A BIOSYSTEMATiC REVIEW OF THE BLOODSUCKING PSYCHODID FLIES OF
COLON-1BIA (DIPTlRA: PHL.EBOTOUMOLAE AND SYCORACfNYE)
DA"ID GRCI-R YOUNG
A DISS1;'RTATrION PRESENTED TO THE GRADUATE COUNCIL OF
THE UN [VERSTTY OF FLORITDA
IN PARTIAL FULFILLMENT OF THE REQUIRiK4FNTS FOR THE
DEGREE OF DOCTOR OF PHILOSOPHY
UL VERSIY OF FLORIDA
I)~dicated to my father,
Howiard G. Young
TABLE OF CONTENTS
LIST O FICURS. .. . . . . . . . . . .
LBSTRAC . . . . . . . . . . . .
INTRODUCTION . . . . . . . . . . . . .
CLASS1FICAT[ON AND TAXONOMIC TREATMENT . . . . . . .
Keys to the Subfamilies of New World Psychodidae. . . .
GE-N' .AL MO)kliOLOGY AND TERMINOLOGY . . . . . . . .
NETiHUDS AND MATERIALS. . . . . . . . . . . .
Field Collections ..... . . . . . .
Laboratory Methods. . . . . . . . . . .
ODSTRIBUTIOX, COLLECTING LOCALITIES, AND CHARACTERISTICS OF
L UN . . . . . . . . . . . . ...
Pacific Coast Region. . . . . . . . . . .
Atlantic Coast Region . . . . . . . . . .
Andean Region . . . . . . . . . . . .
Oriental Region . . . . . . . . . . .
MEDICAL IMPORTANCE . . . . . . . . . . . .
SUliFAMILY SYCORACINAE. . . . . . . . . . . .
Keys to the Sycorax Species . . . . . . . .
1. Sycorax andicola n. sp. . . . . . . .
2. Sycorax colombiensis n. sp. . . . . . .
3. Sauorax fairchildi n. sp. . . . . . .
.. Syiorax tricpinosa n. sp. . . . . . .
S; r,'AMITLY liOLEBOTOMINAE. . . . . . . . . . .
Key to the Now World Genern . . . . . . . .
Genuon W'";.n!. yc ,iortig, 1948 . . . . . . . .
.kevys to Species . . . . . . . . . . .
5. .Vila. (W.) n i,' us . . . . ....
6. W.r, l/ (W. ) r'tudipen.is. . . . . . .
Genus Bruwptomyjia Franca & Parrot, 1921 . . . . . .
Key to Species . . . . . . . . . . . .
7. Brmrptonyia avlari . . . . . . .
8. Brptojnyia beapertuyi . . . . . . .
9. Bruznptromyia galindoi . . . . . . . .
10. Brumrptornjia haata. . . . . . . . .
11. Brwptomya leopoldoi . . . . . . . .
Ge-nus ,utneorryia Franca, 1924. . . . . . . . ...
Keys to Subgenera, Species Groups, and Ungrouped Species. .
Subgenus Lutzomyjia Franca, 1924 . . . . . . . .
Keys to Species . . . . . . . . . . . .
(L.) bifoliata. .
(L.) lihyi . .
(L.) Zongipalpis. .
(L.) gomoi . .
(L.) marinkellei n.
Migonpi Group Theodor, 196b . . .
Keys to Species . . . . . .
. . . . . . 112
. . . . . . 112
Lutzsomyia migonei . .
Lutzomyia mar'joensi .
Lutsoimjia sp. de Baduel .
Lutzormyia walk . .
Saulensis Group Lewis et al., 1978. . . . . . .. 125
21. Lutzomyia saulensis . .
Verrucarvu Group Theodor, 1965. . .
Key to Species. . . . . . .
22. Lutwomyia odax. . . .
23. Lutzomyia serrana . . .
24. Lutzonryia andina. . . .
25. LutsomZia columbiana . .
26. Lutzompia disiunta . .
27. Lutzomyna evansi. . . .
28. Lutsomyia longijfocosa. .
29. Lutsoemyia moralesi n. sp.
30. Lutsonjia nunestovari .
31. Lutzoemia ovallosi .....
32. Lutoryjia quasitownsendi.
33. Lutsomyia sauroida. ....
34. L'itzomsyia spinicrmassa . .
Vespertilionis Group Theodor, 1965. .
Key to Species. . . . . . .
. . . . . . 125
. . . . . .. 130
. . . . . 130
. . . . . . 161
. . . . . 161
35. Lutzomyia isovceprt-"ion s . . . . . . 161
36. Lut:'omrjia vesptiii. . . . . . . 162
Subgenus Dacqpforyiacz Addis, 1945 . . . . . . . 167
37. Lutzomyia (D.) rooabg-li .
Subgenus Pintormia Costa Lima, 1932 .
38. Lutsomyia (P.) spi .osa . . . . . . ..
Subgenus P'asscatia Mangabeira, 1942 . . . . . . .
Keys to Species . . . . . . . . . . . .
39. Lutzorryia (Pr.) eccpoci .
40. LuttoMVia (Pr.) dysponeta .
41. Lutzomryia (Pr.) triceantha.
Baityi Group Theodor, 1965. . . ..
42. Lu toquia baitbjti,. . . ..
. . . . . . 177
. . . . . .. 178
. . . . . .. 180
. . . . . . 186
Subgenus Viannarryia Mangabeira, 1941. . . . . . ... 190
Keys to Species . .. . . . . . . . 190
43. Lutzorryia (V.) capmria. . . . . . . ... 191
44. Lutzomyia (V.) fur-ata. . . . . . . . 192
45. Lutzornyia (V.) tuber -ilata. . . . . . ... 194
Shannoni Group Theodor, 1965. . . . . . . . ... 200
Keys to Species . . . . . . . . . . . 200
46. Lutzomyia abonnenci .
47. Lutzo-yia dasymera . ....
48. Lutzomyia dendrophyla . .
49. Lutzomyia puncOtigeni-ulata.
50. LutzomnPia shannoni. . ....
51. Lutomyia undulata. . ....
Longispina Group Theodor, 1965. . . . . . . .. 224
Keys to Species . . . . . . . . . ... 224
52. Lutzomryia longispina. . .
53. L6utzconvia trirrvela . . .
Ara-gaci Group Theodor, 1965 . . .
Keys to Species . . . . . .
54. Lutzomyia aCa23i .. .....
55. Lut;.omyia barretbti ar.rettoli
56. L T,::on)ia karrettyi z' yuacuia
57. Lot:;;o ia ca,'p n i' .
58. Lut. omic r'nwoids ......
. . . . 225
. . . . . 226
. . . . . 232
. . . . . 232
Dreisbachi Group Lewia et al., 1978 . . . . . . 255
Key to Species. . . . . . . . . . 255
59. Lutzor;ia aolydifeva. . . . . . . .. 255
60. Lut.omyia drisbachhi. . . . . . . ... 257
Subgenus Trichophoro-wia Barretto, 1962 . . . . . 261
Key to Species. . . . . . . . . . . . 261
61. butornyia (T.) auraensis .. . . . . ... 262
62. Lutzorniria (T.) ceZlulana n. sp. . . . . ... 264
63. Lutwornyia (T.) houardi n. sp. . . . . ... 266
64. Lutzomyia (T.) reburra. . . . . . . ... 267
65. Lutzomyia (T.) saltzosa n. sp. . . . . ... 269
66. Ls.tzomyia (T.) ubiquitalis. . . . . . .. 270
Subgenus iNyssomyia Barretto, 1962. . . . . . ... 280
Keys to Species . . . . . . . . . 280
67. Lutzomyia (7l.) antunedi . . . . . . . 283
66. Lutzomyia (f..) flavi.ateilata. . . . . . 285
69. Lutzomjia (!.) olmeric bicolor . . . . . 287
70. Lutzomyia (i. ) trapidoi . . . . . . . 289
71. Lutzomtyia (U7.) urrnbvatilis . . . . . . 292
72. Lutzomyia (N1.) yZephiletor. . . . . . ... 295
73. Lutzomyia (Nl.) yuilli . . . . . . . 297
Subgenus Psychodopygvs Mangabeira, 1941 . . . . . 311
Keys to Species . . . . . . . . . . . 312
74. Lutr.omyia (P.) bera7ei . . . . . . . 317
75. Lutsomyia (P.) chagasi. . . . . . . .. 317
76. Lutzomyia (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.) davisi . . . . . . . 331
82. Lutaomyia (P.) gvyanensis . . . . . . 333
83. Lutzonia (P.) hirsuta.. . . . . . . ... 335
84. Lutzomyia (P.) noctiaola. . . . . . ... 338
85. Lutzorryia (P.) panamcnris . . . . . . 339
86. Lutaromjia (P.) rec:rva. . . . . . . ... 341
87. Lutbomnia (P.) sp. of Tres Esquinas . . . . 342
Vexator Gioup Theodor, 1965 . . . . . . . . 366
Keys to Species . . . . . . . . . . . 367
88. Lutzo' ,jia cirrita . . . . . . . . 369
89. L.i.zojmyia ho tman ri . . . . . . . 370
90. Lutzomia osoroi ... . . . . . 371
91. LutormUia sqhuiria . . . . . . . 372
92. Lutwor';ia scorsai . .
93. Lutzo.nyia ,ctrictivilla. .
94. Lutzomryia sp. of Pichinde
. . . . . . . 374
. . . . . . . 378
. . . . . . . 380
CaUennansis Group Theodor, 1965 . . . . . . . 398
Keys to Species . . . . . . . . . . . 398
95. Lutzomyia cayennensis .
96. Lutzomjyia micropyga .
97. Lutzomyia atroclavata .
98. Lutzomyia venesuelensis
OcsaZdoi Group Theodor, 1965. .
