• TABLE OF CONTENTS
HIDE
 Front Cover
 Title Page
 Table of Contents
 Introduction
 Classification and taxonomic...
 General morphology and termino...
 Methods and materials
 Distribution, collecting localities,...
 Medical importance
 Key to the subfamilies of psychodidae...
 Subfamily sycoracinae
 Subfamily phlebotominae
 Synopsis of new taxa, synonums,...
 Reference
 Systematic index














Group Title: Bulletin - University of Florida. Agricultural Experiment Station ; no. 806
Title: A review of the bloodsucking Psychodid flies of Colombia
CITATION THUMBNAILS PAGE IMAGE ZOOMABLE
Full Citation
STANDARD VIEW MARC VIEW
Permanent Link: http://ufdc.ufl.edu/UF00027135/00001
 Material Information
Title: A review of the bloodsucking Psychodid flies of Colombia (Diptera: Phlebotominae and Sycoracinae)
Series Title: Bulletin University of Florida. Agricultural Experiment Station
Physical Description: 266 p. : ill. ; 23 cm.
Language: English
Creator: Young, David Grier, 1940-
Publisher: Agricultural Experiment Stations, Institute of Food and Agricultural Sciences, University of Florida
Place of Publication: Gainesville Fla
Publication Date: 1979
 Subjects
Subject: Psychodidae   ( lcsh )
Diptera -- Colombia   ( lcsh )
Genre: government publication (state, provincial, terriorial, dependent)   ( marcgt )
bibliography   ( marcgt )
non-fiction   ( marcgt )
 Notes
Bibliography: Biblography: p.241-262.
Statement of Responsibility: David G. Young.
General Note: Cover title.
General Note: Includes index.
Funding: Bulletin (University of Florida. Agricultural Experiment Station) ;
 Record Information
Bibliographic ID: UF00027135
Volume ID: VID00001
Source Institution: University of Florida
Rights Management: All rights reserved by the source institution and holding location.
Resource Identifier: aleph - 001597028
oclc - 06026971
notis - AHM1158

Table of Contents
    Front Cover
        Front Cover
    Title Page
        Title Page
    Table of Contents
        Table of Contents 1
        Table of Contents 2
        Table of Contents 3
        Table of Contents 4
        Table of Contents 5
    Introduction
        Page 1
        Page 2
    Classification and taxonomic treatment
        Page 3
        Page 4
    General morphology and terminology
        Page 5
        Page 6
        Page 7
        Page 8
    Methods and materials
        Page 9
        Page 10
    Distribution, collecting localities, and characteristics of the fauna
        Page 11
        Page 12
        Page 13
        Page 14
        Page 15
        Page 16
    Medical importance
        Page 17
    Key to the subfamilies of psychodidae of the new world
        Page 18
    Subfamily sycoracinae
        Page 19
        Page 20
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    Subfamily phlebotominae
        Page 29
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    Synopsis of new taxa, synonums, female descriptions, and new distribution records
        Page 239
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    Reference
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    Systematic index
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Full Text
L1 .


April 1979


Bulletin 806 (technical)


A Review of the
Bloodsucking Psychodid Flies of Colombia
(Diptera: Phlebotominae and Sycoracinae)


David G. Young



















Agricultural Experiment Stations
Institute of Food and Agricultural Sciences
University of Florida, Gainesville
F. A. Wood, Dean for Research











A Review of the
Bloodsucking Psychodid Flies of Colombia
(Diptera: Phlebotominae and Sycoracinae)

David G. Young
Assistant Research Scientist
Department of Entomology and Nematology
University of Florida, Gainesville, Florida 32611












Research was supported in part by U.S. Army Medical Research and Devel-
opment Command under Contract No. DADA 17-72-C-2139.


This public document was promulgated at an annual cost of $9311 or a
cost of $4.66 per copy to provide information on insects of medical im-
portance. Publication was funded in part through the U. S. Army con-
tract.









CONTENTS
INTRODUCTION ......................................... 1
CLASSIFICATION AND TAXONOMIC TREATMENT ............ 3
GENERAL MORPHOLOGY AND TERMINOLOGY ............... 5
METHODS AND MATERIALS ............................... 9
Field Collections ........................................ 9
Laboratory Methods ....................................... 10
DISTRIBUTION, COLLECTING LOCALITIES, AND
CHARACTERISTICS OF THE FAUNA ...................... 11
Pacific Coast Region ..................................... 11
Atlantic Coast Region .................................... 14
Oriental Region ........................................ 15
MEDICAL IMPORTANCE ................... ............. 17
KEY TO THE SUBFAMILIES OF PSYCHODIDAE OF
THE NEW WORLD ..................................... 18
SUBFAMILY SYCORACINAE ............................. 19
Keys to the Sycorax Species ............................. 20
1. Sycorax andicola n. sp. ........................... 21
2. Sycorax colombiensis n. sp. ......................... 23
3. Sycoraxfairchildi n. sp. ............................ 23
4. Sycorax trispinosa n. sp. ........................... 25
SUBFAMILY PHLEBOTOMINAE ............................. 29
Key to the New World Genera ............................... 29
Genus Warileya Hertig, 1948 ......... .... ............... 29
Keys to Species ..................................... 30
5. Warileya (W) nigrosacculus ....................... 30
6. Warileya (W) rotundipennis ......................... 32
Genus Brumptomyia Franga and Parrot, 1921 ................... 32
Key to Species ........................... ........... 34
7. Brumptomyia avellari ............................. 34
8. Brumptomyia beaupertuyi .......................... 36
9. Brumptomyia galindoi ............................. 36
10. Brumptomyia hamata .............................. 38
11. Brumptomyia leopoldoi ........................... 40






Genus Lutzomyia Franga, 1924 ................. .......... 42
Key to Subgenera, Species Groups, and Ungrouped
Species of Lutzomyia ................................ 42
Subgenus Lutzomyia Franga, 1924 ............................ 51
Keys to Species ....................................... 51
Series longipalpis .......................... ............. 52
12. Lutzomyia (L.) bifoliata ............................ 52
13. Lutzomyia (L.) lichyi .............................. 54
14. Lutzomyia (L.) longipalpis .......................... 56
Series cruciata ........................................ 56
15. Lutzomyia (L.)gomezi ............................. 56
16. Lutzomyia (L.)marinkellei n. sp. .................. ... 59
Migonei Group Theodor, 1965 ............................. 62
Keys to Species ..................................... 63
Series migonei .................. ...................... 63
17. Lutzomyia migonei ................................ 63
Series walker ..................... ...... ............ 63
18. Lutzomyia marajoensis ........................... 63
19. Lutzomyia sp. de Baduel ............. ............. 66
20. Lutzomyia walker .............................. 67
Saulensis Group Lewis et al., 1977 ........................... 69
21. Lutzomyia saulensis ............................... 69
Verrucarum Group Theodor, 1965 .......................... 71
Key to Species ........................................ 71
Series serrana ....................................... 73
22. Lutzomyia odax .................................. 73
23. Lutzomyia serrana ................................ 73
Series verrucarum .................................... 75
24. Lutzomyia andina ............................... 75
25. Lutzomyia columbiana .......................... .. 75
26. Lutzomyia disiuncta .............................. 78
27. Lutzomyia evansi ................................ 78
28. Lutzomyia longiflocosa ............................ 80
29. Lutzomyia moralesi n. sp. ........................... 80
30. Lutzomyia nuneztovari ............................. 83
31. Lutzomyia ovallesi ................................ 84
32. Lutzomyia quasitownsendi .......................... 86
33. Lutzomyia sauroida .............................. 86






34. Lutzomyia spinicrassa ............ ................. 86
Vespertilionis Group Theodor, 1965 ......... .............. 88
Key to Species .................. ......... .......... 88
35. Lutzomyia isovespertilionis ......................... 88
36. Lutzomyia vespertilionis ............................ 90
SubgenusDampfomyia Addis, 1945 ........................ 91
37. Lutzomyia (D.) rosabali ....................... 91
Subgenus Pintomyia Costa Lima, 1932 ........................ 93
38. Lutzomyia (Pi.) spinosa ............................ 93
SubgenusPressatia Mangabeira, 1942 ......................... 95
Keys to Species ................................. 96
39. Lutzomyia (Pr.) camposi ............................ 96
40. Lutzomyia (Pr.) dysponeta ........................ 98
41. Lutzomyia (Pr.) triacantha .......................... 100
Baityi Group Theodor, 1965 ........................... 102
42. Lutzomyia baityi ........... ........... ... ... 102
Subgenus Viannamyia Mangabeira, 1941 ...................... 102
Keys to Species ................................ 103
43. Lutzomyia (V) caprina ........................... 103
44. Lutzomyia (V)furcata ............. ............ 104
45. Lutzomyia (V) tuberculata .......................... 106
Shannoni Group Theodor, 1965 ........................... 108
Keys to Species ............... ..... .. ............ 108
46. Lutzomyia abonnenci ............................ 109
47. Lutzomyia dasymera ............................ 111
48. Lutzomyia dendrophyla ........................ 113
49. Lutzomyiapunctigeniculata ............. ............ 115
50. Lutzomyia shannoni .......... ........ .... 117
51. Lutzomyia undulata .......................... .. 119
Longispina Group Theodor, 1965 .......................... 121
Keys to Species ...................... ............. 121
52. Lutzomyia longispina ............................ 121
53. Lutzomyia triramula ............................. 123
Aragaoi Group Theodor, 1965 .......................... 125
Keys to Species ...................................... 125
Series aragaoi ................. ......... ........... 126
54. Lutzomyia aragaoi ................................ 126
55. Lutzomyia barrettoi barrettoi ........................ 128






56. Lutzomyia barrettoi majuscula n. ssp..........
57. Lutzomyia carpenter ......................
Series brasiliensis .............................
58. Lutzomyia runoides .......................
Dreisbachi Group Lewis et al., 1977 ..................
K ey to Species ..............................
59. Lutzomyia aclydifera ......................
60. Lutzomyia dreisbachi .......................
Subgenus Trichophoromyia Barretto, 1962 ..............
K ey to Species .................................


61. Lutzomyia (T) auraensis ..
62. Lutzomyia (T) cellulana n. sp.
63. Lutzomyia (T) howardi n. sp.
64. Lutzomyia (T) reburra ......
65. Lutzomyia (T) saltuosa n. sp..
66. Lutzomyia (T) ubiquitalis ....
SubgenusNyssomyia Barretto, 1962 ...
Keys to Species ................
67. Lutzomyia (N.) antunesi .....
68. Lutzomyia (N.)flaviscutellata
69. Lutzomyia (N.) olmeca bicolor
70. Lutzomyia (N.) trapidoi .....
71. Lutzomyia (N.) umbratilis ...
72. Lutzomyia (N.) ylephiletor ...


73. Lutzomyia (N.)yuilli ..................
SubgenusPsychodopygus Mangabeira, 1941 ........
K eys to Species ...........................
Series squamiventris .........................
74. Lutzomyia (P) bernalei ................
75. Lutzomyia (P) chagasi ............... .
76. Lutzomyia (P.)fairtigi ................ .
Series arthuri ............................
77. Lutzomyia (P) bispinosa .......... ..
Seriespanamensis ...........................
78. Lutzomyia (P) ayrozai .................
79. Lutzomyia (P) carrerai carrerai .........
80. Lutzomyia (P) carrerai thula n. ssp. ......
81. Lutzomyia (P) davisi ................


. ....... 142
. ....... 144
............ 146
............ 148
. ....... 149
............ 149
............ 150
. ....... 15 1
. ....... 153
. ....... 155
............ 157
............ 160
............ 162
............ 164
............ 166
............ 167
. ....... 170
............ 170
............ 170
............ 170
............ 173
............ 173
............ 175
. ....... 175
............ 178
............ 180
............ 183






82. Lutzomyia (P) guyanensis ...................... 186
83. Lutzomyia (P) hirsuta hirsuta .................... 188
84. Lutzomyia (P) nocticola ................... ... 190
85. Lutzomyia (P) panamensis .......................... 190
86. Lutzomyia (P) recurva ........................... 193
87. Lutzomyia (P.) sp. ofTres Esquinas ................... 195
Vexator Group Theodor, 1965 ................ ...... ... 195
Keys to Species ........................ ............. 196
88. Lutzomyia cirrita ................................. 197
89. Lutzomyia hartmanni ........................... 199
90. Lutzomyia osornoi ............................... 201
91. Lutzomyia sanguinaria ........................ 201
92. Lutzomyia scorzai ................................ 204
93. Lutzomyia strictivilla n. sp. ......................... 206
94. Lutzomyia sp. of Pichinde ................... ... 210
Cayennensis Group Theodor, 1965 ........................... 212
Keys to Species .................................... 212
Series cayennensis ...................... ............. 213
95. Lutzomyia cayennensis ................... .... 213
96. Lutzomyia micropyga .............................. 215
Series atroclavata ..................................... 218
97. Lutzomyia atroclavata ........................... 218
98. Lutzomyia venezuelensis ........................... 220
Oswaldoi Group Theodor, 1965 ............................. 222
Keys to Species ..................................... 223
99. Lutzomyia pia ................................... 223
100. Lutzomyia rorotaensis ............................. 225
101. Lutzomyia trinidadensis ................... ........ 227
Pilosa Group Theodor, 1965 .............................. 230
102. Lutzomyia pilosa ................................. 231
Ungrouped Species ....................... ..... ......... 233
103. Lutzomyia nordestina .................... ........ 233
104. Lutzomyia rangeliana ............................. 235
105. Lutzomyia sp. of Anchicaya ....... ................. 237
SYNOPSIS OF NEW TAXA, SYNONYMS, FEMALE DESCRIPTIONS,
AND NEW DISTRIBUTION RECORDS ...................... 239
REFERENCES CITED .................................... 241
SYSTEMATIC INDEX .................................... 263








INTRODUCTION


Prior to the last decade, we knew very little about the phlebotomine sand fly*
fauna of Colombia. Osorno et al. (1967) in reviewing previous studies by others
and adding records of their own, reported less than 25 species in the Republic.
From 1967 to 1972, the late Dr. E. Osorno-Mesa and his colleagues at Instituto
Nacional de Salud, Bogota, added more records, bringing the total to 77 spe-
cies and subspecies (Osorno et al., 1972a).
In this review 101 species and subspecies of Phlebotominae are reported
from Colombia. This represents about one-third of the described New World
species, but I estimate that at least 40 additional species will be discovered in
the Republic, especially in the little-collected southeastern region.
The small subfamily Sycoracinae, whose members resemble those of Phle-
botominae in being able to take vertebrate blood, is treated here for this reason.
In order to clarify the status of some taxa and to study intraspecific variation
and distributional patterns, an effort was made to examine specimens from as
many Colombian and non-Colombian localities as possible. To this end, I am
very grateful to the following persons for providing specimens, information or
both: Dr. Jorge R. Arias, Instituto Nacional da Pesquisas de Amaz5nia (INPA),
Manaus; Drs. Stephen C. Ayala and Pablo Barreto, Universidad del Valle, Cali;
Drs. Richard D. Ward and Habib Fraiha, Instituto Evandro Chagas, Bel6m; Dr.
David J. Lewis, British Museum (Nat. Hist.); Dr. Cornelis J. Marinkelle, for-
merly 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, for-
merly Systematic Entomology Laboratory (ARS); Dr. Richard C. Wilkerson,
University of Florida; Dr. Thomas H. G. Aitken, Yale University School of
Medicine; Dr. Elisha S. Tikasingh, Caribbean Epidemiology Center (CAREC),
Trinidad; Dr. Harold Trapido, Louisiana State University Medical Center; Dr.
Jorge E. Velasco, La Paz, Bolivia; Dr. Derek A. Duckhouse, University of
Adelaide, South Australia; Maj. John F. Reinert, U.S. Army Medical Research
and Development Command, Washington, D.C.
Others who have supported this study and to whom I am indebted include
members of the Atlantic-Pacific Interoceanic Canal Study Commission, Canal
Zone from 1967-1968. Col. Bruce F. Eldridge of this organization was espe-
cially helpful in developing field programs and in providing data on Choc6 col-
lections.

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






Financial support from U.S. Army Medical Research and Development
Command under contract no. DADA 17-72-C-2139 is gratefully acknowl-
edged. 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. Fairchild, Dale H.
Habeck, and Stephen G. Zam. Miss Stephanie Haney ably mounted thousands
of specimens on slides from 1972 to 1977. For typing the manuscript, I wish
to thank Mrs. Adele Koehler.
Dr. G. B. Fairchild deserves special recognition for his constant encourage-
ment, stimulating discussions, and expert assistance throughout the study. I am
deeply indebted to him. Particular appreciation is expressed also to my wife
Molly who was most understanding during the preparation of this review.
Holotypes and allotypes of new taxa are to be deposited in the U.S. National
Museum (Nat. Hist.). Paratypes, when available, will be held by the following
institutions: Instituto Nacional de Salud, Bogota; Florida State Collection of
Arthorpods, Gainesville; and British Museum (Nat. Hist.), London. A synoptic
collection will be given to the Instituto Nacional de Salud, Bogota.
The remaining 20,000+ specimens, upon which most of this review is based,
are housed in the Department of Entomology and Nematology, University of
Florida.










CLASSIFICATION AND TAXONOMIC
TREATMENT


Of the six recognized subfamilies of Psychodidae (Duckhouse, 1972, 1973),
all but one, Horaiellinae from the Oriental Region, are represented in continental
Colombia. The species ofHoraiella, Trichomyia, and Sycorax and its allies are
united by some in the subfamily Trichomyiinae but this appears to be artificial
and does not "express the wide discontinuities and degrees of difference from
other Psychodidae, especially in the immature stages" (Duckhouse, 1972).
The classification of Phlebotominae is interpreted differently according to
authorss. In a recent paper we reviewed most classificatory schemes and pro-
posed one which is used here for the New World taxa (Lewis et al., 1977). It is
essentially a modification of Theodor's 1965 classification. Three American
genera are recognized Lutzomyia 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. (1977).
The majority of the Lutzomyia spp. were described originally in the genus
Phlebotomus (= Flebotomus) Rondani. Some authors continue to place them
in this genus, ignoring the evidence to the contrary provided by Theodor (1948,
1965) and Christensen et al. (1971).
The large genus Lutzomyia is divided into numerous subgenera, species
groups, and series based upon adult morphology (Lewis et al., 1977). The di-
versity of groups within the genus is attributed to "radiation or even 'explosive
radiation' of some immigrating ancestral stocks in a newly colonized area having
a wealth of ecological niches" (Theodor, 1965). This implies that these ancestral
stocks originated in the Old World a hypothesis which may never be proved.
It would appear that recent speciation in some groups (e.g. subgenus Tricho-
phoromyia) may have been due to climatic changes in the past, especially during
the Pleistocene, which served to isolate conspecific populations in moist refugia
during dry periods (Haffer, 1974).
Other remarks on the classification of New World phlebotomines are given
by Lewis et al. (1977) 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 in-
cluding 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 Instituto Nacional para Programas Especiales de Salud, Bogoti
INPA Instituto Nactional de Pesquisas de Amazonia, Manaus
UV Universidad del Valle, Cali
GML Gorgas Memorial Laboratory, Panama
UF University of Florida
Collectors
C.H.P. Charles H. Porter
C.J.M. Cornelius J. Marinkelle
R.C.W. Richard C. Wilkerson
D.G.Y. David G. Young
Other Abbreviations
ca. about
cf. compared to
classif. classification
descr. description
dist. distribution
morphol. morphology
pop. population
ref. reference







GENERAL MORPHOLOGY AND TERMINOLOGY


This brief discussion focuses on the structures commonly used in taxonomic
studies of phlebotomine sand flies. General accounts of adult anatomy include
those by Fairchild and Hertig (1947a), Kirk & Lewis (1951), Abonnenc & Minter
(1965), Davis (1967), Abonnenc (1972), Lewis (1973), and Forattini (1973).
The terminology varies according to author; that used here applies also to the
sycoracine flies except where otherwise stated.
Although little studied, the immature stages are probably no less important
in systematics but they are not considered here owing to lack of material. For
information on the anatomy of these stages see Barretto (1941), Hanson (1968),
Abonnenc (1972), Carnheiro & Sherlock (1964), Ward (1977), and Zimmerman
et al. (1977). Descriptions of immatures of those species occurring in Colombia
are cited in the appropriate species bibliography.
The sand fly head and its appendages (Fig. 12A) are useful in associating
conspecific males and females and for grouping taxa below the genus level. Head
height, measured from the vertex to the tip of the clypeus, in relation to its great-
est width is of value in distinguishing some of the Lutzomyia species.
As a probable adaptive feature related to host finding, the size of the compound
eyes varies according to species, especially those of Lutzomyia, but may be
characteristic of subgenera and equivalent species groups. Contrary to Davis
(1967), ocelli are lacking in the Phlebotomines and Sycoracines. Illustrations
are by far the best way to indicate eye size, the relative terms such as "small"
or "large" being understood easily with associated figures. The interocular dis-
tance oreye separation is the narrowest distance between the eyes and is measured
directly or may be expressed by the number of facet diameters (or fractions
thereof) needed to connect the eyes.
The interocular suture (= post frontal suture of mosquitoes, Christophers,
1960) is complete in adults of Brumptomyia (Fig. 8B), Warileya (Fig. 6A), and
Sycorax (Fig. 2A) but is incomplete in those of Lutzomyia (Fig. 12A).
In Phlebotominae, the maxillary palpus (Fig. 12A) consists of 5 segments,
the basal one (palp 1) always the smallest and least important in taxonomy
(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 I 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 phle-
botomines and to palp 2 in the Colombian Sycorax spp.






