ascoids on flagellomere II extending beyond the end of flagellomere. Length of palpal segments: 1 (0.04) , 2 (0.11), 3 (0.14), 4 (0.06), 5 missing; palpal sensilla at end of segment 2 and on distal two-thirds of segment 3. Labrum 0.25 long. Cibarium with six needle-like horizontal teeth; 3 in a row on one side, the others irregularly spaced, all more or Jess erect; vertical teeth (ca. 25) subequal in size, in 3 irregular rows; cibarial arch obvious at sides, invisible in middle; pigment patch subtriangular, conspicuous. Pharynx 0.19 long. Pleura with 9 upper and 5 lower episternal setae. Length of wing vein sections: Alpha (0.61), beta (0.30), delta (0.42), gamma (0.17). Legs missing. Abdominal sternites 2-6 entire, without clear or open areas. Genitalia: Style (0.25 long), with 4 major spines at different levels, no subterminal seta. Coxite (0.40 long x 0.12 wide) with 4-6 straight, rather strong setae dis tally. Paramere and aedeagus as shown. Lateral lobe 0.46 long. Genital pump (0.17 long), each filament ca. 0.90 long or 5.3 x length of pump; filament tips simple, very slightly enlarged apically. Cerci as figured.

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Distribution: Colombia (Amazonas) . Material examined: Colombia. 1 b" (holotype no. 615), 17 km W of Letici3 (Amazonas), light trap, 26 July 19 73, D.G.Y. & R.C.W. Discussion: A distinctive species of Iriohophopormjia, L. saltuosa if; easily recognized by the 4-6 straight setae on the distal part of the coxite and by the shape of the parameres. The specific name is based on the Latin r .ccrd, "saltucsus," meaning forested or woody, the unique ho lo type having been captured in a well developed forest about 2 km from the Amazon river. 66. Lutzomyia (T.) ubiquitalis (Fig. 51) Phlebotomus ubiquitalis Mangabeira, 1942a: 158 (6* holotype, Aura, Belem, Para, Brazil). Damasceno et al . , 1949: 833 (Brazilian records). Vargas & Piaz-Najera, 1951a: 22 (mention). Barretto, 1951: 2.25 (Brazilian records). Barretto, 1954: 124 (refs.). Foratt.ini, I960: 476 (Amapa, Brazil). Fairchild & Hertig, 1961b: 248 (cf. to veburrus) . Pifano et al., 1962: 384, 385 (6*, in key to phiebotomines of Venezuela). Phlebotomus aaudhensis Floch & Abonnenc, 1943: 22 (cf, Caux, French Guiana). Barretto, 1947: 192 (full refs.). Floch & Abonnenc, 1952: 104-107 (
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-271Lutzomyia ubiqui talis : Barretto, 1962: 97 (listed). Martins et al., 1963; 335 (Roraima, Brazil). Martins et al . , 1965: 4 (Rondonia, Brazil). Lewis, 1967b: 136 (o cf. to rostrans, refs.). Sherlock & Guitton, 1970: 139 et seq. (classif., 6* keyed, fig.). Lewis et al., 1970: 215 (parous rates). Osorno et al . , 1972a: 37 (Caqueta, Colombia). Lewis, 1975a: 500 et seq. (mouthpart morphol.). Lewis, 1975b: 366 (distrib. of abdominal setae, fig.). Ramirez et al . , 1976: 599 (Amazonas, Venezuela) . Psychodopygus ubiquitalis: Forattini, 1971a: 105 (listed). Forattini, 1973: 417 et seq. (gen. review, figs.). Distribution: Colombia (Caqueta), Brazil, French Guiana, Venezuela. Material examined.'. Colombia. 1 6* (1NPES no. 7099), Finca San Miguel. Milan, Municip. Florencia (Caqueta), tree hole, 25 Aug. 1971, E. Osorno, A. Morales, h J. Ochoa. Brazil. 3 6*cf, Serra do Navio (Amapa) , light trap, 21 Sept. 1972, D.G.Y. & H. Fraiha. 1 6*, 27 km SE of Maraba (Para), light trap, 28 Sept. 1972, D.G.Y. 1 6*, W of Altamira (Para), light trap, 1 Oct. 1972, D.G.Y. 3 oV, Belem (Para), light trap, 18 Oct. 1972, D.G.Y. Discussion: Lutzomyia ubiquitalis, the type species of the subgenus, is widespread in the Amazon Basin, the single Colombian specimen, a male, having been captured in Caqueta Department (Osorno et al., 1972a). The female, not seen by me, was described and figured by Floch and Abonnenc (1942b as Phlebotomus sp. and 1952 as cauchensis) .

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Figure 51 Lutzomyia dreisbaohi male — A. Genitalia. Male: Leticia, Araazonas Comisaria, Colombia Lutzomyia (T.) auraensis male — A. Genitalia. Male: Tres Esquinas, Caqueta Intendencia, Colombia Lutzomyia (T.) ubiquitalis male — A. Genitalia. Male: Belem, Para State, Brazil

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-273-

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Figure 52 Lutzomyia (T.) oellulana — A. Male head, B. Female head, same scale as Fig. 52A, C. Female flagellomere II, D. Female cibariuin, E. Male genitalia, F. Tip of genital filament, G. Genital pump and filaments, H. Spermathecae, I. Female wing, J. Male wing. Male: Tres Esquinas, Caqueta Intendencia, Colombia Female: Same data as male

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-275-

PAGE 288

Figure 53 Lutzomyia (T.) howcwdi, male — A. Genitalia, B. Wing, C. Head, D. Flagellomere II, E. Tip of genital filament, F. Genital pump and filaments, Male: Leticia, Amazonas Comisaria, Colombia Lutzomyia (T.) saltuosa male — G. Wing, H. Genitalia, I. Head, J. Flagellomere II, K. Tip of genital filament, L. Genital pump and filaments. Male: Leticia, Amazonas Comisaria, Colombia

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277-

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Figure 54 Lutzomyia (T.) veburva — A. Male head, B. Male flagellomere II, C. Female head, D. Female flagellomere II, E. Female cibarium and pharynx, F. Male genitalia, G. Genital pump and filaments, H. Spermathecae, I. Female wing, J. Male wing, K. Female cibarium. Male: Curiche, Choco Dept., Colombia Female: Same locality as male

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-2791 i rmrwTH $ ii

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-280Sub genus Ny ssomyia Barr et to, 1962 This subgenus is of special interest to public health workers as most of the included species are anthropophilic and are proven or probable vectors of cutaneous leishmaniasis (Lainson & Shaw, 19 73) and certain arboviruses (Tesh et al., 1974). The group was defined by several authors including Barretto (1962), Theodor (1965, as the intermedia group), and Forattini (1971a, 1973). The latter increased its coverage by including species in other subgenera and species groups recognized as distinct by Theodor (1965) and Lewis et al . (1978). The males of L. antvnesi and Lutzorrryia sp. no. 260.44 (Ward & Killick-Kendrick, 1974) have a definite group of setae on each coxite, lacking in the other Nyssomyia males. Forattini (1973) reviewed the literature pertaining to most of these species and thus I restrict the citations to recent studies or to important earlier works. To date, seven Nyssomyia species have been discovered in Colombia. Keys to Species Males 1. Coxite with a median group of 30+ setae antunesi (Fig. 55) Coxite without a group of setae 2 2. Style with all spines distal to middle of structure. Wing venation with delta shorter than half length of beta. Head and anterior part of mesonotum strongly pigmented 3

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-281Style with basal pair of spines proximal to or at middle of structure. Delta to or longer than beta. Head and mesonotum pigmented or not 4 3. Genital pump much longer than style and nearly as long as lateral lobe. Flagellcmere I (< 0.35 mm long), longer than II + III. . . . flaviscntellata (Fig. 55) Genital pump shorter or equal to length of style and shorter than half length of lateral lobe. Flagellomere I (> 0.40 mm long), longer than II + III olmeca bicolor' (Fig. 56) 4. Style with subterrninal and terminal spines nearly paired, separated by distance = to greatest width of terminal spine ylephiletor> (Fig. 59) Style with distal pair of spines more widely separated 5 5. Genital filament tips slightly inflated, simple. Coxite longer than lateral lobe trapidoi (Fig. 57) Genital filament tips modified. Coxite and lateral lobe subequal in length 6 6. Genital filament tips forked, claw-like. Style over half length of coxite or lateral lobe. Paramere somewhat acute at ventral basal margin. Mesonotum partly pigmented vjnbvatilts (Fig. 58) Genital filament tips otherwise. Style shorter than or = to half length of coxite or lateral lobe. Paramere rounded at lower basal margin. Mesonotum pale yuilli (Fig. 60)

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-282Female s 1. Common sperm duct as long as or longer than individual ducts. ... 2 Common sperm duct, if present, shorter than individual ducts. ... 4 2. Head height subequal to width. Flagellomere I shorter than labrum. Spermatheca with fewer than 8 distinct segments, about 2 x length of individual sperm ducts antunesi (Fig. 55) Head much longer than wide. Flagellomere I = to or longer than labrum. Spermatheca with 9-12 distinct segments, = to or longer than individual ducts 3 3. Flagellomere I = to or longer than II + III. Genital fork stem broad and blade-like. Cibarium with 8-10 (rarely 7) horizontal teeth . . olmeoa bioolor (Fig. 56) Flagellomere I shorter than II + III. Genital fork stem slender. Cibarium with 6-7 horizontal teeth flaviscutellata (Fig. 55) 4. Individual sperm ducts mostly striated and/or with refringent excrescences (papules) 5 Individual sperm ducts smooth-walled 6 5. Mesonotum pale. Individual sperm ducts" with numerous excrescences forming a feather-like pattern yuilti (Fig. 60) Mesonotum pigmented in part. Individual sperm ducts transversely striated but lacking excrescences imbratilis (Fig. 5S)

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-2336. Common sperm duct short but discernible. Spermathecae with segments increasing in size from base to apex ylephiletoi' (Fig. 59) Common sperm duct apparently absent. Spermathecae with segments subequal in size trapidoi (Fig. 57) 67. Lutzomyict (N.) aniunesi (Fig. 55) Phlebotomus centimes £ Coutinho, 1939: 181 (d 7 , Manacapuru, Amazonas, Brazil). Barretto, 1946b: 529-532 (refs., synonyms). Barretto, 194 7: 185 (refs., synonyms). Damasceno et al . , 1949: 818 (distrlb., Brazil). Barretto, 1950a: 108 (keyed). Barretto, 1951: 212 (distrib.). Vargas & Diaz-Najera, 1951a: 22 (mention). Floch & Abonnenc, 1952: 36 (
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-284( mention) . Oyorno et al . , 1972a: 33-34 (Colombian records). Shaw & Lainson, 1972: 710 et seq. (Para, Brazil, negative for flagellates). Shaw et al., 1972: 720 (biting man, Para, Brazil). Llanos, 1973: 33 (in part, o"only, redescr., figs.). Ward & Ready, 1975: 128 et seq. (egg, descr., fig.). Lewis 1975a: 500 et seq. (mouthpart morphol., cibarium fig.). Lewis, 1975b: 366 (hair sockets, figs.). Martins et al., 1976a: 487 (Peru). Martins et al., 1976b: 495 (Peru). Llanos et al., 1976: 480 (Peru). Ramirez et al., 1976: 599 (Venezuela). Ward, 1976: 227 et seq. (larva, descr., fig., keyed). Lainson et al . , 1977 (Mato Grosso, Brazil). Zimmerman et al . , 1977: 575 (egg cf. to that of ylephiletor) . Lutzotnyia maohicouensis: Barretto, 1962: 98 (listed). Theodor, 1965: 184 (= antunesi) . Fsychodopygus antunesi: Forattini, 1971a: 105 (listed). Forattini, 1973: 390 et seq. (gen. review, figs., synonyms). Distribution: Colombia (Caqueta, Guajira, Vichada) , Ver-u, Brazil, French Guiana, Venezuela, Trinidad. Material examined: Colombia. 1 d*, 15 oo, Tres Esquinas (Caqueta), light trap, 10 Nov. 1971, C.J.M. Brazil. 2 oo, Labrea (Amazonas) , light trap, 10 Oct. 1972, D.G.Y. 9 Sd, 4 oo, Serra do Navio (Amapa) , tree trunks, 21 Sept. 1972, D.G.Y. & H. Fraiha. 1 6*, 1 o, Rio Aripuana at Humboldt (Mato Grosso) , tree trunks, 18-20 Aug. 1974, D.G.Y. & J. Shaw. 2 pp, 27 km SE of Maraba (Para), light traps, 26-28 Sept. 1972, D.G.Y. & H. Fraiha. 1 o, Itiatuba (Para), light trap, 4 Oct. 1972, D.G.Y. & H. Fraiha. 3 6V, 7 oo, Belem (Para), tree trunks & flight trap, 29 July5 Aug. 1974, D.G.Y. Trinidad. 25+ do", 30+ oo, various localities, to be treated in a future article.

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-2861976b: 496 (Peru). Ward et al., 1977: 265-266 (bite transmission of Leishmania mexicana amazanensis to hams ters) . Psychodopygus flaviscutellatus • Forattini, 1971a: 105 (listed). Forattini, 1973: 170 et seq. (gen. review, refs., figs.). Gomes, 1975: 9 et seq. (review of disease relationships, ecology). Ward, 1977: 511 (adaptability to hosts and habits) . Distribution: Colombia (Amazonas), Ecuador, Peru, Brazil, Surinam, French Guiana, Venezuela, Trinidad.. Material examined: Colombia. 1
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-237Ecuador; thus i ending additional support to the contention that the taxa are distinct (Fairchild & Theodor, 1971; Lewis, 1976b). Lutzomyia flaviscutellata transmits Leishmanial mexioana amazonensis Lainson and Shaw to small forest mammals and occasionally to man (Ward, 1977), Parasites referable to this subspecies have been recovered from wild flies in Brazil (Lainson & Shaw.. 1968), Trinidad (Tikasingh, 1975), and Venezuela (Pifano et al., 1973). This sand fly is also a vector of Pacui virus in Brazil (Aitken et al., 1975), a disease of small mammals. Our knowledge of the bionomics of flaviscutellata has been gained largely through the efforts of Drs. R. Lainson, J. Shaw, and R. Ward, their papers being cited in the species bibliography. The Disney trap (Disney, 1966) or modifications of it have been effective in capturing large numbers of this and related species. The striking coloration of both sexes (Fairchild & Theodor, 1971) enables one to separate them from other Lutzomyia, species without the aid of a microscope in areas in which L. olmeca bicolov does not also occur. 69. Lutzomyia (11.) olmeca. bicolov (Fig. 56) Lutzomyia olmeca bicolov Fairchild & Theodor, 1971: 157 (cf holotype, Mojinga swamp, Canal Zone; o, Cruces Trail Canal Zone; also Colombian records). Christensen & Fairchild, 1971: 302 (Darien Prov. , Panama). Chaniotis et al . , 1971a: 344 et seq. (pop. dynamics, Panama). Chaniotis et al., 197.1b: 415 et seq. (man biting study). Osorno et al., 1972a: 34-35 (Colombian records). Christensen 1972a: 88 (listed). Christensen et al., 1972: 55 et seq. (collecting data, Panama). Chaniotis et al . , 1972: 95 (resting sites). Christensen & Herrer, 1973: 579 et seq.

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-288(collecting data). Chaniotis, 1974b: 501 (keyed). Lewis, 1975b: 363 et seq. (taxonomy). Rutledge & Ellenwood, 1975a: 73 (breeding site, open forest floor). Herrer & Christe.nsen, 1976a: 62 (collecting data, Panama), Ward, 1976: 238-239 (larva keyed, cf. to flaviscutellata) . Phlebotomus apiaalis (not apicdlis Floch & Abonnenc) : Rodriguez, 1950: 6 (Ecuador). Rodriguez, 1956: 80 (Ecuador). Fairchild &. Hertig, 1959: 121, 123 (in part, Panama only). Johnson & Hertig, 1961: 765 et seq. (rearing data). Thatcher, 1968a: 295 (hosts, Panama). Hanson, 1968: 46 et seq. (larva, pupa, descr., figs.). Lutsomyia flaviscutellata (not flaviscutellata Mang.): Barreto, 1969: 464 (Valle, Colombia). Tesh et al . , 1971a: 153 (blood meals). Osorno et al . , 1972a: 34 (mention). Psychodopygus olmecus: Forattini, 1973: 89 et seq. (gen. review, figs., disease relationships, refs.). Distribution: Costa Rica, Panama, Colombia (Amazonas, Antioquia, Boyaca, Caqueta, Choco, Guajira, Magdalena, Norte de Santander, Santander, To 1 ima , Va 1 1 e ) , Ecuador . Material examined: Colombia. 3 <$<$, 17 km W of Leticia (Amazonas), forest floor, 24 July 1973, D.G.Y. & R.C.W. 80 6V, 22 oo, Rio Anori (Antioquia), light & flight traps, biting man, May-June 1970, C.H.P. 17 oc?, 6 op, same data but light traps, tree buttress, Sept. 1970, D.G.Y. 1 6*, Tres Esquinas (Caqueta), light trap, 10 Nov. 1971, C.J.M. 23 S3, 102 op, Curiche (Choco), light, Shannon & Malaise traps, biting man, April-Dec. 1967, D.G.Y. 2 op, same locality, oil trap baited with Agouti paca, 25 April 1967, D.G.Y. 2 dcT, 16 oo, Alto Curiche (Choco), light & Malaise traps, biting man, July-Dec. 1967, D.G.Y. 21 S3, 42 oq, Teresita (Choco), light, Shannon, oil, & Malaise traps, sweeping, biting

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-289raan, March-Dec. 1967, D.G.Y. 1 a 1 , 4 no, Sautata (Choco) , Malaise trap, tree buttress, Oct. -Nov. 1967, D.G.Y. 8 6*6*, 2 oo, Anchicaya Dam (Valle) , light & flight traps, tree trunks, 9-11 Aug. 1973, D.G.Y. & R.C.W. Costa Rica. 2 dcf, 1 p, N of Marina, forest floor on dead leaves, 13 Feb. 1960, W. Hanson. 1 p, Turriabla, on horse, 2 May 1961, R. Rosabal. Ecuador. 2 $$, Rio Napo at I.imoncocha (Napo) , light trap, 23-24 May 1976, D.G.Y. & T. Rogers. Panama. 8 S3 (holotype and 7 paratypes), 25 pp paratypes, specific data given by Fairchild & Theodor (1971). Discussion: Females of olmeca bieolor, like those of L. flaviscutellata y are not strongly anthropophilic. At Curiche, we captured only 9 oo on human bait from April to Dec, 1967 (141 man hours). We obtained similar results at Alto Curiche where a total of 8 po were tak^n on man from 24 June to 18 Nov., 1967 (76 man hours). Only 1 female was collected on man at Teresita from 9 April to 16 Dec, 1967 (125 man hours). Christensen et al . (1972) studied the bionomics of this subspecies at Sasardi, San Bias Territory, Panama. It was the most common sand fly on the forest floor and in rodent-baited Disney traps. Leishmania mexicana or close ally was recovered from numerous mammals there (Herrer et al . , 1971) but was not detected in olmeca bieolor or other sand flies (Christensen et al . , 1972). 70. Lutzomyia (N.) trapidoi (Fig. 57) Phlebotomus trapidoi Fairchild &. Hertig, 1952: 524 (cf holotype, p. Almirante, Bocas del Toro Prov., Panama). Lutzomyia trapidoi: Barretto, 1962: 98 (listed). Tesh et al., 1971a: 152 et seq . (blood meals). Tesh et al., 1971b: 49.1 et seq .

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-290(virus transmission studies). Chaniotis et al., 1971a: 344 et seq. (pop. dynamics, Panama). Chaniotis et al . , 1971b: 415 et seq. (man biting study). Christensen et al., 1971: 118 (pyloric spines). Christensen & Fairchild, 1971: 301 (Darien Prov. , Panama). Osorno et al., 1972a: 35-36 (Colombian records). Tesh et al . , 1972: 88 (blood meals). Chaniotis et al., 1972: 91 et seq. (resting sites). Christensen, 1972a: 88 (listed). Christensen et al., 1972: 55 et seq. (collecting data, Panama). Christensen & Herrer, 1973: 579 et seq. (collecting data, positive for trypanosomatids) . Chaniotis & Correa, 1974: 115 (collecting data). Chaniotis, 1974a: 73 et seq. (sugar feeding). Chaniotis, 1974b: 501 (keyed). Tesh et al . , 1974: 258 et seq. (summary of virus studies). Chaniotis et al., 1974: 369 (dispersal study). Lewis et al., 1975a: 50.1 et seq. (mouthpart morphol.). Chaniotis, 1975: 183 et seq. (rearing data). Rutledge & Ellenwood, 1975a: 71 et seq. (breeding habitat, open forest floor, Panama), 1975b: 78 et seq. (ecology, breeding sites), 1976c: 83 (ecology, breeding sites). Christensen & Herrer, 1976: 299 et seq. (as host of Etidotrypanum) . Herrer & Christensen, 1976a: 57 (collecting data), 1976b: 62 et seq. (collecting data). Herrer et al . , 1976: 70 (mention). Miles et al . , 1976: 532 (mating aggregation). Zimmerman et al . , 1977: 574 et seq. (egg, descr., figs.). Psyahadopygus trapidoi: Forattini, 1971a: 105 (listed). Forattini, 1973: 123 et seq. (gen. review, refs., figs.). Gomes, 1975: 9 et seq. (gen. review, distrib., ecology, biting habits). DistributionHonduras, Costa Rioa, Panama, Colombia (Antioquia, Boyaca, Choco, Tolima, Valle) , Ecuador. Material examined: Colonbia. 35 c?r?, 4 99, Rio Anori (Antioquia), light traps, May 1970, C.H.P. 4 cjo, same data but Sept. 1970, D.C.Y.