Keys to Species . . . . .
99. Lutzaorgia pia . . .
100. Lutzormy-a rorotaensic .
101. Lutzorrnyia trinidadensis
. . . . . . . 417
. . . . . . . 41 7
. . . . . . . 418
. . . . . . 4 20
. . . . . . . 422
Pilosa Gr"up Theodor, 195..
102. Lutzomyia pilosa. ... . . . . . . 432
Ungrouped Species . . . . .
103. Lutzonrria nordestina. . .
10A. Lutzomyia rangeliana. . .
105. Lutsomyia sp. of Anchicaya.
. . . . . 437
. . . . . . 439
. . . . . . 440
SYNOPSIS OF NEW TAXA, SYNONYMS, FEMALE DESCRIPTIONS, AND
NEW DISTRIBUTION RECORDS . . . . . . . . . . 448
REFERENCES ....... . . . . . . . . 451
BIOGRAPHICAL SKETCH. . . . . . . . . . . .. .iR1
LIST OF FIGURES
1. Map of Colombia showing localities where phlebotomine
and sycoracine flies have been collected . . . . 28
2. Sycooox andicola. . . . . . . . . . . 43
3. Syc-.r-a-. colombiensis. . . . . . . . . .. 45
4. Sycyorax fairchildi. ......... . . . . . 47
5. Soorax trispinosa. . . . . . . . . . .. 49
6. .a' '.,.ya? 7 nig, r.osa j'.. . . . . . . 57
7. Wlarileya rotundipennir.s ...... . . . .. 59
8. Bra.pto'ryia avellari and Brw ptorryia heazupertvyi. .. .... 69
9. Bruptoyia gaZindoi. . . . . . . . . .. 71
10. Brmptoyia hamata. . . . . . . . . . . 73
11 Br.nptomyia Zeopoldoi and Br,',tomryia guim.raesi. . . 75
12. Lutzonpjia (P.) hirsuta nicaragzuensis and Lutzomria
t ,wnsendi . . . . . . . . . . . . 90
13. Lutzomqyia (L.) bifoliata. ... . . . . . 103
14. Lutwsomyia (L.) Zichyi . . . . . . . . . 105
15. Lubzwrm-yia (L.) Zongipalpis. . . . . . . . .. 107
16. Lurzo' -,a (L.) gome.i . . . . . . . . . 109
I 7. LuttomyA a (T.) ,marin elliei. . . . . . . . .. . .11
18. Lursoxaia migonci and Lubvor.ia sp. de Baduel . . . 120
19. Lut;'o ,'.j marajoenai, . .... . . . . . 122
20. Lutomr,ia wa keii . . . . . . . . . 124
21. Latom.pi-a sazulen. is . . . . . . . . . . 129
22. Lutzor.yia servana. . . . . . 148
23. Lutomiyia andina, Lutzomyia disiu.cta, and L1;s:35r,,ia
n ne ov i . . . . . . . . . . . . 150
24. Lutssmyia olwumriana . . . . . . . . . 152
25. Lutzoamyia evansi . . . . . . . . . . . 154
26. Lutzomryia moralesi . . . . . . . . . 156
27. Lutsorjyia ovallesi . . . . . . . . . . 158
28. Lutzom nia sauroida, Lutzoryia quasi townseai, L. tzomyia
longiflocosa, Lutzomyia spinicrassa, and Lutzot: ia odax. . 160
29. Lut;orijyia vespertilionis and Lutzomyia isovesp rtiiionis . 166
30. Lut torniia (D. I rosabali. ... . . . . . . 170
31. Lutzormyza (Pi.) spinosa. . . . . . . . .... 175
32. Lutzomnyia (Pr.) camposi. . . . . . . . ... 183
33. Gu tomyia (Pr.) dysponeta. . . . . . . . .... 185
34. Lutsonayia (Pr.) tricantha and Lutzomyia baityi . . . 189
35. Lutzom.ia (V.) caprina and Lutzonmia (V.) furcata. . . 197
36. Lutzomyia (V.) tuberculata . . . . . . . . 199
37. Tutzonpyia abonnenci. . . . . . . . . . .. 213
38. Lztzomyia dasymera . . . . . . . . . . 215
39. Lutzomyia dendrophy a. . . . . . . . . 217
40. Lutsomrryia punctigeniculata . . . . . . . . 219
41. Lutbawoia a shannoni . . . . . . . . . . 221
42. Lut!'onrjia und lata . . . . . . . . . .. 223
43. Lutcormyia long/ipina .. . . . .. ... 229
44. Lutor,-ia triramula. . . . . . . . . . .. 231
45. Lut ia ao . . . . . . . . . 246
46. Lutsom7rIa barr ttoi majusutila. . . . . . .... 248
47. Lutzor.rj&La barretto; majuscula, LutorrT.jiia baarrettoi
barnethoi, and Lutzomyia texcrna . . . . . . . 250
48. Lutsomnia carpenter. ...... . . . . . 252
49. L'tzognija runoides. ....... . . . . . 254
50. LutzoGmyia aclydifera. . . . . . . . . ... 260
51. ij`utzomyia dreisbachi, Lutzom'yia (T.) anraensis, and
Lutzomyjia ubiquitalis . . . . . . . . 273
52. LL, tzom r ia (T.) celluZana.. . . . . . . . .. 275
53. Lutzonria (T.) howardi and Lutzsomia (T.) saLtzosa. .. .. 277
54. Lutzormia (T.) reburra.. . . . . . . . . .. 279
55. Lutwromyia (N.) antunesi and Lutzomyia (N.) flaviscutella;ta. 300
56. Lutsomy,,a (N.) olmeca bicolor . . . . . . 302
57, Lut.comy-ia (N.) trapidoi . . . . . . 304
58. Lutzojyia (N.) umbratilis . . . . . . . . 306
59. Lutsomyia (N.) ylephiletor. . . . . . . .... 308
60. Lutzomyia (N.) yuili . . . . . . . . . 310
61. Lutzorr-jia (P.) bernalei, Lutzomyia (P.) chagasi, and
Lutzomy a sp. of Tres Esquinas. . . . . . . . 345
62. Lutzorypia (P.) fairtigi . . . . . . . . 347
63. Lutzomyia (P.) bispinosa. . . . . . . . ... 349
64. Lutzojia (P.) ayrozai. ..... . . . . . 351
65. Lutzo'mjia (P.) carrerai carrerai.. . . . . . ... 353
66. Lutzomyia (P.) carrerai thula . . . . . . . 355
67. Lutzormyia (P.) davisi and Lutioeyia (P.) hirsuia hirut'. 357
68. Tut3onrmia (P.) guyanonsis . . . . . . . . 359
69. Lutzonyia (P.) nocticola. . . . . . . . ... 361
70. I,utzsorjia (P.) panamensis . . . . . . . 363
71. Lutaomyia (P.) recurva. .... . . . . . . 365
72. LuLt.omua cerrita. . . .
73. Lutzomyia kai'tmanni. . .
/74. Lutkzomjia osornoi. . . .
75. Lutzomniia sanauinaria. . .
76. Lutzomyia strictivilla . .
77. Lutzotmjia scorzai. . . .
78. Lutzomnyia sp. of Pichinde. .
79. Lutzoryia cayennensis. . .
80. Lutzo'nyia micropyga. . .
81.. Lutzornyia atroclavata. . .
82. Lutzomwjia venezueZensis. .
83. Lutzo Ryia pia. . . . .
84. Lutzomyia rorotaensis. . .
85. Lut-omyia trinidadensis. .
86. Lut;.omria pilosa . . .
87. Lutsomyjia nordestina . .
88. Lutwornjia rangeliana . .
89. Lutzomyia sp. of Anchicaya .
. . . . .
.. . . . .
. . .. . .
. . . . . .
. . . . . .
. . . . . .
. . . . . .
. . . . . .
. . . . . .
. . . . . .
. . . . . .
. . . . . .
. . . . . .
. . . . . .
. . . . . .
.. . . . .