The cylindrical antennal flagellomeres, unlike true segments, are not inde-
pendently musculated (Imms, 1938). For this reason Ijoin the increasing number
of students who use the term" flagellomere' in the place of antennal segment."
The antenna consists of a scape, pedicel, and 14 flagellomeres, each of which
is designated by a Roman numeral (Fig. 12A). In Sycorax the terminal flagel-
lomere is markedly reduced in size (Fig. 2D). The length of flagellomere 1
(= antennal segment 3), the most basal flagellomere, in relation to other flagel-
lomeres, head height, or labrum length is useful as a diagnostic feature at the
species or subspecies level.
The paired antennal ascoids (Fig. 12B), often difficult to observe, are impor-
tant owing to interspecific variation in their distribution, shape, and length. Some
specimens may have but a single ascoid on a particular flagellomere.
The mouthparts of New World phlebotomines in relation to feeding habits and
taxonomy were studied in detail by Lewis (1975a). Following him, I use the
term labrum for the labrum-epipharynx of various authors. Its length is meas-
ured from the apex of clypeus to tip of proboscis.
The cibarium (= buccal cavity) lies within the clypeus and is armed with a
variable number of teeth in the Lutzomyia and Brumptomyia females. Those
in the latter genus are characteristically arranged in 4 longitudinal rows (Fig.
10M). Cibarial teeth are absent in both sexes of Warileya and Sycorax and are
poorly developed or absent in the Lutzomyia and Brumptomyia males.
When viewed ventrally as in Fig. 12E, the horizontal teeth (= hind teeth) of
most Lutzomyia females indeed look like teeth, their tips pointed or not, de-
pending 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.
Two other features of the cibarium are noted in the descriptions and keys.
The chitinous arch (Fig. 12E), when complete, crosses the ventral wall of the
cibarium. Its development varies according to species and may be complete or
not. The pigment patch on the dorsum of cibarium (Fig. 12E) varies in size,
shape, and degree of infuscation (= pigmentation) in the Lutzomyia spp.
The pharynx (Fig. 12C), composed of 3 chitinous plates, is attached to the
cibarium. Many species of Old World phlebotomines (genus Sergentomyia)
have spines on the posterior part of the pharynx but they are relatively rare in
the Lutzomyia spp., occurring mostly in some species in the cayennensis and
oswaldoi groups. The majority of Lutzomyia females have transverse, unarmed
ridges on the wider, posterior area of the pharynx. The length of the pharynx
is given in descriptions although it is flexible and difficult to measure with ac-
curacy (Lewis, 1967a).
I adopt the terminology of Davis (1967) as applied to the thorax. The meson-
otum should correctly include the mesoscutum, mesoscutellum, and postnotum
and should not be restricted to the mesoscutum alone (Saether, 1971).






The degree and distribution of pigmentation of the thorax and other body
regions are important in associating sexes and for distinguishing species of Lut-
zomyia. 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 episternal setae) are present in species of Lutzomyia, Brumptomyia, and
Sycorax. The Warileya spp. and the majority of Old World phlebotomines lack
such setae (Abonnenc & Leger, 1975). Additional pleural setae, not mentioned
in other studies to my knowledge, occur behind the metathoracic spiracle in all
species of Phlebotominae and Sycoracinae examined, those in the latter
subfamily being 4 in number and relatively stout (Fig. 2H). The 4 postspiracu-
lar setae in the phlebotomine species are reduced in size, barely visible (Fig.
12D).
The wing length is measured from the basal costal node (= rudiment of te-
gula; Snodgrass, 1935) to its apex; its width at widest part (Fig. 12G). Certain
wing vein sections (Fig. 12G) are given useful, easily remembered names. Al-
pha (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 R, + R3. Gamma (y)
is the section of R from the r m crossvein to the junction of R5. Delta (8) is
that part of R1 distad of the junction of Rz and R3. It is negative when R, ends
before this junction.
The legs are measured as in Fig. 12H. A few phlebotomine spp. have a row
of short spines on the hind femur (Fig. 31 F). The length of the femur in relation
to that of the tibia and/or basitarsus is sometimes used for distinguishing spe-
cies 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 spe-
cies. 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 useful
in associating male and female sand flies (Hertig & Fairchild, 1950) but the
character state is variable among individuals of conspecific populations (For-
attini, 1954).
The structures of the bilaterally symmetrical male genitalia, labelled in Fig.
2 and 12, are extremely important in systematics. After eclosion, the genitalia
rotates 1800 in the phlebotomine males but not in those of Sycorax. The style,
attached to the apex of the coxite, bears 1 to 6 major spines* and 0 to many


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






additional small 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 Sycorax males. It may or may not have a
setal tuft or other nondeciduous setae on its inner surface. These setae are those
which remain on the coxite after maceration and which usually differ in size or
shape from other setae on the appendage.
The fused, paired aedeagi, each of which is called the aedeagus is subtrian-
gular and well sclerotized in most phlebotomine males. Some species (e.g. L.
walker) have aedeagi with dorsal projections. The aedeagus of the Colombian
Sycorax males is complex as in Fig. 2F There is a basal, laterally flattened
sperm pump within the abdomen, distal paired genital filaments and other
structures mentioned on p. 19. 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.
The phlebotomine parameres lie between the coxites and ventral lateral lobes
when viewed laterally. Their shape and station are important 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. 44H. The cerco-
pods of the Sycorax males appear to correspond to the parameres of Phlebotom-
inae and are very complex.
The ventral, nonsegmented lateral lobes are more or less uniform in the phle-
botomine males, although their length, width, and station may offer good
characters in some species. Corresponding structures are wanting in the syco-
racines.
The shape of the setose cerci may help associate conspecific sexes but in gen-
eral these structures are of limited use in species diagnosis.
The size and structure of the internal paired spermathecae and their ducts
(Fig. 12F) are particularly important in the classification and identification of
female sand flies. An idea of the variety of spermathecae can be gained by scan-
ning 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 Choc6
Department and elsewhere to a lesser extent, we used the following methods to
capture these flies.
Battery powered light traps (Sudia and Chamberlain, 1962) attract many psy-
chodids, the numbers varying according to weather conditions, location, species
composition, etc. We secured these traps to tree branches 2 meters above ground
level but did not use carbon dioxide as an adjunct attraction. At Curiche, where
at least 1 trap per week was operated throughout the night (1800 hrs.-0700
hrs.), we collected a total of 23 phlebotomine spp. (378 dc, 789 99) from April
to Dec., 1967 (39 trap nights). We recorded 36 sand fly species from Curiche
based on all collection techniques.
Shannon traps (Shannon, 1939) made from muslin bedsheets do not trap in-
sects per se but are collecting devices similar to those used by lepidopterists in
"sheeting" for moths at night. A gasoline lantern provides a light source ena-
bling collectors to readily aspirate the psychodids which land on the illuminated
cloth. They are attracted to the light, to the collectors, or to a combination of
both. The species composition of light and Shannon trap captures is generally
similar.
Malaise traps (Townes, 1962) andflight traps (similar to the design of Gressitt
and Gressitt, 1962) were placed in forest clearings, across trails or other 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, these traps do not capture large numbers of psy-
chodids 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
(316 6, 482 99) 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 bur-
rows, and litter on the forest floor. Tree trunks are favored resting sites for several
species, specimens of which are easily captured with a simple aspirator.
During the survey in Choc6 Dept., especially at Curiche and Teresita, we
collected large numbers of phlebotomines on human bait. The majority of col-
lections were made by two men sitting on the forest floor between 1830 and 2100
hours. Phlebotomines were captured with aspirators as they attempted to feed,


*One trap day equals 24 hours.






and the results were computed on a man-hour basis, i.e., the total number cap-
tured in one hour divided by the number of collectors.
Specimens were preserved dry in cardboard pill boxes rather than in alcohol
to prevent hardening of muscle tissue.


LABORATORY METHODS
For routine identification of phlebotomines and sycoracines, undissected
specimens are first macerated (i.e., cleared) in 10-20% NaOH. The loss of
pigmentation can be reduced by heating this solution (containing the flies) to the
boiling point rather than macerating them at room temperature for 12 hours or
more. The flies should be left in the hot NaOH for 5 minutes, then placed directly
into a drop of 80-95% liquid phenol (C6H5OH) in a depression microslide.
Within 5 minutes, the important internal structures such as the spermathecae
and ducts are clearly visible and should be drawn if necessary. Once identified,
the flies can be discarded, preserved in vials of 70% alcohol, or processed further
for slide mounting. I generally follow the procedure of Fairchild and Hertig
(1948c) except that Canada balsam or Euparal is substituted for copal.
One part of balsam is mixed with about five parts of phenol in a depression
microslide. Stirring with a glass rod hastens the process. One to 12 undissected
flies are then placed into this mixture where they remain for about one week at
room temperature. As the phenol evaporates, the balsam or Euparal will become
increasingly viscous or tacky.
Before the medium becomes completely hardened, the head and wings are
removed from the specimen. These parts, as well as the body, are then lifted into
small drops of the tacky medium on a coverslip. Small cut pieces from another
coverslip are placed in each corner to prevent later crushing.
Finally, after the drops of medium with the specimen parts have hardened in
a dust-free cabinet, the coverslip is placed on top of a large drop of balsam on
a microslide and is allowed to dry.
Other slide-mounting procedures are discussed by Osorno et al. (1966), Quate
& Steffan (1966), and Lewis (1973) and others. The choice of which to use is
a matter of personal preference, but in all cases the best preparations are made
from freshly killed flies.
Sometimes it is necessary to remount specimens because of undesirable po-
sition and/or shrinkage. For those embedded in Canada balsam or other xylene-
soluble media, it is advisable to submerge the entire slide in liquid phenol in a
suitably closed container such as a petri dish. After 3 to 7 days, the specimens
become soft and are easily handled. Xylene as a solvent should be avoided be-
cause the specimens tend to become brittle and are easily damaged.
Specimens were drawn with the aid of a camera lucida and Bausch and Lomb
microprojector. I measured specimens with a calibrated ocular micrometer. All
such measurements are given in millimeters throughout the text and figures.








DISTRIBUTION, COLLECTING LOCALITIES,
AND CHARACTERISTICS OF THE FAUNA


The distribution of phlebotomines and sycoracines can be understood by pres-
ent 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 per-
iods. During times of drought, populations of forest birds and other organisms
were restricted to isolated forest refugia which acted as core areas of speciation.
As conditions became wetter, these forests enlarged in area, often merging with
others to form zones of secondary contact. The presumed forest refugia in Col-
ombia and elsewhere were discussed and mapped by Haffer (1967, 1974) and
Brown (1975), the latter author studying speciation in forest butterflies in re-
lation to these presumed refugia.
Although more data are needed, the refuge theory offers a plausible expla-
nation for understanding recent speciation in the Phlebotominae and Sycora-
cinae. 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 re-
gions. D'Allesandro & Barreto (1971) delineate 7 such areas, their large "Ori-
ental Region" consisting of both Amazonian forest and eastern natural savannah
(llanos). Within each region there are one or more life zones, each defined by
a combination of biotemperature, annual precipitation, humidity, and vegetation
(Holdridge, 1967). The life zones in Colombia were studied by Espinal and
Montenegro (1963). Their "bosque humedo" translates to "tropical moist for-
est" (2000- 4000 mm of rain per year) and "bosque muy humedo" corresponds
to "tropical wet forest" (4000- 8000 mm of rain per year). The "bosque plu-
vial" (= "tropical rain forest"), the wettest lowland life zone, receives over
8000 mm of rain per year. A good review of these life zones, climate, soils,
topography, and geologic history of Colombia is given by Mend6z (1977).
Figure 1 depicts where phlebotomines and sycoracines have been collected
in Colombia. Osorno et al. (1972a) provide specific data for most localities;
others from which collections were made by me or my colleagues are discussed
below in relation to faunal regions (D'Allesandro & Barreto, 1971) and distri-
bution centers (Haffer, 1974).

PACIFIC COAST REGION
Haffer (1967) attributes the "high concentration of endemic species in the
tropical lowland forests of western Colombia and Central America" to a gradual





















































-40


FIGURE 1
Map of Colombia showing localities where phlebotomines and sycoracine flies have
been collected (records based on Osorno et al., 1972a and this review). Shaded sections
represent areas 1000 m+ above sea level.


= Phlebotomine record
= Sycoracine record
= Phlebotomine and sycoracine record
= Atlantico Dept.
= Magdalena Dept.


3 = Norte de Santander Dept.
4 = Caldas Dept.
5 = Risaralda Dept.
6 = Quindio Dept.
7 = Cundinamarca Dept.


Legend
*
x


2






accumulation of isolates in the trans-Andean forest region.* The dominant re-
fugium on the Colombian Pacific coast is the Choc6 refugium extending from
Lago Calima (Valle) to north of Quibdo (Choc6) 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 of
the Andes (cis-Andean region). Most of these reached the Pacific coast forests
by advancing around, not across, the northern Andes. These species which now
occur on both sides of the mountains include: Lutzomyia ayrozai, L. guyanensis,
L. bispinosa, L. gomezi, L. nordestina, L. serrana, L. pilosa, L. spinosa, L.
trinidadensis, L. shannoni, L. tuberculata, L. triramula, and others.
Nine of the 41 species and subspecies do not occur in other regions of the
Republic. Of these, only Lutzomyia sp. of Anchicaya is precinctive, the others
occurring also in the trans-Andean regions of Northwestern Ecuador and/or
Central America. These species include: Brumptomyia hamata, Warileya
nigrosacculus, L. reburra, L. recurva, L. isovespertilionis, L. ylephiletor, L.
sanguinaria, 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. barrettoimajuscula 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 possibility, although slight, that some phlebotomines were able to
cross the Andes in southern Ecuador and northern Per6 in times past when wet
forests occupied the now dry valleys. Some Amazonian birds apparently colo-
nized Pacific coast forests in such a manner (Haffer, 1967). Phlebotomines, how-
ever, 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. vespertilionis and its sister species, L. isoverperti-
lionis probably invaded the Pacific coast region of Colombia from Central
America, the former species also occurring east of the Andes, at least as far south
as Bolivar Department.

Department of Chocd
Three collecting sites within this Department were described by Eldridge &
Fairchild (1973) and Eldridge et al. (1973). Maps accompany their descrip-
tions.

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






Curiche. On narrow coastal plain between western slopes of Serrania del
Baud6 and Humboldt Bay on the Pacific Coast; "tropical wet forest;" eleva-
tion* less than 10 m. Sand flies were collected from April 1967, to Dec. 1967,
in disturbed and undisturbed evergreen forests and near a mangrove swamp.
Alto Curiche. About 3 km inland from Curiche on western ridge of Serrania
del Baud6; "tropical wet forest;" elevation 302 m. The sand fly fauna is similar
to that of nearby Curiche.
Teresita. At eastern foothills of Serrania del Baud6, near Rio Truando; tran-
sitional life zone, "tropical moist/tropical wet forest;" elevation 35 m. Collec-
tions made in semi-disturbed forests on flat or gently rolling terrain from
March-Dec. 1967.

Department of Valle
Lower Anchicayd Dam. About 35 km inland from Pacific Ocean, on the Rio
Anchicayi; "tropical wet forest;" elevation ca. 560 m. This site is similar to
that near the Rio Anori (Antioquia Dept.) in terrain, elevation, and climate.
Sycoracines were recorded from both localities and from one other in Cauca
Department.
Twenty-five km E of Buenaventura. "Tropical rain forest" according to Es-
pinal (1968); elevation ca. 50 m. Phlebotomines were captured in a patch of
disturbed forest surrounded by cultivated fields.

ATLANTIC COAST REGION
This area encompasses not only the Caribbean lowlands but also the north-
ern foothills of the western and central Cordilleras of the Andes. The presumed
Nechi refugium is located here, a center "heavily influenced by the Choc6 cen-
ter 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 col-
leagues near the Rio Anori (1970- 1971) yielded 37 species and subspecies.
Thirty of these also occur in Pacific coast forests, a fact not surprising based on
the distribution of other organisms (Haffer, 1967, 1974). Lutzomyia strictivilla
may be endemic. Lutzomyia yuilli and L. nocticola are cis-Andean taxa. The
former reaches its northernmost limits in northern Colombia; whereas L. noc-
ticola invaded extreme western Panama but populations between there and
northern Colombia have since disappeared.

Department ofAntioquia
Rio Anori (7 19'N; 750 04'W), about 24 km SW of Zaragoza; "tropical wet
forest;" elevations from 500-700 m. Located in the foothills of the central

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






Cordillera of the Andes, this site is mostly well forested with steep, well drained
slopes.

Department ofMagdalena
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 Caribbean Sea; "tropical
wet forest;" elevation ca. 3 m.
Between Santa Marta and Minca, near a small stream; probably a trans-
ition zone between "tropical wet forest" and "tropical dry forest"; elevation
ca. 80 m.
Minca, SE of Santa Marta; life zone indeterminate but similar to preceding
site; elevation 200 m.

ANDEAN REGION
Nearly all records of phlebotomine species occurring above 1000 m are given
by Osorno et al. (1972a). On the basis of these records and my limited collec-
tions from the Western Cordillera, it is apparent that the Lutzomyia species in
the verrucarum and vexator groups dominate the rather depauperate phlebo-
tomine fauna.

Department of Valle
Three sites W or SW of Cali 1. Near Pichinde, 2. In forest at the television
tower just E of Saladito, and 3. Near Rio Pance, SW of Cali; "lower montane
very wet forest;" elevations from 1570- 1700 m. Limited collections from 31
July, 1973, to August, 1973.

Department of Cauca
Eastern slope, near summit of Cerro Munchique, west of Popoyin; "lower
montane 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 which forms part of the Oriental Region
(Haffer, 1967, fig. 1). Many of the species known from the Amazonian forests
in Brazil, Peru, and Ecuador are expected to occur also in Colombia.






Comisaria ofAmazonas
About 17 km W of Leticia; "tropical moist forest;" elevation 84 m. Phle-
botomine 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.


Intendencia of Caqueta
Tres Esquinas; transitional life zone, "tropical moist forest"/'tropical wet
forest;" elevation ca. 200 m. Using light traps, Dr. C. J. Marinkelle collected
numerous phlebotomines in November 1971.