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-29174 oc?, 120 oo, Curiche (Choco) , light, Shannon & Malaise traps, tree trunks, biting man, May-Dec. 1967, D.C.Y. 5 dtf, 63 ^>o, Alto Curiche (Choco), light Shannon traps, biting man, tree trunks, June-Dec. 1967, D.G.Y. 12 6*0*, 6.1 oo, Teresita (Choco), light, Malaise & Shannon traps, tree trunks, biting man, March-Dec. 1967, D.G.Y. 3 53, Rio Atrato at Sautata (Choco), Malaise trap, Jan. 1968, D.G.Y. 1 6*, 8 oo, Anchicaya Dam (Valle), light flight traps, tree trunks, 9-11 Aug. 1973, D.G.Y. & R.C.W. 52 &?, 39 oo, 25 km £ of Buenaventura (Valle), light & flight traps, tree trunks, 11-12 Aug. 1973, D.G.Y. h R.C.W. Costa Rioa. 4 oo , Moravia, Chirripo (Cartage), Shannon trap, 4 March 1966, R. Zeledon. Ecuador. 84 6*0*,, 33 00. 17 km E of Santo Domingo de los Colorados (Pichincha), light & flight traps, tree trunks, May 1976, D.G.Y., T. Rogers, & G. Fairchild . Honduras. 3 op, Tela, Lancetilla Valley, light trap, 24 Feb. 1954, W. Mis. Panama. 1 o* (holotype no. 3291), type locality, 19 June 1951, A. Quinones & R. Hartmann. 1 (j> (allotype no. 2474), Finca Nievecita, Almirante (Bocas del Toro) , tree buttress, 21 June 1950, R. Hartmann. 100+ 6*0", 100+ op, various localities, most of which were listed by Fairchild and Hertig (1952). Discussion: Specimens of L. trapidoi from Valle Dept., Colombia and Pichincha Prov., Ecuador are darker than those from other localities, the heads and mesouota of both sexes being moderately infuscated. Structurally, however, they agree with the holotype and allotype of trapidoi. 1 have cited only recent references to this species, Forattini (19/3) having revisited the important papers published prior to 1971. At Curiche from April to Dec, 1967, we secured 52 00 of trapidoi in routine human bait collections. This represents 1.4% of the total catch

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-292of J5 LutRomyia spp. (3763 go, 141 nan hours). At Teresita less than 1% of i'.he biting sand flies captured during an eight month period represented this species. We did not attempt, however, to collect sand flies in the forest canopy where trapidoi is likely to be more abundant (Fairchild & Hertig, 1952; Chaniotis et al., 1971b). During the day, trapidoi adults rest commonly on the forest floor, less frequently on living plants or on tree trunks (Chaniotis et al . , 1972). The larvae live on the open forest floor (Rutledge & Ellenwood, 1975a, b,c) where they tend to be surface feeders (Hanson, 1968). Flagellates, believed to be leishmanial promastigotes, were recovered from wild caught trapidoi females in Panama (Johnson et al. , 1952; McConnell, 1963) . This species may also play an important role in the epidemiology of certain arboviruses in that country, Tesh et al. (1974) having isolated a number of strains from males and females. There is presumptive evidence that vertical (transovarial) transmission of some viruses occurs in L. trapidoi and L. ylephiletor (Tesh & Chaniotis, 1975). 71. Lutzomyia (N.) umbratilis (Fig. 58) Lutzomyia wrbvatilis Ward & Fraiha, 1977 (o ho lo type, Monte Dourado, Rio Jari, Para, Brazil). Phlebotomus intermedia (not intermedia, Lutz & Neiva, 1912) : Floch & Abonnenc, 1941a: 2 (cf, descr. , figs.). Floch & Abonnenc, 1942a: 9 (o, descr. , figs .) . Phlebotomus anduzei (not andazei Rozeboom, 1942): Floch & Abonnenc, 1944c: 11 (o", 0, taxonomy, figs.). Floch & Abonnenc, 1945c: 4 (o ,

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-293keyed, figs.). Floch & Abonnenc, 1947a: 18 (d 1 , keyed). Fairchild & Hertig, 1952: 523 et seq. (cf. to ylephiletor & bra.pid.oi). Floch & Abonnenc, 1952: 37, 43 (6", o, keyed), 121 (cf, o, redescr., figs.). Fairchild & Hertig, 1959: 121 (distrib. , in part). Forattini, 1960: 476 (Amapa, Brazil). Wijers & Linger, 1966: 501 et seq. (Surinam, as probable vector of Leishmanial . Lutzomyia anduzei (not anduzei Rozeboom, 1942): Martins et al . , 1963: 334 (Roraima, Brazil). Martins et al., 1965 (Rondonia & other Brazilian localities). Almeida, 1970a: 2 et seq. (anomalous , redescr., figs., Peru). Lewis, 1975a: 501 et seq. (mouthpart morphol.). Llanos et al., 1975b: 669 (Peru). Martins et al., 1976a: 487 (Peru). Lainson et al., 1976: 171-172 (as vector of Leishmania in northern Brazil). Ward, 1977: 228 et seq. (larva descr., fig., keyed). Lainson et al., 1977 (Mato Grosso, Brazil). Lutzomyia sp. no. 260.31 Ward et al . , 1973: 178 (collecting data, Para, Brazil). Ward & Killick-Kendrick, 1974: 216, 219 (Para, Brazil). Psyohodopygus anduzei: Forattini, 1973: 388 et seq. (in part, gen. review, figs., refs.). Distribution: Colombia (Amazonas, ?Boyaca, ?Caqueta) , Peru,, Brazil, Surinam, French Guiana. Mati-rrial examined: Colombia. 6 ob*, 1 o, 17 km W of Leticia (Amazonas), tree trunks, 24-26 July 1973, D.G.Y. & R.C.W. Brazil. 15 do", 10 op, Serra do Navio (Amapa), tree trunks, 21 Sept. 1972, D.G.Y. & H. Fraiha. 1 cf, Rio Aripuana at Humboldt (Mato Grosso), tree trunk, 16 Aug. 1974, D.G.Y. & J. Shaw. 1 o, 27 km SE of Maraba (Para), light trap,

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-29426-28 Sept. 1972, D.G.Y. & H. Fraiha. 1 cf, near Altamira (Para), tree trunk, 1 Oct. 1972, D.G.Y. Discvjssioni Lutzomyia urribratilis , confused with L. anduzei (Rozebooui) for many years, differs from it by the darker mesonotum, by the striated sperm ducts (smooth walled in anduzei), by the relative lengths of palpal segments 2 and 5, and by other features discussed by Ward & Fraiha (1977). The male of anduzei has not been described. Fairchild & Hertig (1959) reported the presence of L. anduzei in Costa Rica and Panama. I have not examined Costa Rican specimens but 3 op from Almirante (Bocas del Toro Prov., 19-21 June 1950, R. Hartmann) seem to be conspecific with L. anduzei (Rozeboom) . The fifth palpal segment is shorter than the third, post of the insect is pale and the spermathecae and ducts are similar to those of L. anduzei Rozeboom except for some transverse striations on the individual ducts. There is no doubt that the specimens from near Leticia, Colombia, are conspecific with L. vrrbratilis as described by Floch & Abornienc (1944, 1952, as anduzei) and by Ward & Fraiha (1977). I am assuming, perhaps incorrectly, that the records of anduzei by Osorno et al. (1972a) from Boyaca and Caqueta Departments represent L. wrbratilis , not anduzei P.ozeboom. Lainson et al . (1976) found promastigotes in 4 out of 55 wrbratilis females examined in northern Brazil. These subsequently proved to belong in the genus Leishmania and may be identical to the type discussed by Wijers & Linger (1966) in Surinam (Lainson et al., op. cit.).

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29572. Lutzomyia (N.) ylephiletov (Fig. 59) Ffolebotomus ylephiletor Fairchild 6 Hertig, 1952: 518 (6* holotype, Almirante, Bocas del Toro Prov. , Panama; o, Cacique, Colon Prov. , Panama). Lewis & Garnham, 1959: 82 (Belize). Fairchild & Hertig, 1959: 122 (distrib.) • Forattini, 1960: 474 (not ylephiletor Fairchild & Hertig, 1952). Hanson, 1961: 320 et seq. (larvae on forest floor). Johnson & Hertig, 196i: 765 (rearing data). Hanson, 1968: 95 (larva, pupa, figs.) Lutzomyia ylephiletor (or ylephiletrix) : Barretto, 1962: 98 (listed). Barreto, 1969: 468 (Valle, Colombia). Williams, 1970: 332 et seq. (summary of collecting data, Belize). Tesh et al . , 1971a: 152 et seq. (blood meals). Chaniotis et al., 1971a: 344 et seq. (pop. dynamics, Panama). Chaniotis et al . , 1971b: 415 et seq. (man-biting study). Osorno et al., 1972a: 36 (listed). Tesh et al . , 1972: 88 et seq. (blood meals). Chaniotis et al., 1972: 91 et seq. (resting sites). Christensen, 1972a: 88 (listed). Christensen et al., 1972: 55 et seq. (collecting data). Zeledon & Alfaro, 1973: 416 (naturally infected with Leishmania, Costa Rica). Christensen & Herrer, 1973: 579 et seq. (collecting data, Panama). Chaniotis, 1974a: 73 et seq. (sugar feeding tests). Chaniotis & Correa, 1974: 115 (collecting data). Chaniotis, 1974b: 501 (keyed). Tesh et al., 1974: 258 et seq. (summary of virus studies). Chaniotis et al . , 19 74: 372 et seq. (dispersal study). Lewis, 1975a: 501 et seq. (mouthpart morphol.) Lewis, 1975b: 366 (hair sockets, figs.). Tesh & Chaniotis, 1975: 125 et seq. (virus transmission studies). Herrer & Christensen, 1976a: 62 (collecting data), 1976b: 57 (collecting data). Williams, 1976a: 604 (in caves, Belize). Zimmerman et al., 1977: 574 et seq. (egg, descr., figs.).

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-296Psychodopygus ylephiletor: Forattini, 1971a: 1G6 (listed). Forattini, 1973: 123 et seq. (gen. review, refs., figs.). Gomes, 1975: 9 et seq. (gen. review, distrib.). Distribution: Mexico, Guatemala.;, Belize, Honduras, Nicaragua, Costa Rica, Panama, Colombia (Choco, Valle) . Material examined'. Colombia. 17 oo, Curiche (Choco), Shannon & Malaise traps, biting man, tree trunks, 26 June-25 Nov. 1967, D.G.Y. Mexico. 1 o, Palenque (Chiapas), tree buttress, 1 April 1951, C. Fairchild & R. Hartmann. 2 oo, Teapa (Tabasco), light traps, 19 Aug. & 10 Sevt. 1953. Guatemala. 8 oo, Montana del Mico, Rio Blanco, human bait, no data (but probably March 1956), JBoshell. Honduras. 19 oo, Tela, Lancetilla Valley, human bait, March-April, 1954, W. Hils. Nicaragua. 1 6*, 1 o, Villa Soraoza, 15 June 1953. Costa Rica. 2 oo. Esquinas (Puntarenas) , tree buttresses, 28 Nov. 1949, H. Trapido. Panama. 1
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-297year period revealed that most ylephiZetor females bite in the forest canopy (Chaniotis et al . , 1971b). Adults tend to rest on tree trunks (Chaniotis et al., 1972). Leishmanial pi omastigotes were recovered from wild females in Panama (McConnell, 1963) and Costa Rica (Zeledon & Alfaro, 1973). Adults have been found naturally infected with certain arboviruses in Panama (Tesh et al., 1974), some of which may be transovarially transmitted. 73. Lutzomyia (N.) yuilli (Fig. 60) Lutzomyia yuilli Young & Porter, 1972: 524 (d* holotype, o, Rio Anori, Antioquia Dept., Colombia). Osorno et al. , 1972a: 79 (listed). Ward & Ready, 1975: 128 et seq. (egg descr., fig.). Lewis, 1975a: 501 (mouthpart morphol., fig., Colombia & Brazil). Llanos et al . , 1975b: 671 (listed, Peru). Martins et al . , 1976a: 488 (Peru). Lainson et al., 1977 (infected with epimastigote flagellates, Ma to Grosso, Brazil). Zimmerman et al . , 1977: 575 (egg cf. to that of ylephiletor) . Fraiha et al., 1978 (collecting data, Brazil). ?Lutzomia intermedia: Osorno et al., 1972a: 34 (not intermedia Lutz & Neiva, 1912) . Lutzomyia sp . , near intermedia Shaw et al., 1972: 720 (biting man, Para, Brazil) . Lutzomyia antunesi: Llanos, 1973: 33 (in part, o not antunesi Coutinho, figs., redescr.). Psyehodopygus yuilli: Gomes, 1975: 9 (listed). Distribution: Colombia (Antioquia, Caqueta, Met a) , Ecuador, Peru, Brazi I , Surinam .

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-298Material examined: Colombia. 1 S (holotype no. 163), type locality, light trap, 14 May 1970, C.H.P. 1 p (allotype no. 168), same data but 23 May 1970. 224 6*6*, 244 on (including paratypes) , same locality, May S Sept. 1970, C.K.P. & D.G.Y. 2 op, Tres Esquinas (Caqueta) , light trap, 10 Nov. 1971, C.J.M. Brazil. 5 op, Belem (Para), biting man, 28 Oct. 1970, J. Shaw et al . Ecuador. 3
PAGE 311

Figure 55 Lutzornyia (II.) antunesi — A. Male, genitalia, B. Female cibarium, C. 5p>ermathecae. Male: Trinidad Female: Trinidad jUtzomy-ia (N.) flaviseutellata — D. Spermathecae, £. Genital pump and genitalia, F. Female cibarium. Male: Leticia, Amazonas Comisaria, Colombia Female: Trinidad

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-300P=*> :0^ A W|\\-^%^ v -•>-5 A> ^iMS^f F \\\ j I

PAGE 313

Figure 56 Lutzonvjia (N.) olrneoa bicol-ov — A. Male head, B. Male flagellomere II, C. Female head, D. Female flagellomere II, E. Female cibarium, F. Male genitalia, G. Genital pump and filaments, H. Spermathecae, I. Female wing, J. Male wing, K. Female wing. Male: Curiehe, Choco Dept,, Colombia Female: Same locality as male

PAGE 314

-302-

PAGE 315

Figure 57 Lutzormjia (N.) tvapidoi — A. Male head, B. Male flagellomere II, C. Female head, D. Female flagellomere II, E. Female cibarium and pharynx, F. Male genitalia, G. Cenital pump and filaments, H. Spermathecae, I. Female wing, J. Male wing, K. Female cibarium. Male: Curiche, Choco Dept., Colombia Female: Same locality as male

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-304c^fr%

PAGE 317

Figure 58 Lutzomyia (a.) umbrat-ilis -A. Male head, B. Female head, C. Female cibarium, D. Female wing, E. Male genitalia including genital pump, F. Tip of genital filament, G. Spermathecae. Male: Leticia, Amazonas Comisaria, Colombia Female: Para State, Brazil

PAGE 318

306-

PAGE 319

Figure 59 Lutzomyia (N. .' yZephiletov — A. Male head, B. Male flagellomere II, C. Female head. D. Female flagellomere II, E. Female cibarium and pharynx, F. Male genitalia, G. Spermathecae of female from Costa Rica. H. Female wing, i. Male wing, J. Female cibarium. Male: Aimirante, Bocas del Toro Prov., Panama Female: Panama Canal Zone (except Fig. 59G)

PAGE 320

308|LjI D

PAGE 321

Figure 60 Lutzomyia (N.) yv.illi — A. Male head, B. Male flagellomere II, C. Female head, D. Female flagellomere II, E. Female cibarium and pharynx, F. Different views of paramere, G. Male genitalia, H. Tip of genital filament, I. Spermathecae, J. Female wing, K. Male wing, L. Female cibarium (all figures from Young & Porter, 1972). Male: Rio Anori, Antioquia Dept., Colombia Female: Same locality as male

PAGE 322

31!

PAGE 324

-312incorrectly described as L. davisi (not davisi Root, 1934), by Forattini (1960, 1973) . Only the nominate subspecies occurs in Colombia where it was mis identified as L. paraensis by Osorno et al . (19 72a). The other records of paraensis in Colorbia (Barreto, 1969) represent not that species but L. earrerai thula and/or L. ayvozai. Antunes (1937) reported the presence of L. sqiianrii'entr'is (Lutz and Neiva) in Meta Dept., Colombia but this record is based on females alone and I consider it to be doubtful, believing that the specimens represented another species in the series squamiventris , probably L. fairtigi Martins. Based on male genitalia, the Psychodopygus species are divided into 3 series following Theodor (1965) and Lewis et al. (1978). This agreement, admittedly artificial, is useful nevertheless for discussion and identification purposes. The short fifth palpal segment and imbricated spermathecae, both apomorphic features, are characteristic of all the Psychodopygus species. Studies on feeding habits indicate that most, if not all, females feed on mammals. The male genitalia may be simple. (series arthuri) or very complex (series sqi'jxmiventris and panamensis) . At present, 14 species and subspecies in the subgenus are known to occur in Colombia. Keys to Species Males Series squamiventris 1. Paramere as shown, with dense apical patch of 304long erect setae and with a slender upwardly directed process at apex. . chagasi (Fig. 61) Paramere otherwise, without dense patch of dorsal setae but with a group of 5-20 setae near base of subterminal dorsal arm 2

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2. Main lobe of paramere slender and upturned apically, with a group of about 15 long setae near base of dorsal arm . . . .bernalei (Fig. 61) Main lobe of paramere broad, not upturned apically, with about 5 short stout setae near base of dorsal arm fo.ivtigi (Fig. 62) Series arthuri. The male of L. bispinosa, the sole representative of this series in Colombia, differs from other Psycho dopy gus males by having only 2 spines on the style, both of which are terminal. Series pancavens-is 1. Style with 5 (rarely 6) strong spines. Paramere as figured .... davisi (Fig. 67) Style with 3 strong spines and 1 smaller bristle. Paramere otherwise 2 2. Paramere simple, slender distally, without arms or other extensions ..... guyanensis (Fig. 68) Paramere complex, with ventral and/or dorsal arm 3 3. Paramere with recurved apical setae on main lobe and ventral arm; dorsal arm present. Aedeagus long and slender, reaching to ends of longest recurved setae on main lobe of paramere . . Tsaia-va (Fig. 71) Paramere otherwise, without dorsal arm or recurved setae. Aedeagus rather short and stout 4

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-31 4. Main lobe of paramere with 2 tufts of blade-like setae; ventral arm wirh 2 enlarged setae, one terminal and one sub terminal , panamensis (Fig. 70) Main lobe of paramere with 1 group of setae, blade-like or not; ventral arm with smaller, more numerous setae 5 5. Paramere as shown, ventral arm inserted near base of main lobe. . . hirsuta hivsuta (Fig. 67) Paramere otherwise, ventral arm shorter, inserted at or near apex of main lobe 6 6. Mesonotum and pleura pale ........ car-rerai oawevai (Fig. 65) carrevai thula (Fig. 66) Mesonotum partly or entirely pigmented, contrasting with pale pleura* 7 7. Paramere with a short ventral arm (length subequal to width of coxite) apical setae (20+) shorter, not blade-like. Mesonotum, katepisternum, procoxae, and mesocoxae pigmented ayrozai (Fig. 64) Paramere with a longer ventral arm, its length definitely greater than width of coxite; apical setae (less than 20) blade-like and long. Only mesonotum pigmented noat-icota (Fig. 69) *The katepisternum of L. ayrozoi is faintly pigmented but the rest of the pleura is decidedly paler than the mesonotum.

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-315Females 1. Common sperw duct completely smooth-walled, not rugose* bevaalei ehagasi fairtigi (Fig. 62) Common sperm duct rugose in part and/or with transverse striations. 2 2. Cibarium with small subequal vertical teeth in 1-3 transverse rows, without longitudinal rows reourva (Fig. 71) Cibarium with vertical teeth of varying size, those in the middle enlarged and forming 2 or more longitudinal rows, irregular or not. 3 3. Sparmatheca shorter than individual duct 4 Spermatheca longer than individual duct .... 5 4. Common sperm duct with complete transverse striations guyanensis (Fig. 68) Common sperm duct with incomplete transverse striations appearing as short, thick lines sp . of Tres Esquinas (Fig. 61) 5. Mesonotum and pleura pale 6 Mesonotum partly or entirely pigmented, contrasting with pale pleura 7 *I am not able to separate these females without associated males. The female of L. bermalei, presently unknown, probably resembles the other 2 species, all of which belong in the series sqi'amiventris .

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-316• 6. Lai-rum shorter than 0.30 rum or shorter than combined length of scape and f Lagellomere I carrevai. aavrerai (Fig. 65) Labrum longer than 0.30 mm or greater than combined length of scape and f lagellomere I cavvevai thula (Fig. 66) 7. Cibarinm with short inwardly directed horizontal teeth. Katepisternum, procoxae, and mesocoxae pigmented cyrosai (Fig. 64) Cibarium with longer erect horizontal teeth. Katepisternum, procoxae, and mesocoxae pale 8 8. Common sperm duct completely smooth-walled below rugose portion . . 9 Common sperm duct with transverse striations or dot-like thickenings below rugose portion 10 y. Terminal annulation of spermatheca asymmetrical. Mesoscutellum pigmented panarnensis (Fig. 70) Terminal annulation of spermatheca symmetrical. Mesoscutellum pale noaticola (Fig. 69) 10. Common sperm duct with complete or nearly complete transverse striations. Stem of genital fork broad, blade-like . . bispinosa (Fig. 63) Common sperm duct heavily rugose below junction of individual ducts; rest of duct with dot-like thickenings, not transversely striated. Stem of genital fork more slender, subacute at tip 11 11. Individual sperm duct less than half the length of spermathcea; rugose section of common duct swollen near junction of individual ducts. . hirsute hivsuta (Fig. 67)

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-317Individual sperm duct over half the length of sperniatheca; rugose section of common duct not swollen, more or less equal in width throughout davisi (Fig. 67) Series squamiventris 74. Lutzomyia (P.) bernalei (Fig. 61) Phlebotomus bernalei Osorno, Morales, & Osorno, 1967: 30 (6* holocype, Araracuara, Caqueta, Colombia). Lutzomyia bernalei, '. Martins et al., 1968: 22 (mention). Osorno el: al., 1972a: 45 (listed). Psyehodopygus bernalei: Forattini, 1971a: 104 (listed). Forattini, 1973: 392 et seq. (review, keyed, fig.). Distribution: Colombia (Caqueta) . Material examined:
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318Lutzomyia unisetosa: Barretto, 1962, 94 (listed). Barretto, 1966: 141 (keyed). Martins et al . , 1968: 1 et seq. (as synonym of chagasi) . Llanos, 1973: 31 (, redescr., figs., refs.) . Psychodopygus unisetosus: Forattini, 1971a: 104-105 (listed). Psychodopygus chagasi: Forattini, 1973: 392 et seq. (review, keyed) Fraiha et al., 1974: 89 (cf, figs., biting man). Distribution: Colombia. (Caqueta) , Material examined: Colombia. 4 tfd 1 , Tres Esquinas (Caqueta), light trap, 10 Nov. 1971, C.J.M. Brazil. 1 o. , km 13 Macajai-Caracarai (Roraima) , human bait, 16 July 1973, H. Fraiha. Discussion: This man-biting species, precinctive in the. Amazon Basin, is easily recognized but I am unable to separate the females from those of L. fairtigi, L. squamiventris , and related species in the series squamiventris without associated males. 76. Lutsomyia (P.) fairtigi (Fig. 62) Lutsomyia fairtigi Martins, 1970: 279 (c? holotype, Villavicencio, Meta Dept., Colombia). Martins & Morales, 1972: 366 (listed). Phlebotomus squamiventris: Fairchild & Hertig, 1951a: 414 (6", £, redescr., figs., not squamiventris Lutz & Neiva, 1912). Psychodopygus fairtigi: Fraiha et al., 1971: ^95-496 (cf. to wellccmei) . Forattini, 1971b: 152-154 (Mato Grosso, Brazil). Forattini. 1973: 392 et seq. (distrib., figs., keyed).

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-319 Distribution: Colorrbia (Meta, Caqueta) , ?Brazil (Mato Grosso). Material examined: Colombia, c? holotype, o allotype, 6 o_g paratypes, Vil lavicancia (Meta), stable trap, 10 June 1948, C.Y. Chow. 3 d*d" s Tres Esquinas (Caqueta), light trap, 10 Nov. 1971, C.J.M. Discussion: A little known species^ L. fairtigi was first collected in a donkey-baited stable trap at Villavicencio and later in a light trap at 'Ires Esquinas. Forattini (1971b, 1973) lists this species (females only) from Salto Augusto (Mato Grosso), Brazil, but its presence there can only be confirmed when the male is discovered. Series avthuri 77. Lutzomyia (P.) bispinosa (Fig. 63) Phlebotomus bispinosus Fairchild & Hertig, 1951a: 410 (6* holotype, La Victoria, Cerro Jefe, Panama). Fairchild, 1955: 194 (listed). Lewis & Garnham, 1959: 80-81 (o. figs., Belize). Fairchild h Hertig, 1959: 121, 123 (distrib., Central America). Forattini, 1960: 479 (Amapa, Brazil). Williams et al . , 1965: 65 et seq. (Belize). Lutzomyia bispinosa: Barretto, 1962: 94 (listed). Theodor, 1965: 188 (listed). Barretto, 1966: 141, 144 (keyed). Williams, 1970: 332 et seq. (Belize). Chaniotis et al., 1971a: 344-345 (collecting data, Panama). Christensen, 1972a: 88 (listed). Lewis, 1975a: 500 et seq. (mouthpart morphol.). Lainson et al., 1977 (Mato Grosso, Brazil). Psyc-liod.opygus bispinosus: Forattini, 1971a: 104 (listed). Forattini, 1973: 170 et seq. (distrib., cf, o, keyed, figs.). Distribution: Belize, Honduras, Nicaragua., Panama., Colombia (Choco)., Ecuador, Northern Brazil.