. . 385
. . 387
. . 389
. . 391
. . 393
. . 395
. . 397
. . 410
. . 412
. . 414
. . 416
. . 427
. . 429
. . 431
. . 436
. . 443
. . 445
Abstract of Dissertation Presented to the Graduate Council
f Lhe University of Florida in Partial Fulfillment of the Requirements
for the Degree of Doctor of Philosophy
A BIOSYSTEMATIC REVIEW OF THE BLOODSUCKING PSYCHODID FLIES OF
COLOMBIA (DIPTERA; PFILFBOTOMINAlE AND SYCORACINAE)
David Grier Young
Chairman: Graham B. Fairchild
Major e-partmenr: Entomology and ematology
The psychodid subfamily Sycoracinae, previously unknown in Colombia,
is represented by four new Sycora:c species described and illustrated in
Phlebotominae, a much larger subfamily containing vectors of huian
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 ir the genus Lutacormiia are described as new and are given formal or
intormi names. iYh- previously unknown females of Lutsomyia pilosa, L.
(P.) h ircut nicaraguQenis and L. uifoliata are described. Nineteen
phlebotcr.ine species are reported in Colombia for the first time. Other
new distributional records include those from Ecuador (22 spp.), Panama
(3 szp.), Honduras (2 spp.), Costa Rica, Cuba, Guatemala, and Paraguay
(1 Fp. en--h). In order to clarify the status of some forms, an attempt
t.,j.s naide to study specimens from as many Colombian and non-Colombian
local iti's :13 possible. To this end, over 20,000 adults in both sub-
fanl tie. were examined from 1967 to the present.
An extensive bibliography and synonomy is given for each proviousLy
described taxon. Actual and/or probable misidentifications in the
literature are discussed. Identification keys, previously ur:available
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).
Prior to the last decade, we knew very little about the phlebotomine
sand fly- faun? 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 hii 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 Phlehotominae are re-
ported from Colombia. This representsabout 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-
The small subfamily Sycoracinae, whose members resemble those of
Phlebotomlnae 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
spc.inens from as many Colombian and non-Colombian localities as possible.
To this end, I am very grateful to the following persons for providing
specim.ons, information or both: Dr. Jorge R. Arias, Instituto Nacional
de PeOquisas da Amazonia (INPA), Manaus; Drs. Stephen C. Ayala and Pablo
*l prefer to separate "sand" from "fly" in accordance with the suggestion
of borrer, Delong, and Triplehorn (1976).
Barretto, Unjversidad del Valle, Call; Drs. Richard D. Guard an! Hnbib
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. Wilkeison, 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.
Fiairchild, Dale H. Habeck, and Stephen G. Zam. Miss Stephanie Haney ably
slide mounted thousands of specimens from 1972 to 1977. For typing the
ma.nuscript, 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 ami 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 Nuseum (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,
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 Horaiella, T'richorjia, and
Sycorax and its allies are united by some in the subfamily Trichomyiinae
hut this appears to be artificial and does not "express the wide discon-
Linuities and degrees of difference from other Psychodidae, especially in
The classification of Phlebotominae is interpreted differently
according to authorss. 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 -- Lutzomria Franga
(ca. 290 spp.), Brumptomyia Franga and Parrot (ca. 23 spp.), Warileya
Hertig (5 spp.). The genus Hertigia Fairchild is considered to be a
junior synonym of Warileya by Lewis et al. (1978).
The majority of the Lutzomyia spp. were described originally in the
genus Phlebotomr.u.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 subgenera, species
groups, and series based upon adult morphology (Lewis at al., 1978). Ttie
diversity of groups within the genus is attributed to "radiation or even
'explosive radiation' of some immigrating ancestral stocks in a newly
colu.nized 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 T'richophoromzia) 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 l.orld phlebotomines are
given by Lewis et al. (1978) and in this review under each supraspeciFic
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:
USNM -- United States National Museum (Nat. Hist.), Washington, D.C.
INPES -- aInstituto Nacional para Programns Especiales de Salud, Bogota
INPA -- Instituto Nactional de Pasquisas de Amazonia, Manaus
UV -- Universidad del Valle, Call
GML i(krgas Memo,'ial Laboratory, Panama
UF -- University of Florida
C.H.P. -- Charles H. Porter
C.J.M. -- Cornelius J. Marinkeile
R.C.W. -- Richard C. Wilkerson
D.G.Y. -- David C. Young
ef.-- compared to
classic. -- 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+3 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
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-
lomeres subcylindrical. . . . . . . . . . . .. 4
4. Females with 1 spermatheca. Mandibles absent. Male aedeagus
entire. . . . . . . . . . . . . Bruchomyiinae
Females with 2 spenrmathecae. Mandibles present. Males with bifid
aedeagus. . . . . . . . . . Phlebotominae (Fig. 12F)
GENERAL MORPHOLnGY AND TERMINOLOGY
This brief discussion focuses on the structures commonly used in
taxonoiaic 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
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
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
clypeas, in relation to its greatest width is of value in distinguishing
some of the Lt toyia species.
As a probable adaptive feature related to host finding, the size of
the compound Leyes varies according to species, especially those of
Lt;c'yiia, but may be characteristic of subgenera and equivalent- species
groups. Contrary to Davis (1967), ocelli are lacking in the Phleburcoi;ines
and Sycoiacines. Illustrations are by far the best way to indicate eye
size, the relative terms such as "small" 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 incelocular suture (= post frontal suture of mosquitoes,
Christophers, 1960) is complete in adults of Brumptormyia (Fig. 8B),
Warilcja (Fig. 6A), and Syoorax (Fig. 2A) but is incomplete in those of
Ltsoeeya (Fig. 12A).
InL 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 flagelloTmres, unlike true segments, are
not independently musculated (Imms, 1938). For this reason I join the
increasing number of students who use the term "flagellonnere" in the
placc- of "antennal segment." The antenna consists of a scape, pedicel,
and 1i. flag.llomeves, each of which is designated by a Roman numeral
(F'ip. 12A) In Syco'ax the terminal flagellimere is markedly reduced in
size (Fit. 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
The paired atitennal 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
The mouthparts of New World phlebotomines in relation to feeding
habits and taxonomy were studied in detail by Lewis (1975a) Follow ing
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
.,,ith a variable number of teeth in the Lutnomyjia and Brurnptojyia females.
Those in the latter genus are characteristically arranged in 4 longi-
tudinal rows (Fig. 10M). Cibarial teeth are absent in both sexes of
Wariltya and Sycorax and are poorly developed or absent in the Lutsondia
and Brmptomyjia males.
When viewed ventrally as in Fig. 12E, the horizontal teeth (= bind
teeth) of most Lutzotm7ia 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.
Tvon oth-er features of the clbarium are noted in the descriptions
and keys. The c:iitiinous arch (Fig. 12E), wh-n complete, crosses the
ventral wall of the cibarium. Its development varies according to species
and may be complore or not. The pigment patch on the dorsum of cibarium
(hig. 12E) varies in size, shape, and degree of infuscation (= pigmenta-
tion) in the Lu.zomyia spp.
The pharynx (Fig. 12C), composed of 3 chitinous plates, is attached
to the cibariun. Many species of Old World phlebotomines (genus
Sergentom ryia) have spines on the posterior part of the pharynx but they
arc relatively rare in the Lutsomyjia spp., occurring mostly in some
species in the cayennensts and oswaldoi groups. The majority of Lutzo-iia
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 (1967) as applied to the thorax.
The mesonotum should correctly include the nesoscutum, mesoscutellum, and
postnotum and should not be restricted to the mesoscutum alone (Saether,
The 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 I 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 opisternal setae) are present in species of Lutzo'ciya,
Br'',3ptcimia, and Sye orax. The Warileya spp. and the majority of Old World
phlebotemines lack such setae (Abonnenc & Leger, 1975). Additional
leoural setae, not mentioned in other studies to my knowledge, occur
behind the metathoracic spiraclc in all species of Phlebotominae and
Syecoaci.ic examined, those in the latter subfamily being 4 in number
and relatively stout fig. 2H1) 1ie 4 postspiracular setae in the
bhlebotomine species are reduced in site, barely visible (Fig. 12D).
The wing length is measured from the basal costal node (= rudiment
of tegula; Scodgrass, 1935) to its apex; its width at widest part (Fig.
120G. Ce-tain wing vein sections (Fig. 12G) are given useful, easily
remembered names. Alpha (a) is the length of R2 from its junction with
R3 to the costa. Beta (3) is the length or R from the junction of R4 to
the junction of R2 + R CGrma (y) is the section of R from the r m
crossvein to the junction of R5. Delta (6) is that part of R distad of
the junction of R and R3, It is negative when R1 ends before rhis
The legs are measured as in Fig. 12H. A few phlebotcanine 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 Lutzomyia.
The station 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 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 and 3,
is usefu? in associating male and female sand flies (liertig & Fairchild,
1950) but the character state is variable among individuals of conspecific
p;.p)ulztions (Iorattini, 1954).