MEDICAL IMPORTANCE


The phlebotomine sand flies, unlike the sycoracine flies, are notorious vec-
tors of leishmaniasis (Ward, 1977), bartonellosis (Schultz, 1968), and certain
arboviruses (Tesh et al., 1974), diseases which have caused a great deal of hu-
man suffering in the neotropics and elsewhere.
Leishmaniasis, a collective term referring to several disease entities caused
by Leishmania spp., occurs in Colombia but the vectors are poorly known.
Reyes (1957) provided limited information on the distribution and nature of der-
mal (= cutaneous) leishmaniasis in the Republic, noting that 206 out of 725
human infections involved the naso-pharyngeal region (mucocutaneous leish-
maniasis). Ward (1977), citing figures from Reyes (op. cit.) and Garnham
(1962), stated that nearly 2,000 human cases of cutaneous leishmaniasis were
reported in Colombia from 1948 to 1955. The true incidence, distribution, and
identity of leishmaniasis in the Republic remains largely unknown, however.
Visceral leishmaniasis (kala-azar), a very serious disease, is apparently rare
in Colombia, the few human cases from Santander, Tolima, and Cundinamarca
Departments having been discussed by Arjona et al. (1972).*
An outbreak of Bartonellosis (oroya fever, Carrion's disease, etc.), lasting
from 1935 to the early 1940's, in southwestern Colombia (Cauca and Nariiio),
was reviewed by Jaramillo (1943). From 1940 to 1943, there were 2,241 cases
with a mortality rate of nearly 16% (Samaniego, 1944). The disease, also
known to occur in Peri and Ecuador, was probably introduced into Colombia
"by returning soldiers or by 'colporteurs' who travel from village to village"
(Rozeboom, 1947b). Circumstantial evidence led investigators to believe that
Lutzomyia columbiana was the responsible vector.
Arboviruses were recovered from wild caught phlebotomines in the Pacific
lowlands of Colombia (Barreto, 1969). The strain, designated Co Ar 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 parasites, espe-
cially protozoans. References to studies related to them are cited in the species
bibliographies.




*Also see Camacho-Sanchez (1979, Trans. R, Soc. Trop. Med. Hyg. 72:544-545) for
up-to-date information on kala-azar in Colombia.








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
flagellomeres subcylindrical ................ ................ 4
4. Females with 1 spermatheca. Mandibles absent. Male aedeagus
entire ....................................... Bruchomyiinae
Females with 2 spermathecae. Mandibles present.
Males with bifid aedeagus ................. Phlebotominae (Fig. 12F)







SUBFAMILY SYCORACINAE


Following Duckhouse (1972) I presently recognize 3 extant genera in the
subfamily--Sycorax Haliday (ca. 21 spp.), Parasycorax Duckhouse (2 spp.),
and Aposycorax Duckhouse (1 sp.). Until now, only 4 species have been re-
ported from the New World--Sycorax assimilis Barretto and Parasycorax
satchelli (Barretto) from Sio Paulo State, Brazil, Aposycorax chilensis (Ton-
noir) from Chile, and an unnamed sycoracine, the wing of which was figured
by Fairchild (1955) from Palenque, Colon Prov., Panama.
The feeding habits of most species have not been studied but the females of
Sycorax and Aposycorax at least possess mouthparts adapted for sucking
blood. Sycorax silacea Curtis, the type species of the genus from Europe, feeds
on the blood of frogs and transmits a filarial worm to them (Desportes, 1942).
The four new Colombian species, described here in the genus Sycorax, share
several characteristics which set them apart from the other sycoracine species.
In addition to the paired genital filaments between the parameres (= cercopods)
of the male genitalia, there is a single median process which seems to form part
of the aedeagal complex. It lies above the ducts and the proximal sperm pump
(Fig. 2G). Aposycorax chilensis, the only sycoracine with inverted male geni-
talia (Duckhouse, 1972), has very long genital filaments but it lacks the elon-
gate median process. The stout proximal spines (2 or 3) on each style of the
Colombian and Parasycorax species distinguish them from the extant species
in the subfamily.
Other features, not always repeated in the specific descriptions which follow,
are shared by the 4 Colombian Sycorax species. Coloration: Dusky brown,
mesonotum but slightly darker than pleura. Head: Broader than long; frons
puffed out behind antenna. Palpus of 4 segments, the second with an inner
group of 10-20 sensory rods. Antenna slender, of 16 "segments" (scape, ped-
icel, and 14 flagellomeres), the last reduced in size 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. chilensis; pedicel with 2-3 similar
hairs; ascoids subequal in size, often difficult to examine in available material.
Female mouthparts include toothed mandibles and 4 boot shaped sensilla at tip
of labrum; these features absent in males. Cibarium unarmed. Thorax: Pleura
with 6+ setae mainly on the anepimeron but some may extend downwards to
the 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: Females without
setae on tergites 8 and 10 or sternite 8. Male genitalia not inverted. Style with
a patch of 8- 15 short pointed setae 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 up-






wards, sinuous or not; parameres somewhat resembling those of A. chilensis
but differing in the nature and position of the lobes, setae, and projections. Fe-
male genitalia: Each spermatheca and sperm duct forming a continuous tube
with faint transverse striations, terminating in a heavily sclerotized "button"
which appears shiny black; a tenuous sac usually amorphic following treatment
in KOH, arises from the end of the button; an internal apodeme or furca asso-
ciated with paired spermathecae.
The combined length of the spermatheca and duct varies interspecifically
and probably corresponds to the length of the male genital filaments. This fea-
ture and the distribution of the antennal setae and distance between the com-
pound eyes were also used in associating the sexes 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 spe-
cies in the subfamily have not been discovered.


Keys to the Sycorax Species
Males
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. Aedeagal ducts relatively long, exceeding the length of the sperm
pump. A subtriangular plate, dorsoventrally flattened, with acute
bifurcate tip present below the base of the median process of aedeagus.
Tip of median process of aedeagus rounded in lateral view
............... ....... ...... .......... colombiensis (Fig. 3)
Aedeagal ducts shorter, less than length of sperm pump and not as
sinuous. Subtriangular plate absent. Tip of median process of
aedeagus angular in lateral view ................... fairchildi (Fig. 4)
3. Style of male genitalia with 4 strong spines; median process of
aedeagus more slender (Fig. 2G). A larger species, wing length greater
than 2 mm ........................ .......... andicola (Fig. 2)
Style with 3 strong spines; median process of aedeagus relatively
stout (Fig, 51). Wing length less than 1.5 mm ....... trispinosa (Fig. 5)






Females
1. Flagellomeres I- VIII with paired ascoids, other antennal setae randomly
distributed. Eyes rather narrow, separated by distance = to
7 facet diameters ...... .. .. .............. trispinosa (Fig. 5)
Flagellomeres I- IV with paired ascoids, one group of antennal setae
arranged in whorl-like pattern. Eyes separated by 8 or more facet
diameters ................................ ............. 2
2. Furca spade-shaped apically, its length at least / that of sperm duct
+ spermatheca. Flagellomere I, 0.20 mm, or longer .............
..... .. ............................ colom biensis (Fig. 3)
Furca "v" shaped apically, its length but slightly less than that of
sperm duct + spermatheca. Flagellomere I less than 0.20 mm long
........................................... fairchildi (Fig. 4)


1. Sycorax andicola n. sp.
(Fig. 2)
Male holotypee): Wing length 2.35; width 1.05. Head height 0.31; width,
0.37. Eyes separated by 0.14 or by distance = to 8 facet diameters, Antenna
(including scape and pedicel) 1.61 long; flagellomere 1 (0.26 long), about 1.8X
length of flagellomere II; paired ascoids (Fig. 2C), subequal in size, on flagel-
lomeres 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 fig-
ured. 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: Coxite 0.31 long. Style 0.18 long with 1
terminal spine, slightly beveled at tip and with a proximal row of 3 smaller spines.
Paramere as shown with several setose lobes and projections, the most distal
finger-like with rounded, upturned tips. Aedeagal complex of a laterally com-
pressed sperm pump (0.20 long) within the abdomen; a median distal process
(0.165 long) compressed dorsoventrally, curved and slender in lateral view, up-
turned at tip, basal two-thirds much wider than- distal one-third when viewed
dorsally; paired sinuous and relatively long genital filaments. Cerci as figured.
Material examined: Colombia. 3 holotype (no. 537), Cerro Munchique
(Cauca), 2450 m above sea level, light trap in cloud forest, 6 Aug. 1973, D.G.Y.
& R.C.W. paratype (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. trispinosa 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.












































c D F
Median Distal Proc...
of Aedeag\ Metothoracic Spirac
Sperm Pump \ r T





01 H
1G enital Filament
0.03


FIGURE 2
Sycorax andicola 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: Cerro Munchique, Cauca Dept., Colombia






2. Sycorax colombiensis n. sp.
(Fig. 3)
Male holotypee): Winglength 1.12; width0.47. Headheight0.21; width0.25;
eyes separated by 0.126 mm or by distance = to 10 facet diameters. Flagellomere
I (0.19 mm long), about 2.2X length of flagellomere 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. Genitalia: Coxite (0.177 long); style (0.98 long) with
a terminal spine, a long subterminal bristle and 2 proximal spines. Paramere as
figured, the lower distal projection blade-like, pointed at tip. Aedeagal complex:
Sperm pump (0.10 long); median distal process (0.09 long), slender and sub-
equal in width in dorsal view, with a slender, rounded tip in lateral view, a sub-
triangular plate, dorsoventrally flattened, with acute bifurcate tips, pointing to
the rear, present below the base of the median process of aedeagus. Cerci as
shown.
Female (allotype): Wing length 1.37; width 0.59. Head height, 0.24; width
0.29; eyes separated by 0.15 or by distance = to 10 facet diameters. Antenna
(1.15 long), flagellomere I (0.22 long), about 2.4X length of flagellomere II;
paired ascoids on flagellomeres I-IV; other sensory hairs 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 ep-
isternal seta. Wing venation as shown. Legs partly or wholly missing in all spec-
imens, 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 ap-
parently lacking these openings. Spermatheca 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), Anchicayi Dam (Valle),
elev. ca. 400 m above sea level, light trap, 28 Jan. 1975, R.C.W. 9 allotype (no.
530), same data as holotype except collected 10 June 1975. Paratypes (nos.
531-538), all from type locality in light traps; 1 9, 11 August 1973, D.G.Y.
& R.C.W 1 S, 2 99,28 Jan. 1975; 2 99, 10 June 1975, R.C.W
Discussion: The male of S. colombiensis, 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. colombiensis share several character states with S. fairchildi and
S. trispinosa but may be identified by the characters in the key.

3. Sycoraxfairchildi n. sp.
(Fig. 4)
Male 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










B


/ ,




F


G \h. )


FIGURE 3
Sycorax colombiensis A. Male head, B. Female wing, C. Male wing, D. Female
head, E. Male genitalia, dorsal view, F. Male subtriangular plate, G. Spermathecae, H.
Sperm pump, genital filaments and median dorsal process of aedeagus, lateral view.
Male: Anchicayi Dam, Valle Dept., Colombia
Female: Same locality as male






about 0.79 long; flagellomere I (0.18 long), about 2.5X length of flagellomere
II; paired ascoids visible only on flagellomeres I-III, other sensory hairs in a
whorl-like pattern on flagellomeres I- XV, palpal length 0.10 mm; ratio of seg-
ments: 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 appar-
ently 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 par-
amere terminating in a hook-like projection, acute at tip. Aedeagal complex:
Sperm pump (0.11 long); median distal process (0.068 long), angular at tip and
relatively wide in lateral view, subequal in width when viewed dorsally; aedea-
gal 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. Flagello-
mere I (0.16 long) about 2.3X length of flagellomere II, paired ascoids, sube-
qual in size (but not conspicuous) on flagellomeres 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 episternal setae (n = 3). Wing venation as
shown. Length of femora, tibiae, and basitarsi: Foreleg, 0.39, 0.40, 0.22; mid-
leg, 0.42, 0.47, 0.24; hindleg, 0.47, 0.50, 0.20. Spermatheca + sperm duct
length about 0.08 from base to end of subcircular button; furca somewhat "v"
shaped apically.
Material examined: Colombia 6 holotpye (no. 539), AnchicayL Dam
(Valle) elev. ca. 400 m above sea level, light trap in forest, 11 Aug. 1973,
D.G.Y. & R.C.W 9 allotype (no. 540), same data except collected 28 Jan.
1975, R.C.W Paratypes (nos. 541-547) all collected in light traps, 1 6, Rio
Anori (Antioquia), Colombia, elev. ca. 410 m above sea level, 22 Sept. 1970,
D.G.Y. 1 c, Rio Anori, 23 Sept, 1970. 1 9, Anchicayi Dam, 11 Aug 1973,
D.G.Y. & R.C.W 3 3J, 1 9 Anchicayi Dam, 28 Jan. 1975, R.C.W 1 6,
Anchicayi Dam, 5 March 1976, R.C.W.
Discussion: The short sperm ducts of S. fairchildi serve to distinguish this
taxon from the other species of Sycorax in Colombia.
I take pleasure in naming the species in honor of Dr. G. B. Fairchild who has
contributed so much to our knowledge of medically important arthropods in
the neotropics.

4. Sycorax trispinosa n. sp.
(Fig. 5)
Male 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 1.8X length of flagellomere II. Paired
ascoids on flagellomeres I- IV, absent from remaining flagellomeres; other an-








0.5




B





C


FIGURE 4
Syycoraxfairchildi A. Male head, B. Female wing, C. Male wing, D. Female head,
E. Male genitalia, dorsal view, E Sperm pump, genital filaments, and median dorsal
process of aedeagus, lateral view, G. Spermathecae, H. Female pedicel and flagellom-
eres I and II showing paired ascoids and other setae.
Male: AnchicayA Dam, Valle Dept., Colombia
Female: Same locality as male


CO'









B AD





cf c


F 0.5






G












\)1j '


C`





(?l \


FIGURE 5
Sycorax trispinosa A. Male head, B. Female palpus, C. Female, terminal 4 flagel-
lomeres, D. Female pedicel and flagellomeres I and II showing ascoids, E. Female head,
F. Female wing, G. Male wing, H. Male genitalia, dorsal view, I. Sperm pump, genital
filaments, and median dorsal process of aedeagus, lateral view, J. Tip of furca, dorsal
view, K. Spermathecae.
Male: Anchicaya Dam, Valle Dept., Colombia
Female: Same locality as male






tennal setae mostly deciduous, not arranged in a whorl-like pattern. Palpal
length, 0.12 mm; ratio of segments: 1- 1.15-0.84- 0.76, second segment with
15 rod sensilla. Pleura with 9- 16 setae below wing base and 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. Gen-
italia: 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 pro-
jection also curved at tip, finger-like. Aedeagal complex of a laterally com-
pressed sperm pump (0.11 long), a median distal process (0.10 long), with the
basal two-thirds expanded, greater than twice the width of the terminal third in
dorsal view; aedeagal ducts and cerci as shown.
Female (allotype): Wing length 1.54; width 0.71. Head height, 0.31; width,
0.27; eyes separated by distance of 0. 1 or by distance = to 6.9 facet diameters.
Antenna, 1.0 long; flagellomere I (0.16 long), about 1.7 x length of flagellom-
ere II Paired ascoids on flagellomeres I- VIII, absent from remaining flagello-
meres, other antennal setae mostly deciduous not arranged in a whorl-like pat-
tern. 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 ofspermatheca and sperm
duct, from base to tip of sclerotized button, 0.15. Furca spade-shaped as shown.
Material examined: Colombia. 6 holotype (no. 548), Anchicayi Dam (Valle),
elev. ca. 400 m above sea level, light trap in forest, 5 March 1976, R.C.W. 9
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 9Y, 10 Aug.
1973; 1 6, 1 9, 11 Aug. 1973; 13 &6, 4 9, 28 Jan. 1975; 1 S, 2 99, 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. fair-
childi or S. colombiensis. Flagellomere I relatively short (0.14-0.19 mm,
n = 27); compound eyes more narrowly separated than those of the other sym-
patric 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, although quite similar, differ in
size and by other characters given in the key. The female of andicola, presently
unknown, probably resembles trispinosa in details of the spermathecae and
palpi.
The specific name refers to the 3 strong spines on the style of the male.










SUBFAMILY PHLEBOTOMINAE


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 systematics, especially spe-
cies descriptions and distribution records. Owing to the paucity of bionomic
studies on the Colombian Phlebotomines, I felt that it would be desirable to
provide references to such studies carried out in other countries. The results of
these studies may not always apply to conspecific populations in Colombia. New
information on the biting habits of some species in Choc6 Dept. is provided in
the species accounts.
With few exceptions I have not redescribed known taxa because of adequate
original or subsequent descriptions cited in the taxonomic references.

KEY TO THE NEW WORLD GENERA
I. Wing broad, rounded at tip; R2 + R3 + R4 forks before, on same
level as, or slightly beyond r-m crossvein. Pleura without
espisternal setae. Female cibarium unarmed. Male genitalia
with style longer than coxite ................... Warileya (Fig. 6-7)
Wing pointed at tip; R, + R3 + R4 forks well beyond r-m crossvein.
Pleura with episternal setae. Female cibarium armed with teeth.
Male genitalia with style shorter than coxite ........... .......... 2
2. Interocular suture complete. Female cibarium with 4 longitudinal
rows of horizontal teeth. Male genitalia with 5 large spines,
2 of which (usually basal pair) borne on a common tubercle
................ ............... Brumptomyia (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 WARILEYA HERTIG, 1948
This genus, recently enlarged to accommodate W (H.) hertigi (Fchld.) by
Lewis et al. (1977), contains 5 species, 2 of which in the subgenus Warileya
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) phlebotoman-
ica Hertig, the type species from Per6, 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 .............................. nigrosacculus (Fig. 6)
Palpal segment 5 shorter than combined length of segments 2 + 3.
Intraabdominal rods absent. Genital filaments less than 1.3X
length of pump. Style with a basal bristle. Wing length over
1.8 mm .................................. rotundipennis (Fig. 7)

Females
1. Spermathecae large and sac-like with smooth walls. Palpal segments 5
longer than combined length of segments 2 + 3. Wing length less than
2.0 mm ............................... nigrosacculus (Fig. 6)
Spermathecae cylindrical, somewhat worm-like, with transverse
striations. Palpal segment 5 shorter than combined length of segments
2 + 3. Wing length over 2.0 mm .............. rotundipennis (Fig. 7)



5. Warileya (W) nigrosacculus Fairchild & Hertig
(Fig. 6)
Warileya nigrosacculus Fairchild & Hertig, 195 b; 428 (? holotype, Cerro
Campana, Panama Prov., Panama). Fairchild, 1953: 102 (cf. to Hertigia her-
tigi). Barretto, 1955a: 188 (listed). Forattini, 1971a: (107) (Listed). Tesh et al.,
1971a: 153 (blood meals, Panama). Martins & Morales, 1972: 366 (listed).
Young & Chaniotis, 1972: 97 ( descr.). Chaniotis et al., 1972: 95- 96 (in tree
hollows, Panama). Christensen, 1972a: 88 (listed). Forattini, 1973: 536-538
( fig.). Velasco & Trapido, 1974: (cf. to W yungasi). Lewis, 1975a: 500 et seq.
(mouthpart morphol.). Lewis et al., 1977: 325 classicf.).
Distribution: Colombia (Choc6, Valle), Panamd.
Material examined: Colombia: 1 ', 1 Curiche (Choc6), Malaise trap, 27
April 1977. 1 6, 1 9, Anchicaya Dam (Valle), light trap, 28 Jan. 1975, R.C.W.
1 6, same data but taken 16 July 1975. Panama: 9 holotype, Cerro Campana
(Panama), hollow log, 7 Jan. 1947, M. Hertig. 2 6d, 5 99, near Gamboa,
Canal zone, tree hollow, 10 Sept. 1969, B. Chaniotis. 4 cd 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 re-
cently engorged nigrosacculus females reacted with mammalian and reptile-
amphibian antisera. Four other blood meals were nonreactive due to small vol-
umes of blood and/or to the weakness of the reptile-amphibian antisera.