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-320tiatevial examined: Colombia. 24 oo, Curiche (Choco) , biting man, tree trunk, Malaise, light, & Shannon traps, May-Oct. 196 7, D.G.Y. 1 c. , Alto Curiche (Choco), light trap, 8 Sept. 1967, D.C.Y. 10 6"cT, 66 05, Teresita (Choco), biting man, tree trunks, in Malaise, light, & Shannon traps, May-Dec 1967, D.G.Y. Ecuador. 3 op, Rio Napo at Limoncocha (Napo), flight trap, 22-24 May 1976, D.G.Y. & T.E. Rogers. Panama.
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-321Fairr.hild & Hertig, 1951a: 401-404 (listed, keyed). Barretto, 1951: 212 (distrib.). Forattini, 1960: 471 et seq. (mention). Ortiz & Alvarez, 1972: 140, 143 (listed). Ortiz, 1972b: 222 et seq. (in part, taxonoraic discussion, keyed, figs.). Phlebotomus davisi: Coutinho, 1939: 181 (o, not davisi Root, 1934). Sergentomyia ayrozai: Barretto, 1955b: 249 (cf. to pessoana) . Barretto & Zago, 1956: 178-179 (Petropolis, Brazil). Lutzonvjia ayrozai: Barretto, 1962: 94 (listed). Barretto, 1966: .140 et s?q. (in part, keyed). Barreto, 1969: 463 (Valle, Colombia). Osorno et al . , 1972a: 45 (mention). Young, 1973: 108, 111 (mention). Lutzomyia amazonensis : Aitken et al . , 1968: 254 (Trinidad, not amazonensis Root, 1934). Lutzomyia tintinnabula Christensen & Fairchild, 1971: 301 (cf, o, Darien Frov., Panama). Osorno et al., 1972a: 48 (Caqueta, Colombia). Young, 1973: 108, 111 (mention). Lewis, 1975a: 502 et seq. (mouthpart morphol., keyed, Para, Brazil). NEW SYNONYM. Psycho dopy gus ayrozai: Forattini, 1971a: 104 (listed). Forattini, 1973: 392 et seq. (in part, distrib., figs., keyed, taxonomy). Gomes, 1975: 9 (listed) . Forattini & Galati, 1977: 25 (o , figs., discussion). Phlsbotomus tintinnabulus : Ortiz & Alvarez, 1972: 140 et seq. (listed). Ortiz, 1972b: 222 et seq. (taxonomy, discussion). Lutzomyia "Turure sp." Lewis, 19 75a: 502 et seq. (mouthpart morphol., sansilla, keyed, Trinidad) . Psychodopygus tintinnabula.: Gomes, 1975: 9 (listed). Ward, 19 76: 239 (mention) . Distribution: Colombia. (Antioquia, Boyaca, Caqueta, Choco, Valle) , Trinidad, French Guiana, Brazil, Panama.

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-322Material examined". Colombia. 22.5 do", 246 0.5, Rio Anori (Antioquia) , light traps, May 1970, C.H.P. 1 o*, 4 o<^> , sane data but flight trap, 3 May 1970. 2 o^, same data but biting man, 25 June 1970. 107 eft?, 204 00, same data but light traps, Sept. 1970, D.C.Y. 1 q, same data but tree crunk, 21 Sept. 1970. 2 88, 4 go., Puerto Boyaca (Boyaca) , light trap, 10 Aug. 1971, C.J.M. 4 38, 8 00, Trcs Esquinas (Caqueta) , light trap, 10 Nov. 1971, C.J.M. 2 68, 177 00, Curiche (Choco) , biting man, Malaise, Shannon, & light traps, April-Sept. 1967, D.G.Y. 3 68, 5 5$, Alto Curiche (Choco), light & Shannon traps, July 1967, D.C.Y. 2 00 , Anchicaya Dam (Valle), flight trap, 10 Aug. 1973, D.G.Y. & R.C.W. 1 g, same data but 10 June 1975, R.C.W. 1 o, 25 km E of Buenaventura (Valle), light tr^p, 12 Aug. 1973, D.G.Y. & R.C.W. Brazil. 1 £, Manacapura (Amazonas) , 1936. 1 p, near Manaus (Amazonas), 29 Oct. 1974, J. Arias. 2 68, Mata Prefeitura, Oliveira (Minas Gerais) , 22 March 1960, Silva. 1 d\ 1 o, M.D. dc; Rio Santa Antonio (Minas Gerais), 3 Jan. 1970, Silva. 5 6*0*, 2 00 , Belem (Para), flight trap, 2-5 Aug. 1974, D.G.Y. 1 6 (cotype) , Horto Florestal, Sao Paulo (Sao Paulo), 1941, M. Barretto & J. Coutinho. 1 8, S. Jose Campos (Sao Paulo), at light, 11 Nov. 1940, M. Barretto & J. Coutinho. Panama. 1 6", 1 o (paratypes of tintinnabula) , Cerro Quia, Finogana (Darien Prov.), 14 March 1970, F. Beam. Trinidad. 1 (j>, Bush Bush Forest, Nariva Swamp, 20 Nov. 1962, T.H.G. Aitken. Other specimens, both males & females, from Trinidad, various dates and localities, to be treated in another paper on the phlebotomine fauna of Trinidad. Discussion: My treatment of L. tintinnabula as a junior synonym of L. aypozai is based on the following considerations. The female of L. aupozai, correctly associated with the male and described by Forattini and Galati (1977) , was believed earlier to be that

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-323described by Barretto and Coutinho (1943) but, their 1943 description represented that of L. guyanensis (Floch and Abonnenc) , not L. auvozai (Barretto and Coutinho) . This misled others including Christensen and Fairchild (1971) who described L. iintinnabula as a distinct species primarily on the basis of female characters, notably the different spermathecae and ducts. The true female of ayvozai and that of L. tintinnabu'la are indistinguishable, however. The character states used in separating the males of tintinnahula and ayvozai-, discussed by Christensen and Fairchild (1971), are variable and reflect intraspecif ic , not interspecific, variation. Delta, of wing venation is shorter than, equal to or longer than gamma depending on the individual specimen. The shape of the parameres varies slightly according to geographic locality but not to the extent that it repi-esents a difference at the species level. One feature shared by all the specimens examined is that the procoxae, mesocoxae, and areas just above them on the pleura are moderately infuscated. Lutzomyia ayvozai is a widespread, anthropophilic species which probably occurs in other parts of Colombia, Brazil, and possibly Venezuela, Peru, and Ecuador. At Curiche (Choco) from April to Dec, 1967, we collected 144 females on human bait, the majority of which were taken in June (122 specimens) . The total catch represented 3.8% of all biting specimens taken during this 8 month period (141 man hours) . 79. Lutzorrryia (P.) cavrovai car'vevai (NEW STATUS) (Fig. 65) Phlebotornus cavrevci Barretto, 1946d: 286 (c? holotype, o, Vega Grande, Restrepo, Meta Dept., Colombia). Fairchild & Hertig, 1951a: 401

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324et seq. (listed, keyed). Ortiz & Alvarez, 1972: 140 et seq. (listed, cf. to parimaensis) . Ortiz, 1972b: 222 et seq. (taxonomic discussion, figs. , keyed) . ?Phlebotomus davisi: Root, 19 34: 233 (in part, c. only). Sergentomyia carrerai: Barretto, 1955a: 82 (listed). Barretto, 1955b: 249 (cf. to pessoana) . Sergentomyia pessoana Barretto, 1955b: 247 (
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Sao Paulo, Faculdade de Saude Publica. Brazil. 2 o, Caravelas (Bahia) , • r an .1931, N. Davis. 1 cf, ] o, near Bacuri, N of Maraba (Para), flight trap, 2 C ' Oct. 1974, J.F. Reinert. 1 o, same data but 1 Nov. 1974. 6 3d, 40 oj, near Altamira (km 165) (Para), biting man & Shannon trap, 6 Nov. 1974, J.F. Reinert. 1 o, same data but flight trap, 13 Nov. 1974. 1 d*, 1 o, Itatiaia (Rio de Janeiro), at light, Feb. 1955, M. Barretto. Ecuador. 2 no, Limoncocha, near Rio Napo (Napo) , flight & light traps, 20-24 May 1976, D.G.Y. & T. Rogers. Peru. 1 cf, 2
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326The earrerai carvevai females share other features which should be noted. The base of the rugose section of the common sperm duct is distal ly tapered or "v" shaped. The area within the "v" is non-sclerotized as shown by Root (1934, fig. 3) and Fig. 65 (this paper). This feature, present in all females including the female identified as "pessoana" by Dr. Barretto and seen by me, is seemingly unimportant but it helps to separate the subspecies of L. aarrerai and provides additional evidence that oavrerai and pessoana are conspecific. Other character states believed to be diagnostic by Barretto (1946d, 1966) include those of the spermathecae (12-13 segments in cavverai vs. 10 segments in pessoana) and the position of the horizontal teeth of the cibariuai. These are variable characteristics which, in my opinion, do not reflect specific differences in the absence of other characters. The males of aarrerai from Amazonia and "pessoana" from southern Brazil are indistinguishable. The characters of the genitalia used by Barretto (1966) and Martins et al. (1973) vary according to geographic locality and are not diagnostic by themselves. Present data indicate that L. a. aarrerai, a man-biter, is locally common in parts of the Amazon Basin but extends south into southern Brazil and north into Colombia, East of the Andes (ais-Andean distribution) . 80. Lutzorrrjia (P.) aarrerai thula n. ssp. (Fig. 66) Phlebotomus paraencis: Fairchlld & Hertig, J951a: 404 et seq., not paraensis Costa Lima, 1941 (keyed, figs., descr.). Fairchild & Hertig, 1959: 122, 123 (distrib., Central America only).

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-327Phlebotomus pessoanus: Hanson, 1961: 320 (breeding sites, Panama). Johnson & Hertig, 1961: 765 et seq. (rearing). Thatcher, 1968a: 295 (hosts, Panama). Lutzomyia pessoana: Chaniotis et al., 1971a: 344 et seq. (collecting data, Panama). Chaniotis et al., 1971b: 417 et seq. (biting habits, Panama).. Tesh et al., 1972: 90, 92 (hosts, Panama). Chaniotis et al . , 1972: 94 et seq. (resting sites, Panama). Chaniotis & Correa, 1974: 115-116 (biting activity, Panama). Chaniotis, 1974b: 501 (keyed). Rutledge & E.llenwood, 1975a, b,c: 71 et seq. (breeding sites, pop. dynamics, Panama). Lutzomyia paraensis: Barreto, 1969: 465-466 (Valle, Colombia, not paraensis Costa Lima) . Psyckodopygus pessoanus: Forattini, 1971a: 105 (listed). Forattini, 1971b: 173-176 (as a synonym of paraensis, pessoanus, & fairchildi) . Ward, 1976: 239 (larva, keyed). Distribution: Colombia (Antioquia, Choco, Valle), Honduras, Costa Rica, Panama, Western Ecuador. Male: As described and illustrated by Fairchild and Hertig (1951a, as P. paraensis) with the following additions based on male holotype (no. 2328). Wing length 1.88; width 0.51. Head height 0.33; width 0.37. Eyes large, separated by 0.05 or by distance = to 2.2 facet diameters. Flagellomere I (0.24 long), 1.2 x length of II + III. Length of palpal segments: 1 (0.03), 2 (0.09), 3 (0.12), 4 (0.04), 5 (0.04); palpal sensilla (ca. 10) on distal 3/4 of segment 3. Labrum length 0.20. Pharynx length 0.18. Pleura with 17 upper and 5-11 lower episternal setae. Length of wing vein sections: Alpha (0.43), beta (0.26), delta (0.16), gcwma (0.12). Length of femora, tibiae, and basitarsi: Forleg, 0.78, 1.13, 0.76; midleg, 0.74, 1.26, 0.78; hindleg, 0.88, 1.38, 0.88. Style

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-328(0.12 Long). Coxite (0.19 long x 0.05 wide). Lateral lobe (0.26 long). Genital pump (0.18 long), each filament 0.50 long or 2.8 x length of pump . Female: As described and illustrated by Fairchild and Hertig (1951a, as P. paraensis with the following additions based on female allotype (no. 2228). Wing length 2.15; width 0.66. Head height 0.41; width 0.37. Eyes large, separated by 0.08 or by distance = to 4.5 facet diameters. Flagellomere J (0.25 long), 1.2 x length of II + III length of palpal segments: 1 (0.06), 2 (0.13), 3 (0.17), 4 (0.05), 5 (0.06). Labrum length 0.36. Pharynx length 0.19. Pleura with 13-18 upper and 6-8 lower epioternal setae. Length of wing vein sections: Alpha (0.62), beta (0.25), delta (0.31), gamma (0.18). Length of femora, tibia, and basitarsi: Foreleg, 0.81, 1.15, 0.76; midleg, 0.76, 1.32, 0.80; hindleg, 0.88, 1.47, 0.88. Material examined: Panama. 6 holotype (no 2328), Cerro Camp ana (Panama Prov.), Shannon trap, 24 Aug. 1950, M. Hertig & G. Fairchild. o allotype (no. 2228), same data but biting man, 21 June 1950. 15 go paratypes (nos. 2223-2227 & 2229-2240), same data as allotype. 2 6*0*, 11 jo paratypes (nos. 634-646), Rio Changena (Bocas del Toro) , Shannon trap, 8 Sept. 1961, P. Galindo. 2 dc? paratypes (nos. 647-648), upper Pacora River (Panama Prov.), light trap, 21 Oct. 1953, F.S. Blanton. Colombia. 2 33, 4 op paratypes (nos. 649-654), Rio Anori (Antioquia) , light traps, Sept. 1970, D.G.Y. Honduras. 1 o* paratype (no. 4541), Lancetilla Valley, Tela, light trap, 25 Nov. 1953, W. Hiis. Other specimens, mostly preserved in alcohol but not part of the type series include the following. Colombia. ] p, Rio Anori (Antioquia), light trap, 19 April 1970, C.H.P. 8 dcf, 4 o^ , sane data but May 1970. 7 dtf, 10 o^,

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329same data but Sept. 1970, D.G.Y. 5 tfcf, 161 oo, Curiche (Choco) , JuneJuly, Sept., Nov. 1967. 9 d'o", 89 oo , same data but Teresita (Choco), April-Nov., 1967. 1 6*, 25 km E of Buenaventura (Valle) , light trap, 12 Aug. 1973, D.G.Y. & R.C.W. Costa Rica.. 1 q, N of Marina (Alajuela) under dead leaves, 13 Feb. 1960, W.J. Hanson. Ecuador. 1 o., 16 km E of Santo Domingo de Los Colorados (Pichineha) , flight trap, 28 May 1976, D.G.Y., T. Rogers.. & G. Fairchild. Discussion: Unlike Barretto (1966), I am convinced that L. cawevai (Barretto) is not only conspecific with L. pessoana (Rarretto) but that it differs from the taxon described as Phlebotomus paraensis by Fairchild and Hertig, 1951a, and subsequently identified as L. pessoana. by other workers in Panama. Accordingly, 1 am assigning this taxon a new name L. cawevai thula, the subspecific epitaph being derived from the Latin word, "thule," meaning farthest north. This form and the nominate subspecies are allopatric, the former subspecies occurring west of the Andes from Ecuador to Colombia, then into Panama and Central America (trans-Andean distribution, Haffer, 1967, 1974) . Lutzomyia c. cawevai although widespread in the Amazon Basin and southwards, does not extend into Central America (cis-Andean distributtion) . The following character states, seemingly minor but consistent, are used to separate the females. The length of the labrum of L. c. cawevai (n 20) from various localities ranges from 0.24 mm to 0.27 mm (average length 0.25 mm). The labrum is shorter than the combined length of the pedicel and the first f lagellomere. in contrast, the labrum of L. c. tkula is much longer, ranging from 0.33-0.45 (average length 0.38, n = 21) and is greater than the length of the pedicel + f lagellomere I. The

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-330palpi are longer in thula as well. The "v" shaped clearing at the base of the rugose section of the common sperm duct is conspicuous in females of the nominate subspecies but is absent or nearly absent in females of L. c. thula. I am not able to separate the males of these subspecies without knowledge of locality or in the absence of females. The eggs of each appear to be similar (Ward and Ready, 1975; Zimmerman et al., 1977) but further study is needed to detect possible differences. Hanson (1968) partially illustrated the fourth instar larva of thula (as pessoonus) noting that it and the pupa are almost indistinguishable from those of L, panamsnsis. We collected a total of 135 females of L. a. thula in routine human bait collections at Curiche in May and June 1967 (52 man hours) . One specimen was taken in April (17 man hours), 2 in August (25 man hours) but none was collected in July, Sept. to Nov., 1967 (47 man hours). Based on soil emergence traps (Rutledge and Ellenwood, 1975a), specimens of L. a. thula (as pessoana) were most common from May to August in Panama. Other information on the bionomics of this taxon in Panama include the following. Most adults rest on the undersides of green leaves during the day (Chaniotis et al., 1971a, 1972). Females are anthropophilic, being especially common in mature forests where they bit primarily at ground level (Chaniotis et al., 1971b) during the day and night. Precipitin tests indicate that a variety of mammals are fed upon, the edentates accounting for the majority (72.9%) of 96 reacting blood meals in one study (Tesh et al., 1972). Immature stages have been recovered from the open forest floor (Hanson, 1968, Rutledge and Ellenwood, 1975a).

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81. Lutzomyia (P.) davisi (Fig. 67) Fhlebot.omi.is davisi Root, 1934: 242 (o* holotype, Fordlandia, Para, Brazil; o, not davisi Root). Barretto, 1947: 196 (cataloged). Fair-tchild & Hertig, 1951a: 401 et seq. (discussed, keyed). Floeh & Abonnenc, 1952: 157 (French Guiana, o*, figs.). Forattini, 1950: 468 et seq. (in part, 6 s only, figs.). Ortiz & Alvarez, 1972: 140 (cf. to pai'-imaensis) Ortiz, ]972b: 222 et seq. (discussed, keyed, 6* figs.; o not davisi Root.) . Phlebotomns vooti Mangabeira, 1942a: 112 (6*, Aura, Belem, Para, Brazil). Fairehild & Hertig, 1951a: 402 et seq. (discussed, keyed). Ortiz & Alvarez, 1972: 140 (cf. to parimaensis) . Ortiz, 1972b: 222 et seq. (in part, keyed, figs.). Lutzomy ia davisi: Barretto, 1962: 94 (listed). Barretto, 1966: 142 (keyed; o not davisi Root). Lewis et al . , 1970: 212 et seq. (parous rates, ovariole fig.). Osorno et al . , 1972a: 46 (Caqueta, Colombia). Llanos, 1973: 31 (d\ p, descr., figs., Peru). Martins et al., 1973b: 419 (p descr., ref s , ) . Lewis, 1975a: 502 et seq. (r.outhpart morphol.). Llanos et al . , 1975b: 670 (Peru). Llanos et al . , 1976: 480 (Peru). Lutzomyia, voobi: Barretto, 1962: 94 (listed). Barretto, 1966: 142 (keyed). Martins et al . , 1973b: 419 (as synonym of davisi). Psy chodopygus davisi: Forattini, 1971a: 105 (listed). Forattini, 1973: 163 et seq. (in part, 6" only, gen. review). Ward & Ready, 1975: 128 (egg, figs.). Ward, 1976: 233 (larva descr., fig.). Lainson et al . , 1977 (Rio Aripuana, Mato Grosso, Brazil). Fsu chodopygus rooti: Forattini, 1971a: 105 (as synonym of davisi). Ward et al., 1973: 178 (Para, Brazil). Lainson et al . , 3 973: 190 (Para, Brazil) .

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-33: Distribution: Colombia (Amazonas, Caqueta) , • Ecvjadov, Peru, Bolivia, French Guiana, Brazil. Material examined: Colombia. 3 cfcf, 1 9, 18 km W of Leticia (Amazonas), light trap, 26 July 1973, D.G.Y. & R.C.W. 5 cftf, 4 0.0., Tres Esquinas (Caqueta) , light trap, 10 Aug. 1971, C.J.M. Brazil. 1 d", Rio Aripuana at Humboldt (Mato Grosso) , tree trunk, 20 Aug. 1974, D.G.Y. & J. Shaw. 4 6*0*, 6 o^, Belem (Para), flight trap, 30 July-6 Aug. 1974, D.G.Y. 1 cf, 8 qo, near Bacuri, N of Maraba (Para), biting man, light, Shannon & flight traps, 23-31 Oct. 1974, J.F. Reinert. 11 99, W of Altamira, km 164 (Para), biting man, 6 Nov. 1974, J.F. Reinert. 5 OcT, 12 00, same data but Shannon & flight traps, 6-7, 12 Nov. 1974. Ecuador. 16*, 1 0, Rio Napo at Limoncocha (Napo) , flight trap, 19 May 1976, D.G.Y. & T. Rogers . Discussion: Lutzomyia davisi, an anthropophilic species (Ward et al. , 1973), is widely distributed in the Amazon basin, occurring as far north as central Colombia, east of the Andes. A recently described, but doubtfully distinct, species from Venezuela, L. parimaensis (Ortiz and Alvarez) may be conspecific with davisi but the male is unknown and I have not seen the holotype female or para types. Forattini (1973) refers Phlebotomus sp. no. 780 of Floch and Chassignet, 1948, to davisi but this is questionable based on the description and figures of that taxon (o only) and by the fact that Forattini (1960, 1973) misassociated the sexes of davisi. Phlebotomus sp. no. 780 is more likely the female of L. hirsuta but this is only speculation as J. have not examined material referable to P. sp. no 780 from French Guiana.

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-333The true identity of the davisi female was confused for many years but Martins et al. (1973b) correctly associated the sexes and reviewed the literature, clarifying inisidentif ications of others. Based on their studies and those of my own, I also have no doubt that L. Tooti is conspecific with L. davisi. 82. Lutzomyia (P.) guyanensis (Fig. 68) Phlebotomus guyanensis Floch & Abonnenc, 1941a: 17 (p holotype, Montana Lucifer, French Guiana). Barretto, 1947: 203 (refs.). Fairchild h Hertig, 1951a: 401, 404 (mention, keyed). Floch & Abonnenc, 1952: 173 (o, redescr., figs.). Ortiz & Alvarez, 1972: 140 (mention). Ortiz, 1972b: 221 et seq. (o, keyed, figs., discussion). Phlebotomus geniculatvs Mangabeira, 1941a: 245 (c? holotype, Aura, Belem, Para, Brazil). Fairchild & Hertig, 1951a: 401 et seq. (, descr., figs., Panama). Bruijning, 1957: 229 (Surinam). Lewis & Garnham, 1959: 80-81 (o, figs., Belize). Johnson & Hertig, 1961: 765 (rearing). Hanson, 1968: 58 (larva descr., figs.). Phlebotomus ayvozai: Barretto & Coutinho, 1943: 183 (o not Barretto & Coutinho, 1943) . Lutzomyia geniculate: Barretto, 1962: 94 (listed). Barretto, 1966: 141, 145 (keyed). Williams, 1970: 332 (Belize). Osorno et al . , 1972a: 46 (Caquata, Colombia). Lewis, 1975a: 502 et seq. (mouthpart morphol.) . Llanos et al . , 1975b: 670 (Peru). Lutzomyia guyanensis: Barretto, 1962: 94 (listed). Barretto, 1966: 145 (keyed). Lewis, 1975a: 502 et seq. (mouthpart morphol.).

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-334Psyohodopygus geniculatus: Forattini, 1971a: 105 (listed). Forattini, 19 73: 124 et seq. (as synonym of guyanensis). Shaw et al., 1972: 720 (biting man). Ward et al., 1973: 178 (biting man. Para, Brazil). Lainson et al . , 1973: 190 (Para, Brazil). Ward, 1976: 239 (larva keyed) . Psyohodopygus guyanensis: Forattini, 1971a: 105 (listed). Forattini, 1973: 124 et seq. (gen. review). Forattini & Galati, 1977: 25 (o, of. to ayrozai) . Distribution: Belize, Panama, Colombia. (Caqueta, Choco) , Ecuador, Peru, SuY-lnam, Brazil. Material examined'. Colombia. Id 1 , 2 ra, Tres Esquinas (Caqueta), light trap, 10 Nov. 1971, C.J.M. 117 op, Curiche (Choco), biting man, light & Shannon traps, June-Sept. 1967. 16 oo, Alto Curiche (Choco), biting man, light traps, June-Sept. 1967. Brazil. 6 oo, Belem (Para), light trap, 18 Oct. 1972, D.G.Y. 1 o, Cabeca da Anta (Sao Paulo), at light, 1941, M. Barretto & J. Coutinho. Ecuador. 6 S3, 208 o^> , Rio Napo at Limoncocha (Napo), light traps, 19-23 May 1976, D.G.Y. & T. Rogers. 5 qq , same data but flight traps. Panama. 8 op, Almirante (Bocas del Toro), Shannon trap, 22 June 1951, W. Hils . 1 o*, same data but light trap, 30 March 1953, A. Quinones. Discussion: Although Forattini (1973) is probably correct in treating L. genicula.ta as a junior synonym of L. guyanensis, there remains the possibility that they are distinct taxa. Additional material, particularly males, from or near the type locality of guyanensis , is needed to confirm the synonomy. At Curiche in 1967, we collected 102 guyanensis females biting man, the majority of which were taken in June (98 specimens, 24 man hours) .