The structures of the bilaterally symmetrical male genitalia,
lJhbelnd i. Fig. 2 a-ed 12. are extremely ir.portant ir systematics. After
eclosion, the genitalia rotates 1800 in the phlebotoaine males but not in
thoseir of Syo'r.. The style, attached to the apex of the coxite, bears
1 to 6 major spines* and C to many additional snwall setae. The presence
of numerous strong spines is presumably a plesiomorphic feature (Theodor,
The coxite is the dorsal basal appendage in the phlebotomine males
and, of course, is the ventral basal one in the Sdlcorax males. It may or
may not have a setal tuft or other nondeciduous setae on its inner sur-
Cace. These setae are those which remain on the coxite after maceration
aid which usually differ in size or shape from other setae on the
The fused, paired aedeagi, each of which is called the aedeagus is
subtriangular and well sclerotized in most phlebotomLne males. Some
species (e.g. L. walker) have aedeagi with dorsal projections. The
aedeagns of the Colombian Siqcorax 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 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
ter- referring here to size. One author's interpretation may differ from
that of another. In doubtful cases, the illustrations should be studied
to und c sand the meaning of statements in the text.
The phlebotomine parameres lir: between the coxites and ventral
lateral lobes when viewed laterally. Their shape and station 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. A4H. The cercopods of the Syc-orax males appear to
correspond to the parameres of Phlebotominae and are very complex.
The ventral, nonsegmented lateral lobes are more or loss uniform in
the phlebotomine males, although their length, width, and station 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 !;ize and structure of the internal paired speimathecae and their
ducts (Fig. 12F) are particularly important in the classification and
identification of female sand flies. An idea of the variety of sperma-
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 MATERIALS
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.
Battf.ery pcwcred 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 I trap per week was operated
throughout the night (1800 hrs.-0700 hrs.), we collected a total of 23
phlebotomine spp. (378&6. 78999) from April to Dec., 1967 (39 trap nights).
We recorded 36 sand fly species from Curiche based on all collection
Shannon traps (Shannon, 1939) made from muslin bedsheets do not trap
insects per s6e but are collecting devices similar to those used by
lepidopzcrists in "sheeting" for moths at night. A gasoline lantern pro-
vides 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
Jight !Lnd Shannon trap captures is generally similar.
Aa'.[se traps (Townes, 196?) and flight brap (similar to the design
of Grassi't and Gressitt, 1962) were placed in forest clearings, across
tr;: is -t oth:-Il 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,
then traps 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
ground level, one Malaise trap captured 24 Phlebotomine species (3166s,
48299) 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 Jn 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
I830 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 ;ere preserved dry in cardboard pill boxes rather than in
alcohol to prevent hardening of muscle tissue.
*One trap day equals 24 hours.
For routine identification of phlebotomiine and sycotiacines, un-
dissecLed 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 lacerating them at
room temperature for 12 hours or more. The flies should be left in the
hot NaOH for 3 minutes, then placed directly into a drop of 80-95% liquid
phenoi (C 6H5OH) 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
fIIl is ca- be discarded, preserved in vials of 70% alcohol, or pLocesEcd
further for slid mounting. I follow the procedure of Fairchild aud
Hoitig (1948c) except that Canada balsam or Euparal is substituted for
Other slide-mounting procedures are discussed ty Osorno et sl. (1966),
Quate & Stoffra (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 phan-; 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
Specimens were drawn with the aid of a camera lucida and Bausch and
Lomb miTroprojector. I measured specimens with a calibrated ocular
ficromtetpr, 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 drought, popu-
latoans 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
Sycoraciiae. 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 (llanos). Within each region there are one or
more life zones, each defined by a combination of bioremperature, annual
li-.;ipitation, humidity, and vegetation (Holdridge, 1967). The life
.onrs in Colom';ia were studied by Espina] and Montonegro (1963). Their
ucs.que ouredo" translates to "tropical moist forest" (2000-4000 mm of
r-tin per year) and "bosque muy humedo" corresponds to "tropical wet
foref-s" (4000--8000 mm of rain per year). The "bosque pluvial" (= tropicall
rain forest"), tne wettest lowland life zone, receives over 8000 num of
rai. p-'r year.
Figure 1 depicts where phlebotomines and sycoracines have been col-
lectted in Colombia. Osorno et al. (1972a) provide specific data for most
l ai ties; others from which collections were made by me or my colleagues
arc- discussed below in relation to faunal regions (D'Allesandro & Barreto,
1971) and distribuLion 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"
tc a gradual accumulation of isolates in the trar s-Andean forest region.*
T"ie 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 (c&s-Andean region). Most of these reached the
*The btr wn-Andean region comprises "Middle America and the narrow Pacific
lowlands along the western base of the Andes" (Haffer, 1967). Cia-Andean
refers Io lowland forests east of the Andes.
Pacific coast forests by advancing around, not across, the northern Andes.
Ihese species which now occur on both sides of the mountains include:
L;uonqiea ayrozai, L. gu~anensir, L .biopin;sa, L. cN7ezi, L. nordestina,
L. seirna, L. pilosa, L. spinosa, L. trinidadinsis, L. shannoni, L.
tulacuvata, L. trinuelac, and others.
Nine of the 41 t-pecies and subspecies do nol occur in other regions
of the Republic. Of these, only LutAotWaia 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: BrunptomCyia
habjata, Warileya nigrosacculus, L. rebutra, L. recurva, L. isovesperti-
lionis. L. ylephiletor, L. sanguinavia, and possibly L. odax. Two forms
which probably evolved in Pleistocene forest refugia within the trans-
Andean region include L. carrerai thula n.spp. and L. barrettoi majuscua
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. reburra -- a species whose closest allies occur entirely in
the cis-Andean region, especially in Amazonian forests.
A few species such as L. vespertilioni- and its sister species,
A. isoveperit-ioris probably invaded the Pacific cost region of Colombia
from Central America, the former species also occurring east of the Andas,
at least as far south as Bolivar Department.
departmentt of Chocc
'Tirea collecting sites within this Department were described by
Eldridge & Fairchild (1973) and Eldridge et a]. (1973). Maps accompany
Cur'iche. On narrow coastal plain between western slopes of Serrania
del Baudo and Ilumboldt Bay on the Pacific coast; "tropical wet forest;'
e.levrtion* less than 10 m. Sand flies were collected from April 1967, to
Dec. 196], in disturbed and undLsturbed evecgreeu forests .ad near a
Alto 'uricho. 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.
Department of Valle
An(ichaya Dam. About 35 km inland from Pacific Ocean, on the Rio
Anchicaya; "tropical wet forest;" elevation ca. 560 m. This site is
saillar to that near the Rio Anori (Antioquia Dept.) in terrain, elevation,
*Elevation is expressed here and elsewhere in meters above sea level.
andt cIL!riate. Sycora.cines were recorded fror., both localities and from one
other in C;.ucr Department.
265 k: E of Buenaventura. "Tropical rain forest" according to Espinal
(196;S); elevation ca. 50 m. Phlebotomines were captured in a patch of
disturbed forest surrounded by cultivated fields.
Atlantic Coast Region
Thi-' 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 Lhe Choco center to the west" (Haffer, 1967). The present life zones
are more 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 (Haffer, 1967,
1974). Lutzoymjia strictivitZa may be endemic. Lutzoryia yuilli and
L. nocticola are cis-Andean taxa. The former reaches its northernmost
limits in northern Colombia; whereas L. nocticola invaded extreme western
Panaea but populations between there and northern Colombia have since
Department of Antioquia
Rio Anori (70 19'N; 750 0''W), about 24 km SW of Zaragoza; "tropical
wer forest;" elevations from 500-700 m. Located in the foothills of the
central Cordillera of the Andes, this site is mostly well forested with
step. well (drained slopes. A detailed description is given by Porter
& "Defoliiar- (MS).
Department of Magdalena
Three sites near Santa Marta yielded sand flies from tree trunks or
under loose bark of standing trees in cultivated or semicleared areas
Rio Don Diego, E of Santa Marta within 1 km of Carribean Sea; "tropi-
cal wet forest;" elevation ca. 3 Tm.
Pretzoe n S 'nt-a Marta and Mi -na; near a small stream; probably a
transition zone between "tropical wet forest" and "tropical dry forest;"
elevation ca. 80 m.
t4inea, SE of Santa Marta; life zone indeterminate but similar to
preceding site; elevation 200 m.
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 Lutzormyia species
in the verra'carum and vexator groups dominate the rather depauperate
Department of Valle
Three sites W or SW of Caili -- 1. Near Pichinde, 2. In forest at the
television tower, just E of Salidito, and 3. Near Rio Punce, SW of Call;
"'lower montane very wet forest;" elevations from 1570-1700 Ii. Limited
collections fro- 31 July, 1973, to August, 1973.