B
B


D


( I
, ,


0.05


G 005


1.0


H



I'--


FIGURE 6
Warileya nigrosacculus A. Male head, B. Male flagellomere II, C. Female head, D.
Female flagellomere II, E. Female cibarium and pharynx, E Male genitalia, dorsal, G.
Tips of genital filaments, H. Female wing, I. Male wing, J. Spermathecae.
Male: Curiche, Choc6 Dept., Colombia
Female: Same locality as male


J






6. Warileya (W.) rotundipennis Fairchild & Hertig
(Fig. 7)
Warileya rotundipennis Fairchild & Hertig, 195 lb: 424 ( holotype, 9, Cerro
Campana, Panama Prov., Panama). Fairchild, 1953: 102 (cf. to Hertigia her-
tigi). Barretto, 1955a: 188 (listed). Fairchild & Hertig, 1959: 122 (Costa Rica).
Johnson & Hertig, 1961: 765, 775 (rearing). Hanson, 1968: 93 (larval fig.).
Forattini, 197 a: 107 (listed). Young & Chaniotis, 1972: 366 (listed). Christen-
sen, 1972a: 88 (listed). Forattini, 1973: 536-538 (d, 9 fig.). Velasco & Trap-
ido, 1974: 436 (cf. to W yungasi). Lewis, 1975a: 500 et seq. (mouthpart
morphol.). Lewis, 1975b: 366 (hair sockets, fig.). Lewis et al,: 1977: 325
(classif.).
Distribution: Colombia (Antioquia, Choc6, Valle), Costa Rica, Panamd.
Material examined: Colombia. 2 99, Teresita (Choc6), tree buttress, 15 June
1967. 27 99, Rio Anori (Antioquia), light traps, Sept. 1970. 5 99, same data
but 29 May 1970, C.H.P. 8 dc, 2 99, Anchicaya Dam (Valle), light trap, 28
Jan. 1975, J. E. Browne. 6 dd, 6 9, same data but taken 10 June 1975, R.C.W.
5 d 4 Y9 same data but taken 16 July 1975. Panamd. S holotype (no. 2335),
9 allotype (no. 2312), Cerro Campana (Panama), Shannon trap, 24 Aug. 1950,
M. Hertig & G. Fairchild. 2 SS, Rio Changena (Bocas del Toro), Shannon trap,
8- 11 Sept. 1961, R. Hartmann & P. Galindo. CostaRica. 2 SS, 1 9, Turrialba
(Cartago), Shannon trap, 20 Aug. 1961, G. Fairchild & M. Hertig.
Discussion: Warileya rotundipennis, the only Warileya species lacking intra-
abdominal rods (Lewis et al., 1977), has been reported feeding on man in
Panama (Fairchild & Hertig, 1951 b), but its role, if any, in disease transmission
is not known. Dr. C. H. Porter also has collected man-biting females at the Rio
Anori (Antioquia) locality, mostly in forest clearings (pers. comm.).
The records of W rotundipennis and W nigrosacculus from Choc6 and Valle
Depts. indicate that both taxa probably occur in other areas of the trans-Andean
region perhaps as far south as Guayas Prov., Ecuador.


GENUS BRUMPTOMYIA FRANCA AND PARROT, 1921
Characterized by Theodor (1965), Lewis et al. (1977), and others, the genus
Brumptomyia now contains over 21 species, 5 of which occur in Colombia. Ex-
cept for the female of B. hamata, which has very short sperm ducts (Fig. 101),
the females of the other species in Colombia are virtually impossible to identify
in the absence of males.
Brumptomyia avellari, one of the Colombian species, was reported feeding
on armadillos in Brazil (Mangabeira, 1942b). This and other Brumptomyia
spp. often rest in burrows dug by these animals. They do not bite man, and the
preferred hosts of most species remain unknown.














B


1.0


GP~ c


HC


FIGURE 7
Warileya rotundipennis A. Male head, B. Male flagellomere II, C. Female head, D.
Female flagellomere II, E. Female cibarium and pharynx, E Male genitalia, G. Female
wing, H. Male wing, I. Spermathecae.
Male: Cerro Campana, Panama Prov., Panama
Female: Rio Anori, Antioquia Dept., Colombia


E 'I






(U






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. Parameres
more slender as shown .......................... galindoi (Fig. 9)
Coxite tuft of fewer than 20 setae, usually about 12, not implanted
on a tubercle. Paramere broader as shown ............ avellari (Fig. 8)
4. Style with isolated spine well distad of proximal pair. Coxite tuft
implanted on a distinct raised tubercle ............ beaupertuyi (Fig. 8)
Style with isolated spine at or near level of proximal spines. Coxite
tuft implanted on a slightly raised,
raspberry-like (circular) base .................... leopoldoi (Fig. 11)


7. Brumptomyia avellari (Costa Lima)
(Fig. 8)
Phlebotomus avellari Costa Lima, 1932: 48 (5, Lassance, Minas Gerais,
Brazil). Mangabeira, 1942b, 225 et seq. (immatures, adults, descr., fig.). Bar-
retto, 1947:187-188 (full refs.). Fairchild & Hertig, 1947a: 615-616 (cf. to gal-
indoi). Barretto, 1951: 212 (dist.). Floch & Abonnenc, 1952: 39, 45 (C, 9,
keyed). Rodriguez, 1953b: 55 (mention). Forattini, 1954: 214-217 (second
sternite, fig.). Forattini & dos Santos, 1955: 17 (Brazilian record). Ortiz, 1963:
320 (9, keyed).
Brumptomyia avellari: Barretto, 1955a: 187 (listed). Martins et al., 1961b:
309 (mention). Martins et al., 1962d: 380 (Goias, Brazil). Sherlock, 1962:
332, 335 (mention). Carnheiro & Sherlock, 1964: 315 (pupa, keyed, fig.).
Fraiha et al., 1970a: 468 (6 keyed). Christensen, 1972a: 88 (Panama). Osorno
et al., 1972a: 14 (Boyaca, Colombia). Forattini, 1973: 122 et seq. (gen. review,
fig.). Llanos et al., 1976:480 (Peru). Martin et al., 1976b: 496 (Per6). Ramirez
et al., 1976: 599 (Venezuela).
Distribution: Panama, Colombia (Boyaca), Pera, Venezuela, Brazil, Para-
guay.
Material examined: Colombia. 2 &T, 1 9, Puerto Boyaca (Boyaca), light
trap, 6 May 1973, C.J.M. Brazil. 1 6, Coqueiros (Si~o Paulo), armadillo bur-
row, Oct. 1953, Albertin. Panama. 1 6, Tocumen Airport (Panama), light
trap, 23 March 1953, F Blanton. Paraguay. 5 dd, Aca-Poi, San Pedro, bur-
row, 18 April 1950, M. Hertig.














A 0.
.0


.-.. `


CI
I'



c


0'05 E
D





F


~I


G

v\)Y,^


FIGURE 8
Brumptomyia avellari male A. Genitalia.
Male: Puerto Boyaca, Boyaci Dept., Colombia

Brumptomyia beaupertuyi male B. Head, C. Flagellomere II, D. Paramere, E. Wing,
F Genital pump and filaments, G. Genitalia, H. doxite tuft, same scale as Fig. 8G.
Male: Lara State, Venezuela


0o






Discussion: Brumptomyia avellari and B. brumpti (Larousse), with sym-
patric populations in Brazil and Paraguay, closely resemble one another, the
males differing chiefly by the shape of the parameres. I regard the Colombian
specimens as being conspecific with the former species.

8. Brumptomyia beaupertuyi (Ortiz)
(Fig. 8)
Phlebotomus beaupertuyi Ortiz, 1954: 235 (d holotype, Duaca, Lara State,
Venezuela). Scorza & Ortiz, 1960: 434 et seq. (ecology). Pifano et al., 1962:
383, 387 (8, 9, keyed), 411-412 (d, 9, descr., fig.). Ortiz, 1963: 322 (9
descr.). Ortiz & Scorza, 1963: 350 (listed), 354 (d keyed). Ortiz, 1965a: 208
(mention). Scorza et al., 1967: 193, 195 (d, 9 keyed). Mirmol Le6n, 1968:
30 (listed).
Phlebotomus galindoi (not galindoi of Fairchild & Hertig, 1947a): Pifano
& Ortiz, 1952: 138 (listed, Venezuela). Marmol Le6n, 1968: 30 (listed).
Brumptomyia beaupertuyi: Barretto, 1955a: 187 (listed). Sherlock, 1962:
321 et seq. (d, fig., Colombia). Fraiha et al., 1970a: 468 (8, keyed). Forattini,
1971a: 98 (listed). Forattini, 1973: 522 et seq. (gen. review, fig.).
Distribution: Colombia (Santander), Venezuela.
Material examined: Venezuela. 2 &d (topotypes), Duaca (Lara), no date, I.
Ortiz.
Discussion: At present, this little-known species is represented in Colombia
by a single male, skillfully illustrated by Dr. O. Mangabeira (in Sherlock,
1962). Ortiz (1963) described the female based on Venezuelan specimens.

9. Brumptomyia galindoi (Fairchild & Hertig)
(Fig. 9)
Phlebotomus galindoi Fairchild & Hertig, 1947a: 615 (c holotype, near Bo-
quete, Chiriqui Prov., Panama). Floch & Abonnenc, 1952: 40 (d keyed). Or-
tiz, 1954: 238 (cf. to beaupertuyi). Rosabal, 1954: 30 et seq. (8 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 descr., Mexico, fig.). Ortiz & Scorza,
1963: 354 (d keyed). Biagi et al., 1966: 149 (Mexican records), 151 (S fig.).
Strangways-Dixon & Lainson, 1966:193 (Belize). Hanson, 1968: 56-58 (larva,
pupa, descr., fig.). Marmol Le6n, 1968: 30 (listed).
Brumptomyia galindoi: Barretto, 1955a: 187 (listed). Osorno et al., 1972a:
14-15 (Colombian records). Forattini, 1973: 122 et seq. (gen. review, fig.).
Lewis, 1975a: 500 et seq. (mouthpart morphol.).
Brumptomyia mesa Sherlock, 1962: 332 (d, San Vicente de Chucuri, San-
tander, 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).

















D


E


G


\


0.05


K ..


L 9


0


FIGURE 9
Brumptomyia galindoi A. Male head, B. Male flagellomere II, C. Female head, D,
Female flagellomere II, E. Female cibarium and pharynx, E Genital pump, G. Genital
pump and filaments, H. Male genitalia, I. Paramere, J. Coxite tuft, K. Spermathecae, L.
Body of spermatheca, M. Female wing, N. Male wing, 0. Female cibarium.
Male: Teresita, Choc6 Dept., Colombia
Female: Same locality as male


3


~ ,,


M9



NCT






Distribution: Mexico, Belize, Honduras, Costa Rica, Panama, Colombia
(Caqueta, Choc6, Bolivar, Boyaca), Ecuador, Paraguay.
Material examined: Colombia. 1 6, Puerto Boyaca (Boyaca), light trap, 6
May 1973, C.J.M. 1 c, Tres Esquinas (Caqueti), light trap, 11 Nov. 1971,
C.J.M. 27 d3, 11 9, Teresita (Choc6), light & Malaise traps, tree trunks,
April-Aug. 1967, D.G.Y. Costa Rica. 1 S, San Carlos (Alajuela) tree but-
tress, 23 June 1955, R. Rosabal. Ecuador. 5 6d, 4 99, Rio Napo at Limon-
cocha (Napo), light & flight traps, tree buttresses, 20-24 May 1976, D.G.Y.
& T. Rogers. Honduras. 1 6, Tela, Lancetilla Valley, tree buttress, 24 July
1953, W. Hils. 1 S, same data but light trap, 13 Jan. 1954. Mexico. 2 CS, 2
Y9, Ocoscoautla (Chiapas), tree hollows, 8 April 1951, G. Fairchild & R.
Hartmann. 2 &T, Palenque (Chiapas), tree buttresses, 30- 31 March 1951, G.
Fairchild & R. Hartmann. Paraguay. 1 S, Sommerfeld, Yhu, tree cavity, 15
March 1950, V. Zelada. 1 6, same data but 18 March 1950, M. Hertig. Pan-
amd. 1 & holotypee no. 814), type locality, 17 Feb. 1947, P Galindo. 1 Y,
same data but 26 March 1948. 1 3, 1 Y, Mojinga Swamp near Ft. Sherman,
Canal Zone, light trap, 19 Nov. 1951, F. Blanton.
Discussion: The holotype of B. galindoi is a large specimen, wing length
nearly 3.0 mm, from Chiriqui Province, Panama (ca. 1200 m above sea level).
Smaller specimens from lower elevations in Santander Dept., Colombia were
later named B. mesa 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 Panami.
Aside from size and the number of distal coxite setae (7 in the galindoi hol-
otype, 5-6 in males from the lowlands), I can detect no marked differences
among the specimens listed above. The number of distal setae on the coxites
probably varies according to the size of the insect, and in this case, apparently
represents infraspecific variation. It remains to be determined whether or not
the size of galindoi individuals is correlated with altitude; i.e., is there a con-
tinuous increase in size of specimens from the lowlands to the highlands? Be-
lieving this to be the case but without evidence to support it, I tentatively
consider B. mesa and B. galindoi to be conspecific following Fraiha et al.
(1970a).

10. Brumptomyia hamata (Fairchild & Hertig)
(Fig. 10)
Phlebotomus hamatus Fairchild & Hertig, 1947a: 614 (< holotype, Chili-
brillo bat caves, near Chilibre, Panama). Barretto, 1951: 217 (mention). Floch
& Abonnenc, 1952: 39 (6 keyed). Rodriguez, 1953b: 53 (6, Y measured, Ec-
uador), 55 (cf. to leopoldoi). Ortiz, 1954: 237 (listed). Lewis & Garnham,
1959: 83-84 (6, Belize, fig.). Garnham & Lewis, 1959: 24 (Belize). Hanson,
1961: 320 et seq. (breeding sites, Panama). Johnson & Hertig, 1961: 765 et

















D
B

C E
A









H


00o













S. M
L

FIGURE 10
Brumptomyia hamata A. Male head, B. Male flagellomere II, C. Female head, D.
Female flagellomere II, E. Female cibarium and pharynx, F Genital pump and filaments
of male from Teresita, Choc6 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 cibarium.
Male: Barro Colorado Island, Panama Canal Zone (except Fig. 10F)
Female: Teresita, Choco Dept., Colombia






seq. (rearing data). Diaz-Najera, 1963: 193 et seq. (Mexico). Ortiz & Scorza,
1963: 353 (d, keyed). Disney, 1966: 449 (66 in rodent-baited traps, Belize).
Hanson, 1968: 58 (larvae identical to those of galindoi).
Brumptomyia hamata: Barretto, 1955a: 187 (listed). Sherlock, 1962: 332
(cf. to mesai. Williams, 1970: 331 (listed, Belize). Fraiha et al., 1970a: 468
(6 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, Panamd, Colombia (Choc6), Ecuador.
Material examined: Colombia. 1 d, Teresita (Choc6), hollow tree, 25 March
1967, D.G.Y. 1 9 same data but Malaise trap, 17 June 1967. Panamd. 1 &
holotypee no. 112), type locality, 4 Dec. 1943. 4 dd, 5 99, lab reared from
larvae collected in Canal Zone, 1957- 1958, W. Hanson.
Discussion: Although the adults of B. hamata are rarely encountered, the
larvae may be quite common in soil at the base of trees. Hanson (1961) and
Rutledge & Mosser (1972) in Panami found that the larvae of this species far
outnumber those of other phlebotomine species in this microhabitat. Like other
Brumptomyia spp., the fourth instar larva has only two caudal setae.

11. Brumptomyia leopoldoi (Rodriguez)
(Fig. 11)
Phlebotomus leopoldoi Rodriguez, 1953b: 52 (d holotype, Quevedo, Los
Rios Prov., Ecuador; 9, Naranjal, Guayas Prov., Ecuador). Ortiz, 1963: 320
(9 keyed).
Brumptomyia leopoldoi: Barretto, 1955a: 187 (listed). Sherlock, 1962: 332
(cf. to mesai. Williams, 1970: 331 (Belize). Fraiha et al., 1970a: 468-469
(S keyed, as possible synonym of guimaraesi). Forattini, 1971a: 98 (listed).
Christensen, 1972a: 88 (listed, Panama). Forattini, 1973: 530 (as synonym of
guimaraesi). Williams, 1976a: 603 (in caves, Belize).
Distribution: Belize, Panamd, Colombia (Antioquia, Choc6, Valle),
Ecuador.
Material examined: Colombia. 3 d~, Rio Anori (Antioquia), light traps, May
1970, C.H.P. 1 6, Curiche (Choc6), light trap, 26 May 1967, D.G.Y. 2 ~d, 5
99, ca. 10 km W of Cali near Pichind6 (Valle), rock crevice, tree trunks and
light trap, 31 July 1973-4, Aug. 1973, D.G.Y. & R.C.W. Ecuador. 1 5 (para-
type no. 4314), Quevedo (Los Rios), no date, J. Rodriguez. 2 d6, 2 99, 17 km
E of Santo Domingo de Los Colorados (Pichincha), mammal burrow, 4 May
1976, D.G.Y. & T. Rogers. 1 c, same data but 27 May 1976. 1 S, same data
but light trap, 27 May 1976. Panamd. 1 & (paratype no. 4292), Almirante (Bocas
del Toro), animal burrow, 22 Jan. 1953, W. Hils. 1 ~ same data but tree buttress,
27 Jan. 1956, R. Hartmann.
Discussion: B. leopoldoi, considered to be conspecific with B. guimaraesi
(Coutinho and Barretto) by Forattini (1973), is specifically distinct, judged by
the following considerations:



















6 C
B C


E


F
-. ( l.


IL 7/


G C


FIGURE 11
Brumptomyia leopoldoi male A. Head, B. Flagellomere II, C. Coxite tuft, paramere,
and aedeagus of male from near Santo Domingo de Los Colorados, Ecuador, D. Wing,
E. Genital pump and filaments, E Genitalia, G. Genital pump of male from Fig. 11C
locality.
Male: Rio Anori, Antioquia Dept., Colombia (except for Figs. 11 C and G)
Brumptomyia guimaraesi male H. Genital pump, I. Coxite tuft, paramere, and ae-
deagus, same scale as 11C.
Male: Itaporanga, Sio Paulo State, Brazil


1,0



D


0r



"^






1. The 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
1 I). The original figure of leopoldoi (Rodriguez, 1953b) shows a rather
broad paramere, but I believe this is a distortion due to mounting tech-
nique, not a reflection of normal 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 mm) than that of leopoldoi which is 0.22-0.24 mm long
(cf. Fig. 11G and 11H).
3. The isolated spine of the style of guimaraesi is inserted at 0.59-0.61 of
the segment whereas that of leopoldoi is located at 0.53.
In addition to the leopoldoi specimens listed above, I examined 4 guimaraesi
males from southern Brazil: 2 dd Faz. Ribeirao de Baizo, Patos, Minas Gerais;
1 6, Coqueiros, Sao Paulo; 1 & reared from egg laid by 9 captured at the type
locality (Itaporanga, Sao Paulo, Brazil). I conclude that these allopatric species
are distinct, B. guimaraesi occurring only in southeastern Brazil as far as is
known.


GENUS LUTZOMYIA FRANCA, 1924
As mentioned earlier, the classification of the Lutzomyia sand flies is based
upon the scheme of Lewis et al. (1977). It should be emphasized that this rep-
resents a flexible classification, subject to change as new species, new character
states, or unknown sexes are discovered.
It is hoped that the following keys and discussions will be helpful to those
interested in sand flies and disease in Colombia and elsewhere in the neotropics.