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-335The remaining 4 man-biting females were captured in July and August (43 nan hours) . Williams (1970) in Belize dissected 19 females but. found no flagellates. He captured specimens on dead leaves on the forest floor, on man, in light traps, and in a rodent-baited Disney trap. 83. Lutzomyia (P.) hirsute hirsute NEW STATUS (Fig. 67) Phlebotomus hir-sutus Mangabeira, 1942a: 116 (
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Lutzomyia paraensis: Osorno et al., 19 72a: 48 (not paraensis Costa Lima; Caqueta, Colombia) . Pcychodopygus amazonensis : Ward ct al . , 1973: 178 (in part, 6" only) . Psycho dopy gus davisi: Forattini, 1973: 407-408 (in part, d*only, figs.). Lutzomyia sp. "c." Velasco, 1973: 84 (2, Cerro de Polini, Bolivia). Distribution: Colombia (Caqueta), Ecuador, Peru., Bolivia, French Guiana, Brazil. Material examined: Colombia. 6 eft/, 13 op, Tres Esquinas (Caqueta), light trap, 10 Nov. 1971, C.J.M. 1 o", 2 go., El Palmar, Solano (Caqueta), light trap, 31 March 1972, C.J.M. (INPES) . Bolivia. 1 o., Cerro de Polina (Nor Yungas) , tree buttress, 30 June 1971, J. Velasco. Brazil. 2 90, Serra do Navio (Amapa), light trap, 21 Sept. 1972, D.G.Y. & H. Fraiha. 1 o, near Bac.uri, N of Maraba (Para), flight trap, 22 Oct. 1974, J. Reinert. 1 6 1 , 1 o, W of Altamira, km 164 (Para), Shannon trap, 6 Nov. 1974, J. Reinert. Ecuador. 15 S<$ , 36 o
PAGE 349

-337 Psychorbpygies females by the shape of the common sperm duct, i.e., the rugose section is relatively narrow at the base but becomes noticeably swollen apically where the individual duett; join. In other words, the shape of the rugose section is similar to that of a tapered carrot. The remaining section of the common duct is partially striated, the. thickenings forming incomplete lines or distinct dots. In this respect the common duct of L. davisi is similar but the rugose part of the sperm duct of that species is not swollen apically and the individual ducts are longer (Fig. 67). Fhlebotomus sp. no. 780 of Floch and Chassignet, discussed earlier under davisi, resembles hirsute, in the shape of the common sperm duct but it appears to be heavily rugose throughout ics length (Floch and Chassignet. 1948, fig. 1). If this is actually the case, then sp. no. 780 is a distinct species but, if not, then it probably represents I. li. hivsuta, the male of which was taken also at Baduel, French Guiana (Floch and Chassignet, 1947a) . It seems appropriate here to describe the previously unknown female of L. h. nicaraguensis based on 5 specimens from Bocas del Toro Prov. , Panama. Fhlebotomus nicavaguensis Fairchild & Hertig, 1961a: 26 (6*, holotype, Villa Somoza, Nicaragua) . Female (Fig. 12): Wing length 2.10-2.42; width, 0.68-0.73. Head, mesonotum, and abdominal tergites strongly pigmented, contrasting with pale pleura. T'rocoxae nearly as dark as mesonotum, other coxae pale (all coxae pale in L. hivsuta. hirsuta) . Head height 0.40-0.43; subequal to width. Eyes separated by 0.12 or distance = to ca. 6 facet diameters. Flagellomere I (0.27-0.29 long), 1.3 x length of II + III; uscoids nearly reaching end of flagellomeres, on all except last 3. Length of palpal

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-338segments: 1 (0.05-0.06), 2 (0.17-0.20), 3 (0.19-0.20), 4 (0.06), 5 (0.070.09); palpal sensilia (ca. 12) on distal 1/3 of segment 3. Labrum 0.370,42 long. Cibarium with 4 horizontal teeth, spaced as shown, the median pair inclined towards middle; pigment patch as shown, moderately pigmented; pharynx 0.18-0.20 long, unarmed. Pleura with 12-30 upper and 6-11 lower episternal setae. Length of wing vein sections: Alpha (0.230.29), beta (0.11-0.13), delta (0.08-0.12), -ap-.ma (0.06-0.10). Length of femora, tibiae, and basitarsi of slide 7943: Foreleg, 0.88, 1.42, 0.95; midleg, 0.83, 1.55, 1.05; hindleg, 0.98, 1.85, 1.11. Abdominal sternite 2 with a pair of lateral clear spots posteriorally . Tergite 8 with 10-30 setae on each side. Spermathecae and ducts as in Fig. 12. Material examined: Panajva. 4 do", 5 oo, Rio Changena (Bocas del + T Toro), Shannon trap, 7-11 Sept. 1961, P. Galindo&R. Hartmann. Nicaragua. d'holotype, Villa Somoza, 15 June 1953, P. Galdndo & H. Trapido. Discussion: I associated the sexes on the basis of collecting data, similar coloration, and by the fact that no other possible mates were taken at Rio Changena. Except for the dark procoxae (pale in h. hirsuta) , the female of nioaraguensis is indistinguishable from that of the nominate subspecies. The males are separated by this same feature and by the genitalic characters discussed by Fairchild and Hertig (1961a) . 84. Lutzomyia (P.) nocticola (Fig. 69) Lutzomyia nocticola Young, 1973: 109 (6* holotype, o_, Rio Anori, Antioquia, Colombia). Distribution: Panama, Colombia (Antioquia) , Ecuador. Material examined: Colombia. 1 cf, 1 oo (including holotype & allotype), Rio Anori (Antioquia), light traps, May 1970, C.H.P. 6 6*a*,

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-3396 oo, same data but Sept. 1970, D.G.Y. Ecuador. 28 , Cano Saddle, near Gatun, Canal Zone). Barretto, 1947: 216 (complete refs.). Fairchild & Hertig, 1951a: 399 et seq. (o*, o_, descr., discussed, figs.). Osorno et al., 1967: 28 (Meta, Colombia). Morales et al . , 1969a: 381 (Meta, Colombia). Ortiz, 1972b: 22 et seq. (cf, o_ , keyed, figs.). Ortiz & Alvarez, 1972: 140 (listed). Lutzomyia panamensis : Barretto, 1962: 94 (listed). Theodor, 1965: 188 (listed, figs.). Barretto, 1966: 142, 144 (keyed). Barreto, 1969: 465 et seq. (Valle, Colombia). Tesh et al., 1971a: 153 (blood meals). Osorno et al., 1972a: 47-48 (Colombian records). Tesh et al., 1972: 90 (blood meals). Young, 1973: 108 (mention). Chaniotis & Correa, 1974: 115 (biting habits, Panama). Chaniotis, 1974b: 501 (keyed). Lewis, 1975a: 502 et seq. (mouthpart morphol.). Rutledge & Ellenwood, 1975a:

PAGE 352

-34012 et seq. (breeding sites, refs. on bionomics). Rutledge et al. , 1975: 179 (ecology, Panama), Williams, 1976a: 604 (in caves, Belize). Phlebotomus peasoanus: Strangways-Dixon & Lainson, 1966: 192 (not pessoana Barretto, 1955, Belize). Psychodopygus panamensis : Lewis, 1965: 376 et seq. (internal morphol.). Forattini, 1973a: 89 et seq. (gen. review, figs., cf, p) . Ward, 1976: 239 (larva keyed) . Distribution: Mexico, Central America, Colombia (Antioquia, Boyaca, Caldas, Caqueta, Choco, Guajira, Meta, Norte de Santander, Santander, Valle) , Ecuador, Peru, Brazil, Venezuela. Material examined: Colombia. 40 do", 45 op, Rio Anori (Antioquia), light traps, May 1970, C.H.P. 5 do*, 8 o^, same data but Sept. 1970, D.G.Y. 4 on, Puerto Boyaca (Boyaca), light trap, 10 Aug. 1971, C.J.M. 1 d", 4 op, same data but 6 May 1973. 5 do", 6 go, Tres Esquinas (Caqueta), light trap, 10 Nov. 1971, C.J.M. 65 do*, 2386 o^ , Curiche (Choco), biting man, light, Shannon, & Malaise traps, April-Dec. 1967, D.G.Y. 12 do*, 474 op, same data but Alto Curiche, June-Dec, 1967. 72 dcf, 1528 (jig, same data but Teresita, March-Dec. 1967. Ecuador. 1 o, 16 km E of Santo Domingo de Los Colorados (Pichincha) , flight trap, 28 May 1976, D.G.Y., T. Rogers, & G. Fairchild. Panama. 3 dcf, Almirante (Bocas del Toro) , tree buttress, 21 June 1950, R. Hartmann. 5 oo, Juan Mina (Canal Zone) biting man in daytime, Sept. 1949, H. Trapido. Numerous other specimens from Panama, Mexico, Nicaragua, and Venezuela in collection at UF. Discussion: Lutzomyia panamensis , a widely distributed, strongly anthropophilic species, was the dominant man biter at Curiche, Alto Curiche, and Teresita during medical ecology studies in the Choco in 1967, At Curiche, we captured a total of 1751 panamensis females on human bait

PAGE 353

341from April Lo December (141 man hours), over half of which were taken in June (624 oo, 24 man hours) . One extraordinary biting collection on 8 June from 2000 to 2200 hours (4 man-hours) yielded 1151 Lutzomyia females of 11 species. Lutzomyia panamensis accounted for 250 of these. Most of the information on the bionomics and disease relationships of this important species is summarized by Forattlni (1973) . A few additional papers, published since then, are cited here in the species bibliography . 86 . Lutzomyia (P. ) reourva (Fig. 71) Lutzomyia reourva Young, 1973: 106 (6* holotype, o, Curiche, Choco, Colombia). Lewis, 1975a: 505 (listed). Distribution: Panama (Darien), Colombia (Choco). Material examined: Colombia. 11 3>3, 480 oo, Curiche (Choco), biting man & light, Shannon, & Malaise traps, April-Dec. 1967, D.G.Y. Panama. 1 6*, 2 op, Altos de Quia (Darien), 1970, F. Beam (GML collection). Discussion: Lutzomyia reourva and L. amazonensis (Root, 1934) , both anthropophilic, apparently are closely related. The male of the latter species, unknown for many years, was recently described by Llanos et al . (1975b). It and reourva, unlike the other series panamensis males, possess a slender, arched dorsobasal arm on each paramere but differ from each other in the shape and setation of the paramere. Lutzomyia reourva, known only from Choco Dept., Colombia, and Darien Prov., Panama, occurs well outside the range of amazonensis. Specific data on human biting collections of reourva were given in an earlier paper (Young, 1973) .

PAGE 354

17. Lutzomyia (P.) sp. of Tres Esquinas (Fig. 61) Female (description based on 3 09) : Wing length 2.0-2.3; width 0.6. Mesonotum and abdominal tergites well pigmented, head less so, rest of insect pale. Head height 0.41-0.45; width 0.38-0.40. Eyes separated by 0.10-0.11 or by distance = to ca . 5.2 facet diameters. Flageliomare I (0.23-0.25 long), 1.2 x length of II + III; ascoids simple, reaching to or beyond ends of their respective segments, on all but last 3 flagellomeres. Length cf palpal segments: 1 (0.05), 2 (0.13-0.14), 3 (0.150.16), 4 (0.05), 5 (0.08); palpal sensilla (8-12) on distal 1/3 or segment 3. Labrum 0.30-0.33 long. Cibarium with 4 horizontal teeth arranged as shown (1 o has 5) and 15-20 vertical teeth, those in middle enlarged; chitinous arch not visible; pigment patch well defined, moderately pigmented. Pharynx 0.20 long, unarmed. Pleura with 10-19 upper and 6-7 lower episternal setae. Length of wing vein sections: Alpha (0.45-0.61), beta (0.21-0.31), delta (0.05-0.16), gamma (0.18-0.20). Length of femora, tibia, and basitarsi of slide 658: Foreleg (missing), midleg, 0.83, 1.4, 0.88; hindleg, 0.93, 1.6, 0.98. Abdominal sternites entire, lacking clear areas. Tergite 8 with 10+ setae on each side. Spermatheca with 6-8 well formed annuli, the terminal one symmetrical; individual sperm duct over 2 x length of spermatheca; common duct as shown, the apical 1/5 rugose and with transverse striations appearing as complete lines, the remainder with incomplete striations more pronounced and numerous on the basal half of duct. Cerci unremarkable. D-is tpibution: Colombia (Caqueta) . Material examined: Colombia. 3 09, Tres Esquinas (Caqueta), light trap, 10 Nov. 1971, C.J.M.

PAGE 355

-343Discussion: I hesitate to formally name this species because the male is unknown and because I am not entirely convinced that it and L. guyanensis are distinct taxa. Except for differences in the sperm ducts and slightly longer individual ducts of the Tres Esquinas specimens, the females appear to be indistinguishable.

PAGE 356

Figure 61 Lutzomyi-a (P.) hevaalei — A. Male genitalia (after Osorno et al . , 1967). Male: Araracuara, Caqueta Intendencia, Colombia Lutzomyia (P.) ohagasi — B. Male genitalia. Ma.le: Tres Esquinas, Caqueta Intendencia, Colombia Lutzomyi.a sp. of Tres Esquinas, female — C. Cibarium, D. Head, E. Flageliomere II, F. Wing, G. Spermathecae .

PAGE 357

•34b-

PAGE 358

Figure 62 Lutzomyia (P.) failrtigi — A. Male head, B. Male f lagelloinere II, C. Female head, D. Male flagellomere II, E. Sperrnathecae, F. Paramere, G. Male genitalia, H. Female cibarium, I. Female cibarium and pharynx, J. Female wing, K. Male wing. Male: Villavicencio, Meta Dept., Colombia Female: Same locality as male

PAGE 359

347-

PAGE 360

Figure 63 Lutzomyia (P.) bispinosa -A. Male head, B. Male flagellomere II, C. Female head, D. Female flagellomere II, E. Female cibarium and pharynx, F. Male genitalia, G. Genital pump and filaments, same scale as ?'ig. 63F, H. Body of spermatheca, I. Spermathecae, J. Female wing, K. Male wing, L. Female cibarium. Male: Cerro Jefe, Panama Prov. , Panama Female: Curiche, Choco Dept., Colombia

PAGE 361

349ra J i/imk 4 IU .-;'^ > //I V ^. ft /f t ^

PAGE 362

Figure 64 Lutzomy-ia (P.J ayvozai — A. Male head, B. Male flagellomere II, C. Female head, D. Female flagellomere II, E. Female cibarium and pharynx, F. Genital pump and filaments, G. Male genitalia, H. Body of spermatheca, I. Spermathecae, J. Female wing, K. Male wing, L. Female cibarium. Male: Curiche, Choco Dept., Colombia Female: vSame locality as male

PAGE 363

-351-

PAGE 364

Figure 65 Lubzormjia (P.) eavvevax carrerai — A. Male genitalia, B. Spermathecae, C. Female head. Male: Tres Esquinas, Caqueta Intendencia, Colombia Female: Same locality as male

PAGE 365

353-

PAGE 366

Figure 66 Lutzomyia (P.) carvevai tkula — A. Male genitalia, B. Female head, Panamanian specimen, C. Spermathecae, D. Female cibarium. Male: Curiche, Choco Dept., Colombia Female: Same locality as male (except Fig. 66B)

PAGE 367

-355-

PAGE 368

Figure 67 Lutzonryia (P.) davisi — A. Spermathecae of female from Para State, Brazil. B. Male genitalia, C. Female cibarium. Male: Tres Esquinas, Caqueta Intendencia, Colombia. Female: Same locality as male (except Fig. 67A) Lutzomyia (P.) kirsuta hirsuta — D. Spermathecae, E. Male genitalia, F, Female cibarium. Male: Tres Esquinas, Caqueta Intendencia, Colombia Female: Same locality as male

PAGE 369

-357rj [V3 M J tx .\ i w

PAGE 370

Figure 68 Lutzomyia (P.) guyanensis — A. Male head, B. Male flagellomere II, C. Female head, D. Male flagellomere II, E. Female cibarium and pharynx, F. Male genitalia, G. Body of spermatheca, H. Spermathecae, I. Female wing, J. Male wing, K. Female cibarium. Male: Almirante, Bocas del Toro Prov. , Panana Female: Curiche, Choco Dept., Colombia

PAGE 371

rft«S\ f\ Gj

PAGE 372

Figure 69 Lutzomyia (P.) noctieola — A. Male head, B. Male flagellomere II, C. Female head, D. Female flagellomere II, E. Female cibarium and pharynx, F. Aedeagus and paramere, G. Male genitalia, H. Spermathecae, I, Proximal end of genital pump, J. Genital pump and filaments, K. Female wing, L. Male wing, M. Female cibarium (all figures from Young, 1973). Male: Rio Anori, Antioquia Dept., Colombia Female: Same locality as male

PAGE 373

-361-^ it yftvs 7> ii'l B W » u

PAGE 374

Figure 70 Lutzormjia (P.) panamensis — A. Male head, B. Male flagellomere II. C. Female head, D. Female flagellomere II, E. Female clbarium and pharynx, F. Genital filaments, G. Male Genitalia, H. Spermathecae, I. Female wing, J. Male wing, K. Female cibarium. Male: Curiche, Choco Dept., Colombia Female: Teresita, Choco Dept., Colombia

PAGE 375

-363-

PAGE 376

Figure 71 Lutzowjia (P.) recurva — A. Male head, B. Male flagellomere II, C. Female head, D. Spermathecae, E. Genital pump and filaments, F. Female cibarium, G. Aedeagus and paramere, H. Female wing, I. Male •ring, J, Male genitalia. Male: Curiche, Choco Dept., Colombia Female: Same locality as male

PAGE 377

-365-

PAGE 378

-366Vexa tor Group The od or 1965 Defined and discussed by Fairchild and Hertig (1957) and Theodor (.1965), this group is divided into 2 series — the series vexa£or(/i+ species, Central America and Nearctic Region) and the periiensis (14+ species, Central and South America). Most of the described species in the latter series occur in mountainous areas but a few including L. hartmanni, L. sanguinaria, and an undescribed Amazonian species from Ecuador inhabit lowland forests where they are strongly anthropophilic. Forattini (1971a, 1973) places hcrtmanni and sanguinaria in the subgenus Triahophoromyia Barretto, gsnus Psychodopygus Mang., but this does not seem to reflect a natural arrangement. The females, unlike those of the Triahophoromyia spp. (in the sense of Sherlock and Guitton, 1970) have only 4 horizontal teeth in the cibarium and the males have 5 major spines on each style. The length of palp 5, variable in the vexator group spp., is relatively short in the Triahophoromyia spp. The single known fossil phlebotomine in the New World, L. paterna (Quate, 1963), was discovered in Mexican amber (late Oligocene or early Miocene). It, a male, differs very little from some extant species in the vexator' group (Quate, op. cit.). For the most part, the females in this group are keyed on the basis of external characters owing to similar cibaria and spermathecae, the latter being difficult to observe in permanent mounts. Not being completely satisfied with this key, I recommend that associated males be on hand before confirming specific identifica-

PAGE 379

367Keys to Species Kale s 1. Coxite with 20+ setae forming a tuft or loosely arranged patch. . . 2 Coxite with fewer than 5 setae at base .4 2. Coxite setae loosely arranged. Genital filaments ca. 5X length of pump. Palpal segment 5 shorter than third. Flagellomere I shorter than or = to head height airrita (Fig. 72) Coxite setae forming a compact tuft on a subcircular base. Genital filaments shorter than 4X length of pump. Palpal segment 5 longer than third. Flagellomere I longer than head height 3 3. Palpal segment 5 shorter than segments 2-1-3. Paramere slender. Coxite tuft on a smaller subcircular base, its diameter less than greatest width of cercus striativilla (Fig. 76) Palpal segment 5 longer than segments 2+3. Paramere broader. Coxite tuft on a larger subcircular base, its diameter = to greatest width of cercus osomoi (Fig. 74) 4. Aedeagus and genital filaments with blunt tips . sanguinaria (Fig. 75) Aedeagus and genital filaments with pointed tips 5 5. Coxite tuft of 2-3 setae, one markedly thickened, over twice length of others and much wider at base. Style with all spines on apical half of structure soorzai (Fig. 77) Coxite tuft of 3-5 setae, none markedly thickened as above. Style with basal spine usually on proximal half of structure 6

PAGE 380

-3686. Coxite tuft of 3-5 setae, one or more distinctly longer than others. Eyes separated by distance = to 5 or more facet diameters. Lab rum or palp 5 shorter than 0.28 mm hartmanni (Fig. 73) Coxite tuft of 3 setae subequal in length. Eyes closer together, separated by less than 4.5 facet diameters. Labrum or palp 5 longer than 0.30 mm sp. of Pichinde (Fig. 78) Females 1. Spermathecae carrot-shaped and large, each with a relatively broad terminal knob. Basitarsus of hind leg shorter than or = to length of hind femur • sanguinavia (Fig. 75) Spermathecae slender and smaller, each with a narrow, more protruding terminal knob. Basitarsus of hind leg longer than hind femur. . . 2 2. Labrum (< 0.43 mm long) shorter than head width. Combined length of palpal segments 1+2+3 shorter than 0.50 mm 3 Labrum (> 0.43 mm long) longer than head width. Combined length of palpal segments 1+2+3 over 0.50 mm 4 3. Flagellomere I (> 0.40 mm long) much longer than labrum. Palpal segment 5 (> 0.25 mm long) = to or longer than segments 3+4 stvictivi-lla (Fig. 76) Flagellomere I (< 0.40 mm long) shorter, = to or but slightly longer than labrum. Palpal segment 5 (< 0.25 mm long) shorter than segments 3 + 4 havtmanni (Fig. 73)

PAGE 381

-3694. Flagellomere I longer than labrum osornci (Fig. 74) Flageliomere I shorter than labrum 5 5. Genital fork stem markedly enlarged at tip .... cirrita (Fig. 72) Genital fork stem subequal in width throughout sp. of Pichinde (Fig. 78) scorzai* (Fig. 77) 88. Lutzonvjia cirrita. (Fig/ 72) Lutzomyia cirrita Young & Porter, 1974, 321 (cf holotype, o, Rio Anori, Antioquia Dept,, Colombia). Distribution: Colombia (Antioquia, Valle) . Mats-rial examined: Colombia. 2 o*c?, 9 o (including holotype and allotype), type locality, other specifics by Young & Porter (1974). 1 <7, Anchicaya Dam (Valle), flight trap, 10 June 1975, R.C.W. Discussion: Other than biting man and being attracted to light traps (Young and Porter, op. cit.), this species is little known. The male, with the scattered coxite setae, is readily distinguished from other Colombian males in the vexator group. ''According to Ortiz (1965b), the length of palp 5 in Venezuelan scorzai females ranges from 0.28-0.33 mm. In the Colombian females it is generally shorter, ranging from 0.22-0.29 mm, average length 0.26 mm, n = 9. The length of this segment in Lutzomyia sp. of Pichinde ranges from 0.310.36 mm, average length 0.35 m, n = 9 so that in Colombia, at least, this difference may be helpful in separating the females of scorzai and Lutzomyia sp. of Pichinde, both of which occur together in Valle Dept., near Cali.