DcoartmenL of Cauca
faster slope, near summit of Cerro Munchique, west of Popoyan;
"lower uioteane rain forest;" elevation ca. 2500 m. Briefly surveyed on
5-6 August, 1973. One Sycorax sp. taken but no phlebotomines.
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 Uhich 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.
Phleboto.m.ne 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.
.ntuc'd.ncia of Caquota
"rs 'I,^qu.':iZ ; transitional life zone, "tropical moist forest"/
"rrop:ical waot forest;" elevation ca. 200 m. Using light traps, Dr. C.J.
M.arinkelle collected numerous phleboto.mines in November, 1971.
Map of Colombia showing localities where phlebotomies 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.
o = Phlebotomine record
X = Sycoracine record
4= Phlebotomi[ne and sycoracine record
1 = Atlantico Dept.
2 = LMagdalena Dept.
3 = Norte de Santander Dept.
4 = Caldas Dept.
5 = Risaralda Dept.
6 = Quindio Dept.
7 = Cundinamarca Dept.
.... .... ....
c: f, .- ....
-a Chic z O
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
tave caused a great deal of human suffering in the neotropics and
Leishmaniasis, a collective term referring to several disease
entities caused by Leishmania spp., occurs n Colombia but the vectors
are poorly known. Reyes (1957) provided limited information on the dis--
tribuAtion 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
FRepublic 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 Cundinamarca Departments having been discussed by Arjona ec al.
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
&ere 2,24L cases eith a mortality rate of nearly 16% (Samaniego, 1944).
The disease, also known to occur in Peru and Ecuador, was probably intro-
duced into Colombia "by returning soldiers or by 'colporteurs' who travel
froi'm village to village" (Rozeboom, 1947b) Circumstantial evidence led
investigators to believe that Lutzonia coleiTmbiana was the responsible
Arboviruses were recovered from wild caught phlebotomines in the
Pacific lowlands of Colombia (Barreto, 1969). The strain, designated
Co Ar 3319, was discussed 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.
Following Duckhouse (1972) 1 presently recognize 3 extant genera in
th- subfamily -- Syoorax 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
Par-ayoorai satchelli (Barretto, 1956) from Sao Paulo State, Brazil,
ApoeUysorax chilensis (Tonnoir, 1929) from Chile, and an unnamed sycoracine,
the wing of which was figureC4 by sairchi-ld (1955) from Palenque, Colon
The feeding habits of most species have not been studied but the
females of Syoorcy and Aposycorax at least possess mouthparts adapted for
sucking blood. Sycorax silaoea Curtis, the type species of the genus from
Europe, feeds on frogs and transmits a filarial worm to them (Desportes,
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). Aposyoorax 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
U .r c.ra'x ,pecies distinguish them from the extant species in the sub-
Other features, not always repeated in the specific descriptions
which follow, are shared by the 4 Colombian Sycorax species. Coloration:
Dusky brown, miesonotum but slightly darker than pleuron. Head: Broader
than long; frons pufftted 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
siza' and terminating in a cone shaped peg sensilla (two similar sensilla
on flagellomere 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;
par.n ieres somewhat resembling those of A. chilsen;is but differing in the
[acu-e and position of the lobes, setae, and projections. Female geni-
talia: Each spermatiheca and sperm duct forming a continuous tube with
faint transverse striations, terminating in a heavily sclerotized "button"
which nppeasrs shiny black; a tenuous sac usually amorphic follo.''ing
treatment in KOH, arises from the end of the button; an internal apodeme
or furca associated with paired spermatherae.
The combined length of the spermatheca and duct varies interspeci-
fically an:( p-obably correspondsi to the length of the male genital
filaments. This feLture and the distribution of the antennal setae and
distance between the compound eyes were also used in associating the
se-xes 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
Keys to the Syeorax Species
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
2. Aeoceagal die's relatively loug, exceeding the length of the sperm
pump. A subtrlangular plate, dorsovenrrally flattened, with acute
biFur-at a 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 ducts shorter, less than length of sperm pump and not as
sinuous. Subtriangular plate absent. Tip of median process of
cedeagus angular in lateral view. . . . ... fairchildi (Fig. 4)
3. Style of male genitalia with 4 strong spines; median process of
aedeagus more slender (Fig. 2G). A larger species, wing length greater
than 2 mn . . . . . . . . . . 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)
1. Flagellomeres I-VIII with paired ascoids, other antenna 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
2. Fi.rca spede-shaped apically, its length at least s that of sperm duct
+ speratheca. Flagellomare, 1, 0.20 .m, or longer . . . . .
. . . .. . . . . . . . co o i rsis (Fig. 3)
Furca "v" shaped apically, its length but slightly less than that of
sperm duct + spemrratheca. Flagellomere I less than 0.20 mm long
. . . . . . . . . . . . fa rchildi (Fig. 4)
1. Sycorax andicola n. sp.
Male holotypee): Wing length 2.35; width 1.05. Head height 0.31;
widL;. C.. Eyes separated by 0.14 or by distance to 8 facet dia-
meters. Antenna (including scape and pedicel) 1.61 long; Elagellomere I
(0.26 long), about 1.8 length of flagellomere II; 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.
Stryle 0.18 long with I terminal spine, slightly beveled at tip and with a
proximal cow of 3 smaller spines. Parameie as shown with several setose
libes aind projections, the most distal finger-like with rounded, upturned
tips. Aedeagal complex of a laterally compressed sperm pump (0.20 long)
within the abdomen; n median distal process (0.165 long) compressed dorso-
ventrally, curved and slender in lateral view, upturned at tip, basal
t-o-thirds much wider than distal one-third when viewed dorsally; paired
sinuos, relatively long genital filaments. Cerci as figured.
Xateri-al cxarined: 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. dparatype (no. 538), same data except collected
by R.C.W., 8 Aug. 1975.
Discussion: S. andicola, the largest Sycorax known from Colombia,
closely resembles S. tinispinosa n. sp. in details of the male genitalia,
station of the antenna and in other character states. The two species,
apparently allopatric, are separated by the characters given in the key.
2. Sqcorax colombiensis n. sp.
mate holotypee): Wing length 1.i2; 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 length 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
holotype. Wing venation as figured. Legs missing. Nature of sternites
not determinable. Cenitalia: 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
flattened, with acute bifurcate tip, pointing to the rear, present below
the .ase of the median process nf aedeagus. Cerci as shown.
Female (ailotype): 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 length 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.
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. d' holotype (no. 529), Anchicaya Dam
(Valle), elev. ca. 400 m above sea level, light trap, 28 Jan. 1975, R.C.W.
? allotype (no. 530), same data as holotype except collected 10 June
1975. Paratypes (nos. 531-538), all from type locality in light traps;
1 T, 11 Aug. 1973, D.G.Y. and R.C.W. 1 6', 2 oo, 28 Jan. 1975; 2 oo, 10
June 1975, R.C.W.
discussion : The male of S. cotombiensis, the only Colombian species
with a 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. coZom.biensis share
several character states with S. fair'chiLdi and S. trispinosa but may be
identified by the characters in the key.
3. SZcorax fairchild[i sp.
!oZice holotypee): 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 'I; paired ascoids visible only on flagello-
meres I-IIi, uther sensory hairs in a whorl-like pattern on flagellomeres
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 as tip. Aedeagal
complex: Sperm pump (0.11 long); median disLal 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
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 length of flagellomere
II; paired ascoids, subequal in size (but not conspicuous) on flagello-
meres i-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
spisLernal setae (n = 3). Wing venation as shown. Length of femora,
tibias, and basitarsi: Foreleg, 0.39, 0.40, 0.22; iridleg, 0.42, 0.47,
0.24; hindlcg, 0.47, 0.50, 0.20. Sperinathcca + sperm duct length about
0.03 from base to end oi subcircular button: furca somewhat "v" shaped
Matetz'al exaarined: 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. y allotype (no. 540), same data except collected
28 Jan. 1975, R.C.W. Paratypes (nos. 541-547) all collected in light
traps, 1 3, Rio Anori (Antioquia), Colombia, elev. ca. 410 m above sea
level, 22 Sept. 1970, D.G.Y. 1 d', Rio Anoci, 23 Sept. 1970. 1 o,
Anchicaya Dam, 11 Aug. 1973, D.C.Y. and R.C.W. 3 MS, 1 T 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 Sycorax in Colombia.
I take pleasure in naming the species in honor of Dr. C.B. Fairchild
who has contributed so much to our knowledge of medically important
arthropods in the neotropics.