Keys to Subgenera, Species Groups, and Ungrouped
Species of Lutzomyia
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 Lutzomyia subgenera and equivalent species groups
are defined by the characters given here. It is important to note, however, that
species in the same groups, presently unknown in Colombia, may not exactly
"fit" into a subgenus or species group as characterized by these keys. For ex-
ample, L. nevesi (Damasceno and Arouck) belongs in the verrucarum 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 Putu-
mayo Intendencia, Colombia.
As additional species are discovered in the Republic, these keys obviously
will have to be revised. For the present, they are intended to be used with cau-
tion, applicable only to the Colombian fauna.
42






Males
1. Palp 5 very short, less than half length of third and less than twice
length of fourth ......... subgenus Psychodopygus (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 squamiventris (Fig. 61)
Style with 2-6 major spines. Coxite not indented near middle ....... 3
3. (2) Style with 2 major spines, both terminal. Paramere
simple ...................... series arthuri, L. bispinosa (Fig. 63)
Style with 3-5 (sometimes 6) major spines. Paramere simple or
complex ......................... series panamensis (Fig. 64-71)
4. (1) Lateral lobes markedly inflated ...... vespertilionis group (Fig. 29)
Lateral lobes not inflated ....................... ............ 5
5. (4) Lateral lobes with spatulate setae at
tips ................... subgenus Evandromyia, series infraspinosa*
Lateral lobes without spatulate setae ........................... 6
6. (5) Style deeply forked. Parameres with 2-4 modified apical setae as
shown ....................... subgenus Viannamyia (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 dorsal setiferous arm. Coxite without nondeciduous
setae ................. subgenus Dampfomyia, L. rosabali (Fig. 30)
Paramere simple, without a dorsal arm. Coxite with a basal tuft or
group of setae ........... verrucarum group, series serrana (Fig. 22)
9. (7) Style with 3 major spines ................................ 10
Style with 4-6 major spines ................................. 11
10. (9) Coxite with a basal tuft of simple and modified setae. Style with a
small subterminal and a small median seta in addition to the major
spines ......................... subgenus Pressatia (Fig. 32-34)
Coxite without a basal tuft but with a distal group of 15 simple setae.

*Species of Evandromyia in this series have not been found as yet in Colombia, but it is
probable that at least one species occurs in the southeastern part of the Republic. The
subgenus was reviewed by Young and Arias (1977).






Style with only a small median seta and 3 major spines, no subterminal
seta ......................... .. pilosa group, L. pilosa (Fig. 86)
11. (9) Style with 4 major spines ............... ............ 12
Style with 5, sometimes 6, major spines* .................. .... 32
12. (11) Antennal ascoids with very long pointed or short blunt basal spurs.
Style without a subterminal seta ............................ 13
Antennal ascoids with or without basal spurs, but if present they are
very short and pointed. Style with or without a subterminal seta ..... 14
13. (12) Coxite with 5-8 strong distal setae. Style with isolated basal
spine ... ...................... dreisbachi group (Fig. 50-51)
Coxite without distal setae. Style with basal spines more or less
paired ............................ shannoni group (Fig. 37-42)
14. (12) Palp 5 shorter than palp 3. Style without a subterminal seta..... 15
Palp 5 as long as or longer than palp 3. Style with or without a
subterminal seta ....................................... 18
15. (14) Coxite with 2 or more basal and/or median setae ............. 16
Coxite without nondeciduous setae ........................... 17
16. (15) Style with basal spine isolated. Antennal ascoids with very short
pointed basal spurs. Flagellomere I shorter than flagellomeres II +
III ...................... subgenus Trichophoromyia (Fig. 51- 54)
Style with paired basal spines. Antennal ascoids simple,
no basal spurs visible. Flagellomere I
longer than II + III subgenus Nyssomyia (in part), L. antunesi (Fig. 55)
17. (15) Antennal ascoids with short, pointed basal spurs.
Paramere with a dorsobasal hump. Flagellomere I
shorter than II + III ................... aragaoi group (Fig. 45-49)
Antennal ascoids simple. Paramere lacking a dorsobasal hump.
Flagellomere I longer than II + III ...........................
................. subgenus Nyssomyia (in part), (Fig. 55-60)
18. (14) Coxite with 3- basal and/or median setae in a tuft or group;
distal setae may also be present ............................ 19
Coxite without a basal and/or median group of setae but long ventral
setae may be present ...................................... 28
19. (18) Style with 4 short stubby spines, 2 of which are terminal;
subterminal seta present. Paramere simple, undivided. Coxite with
4-8 strong distal setae ............. baityi group, L. baityi (Fig. 34)

*Males of L. rorotaensis usually have 5 major spines, but those examined from Valle
Dept. have but 4 (see p. 227).






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 Lutzomyia, series longipalpis (Fig. 13- 15)
Paramere without such setae ............................... 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 distinct
inner "tooth" ........... migonei group, series walker (Fig. 19-20)
Genital filament tips enlarged or not but lacking an inner tooth ...... 24
24. (23) Coxite with setae of basal tuft shorter than width of coxite ....
......................... migonei group, L. migonei (Fig. 18)
Coxite with setae of basal tuft longer than width of coxite .......... 25
25. Mesonotum & pleura pale ...... saulensis group,*L. saulensis (Fig. 21)
Mesonotum distinctly darker than pleura .....................
........ verrucarum group, series verrucarum (in part), (Fig. 23-24)
26. (22) Coxite with basal setae inserted on a raspberry-like, subcircular
base .............. subgenusLutzomyia, series cruciata (Fig. 16- 17)
Coxite with basal setae, if present, not implanted on such a base ..... 27

27. (26) Style with paired basal spines. Coxite with basal tuft of slender,
subequal setae ..................... .................
verrucarum group, series verrucarum (in part), L. nuneztovari (Fig. 23)
Style with basal spine isolated. Coxite with either basal-median setae,
1 at least larger than others, or with a patch of slender distal
setae ............ cayennensis group, series atroclavata (Fig. 81- 82)
28. (18) Style with a small subterminal seta ...................... 29
Style without a small subterminal seta ................... 31
29. (28) Paramere divided. Coxite with long ventral setae ............
....... ...................... longispina group (Fig. 43-44)
Paramere simple, undivided. Coxite without long ventral setae ...... 30

*I am unable to separate the males on the group level using structural characters.






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. nordestina (Fig. 87)
Style with paired basal spines. Flagellomere I shorter than head height;
ascoids without visible basal spurs. Palp 5 longer than segments 3 +
4 .............. cayennensis group, series 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 ....................
..................... vexator group, seriesperuensis (Fig. 72-78)
Coxite without setae at base but with a few scattered setae near middle
of coxite ................... oswaldoi group (in part), (Fig. 84- 85)

Females
1. Spermathecae associated with.paired sclerotized structures,
intracellular ducts (= "hairs") covering most of spermathecae ....
....................... subgenus Viannamyia (Fig. 35-36)
Spermathecae without associated sclerotized structures, intracellular
ducts confined to small area, usually but not always, to terminal
knob.................. ............................ 2
2. (1) Pharynx with posterior spines ............................. 3
Pharynx without spines ...................................... 6
3. (2) Cibarium with inner pair of horizontal teeth pointing inwards ...
........ ....... oswaldoi 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 ........
cayennsis group, series cayennensis (in part), L. cayennensis (Fig. 79)
Cibarium with 4 horizontal teeth not in a comb-like row ............ 5
5. (4) Foreleg with femur longer than tibia. Pharynx markedly enlarged
with prominent, subequally spaced transverse ridges and conspicuous
spines. Pleura pale.. cayennensis group, series atroclavata (Fig. 81- 82)






Foreleg with femur shorter than tibia. Pharynx more slender, the
transverse ridges and spines less defined. Pleura dark ............
cayennensis group, series cayennensis (in part), L. micropyga (Fig. 80)
6. (2) Spermathecae with bubble-like expansions ................... 7
Spermathecae without bubble-like expansions .................... 8
7. (6) Cibarium with 4 broad and blunt horizontal teeth and conspicuous
lateral teeth. Mesonotum well pigmented, darker than pleura. Eyes
very small ............. subgenus Dampfomyia, L. rosabali (Fig. 30)
Cibarium with 4 slender, sharp horizontal teeth, without obvious lateral
teeth. Mesonotum and pleura pale. Eyes larger .................
.......................... saulensis group, L. saulensis (Fig. 21)
8. (6) Antennal ascoids with long pointed or short blunt basal spurs ..... 9
Antennal ascoids simple or with short pointed basal spurs ......... 10
9. (8) Cibarium with 10+ horizontal teeth. Spermathecae distinctly
annulated ....................... dreisbachi group (Fig. 50-51)
Cibarium with 4- 8 horizontal teeth. Spermathecae annulated or
not ............................ shannoni group (Fig. 37-42)
10. (8) Hind femur with row of 3-5 short spines ...................
........................ subgenusPintomyia, L. spinosa (Fig. 31)
Hind femur without spines ............................ 11
11. (10) Cibarium with 6 or more horizontal teeth ................... 12
Cibarium with 4 horizontal teeth ........................... 17
12. (11) Individual sperm ducts markedly convoluted, twisted .........
............ aragaoi group, series brasiliensis, L. runoides (Fig. 49)
Individual sperm ducts not convoluted as above .................. 13
13. (12) Palp 5 equal to or shorter than segments 3 + 4 ............... 14
Palp 5 longer than segments 3 + 4 ............................ 16
14. (13) Spermathecae larger, sac-like, without annuli. Ascoids with short
pointed basal spurs ....... aragaoi group, series aragaoi (Fig. 45-48)
Spermathecae smaller, with incomplete or complete annuli. Ascoids
with or without short basal spurs ............................. 15
15. (14) Flagellomere I longer than flagellomeres II + III. Ascoids
with short basal spurs ....... subgenus Trichophoromyia (Fig. 51-54)
Flagellomere I shorter than II + III. Ascoids without visible basal
spurs .................. ...... subgenus Nyssomyia (Fig. 55-60)
16. (13) Cibarium with 6 horizontal teeth. Spermatheca with terminal
segment smaller than preceding segment. Common sperm duct longer






than individual ducts ....... oswaldoi group (in part), L. pia* (Fig. 83)

Cibarium with 8+ horizontal teeth. Spermatheca with terminal
segment as large as others. Common sperm duct much shorter than
individual ducts .......................... ...........
subgenus Lutzomyia, 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)
Spermathecae not imbricated, annuli, if present, not semi-telescoped.
Individual sperm ducts non-rugose, smooth walled or with slight
thickenings ........ ............... .................... 19
19. (18) Cibarium with inner pair of horizontal teeth pointing inwards.
Flagellomere I extremely long, subequal to or longer than combined
lengths of palpal segments. Ascoids with very short basal
spurs ................... ...... ungrouped, L. nordestina (Fig.87)
Cibarium with horizontal teeth pointing toward pharynx, not slanted
inwards. Flagellomere I shorter than palpus. Ascoids without visible
basal spurs. .......... ......... ... .... ............... 20
20. (19) Spermathecae as shown, longer than individual sperm ducts.
Abdominal tergite 8 without lateral setae ................ ....
............ ..vespertilionis group, series vespertilionis (Fig. 29)
Spermathecae otherwise, shorter than individual sperm ducts.
Abdominal tergite 8 with lateral setae ........................ 21
21. (20) Spermathecae less than twice as long as wide, terminal segment
smaller than preceding segment. Common sperm duct longer than
individual ducts ............ oswaldoi group (in part), L. pia (Fig. 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, series peruensis (in part), (Fig. 72-74; 76-78)
22. (17) Individual sperm ducts heavily pigmented in part or whole .....
.............................. subgenus Pressatia (Fig. 32- 34)
Individual sperm ducts not pigmented ....................... 23
23. (22) Cibarial arch absent or conspicuous only at sides ............. 24
Cibarial arch complete or nearly so ........................... 26

*L. pia is keyed twice because of intraspecific variation in the number of horizontal teeth
(seep. 225).






24. (23) Cibarium with horizontal teeth like flattened plates viewed almost
edgewise. Common sperm duct absent ................. ......
.............................. pilosa 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. Cibarium with
horizontal teeth pointing towards pharynx, not inwardly slanted .....
................ vexator group, series peruensis (in part), (Fig. 75)
Wing venation with beta over half length of alpha. Cibarium with
horizontal teeth pointing inwards .............................
................ oswaldoi group (in part), L. rorotaensis (Pig. 84)
26. (23) Spermathecae completely smooth-walled ................... 27
Spermathecae wrinkled, with transverse striations or indentations
forming complete annuli or not ............................ 28
27. (26) Spermathecae tubular, much longer than wide and only slightly
wider than individual ducts. Common sperm duct short, not exceeding
one-third length of individual ducts ...........................
.............. migonei group, series migonei, L. migonei (Fig. 18)
Spermathecae capsular, about as long as wide, distinctly wider than
individual ducts where they enter spermathecae. Common duct over
one-third length of individual ducts ..........................
..................... migonei group, series walker (Fig. 19-20)

28. (26) Spermathecae as shown, with some clearly defined annuli,
terminal one hemispherical and larger than others ................
..................................... .. subgenus Lutzomyia,
series longipalpis (in part), and series cruciata (Fig. 13- 14; 16- 17)
Spermathecae otherwise, with transverse striations or wrinkles,
terminal segment, if present, not hemispherical .................. 29
29. (28) Spermathecae as shown, pear-shaped with fine transverse
striations, complete or not ........... longispina group (Fig. 43-44)
Spermathecae otherwise, sac-like and wrinkled .................. 30
30. (29) Individual sperm ducts nearly absent, spermathecae seemingly
joined directly to common duct. Eyes very small .................
......... ......... ungrouped, L. rangeliana (Fig. 88)
Individual sperm ducts at least as long as half length of spermathecae.
Eyes large ....................... verrucarum group (Fig. 24- 27)


























Length


H i


ina PF J

enltal Pump Genital Filamenfr


FIGURE 12
Lutzomyia (P) hirsuta nicaraguensis female A. Head with palpal segments marked
in Arabic numerals and flagellomeres designated by Roman numerals, B. Flagellomere
II showing ascoids, C. Pharynx attached to cibarium, D. Metathoracic spiracle and post-
spiracular setae, E. Cibarium, E Spermathecae and associated structures, G. Wing, H.
Femur, tibia, and basitarsus of front leg.
Female: Rio Changena, Bocas del Toro Prov., Panama

Lutzomyia townsendi male I. Male genitalia; small subterminal seta on style is present,
J. Genital pump and filaments, same scale as Fig. 121.
Male: Rancho Grande, Aragua State, Venezuela


Ph-~n -- ---

C






SUBGENUS LUTZOMYIA FRANCA, 1924
This subgenus, divided into 3 series (Lewis et al., 1977), contains 21 species
including L. marinkellei n. sp. described in this review. Seven of these are an-
thropophilic, and others probably share the same habit. Lutzomyia longipalpis,
the principal vector of visceral leishmaniasis in the New World (Lewis, 1974),
ranges from southern Mexico to central Argentina, where it is locally common
in dry, nonforested areas. Lutzomyia gomezi, also reported in Colombia, has
been implicated as a vector of dermal leishmaniasis in Panama (Christensen et
al., 1969).
In Colombia the series longipalpis is represented by 3 species-L. longi-
palpis, L. lichyi, and L. bifoliata. Two species, L. gomezi and L. marinkellei,
are included in the series cruciata. Except for longipalpis, the females of these
species are remarkably similar and are separated only by minor nonsexual char-
acteristics.

Keys to Species
Males
1. Paramere with long dorsobasal arm bearing apical setae. Coxite with
2 very broad fan-like setae at inner base .......................... 2
Paramere without dorsobasal arm. Coxite with 4 or more simple
setae at inner base ......................................... 3
2. Paramere with dorsobasal arm bearing 2 slender, recurved setae. Coxite
with patch of long setae distad of fan-like setae ...... bifoliata (Fig. 13)
Paramere with dorsobasal arm bearing 2 fan-like setae. Coxite
without a distal patch of long setae ................... lichyi (Fig. 14)
3. Paramere with 2 strongly developed, dorsobasal setae. Coxite tuft of
4 setae. Style with subterminal seta ............. longipalpis (Fig. 15)
Paramere without such setae. Coxite tuft of 10+ setae. Style without
subterminal seta ......................................... 4
4. Pleura entirely pale. Style over half length of lateral lobe. Paramere
with dorsal setae on distal half of structure .......... gomezi (Fig. 16)
Pleura mostly dark. Style shorter than half length of lateral lobe.
Paramere with short dorsal setae confined to distal third of structure .
....... ............................... m arinkellei (Fig. 17)
Females
1. Cibarium with at least 8 horizontal teeth. Spermathecae as shown,
terminal annulation not markedly larger than others ...............
........................................ longipalpis (Fig. 15)
Cibarium with 4 horizontal teeth. Spermathecae otherwise, terminal
annulation spherical much larger than others ................... ... 2






2. Pleura pale, contrasting with dark mesonotum. Flagellomere I shorter
than or equal to length of labrum .................. gomezi (Fig. 16)
Pleura mostly dark. Flagellomere I longer than labrum .............. 3
3. Flagellomere I less than 0.32 mm long ........... marinkellei (Fig. 17)
Flagellomere I greater than 0.32 mm long .................. ..... 4
4. Terminal flagellomere longer than preceding flagellomere (XV) ......
................. ........................... lichyi (Fig. 14)
Terminal flagellomere shorter than preceding flagellomere (XV) .....
........................................ bifoliata (Fig. 13)


SERIES LONGIPALPIS
12. Lutzomyia (L.) bifoliata Osorno, Morales, Osorno & Hoyos
(Fig. 13)
Lutzomyia bifoliata Osorno, Morales, Osorno, & Hoyos, 1970: 8 (6, El Ter-
minal, Municip. Puerto Boyaca, Boyach, Colombia). Osorno et al., 1972a: 15
(listed). Forattini, 1973: 248 (gen. review, 6 fig.).
Distribution: Colombia (Antioquia, Boyach).
Material examined: Colombia. 13 66, 1999, Rio Anori (Antioquia), tree
buttresses, Sept. 1970, D.G.Y. 3 (, 1 light traps, same locality, May 1970,
C.H.P.
Discussion: Lutzomyia bifoliata males from Rio Anori generally agree with
the description and figures given by Osorno et al. (1970), but the strong erect
seta on the dorsum of the paramere is lacking in our material. The female, de-
scribed below, closely resembles L. lichyi, but the terminal flagellomere is
shorter, not longer than the preceding flagellomere. The longer labrum of bi-
foliata females (0.37-0.42 mm as opposed to0.25-0.36 mm, n = 17, in lichyi)
also may be useful in distinguishing the species if it proves to be a consistent,
nonoverlapping feature.
The following description is based on 10 females captured near the Rio Anori.
Female: Wing length 2.35-2.57; width 0.73. Head, mesonotum, and ab-
dominal tergites strongly pigmented: Rest of insect, including most of pleura,
paler. Head height 0.46; width 0.44. Eyes separated by 0.14 or distance = to
ca. 7.8 facet diameters. Flagellomere I (0.40-0.50 long), 1.2 x length of II +
III; ascoids longer than those of 6, but not reaching ends of flagellomeres, with
almost indiscernible posterior spurs, present on all but last flagellomere. Length
of palpal segments: 1 (0.06), 2 (0.15-0.18), 3 (0.19-0.21), 4 (0.09-0.12, 5
(0.30-0.37); palpal sensilla on segments 2 and 3. Labrum length = 0.37-
0.42. Cibarium with 4 nearly straight, sharp horizontal teeth and 20-30 verti-
cal teeth distributed as shown; chitinous arch complete, well defined; pigment
patch subtriangular, darker, and broader posteriorally. Pharynx (0.20 long) un-
armed. Pleura with 14-22 upper and 3-5 lower episternal setae. Length of






-,I


Id





B




A







F -oos


Y







D


H


J Y


L


FIGURE 13
Lutzomyia (L.) bifoliata A. Male head, B, Male flagellomere II, C. Female head, D.
Female flagellomere II, E. Female cibarium and pharynx, E Dorsal arm ofparamere, G,
Genital pump and filaments, H. Male genitalia, I. Spermathecae, J. Female wing, K.
Male wing, L. Female cibarium.
Male: Rio Anori, Antioquia Dept., Colombia
Female: Same locality as male






wing vein sections: Alpha (0.64-0.71), beta (0.27-0.34), delta (0.13-0.19),
gamma (0.27-0.34). Length of femora, tibiae, and basitarsi of slide 618: For-
eleg, 0.95, 1.22, 0.74; midleg, 0.91, 1.47, 0.81; hindleg, 0.98, 1.67, 1.27.
Abdominal sternite 2 horseshoe-shaped, hollow in middle. Tergite 8 without
setae. Spermathecae and sperm ducts as figured.