PAGE 382

-37089. Lutzomyia havtmanni (Fig. 73) Fhlehotomus havtmanni Fairchild & Hertig, 1957: 328 (6* holotype, o., Cerro Campaaa, Panama Prov., Panama). Johnson & Hertig, 1961: 765 et seq. (rearing data). Ortiz & Alvarez, 1963b: 321 (listed). McCcnnell &. Correa, 1964: 526 (infected with gregarines) . Ortiz, 1965b: 25 et seq. (listed, cf. to soovzai , measurements). Hanson, 1968: 63 (larva cf. to sariguinaria , figs.). Lutzomyia hartmanni: Barretto, 1962: 96 (listed). Barreto, 1969: 464 (Valle, Colombia). Christensen & Fairchild, 1971: 302 (Darien, Panama). Osorno et al., 1972a: 31 (Colombian records). Christensen, 1972a: 88 (listed). Young, 1973: 109 (Choco, Colombia). Young & Porter, 1974: 323 (cf. to civvita) . Psychodopygus havtmanni: . Forattini, 1971a: 105 (listed). Forattini, 1973: 123 et seq. (gen. review, figs.). Distribution: Panama, Colombia (Antioquia, Choco, Valle) , Ecuador. Material examined: Colombia. 16 6*6", 8 oo, Rio Anori (Antioquia), light traps, May 1970, C.H.P. 1 cf, 8 oo , same data but Sept. 1970, D.G.Y. 8 cW, 1,066 yy, Curiche (Choco), biting man, tree trunks, light, Malaise & Shannon traps, March-Dec. 1967, D.G.Y. 1 cf, 114 c^o. , same data but Alto Curiche (Choco), June-Nov. 1967. 2 cfcf, 13 oo., Teresita (Choco), biting man, tree trunks, Shannon traps, May-Oct. 1967, D.G.Y. 1 cf, Rio Raposo (Valle), light trap. y 4 May 1964, P. Barreto. 3 oV, 6 o^, Anchicaya Dam (Valle), light, flight, traps, tree trunks, 9-11 Aug. 1973, D.G.Y. & R.C.W. 8 op, 25 km E of Buenaventura (Valle), light traps, 12 Aug. 1973, D.G.Y. & R.C.W. Ecuador. 2 6'cf, 3 oo , 17 km E of Santa Domingo de los Colorados (Pichincha) , light and flight traps, May 1976, D.G.Y., T. Rogers,

PAGE 383

-371(t. Falrchild. Panama. 1 6" (holotype no. 3997), type locality, light trap, 24 April 1952, F. Blanton. 1 p (allotype no. 2904), type locality, Shannon trap, 10 Feb. 1951, M. Hertig et al. 2 S3, 8 oc_, typa locality; Almiraute (Bocas del Toro) ; Canal Zone; Cerro Azul (Panama), various dates and collectors. Discussion: At Curiche (Choco) , this species represented nearly 20% (733 po) of all man-biting sand flies taken from April to November, 1967 (141 man hours). Only L. panamensis was more abundant in human bait collections, accounting for 46.5% (1751 ) of 3707 total females. According to Hanson (1968) , the larvae of hartmanni and sanguinaria are very close, differing only in the length of certain setae on the thorax and abdomen. The breeding sites of the havtmanni immatures and the diurnal resting sites of the adults remain virtually unknown. At Curiche, we collected 1 &* 2 op in tree buttresses but other resting sites were not sampled thoroughly. 90. Lutzomyia osornoi (Fig. 74) Phlebotomus osornoi Ristorcelli & Van Ty, 1941: 260 (o holotype, Valle de Capuli, Narino Dept., Colombia). Rozeboom, 1947a: 177 (6", dascr . ) Phlebotomus montoyai Sherlock, 1962: 328 (
PAGE 384

-372Distvibution: Colombia (Narino) . Materi-al examined: Colombia. 1 6", 1 o (USNM) , Chirristes (Narino) 15 Aug. 19 44, J. A. Montoya. Visujssion: See L. striotivilla n. sp. 91. Lutzonvjia sanguinaria (Fig. 75) Phlebotomus sanguinarius Fairchild & Hertig, 1957: 332 (d* holotype, o, Alrairante, Bocas del Toro Prov. , Panama. Fairchild & Hertig, 1959: 122 (Central American records). Johnson & Hertig, 1961: 764 et seq. (rearing data). Johnson et al., 1962: 158 (infected with promastigotes) . Ortiz & Alvarez, 1963b: 312 (listed). Hertig & McConnell, 1963: 93 et seq. (infected with leishmania in laboratory, fig.). McConnell, 1963: 124 et seq. (naturally infected with promastigotes). Shaw, 1964: 418 (feeding on sloths, Panama). McConnell & Correa, 1964: 526-528 (infected with gregarines, fungi, and nematodes). Thatcher & Hertig, 1966: 46 et seq. (hosts). Hanson, 1968: 74 et seq. (larva, pupa, descr., figs.). Thatcher, 1968a, 293 et seq. (hosts). Lutzomyia sanguinaria: Barretto, 1962: 96 (listed). Johnson & Hertig, 1970: 281 et seq. (infected with Leishmania in laboratory). Tesh et al., 1971a: 153 (blood meals). Christensen et al., 19 71: 118 (pyloric spicules). Christensen & Fairchild, 1971: 302 (Darien Prov., Panama). Chaniotis et al., 1971a: 344 et seq. (pop. dynamics, Panama). Chaniotis et al., 1971b: 415 et seq. (biting habits, Panama). Tesh et al., 1972: 90 et seq. (blood meals). Christensen, 1972a: 88 (listed). Chaniotis et al., 1972: 95 (resting sites). Young, 1973: 109 (Choco, Colombia). Christensen & Herrer, 1973: 579 et seq. (collecting data, as

PAGE 385

-373probablc vector of Leishmania) . Chaniotis, 1974b: 501 (keyed). Chaniotis & Correa, 1974: 115 (collecting data). Young L Porter, 1974: 323 (ct. to cirri to) . Ward 6 Killick-Kendric.k, 1974: 214 (mention). Lewis, 1975a: 501 et seq. (mouthpart morphol.). Rutledge & Mosser, 1975: 410 (mention). Christensen & Herrer, 1976: 299 (as host for Endotrypanum schaudinni) . Herrer & Christensen, 1976c: 62 (collecting data, Panama). Herrer & Christensen, 1976b: 57-58 (on man, horse, and sloths). Miles et al., 1976: 532 (mating aggregation, Panama). Zimmerman et al., 1977: 5/5 (egg, oescr., figs.). Pcychodopygvs sanguinarius : Forattini, 1971a: 105 (listed). Forattini, 1973: 123 et seq. (gen. review, refs., figs.). Distribution: Honduras, Costa Rica, Panama, Colombia (Choco, Valle) . Material examined: Colombia. 7 53, 568 cq, biting man, light, Shannon, Malaise traps, tree trunks, Curiche (Choco), April-Oct. 1967, D.G.Y. 1 6", 81 o^, Alto Curiche (Choco), biting man, light traps, JulyNov. 1967. 3 S3, 2 op, Teresita (Choco), light, Malaise traps, tree trunks, March, May-July 1967, D.G.Y. 1 o, Anchicaya Dam (Valle), light trap, 11 Aug. 1973, D.G.Y. & R.C.W. Panama. 4 S3, 9 oo, laboratory reared at GML. Discussion: A highly anthropophilic species which has been incriminated as a vector of leishmaniasis in Panama (Christensen & Herrer, 1973) , L. sanguinaria was the third most common sand fly in human bait collections at Curiche from April to Nov., 1967. Nearly 12% (432 oo) of 3763 females taken during this time belonged to this taxon. The sanguinaria female is easily identified by the characteristic sperraathecae (Fig. 75H) , the structure of which resembles those of the verrucarum group species (Theodor, 1965) . For further information on the

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-374habits and disease relationships, the reader is directed to the above references, especially Forattini (1973) who reviews most of them prior to 1972. 92. Lutzomyia seorzai (Fig. 77) Phlebotomus scovzai Ortiz, 1965b: 28 (o* holotype, q, Rancho Grande, Aragua State, Venezuela). Scorza et al . , 1967: 191 et seq. (listed,
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-375Accordiug to Ortiz (1965b) , the scovzai males have 2-3 (usually 2) basal coxite setae, one of which is markedly enlarged. The Colombian males also have an enlarged seta plus at least 2 other smaller setae at the coxite base. The parameres are generally similar although those of the Colombian males are slightly more clubbed apically and the ventral margin, below the aedeagus, is less acute. Other character states of the males, and seemingly all those of the females, are quite similar except for the longer palp 5 of some of the scovzai females from Venezuela. In Colombia, L. scovzai occurs with Lutzomyia sp. of Pichinde at Pichinde (Valle) and with L. havtmanni _, L. sanguinaviaj and L. civvita at Anchicaya Dam (ca. 400 m above sea level). Both the scovzai male and female , the only specimens available from the latter site, are smaller than any of those from higher localities in Valle and Cauca Depts. (12002180 m) . The wing length of the male = 2.1 mm; that of the female = 2.5 mm as opposed to 2.6-3.0 mm for the others from higher elevations. A similar variation in size as related to altitude is that observed in specimens of B. galindoi discussed earlier. Thus far, L. scovzai has been taken only in light traps. I associated the sexes of L. scovzai on the basis of several features, notably the length of palp 5 which ranges from 0.20 to 0.31 mm in males; 0.22 to 0.29 mm in females (n = 10 for each sex). The interocular distance is rather narrow when compared to that of sympatric species in the vexatov group. Light trap collections near Cali, where a large television tower is situated, yielded scovzai males and females but no others in the vexatov group . The scovzai male, characterized by the enlarged seta of the coxite tuft, is readily distinguished from the males of havtmanni, Lutzomyia sp .

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-376of Pichinde, and others by this character and by others in the key. The females of soorzai , havtmanni, and sp. of Pichinde are difficult to separate, the spermathecae, cibaria, and palpal proportions being of little diagnostic value. I therefore must rely on direct measurements to distinguish them. The labruin of hartmanni females ranges from 0.32 to 0.42 mm in length (n = 43 from various localities in Panama and Colombia). Flagellomere I is shorter than 0.40 mm. Both structures are loager in scorzai females, the labrum ranging from 0.47 to 0.54 mm in length; the first flagellomere 0.44 to 0.50 mm long. The females of scovza'l and sp. of Pichinde are nearly indistinguishable, only the length of palp 5 being helpful in diagnosis (see footnote on p. 369) . A male and female of scorzai from west of Cali at the television tower locality are described as follows. Male: Wing length 2.62; width 0.80. Except for middle part of pleuron which is paler, whole insect well pigmented, dark. Head height 0.40; width 0.39. Eyes separated by 0.12 or by distance = to 6.3 facet diameters. Flagellomere I (0.47 long), 1.2X length of II + III; ascoids simple, short, those on flagellomere II ending near middle, on all flagellomeres except last. Length of palpal segments: 1 (0.05), 2 (0.15), 3 (0.16), 4 (0.08), 5 (0.21); palpal sensilla (ca. 14) on distal twothirds of segment 3. Labrum length 0.29. Cibarium with 15+ dot like vestiges of teeth; chitinous arch indiscernible in middle; pigment patch invisible. Pharynx 0.18 long, unarmed. Pleura with 13 upper and 4 lower episternal setae. Length of wing vein sections: Air ha (0.75), beta (0.27), delta (0.10), garrma (0.29). Length of femora, tibia, and basitarsi:

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-377Foreleg, 0.93, 1.41, 1.02; midleg, 0.89, 1.60, 1.10; hindleg, 1.00, 1.90, 1.20. Abdominal sternite 2 entire, others (3-5 at least) with barely visible, paired circular openings laterally. Cenitalia: Style 0.20 long, with 5 major spines, all beyond middle of structure, no subterminal seta. Coxite (0.32 long x 0.06 wide) with a basal tuft of 3 setae, one much thicker and longer than others, its length exceeding width of coxite. Paramere simple as figured. Aedeagus subtriangular, pointed at tip. Lateral lobe 0.30 long. Genital pump 0.14 long, each filament 0.39 long or nearly 2.8X length of pump. Cercus as shown. Female: Wing length 3.06; width 1.03. Coloration as in 6*. Head height 0.52; width 0.47. Eyes separated by 0.16 or by distance = to 8 facet diameters. Flagellomare I (0.47 long), nearly 1.2X length of II + ill; ascoids as in 6* but slightly longer. Length of palpal segments: 1 (0.07), 2 (0.24), 3 (0.26), 4 (0.09), 5 (0.24); palpal sensilla (22+) as in o. Labrum 0.52 long. Cibarium with 4 sharp, equidistant horizontal teeth, ca . 14 vertical teeth, some larger than others; chitinous arch hardly visible, apparently incomplete; pigment patch faintly pigmented. Pharynx 0.27 long, unarmed. Pleura with 20 upper and 5 lower episternal setae. Length of wing vein sections: Alpha (0.88), beta (0.37), delta (0.18), gamma (0.33). Length of femora, tibiae, and basitarsi: Foreleg, 1.13, 1.50, 1.10; midleg, 1.01, 1.79, 1.16; hindleg, 1.15, 2.11, 1.35. Abdominal sternites as in
PAGE 390

93. Lutzomyia str>ictivilla n. sp. (Fig. 76) Male (holotype) : Wing length 2.10; width 0.67. Whole insect dusky except for darker head, part of mesonotum, procoxae, and abdominal tergites 1-3. Head height 0.42; width 0.37. Eyes separated by 0.12 or by distance = to 6 facet diameters. Flageliomere I (0.50 long), slightly longer than II + III; ascoids simple, short, those on II not extending beyond middle of structure, on all f lagellomeres except last. Length of palpal segments: 1 (0.04), 2 (0.13), 3 (0.17), 4 (0.09), 5 (0.25); palpal sensilla (ca. 20), on distal two-thirds of segment 3. Labrum 0.25 long. Cibarium with ca. 20 vestigial teeth; pigment patch slender, it and chitinous arch nearly invisible, the latter seemingly complete. Pharynx 0.23 long, unarmed. Pleura with 11-12 upper and 5-6 lower episternal setae. Length of wing vein sections: Alpha (0.67), beta (0.17), delta (0.22), gamma (0.40). Length of femora, tibiae, and basitarsi: Foreleg, 1.12, 1.71, 1.20; midleg, 1.0, 1.94, 1.32; hindleg, 1.23, 2.33, 1.47. Abdominal sternites entire, no openings visible. Genitalia: Style 0.24 long, with 5 major spines, basal pair on different levels, not quite paired, no subterminal bristle. Coxite (0.41 long x 0.08 wide), with basal tuft of ca. 25 slender setae inserted on a circular, raspberry-like base. Paramere simple, rather slender as shown. Aedeagus sub triangular, pointed at tip. Lateral lobe 0.33 long. Genital pump 0.12 long, each filament very thin, 0.43 long or nearly 3.6X length of pump. Cerci unremarkable. Female (allotype): Wing length 2.30; width 0.78. Coloration as in male. Head height 0.47; width 0.40. Eyes separated by 0.13 or by distance = to 6.3 facet diameters. Flageliomere I (0.4.1 long), slightly

PAGE 391

-379longer than II + III; ascoids simple, longer than in 6* but not reaching end of flagellomere II, on all but last flagellomere. Length of palpal segments: 1 (0.05), 2 (0.15), 3 (0.19), 4 (0.09), 5 (0.27); palpal sensilla as in d 1 . Labrum 0.32 long. Cibarium with 4 straight, equidistant horizontal teeth, ca, 15 small vertical teeth in an irregular transverse row; pigment patch slender, wider near vertical teeth; chitinous arch complete but hardly visible in middle. Pharynx 0.21 long. Pleura. with .10-11 upper and 3-4 lower episternal setae. Length of wing vein sections: Alpha (0.77), beta (0.16), delta (0.33), gamma (0.37). Length of femora, tibiae, and basitarsi: Foreleg, 0.98, 1.40, 0.95; midleg, 0.88, 1.50, 1.05; hindleg missing. Abdominal sternites as in d*. Tergite 8 with 2-5 setae on each side. Spermathecae and ducts as shown. Cerci unremarkable . Distribution: Colombia (Antioquia) . Material examined: Colombia.
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380(n 7) as opposed to less than 0.21 mm for havtmarmi (n = 16 from various localities including Rio Anori) . Similarly, f lagellomere I is shorter in hai'tnwwii> females, not exceeding 0.37 mm in 18 oo measured; whereas in stria Hvilla females it ranges from 0.41-0.48 mm in length (n = 7) . Of the vexator group species, L. otrietiviVla most closely resembles L. osowoi but that species is very large, the wing being 3.0 mm or longer in both sexes. In contrast to the osovnoi male, L. stvictivilla has relatively slender parameres, the coxite tuft is smaller and consists of fewer .setae. Other than size differences, I am unable to separate the females of stvictivilla and osornoi . The labrum of the latter exceeds 0.47 mm in length; whereas in stvictivilla females, the labrum is less than 0.39 mm long (0.32-0.38 mm, n = 7) . Future studies based on material from other localities may reveal that these taxa are distinct as subspecies or geographic variants but until, or unless, evidence to support this becomes available, 1 will continue to treat them as distinct species. The specific name means "vile woman" and refers to the man-biting habit of the females, Dr. C.H. Porter having collected over 20 females attacking man at Rio Anori in 1970 and 1971 (pers. comra.) . 94. Lutzomyia sp. of P ichinde (Fig. 78) Distribution: Colombia (Valle) . Material examined: Colombia. 6 6V, 2 op, ca. 10 km W of Cali (1750 m) (Valle), tree trunks, rock crevice, 31 July 1973, D.G.Y. & R.C.W. 3 &f, 1 q, same data but 2 Aug. 1973. 1 d", 7 on, same locality

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381but light trap, 2 Aug. 1973. 2 6*6*, ca. 15 km SW of Cali near Rio Pance (1600 m) (Valle), tree trunk, 7 Aug. 1973, D.G.Y. & S. Ayala. Discussion: Owing to the uncertain status of this taxon, I prefer to give it an informal name, hoping that future studies will firmly establish its position as a distinct species or as a geographic variant of L. aavtmanni . Lutzomyia sp. of Pichinde is larger than hartnanni, possibly reflecting altitudinal differences as noted for B. galindoi and L. soovzai. Thus far, specimens of sp. of Pichinde have been taken only near Cali (Valle) above 1200 in. It has not been determined whether or not the females are anthropophilic. The coxite tuft of the male consists of 3 setae, all subequal in size, and palp 5 is always longer than palp 3. Both sexes were collected together in forests on tree trunks, in a rock crevice and once in a light trap. Lutzomyia hartmanni, an anthropophilic species most common in the lowlands of Colombia, has been taken as high as 950 m at Cerro Campana, Panama Prov. , Panama --the type locality. Specimens from there, including the holotype and allotype, are larger than those from the lowlands but do not differ in structure. One seta of the coxite tuft is larger than the others. Palp 5 is variable, being less than, equal to, or but slightly longer than palp 3. This species is rarely found resting on tree trunks. A description of Lutzomyia sp. of Pichinde is given below. Both the male and female were taken together on a tree trunk near Pichinde on 31 July 1973. Male (no. 593): Wing length 3.09; width 0.98. Head, mesonotum and genitalia well pigmented, contrasting with rest of insect which is faintly pigmented. Head height 0.49; width 0.44. Eyes narrowly separated,

PAGE 394

-382interocuiar distance = 0.76 or 3.9 facet diameters. Flagellomere I (0.50 long), 1.35X length of IT + III; ascoids simple, tips extending beyond middle of II, on all f lagelloraeres except last. Length of palpal segments; 1 (0.06), 2 (0.18), 3 (0.22), 4 (0.11), 5 (0.35); palpal sensilla (ca. 15) on distal two-thirds of segment 3. Labrum length 0.35. Cibarium with ca. 20 small remnants of teeth. Pigment patch slender, hardly visible; chitinous arch indefinite. Pharynx 0.24 long, unarmed. Pleura with 15 upper and 6 lower episterna] setae. Length of wing vein sections: Alpha (0.84), beta (0.35), delta (0.26), gamma (0.38). Length of femora, tibia, and basitarsi of slide no. 600: Foreleg, 1.07, 1.83, 1.24: midleg, 1.02, 1.93, 1.34; hindleg, 1.20, 2.30, 1.57. Abdominal sternite 2 not visible (other cfo* have paired lateral openings). Genitalia: Style 0.20 long, with 5 major spines, the most basal on proximal one-half of structure, no subterminal seta. Coxite (0.36 x 0.08 wide) with 3 monomorphic, thin setae at base, none longer than greatest width of coxite. Aedeagus subtriangular, pointed at tip. Lateral lobe 0.31 long. Genital pump 0.15 long, each filament 0.51 long or 3.4X length of pump. Cercus as shown. Female (no. 599): Wing length 3.29; width 1.1. Coloration as in 3. Head height 0.60; width 0.51. Eyes separated by 0.13 or by distance = to 6.2 facet diameters. Flagellomere I (0.53 long), 1.32X length of II + III; ascoids as in o but slightly longer as shown. Length of palpal segments: 1 (0.07), 2 (0.28), 3 (0.31), 4 (0.13), 5 (0.35); palpal sensilla as in
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-383epiccernal setae. Length of wing vein sections: Alpha (0.98), beta (0.32). delta (0,32), gamma (0.40). Length of femora, tibiae, and basitarsi: Foreleg, 1.24, 1.91, 1.27; midleg, 1.15, 2.11, 1.40; hlndleg, .1.35, 2.45, 1.40. Abdominal sternite 2 entire (other on show barely visible paired lateral openings) . Tergite 8 with 1 seta on each side (other 92 have 4+ setae on each side). Spermathecae and ducts as shown. Cere i normal.

PAGE 396

Figure 72 Lutzomyia eiwita — A. Male head, B. Female wing, C. Male wing, D. Female head, E. Female cibarium and pharynx, F. Male genitalia, G. Male flagellcrnere II, H. Female flagellomere II, I. Aedeagus and paramere, J. Genital pump and filaments, K. Spermathecae. Male: Rio Anori, Antioquia Dept . , Colombia Female: Same locality as male

PAGE 397

385-

PAGE 398

Figure /3 Lutzomyia kartmanni — A. Male head, B. Male flagellomere II, C. Female bead, D. Female flagellomere II, E. Female cibarium and pharynx, F. Genital pump and filaments, same scale as Fig. 73G, G. Male genitalia, H. Spermathecae, I. Female wing, J. Male wing, K. Female cibarium. Male: Valle Dept., Colombia Female: Curiche, Choco Dept., Colombia

PAGE 399

-387-

PAGE 400

Figure 74 Lutzomyia osoimoi — A. Male head, B. Male flagellomere II, C. Female head, D. Female flagellomere II, E. Female cibarium and pharynx, F. Genital pump and filaments, G. Spermathecae, H. Body of spermatheca, I. Male genitalia, J. Genital pump, K. Aedeagus and paramere, L. Female wing, M. Male wing. Male: Chirristis, Nariho Dept., Colombia Female: Same locality as male

PAGE 401

-369-

PAGE 402

Figure 75 Lutzomyi-a sanguinaria — A. Male head, B. Male flagellomere II, C. Female head, D. Female flagellomere II, E. Female cibarium and pharynx, F. Genital pump and filaments, same scale as Fig, 75G, G. Male genitalia, H. Spermathecae, I. Female wing, J. Male wing, K. Female cibarium. Male: Panama Canal Zone Female: Curiche, Choco Dept., Colombia

PAGE 403

-391-

PAGE 404

Figure 76 Lutzomyia striotivilla — A. Male head, B. Female wing, C. Male wing, D. Female head, E. Male genitalia, F. Female cibarium, G. Male flagellomere II, H. Female flagellocnere II, I. Spermatheca. Male: Rio Anori, Antioquia Dept., Colombia Female: Same locality as male

PAGE 405

393

PAGE 406

Figure 77 Lutzomyia scorzai -A. Female head, B. Female flagellomere II, C. Genital pump, D. Female cibarium, E. Male genitalia, F. Male wing, G. Female wing, H. Male wing, I. Spermathecae. Male: East of Salidito, Valle Dept., Colombia Female: Same locality as male

PAGE 407

39.

