Group Title: biosystematic review of the bloodsucking psychodid flies of Colombia (Diptera: Phlebotominae and Sycoracinae) /
Title: A biosystematic review of the bloodsucking psychodid flies of Colombia (Diptera: Phlebotominae and Sycoracinae) /
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 Material Information
Title: A biosystematic review of the bloodsucking psychodid flies of Colombia (Diptera: Phlebotominae and Sycoracinae) /
Physical Description: xiii, 481 leaves : ill. ; 28 cm.
Language: English
Creator: Young, David Grier, 1940-
Publication Date: 1977
Copyright Date: 1977
 Subjects
Subject: Sand flies -- Colombia
Psychodidae   ( lcsh )
Insects -- Colombia   ( lcsh )
Entomology and Nematology thesis Ph. D
Dissertations, Academic -- Entomology and Nematology -- UF
Genre: bibliography   ( marcgt )
non-fiction   ( marcgt )
 Notes
Thesis: Thesis--University of Florida.
Bibliography: Bibliography: leaves 451-480.
Statement of Responsibility: by David Grier Young.
General Note: Typescript.
General Note: Vita.
 Record Information
Bibliographic ID: UF00099259
Volume ID: VID00001
Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: All rights reserved by the source institution and holding location.
Resource Identifier: alephbibnum - 000012430
oclc - 03983178
notis - AAB5221

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A BIOSYSTEMATiC REVIEW OF THE BLOODSUCKING PSYCHODID FLIES OF
COLON-1BIA (DIPTlRA: PHL.EBOTOUMOLAE AND SYCORACfNYE)










By

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. . . . . . .


viii

xii

1

4

6

8

15

15
17










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 . . . . . . . . . . . .


Lu:tzomyia
Lutwmzoia
Lutzomyia
Lutzomyia
Lutzomrjia


(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








Page

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 .


. 167


. 171


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


i86


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


n. ssp.








Page

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


. . . . . 437
. . . . . . 439
. . . . . . 440


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

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

BIOGRAPHICAL SKETCH. . . . . . . . . . . .. .iR1


vii














LIST OF FIGURES


Feigur Pae

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








Figure Page

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




.ix










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










Figure


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)

By

David Grier Young

December, 1977

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

this review.

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).


xiii















INTRODUCTION


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.,

1972a).

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-

eastern region.

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).


--1-








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,

Bogota.














CLASSIFICATION AND TAXONOMIC TREATMENT


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

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

in continental Colombia. The species of 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

taxon.

An extensive bibliography and synonomy is given for most taxa treated

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

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

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

of discussions related to classification and distribution. For each

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

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

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

for full references.

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

to some readers are given below:

Institutions

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


Collectors

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

C.J.M. -- Cornelius J. Marinkeile

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

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


Other Abbreviations

ca.-- about

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

Staz Lted.

Although little studied, the immature stages are probably no less

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

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

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

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

species occurring in Colombia are cited in the appropriate species

bibliography.

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

associating conspecific males and females and for grouping taxa below the

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

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,






-10-


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

subspecies level.

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

particular flagellcmere.

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,

1571),

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

juncr ion.

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,

1965).

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

appendage.

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


Field Collections


For sampling phlebotomine and sycoracine populations, several methods

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

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

methods to capture these flies.

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

techn iques.

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.





-.1b-


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.






- I/-


Laboratory Methods


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

copal.

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

easily damaged.






-18-


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





-20-


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

their descriptions.

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

mangrove swamp.

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.





-23-


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

disappeared.



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





-24-


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

(August, 1973).

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.



Andean Region


Nearly all records of phlebotomine species occurring above 1000 m are

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

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

in the verra'carum and vexator groups dominate the rather depauperate

phlebotomine fauna.



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.



Oriental Region


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

Colombia is apparently rich in phlebotomine species but few collections

have been made there or in the more northern Llanos 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.





-26-



.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.

























Figure 1

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.

Legend

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.






-28


spa-







00

















.... .... ....

c: f, .- ....



-a Chic z O







Ecuador(




AroGizannig

C20c0!t -















MEDICAL IMPORTANCE


The phlebotomine sand flies, unlike the sycoracine flies, are

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

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

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

else-where.

Leishmaniasis, a collective term referring to several disease

entities caused by Leishmania spp., occurs 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.

(1971).

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

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

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





-30-


&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

vector.

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.















SUBFAMILY SYCOR^ACENAE


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

Prov.. Panama.

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,

1942).

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

share several characters which set them apart from the other

sycoracine species. In addition to the paired genital filaments between

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

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

the ducts and the proximal sperm pump (Fig. 2G). 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-

fam:ily.

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

been discovered.



Keys to the Syeorax Species


Mal es

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

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

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

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

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


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

median process of aedeagus with basal two-thirds

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

projection of paramere rounded at tip. Antennal sensory setae,

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

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









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)



Fl males

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.
(Fig. 2)


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.
(Fig. 3)


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.
(Fig. 4)


!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

as shown.