13. Lutzomyia (L.) lichyi (Floch & Abonnenc)
(Fig. 14)
Phlebotomus lichyi Floch & Abonnenc, 1950a: 1 (9 holotype, Rio Borbur-
ata, Carabobo, Venezuela). Morales et al., 1969a: 381 (Meta, Colombia).
Phlebotomus vexillarius Fairchild & Hertig, 1952: 514 (C, 9, Panama). Pi-
fano et al., 1960: 65 (Miranda State, Venezuela). Pifano et al., 1962: 385
(keyed). McConnell & Correa, 1964: 527 (infected with fungi). Floch & Kra-
mer, 1965: 1 (as synonym of lichyi). Hanson, 1968: 90 (larva & pupa). Scorza
et al., 1968: 35 bionomicss). Calderon, 1973: 87 (Merida State, Venezuela).
Phlebotomusfoliatus Mirsa & Ortiz, 1952: 249 (6, Venezuela). Fairchild
& Hertig, 1958b: 205 (as synonym of vexillarius).
Lutzomyia vexillaria: Martins et al., 1963: 335 (Roraima, Brazil).
Lutzomyia lichyi: Barretto, 1962: 98 (listed). Osorno et al., 1970: 12 (cf. to
L. bifoliata). Arjona et al., 1971: 93 (biting record, Tolima, Colombia). Osorno
et al., 1972a: 16 (Colombian records). Forattini, 1973: 213 (Trinidad), 258
(figs., redescr.). Lewis, 1975a: 500 et seq. (mouthpart morphol.).
Distribution: Panamd, Colombia (Antioquia, Caldas, Choc6, Huila, Mag-
dalena, Santander, Tolima, Valle), Venezuela, Trinidad, Brazil.
Material examined: Colombia. 1 9, Curiche (Choc6), light trap, 16 Aug.
1967, D.G.Y. 4 99, Alto Curiche (Choc6), biting man, July-Aug. 1967,
D.G.Y. 1 6, Alto Curiche, tree buttress, 4 April 1967, D.G.Y. 1 6, Alto Cur-
iche, Shannon trap, 13 Sept. 1967, D.G.Y. 1 9, Miraflores, Guatape (Antio-
quia), light trap, 12 March 1969, P. Barreto. 3 99, Suaza (Huila), horse bait,
9 March 1971, C. Aguila. 3 99, 25 km E of Buenaventura (Valle), tree trunks,
11 Aug. 1973, D.G.Y. & R.C.W. 27 d6, 5 99, 14 km SE of Santa Marta at
Minca (Magdalena), tree trunks, 17 Aug. 1973, D.G.Y. & R.C.W. 2 99, An-
chicaya Dam (Valle), light trap, 28 Jan. 1975, J. E. Browne. Venezuela. 1 S,
2 99, no other data, I. Ortiz. Trinidad. 1 9, Perserverance cave, Maraval, 22
Nov. 1974, E. S. Tikasingh. Panamd. 1 6, Serrania Maje, Cerro Chucanti
(Panama), tree buttress, 5 March 1950, R. Hartmann. 1 9, Cerro Campana
(Panama), Shannon trap, 25 Aug. 1950, M. Hertig.
Discussion: Man biting records for this species include those from Panama
(Fairchild and Hertig, 1952), Venezuela (Scorza et al., 1968), and Colombia
(Osorno et al., 1972a). During the interoceanic canal survey in Choc6 Dept. at
Alto Curiche, we collected only 4 females on human bait during a 5 month pe-
riod. Each was a daytime catch, although routine biting collections were carried
out at night as well. Dr. Pablo Barreto (pers. comm.) has collected biting fe-
males within the city limits of Cali, Colombia. In Panama, McConnell (1963)





















A
c E










0.
Fjr \















I








FIGURE 14
Lutzomyia (L.) lichyi A. Male head, B. Male flagellomere II, C. Female head, D.
Female flagellomere II, E. Female cibarium and pharynx, E Spermathecae, G. Female
cibarium, H. Female wing, I. Male wing, J. Male genitalia.
Male: Alto Curiche, Choc6 Dept., Colombia
Female: Curiche, Choc6 Dept., Colombia






found non-mammalian nucleatedd) red blood cells in the gut of a resting female,
and Thatcher and Hertig (1966) captured females feeding on Didelphis and Po-
tos.
According to Hanson (1968), the larvae of L. lichyi (as vexillarius) and L.
gomezi share certain characters not observed in other species. This, along with
adult morphology, supports the placement of L. gomezi in the subgenus Lut-
zomyia.

14. Lutzomyia (L.) longipalpis (Lutz & Neiva)
(Fig. 15)
Phlebotomus longipalpis Lutz & Neiva, 1912: 90 (6, 9, Brazil, definite
type locality not given). Barretto, 1947: 208 (full references). Guitton & Sher-
lock, 1969: 383 (immature stages).
Lutzomyia longipalpis: Osorno et al., 1969: 379 (redescr., 6, 9, Tolima
Dept., Colombia), Osorno et al., 1972a: 17 (Colombian records). Forattini,
1973: 213 (gen. review, figs.). Lewis, 1975a: 500 (mouthpart morphol.).
Killick-Kendrick et al., 1977: 429 (rearing).
Distribution: Southern Mexico to central Argentina. Colombia (Caldas,
Cundinamarca, Huila, Tolima).
Material examined: Colombia. 1 6, 2 99, Baraya (Huila), rock crevices,
March 1945, M. Hertig. Venezuela. 7 od, 13 99, near Calabozo (Guarico),
outside wall of house, 17 July 1965, D.G.Y. Bolivia, Brazil, Costa Rica, El
Salvador, Mexico, & Paraguay. Numerous specimens from various localities
within these countries.
Discussion: Lutzomyia longipalpis, first reported in Colombia by Osorno et
al. (1969), has received much attention because of its role as a vector of kala-
azar in the New World. It would be redundant to cite all important references
to this species, the type species of the genus, because Forattini (1973) has re-
cently reviewed most of them.
Phlebotomus otamae Nunez-Tovar and Phlebotomus almazani Galliard are
well established junior synonyms of L. longipalpis.

SERIES CRUCIATA
15. Lutzomyia (L,) gomezi (Nitzulescu)
(Fig. 16)
Phlebotomus gomezi Nitzulescu, 1931: 247 ( San Cristobal, Tachira State,
Venezuela). Barretto, 1947: 202 (full references). Fairchild & Hertig, 1948a:
252 (taxonomy). Fairchild & Hertig, 1953b: 382 (6, 9, redescr., refs., figs.).
Hanson, 1968: 60 (larva & pupa). Barreto, 1969: 464 (Meta & Valle, Colom-
bia). Morales et al., 1969a: 378 (Meta, Colombia).
Phlebotomus suis Rozeboom, 1940: 8 (6, 9, Panama). Barretto, 1946a:
1 (as synonym of gomezi). Fairchild & Hertig, 1948a: 252 (as synonym of
gomezi).





















FFt

G'Li'
B-- C__


E


J


L


M


FIGURE 15
Lutzomyia (L.) longipalpis A. Male head, B. Male flagellomere II, C. Female head,
D. Female flagellomere II, E. Female cibarium and pharynx, E Female wing, G. Male
wing, H. Genital pump and filaments, I. Male genitalia, J. Body of spermatheca, K.
Spermathecae, L. Genital pump, same scale as Fig. 151, M. Paramere of male from Huila
Dept., Colombia, same scale as Fig. 151, N. Female cibarium.
Male: Calabozo, Guarico State, Venezuela
Female: Same locality as male (except for Fig. 15M)


i u

\e ~ ,
Z___C--ylXL-Y.-I




















B

E














I \
A C































FIGURE 16
Lutzomyia (L.) gomezi A. Male head, B. Male flagellomere II, C. Female head, D.
Female flagellomere II, E. Female cibarium and pharynx, E Male genitalia, G. Sper-
mathecae, H. Female wing, I. Male wing, J. Female cibarium.
Male: Trinidad
Female: Trinidad






Phlebotomusjapignyi Floch & Abonnenc, 1944c: 2 (6, 9, French Guiana).
Fairchild & Hertig, 1948a: 252 (as synonym of gomezi).
Lutzomyia gomezi: Martins et al., 1962b: 90 (cf. to L. gasparviannai).
Osorno et al., 1972a: 19 (Colombian records). Forattini, 1973: 240 (gen. re-
view). Lewis, 1975a: 500 (mouthpart morphol.). Intermill & Muller, 1976:
543 (Honduras). Miles et al., 1976: 531-532 (mating behavior). Ward &
Ready, 1975: 128 (egg, figs.). Zimmerman et al., 1977: 575 (egg, figs.).
Distribution: El Salvador to southern Brazil. Colombia (Amazonas, Antio-
quia, Bolivar, Boyaca, Caldas, Caqueta, Cesar, Choc6, Guajira, Huila, Mag-
dalena, Meta, Norte de Santander, Santander, Tolima, Vichada).
Material examined: Colombia. 19 c6, 14 99, Curiche (Choc6), Malaise
trap, May-Nov. 1967, D.G.Y. 23 99, same data but biting man. 2 6 same
data but on tree trunk, 4 April 1967. 1 9 Sautata, Rio Atrato (Choc6), Malaise
trap, 20 Jan. 1968, D.G.Y. 8 d, 3 99, Rio Anori (Antioquia), light traps,
May 1970, C.H.P. 1 9, same data by 15 Sept. 1970, D.G.Y. 7 99, Puerto
Boyaca (Boyaca), light trap, 6 May 1973, C.J.M. 6 d>, 2 99, Tres Esquinas
(Caqueta), light trap, 10 Nov. 1971, C.J.M. 1 S, 17 km W of Leticia (Ama-
zonas), light trap, 26 July 1973, D.G.Y. & R.C.M. Brazil, Costa Rica, Ecua-
dor, Nicaragua, Panamd, Venezuela. Numerous specimens on slides and in
alcohol, in collection at UF
Discussion: Lutzomyia gomezi, a widespread species in Colombia and else-
where, is highly anthropophilic. Some of the many man-biting records include
those from Panami (Fairchild and Hertig, 1948a; Chaniotis et al., 1971b), Col-
ombia (Osorno et al., 1972a), Venezuela (Pifano and Ortiz, 1965), Trinidad
(Callan, 1947, as trinidadensis), and Brazil (Ward et al., 1973). In Panama,
Chaniotis et al. (1971b) observed that females were more commonly taken in
the canopy than at ground level in human biting collections. This species is one
of the suspected vectors of dermal leishmaniasis in that country (Schneider and
Hertig, 1966; Christensen and Herrer, 1973).
Forattini (1973) provides additional references and information on the biol-
ogy and disease relationships of L. gomezi.

16. Lutzomyia (L.) marinkellei n. sp.
(Fig. 17)
Male: Wing length 1.7; width 0.42. Head, mesonotum, abdominal tergites,
and genitalia strongly pigmented; rest of insect, including most of pleura, mod-
erately pigmented. Head height 0.34; width 0.33. Eyes large, separated by 0.09
or by distance = to 5.3 facet diameters. Flagellomere I (0.33 long), 1.1 x
length of II + III; ascoids with very short posterior spurs, their distal tips ending
before apical third of flagellomere II, on all flagellomeres except last 2. Length
of palpal segments: 1 (0.04), 2 (0.12), 3 (0.15), 4 (0.12), 5 (0.32); palpal sen-
silla, fewer than 10, on middle third of segment 3. Labrum length 0.195. Ci-
barium unarmed except for vestiges of vertical teeth; chitinous arch complete,
diffuse in middle; pigment patch subtriangular, more narrow than 9. Pharynx






.0


B


FIGURE 17
Lutzomyia (L.) marinkellei A. Spermathecae, B. Female wing, C. Male wing, D.
Female flagellomere II, E. Base of ascoid, E Female head, G. Female cibarium, H. Male
head, I. Male genitalia.
Male: Tres Esquinas, Caqueti Intendencia, Colombia
Female: Rio Aripuana, Mato Grosso State, Brazil






length 0.146. Pleura with 19 upper and 5 lower episternal setae. Length of wing
vein sections: Alpha (0.30), Beta (0.21), delta (0.07), gamma (0.30). Legs
missing but length of femora, tibiae, and basitarsi of paratype no. 631: Foreleg,
0.71,0.70,0.39; midleg, 0.71, 0.93, 0.46; hindleg, 0.74,0.98,0.56. Abdom-
inal sternite 2 horseshoe-shaped, the median opening not enclosed. Style short
(0.13 long) with 4 strong spines as shown, no subterminal bristle. Coxite stout
(0.25 long x 0.10 wide at greatest width), basal tuft of 12-13 long setae on
individual tubercles which form a raspberry-like base. Paramere as shown, up-
turned apically, dorsal setae few and rather short, these and lateral setae re-
stricted to apical third of structure, ventral setae on distal half. Aedeagus (ca.
0.11 long) slender apically. Lateral lobe (0.28 long), wider distally, perhaps
due to mounting. Genital pump (0.126 long); each filament ca. 0.43 long or
3.4 x length of pump, tip simple, not significantly enlarged. Cercus as shown.
Female: Wing length 1.85; width 0.57. Coloration same as 6. Head height
0.38, width 0.37. Eyes large, separated by 0.98 or distance = to 5.5 facet di-
ameters. Flagellomere I(0.28 long), 1.2 x length of II +III; ascoids with short
posterior spurs, longer than those of 3 but not reaching end of flagellomere II,
last 7 flagellomeres missing. Length of palpal segments: 1 (0.04), 2 (0.13), 3
(0.16), 4 (0.10), 5 (missing); palpal sensilla as in 6. Labrum length 0.23. Ci-
barium with 4 sharp horizontal teeth, 11-15 small vertical teeth in a single
row; chitinous arch complete, diffuse in middle; pigment patch as shown,
darker and wider posteriorally. Pharynx length 0.17. Pleura with 19 upper and
7 lower episternal setae. Length of wing vein sections: Alpha (0.44), beta
(0.25), delta (0.11), gamma (0.30). Length of femora, tibiae, and basitarsi of
slide 632: Foreleg, 0.78, 0.71, 0.39; midleg, 0.75, 0.90, 0.46; hindleg, 0.83,
1.1, 0.56. Abdominal sternite 2 apparently open in middle as in 3 but hardly
noticeable. Tergite 8 lacking setae. Tergite 9 without dorsolateral papillate
lobes. Spermathecae and sperm ducts as shown.
Material examined: Colombia. 3 holotype (no. 628), Tres Esquinas (Ca-
queti), light trap, 10 Nov. 1971, C. J. Marinkelle. 9 allotype (no. 629), same
data. 1 3, 1 9 paratypes (no. 630, 633) same data. Brazil. 1 6, 1 9 paratypes
(nos. 631-632), near Rio Aripuana (Mato Grosso), J. Arias.
Discussion: Lutzomyia marinkellei is closely allied with L. cruciata (Coq.),
L. evangelistai Martins and Fraiha, L. gomezi, and L. sherlocki Martins, Silva,
and Falcao, all of which form the series cruciata. From these, the male of mar-
inkellei is separated by the following combination of characters: both mesono-
tum and pleura infuscated; style short, its length less than or equal to half the
length of lateral lobe; paramere with few dorsal setae, these and lateral setae
confined to apical third of structure; coxite tuft of fewer than 20 setae.
The dark pleura differentiates the female from evangelistai, gomezi, and
sherlocki species with pale pleura which are (or may be) sympatric with mar-
inkellei. The status of L. cruciata and the form described as Phlebotomus dia-
bolicus Hall is still uncertain, but the latter taxon is probably a subspecies of
cruciata as treated by Lewis (1975a). The females of marinkellei and c. dia-






bolica, unlike c. cruciata, lack papillate lobes of tergite 9 and have well pig-
mented pleura. It would be difficult to separate these females in the absence of
males were it not for the fact that c. diabolica occurs well outside the range of
marinkellei in Mexico and Texas.
The length of the first flagellomere, the character used in the key to separate
marinkellei from females of lichyi and bifoliata, may prove to be variable as a
diagnostic character. With these taxa specific identification should be based on
associated males.
I am pleased to name this species after Dr. C. J. Marinkelle, who was most
helpful in providing material for this study.

MIGONEI GROUP THEODOR, 1965
This group is divided into 3 series following Lewis et al. (1977). The series
are migonei, walker, and costalimai (= subgenus Barrettomyia Martins and
da Silva, 1965). Species in the latter series occur in Brazil, but some of those
in the other series are more widespread with 4 species occurring in Colombia.
Barretto (1962) and Forattini (1971a, 1973) include some of the migonei group
taxa in the subgenus Coromyia Barretto, 1962.
The only species in this group known to be man-biting is L. migonei, here
reported in Colombia for the first time.


Keys to Species
Males
1. Genital filaments over 3 x length of pump, with simple tips. Coxite
with basal tuft of 4- 6 short setae .................. migonei (Fig. 18)
Genital filaments less than 3 x length of pump, their tips spherically
enlarged with an inner "tooth." Coxite with basal tuft of 10+ longer
setae and a median ventral group of 6+ setae ........ ............ 2
2. Aedeagus simple, without a dorsal projection. Genital filaments
0.30 mm or longer ............. ...... ... marajoensis (Fig. 19)
Aedeagus complex, with a dorsal cylindrical projection. Genital
filaments less than 0.30 mm ................... ... walker (Fig. 20)

Females
1. Spermathecae narrow and tubular, at least 3 x longer than wide ......
................... ........................... migonei (Fig. 18)
Spermathecae capsule-shaped, less than 2 x as long as wide ......... 2
2. Individual sperm ducts over 3 x length of common duct, wider and
with fine transverse striations basally ........... sp. de Baduel (Fig. 18)

Individual sperm ducts about /2 as wide as common duct ..........
.......... ........................... m arajoensis (Fig. 19)






Individual sperm ducts more slender, about /3 as wide as common
duct ....................................... walker (Fig. 20)


SERIES MIGONEI
17. Lutzomyia migonei (Franga)
(Fig. 18)
Phlebotomus migonei Franga, 1920: 230 (d holotype, Asunci6n, Paraguay).
Barretto, 1947: 211-213 (full refs., synonyms). Forattini, 1954: 214 et seq.
(sternites, figs.). Forattini, 1960: 479 (Amapa, Brazil). Pifano et al., 1962:
387, 388 (6, 9, keyed). Calderon, 1973: 87 (Merida State, Venezuela).
Lutzomyia migonei: Theodor, 1965: 182 (d, 9, figs.). Forattini, 1973: 122
et seq. (gen. review, refs., immatures, figs.). Lewis, 1975a: 500 et seq. (mouth-
part morphol.).
Distribution: Colombia (Magdalena), Venezuela, Trinidad, Brazil, Para-
guay, Argentina.
Material examined: Colombia. 2 6d, Rio Don Diego, E of Santa Marta
(Magdalena), tree trunks, 15 Aug. 1973, D.G.Y & R.C.W Brazil. 2 3S, La-
phina (Minas Gerais), flight trap near cave, 28 Aug. 1974, D.G.Y. & P.
Williams. Trinidad. 2 d6, Bush Bush Forest, Nariva Swamp, chicken-baited
trap, 17- 18 May 1961, T. H. G. Aitken. 1 9, same data but 22- 23 May.
Discussion: Ranging from Argentina to northern Colombia, L. migonei has
been found naturally infected with flagellates of uncertain identity in Venezuela
and Brazil (Johnson et al., 1963; Forattini, 1973).
Other information dealing with the biology of this anthropophilic species
was reviewed by Forattini (1973) and need not be repeated here.