PAGE 408

Figure 78 Lutzomyia sp. of Pichinde — A. Female head, B. Female flagellomere II. C. Genital pump, D. Female cibarium, E. Male flagellomere II, F. Male genitalia, G. Female wing, H. Male wing, I. Spermathecae. Male: Pichinde, Valle Dept., Colombia Female: Same locality as male

PAGE 409

s~\

PAGE 410

Cayennensi s Group Theodor, 1965 The cayennsnsis group with over 17 species and subspecies, depending on author, corresponds to the subgenus Micropygomyia Barretto, 1S62. The species are divided into 3 subgroups — the series co.yennensis , chiapanensis, and atroalavaba (Lewis et al . , 1978). There is circumstantial and direct evidence indicating that some (and probably most) of these species feed on cold-blooded vertebrates (Fairchild, 1955; Williams et al., 1965; Lewis, 1975a). Four species in the cayennensis group are presently known to occur in Colombia. Keys to Species Kales 1. Coxxte tfith a tuft or group of 4+ setae 2 Coxite without nondeciduous setae 3 2. Coxite tuft of 4 strong setae L. atroelavata (Fig. 81) Coxite tuft of 7-11 rather slender, straight setae venezuelensis (Fig. 82) 3Profemur shorter than protibia. Cibarium with a comb-like row of vestigial horizontal teeth, similar to Fig. 79J. Pleura pale. Wing broader cayennensis (Fig. 79) Profemur longer than protibia. Cibarium without horizontal teeth in a comb-like row. Pleura dark. Wing very slender micropyga (Fig. 80)

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-399F emales i. Cibarium with a comb-like row of 12+ horizontal teeth cayennensis (Fig. 79) Cibarium with fewer than 6 horizontal teeth, not in a comb-like row ......... 2 2. Profemur longer than protibia. Pharynx shorter with inconspicuous posterior spines. Pleura as dark as mesonotum. Cibarium with protuberance between inner pair of horizontal teeth miavopyga (Fig. 80) Profemur shorter than protibia. Pharynx longer and wider with prominent spines and transverse ridges. Pleura obviously paler than mesonotum. Cibarium without a protuberance between inner pair of horizontal teeth. . 3 3. Cibarium with very small horizontal teeth. Pigment patch rather broad atroolavata (Fig. 81) Cibarium with much larger horizontal teeth. Pigment patch more slender venezuslensis (Fig. 82) Series cayennensis 95. Lutzornyia cayennensis (Fig. 79) Phlebotomus cayennensis Floch & Abonnenc, 1941a: 13 (cTholotype, Montjoly, Cayenne, French Guiana). Barretto, 1947: 192 (refs.). Fairchild & Hertig, 1948b: 460 et seq . (
PAGE 412

-400Panaroa). Floch & Abonnenc, 1952: 37, 46 (6\ o., keyed), 138-142 (6\ £, redescr., figs.). Pifano & Ortiz, 1952: 143, 148 (<5\ o., keyed, Venezuela). Rosabal, 1954: 10 et seq. (d*, redescr., figs., Costa Rica). Fairchiid & Hertig, 1959: 121-123 (distrib.). Arzube, 1960: 155 (Ecuador). Johnson & Hertig, 1961: 773 (rearing data). Pifano et al., 1962: 385,388 (6", y, keyed), 400-401 (d*, ^, redescr., figs., refs.). Rosabal & Trejos, 1964: 167 (El Salvador). Williams et al., 1965: 70 (reptile blood in gut). Ortiz, 1965a: 205 et seq. (g, keyed, cf. to yencanensis) . Osorno et al., 1967: 28 (Colombian record). Hanson, 1968: 53-55 (larva, pupa, descr., figs.). Morales et al . , 1969a: 378 (Colombian record). Henriquez et al., 1970: 768 (Venezuelan record). Lutzomyia eayennensis: Barretto, 1962: 95 (as type species of Micropygomyia) . Martins et al., 1962a389 (cf. to acantho pharynx) . Theodor, 1965: 186 (6*, y, figs.). Lewis, 1967a: 76 et seq. (classif . , o keyed, refs.). Aitken et al., 1968: 264 (Trinidad). Williams, 1970: 332 et seq. (summary of collecting data, Belize). Christensen et al., 1971: 118 (mention). Christensen, 1972a: 88 (listed). Christensen et al., 19 72: 57 (collecting data, Panama). Osorno et al . , 1972a: 38 (Colombian records). Forattini, 1973: 122 et seq. (gen. review, figs.). Christensen & Herrer, 1973: 579 (Panamanian record) . Lewis, 1975a: 502 et seq. (mouthpart morphol., figs.). Distribution: Mexico, Central America, Colombia. (Antioquia, Bolivar, Caldas, Cesar, Cundinamarca, Guajira, Huila, Magdalena, Norte de Santander, Santander, Tolima) , Ecuador, Venezuela, Trinidad, West Indies, French Guiana. Material examined: Colombia. 2 dcf, Caucasia (Antioquia), light trap, 15 Feb. 19 72, J. Longridge. 1 cf, 2 oo., ca. 10 km SE of Santa Marta

PAGE 413

-401(Magdalena) , tree trunks, 17 Aug. 1973, D.G.Y. & R.C.W. Costa Rica. 4 ocf, Barranca (Puntarenas) , tree buttresses, 17 Dec. 1951, R. Rosabal. Panama. 19 od*, 30 oo, various localities in the Canal Zone & Panama Prov. Trinidad. 7 do*, 2 05, Bush Bush Forest, Nariva Swamp, 1961-1962, T.H.G. Aitken. Venezuela. 1 cf, 1 o, Mene Mauroa (Falcon), in house, I.Ortiz. 1 o, near Calabozo (Guarico) , outside wall of house, July 1965, D.G.Y. Discussion: The cayennensis specimens from near Santa Marta agree with the descriptions of Floch and Abonnenc (1941a, 1952) and thus, I believe, represent L. cayennensis co.yenner.sis y the nominate subspecies. The other cayennensis subspecies, now 6 in number, were discussed and keyed by Lewis (1967a). The status of some of these, as noted by him, can be established only when additional material become available. It may be noteworthy here to report the presence of L. cayennensis in Cuba. The single female, captured in a light trap at Guantanamo Bay, U.S. Naval base, Jan. 1970, J.E. Tisdale, is slightly damaged but it resembles L. c. jamaicensis (Fairchild & Trapido) in details of the cibarial armature and spermathecae. The cayennensis female is easily distinguished from other Lutzomyia females in Colombia by the comb-like row of horizontal teeth and armed pharynx. 96 . Lutzomyia micvovygc (Fig. 80) Phlebotomus micropygus Mangabeira, 1942a: 132 (cf, Aura, Belem, Para State, Brazil). Damasceno et al., 1949: 829 (Para, Brazil record). Barretto, 1950a: 107 (6* keyed) . Floch & Abonnenc, 1952: 37 (6" keyed) .

PAGE 414

-402Scorza & Ortiz, 1960: 434 et seq. (ecologic study). Pifano et al., 1962: 385 (6* keyed) , 402-403 (6", o, descr., figs.). Scorza et al., .1963: 441 (mention). Ortiz & Scorza, 1963: 342 (collecting data, Venezuela). Ortiz, 1965a: 209 (o. keyed) . Scorza et al., 1967: 190 et seq. (distrib., cf, o, keyed, collecting data). Lucena, 1967: 274276 (a* redescr. , figs., Peru). Leon, 1968: 30 (listed). Morales et al., 1968a: 380-381 (Colombian records). Lutzomyia micropyga: Barretto, 1962: 96 (listed). Martins et al., 1963: 334 (Roraima, Brazil). Martins et al . , 1965: 3 (Rondonia, Brazil). Aitken et al . , 1968: 264 (Trinidad). Thatcher, 1968b: 11421143 (arboreal breeding sites, Panama). Tesh et al . , 1971a: 153 (blood meals). Chaniotis et al . , 1971a: 344 et seq. (pop. dynamics, Panama). Christensen et al., 1972: 57 (collecting data). Chaniotis et al., 1972: 95 (resting sites, Panama). Christensen, 1972a: 88 (listed). Llanos, 1973: 34 (6*, £, redescr., 6* figs.). Christensen & Herrer, 1973: 579 (collecting data). Forattini, 1973: 139 et seq. (in part, figs., most information pertains to L. sehreiberi , distrib.). Martins et al . , 1975: 766 et seq. (refs., cf. to sehreiberi, distrib.). Llanos, 1975b: 670 (Peru). Herrer & Christensen, 1976b: 62 (collecting data). Llanos et al., 1976: 480 et seq. (p, descr., figs., refs., Peru). Martins et al., 19 76b: 496 (Peru). Lainson et al . , 1977 (Mato Grosso, Brazil). Distribution: Panama., Colombia (Bolivar, Boyaca, Caldas, Choco, Cundinamarca, Guajira, Magdalena, Meta, Norte de Santander, Santander, Tolima), Ecuador, Peru, Brazil, Venezuela, Trir.id.aj5.. Material examined'. Colombia. 16", 1 ^, Curiche (Choco), tree trunk, 5 May 1967, D.G.Y. ] £, Teresita (Choco), tree trunk, 23 June 1967, D.O.Y. 1 6*, Rio Don Diego E of Santa Marta (Magdalena), tree trunk, 15

PAGE 415

-403Aug. 1973, D.G.Y. & R.C.W. 7 cTa", 6 00, same data but ca. 10 km SE of Santa Marta, 17 Aug. 1973. Brazil. 3 cW, 5 yg, Labrea (Amazonas) , tree trunks, 10 Oct. 1972, D.G.Y. 1 cf, Rio Aripuana at Humboldt (Mato Grosso) , tree trunk, 17 Aug. 1974, D.G.Y. Ecuador. 1 6*, Rio Napo at Limoncocha (Napo) , tree trunk, 23 May 1976, D.G.Y. L T. Rogers. Panama. 12 6Y?, 2 09, localities in Panama and Colon Provinces & Canal Zone. Trinidad. 3 o'd 1 , 4 QQ, Bush Bush Forest, Nariva Swamp 1962-1964, T.H.G. Aitken. Venezuela. 2 go, Agua Santa (Trujillo), tree trunk, 10 Sept. 1965, D.G.Y. Discussion: Several references to "L. micropyga" including those bySherlock £< Carnheiro (1962, 1964), Carnheiro & Sherlock (1964), and Sherlock & Pessoa (1966) refer not to that species but, instead, to L. sckreiberi — a closely related taxon from Brazil (Martins et al., 1975) . The males of both species have remarkably small genitalia but are distinguished by the position of the apical spines on the style (1 terminal spine in schreiberi, 2 terminal spines in micropyga) ; by the pigmentation of the pleura (pale in sc'nreiberi, dark in micropyga) and by the fact that the profemora of both sexes of micropyga, unlike those of schreiberi, are shorter than the protibiae. This latter feature appears to be unique among the Lutzonvyia species as pointed out by Martins et al. (1975) and Llanos et al. (1976). Both L. minasensis (Mang.) from Brazil and L. quadrispinosa (Floch & Chassignet) from French Guiana are treated as junior synonyms of L. micropyga by Forattini (1973). Unlike the micropyga male, however, that of minasensis has pale pleura, the genitalia are larger, the style has but 1 terminal spine, and the profemur is longer than the protibia. 1 examined 3 6o of minasensis from Minas Gerais, Brazil, which were identified by Prof. A.V.

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-404 Martins and which agree with Mangabeira's original description of the species . According to Martins et al. (1970), the minasensis feraala is nearly indistinguishable from that of L. otiveiro.i Martins, da Silva & Falcao, (1970), The individual sperm ducts markedly widen towards the junction with the common duct unlike those of micropyga which are subequal in width throughout (Fig. 80G) . The pharynx of the micropyga female is armed with short spines (Fig. 80E) unlike minasensis in which they are lacking. I should point out that Forattini's concept (1973) of L. micropyga was based partly upon the studies of Sherlock & Carnherio (1964, 1966) buL, as noted above, they were dealing with L. schreiberi , not L. mtCTOpygc (Mang.). It is thus understandable why Forattini (op. cit.) considered it and minasensis to be conspecific, the males of each having 1 terminal spine on the style and with pale pleura. The larger size of the minasensis genitalia and differences in the females (similar to those between micropyga and minasensis) indicate, however, that schreiberi is distinct from minasensis . Based solely on the descriptions of L. quadri spinas a by Floch & Chassignet (1947b) and Floch & Abonnenc (1952), I am not convinced that it and micropyga are conspecific. Only the male of the former taxon is known. It has very small genitalia, like micropyga, but the parameres are more slender apically. I have not observed such slender parameres in any of the micropyga males examined. The length of the profemur and protibia was not mentioned in the descriptions of quadrispinosa . It seems, then, that additional material of this taxon is needed from French Guiana to firmly establish its status.

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-405The female of micvopyga was described by Pifano et al. (1962) and Llanos et al. (1976), tbe latter authors noting that the cibarium generally lias 4 or 5 horizontal teeth but illustrating a specimen which has 7. Ail of the females examined by me from Colombia and elsewhere have 4 distinct teeth (Fig. 80J) . They agree with the descriptions cited above in all other character states. Flagellates of uncertain identity were recovered from a micvopyga female in Panama (Christensen et al . , 1972). These parasites were found posterior to the midgut, similar in position to those observed in L. schveiber-i females by Sherlock & Pessoa (1966, as micvopyga) . According to Tesh et al. (1971a), L. micropyga in Panama feeds principally on coldblooded vertebrates. Series atvoclavata 97. Lutzomyia atvoclavata (Fig. 81) Phlebotomus atvoclavatus Knab, 1913: 135 (6*, o., Gasparee Island, Trinidad). Barretto, 1947: 186-187 (full refs., synonyms). DampE, 1947 296-305 (refs., a" redescr., full refs., figs.). Fairchild & Hertig, 1948b: 455 et seq. (6*, o, redescr., figs., refs., synonyms). Fairchild & Trapido, 1950: 405, 409 (<5\ o., keyed). Barretto, 1951: 212 (distr.) . Floch & Abonnenc, 1952: 34, 47 (6*, o., keyed), 107-112 (6*, o, redescr., figs.). Scorza & Ortiz, 1960: 434 et seq. (biology). Pifano et al., 1962: 387, 389 (6*, g, keyed), 398-400 (6", ^, redescr., figs., synonyms, refs.). Fauran et al., 1966: 904-908 (cf, redescr., figs. Martinique). Osorno ct al., 1967: 28 (Colombian record). Henriquez et al., 1970: 768 (Venezuelan record) .

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-40G'Phlebotonrus tejerae Larrousse, 1922: 71, 73 (6*, o, Zulia State, Venezuela). Dyar, 192.9: 120 (as synonym of atroclavatus) . Barret co, 1947: 137 (rets.) . Pkelbotomus guadeloupensis F.loch & Abotmenc, 1945a: 1 (6", o, Guadeloupe). Dampf, 1947: 302 (as synonym of atrocZavatiAs) • Fairchild & He-tig, 1948b: 456-460 (as synonym of atroelavatus) . Courmes et al . , .1966: 217 et seq. (as possible vector of kala-azar in Guadeloupe). Lutzomyi-a atroalavata: Theodor, 1965: 195 (g, figs.). Forattini, 1971a: 101 (listed). Christensen, 1972a: 88 (listed). Osorno et al., 1972a: 63-65 (Colombian records). Forattini, 1973: 211 et seq. (in part, gen. review, figs.). Distribution: Panama:, Colombia (Boyaca, Caldas, Cundinamarca, ?Guajira, Huila, Magdalena, ?Meta, Norte de Santander, Santander, Tolima) , Venezuela, Trinidad, and other islands in the West Indies. Material examined: Colombia.. 2 oo, SE of Santa Marta near Minca (Magdalena), tree trunk, 17 Aug. 1973, D.G.Y. & R.C.W. Panama, Trinidad, Venezuela, and West Indies (St. Croix, Virgin Islands, Granada, St. Lucia). 51 cf6' f , 19 oo. Discussion: Although generally having a maritime distribution (Fairchild & Hercig, 1948b), L. atroc.lavota was reported from several inland localities in Colombia by Osorno et al. (1972a). The records based upon males are undoubtedly correct but the females from Guajira and Meta Depts. may represent, instead, those of venezuelensis (zuliaensis) , the males of which were taken with the females in these two Departments. The females are very similar in structural details. At present there is no evidence to support the suggestion by Courmes et al. (1966) that L. atvoatavata may be the vector of kala-azar in Guade loupe, W.I.

PAGE 419

407Specimens are usually found resting in hollow trees, on tree trunk or in dark crevices. The feeding habits remain virtually unknown. 98. Lutzomyia venezuelensis (Fig. 82) Phlebotomus VenszuelensJ S Floch & Abonnenc, 1948b: 1 (6* hoiotype, Selva de Tamanaco, Zulia State, Venezuela). Pifano & Ortiz, 1952: 145 (distrib.). Scorza & Ortiz, 1960: 434 et seq. (ecologic study). Pifano et al., 1962: 387, 389 (
PAGE 420

-408I'azevia.l examined'. Colombia. 2 09, Caucasia (Antioquia) , light trap, 29 April 1972, J. Longridge. 1 6", Minca, SE of Santa Marta (Magdalena), tree trunk, 17 Aug. 1973, D.G.Y. & R.C.W. 6 33, same data but 10 km SE of Santa Marta. D-iscussion: Lutzornyia zutiaensis (Floch & Abonnenc) from Venezuela is probably the female of L. venezuelensis (Floch & Abonnenc) as stated by Pifano et al. (1962). This association is based upon collecting data and morphology. The pharynx of the venezuelensis male, although smaller and lacking spines, is generally shaped like that of zutiaensis and atvoclavaia. The males of atvoclavata and venezuelensis are nearly identical in most character states but differ readily by one feature — the number and nature of the coxite setae (cf. Fig. 81F and Fig. 82). The larger horizontal teeth and smaller pigment patch in the cibarium of the venezuelensis (Fig. 82) female contrasts with those observed in that of atroalavata (Fig. 81J) . Forattini (1973) treats L. venezuelensis as a junior synonym of L. oswaldoi (Mang.) but the latter male has 5 major spines on the style and the female pharynx is unarmed.

PAGE 421

Figure 79 LutzomyLa cayennensis — A. Male head, B. Male flagellomere II, C. Female head, D. Female flagellomere II, E. Female cibarium and pharynx, F. Male genitalia, G. Spermathecae of female from Trinidad, H. Female wing, I. Male wing, J. Female cibarium. Male: Trinidad Female* Falcon State, Venezuela (except Fig. 79G)

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-41C

PAGE 423

Figure 80 Lutzomyia mieropyga — A. Female head, B. Female flagellomere II, C. Male, head, D. Male flagellomere II, E. Female cibarium and pharynx. F. Male genitalia, G. Sperraathecae, H. Female wing, I. Male wing, J. Female cibarium. Male: Curiche, Choco Dept., Colorrbia Female: Same locality as male

PAGE 424

(12-

PAGE 425

Figure 81 Lutzomyia atroclavata — A. Male head, B. Male flagellomere II, C. Female head, D. Female flagellomere II, E. Female cibarium and pharynx, F. Male genitalia, G. Spermathecae, H. Female wing, I. Male wing, J. Female cibarium. Male: Trujillo State, Venezuela Female: Same locality as male

PAGE 426

-414*

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Figure 82 Lutzomxjia venezuelensis — A. Female cibarium and pharynx, B. Male genitalia, C. Spermathecae. Male: Minca, Magdalena Dept., Colombia Female: Caucasia, Antioquia Dept., Colombia

PAGE 428

•416V* I

PAGE 429

-417Oswa Idoi Group Theodor, 1965 This group of over 16 described species was defined by Theodor (1965) and by Barre.tto (1962) as group 2 (oswaldoi) of the subgenus Helcoeyvtorryia Barretto, 1962. Three species are known from Colombia. The majority of species occur in South America, especially Brazil, where they are found most commonly on tree trunks and sometimes in light traps. The females, except for L. pia which may not be closely allied with the others, can not be regarded as anthropophilic . A few species such as L. oswaldoi (Mang.), L. rorotaensis, and L. trinidadensis are known to feed on poikilo thermic vertebrates. The latter species, placed with the ungrouped species by Theodor (1965) , is treated here as a member of the oswaldoi group based upon the female, cibarium, male genitalia, and palpi of both sexes. The status of some taxa remains to be clarified. Keys to Species M ales I. Style with 1 terminal spine and a subterminal seta. Palp 5 subequal in length to 3 + 4 pia (Fig. 83) Style with 2 terminal spines* but lacking a subterminal seta. Palp 5 much longer than 3 + 4 2 *As discussed under L. vovotaensis (p. 42.1), some males have only 4 spines on the style; in which case, only 1 is terminal. The subterminal seta is absent.

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2. Head and body markedly infuscated, lover part of pleura with faint pigmentation. Style with 1 terminal spine much smaller than the other rorotaensis (Fig. 84) Head and body faintly infuscated, pleura entirely pale. Style with terminal spines subequal in size trinidadensis (Fig. 85) Females 1. Fharynx armed with conspicuous spines. Spermathecae cylindrical (sausage-shaped) and smooth-walled tvinidadensis (Fig. 85) Pharynx unarmed. Spermathecae otherwise, with complete or incomplete ambulations 2 2. Spermathecae with terminal annulation much smaller than others. Cibarium with 4-6 sharp, nearly straight horizontal teeth. Labrum length subequal to head height pia. (Fig. 83) Spermathecae with large hemispherical head. Cibarium with 4-6 horizontal teeth, inner pair bent inwards. Labrum much shorter than head height vovotaensis (Fig. 84) 99. Lvtzorrvjia pia (Fig. 83) Phlebotorrrus pius Fairchild & Hertig, 1961b: 248 (6* holotype, Santa Clara, El Volcan, Chiriqui Prov., Panama; o, Palo Santo, Chiriqui Prov., Panama). Ortiz & Alvarez, 1963b: 313 (mention). McConnell & Correa, 1964: 528 (parasitized with mites).

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419Lutzomyia pia: Theodor, 1965: 187 (listed). Forattini, 1971a: 102 (listed). Osorno et al., 1972a: 45 (Boyaca, Colombia). Forattini, 1.973: 189 et seq. (gen. review, figs.). Distribution: Costa Rica, Panama, Colombia (Boyaca) . Mate-Hal examined: Colombia. 1 d\ 1 c_ (nos. 1925 and 1929, INPES) , Yareda Umbabita, Municip. Almeida (Boyaca), tree hollow, 4 July 1968, E. Osorno, A. Morales, & J. Ochoa . Costa Rica. 1 p, Moravia (Chirripo) , Shannon trap, 4 March 1966, R. Zeledon. Panama. 1 c? (holotype no. 2428), type locality, tree buttress, 11 Aug. 1950, R. Hartmann. 1 p (allotype no. 2269), Palo Santo (Chiriqui) , tree buttress, 19 May 1950, R. Hartmann. 18 op, Rio Changena (Bocas del Toro) , biting man, 8 Sept. 1961, V. Tipton. 3 def, 97 oo, same data but Shannon trap, 12 Sept. 1961. Discussion: Both sexes of L. pia differ from other oswaldoi group species in several features, the most notable being: 1) The presence of a subterminal seta on the male style. 2) The female cibarium with 4-6 sharp, nearly straight horizontal teeth and a prominent, complete cibarial arch. 3) The man-biting habit of the females. 4) The unusual spermathecae and 5) The relatively short palp 5 of both sexes. The species is placed here chiefly on the basis of the male genitalia. In the original description (Fairchild and Hertig, 1961b) , the authors stated that there are 6 horizontal teeth in the female cibarium. One of the paratypes figured by them shows this number but the majority of females, including the allotype and a Colombian specimen, have only 4 such teeth. Of 103 females examined for this feature, 100 had 4 teeth, the remainder had 5. The Columbian specimens agree closely with the holotype and allotype of L. pia and are undoubted]. y conspecific. The present, apparently

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.20disjunct distribution of this species suggests that populations were continuous in times past from northern Colombia to Costa Rica. 100. Lutzomyia vorotaensis (Fig. 84) Phlebotomies rorotaensis Floch & Abonnenc, 1944d: 4 (6* holotype, o, Rorota, French Guiana). Barretto, 1946c: 431 (d* keyed) . Barretto, 1947: 216 (as synonym of oswaldoi, in part); 221 (refs.). Floch & Abonnenc, 1950b: 9 (cf. to related spp., refs.). Barretto, 1950b: 145 (cf. to zikani) . Barretto, 1951: 223 (distrib.). Vargas & Diaz-Najera, 1952: 72-73 (cf. to durani) . Floch & Abonnenc, 1952: 41, 45 (6", g, keyed), 167-170 (6*, 5, redescr., figs.). Lutzomyia vorotaensis: Martins et al., 1961a: 300 (mention). Barretto, 1961: 1 (as synonym of villelai Mang.). Martins et al . , 1963: 335 (as a distinct species, Roraima, Brazil). Lewis et al . , 1970: 215 (parous rates). Forattini, 1971a: 102 (as a synonym of trinidadensis) . Tesh et al., 1971a: 152 et seq. (reptile and/or amphibian blood meals). Chaniotis et si., 1971a: 344 (pop. dynamics, Panama). Chaniotis et al., 1972: 93 et seq. (tree trunks as resting sites). Christensen, 1972a: 88 (listed). Osorno et al., 1972a: 45 (Colombian records). Forattini, 1973: 312 (as synonym of oswaldoi). Rutledge & Ellenwood, 1975a: 71 et seq. (breeding on open forest floor); 1975b: 82; 1975c: 83 et seq. Lewis, 1975a: 502 et seq. (mouthpart morphol.). Distribution: Panama, Colombia (Antioquia, Choco, Valle) , French Guiana . Material examined: Colombia. 1 c?, 2 90, Rio Anori (Antioquia), tree buttresses, Sept. 1970, D.G.Y. 1 0, Alto Curiche (Choco), tree buttress,

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-4214 April 1967, D.G.Y. 1 6\ 3 oo, Anchicaya Dam (Valle) , tree trunks, 9 Aug. 1973, D.G.Y.. & R.C.W. 14 <3c?, 8 p£, 20 km E of Buenaventura (Valle), tree trunks, 11 Aug. 1973, D.G.Y. & R.C.W. Brazil. 4 $<$ , 2 go, Serra do Navio (Amapa) , tree trunks, 21 Sept. 1972, D.G.Y. & II. Fraiha. Panama. 13 6o, 8 oo, various localities in Panama & Colon Provinces & Canal Zone. Discussion: Lutzomyia vovotaensis males from Valle Dept., Colombia, are unique among the oswaldoi group species in having 4 instead of 5 spines on each style. This reflects intraspecific variation in my opinion, because other features of both sexes from there and elsewhere are nearly identical . Where they occur together in northern Colombia, Panama, and French Guiana, this species and L. tvinidadensis are found often on tree trunks, the latter species almost always more abundant. Females of both species feed on cold blooded vertebrates (Tesh et al . , 1971a). The immatures are known to live on the open forest floor (Rutledge and Ellenwood, 1975a). Lutzomyia oswaldoi (Mang.) a Brazilian species is treated as a senior synonym of L. vovotaensis and L. venezuelensis (Floch and and Abonnenc) by Forattini (1973) . The latter taxon, obviously distinct from the others, is actually conspecific with L. zuliaensis (see p. 408). The differences between oswaldoi and vovotaensis are not as salient, however, and without material of oswaldoi, 1 am unable to judge whether or not they are conspecific. For the present, I prefer to separate them, pointing out that Panamanian and Colombian specimens (except those from Valle Dept,, Colombia) agree closely with a male (no. 2295) and female (no. 2277) from Panama which were compared with the types from French Guiana by Dr. H. Floch in 1952 (Dr. G.B. Fairchild, pers. coram.).