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

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

diameters. Flagellomere I (0.16 long) about 2.3 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

apically.

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.
(Fig. 5)


.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

shown.

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


mile.

























Figure 2

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
postspiracular setae.

Male: Ccrro Munchique, Couca Dept., Colombia








































ff 1




Ao


G0C

























Figure 3

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





















B >~
>~-~


Cr


I.


4.

,~~2 II,

H

























Figure 4

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















(T N_

Jl -,2 -

A ___ ^
IID
C




N\. 0,. <._<
-< -'/ / ""c

\. (> 1.7n
~ t E

K IH \


1: (-K .<

























Figure 5

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










W, ,-.



; of c. *-.


F
.~ -"-oryi.

." <" : '.^. ,,- ..._ _'' c-s ^ r.-
/*r- .
G
^ f ]






"- : !'"
..

/CF
\\% >1<

(^^^ G














SUBFAMILY PHILEIOTOMINAE


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

accounts.

With few exceptions I have not redescribed known taxa because of

adequate original or subsequent descriptions. Hopefully, the figures

and keys will serve to distinguish the taxa.



Key to the New World Genera


1. Wing broad, rounded at tip; R2 + R3 + R4 forks before, on same level

as, or slightly beyond r-m crossvein. Pleura without episternal

setae. Female cibarium unarmed. Male genitalia with style longer

than coxite . . . . . . . . ... 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


-50-





-51-



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

anthropophilic.



Keys to Species


Males

1. Palpal segment 5 longer than combined length of segments 2 + 3.

Paired intraabdominal rods present. Genital filaments about 1.8X

length of pump. Style lacking a basal bristle. Wing length less

than 1.5 mm. . . . . . . . . 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)



Females

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
(Fig. 6)


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.





-52-


6. Warileya (W. ) rotundipsnnis
(Fig. 7)


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






-55-


females at the Rio Anori (Antioquia) locality, mostly in forest clearings

(pers. comm.).

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.

























Figure 6

'/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





-57-






7
(N 7 ~1 ~
Ic\ C,

I
; a I; I::
U
11 D E


N B ~

A N K 1/




005 -
~.z ~" ~
F
~ _______

~

2

10 U



~ 7
"N

-ci ~-'
__ ---*---- I /
[I

























Figure 7

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-
thecae.

Male: Cerro Campana, Panama Prov, Panama
Female: Rio Anori, Antioquia Dept., Colombia












~ i

~ 7 Y ~\ ~(i K~'

i~l I
B '~I i~




1K









'\ %j~\\ /--~
GO


si/I
H C









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


Males

1. Genital filaments less than 4X length of pump. . hamata (Fig. 10)


Genital filaments greater than 4X length of pump . . . . 2


2. Coxite tuft of mostly stout, spine-like setae. . . . . . 3


Coxite tuft of slender, hair-like setae. . . . . . . 4


3. Coxite tuft of 20+ setae implanted on a distinct tubercle. Paramercs

more slender as shown. . . . . . . . galindoi (Fig. 9)


Coxite tuft of fewer than 20 setae, usually about 12, not implanted

on a tubercle. Paramere broader as shown. . ... 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
(Fig. 8)


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:

599 (Venezuela).

Distribution: Panaar, Colombia (Boyaca), Peru, Venezuela, Brazil,

Paraguay.

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

species.



8. Brnmptoryia beaupertuyi
(Fig. 8)


'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

specimens.








9. Bruaptcr!'ia yali'.doc
(Fig. 9)


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,

descr., fig.).

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
(Fig. 10)


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
(Fig. 11)


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,

Belize).

Distribution: Belize, Panama, Colombia (Antioquia, Choco, Valle),

.~uador.

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





-67--


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

at 0.53.

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.

























Figure 8

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



























- >3)


D


F.y


F





w


Th2.


2


I J






8

C


/
/
(3


N

























Figure 9

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






















B




U




-if
G

H












I L)
I*


MY

C- - - -
-~
NO


h





V

F I



/~~

V






I I


ii




I)
D ii
F

t\






N
L~- ;1~2)
~, ----a-

C 9~



N'

























Figure 10

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














fK\

.1"
r

\ i~7j


B


i~l
Y
4;
.2]

B


A.
1A


V
G '~~



F)









K
I 'N:f


10

~
C)
1K

-1::----)
2

























Figure 11

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






























B
03


tk,,
(J
2


7 -H
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It '2
~2


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'1''~i


.7 I.,


B C


--


C ____










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.



Males

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
characters.









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)





-a S-


Coxite without setae at base but with a few scattered setae near

,.Ldit of coxite. . . ... .csuwadoi group (in part), (Fig. 84-85)




Females

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


-84-









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).





-85-


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)




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