SERIES WALKER
18. Lutzomyia marajoensis (Damasceno & Causey)
(Fig. 19)
Phlebotomus marajoensis Damasceno & Causey, 1944: 339 (d holotype,
Ilha do Maraj6, Brazil). Fairchild & Hertig, 1961b: 250-254 (T, 9, redescr.,
figs., refs., dist.). Pifano etal., 1962:386 et seq. (J, 9, redescr., figs., keyed).
Sherlock, 1962: 327- 328 (cf. to dubitans).
Lutzomyia marajoensis: Barretto, 1962: 98 (listed). Martins et al., 1962a:
381 (Goias, Brazil). Martins et al., 1963: 334 (Roraima, Brazil). Lewis, 1967b:
132 refss., cf. to walkeri. Barreto, 1969: 461,469 (mention). Forattini, 1971a:
101 (listed). Osorno et al., 1972a: 23 (Colombian records). Christensen, 1972a:
88 (listed). Forattini, 1973: 292 (as synonym of walkeri. Llanos, 1973: 31
(6, 9, redescr., figs., Peru). Christensen & Herrer, 1973: 579 (Panama rec-
ord). Lewis, 1975a: 500 et seq. (mouthpart morphol.).
Phlebotomus dubitans Sherlock, 1962: 324 (d holotype, San Vicente de
Chucuri, Santander, Colombia). Orson et al., 1967: 28 (mention).





































S I,



D E










FIGURE 18
Lutzomyia migonei A. Male genitalia, B. Female cibarium, C. Spermathecae.
Male: Trinidad
Female: Trinidad
Lutzomyia sp. de Baduel female D. Cibarium, E. Spermathecae.
Female: Cumariana, Vichada Dept., Colombia











B




F


D


ML
M (


NJ


FIGURE 19
Lutzomyia marajoensis A. Male head, B. Male flagellomere II, C. Female head, D.
Female flagellomere II, E. Female cibarium and pharynx, E Body of spermatheca, G.
Male genitalia, H. Genital pump, I. Genital pump and filaments, J. Tips of genital fila-
ments, K. Spermathecae, L. Female wing, M. Male wing, N. Female cibarium.
Male: San Jose, Los Santos Prov., Panama
Female: Puerto Mensabe, Los Santos Prov., Panama


' "i



p 2 .' '


b


01






Lutzomyia dubitans: Theodor, 1965: 182 (listed). Lewis, 1967b: 131 (men-
tion). Forattini, 1971a: 103 (listed). Osorno et al., 1972a: 23, 78 (mention).
Martins & Morales, 1972: 366 (listed). Forattini, 1973: 292 (as synonym of
walker and marajoensis).
Distribution: Panama, Colombia (Boyaca, Huila, Santander), Brazil, Ven-
ezuela, Trinidad.
Material examined: Colombia. 1 3, Soledad, San Vicente de Chucuri (San-
tander), chicken coop, 16 Aug. 1944, A. Gast, E. Osorno, & O. Mangabeira.
Panama, Venezuela, and Trinidad, those specimens listed by Fairchild & Her-
tig (1961b).
Discussion: A L. marajoensis male, the one listed by Fairchild and Hertig
(1961b) from San Vicente de Chucuri, Santander, Colombia, was given to Gor-
gas Memorial Laboratory by Dr. O. Mangabeira in the 1940's (Dr. G. B. Fair-
child, pers. comm.). With some reservations, Sherlock (1962) later described
L. dubitans based on other males taken at the same time and place as this male.
Forattini (1973) treated both dubitans and marajoensis as junior synonyms of
L. walker, but only the first two are conspecific, the latter species being easily
separated from marajoensis by the characteristics given in the key and by the
shape of the parameres (cf. Figs. 19G and 20C).
Although L. walker appears to be more widely distributed than marajoen-
sis, both species occur together in some localities in Colombia and Trinidad.


19. Lutzomyia sp. de Baduel (Floch & Abonnenc)
(Fig. 18)
Phlebotomus sp. de Baduel Floch & Abonnenc, 1945b: 1 ( Baduel, French
Guiana). Floch & Abonnenc, 1945c: 12 (9 keyed). Floch & Abonnenc, 1952:
21 et seq. ( keyed, redescr., figs.). Forattini, 1960: 480 (Amapi, Brazil).
Lutzomyia sp. de Baduel: Theodor, 1965: 196 (listed). Martins et al., 1965:
4 (Rond6nia & MaranhAo, Brazil). Osorno et al., 1972a: 66 (Vichada, Col-
ombia).
Distribution: Colombia (Vichada), Brazil, French Guiana.
Material examined: Colombia. 1 9 (no. 652 INPES), Cumariana (Vichada),
light trap, 15 May 1967, C.J.M.
Discussion: The male of this informally named taxon remains unknown,
but it is certainly possible that it may be conspecific with one of the described
males in the series, possibly L. sericea (Floch and Abonnenc) or L. williamsi
(Damasceno, Causey, and Arouck).
The single known female from Colombia, seen by me, compares favorably
with the description and figures given by Floch and Abonnenc (1945b, 1952).
I have no reservations about placing it in the series walker of the migonei group
on the basis of spermathecae, cibarial structure, and palpi.






20. Lutzomyia walker (Newstead)
(Fig. 20)
Phlebotomus walker Newstead, 1914:188 ( 6, 9, Rio Abund, Bolivia-Brazil
boundary). Barretto, 1947: 230 (full refs.). Fairchild & Hertig, 1961b: 250, 254
(cf. to marajoensis).
Lutzomyia walker: Barretto, 1962: 98 (listed). Martins et al., 1965: 4 (Ron-
donia, Brazil). Lewis, 1967b: 132 et seq. (6, 9, redescr., figs.). Forattini,
1971a: 101 (listed). Osorno et al., 1972a:23 (Colombian records). Llanos, 1973:
31 (6, 9, redescr., fig.). Forattini, 1973: 274 et seq. (in part, gen. review, figs.).
Lewis, 1975a: 500 et seq. (mouthpart morphol.). Llanos et al., 1975b: 671
(Peri).
Phlebotomus gasti Sherlock, 1962: 326 (6, San Vicente de Chucuri, San-
tander, Colombia). Osorno et al., 1967: 28 (mention). NEW SYNONYM.
Lutzomyia gasti: Theodor, 1965: 182 (listed). Lewis, 1967b: 131 (listed).
Forattini, 1971: 103 (listed). Christensen, 1972a: 88-89 (Panama). Osorno et
al., 1972a: 23 (Colombian records). Forattini, 1973: 348 et seq. (gen. review,
fig.).
Distribution: Panamd, Colombia (Antioquia, Caqueti, Santander), Ecuador,
Peru, Brazil, Trinidad.
Material examined: Colombia. 1 S (no. 3924 INPES), La Nevera, Solano
(Caqueti), tree hole, 25 June 1969, A. Gast & J. Ochoa. 3 99, Tres Esquinas
(Caqueta), light trap, 10 Nov. 1971, C.J.M. 1 6, labelled P gasti Mang. MS,
no. 1120, Colombia (probably from the type locality of gasti). Brazil. 1 6 from
type series, Abuna River, Bolivia-Brazil boundary, Dr. F D. Walker, the Elders
Brixham, S. Devon. 2 66, Labrea (Amazonas), tree trunk, 9 Oct. 1972. D.G.Y.
1 6, 1 9, same data but light trap, 10 Oct. 1972. Ecuador. 1 9, Rio Napo at
Limoncocha (Napo), flight trap, 22 May 1976, D.G.Y. & T. Rogers. Trinidad.
150+ specimens of both sexes from various localities to be discussed in a forth-
coming paper.
Discussion: Both Lewis (1967b) and Llanos (1973) carefully redescribed the
male and female of L. walker, the former author studying material from the
type series, the latter basing her description on Peruvian specimens. I re-
mounted a male from the type series, the one mentioned by Lewis (op. cit.) as
being in the collection at Gorgas Memorial Laboratory but which is now at the
University of Florida. I also examined a male from Santander Dept., Colombia,
which Mangabeira collected and which bears the name P gasti MS. These two
males are identical in all aspects, including the presence of the dorsal projection
of the aedeagus as figured by Sherlock (1962) for L. gasti and by Llanos (op.
cit.) for L. walker. I am convinced therefore that L. gasti is a junior synonym
of L. walker. I should point out that the dorsal projection of the aedeagus is
lightly sclerotized, less so than Sherlock (1962) depicts, and this probably is the
reason why Lewis (op. cit.) did not observe this character state in the specimens
at the British Museum (Nat. Hist.).
I consider the Ecuadorian female to be conspecific with walker on the basis
of morphology and distribution.














/ / ,



A B









D
,- -.^ ^













FIGURE 20
Lutzomyia walker A. Spermathecae, B. Female cibarium, C. Male genitalia, D. Tip
of genital filament, greatly enlarged.
Male: Trinidad
Female: Trinidad






SAULENSIS GROUP LEWIS et al., 1977
This group was created to accommodate two closely related species-L. sau-
lensis and L. wilsoni (Damasceno and Causey). Forattini (1971a, 1973) places
the former species in the subgenus Dampfomyia Addis, but the spermathecae
are structurally different (Theodor, 1965); the male parameres lack a dorsal arm,
and the style has no subterminal bristle.
The females of saulensis and wilsoni are nearly identical (Vianna Martins,
pers. comm.), their populations probably overlapping in northern Brazil.

21. Lutzomyia saulenis (Floch & Abonnenc)
(Fig. 21)
Phlebotomus saulensis Floch & Abonnenc, 1944a: 11 (6 holotype, Saiil,
Haute Mana, French Guiana). Barretto, 1947: 222 (listed), Damasceno et al.,
1949: 831 (Brazilian records). Floch & Abonnenc, 1952: 35, 112-114 (d
keyed, redescr., figs.). Fairchild & Hertig, 1958b: 204-205 refss. taxonomic
discussion), Fairchild & Hertig, 1959: 122 (dist.). Sherlock, 1962: 321, 330
(Santander, Colombia), Pifano et al., 1962: 386, 389 ( 9, keyed, Venezuela).
Osorno et al., 1967: 28 (mention).
Phlebotomus pinealis Floch & Abonnenc, 1944b: 11 ( holotype, Crique
Anguille, French Guiana). Fairchild & Hertig, 1958b: 204- 205 refss. as syn-
onym of saulensis). Johnson & Hertig, 1961: 765, 774 (rearing attempt). Han-
son, 1968: 69 (1st instar larva, descr., fig.).
Lutzomyia saulensis: Martins et al., 1963: 335 (Roraima, Brazil). Martins et
al., 1965: 3 (Rond6nia, Brazil). Theodor, 1965: 196 (6, 9, figs.). Barreto,
1969: 466-467 (Valle, Colombia). Lewis et al., 1970: 215 (parous study).
Fraiha et al., 1970b: 215 (mention). Christensen & Fairchild, 1971: 301 (Pan-
ami), Chaniotis et al., 1971a: 344 et seq. (pop. dynamics, Panami). Osorno et
al., 1972a: 66 (Colombian records). Christensen, 1972a: 88 (listed). Chaniotis
etal., 1972:95 et seq. (resting sites). Shaw& Lainson, 1972:710et seq. (infected
with nonleishmanial flagellates, collecting data, Brazil). Llanos, 1973: 34 (C,
Y, redescr., figs., Peri). Forattini, 1973: 327 et seq. (gen. review, figs.). Chris-
tensen & Herrer, 1973: 579 (listed). Ward et al., 1973: 178 (attracted to rodents
and man, Brazil). Lainson et al., 1973: 190 (negative for Leishmania). Lewis,
1975a: 504 et seq. (mouthpart morphol.). Llanos et al., 1975b: 671 (Peri).
Llanos et al., 1976: 480 (Peri). Martins et al., 1976b: 496 (Peru).
Distribution: Costa Rica, Panama, Colombia (Antioquia, Caqueti, Choc6,
Santander, Valle), Peru, Brazil, French Guiana.
Material examined: Colombia. 2 99, Rio Anori (Antioquia), light traps, 3
May 1970, C,H.P. 8 66, 11 2Y, same data but some in tree hollows, Sept.
1970. D.G.Y. 14 6, 55 9? Curiche (Choc6), light, Shannon & Malaise traps,
tree trunks, May-Dec. 1967, D.G.Y. 3 YY, Alto Curiche (Choc6), Malaise
& light traps, Aug., Sept., Nov. 1967, D.G.Y. 1 Teresita (Choc6), Malaise
trap, 22 April 1967, D.G.Y. 4 9 Rio Atrato at Sautata (Choc6), Malaise traps,










6T ;



B


E


I.0


6- -


FIGURE 21
Lutzomyia saulensis A. Male head, B. Male flagellomere II, C. Female head, D.
Female flagellomere II, E. Female cibarium and pharynx, E Male genitalia, G. Genital
pump and filaments, H. Spermathecae, I. Female wing, J. Male wing, K. Female cibar-
ium.
Male: Curiche, Choc6 Dept., Colombia
Female: Same locality as male


P






Nov.- Dec. 1967, Jan. 1968,D.G.Y. 2 ~d ,Anchicayi Dam (Valle), tree trunks,
9 Aug. 1973, D.G.Y. & R.C.W. 1 3, same data but light trap, 10 Aug. 1973.
2 99, 25 km E of Buenaventura (Valle), flight trap, 12 Aug. 1973, D.G.Y. &
R.C.W Brazil. 1 o, Belem (Para), 1 Feb. 1944, R. Damasceno & O. Causey.
Discussion: Although widely distributed in lowland forests, L. saulensis is
a little known species. Ward et al. (1973) collected specimens on rodent and
human bait in Pari, Brazil, but this species can not be regarded as being anthro-
pophilic. Both sexes are readily identifiable.

VERRUCARUM GROUP THEODOR, 1965
This group of 22 species is divided into 2 series based on male characteristics
(Theodor, 1965)-the series verrucarum (14 spp.) and the series serrana (7
spp.). At present, L. bursiformis (Floch & Abonnenc) from French Guiana can-
not be placed because the male remains undiscovered. Some authors include
these species in the subgenus Lutzomyia Franga (Forattini, 1971a, 1973) or in
the subgenus Coromyia Barretto, 1962.
Two species in the series serrana and 11 in the series verrucarum occur
within or very near the borders of Colombia. The majority of species in the
latter series are Andean, although L. ovallesi and L. evansi are widespread in
lowland areas. The Andean taxa are morphologically similar and for the most
part have limited, allopatric distributions.
Unfortunately, the females of most species are difficult, if not impossible, to
identify without associated males. For this reason I am omitting keys to the
females but will point out, when possible, the distinguishing characteristics for
some of the species. It may be appropriate to add that the presence of L. ver-
rucarum (Townsend) in Venezuela (Anduze et al., 1947; Floch and Abonnenc,
1950a; Forattini, 1973) can be confirmed only when the male is discovered in
that country.

Key to Species
Males
1. Style of male genitalia with 2 or 3 strong spines (series serrana) ..... 2
Style with 4 strong spines (series verrucarum) ................... 3
2. Coxite tuft of 5- 6 setae, upturned but not strongly recurved at tips,
implanted in a straight row ................... ... serrana (Fig. 22)
Coxite tuft of 6- 7 setae, markedly recurved apically, not implanted in a
straight row ............................ .. ... odax (Fig. 28)
3. Basal spine of style isolated ................................ 4
Basal spines of style paired, inserted more or less on the same level ... 7
4. Terminal spine of style very thick and crooked. Paramere as in
Fig. 28E. ................................. spinicrassa (Fig. 28)






Terminal spine of style relatively slender, not crooked. Paramere
otherwise ................. ........................... 5

5. Middle third of paramere with dorsal and ventral setae ............
..... ............................. sauroida (Fig. 28)
Middle third of paramere without dorsal or ventral setae ............ 6
6. Paramere club-like, angular at tip and with dorsal subspical row of
13- 16 short spine-like setae ................ longiflocosa (Fig. 28)
Paramere more or less rounded at end, less club-like, dorsal subapical
setae longer, not spine-like ................ quasitownsendi (Fig. 28)
7. Coxite with subapical group of long hairs. Basal spines of style inserted
on well marked rather long tubercles ........................... 8
Coxite without subapical group of long hairs. Basal spines of style not
implanted on tubercles ........................ ............ 9
8. Setae of basal coxite tuft thicker. Paramere as shown, with distal
lobe on the ventral margin and with dorsal setae confined to
extreme tip. Style with simple basal spines, no subterminal
bristle ................................ moralesi n. sp. (Fig. 26)
Setae of basal coxite tuft slender. Paramere more clubbed at end,
without ventral lobe but with numerous dorsal setae in a subapical
patch. Style with shorter basal spine modified as shown,
subterminal bristle present ....................... andina (Fig. 23)
9. Dorsal setae of paramere restricted to distal third of structure ....... 10
Dorsal setae of paramere covering at least distal half of structure .... 11
10. Paramere strongly clubbed. Coxite tuft of 10+ setae. Flagellomere I at
least 20% longer than labrum ................. columbiana (Fig. 24)
Paramere not clubbed. Coxite tuft of fewer than 7 setae. Flagellomere I
short, subequal in length to labrum ............... ovallesi (Fig. 27)
11. Basal coxite tuft of 18+ setae. Distal third of paramere wider than middle
third............................ .......... disiuncta (Fig. 23)
Basal coxite tuft of 10 or fewer setae. Distal third of paramere not
enlarged ............. ...................... ........ 12
12. Genital filaments over 4 x length of pump. Style with a subterminal
seta ...................................... evansi (Fig. 25)
Genital filaments less than 4 x length of pump. Style without a
subterminal seta .......................... nuneztovari (Fig. 23)






SERIES SERRANA
22. Lutzomyia odax (Fairchild & Hertig)
(Fig. 28)
Phlebotomus odax Fairchild & Hertig, 1961b: 230 (J holotype, 2, Almi-
rante, Bocas del Toro, Panami).
Lutzomyia odax: Barretto, 1962, 97 (listed). Martins et al., 1965: 7 (cf. to
dubia). Christensen & Fairchild, 1971: 302 (Darien, Panama). Christensen,
1972a: 88 (listed). Christensen et al., 1972: 57 (PanamA). Forattini, 1973: 213
et seq. (gen. review, figs.).
Distribution: Guatemala, Honduras, Nicaragua, Panamd.
Material examined: Panamd. S holotype (no. 4168), Almirante (Bocas del
Toro), light trap, 15 Oct. 1952, A. Quinones. Y allotype (no. 3327), same data,
but human bait collection, 20 June 1951, A. Vivanco. 1 d, same data but 17
June 1950, R. Hartmann.
Discussion: There is little doubt that this species occurs in Colombia, Chris-
tensen and Fairchild (1971) having reported its presence within 4 km of the
Colombian border in Darien Province, Panama. Females closely resemble
those of L. serrana but differ in the lengths of the individual sperm ducts (Fair-
child & Hertig, 1961b). Both species occasionally bite man but are more com-
mon in light trap and resting collections.
Forattini (1973) may be correct in treating L. dubia Martins, Falcao, & da
Silva as a junior synonym of L. odax, but more evidence is needed to support
this view. Martins et al. (1965) point out differences between the taxa, expe-
cially those of the male genitalia.