4. Sycorax trispinosa n. sp.
.Vale holotypee): 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 length of flagello-
mere II. Paired ascoids on flagellomeres I-IV, absent from remaining
flagellom0eres; other antenna setae mostly deciduous, not arranged in a
whorl-like pattern. Palpal length, 0.12 mra; ratio of segments: 1-1.15-
0.8,`-0.76, second segment with 15 rod sensilla. Pleura with 9-16 setae
below ving base ane 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
spenn 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
fact diameters. Antenna, 1.0 long; flagellomere 1 (0.16 long), about
1.7 x length of fl.agellomere 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
MaIteial examined: Colombia. & holotype (no. 548), Anchicaya Dam
(Valle), elev. ca. 400 m above sea level, light trap in forest, 5 March
1976, R.C.W. T allotype (no. 549), same data. Paratypes (nos. 550-573),
all collected from type locality in light or flight traps by D.G.Y.
and/or R.C.W. 2 o9, 10 Aug. 1973; 1 6d, 1 o, 11 Aug. 1973; 13 oSo, 4 o,
28 Jan. 1975; 1 ', 2 oo, 5 March 1976.
Discussion: I associated the male and female of S. trispinosa on
the basis of the following characteristics which, in combination, are not
shared by S. fairehiidi or S. cotomrbiensis. Flagellomere I relatively
short (0.14-0.19 mm, n = 27); compound eyes more narrowly separated than
those 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
am)dicola, presently unknown, probably resembles trispinosa in details of
thie sparmathecae and palpi.
The specific name refers to the 3 strong spines on the style of the
S5'oroz oRndiola 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
Male: Ccrro Munchique, Couca Dept., Colombia
SyCoraix colorabiensis -- 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.
Male: Anchicaya Dam, Valle Dept., Colomba
Female: Same locality as male
Sycoa.r fairildid -- 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 flagellomeres I and II showing
paired ascoids and other setae.
Male: Anchicaya Dam, Valle Dept., Colombia
Female: Same locality as male
Jl -,2 -
A ___ ^
N\. 0,. <._<
-< -'/ / ""c
\. (> 1.7n
~ t E
K IH \
1: (-K .<
SyCorPt: t"i.spizosa -- A. Male head, B. Female palpus, C. Female,
t-rminal 4 flagellomeres, D. Female pedicel and flagellomeres T and
II showing ascoids, E. Female head, F. Female wing, G. Male wing,
H. Kale genitalia, dorsal view, I. Sperm pump, genital filaments,and
median dorsal process of aedeagus, lateral view, J. Tip of furca,
dorsal view, K. Spermath1caeo
Male: Anchicaya Dam, Valle Dept., Colombia
Female: Same locality as male
; of c. *-.
." <" : '.^. ,,- ..._ _'' c-s ^ r.-
^ f ]
"- : !'"
Previous studies on this subfamily in Colombia were reviewed by
Osorno et al. (1972a) and will not be repeated here except for literature
citations in the species accounts. Nearly all of these studies involve
vsstemntics, especially species descriptions and distribution records.
Owing to the paucity of bionomic studies on the Colorbian phlebotomines,
I felt that it would be desirable to provide references to such studies
carmied 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
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 . . . . . . . . ... Wari.eya (Fig. 6-7)
Wing pointed at tip; R2 + R3 + R4 forks well beyond r-m crossvein.
Pleura with episternal setae. Female cibnrium armed with teeth.
Male genitalia with style shorter than coxite . . . . . 2
2. Interocular suture complete. Female cibarium with 4 longitudinal
rows of horizontal teeth. Male genitalia with 5 large spines, 2 of
whizh (usually basal pair) borne on a common tubercle . . . .
. . . . . . . ...... . . . Brumnptomyria (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)
Genus ,arflea Hertig, 1948
This genus, recently enlarged to accommodate W. (H.) hertij (Fchld.)
by Lewis et al. (1978), contains 5 species, 2 of which in the subgenus
Warit.eya 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
Keys to Species
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. . . . . . . . . nigrosaccuius (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)
L. Speruatheca2 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. . . . . . . . . nigrosacculu. (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. . . . rotendipennis (Fig. 7)
5. Warileya (W.) nigros3;cuZus
Warilcya nigrosacculus Fairchild & Hertig, 1951b: 428 (n holotype.
terro Campana, Panama Prov., Panama). Fairchild, 1953: 102 (cf, to
Pertiyia 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 (5d, descr.).
Chaniotis et al., 1972: 95-96 (in tree hollows, Panama). Christensen,
1972a: 88 (listed). Forattini, 1973: 536-538 (T, 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 examined: Colombia: 1 6T, 1 T, Curiche (Choco). Malaise
trap, 27 April 1977. 1 e, 1 o, Anchicaya Dam (Valle), light trap, 28 Jan.
1975, R.C.W. 1 85, same data but taken 16 July 1975. Panama: o holotype,
Cerro Campana (Panama Prov.), hollow log, 7 Jan. 1947, M. Hertig. 2 Sf',
5 TV, near Gamboa, Canal zone, tree hollow, 10 Sept. 1969, B. Chaniotis.
4 5d', 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.
6. Warileya (W. ) rotundipsnnis
Wac-ileya rotzidipennis Fairchild & Hertig, 1951b: 424 (e holctype,
o, Cerro Campana, Panama Prov., Panama). Fairchild, 1953: 102 (cf. to
Pertigia hb'r-tigi). Barretro, 1955a: 188 (listed). Fairchild & Hertig,
1959: J22 (Costa 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 (6', 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 (class-if.).
Distribution: Colombia (Antioquia, Choco, Valle), Costa Rica, Panama.
Material examined: Colombia. 2 ??, Teresita (Choco), tree buttress,
15 June 1967. 27 oo, Rio Anori (Antioquia), light traps, Sept. 1970.
5 ?c, same data but 29 May 1970, C.H.P. 8 &', 2 oo, Anchicaya Dam (Valle),
light trap, 28 Jan. 1975, J.E. Browne. 6 dJ, 6 7o, same data but taken
10 June 1975, R.C.W. 5 3'd, 4 oo, same data but taken 16 July 1975.
Panama. S holotype (no. 2335), p allotype (no. 2312), Cerro Canpana
(Panama), Shannon trap, 24 Aug. 1950, M. Hertig & G. Fairchild. 2 s3',
Rio Changena (Bocas del Toro), Shannon trap, 8-11 Sept. 1961, R. Hartmann
& P. Galindo. Costa Rica. 2 Se, 1 T, 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. Dr. C.H. Porter also has collected man-biting
females at the Rio Anori (Antioquia) locality, mostly in forest clearings
The records of W. rotundipennis and 5'. rigrosacculus from Choco and
Valle Depts. indicate that both taxa probably occur in other areas of the
trans-Auidean region perhaps as far south as Guayas Prov., Ecuador.
'/ariicya nipzosacculus -- A. Hale head, B. Male flagellormere II,
C. Female head, D. Female flagellomere If, 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
(N 7 ~1 ~
; a I; I::
11 D E
N B ~
A N K 1/
~.z ~" ~
__ ---*---- I /
armieyaa rotwd.ipennis -- 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-
Male: Cerro Campana, Panama Prov, Panama
Female: Rio Anori, Antioquia Dept., Colombia
~ 7 Y ~\ ~(i K~'
B '~I i~
Genus Brwptorm'ga Franca and Farrot, 1921
Characterized by TlIcodor (1965), Lewis et al. (1978), and others,
the genus Brtu.ptomuia now contains over 21 species, 5 of which occur in
Colonbia. Except for the female of B. hapata which has very short sperm
ducts (Fig. 101), the females of the other species in Colombia are
virtuallv impossible to identify in the absence of males.
Rrzmrptosrjia avellari, one of the Colombian species, was reported
feeding on armadillos in Brazil (Mangabeira, J942b). This and other
Bramptmyjia 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
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. . ... avezllavi (Fig. 8)
4. Style with isolated spine well distad of proximal pair. Coxite tuft
implanted on a distinct raised tubercle. . ... beaupert'qyi (Fig. 8)
Style with isolated spine at or near level of proximal spines. Coxite
tuft implanted on a slightly raised, raspberry-like (circular)
ba e. . . . . . . . . . . . eopoldoi (Fig. 11)
7. Brwaptomyia avelluri
Fhtebotomus avellari Costa Lima, 1932: 48 (0, 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 galinloi). Barretto, 1951: 212 (dist.).
Floch & Abonnenc, 1952: 39, 45 (J, o, keyed). Rodriguez, 1953b: 55
(mention). ForattJni, 1954: 214-217 (second sternite, fig.). Forattini
& dos Santos, 1955: 17 (Brazilian record). Ortiz, 1963: 320 (o, keyed).
Brumptovmyia 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 (c? 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:
Distribution: Panaar, Colombia (Boyaca), Peru, Venezuela, Brazil,
Mate-al examined: Colombia. 2 Vd', 1 o, Puerto Boyaca (Boyaca),
light trap, 6 May 1973, C.J.M. BraziZ. 1 6, Coqueiros (Sao Paulo),
armadillo burrow, Oct. 1953, Albertin. Pancaa. 1 d, Tocumen Airport
(Panama), light trap, 23 March 1953, F. Blanton. Paraguay 5 6o6, Aca-
Poi, San Pedro, burrow, 18 April 1950, M. Hertig.
btisc2ei;on': Brh'nmptoryia awclicri and B. rzz,'pti (Larousse) with
sympatric populatLons 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
8. Brnmptoryia beaupertuyi
'Phlebotor.us beaupertui Ortiz, 1954: 235 (d holotype, Duaca, Lara
State, Venezuela). Scorza & Ortiz, 1960: 434 et seq. (ecology). Pifano
et al., 1962: 383, 387 (o, q, keyed), 411-412 (6, ,, descr., fig.).