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-422101. Lutzomyia tvinidaderisis (Fig. 85) Phlebotomus ti'inidadensis Newstead, 1922: 4 (6*, o, Trinidad). Fairchiid & Hertig, 1948a: 253 et seq. (full refs., synonyms, 6*, o, redescr., figs.). Hertig & Fairchiid, 1950: 91 et seq. (sternites, figs.). Barretto, 1951: 224 (distrib.). Rosabal, 1954: 25 et seq. (cf, o, measurements, Costa Rica, figs.). Deane & Deane, 1957: 225 et seq. (Ceara, Brazil). Fairchiid & Hertig, 1959: 122-124 (Central American records). Lewis & Garnham, 1959: 83 (Belize, figs.),Hanson, 1961: 320 (larvae in soil between tree buttresses, Panama). Johnson & Hertig, 1961: 765 et seq. (rearing data). Pifano et al., 1962: 384, 388 (6*, p, keyed), 407 et seq. (cf, o, redescr. , refs., figs.). McConnell & Correa, 1964: 523 et seq. (infected in nature with trypanosomes , gregarines, and fungi). Thatcher & Hertig, 1966: 46 et seq. (host preferences, Panama). Osorno et al., 1967: 29 (Colombian records). Hanson, 1968: 84 et seq. (larva, pupa, descr., figs.). Thatcher, 1968a: 296 (in castor oil traps) . Phlebotomus yucatanensis Galliard, 1934: 1 (6", o, near Chichenitza, Yucatan, Mexico). Barretto, 1947: 232 (full refs.). Fairchiid & Hertig, 1948a: 255 (as synonym of trinidadensis) • Mirsa, 1953: 63 et seq. (rearing data) . Phlebotomus yucatanensis var. baduelensis Floch & Abonnenc, 1941b: 4 (cf, Baduel, French Guiana). Fairchiid & Hertig, 1948a: 225 (as synonym of ti'inidadensis) . Phlebotomus longipalpis: Ristorcelli & Van Ty, 1941: 252 et seq. (o, Narino, Colombia, figs.). Not longipalpis Lutz & Neiva, 1912, as noted by Fairchiid & Hertig (1948a).

PAGE 435

423Phlebotomus villelai Mangabeira, 1942a: 196 (6*, Ceara S> Para States. Brazil). Barretto, 1946b: 527 (as synonym of baduelensis, refs.). Fairchild & Hertig, 1948a: 255 (as synonym of trinidadensis) PhZebotomus baduelensis Floch & Abonnenc, 1944d: 1 (
PAGE 436

Lutzorryia villelai: Barretto, 1962: 96 (listed). Martins et al . , 1962a: 393 (listed). Martins & Morales, 1972: 367 (listed). Brumptorryia tr-inidadensis : Lewis, 1965: 376 et seq. (internal structures, figs.) . Lutzorrryia baduelensis : Christensen et al., 1971: 118 (mention). Distribution'Mexico to Brazil, Colombia (Antioquia, Bolivar, Boyaca, Caldas, Cesar, Choco , Cundinamarca, Guajira, Huila, Magdalena, Nari.no, Norte de Santander, Santander, Tolima) . Material examined: Colorrbia. 12 33, 3 on, Rio Anori (Antioquia), light traps & tree trunks, May 19/0, C.H.P. 74 33, 52 oo, same data but Sept.. 1970, D.G.Y. 113 cfd*, 74 go, Curiche (Choco), Malaise, Shannon & light traps, April-Dec. 1967, D.G.Y. 333 33, 301 oo, same data but tree trunks only. 12 6*0", 1 0, Alto Curiche (Choco), tree trunks, April, May, July, Sept. 1967, D.G.Y. 451 33, 206 00, Teresita (Choco), Malaise, Shannon & light traps, tree trunks, MarchDec. 1967, D.G.Y. 3 33, 3 00, Rio Atrato at Sautata (Choco), Oct. 1967, D.G.Y. 3 09, Baraya (Huila), March 1945, M. Hertig. 35 6*0", 33 gg, Rio Don Diego, E of Santa Marta (Magdalena), tree trunks, 15 Aug. 1973, D.G.Y. & R.C.W. 6 33, 2 oo, Minca, SE of Santa Marta (Magdalena), tree trunks, 17 Aug. 1973, D.G.Y. & R.C.W. 43 33, 36 go, 10 km SE of Santa Marta (Magdalena), tree trunks, 1973. Bolivia. 1 o, Teoponte, 400 m above sea level (Larecaja) , tree hole, 12 July 1971, J. Velasco. Costa Rica, French Guiana, Nicaragua, Panama, Trinidad, & Venezuela. 300+ 33, 300+ go, various localities, dates and collectors, stored mostly in alcohol. Discussion: Because of its abundance in lowland forests, L. trinidadensis is a familiar sand fly to specialists working in Central America and northern South America. In Choco Dept., Colombia, it was the dominant

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-425species on tree trunks, accounting for over 50% of all phlebotomina specimens captured. Hanson (1961) recovered a few larvae from soil between tree buttresses. Rutledge and Ellenwood (1975a) secured adults in soil emergence traps on the open forest floor. In neither study were the immatures plentiful. The females feed principally on lizards, especially geckoes, but are also known to feed on or be attracted to mammals (including man) and possibly birds (McConnell and Correa, 1964; Aitken et al., 1968; Williams, 1970, and Rutledge & Mosser, 1975) . This species and L. cruciata (Coq.) were confused for many years but Fairchild and Hertig (1948a) distinguished the two and provided references to the misidentifications of others. The distribution of trinidadensis in Colombia is rather interesting. It is very common in northern Choco but appears to be absent, or at least very rare, in forests along the Pacific coast in Valle Dept . Neither Earreto (1969) nor ourselves noted its presence there. I have not seen specimens referable to trinidadensis from the Amazon basin in Colombia or elsewhere. The status of L. vitlelai, considered by most to be a junior synonym of trinidadensis, is not settled at present. This is also true for L. goiana Martins, Falcao, and da Silva, a species treated as a junior synonym of trinidadensis by Forattini (1973) . The females differ in the size and distribution of the pharyngeal spines, those of goiana being reduced in size and number (Martins et al., 1962a). According to the describers, the males of goiana and trinidadensis are indistinguishable. Lewis (1975a) illustrated the o cibaria of both species.

PAGE 438

Figure 83 Lutzomyia pia — A. Male genitalia, B. Female cibarium, C. Spermathecae of female from Palo Santo, Chiriqui Prov., Panama. Male: Boyaca Dept., Colombia Female: Same locality as male (except Fig. 83C)

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-427u iw

PAGE 440

Figure 84 Lutzomyia rorotaensis — A. Male head, B. Male flagellomere II, C. Female head, D. Female flagellomere II, E. Female cibarium and pharynx, F. Genital pump and filaments, G. Gential pump, H. Male genitalia, I. Spermathecae, J. Female wing, K. Male wing, L. Female wing. Male: Colon Prov., Panama Female: Rio Anori, Antioquia Dept., Colombia

PAGE 441

-429-

PAGE 442

Figure 85 butzomyia trinidadensis — A. Male head, B. Male flagellomere II, C. Female head, D. Female flagellomere II, E. Genital pump and filaments, F. Female cibarium and pharynx, G. Spermathecae, H. Male genitalia, I. Female wing, J. Male wing. Male: Curiche, Choco Uept., Colombia Female: Same locality as male

PAGE 443

-431-

PAGE 444

-432 Pilosa Group The odor, 1965 Characterized by Theodor (1965) , this group of 3 species appears to be distinct although the females resemble those in the oayennensis group in some aspects, notably the cibarial armature. Forattini (1971a, 1973) includes species in both groups in the subgenus Mioropygomyia Barretto, 1962, a category established for L. oayennensis and its allies which is equivalent to the oayennensis group of Theodor (op. cit.). At present, L. pilosa is the sole representative of the pilosa group in Colombia. Lutzomyia chassignebi (Floch and Abonnenc, 1944) occurs in French Guiana and Brazil and L. mangabeivana. (Martins, Falcao and da Silva, J 963) is presently known from northern Brazil. 102. Lutzomyia pilosa (Fig. 86) Phlebotomies pilosus Damasceno & Causey, 1944: 342 (o* holotype, Santa Isabel, Belem, Para, Brazil). Barretto, 1947: 219 (ref.). Damasceno et al., 1949: 830 (Brazilian records). Damasceno & Arouck, 1949: 845 (cf. to serrana) , Fairchild & Trapido, 1950: 410 (mention). Barretto, ; L 950a: 102 (d" keyed) . Barretto, 1951: 222 (distrib.). Ortiz, 1952: 155 (cf. to range liana) . Floch & Abonnenc, 1952: 30 (cf keyed) . Fairchild & Hertig, 1959: 122 (Central American records). Pifano et al., 1962: 385 (a*, in key to Venezuelan sand flies). Lutzomyia pilosa: Theodor, 1965: 194 (cf, genitalia figs.). Aitken et al,, 1968: 264 (Trinidad). Osorno et al . , 1969: 380 (Colombian record). Forattini, 1971a: 101 (listed). Osorno et al., 1972a: 62-53 (Colombian records). Christensen, 1972a: 88 (listed). Forattini, 1973: 209 et seq. (gen review, 6*, figs.).

PAGE 445

Distribution: Costa Rica, Panama., Colombia (Antioquia, Bolivar, Boyaca, Caqueta, Choco, Huila) , Brazil, Venezuela, Trinidad. Material examined: Colombia. 16*, 1 o, Rio Anori (Antioquia), light trap, 3 May 1970, C.H.P. 2 6*o*, 3 09, same locality, three buttresses, Sept. 1970, D.G.Y. 1 o, Puerto Boyaca (Boyaca), light trap, 6 May 1973, C.J.M. 11 (?(?, 14 w, Alto Curiche (Choco), light & Shannon traps, tree trunks, 31 May-2 Aug. 1967, D.G.Y. Costa Rica. 1 9, Wauchope (Limon) , tree buttress, 9 Aug. 1951, P. Galindo & H. Trapiclo. Panama. 3 cjcj>, Almirante (Bocas del Toro) , light traps, Nov. 1952-Jan. 1953, W. Hils. coll. 1 6", Rio Platanar (Panama), hollow tree with bats, 20 July 1950, R. Harmann. Trinidad. 5+ 6V, 1 Q_, Nariva Swamp, Bush Bush Forest, various dates, T.H.G. Aitken. Discussion: Lutzomyia pilosa and L. chassigneti are very similar, the male of the former species having a distal patch of coxite setae, lacking in chassigneti. The pilosa female, known but undescribed, is described here based on 2 specimens from Alto Curiche (Choco) Colombia and 1 from Almirante (Bocas de Toro), Panama. As Theodor (1965) mentioned, the yilosa and chassigneti females are indistinguishable. Female: Wing length 1.64-1.80; width 0.49-0.57. Head, mesonotum, abdominal tergites moderately pigmented, rest of insect dusky, pleura slightly paler than mesonotum. Head height 0.32-0.35; width 0.30-0.32. Eyes separated by 0.11 or by distance = to ca. 5.5 facet diameters. Rlagellomere I (0.25-0.27 long), nearly 1.2 x length of II + III; ascoids sinple, those on flagellomere II reaching to or beyond end, on all flagellomeres except last. Length of palpal segments: 1 (0.03), 2 (0.09-0.11), 3 (0.14-0.16), 4 (0.10), 5 (0.34-0.37); palpal sensilla at

PAGE 446

-434apex of segment 3. Labrum 0.15-0.16 long. Cibarium as figured. Pharynx 0.13-0.14 long, unarmed. Pleura with 5-10 upper and 3-6 lower episternal setae. Length of wing vein sections: Alpha (0.34-0.47), beta (0.29-0.31) delta (0.15-0.25), g aroma (0.18-0.20). Length of femora, tibiae, and basitarsi of slide 4276 (Almirante) : Foreleg, 0.71, 0.71, 0.39; midleg, 0,67, 0.83, 0.46; hindleg, 0.73, 0.98, 0.54. Abdominal sternite 2 entire, without openings, sternite 2, at least, with paired circular openings at sides. Tergite 8 with 5-10 setae on each side. Spermathecae, ducts, and cerci as shown.

PAGE 447

Figure 86 LubzomyLa pilosa — A. Male head, B. Male f lagellomere II, C. Spermathecae, D. Female head, E. Female flagellomere II, F. Male genitalia, G. Genital pump and filaments, H. Tip of genital filament, I. Female cibarium and pharynx, J. Female wing, K. Male wing, L. Female cibarium. Male: Alto Curiche, Choco Dept., Colombia Female: Same locality as male

PAGE 448

-436-

PAGE 449

-437Ungr ouped Speci es 103. Lutzomyia nordestina (Fig. 87) Phlebotomus novdestinus Mangabeira, 1942e: 327 (o holotype, Nova Olinda, Ceara, Brazil). Barretto, 1947: 215 (refs.). Damascene et al . , 1949: 830 (Brazilian records). Barretto, 1951: 221 (distrib.). Floch & Abonnenc, 1952: 37, 45 (o*, o keyed), 145 (d*, 9, redescr., figs.). Rodriguez, 1956: 76 et seq. (Ecuador). Hanson, 1961: 321 (breeding site, Panama). Johnson & Hertig, 1961: 765 et seq. (rearing data. Thatcher & Rertig, 1966: 52 (in burrows, Panama). Pklebotonras longieovnutus Floch & Abonnenc, 1943: 6 (d holotype, p, Montabo, French Guiana). Barretto, 1946b: 534 (as synonym of nordestina). Forattini, 1960: 478 (Amapa, Brazil). Lutzomyia nordestina: Barretto, 1962: 96 (listed). Martins et al. , 1962a: 381 (Goias, Brazil). Theodor, 1965: 195 (characterized). Martins et al., 1965: 3 (Rondonia, Brazil). Martins et al., 1970: 553 (mention). Tesh et al., 1971a: 152 (blood meals). Chaniotis et al., 1971a: 344 et seq. (pop. dynamics, Panama). Chris tensen & Fair child, 1971: 301 (Darien, Panama). Osorno et al., 1972a: 65-66 (Colombian records). Christensen, 1972a: 83 (listed). Christensen et al . , 1972: 57 (collecting data). Chaniotis et al . , 1972: 95 (resting sites). Christensen & Herrer, 1973: 579 (collecting data, Panama). Lewis, 1975a: 504 et seq. (mouthpart morphol . ) . Llanos et al . , 1976: 480 (Peru). Martins et al., 1976b: 496 (Peru).

PAGE 450

-438Psychodopygus noraestina: Forattini, 1971a: 105 (listed). Forattini, 1973: 140 et seq. (gen. review, figs.). Distribution: Panama, Colombia (Amazonas, Antioquia, Boyaca, Caqueta, Choco, Tolima, Valle) , Ecuador, Panama, Peru, French Guiana, Trinidad, Brazil. Material examined: Colombia. 1 o, 17 km W of Leticia (Amazonas), burrow, 26 July 1973, D.G.Y. & R.C.W. 1 o_, Rio Anori (Antioquia), light trap, 3 May 1970, C.H.P. 4 do*, same locality, tree trunks, Sept. 1970, D.G.Y. 1 0, Puerto Boyaca (Boyaca), light trap, 10 Aug. 1971, C.J.M. 6 $6\ 16 00, Curiche (Choco), light, Malaise & Shannon traps, May-Oct. 1967, D.G.Y. 1 6*, 7 OO, Alto Curiche (Choco), light & Shannon traps, tree trunk, June-Sept. 1967, D.G.Y. 1 6*, Anchicaya Dam (Valle), tree trunk, 9 Aug. 1973, D.G.Y. & R.C.W. 2 6*0*, 1 o., 25 km E of Buenaventura (Valle), light & flight traps, 12 Aug. 1973, D.G.Y. & R.C.W. Brazil. 2 do, 5 00, 27 km SE of Maraba (Para), light traps, 26-28 Sept., 1972, D.G.Y. & H. Fraiha. 5 cfcf, 1 o, same locality, burrow in leaf-cutting ant nest, 26 Sept. 1972, D.G.Y. 1 6*, near Altamira (Para), tree trunk, 1 Oct. 1972, D.G.Y. 1 o_, Belem (Para), flight trap, 2 Aug. 1974, D.G.Y. 1 o, near Bacuri, N of Maraba (Para), light trap, 28 Oct. 1974, J. Reinert. 2 6*6*, 5 00, Rio Napo at Limoncocha (Napo) , light & Flight traps, 19-24 May 1976, D.G.Y. & T. Rogers. 2 00., 17 km E of Santa Domingo de los Colorados (Pichincha) , burrow, 4 May 1976, D.G.Y. & T. Rogers. Panama. 12 c?cf, 24 00, various dates and localities in the Canal Zone, Colon, 4 Panama Provinces. Trinidad. 1 o_, Nariva Swamp . Bush Bush Forest, light trap, 2-3 Nov. 1965, T.H.G. Aitken. Discussion: Lutzomyia nordestina, a widespread species occurring on both sides of the Andes in Colombia and Ecuador, is often found resting

PAGE 451

-439in animal burrows. There is some evidence indicating that females feed on cold-blooded vertebrates (Tesh et al., 1971a). 1 have examined females of an undescribed Lutzomyia sp. from Aitamira (Para), Brazil (km 164), which resembles that of nordestina in most nonsexual and sexual characters but the cibarial armature is quite different. Other than this species, L. nordestina does not seen to be closely allied to any other described taxon. 104. Lutzomyia rangeiiana (Fig. 88) Phlebotomus rangelianus Ortiz, 1952: 153 (6* holotype, Duaca, Lara State, Venezuela). Pifano et al., 1962: 384, 389 (6\ o_, keyed), 406407 (
PAGE 452

-440absetit, individual sperm ducts. The sac-like spermathecae, cibarial armature and pigment patch of the females, the male genitalia, and palpi of both sexes, however, resemble those of the varruoarum group spp . , indicating perhaps that vangeliana and these species may have evolved from common stock. Except for L. nevesi (Damasceno & Arouck) , the males of the verruoarum group possess a coxite tuft or tufts although this feature may not always be present in taxa which appear to be otherwise closely related. This suggests that vangeliana may belong with them when all characters are considered, the male lacking a coxite tuft but having simple parameres, reduced spines on the style, and a subterminal seta like those in the series sevvana (vevrucarvm group) . 105. Ly.tzomyia sp. of Anchicaya (Fig. 89) Male (n = 1): Wing length 1.98; width 0.56. Head, mesonotum, lower half of pleura, coxae, and abdominal tergites well pigmented, rest of insect paler. Head height 0.36; width 0.36. Eyes large, separated by 0.11 or by distance = to 6.2 facet diameters. Flagellomere I (0.28 long), 1.1 x length of II + III; ascoids simple, no posterior spurs, on all flagellomeres except last, although difficult to see on two preceding flagellomeres. Length of palpal segments: 1 (0.04), 2 (0.09), 3 (0.14), 4 (0.07), 5 (0.14); palpal sensilla (5) on distal fourth of segment 2. Labrum length 0.21. Cibarium with ca. 20 subequal vertical teeth in 2 irregular rows, some lateral teeth, vestiges of horizontal teeth on sides with a space in middle; chitinous arch nearly complete, diffuse in middle; pigment patch sub triangular , faintly pigmented. Pharynx length

PAGE 453

0.19. Pleura with 20 upper and 6-7 lower episternal setae. Length of wing vein sections: Alpha (0.54), beta (0.19), delta (0.15), gamma (0.19). Length of femora, tibiae, and basitarsi: Foreleg, 0.70, 0.99, 0.54; midleg, 0.78, 1.28, 0.74; hindleg, missing. Abdominal sternites apparently entire, lacking nonsclerotized openings. Genitalia: Style (0.12 long) with a proximal isolated spine, 3 large spines at apical third of structure and a small sub terminal bristle. Coxite (0.19 long x 0.06 wide), lacking a tuft, deciduous setae mostly strap-like. Paramere simple. Aedeagus (0.10 long), sub triangular, more slender distally. Lateral lobe (0.21 long). Genital pump (0.17 long); each filament 0.37 long or nearly 2.2 x length of pump, filament tips simple, angular, not pointed. Cerci as shown. Distribution: Colombia (Valle) . Material examined: Colombia. 1 d\ Anchicaya Dam (Valle), tree trunk, 9 Aug. 1973, D.G.Y. & R.C.W. Discussion: Although the coxite tuft is lacking, this male may belong in the subgenus Pintomyia, confirmation being possible only when complete males and females are discovered. The hind legs are missing from the male on hand; therefore it is impossible to determine whether or not short spines are present on the hind femora.