Ortiz, 1963: 322 (o descr.). Ortiz & Scorza, 1963: 350 (listed), 354
(0' keyed). Ortiz, 1965a: 208 (mention). Scorza et al., 1967: 193,
195 (6, keyed). Leon, 1969: 30 (listed).
Phlebotomnus galindoi (not galindoi Fairchild & Hertig, 1947) : Pifano
& Ortiz, 1952: 138 (listed, Venezuela). Leon, 1969: 30 (listed).
Brunentomyia beaupertuyi: Barretto, 1955a: 187 (listed). Sherlock,
1962: 321 et seq. (3', 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 Jo' (topotypes), Duaca (Lara), no
date, 1. Ortiz.
Discu..uhsion: 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
9. Bruaptcr!'ia yali'.doc
Phlebotoams galinroi Fairchild & Hertig, 1947a: 615 (S holotype,
near hBquete, Chiriqui Prov., Panama). Floch & Abonnenc, 1952: 40 (6
keyed). Ortiz, 1954: 238 (cf. to beauperthyi). Rosabal, 1954: 30 er
seq. (5 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 (6'keyed).
Biagi at al., 1966: 149 (Mexican records), 151 (c fig.). Strangways-
Dixon & Lainson, 1966: 193 (Belize). Hanson, 1968: 56-58 (larva, pupa,
Bruptozmyia 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.).
BpYrptomyia mesa Sherlock, 1962: 332 (dr, 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).
Di-.sribu tion: Mexico, Belize, Honduras, Costa Rica, Panama, Colombia
(Caqueta, Choco, Bolivar, Boyaca), Ecuador, Paraguay.
Material exceined: Colormbia. 1 6', Puerto Boyaca (Boyaca), light
trap, 6 May 1973, C.J.M. 1 6', Tres Esquinas (Caqueta), light trap, 11
Nov. 1971, C.J.M. 27 S6, 11 ?q, 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 6',
4 w, Rio Napo at Limoncocha (Napo), light & flight traps, tree buttresses,
20-24 May 1976, D.C.Y. & T. Rogers. Horndwas,. 1 d, Tela, Lancetilla
alJ.ey, tree buttress, 24 July 1953, W. Iils. 1 (6, same data but light
trap. 13 Jan. 1934. iexico. 2 (To, 2 ?y, Ocoscoautla (Chiapas), tree
hollows, 8 April 1951, G. Fairchild & R. Hattnann. 2 63, Palenque
(Chiapas), tree buttresses, 30-31 March 1951, G. Fairchild & R. Hartmann.
Paurafuay. 1 6, Sommerfeld, Yhu, tree cavity, 15 March 1950, V. Zelada.
1 d', same data but 18 March 1950, M. Hertig. Panama. 1 o' holotypee no.
814), type locality, 17 Feb. 1947, P. Galindo. 1 T, same data but 26
March 1948. 1 d6, 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 rmm, from Chiriqui Province, Panama (ca. 1200 m above
sea level). Smaller specimens from lower elevations in Santander Depart-
ment, Colombia were later named B. rresai by Sherlock (1962) who separated
the males on size, number of distal setae on the coxites, shape of the
aedeagi and station 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
galZindoi 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 alLitude, i.e., is there a continuous increase in size of specimens
from the lowlands to the highlands? Believing this to be the case but
without evidence to support i t, I tentatively consider B. mesai and
B. ialindoi to be conspecific following Fraiha et al. (1970a).
10. Brumptomyia hcramta
Phlboto mn s hawnatus Fairchild & Hertig, 1947a: 614 (1 holotype,
Chilibrcil.o bat caves, near Chilibre, Panama). Barretto, 1951: 217
(mention). Floch & Abonnenc, 1952: 39 (1 keyed). Rodriguez, 1953b: 53
(6, o measured, Ecuador), 55 (cf. to Zeopoltdi). Ortiz, 1954: 237
(listed). Lewis & Garnham, 1959: 83-84 (d, 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 (6,
keyed). Disney, 1966: 449 (l'e1 in rodent-baited traps, Belize). Hanson,
1968* 58 (larvae identical to those of galir.doi).
Bruwptomyji hamata: Barretto, 1955a; 187 (listed). Sherlock, 1962:
332 (cf. to mesai). Williams, 1970: 331 (listed, Belize). Fraiha et
al., 1970a: 468 (3 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: Mexico, Belize, Panama, Colombia (Choco), Ecuador.
Ia2teri'al examined: Colombia. 1 e, Teresita (Choco), hollow tree,
25 March 1967, D.G.Y. 1 ? same data but Malaise trap, 17 June 1967.
Panawia. 1 6 holotypee no. 112), type locality, 4 Dec. 1943. 4 So, 5 p,
lab reared from larvae collected in Canal Zone, 1957-1958, W. Hanson.
Dirc ussio: Although the adults of B. hanaata are rarely encountered,
t'ha larvae may be quite common in soil 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
microhabttat. Like other Brzpcpto7tia spp., the fourth instar larva has
only two caudal setae.
11. Brunptov yia leopoldoi
PhZebotomus leopoldoi Rodriguez, 1953b: 52 (d"holotype, Quevedo,
Lcs Rios Prov., Ecuador; 9, Naranjal, Guayas Prov., Ecuador). Ortiz,
1963: 320 (7 keyed).
BrT-mptomyia leopoldoi: Barretto, 1955a: 187 (listed). Sherlock,
1962: 332 (cf. to mesai. Williams, 1970: 331 (Belize). Fraiha et al.,
197Ca: 468-469 (d keyed, as possible synonym of guimaraesi). Forattini,
1971a: 98 (listed). Christensen, 1972a: 88 (listed, Panama). Forattini,
1973: 530 (as synonym of guimaraesai). Williams, 1976a: 603 (in caves,
Distribution: Belize, Panama, Colombia (Antioquia, Choco, Valle),
Material examined: Colombia. 3 S'5d, Rio Anori (Antioquia), light
traps, May 1970, C.H.P. 1 3, Curiche (Choco), light trap, 26 May 1967,
D.G.Y. 2 Vd', 5 QQ, 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 S (paratype no. 4314), Quevedo (Los Rios), no date,
J. Rodriguez. 2 (ro, 2 yo, 17 km !: of Santo Domingo de Los Colorados
(Pichincha), mammal burrow, 4 May 1976, D.G.Y. & T. Rogers. 1 6', same
data but 2/ May 1976. 1 e, same data but liht trap, 27 May 1976.
Panama. 1 ,V (paratype no. 4292), Almirante (Bocas del Toro), animal
burrow, 22 Jan. 1953, W. Hils. 1 ', same data but tree buttress, 27 Jan.
1956, R. iartmanu.
Discaistaon: 3. leopoldoi, considered to be conspecific with
B. guimaraesi (Coutinho and Barretto) by Forattini (1973), is specifically
distinct based upon the following considerations:
1. Thp nature and size of the basal coxite tufts and the structure
and station 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 norn-l structure. Paratype no. 4314 is obviously flattened.
2. The genital pump of the guimaraesi male is less flared and
notably shorter (0.12 mm-0.15 me.) than that of Zeopoldoi which
is 0.22-0.24 mm long (cf. Fig. 11G and 11H).
3. The isolated spine of the style of :uirmaraesi is inserted at
0.59-0.61 of the segment whereas that of Zeopoldoi is located
In addition to the leopoldoi specimens listed above, I examined 4
guimararsi males from southern Brazil: 2 d'd, Faz. Ribeirao de Baizo,
Patos, Minas Gerais; 1 d6, Coquejros, Sao Paulo; 1 cd reared from egg laid
by $ captured at the type locality (Itaporanga, Sao Paulo, Brazil). I
conclude that these allopatric species are distinct, B. Jg"imaraezsi
occurring only in southeastern Brazil as far as known.
3-ruwtomy .la a.ellari male -- A. Genitalia.
Male: Puerto Boyaca, Boyaca Dept., Colombia
BhR'vrtowyia becapertuyi male -- B. Head, C. Flagellomere II, D. Parainere,
E. Wing, F. Genital pump and filaments, G. Genitalia, H. Coxite tuft,
.ame scale as rig. 8A.
Mile: Lara State, Venezuela
prwr,po'nyia gatindoi -- A. Male head, B. Male flagellomere 11,
C. Female head, 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
spermnatheca, I. Female wing, N. Male wing, 0. Female cibarium.