PAGE 454

Figure 87 Lutzomyia nordestina — A. Male head, B. Male flagellomere II, C. Female head, D. Female flagellomere II, E. Female cibarium and pharynx, F. Genital pump and filaments, G. Male genitalia, H. Spermathecae, I. Female wing, J. Male wing, K. Female cibarium. Male: Guriche, Choco Dept., Colombia Female: Same locality as male

PAGE 455

-44;

PAGE 456

Figure 88 Lutzomyia vangeliana — A. Male head, B. Female cibarium, C. Spermar.hecae, D. Male genitalia. Male: Trinidad Female: Trinidad

PAGE 457

-445v>M V V-

PAGE 458

Figure 89 Lutzomyia sp. of Anchicaya male — A. Head, B. Wing, C. Genital pump and filaments, D. Flagc-;] ioroere. II, E. Genitalia. Kale: r>:ich i.caya Darn, Val le Dept., Colonbia

PAGE 459

-film # » \ \ i . .» ! U 1 t — ? M i i / : _y" // v\-

PAGE 460

SYNOPSIS OF NEW TAXA, SYNONYMS, FEMALE DESCRIPTIONS, AND NEW DISTRIBUTION RECORDS Taxa Sycorax andicola Sycorax aolombiensis Sycorax faivchi Idi Sycorax trispinosa Luizomyia (L.J marinkellei Lutzomyia moralesi Lutzomyia, barrettoi majuscula Lutzomyia (T. ) howardi Lutzomyia (T.) cellulana Lutzomyia (T. ) saltuosa Lutzomyia (P. ) carrerai thul-a Lutzomyia. (P. ) of Tres Esquinas Lutzomyia sp. of Pichinde Lutzomyia sp. of Anchicaya Lutzomyia tintinnabula = L. ayrozai Lutzomyia gasti L. waVkeri male Descriptions Lutzomyia (L.) bifoliata Lutzomyia (P. ) hirsuta r.icaraguensis Lutzomyia pilosa New Distribution Records COLOMBIA Brumptomyia hamata Brumptomyia leopo~ldx)i Lutzomyia baityi Lutzomyia (P.) bispinosa Lutzomyia carpenteri Lutzomyia dreisbachi Lutzomyia (Pr.J dysponeta Lutzomyia isovesp>e.rti lionis Lutzomyia rnigonei Lutzomyia nunsztovari Lutzomyia scorzai

PAGE 461

-449COSTA RICA Lutzomia piu CUBA Lutzomyia oayennensis ECUADOR Bvumptomyia galindoi Lutzomyia abunaensis Lutzomyia aalydifeva Lutzomyia baity i Lutzomyia (P. ) bispinosa Lutzomyia (P. ) cavrerai oarrerai Lutzomyia (P. ) cavrevai thula Lutzomyia (P, ) davisi Lutzomyia dendrophy'uz Lutzomyia (II.) flavisoutellata Lutzomyia guyanensis Lutzomyia hartmanni Lutzomyia (P.) hirsuta kirsuta Lutzcrryia micropyga Lutzomyia nevesi Lutzomyia (P.) nocticola Lutzomyia (S. ) olmeaa bicolor Lutzomyia (P. ) pananensis Lutzomyia (T. ) rebwra Lutzomyia ur.dulata Lutzomyia walker I Lutzomyia (N.) yuilli GUATEMALA Lutzomyia (II.) ylepkiletov HONDURAS T'Utzomyia evansi Lutzomyia texawa

PAGE 462

-450PANAtlA Lutzomyla (P. ) noa Lutzomyia vangeVLa. Lutzomyia (T . ) vec PARAGUAY Brvjnptomqia gatindoi

PAGE 463

REFERENCES Aboimenc , E . 1948. Phlebo tomes de la Guyane Francaise (YJOJ) . Sur P. ayvozai Barretto er Coutinho, 1940 et P. varaensis Costa Lima 1941. Inst. Pasteur Guyane, Pub. No. 177, 4 p. 1972, Les Phlebotomes cle la Region Ethiopienne (Dxptera, Psychodidae) . Mem. O.R.S.T.O.M., No. 55, 289 p. Abonnenc, E. and N. Leger 1975. Sur une classification rationnelle des dipteres Phlebotomidae. Call. O.R.S.T.O.M. Ser. Ent. Med. Paraslt. 14:69-73. Abonnenc, E. and D.M. Minter 1955. Bilingual keys for the identification of sandflies of the Ethiopian Region (French and English). Cah. O.R.S.T.O.M., Ser. Ent. Med., No. 5, 63 p. Aif-ken s T.H.G., J. P. Woodall, A. P.P. de Andrade, G. Bensabath, and R.F-. Shope 1975. Pacui virus, phlebotomine flies and small mammals in Brazil: An epidemiological study. Amer J, Trop . Med. Hyg. 24: 358-368. Aitken , T . H . G . , C . B . Worth , and E . 5 . Hi kasingh i960. Arbovirus studies in Bush Forest, Trinidad W.I., September 1959-December 1964. Amer. J. Trop. Med. Hyg. 17:253-268. Almeida, F.B. 1970a. Flebotomos da Amazonia II — Sobre ocorrencia de anomalias no genitalia masculina em Lutzomyia anduzei (Rozeboom, 1942) e Lutzomyia vovataensis (Floch and Abonnenc, 1944) (Diptera, Psychodidae). Bol. INPA Path. Trop. 1:1-4. 1970b. Flebotomos da Amazonia I — Sobre a presence de Lutzomyia anduzei (Rozeboom, 1942) no Brasil (Diptera, Psychodidae). Bol. INPA Path. Trop. 3:1-16. Anduze, P.J., F. Pifano, and E. Vogelsang 3 947. Homina de los atropodos vulnerantes conocidos actualmente en Venezuela. Bol. Ent. Venez. (num. extra), 1-16. An l ur.es , P . C . A . 1937. Informe sobre una investigation entomologica realizada en Colombia. Rev. Fac . Med. Univ. Nac . Colomb. 6:3-29. -451-

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Arjona, A.C., E. Osorno-Mesa, and J. P. Marquez 1.971. Contribucion al estudio epideraiologico del kala-azar en Colombia. "Rev. Fac. Med. Univ. Nac. Colomb. 37:90-94. Arzube, R. 1960. Los Phlebotomua del Ecuador. Rev. Ecuat. Hig. Med. Trop . 17:155-159. Barreto, P. 1969. Artropodos hematcff agos del Rio Raposo, Valle, Colombia. IV. Psychodidae. Caldasia 10:459-472. Barretto, M.P. 1941. Morfologia dos ovos, das larvas e pupas de alguns fiebdtomos de Sao Paulo. Anais Fac. Med. Univ. S. Paulo 17:357-427. 1046?. Sobre a sinonimia de flebotomos Americanos (Diptera, Psychodidae). Anais Far. Med. Univ. S. Paulo 22:1-27. 1946b. Sobre a sinoniraia da flebotomos Americanos (Diptera, Psychodidae). Primeira nota. Rev. Brasil. Biol. 6:527-536. 1946c. Uma nova especie de Ilebotorao do Estado de Goias, Brasil, e chave para determinac,ao das esyecies afins (Diptera: Psychodidae). Rev. Brasil. Biol. 6:427-434. 1946d. Urca nova especie de flebotomo da Colombia e chave para a deterrainacao das especies afins. Anais Fac. Med. Univ. S. Paulo 22:279-293. 1947. Catiilogo dos flebotomos Americanos. Arch. Zool . Est. S. Paulo 5:177-242. 1950a. Chaves para os flebotomos Brasileiros (Diptera, Psychodidae) Ar.ais Fac. Med. Univ. S. Paulo 25:101-120. 1950b. Duas novas especies de flebotomos Brasileiros (Diptera, Psychodidae). Folia Clin. Biol. 16:143-149. 1951. Nova contribuiQao para o estudo da distribuigao geographica dos flebotomos Americanos (Diptera, Psychodidae). Arch. Hyg. Saude Publ. 15:211-226. 1953. Sobre a sistematica da subfamiiia Phelbotominae Rondani (Diptera, Psychodidae). Terceira nota. Folia Clin. Biol. 20:209-214. 1954. Sobre a sinonimia de flebotomos Americanos (Diptera, Psychodidae). Quarta nota. Folia Clin. Biol. 21:121-126. 1955a. Sobre a sistematica da subfamiiia Phlebotominae Rondani. Rev. Brasil. Ent. 3:173-190.

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Barretto, M.P. 1955b. Nova espe"cie de flebdtomo do Brasil (Diptera, Psychodidae) . 8ev. Hosp. Clin. S. Paulo 10:247-249. 1956. Sob re o genero Sycovax Hal., coir, as describes de duas novas especies do Rrasil (Diptera, Psychodidae). Folia Clin. Biol. 26:71-78. 1961. Introducao ao estudo systema'tico dos f iebctomlneos Americanos (Diptera, Psychodidae). Thesis, Fac. Med. Kebeirao Pretc, 171 p. .1962. Novas subgeneros de Lutzomyia Franca, 1924 (.Psychodidae, subfamilia Phlebotominae) . Rev. Inst. Med. Trop. S. Paulo 4:01-100. 1966. Estudes sobre flebotomineos Americanos. I. Notas sobre especies de Lutzomyia Franca, subgenero Psycho dopy gixs Mar.gabeira, 1941, com a descricao de una nova especie (Diptera, Psychodidae). Papeis Dep . Zool. S. Paulo 18: 133-146. Baretto, M.P. and J.O. Coutinho 1940. Contribuicao ao conhecimento dos flebotomos de Sao Paulo. II. Descrigao do macho de PhelbotorratS lirr.ai Fonseca, 1935 e de duas novas especies: Phlebotomus ayrozai e P. amarali (Diptera, Psychodidae). Anais Fac. Med. Univ. S. Paulo 16:127-139. 1943. Contribuicao para o conhecimento dos flebotomos de Sao Paulo. VIII. Descricao de femea de P. ayrozai Barretto and Coutinho, 1940 e do macho de P. basispinosus n.sp. (Diptera, Psychodidae). Rev. Brasil. Biol. 3:183-189. Baretto, M.P. and J. P. Duret 1953. Uica nova especie de flebotomo da Republica Argentina: Phlebotomus microcephalics, n.sp. (Diptera, Psychodidae). Papeis Dep. Zool. S. Paulo 11:341-344. Barretto, M.P., A.V. Martins, and J. Pellegrino 1956. Notas sobre flebotomineos do nordeste do Estado de Minas Gerais, coin as descricoes de duas espe'cies novas (Diptera, Psychodidae). Folia Clin. Biol. 26:47-56. Barretto, M.P. arid H. Zago Filho 1956. Flebotomos encontrados em Petropolis, R.J., com a descricao de utna nova especie (Diptera, Psychodidae). Rev. Rrasil. Ent. 5:177-135. Biagi, A.M., F.F. Beltran, and F.F. Biagi 1966. Nuevos conocimientos sobre los flebotomos del area endemica de leishmaniasis cutanea en Yucatan. Rev. Invest. Salud Publ., Mex. 26:139-153.

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-454Bl.ancas, I. 1959-1960. Notas sobre flebotomos Pei'uanos. Enuraeracidn de las especies de flebotomos encov. trades pii el Peru y descripcidn de una e.specie nueva, Phlebotomus govbitzi n.sp. (Diptera, Psyenodidae) . Rev. Med. Exp., Lima 13:125-134.. Sorror, O.J... D.M. DeLong, and C.A. Tripl^horn 1976. An introduction to the study of insects. Holt, Rinehart and Winston, N.Y. 852 p. nrora, K.S., Jr. 1975. Geographical patterns of evolution in Neotropical Lepidoptera. Systenatics and derivation of known and new Helieoniini (Nymphalidae: Nymphalinae) . J. Ent. (3) 44:201-242. Eruijning, C.F.A. 1957. Man-biting sandflies in endemic leishmaniasis area of Surinam. Doc. Med. Ceogr. Trop. 9:229-236. Buck, A. A., Sasaki, T.T., and R.I. Anderson 1968. Health and disease in four Peruvian villages. Contrasts and Epidemiology. The Johns Hopkins Press, Baltimore. 142 p. Calderon, L. .1973. Phtebotomus townsendi, Oritz, 1959, en una zona endemica de leishmaniasis tegumenteria del Estado Merida, VenezuelaAc La Cient. Zenez. 24:87. Calisher, C.H., E.G. McLean, G ,C . Smith, D.M. Szmyd, D.J. Muth, and J.S. Laauick .19/7. Rio Grande -A new phlebotomus fever group virus from south Texas. Araer. J. Trop. Med. Hyg. 26 (in press). Callan, E. McC 1947. A rote on Phlebotomus tri-nidadensis Nevstead (Dipt., Psychodidae) . Rev. Ent. 18:215-218. CarnheJ.ro, M. end l.A. Sherlock 1964. Escuco morfologico sobre as pupas de Phlebotominae. (Diptera, Psychodidae). Rev. Brasil. Malar. Doeng. Trop. 16:311-329. Causey, O.R. and Pv.C. Damasceno 1945. Estudo sobre Flebotomus no Vale Anazonico. Parte IV. Descrigao de F. cevqueivai-, F. dveisbaehi 3 F. meivai e F. fevrelvai (Dtptera-Psychodidae) . Mem. Inst. Oswaldo Cruz 42:645-660. L948. Estudo sobre Flebotomus no Vale Amazonico. Parte II. Cescricao de F. dunharni, F. melloi e F. waglevi. Rev. Ser. Esp. Sauce Publ. 1:625-640. Ch-jnioMs, B.N'. 1971. Female gynandromorphs of Lutzomyia. trinidadensis Newstead from Panama. J. Med. Ent. 8:459.

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Rutledge, L.C. and D.A. Ellenwood 1.975b. Production of phlebotomine sandflies on the open forest floor in Panama: hydrologic and physiographic relations. Environ. Ent. 4:78-82. 1975c. Production of phlebotomine sandflies on the open forest floor in Panama: Phytologic and edapbic relations. Environ. Ent. 4:83-89. Rutledge, L C, D.A. Ellenwood, and L. Johnston 19 75. An analysis of sandfly light trap collections in the Panama Canal Zone (Diptera: Psychodidae) . J. Med. Ent. 12:179-183. Rutledge, L.C. and H.L. Mosser 1972. Biology of immature sandflies. (Diptera: Psychodidae) at the bases of trees in Panama. Environ. Ent. 1:300-309. 1975. Notes on the host ranges of some phlebotomine sandflies in Panama. Mosquito News 35:410-411. Saether, O.A. 1971, Notes on the eepprnl morphology and terminology of the Chironomiriae (Diptera). Can. Ent. 103:1237-1260. Samaniego, B.M. 1944. Comision de estudios sobre bartonellosis. Pasto-NarinoColombia. Situacion actual y programa. Rev. Fac . Med. Univ. Nac. Colomb. 13:523-530. Schneider, C.R. and M. Hertig 1965. Immunodiffusion reactions of Panamanian Leishmania. Exp. Parasit. 18:25-34. Schultz, M.G. 1968. A history of bartonellosis (Carrion's disease). Araer. J. Trop. Med. Hyg. 17:503-515. Scorza, J.V., I. Gomez, and M. Ramirez 1968. Observaciones biologicas sobre algunos flebotomos de "Rancho Grande" (Venezuela). 3. Notas bionomicas sobre Phlebotomus townsendi Ortiz, 1959 y P. vexillarn-us Fairchild and Hertig, 1952. Acta Biol. Venez. 6:28-40. Scorza, J.V. and I. Ortiz 1960. On microclimatic conditions of the habitats of certain sandfly species (Diptera, Psychodidae) from Venezuela. Z. Tropenmed. Parasit. 11:433-440. Scorza, J.V., 1. Ortiz, and M.T. McLure 1963. Ecologia de las formas estacionales de Phlebotomus townsendi Ortiz, 1960 y Phlebotomus oayennensis Floch & Abonnenc, 1941 (Diptera, Psychodidae) en el centro-norte de Venezuela. Acta Biol. Venez. 3:437-45 3.

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Scorza, J.V., I. Ortiz, and M. Ramirez 1967. Obsecvaciones biologicas sobre algunos flebotomos de Rancho Grande (Venezuela). I. Descripcion de un area restringida tie la selva nublada y taxonoraia de los Fh'le.botonvAS (Diptera, Psychodidae). Acta Biol. Venez. 5:179-200. Shannon, R.C. 1926. The occurrence of Vhleboloraua in Panama. J. Wash. Acad. Sci. 16:190-193. 1939. Methods for collecting and feeding mosquitoes in jungle yellow fever studies. Auier. J. Trop . Med. 19:131-138. Shaw, J.J. J 964. A possible vector of Endotrnjpanum schaudini of the sloth Cholsopus hoffmanni, in Panama. Nature 201:417-418. Shaw, J.J. and R. Lainson 1972. Leishmaniasis in Brazil. VI. Observations on the seasonal variations of Lutzomyza fl.avisoutellata in different types of forest and its relationship to enzootic rodent leishmaniasis (Leishmania rnexioana amazonensis) . Trans. Roy. Soc . Trop. Tied. Hyg. 66:709-717. Shaw, J.J., R. Lainson, and R.D. Ward 1972. Leishmaniasis in Brazil. VII. Further observations on the feeding habits of Lutzomyia flaviscutsllata (Mangabeira) with particular reference to its biting habits at different heights. Trans. Roy. Soc. Trop. Med. Hyg. 66:718-723. Sherlock, I. A. 1962. Sobre alguns Phlebotomus e Brurnptomyia da Colombia (Diptera, Psychodidae). Mem. Inst. Oswaldo Cruz 60:321-336. Sherlock, I. A. and M. Carnheiro 1962. Algumas femeas de Phlebotomos do Brasil (Diptera, Psychodidae). Mem. Inst. Oswaldo Cruz 60:423-435. 1964. Observacoes sobre o aparelho genital interno do macho de alguns "Phlebotominae" (Diptera, Psychodidae). Rev. Brasil. Biol. 24:203-210. Sherlock, I. A. and N. Guitton 1970. Notas sobre o subgenero Trichophovorrtyia. Barretto, 1961 (Diptera, Psychodidae, Phlebotominae). Rev. Brasil. Biol. 30:137-149. Sherlock, I. A. and S.B. Pessoa 1964. Metodos praticos para a captura de flebdtomos. Rev. Brasil. Biol. 24:331-340. 1966. Lepi;omonas infectando naturalmente Fhlebotomis em Salvador (Bahia, Brasil). Rev. Lat.-Amer. Microbiol. 8:47-50.

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-477Snodgrass, R.E. 1935. Principles cf insect morphology. McGraw-Hill, New York and London. 667 p. Strangways-Dixon, J. and R. Lainson 1962. Dermal leishmaniasis in British Honduras: Transmission of L. brasitiensis by Phtebotomus species. Brit. Med. J. 1:297-299. 1966. The epidemiology of dermal leishmaniasis in British Honduras. Part III. The transmission of L. mexioana to man by Phtebotomus pessoanus with observations on the development of the parasite in different species of Phtebotomus. Trans. Roy. Soc. Trop. Med. Hyg. 60:192-201. Sudia, W.D. and R.W. Chamberlain 1962. Battery light trap, an improved model. Mosquito News 22: 125-129. Tesh, R.B. and B.N. Chaniotis 1975. Transovarial transmission of viruses by phlebotomine sandflies. Ann. N.Y. Acad. Sci. 266:125-134. Tesh. R.E., B.N. Chaniotis, M.D. Aronson, and K.M, Johnson 1971a. Natural host preferences of Panamanian phlebotomine sandflies as determined by precipitin test. Amer. J. Trop. Med. Hyg. 20:150-156. Tesh, R.B., B.N. Chaniotis, B.R. Carrera, and K.M. Johnson 1972. Further studies on the natural host preferences of Panamanian phlebotomine sandflies. Amer. J. Epidem. 95:88-93. Tesh. R.B., B.N. Chaniotis, and K.J. Johnson 1971b. Vesicular stomatis virus, Indiana serotype: Multiplication in and its transmission by experimentally infected phlebotomine sandflies (Lutzomyia trapidoi) . Amer. J. Epidem. 93:491-495. Tesh, R.B., B.N. Chaniotis, P.H. Peralta, and K.M. Johnson 1974. Ecology of viruses isolated from Panamanian phlebotomine sandflies. Amer. J. Trop. Med. Hyg. 23:258-269. Thatcher, V.E. 1968a. Studies of phlebotomine sandflies using castor oil traps baited with Panamanian animals. J. Med. Ent. 5:293-297. 1968b. Arboreal breeding sites of phlebotomine sandflies in Panama. Ann. Ent . Soc. Amer. 61:1141-1143. Thatcher, V.E. and M. Hertig 1966. Field studies on the feeding habits and diurnal shelters of some. Phtebotorwxs sandflies (Diptera: Psychodidae) in Panama. Ann. Ent. Soc. Amer. 59:46-52.

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Thai ;<-•;: , M.T. and W.G. Downs 19 73, The arthropod-borne viruses of vertebrates. Yale Univ. Press, New Baveu. 57S p. Tneodor, 0. 1948. Classification of the Old World species of the. subfamily Phlebotominae (Diptera, Psychodidae) . Bull. Ent. Res. 39:85-115. 1965. On the classification of American Phlebotominae. J. Med. Ent. 2:173-197. Tikasiogh, E.S. 1973, Observations on Lutzomyia f'LaviscutetZaza (Mangabeira) (Diptera: Psychodidae), a vector of enzootic leishmaniasis in Trinidad, West Indies. J. Med. Ent. 12:228-232. Tovnes, H. 1962. Design for a Malaise trap. Proc. Ent. Soc. Wash. 64:253262. Vargas, L. and k. Diaz-Najera 1951a. Pklebotomus pvatti, n.sp de Iguala, Estado de Guerrero (Insecta: Deptera) . Bo.l. . Inst. Estud. Med. Biol, Univ. Nac. Mex. 9:19-23. 1951b. Phlebotomus beltrcni n.sp. y Fhlebotomus wivthi n.sp. del Estado de Veracruz (Diptera: Psychodidae). Rev. Inst. Saluhr. Enferm. Trop. Mex. 12 : 101-106. 1952. Dos flebotomos nuevos de Mexico (Diptera, Psychodidae). Rev. Soc. Mex. Hist. Nat. 13:63-74. 1953a. Nuevas especies de Flebotomus de Mexico. Rev. Inst. Salubr. Enferm. Trop., Mex. 13:41-52. 1953b. Lista de flebotomos Mexicanos y su distribucion geografica (Diptera: Psychodidae). Rev. Inst. Salubr. Enferm. Trop. Max. 13:309-314. 1959. Phlebotomus farilli n.sp., Ph. hvntboldti n.sp. y Ph. olmeous f:.sp. de Mexico (Diptera: Psychodidae). Rev. Inst. Salubr. Enferm. Trop., Mex. 19:141-159. Ve ' asco, J . E . 1973. The phlebotomine sandflies of the Los Yungas region of Bolivia. M.S. Thesis, Lousiana State Univ., Dep. Trop. Med. & Med. Parasit. , 204 p . Velasco, J. and H. Trapido 1974. Two new phlebotomine sandflies from Bolivia, Lutzomyia boliviano, n.sp. and Warileya yungasi n.sp. (Diptera, Psychodidae). J. Med. Ent. 11:433-436.

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-479Word, R.D. 1976. The immature stages of some phlebotomine sandflies from Brazil (Diptera: Psychodidae) . Syst. Ent. 1:227-240. 1977. New World leishmaniasis: A review of the epidemiological changes in the last three decades. Proc. XV Int. Congr. Ent., Washington, 505-522. Ward, R.D. and H. Fraiha 1977. Lutzomyia urrbratitis n.sp. — A sandfly previously identified as L. andlizei (Rozeboom, 1942) (Diptera: Psychodidae).. J. Med. Ent. 14 (in press). Ward, R.D. and R v Killick-Kendrick 1974. Field and laboratory observations on Psyohodopygus lainsoni Fraiha & Ward and other sandflies (Diptera, Phlebotomidae) from the transamazonica highway, Para State, Brazil. Bull. Ent. Res. 64:213-221. Ward, R.D., R. Lainson, and J.J. Shaw J 977, Experimental transmissions of Leishmania mexicana amazonensis Lainson & Shaw, between hamsters by the bite of Lutzomyia flavisoutellata (Mangabeira) . Trans. Roy. Soc. Trop . Med. Hyg. 71:265-266. Ward, R.D. and P. Ready 1975. Chorionic sculpturing of some sandfly eggs (Diptera: Psychodidae). J. Ent. (A) 50:127-134. Ward, R.D., J.J. Shaw, and R. Lainson 1973. Leishmaniasis in Brazil. VIII. Observations on the phlebotomine fauna of an area highly endemic for cutaneous leishmaniasis, in the Serra dos Carajas, Para State. Trans. Roy. Soc. Trop. Med. Hyg. 67:174-183. Wijers, D.J.B. and J. Huisenga 1.967. A new species of Lutzomyia from Surinam (Diptera: Psychodidae). Bull. Ent. Res. 57:395-397. Wiyers, D.J.B. and R. Linger 1966. Man-biting sandflies in Surinam (Dutch Guiana). Phlebotomus anduzei as a possible vector of Leishrnania brazitiensis . Ann. Trop. Med. Parasit. 60:501-508. Williams, P. 1970. Phlebotomine sandflies and leishmaniasis in British Honduras (Belize). Trans. Roy. Soc. Trop. Med. Hyg. 64:317-368. 1976a. The phlebotomine sandflies (Diptera, Psychodidae) of the caves in Belize, Central America. Bull. Ent. Res. 65:601-614. 1976b. Flagellate infections in cave-dwelling sandflies (Diptera, Psychodidae) in Belize, Central America. Bull. Ent. Res. 65:615-629.

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".J 'i 1.

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BIOGRAPHICAL SKETCH David Grier Young was born May 9, 1940, at Dayton, Ohio. In June, J.953, he was graduated from Oakwood High School, Dayton, Ohio. In December, 1962, he received the degree of Bachelor of Science with a major in entomology from the University of Florida. From i.963 to 1968, he served as a commissioned officer in the Medical Service Corps, U.S. Army at Ft. Sam Houston, Texas, and the Panama Canal Zone. Following his discharge, he worked on a research project involving siudli mammals and ectoparasites in Ohio (Dayton Museum of Natural History) In 1968, be enrolled in the Graduate School of the University of Florida, receiving the degree of Master of Science in June, 1971, with a major in entomology, From 1972 to December, 1977, he Las been co-investigator and principal investigator of a U.S. Army Medical Research and Development contract dealing with the systematics of phlebotomine sand flies. The author is a member of the American Association for the Advancement of Science, the Entomological Society of America, and the Florida Entomological Society. He is a research associate of the Florida State Collection of Arthropods. David Young is married to the former Marianna Brown. They have twin sons, Alexander and Mark. -48.1

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1 certify that I have read this study and that in my opinion it coniorns Co acceptable standards of scholarly presentation and is fully adequate, i;i scope and quality, as a dissertation for the degree of Doctor of Philosophy. ^7 ,/X
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1 certify that I have read this study and that in ray opinion it conforms tn acceptable standards of scholarly presentation and is fully adequate, in scope and quality, as a dissertation for the degree of Doctor of Philosophy. 9/V? Dr. S.G. Zam f> Associate Professor of Microbiology This dissertation was submitted to the Graduate Faculty of the College of Agriculture and to the Graduate Council and was accepted as partial fulfillment of the requirements lor the degree of Doctor of Philosophy. December 1977 vok dC.
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UNIVERSITY OF FLORIDA 3 1262 08553 1670


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