Male: Teresita, Choco Dept., Colombia
Female: Same locality as male
C- - - -
BVrnWotomyia hanata -- A. Male head, B. Male flagellomert 11, 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 cibariumn
Male: Barro Colorado Island, Panama Canal Zone (except Fig. 10F)
Female: Teresita. Choco Dept., Colombia
Brvmptomya 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, h. Genital pump and filaments, F. Genitalia,
G. Genital pump of male from Fig. 11C locality.
Male: Hio Anori, Antioquia Dept., Colombia (except f-r Figs.
11C and G)
Brumptomyia guimaraesi male -- I. Genital pump, I. Coxite tuft, paramere
and aedeagus, same scale as 11C.
Male: Itaporanga, Sao Paulo State, Brazil
Genus LutKasia Franca, 1924
As mentioned earlier the classification of the Lutzonyia sand flies
is hasfed 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.
'eys_ to Subgenera, Species Croups, and Ulngrouped
Species of Lutzon, jia
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 Lutszomyia subgenera and equivalent species
groups are defined by the characters given here. It is important to
nore, 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 ver rcoarum 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.
As additional species are discovered in the Republic, these keys
obviously will have to bh revised. For the present, they are intended
to be used with caution, applicable only to the Colombian fauna.
1. Palp 5 very short, less than half length of third and less than twice
length of fourth. . . subgenus Pse'uhodopygus (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 squamiventr-i (Fig. 61)
St)le 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 .vsspertilionis group (Fig. 29)
Lateral lobes not inflated. ..... . . . . . . . 5
5. (4) Lateral lobes with spatulate setae at tips . . . . .
. . . . . .. subgenus Evadromio, series infra.,pin-oa^
*Species of Evandzomyia in this series have not been found as yet in
Colomribia 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).
Lateral lobes without sp.itulate setae . . . . . . 6
6. (5) Style deeply forked. Parameres with 2-4 modified apical setae as
shown . . . . . . . subgenus Viannaryia (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
state . . . . subgenus Damprolmia, L. rosabati (Fig. 30)
Paramnere 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
a small subterminal and a small
spines. . . . . . .
Coxite without a basal tuft but
setae. Style with only a small
subterminal seta. . . . .
simple and modified setae. Style with
median seta in addition to the major
. . subgenus Pressatia (Fig. 32-34)
with a distal group of 15+ simple
median seta and 3 major spines, no
. 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. rovotaensis usually have 5 major spines but those examined
from Valle Dept., have but 4 (see p. 421).
12. (11) Antennal ssc(,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 distaj setae. Style with isolated basal
spine . . . . . . . . dreisbachi group (Fig. 50-51)
Coxite without distal setae. Style with basal spines more or less
paired. . . . . . . . . shnnoni 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 flagellomeres
II + III. . . . . . .subgenus Trhihophoro,,yia (Fig. 51-54)
Style with paired basal spines. Antennal ascoids simple, no basal
spurs visible. Fingellomere I longer than II + III . . . .
. . . . .subgenus [.yssomYia (in part), L. actunesi (rig. 55)
17. (15) Atntennal ascoids with short, pointed basal suprs. Paramere with
r dorsobasal hump. Flageliomere I shorter than II + III . . ..
. . . . . . . . . . aragaoi group (Fig. 45-49)
Aitrennal ascoids simple. Paramere lacking a dorsobasal hump. Flagel-
lomera I longer than II + III . . . . . . . . . .
. . . . . . subgenus N .ssomyia (in part), (Fig. 55-60)
I8. (14) Coxite with 3+ basal and/or median setae in a tuft or group,
discal setae may also be present. . . . . . . . . 19
Co-x zie without a basal and/or median group of setae but long ventral
etae may be present. . . . . . . . . 2S
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 . . . .baityi 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 LutW'aylia, series longipalpi- (Fig. 13-15)
Paramere without such sotae . . . . . . . . 22
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" . rrmigonri group, series walkeri (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 coxita . 25
25. Mesonotum & pleura pale . saulensis group,* L. sauZensis (Fig. 21)
Mesonotum distinctly darker than pleura . . . . . . .
verru oarum group, series verricar w (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 such base. 27
27. (26) Style with paired basal spines. Coxite with basal tuft of
slender, subequal setae . . . . . .. . . . ... ver-
rucarum group, series verrusaram (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
sotae . . . .eaynnenois group, series at-eoclavata (Fig. 81-82)
*T im unable to separate the males on the group level using structural
28. (18) Style jith a small subterminal seta. . . . . . . 29
Style without a small subterminal seta. . . . . . . 31
29. (28) Paramere divided. Coxite with long ventral setae. . . .
.. ..................... onispir.a 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. Flagellomere I longer than
head height; ascoids with very short basal spurs. Palp 5 shorter
than segments 3 + 4 . . . ungrouped, L. rordestina (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. . . . . . .
. ........... v.oxator group, series peruenwis (Fig. 72-78)
Coxite without setae at base but with a few scattered setae near
,.Ldit of coxite. . . ... .csuwadoi group (in part), (Fig. 84-85)
1. Spermathecae associated with paired sclerotized structures, intracel-
lular ducts (= "hairs") covering most of spermathecae . . . .
. . . . . . . . . subgenus Viannrzajyia (Fig. 35-36)
Spernathecae without associated sclerotized structures, intracellular
ducts confined to small area, usually but not always, to terminal
knol' . . . . . . . . * . . ............ 2
2. (1)Pharynx with posterior spines. . . . . . . . . 3
Pharynx without spines. . . . . . . . . . . 6
3. (2) Cibarium with inner pair of horizontal teeth pointing inwards
. . . .osualoi 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 eaeennensis (in part),L. cayennensis (Fig. 79)
Cibarium with 4 horizontal tech not in a comb-like row . . 5
5. (4) Foreleg with femur longer than tibia. Pharynx markedly enlarged
with prominent, subequal ly spaced transverse ridges and conspicuous
spines. Pleura pale. . . . . . . . . . . . .
........ cayen'newf group, series atroclaoata (Fig. 81-82)
Foreleg with femur shorter than tibia. iharynx nore slender, the
tranasverse ridges and spines less defined. Pleura dark .......
zzcaenensis group, series ayeranre.is (in part), L. tmir-opyga (Fig. 80)
6. (2) Spermiathecae with bubble-like expansions. . . . . . 7
SperiGathecae without bubble-like expansions . . . . . . 8
7. (6) Cibariu-n with 4 broad and blunt horizontal teeth and conspicuous
lateral teeth. Mesonotum well pigmented, darker than pleura. Eyes
very small. . . . .subgenus Datpfomeia, L. rosabati (Fig. 30)
Cibarium with 4 slender, sharp horizontal teeth, without obvious
lateral teeth. Mesonotum and pleura pale. Eyes larger . . .
. . . . . . .. soauensis 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
annolated . . . . . . . drisabachi group (Fig. 50-51)
Cibarium with 4-8 horizontal teeth. Spermathecae annulated or not
. . . . . . . . . . hawnn.ni group (Fig. 37-42)
10. (8) Hind femur with row of 3-5 short spines . . . . . . .
. . . . . ... .subgenus PiLtosyia, L. spinosc (Fig. 31)
Hind femur without spines . . . . . . . . . 11
11. (10) Cibarium with 6 or more horizontal teeth . . . . . 12
Cibarium with 4 horizontal teeth. .. . . . . . 17
12. (11) Individual sperm ducts markedly convoluted, twisted. . . .
I . avaao'i group, series bra.i.liensis, 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. araCaoi 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 Nyssoomyia (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. 419).
d ts . . . . . . . . . . . . . . subge-
nLsL',tzcomlyia, series longipalpis (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 Psychodopygus (Fig. 61-71)
Spern.athecae 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 setae . . ............
. . . .. vespertilionis group, series vespertilionis (Fig. 29)
Spermathecae otherwise, shorter than individual sperm ducts. Ab-
dominal tergite 8 with lateral setae. . . . . . . 21
21. (20) Sperrmathecae less than twice as long as wide, terminal segment
smaller than preceding segment. Common sperm duct longer than in-
dividual ducts. . .. oe ualdoi group (in part), L. pia (Fig. 83)
Spermathecae cylindrical, its length at least 3 times width, terminal
segment subequal to or smaller than others. Common sperm duct
shorter than individual ducts . . . . . . . . . .
. vexator group, seriesperUensis (in part), (Fig. 72-74; 76-78)
22. (17) Tndividual sperm ducts heavily pigmented in part or whole. .
. . . . . . . . . .subgenus Pressatia (Fig. 32-34)
Individual sperm ducts not pigmented. . . . . . . . 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 . . . . . ..
. . . . . . . .pilos. group, L. pilosa (Fig. 86)
Cibarium with horizontal teeth otherwise. Common sperm duct
present . . . . . . . . . . . 25
25. (24) Wing venation with beta less than half alpha. Cibariumr with
horizontal teeth pointing towards pharynx, not inwardly slanted .
S. . .... vexator group, series pe-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. rorotaen ai's (Fig. 100)