23. Lutzomyia serrana (Damasceno & Arouck)
(Fig. 22)
Phlebotomus serranus Damasceno & Arouck, 1949: 843 (c holotype, Serra
da Piriabas, Para, Brazil). Forattini, 1960: 479 (Amapi, Brazil). Blancas,
1959- 1960: 125 (Per6). Fairchild & Hertig, 1961b: 237 refss., descr., figs.).
Hanson, 1961; 320 (breeding sites). Johnson & Hertig, 1961: 765 (rearing
data). Sherlock, 1962; 330 (Santander, Colombia). Rosabal, 1966: 1 (Costa
Rica). Hanson, 1968: 76 (larva, pupa, descr.).
Phlebotomus guayasi Rodriguez, 1956: 76 (S, 9, Guayas, Ecuador). Fair-
child & Hertig, 1961b: 237 (as synonym of serranus).
Lutzomyia serrana: Barretto, 1962: 97 (listed). Martins et al., 1965: 3 (Ron-
d6nia, Brazil), 6 (cf. to dubia). Lewis, 1967a: 74 (listed). Forattini, 1971a: 100
(listed). Tesh et al., 1971a: 152 (blood meals). Chaniotis et al., 1971a: 345
(pop. dynamics, PanamA). Osorno et al., 1972a: 30 (Colombian records). Tesh
et al., 1972: 90 (blood meals). Chaniotis et al., 1972: 95 (listed). Christensen,
1972a: 88 (listed). Forattini, 1973: 123 et seq. (gen. review, figs.). Llanos,
1973: 32 (Peri, figs.). Velasco, 1973: 77 (Bolivia). Llanos et al., 1975b: 668
(unusual spermathecae, fig.), 671 (Peri). Herrer & Christensen, 1976a: 62














B


D
D


H Qq5'










(iit 'i


FIGURE 22
Lutzomyia serrana A. Male head, B. Male flagellomere II, C. Female head, D. Female
flagellomere II, E. Female cibarium and pharynx, E Male genitalia, G. Paramere, H.
Spermathecae, I. Genital pump and filaments, J. Female wing, K. Male wing, L. Female
cibarium.
Male: Alto Curiche, Choc6 Dept., Colombia
Female: Rio Anori, Antioquia Dept., Colombia


E
*


G i\\


1.0


Kd-


01


~
u
~






(Panami). Llanos et al., 1976: 480 (Peri). Martins et al., 1976a: 488 (Peri).
Martins et al., 1976b: 496 (Peri).
Distribution: Mexico, Honduras, Nicaragua, Costa Rica, Panamd, Colom-
bia (Antioquia, Bolivar, Boyaca, Caldas, Choc6, Cundinamarca, Santander),
Ecuador, Peru, Bolivia, Brazil.
Material examined: Colombia. 1 6, Alto Curiche (Choc6), tree buttress, 21
May 1967, D.G.Y. 1 6, Rio Anori (Antioquia), tree buttress, 6 Feb. 1970,
C.H.P. 1 9, same data but at light, 21 Sept. 1970, D.G.Y. Panamd. 1 6,
1 9, Rio Platanar (Panami), hollow tree, 20 July 1950, R. Hartmann.
Discussion: In Panama, Tesh et al. (1971a, 1972) identified the blood meals
of 14 females using the precipitin test. Ten had fed upon rodents, 2 on edentates,
and 2 on marsupials. Dr. C. H. Porter (in litt.) collected less than 10 specimens
on human bait over a one year period at Rio Anori (Antioquia), Colombia.


SERIES VERRUCARUM
24. Lutzomyia andina Osorno, Osorno & Morales
(Fig. 23)
Lutzomyia andina Osorno, Osorno, & Morales, 1972b: 2 (6 holotype, 9,
El Chariquito, Cundinamarca, Colombia). Osorno et al., 1972a: 24 (Colom-
bian records).
Phlebotomus verrucarum: Osorno et al., 1967: 29 (not verrucarum Town-
send).
Distribution: Colombia (Cundinamarca).
Material examined: 1 6, paratype, type locality, 2550 m above sea level,
Aug. 1966. 1 9, paratype, same data but Sept. 1966.
Discussion: Specimens of this recently described species, known only from
the type locality, were taken on human bait and in tree holes. Females bite man
during the day and night.
The shorter of the paired basal spines of the male style is curiously modified,
the apical part widened and with a flange (Fig. 23A). This characteristic, if
shared by other andina males, will serve to separate it from other males in the
verrucarum group.

25. Lutzomyia columbiana (Ristorcelli & Van Ty)
(Fig. 24)
Phlebotomus columbianus Ristorcelli & Van Ty, 1941: 263 (9 holotype,
Valle de Capuli, Narifio, Colombia). Rozeboom, 1947b: 705 (6, figs.). Sher-
lock, 1962: 322 (6, 9, figs.). Osorno et al., 1967: 28 (mention).
Phlebotomus monticolus var. incarum Ristorcelli & Van Ty, 1941: 266 (9,
Valle de Capuli, Narifio, Colombia). Rozeboom, 1947b: 705 (as synonym of
columbiana).
Lutzomyia columbiana: Barretto, 1962: 97 (listed). Osorno et al., 1972a:
24-26 (Colombian records). Forattini, 1973: 206 et seq. (gen. review, figs.).

























0-5s B







C


\ 3,- / /




o* E


FIGURE 23
Lutzomyia andina male A. Male genitalia.
Male: El Charquito, CundinamarcaDept., Colombia
Lutzomyia disiuncta male B. Male genitalia.
Male: Bogoti, Municip. de Suba, Cundinamarca Dept., Colombia
Lutzomyia nuneztovari C. Female cibarium, D. Male genitalia, E. Spermatheca; the
common duct and part of individual duct not visible.
Male: Timana, Huila Dept., Colombia
Female: Same locality as male












E


H


~Q05


1 0
K
6:1

L2-- --
L -ZTra


FIGURE 24
Lutzomyia columbiana A. Genital pump and filaments, B. Male head, C. Male fla-
gellomere II, D. Female head, E. Female flagellomere II, F Female cibarium and phar-
ynx, G. Male genitalia, H. Female cibarium, I. Spermathecae, J. Body of spermatheca,
K. Female wing, L. Male wing, M. Spermathecae.
Male: San Pedro, Narifio Dept., Colombia
Female: Same locality as male


Ii I




I')))


1.1






Distribution: Colombia (Cauca, Guajira, Nariiio, Valle).
Material examined: Colombia. 5 d6, 3 $9, Bolivar (Cauca), Jan. 1944,
J. Burbano. 1 6, 3 99, San Pedro (Narifio), Jan. 1944, J. Burbano. 2 &d,
ca. 30 km NW of Cali, near El Carmen (Valle), tree trunks, 12 Aug. 1973,
D.G.Y. & R.C.W
Discussion: This precinctive Colombian species was the suspected vector of
bartonellosis during the outbreak in southeastern Colombia over 40 years ago
(Rozeboom, 1947b).
The record of columbiana from Montes de Oca and Serra Mucuiru in Guajira
(Osorno et al., 1972a) is very interesting, these localities being widely sepa-
rated from southeastern Colombia where columbiana was first discovered. The
Andes, especially the eastern chain, may have acted as a corridor for the dis-
persal of this species.
I have seen other specimens of columbiana in the collections of the Califor-
nia Academy of Sciences and University of Florida. All were captured in south-
eastern Colombia and were identified by 0. Mangabeira and P. Galindo.


26. Lutzomyia disiuncta Morales, Osorno & Osorno
(Fig. 23)
Lutzomyia disiuncta Morales, Osorno, & Osorno, 1974: 446 (J holotype,
9, Hacienda La Conejera, Municip. de Suba, Bogota, Cundinamarca, Col-
ombia).
Distribution: Colombia (Cesar, Cundinamarca).
Material examined: Colombia. 8 6d, 21 99, Municip. Agustin Cadazzi
(Cesar), Sept. 1969 (INPES). 3 c6 (including holotype), 9 99, type locality,
Aug.- Sept. 1969 (INPES).
Discussion: This is another Andean species which is anthropophilic, speci-
mens having been taken on human bait during the day and night (Morales et
al., 1974). Other specimens were captured in tree holes in Cesar Department.

27. Lutzomyia evansi (Nunez- Tovar)
(Fig. 25)
Phlebotomus evansi: Nunez-Tovar, 1924: 44 (d, Mariara, Carabobo,
Venezuela). Barretto, 1947: 198 (full refs.). Mirsa, 1953: (rearing data). Fair-
child & Hertig, 1959: 122 (dist.). Pifano et al., 1962: 412 refss., redescr.,
figs.). Rosabal & Trejos, 1964: 167 (El Salvador). Rosabal & Trejos, 1965: 222
(El Salvador). Osorno et al., 1967: 28 (Nariio, Colombia).
Lutzomyia evansi: Theodor, 1965: 183 (listed). Morales et al., 1969b: 385
(mention). Osorno et al., 1972a: 27 (Colombian records). Llanos, 1973: 34
(6, 9 redescr., figs., Peri). Forattini, 1973: 122 et seq. (gen. review, figs.).
Llanos et al., 1975b: 670 (Peri).
Distribution: Guatemala, El Salvador, Honduras, Nicaragua, Costa Rica,
Colombia (Bolivar, Guajira, Magdalena, Narifio, Santander), Peru, Venezuela.










D


E
J


H
\\


G






I


1.0





Lc


FIGURE 25
Lutzomyia evansi A. Male head, B. Male flagellomere II, C. Female head, D. Female
flagellomere II, E. Female cibarium and pharynx, F Genital pump and filaments, G.
Genital pump, same scale as 25H, H. Male genitalia, I. Paramere, outer aspect, J. Sper-
mathecae, K. Body of spermatheca, L. Female wing, M. Male wing, N. Female cibar-
ium.
Male: Guapilonar, Carrazo, Nicaragua
Female: Same locality as male


B:


f~--L--






Material examined: Colombia. 1 9, Arjona (Bolivar), 27 May 1940. 1 9,
10 km SE of Santa Marta (Magdalena), tree trunk, 17 Aug. 1973, D.G.Y. &
R.C.W Honduras: 30+ dS, 100+ 99, Chumbagua, Santa Barbara, light
traps, July 1966, J. Matta. Other specimens from Central America and Vene-
zuela in collection at UE
Discussion: The female of evansi has very long sperm ducts and relatively
small spermathecae (Fig. 25J), features which separate it from other verruca-
rum group females in Colombia. This species has been reported biting man in
several countries including Colombia (Osorno et al., 1972a). It and L. longi-
palpis often, but not always, occur together in Central America, Colombia, and
Venezuela. The apparent absence of this species from Panama is most peculiar.

28. Lutzomyia longiflocosa Osorno, Morales, Osorno & Hoyos
(Fig. 28)
Lutzomyia longiflocosa Osorno, Morales, Osorno, & Hoyos, 1970: 1 (3 hol-
otype, Municip. de Tello, Huila, Colombia. Osorno et al., 1972a: 28 (Colom-
bian records). Osorno et al., 1972b: 18 (cf. to andina and quasitownsendi).
Distribution: Colombia (Huila).
Material examined: Colombia. 1 d (paratype no. 1641), type locality, 9 Feb.
1968, E. Osorno et al.
Discussion: Other than resting in tree holes at the type locality, nothing is
known about the habits of this species. The female is unknown.


29. Lutzomyia moralesi n. sp.
(Fig. 26)
Male: Wing length 1.96; width 0.58. Head, mesonotum, and genitalia
strongly pigmented; rest of insect pale. Head height 0.35; width 0.32. Eyes sep-
arated by 0.11 or by distance = to 5.9 facet diameters. Flagellomere I (0.33
long), 1.2 x length of II + III; ascoids simple, those on flagellomere II ending
near middle of structure, on all flagellomeres except last. Length of palpal seg-
ments:: 1 (0.04), 2 (0.15), 3 (0.16), 4 (0.11), 5 (0.32); palpal sensilla (ca. 10)
grouped loosely at middle of segment 3. Labrum length 0.21. Cibarium with
30+ vestigial, dot-like teeth; chitinous arch complete, diffuse in middle; pig-
ment patch invisible. Pharynx 0.15 long, unarmed. Pleura with 8-9 upper and
4 lower episternal setae. Length of wing vein sections: Alpha (0,53), beta
(0.18), delta (0.17), gamma (0,32). Length of femora, tibiae, and basitarsi:
Foreleg, 0.78, 0.90, 0.54; midleg, 0.74, 1.07, 0.61; hindleg 0.83, 1.22, 0.69.
Abdominal sternites hardly visible some with lateral, circular openings. Geni-
talia: Style 0.13 long, shaped as shown, the proximal 2 spines inserted at the
same level, the longer on a long process, no subterminal seta. Coxite (0.32 long
x 0.12 wide) with a distal patch of 30+ straight or nearly straight setae, a dense
tuft of ca. 30 thick sinuous setae at base. Paramere as shown but variable ac-
cording to angle of view; the apical part with an incipient lateral lobe and a





































H


FIGURE 26
Lutzomyia moralesi A. Female head, B. Female wing, C. Male wing, D. Male head,
E. Male flagellomere II, E Female flagellomere II, G. Female cibarium, H. Genital pump,
same scale as Fig. 26J, I. Tip of genital filament, same scale as Fig. 26G, J. Male genitalia,
K. Spermathecae.
Male: Near Rio Pance, Valle Dept., Colombia
Female: Near Pichind6, Valle Dept., Colombia


B





C






ventral heel. Aedeagus subtriangular, ummodified. Lateral lobe relatively long
(0.42). Genital pump (0.155 long), each filament 0.63 long or 4 x length of
pump, tip enlarged. Cercus as shown.
Female: Wing length 2.45; width 0.76. Coloration as in d. Head height
0.43; width 0.39. Eyes separated by 0.13 or distance = to ca. 6.6 facet diame-
ters. Flagellomere I (0.31 long) nearly 1.3 x length of II + III; ascoids longer
than those of o but not reaching end of flagellomere II; on all flagellomeres ex-
cept last. Length of palpal segments: 1 (0.014), 2 (0.20), 3 (0.19), 4 (0.12), 5
(0.39); palp 2 with ca. 5 apical sensilla, palp 3 with 25+ sensilla on apical half.
Labrum 0.36 long. Cibarium with 4 sharp horizontal teeth evenly spaced, 2
transverse rows of subequal vertical teeth, ca. 13 teeth in each row; chitinous
arch complete; pigment patch subtriangular, well defined. Pharynx 0.20 long,
unarmed but with distinct ridges. Pleura with 9-10 upper and 2 lower epister-
nal setae. Length of wing vein sections: Alpha (0.67), beta (0.20), delta
(0.26), gamma (0.41). Length of femora, tibiae, and basitarsi: Foreleg, 0.88,
0.99, 0.59; midleg, 0.83, 1.18, 0.66; hindleg, 0.87, 1.34, 0.78. Abdominal
sternite 3 with paired circular openings laterally. Tergite 8 with 1 or 2 setae on
each side. Spermathecae sac-like, as figured; common duct ca. 3 x length of
individual duct.
Distribution: Colombia (Valle).
Material examined: Colombia. d holotype (no. 512), ca. 15 km SW of Cali
(1600 m above sea level) near Rio Pance (Valle), on tree trunk, 8 Aug. 1973,
D.G.Y. & S. Ayala. Y allotype (no. 513), ca. 10 km W of Cali (1800 m above
sea level), near Pichinde (Valle), rock crevice, 2 Aug. 1973, D.G.Y. & R.C.W
Paratypes (nos. 514-526), 1 5, 3 9 same data as allotype except taken on
31 July 1973. 3 9 Y, same data as allotype. 1 S, 5 99 same data as holotype.
Discussion: The sexes of L. moralesi were associated on the basis of collect-
ing data, coloration, and metrical characters. No other verrucarum group spe-
cies were taken at either of the two localities listed above.
The male of this species resembles that of L. andina in details of the male
genitalia, but the setae of the basal coxite tuft are thicker, the paramere and its
station are different, the style lacks a small subterminal seta, and the shorter of
the two proximal spines is unmodified. The male of L. verrucarum (Townsend),
an allopatric Peruvian species, is similar to these species, but the proximal spines
of the style are not inserted on long or well marked tubercles and the parameres
are different.
Without males, I am unable to separate the females of L. moralesi from those
of most other verrucarum group species. Identifications based on this sex alone
require confirmation.
I am pleased to name this species after Dr. Alberto Morales-Alarcon, not only
for his valuable assistance in this study, but also to acknowledge his significant
contribution to our knowledge of Colombian phlebotomines.






30. Lutzomyia nuneztovari (Ortiz)
(Fig. 23)
Phlebotomus nunez-tovari Ortiz, 1954: 232 (6 holotype, Duaca, Lara, Vene-
zuela). Scorza & Ortiz, 1960: 434 et seq. (ecology). Pifano et al., 1962: 387
(6 keyed). Ortiz & Scorza, 1963: 344 (Venezuela). Scorza et al., 1967: 194
(d keyed), 195 (9 keyed), 196 (collecting data).
Lutzomyia nuneztovari: Theodor, 1965: 183 (mention), Martins & da Silva,
1965: 275 (mention). Martins et al., 1965: 9 (mention). Morales et al., 1969b:
385 (mention). Forattini, 1971a: 103 (listed). Martins & Fraiha, 1971: 364 (cf.
to evangelistai)., Young, 1972a: 312 (mention). Osorno et al., 1972b: 18 (men-
tion). Forattini, 1973: 265 (as synonym of ovallesi).
Distribution: Venezuela, Colombia (Huila).
Material examined: Colombia. 1 3, Timana (Huila), Shannon trap, 21 July
1971, C. Aguila. 1 same data but 1-2 Aug. 1971. 1 9, same data but light
trap, 2- 3 Aug. 1971. 1 2 same data but Shannon trap, 12- 13 Aug. 1971.
Discussion: Unlike Forattini (1973), I consider L. nuneztovari and L. ovallesi
to be distinct species, the male of the latter having a subterminal seta on the style
(lacking in nuneztovari) and with different parameres. The female of ovallesi
has elongate spermathecae, slightly constricted in the middle (Fig. 27J) unlike
those of nuneztovari, which are stout and non-constricted (Scorza et al., 1967,
in key and Fig. 23E this paper).
* Fairchild (pers. comm.) examined the holotype of nuneztovari in 1963, and
noted the following: Tips of genital filaments slightly expanded. Subterminal
seta of style lacking. Parameres long and slender, very slightly clubbed, not
pointed as shown by Ortiz (1954) but blunt, slightly shorter than lateral lobes.
Alpha of wing venation very long, at least equal to beta + gamma, not as shown
by Ortiz (1954).
The males from Huila, Colombia, agree with the description of the holotype
by Ortiz (1954) (as slightly emended by Fairchild), and I therefore have no doubt
that they are conspecific. The Colombia females of nuneztovari are in poor con-
dition with partially missing palpi and antennae, but other structures are intact
and are described as follows.
Female (n = 2): Wing length 2.25; width 0.74. Head, mesonotum, and ab-
dominal tergites strongly pigmented, rest of insect, including pleura, pale or
dusky, but the degree of pigmentation difficult todetermine due to overstaining.
Head height 0.42-0.43. Eyes separated by 0.11-0.12 or by distance = to ca.
6 facet diameters. Length of palpal segment 1 (0.05), 2 (0.19), others missing;
2 palpal sensilla on apical part of segment 2. Labrum 0.32 long. Cibarium with
4 sharp equidistant horizontal teeth and 12-20 small vertical teeth in a row;
chitinous arch complete; pigment patch subtriangular, the posterior (wider) half
more heavily pigmented. Pharynx 0.20 long, unarmed. Pleura with 12 upper
and 4 lower episternal setae. Length of wing vein sections: Alpha (0.71), beta
(0.22), delta (0.32), gamma (0.31). Length of femora, tibiae, and basitarsi:
Foreleg, 0.78, 0.86, 0.54; midleg, 0.83, 1.05, 0.64; hindleg, 0.88, 1.27, 0.71.






Abdominal sternite 2 with a median clear area not enclosed posteriorally. Te rg ite
8 with 5-7 setae on each side. Spermatheca wrinkled and sac-like, terminal
knob deeply recessed; common sperm duct and most of individual ducts
invisible.

31. Lutzomyia ovallesi (Ortiz)
(Fig. 27)
Phlebotomus ovallesi Ortiz, 1952:155 (d holotype, Duaca, Lara, Venezuela,
not San Felipe, Yaracuy, Venezuela). Ortiz, 1954: 239 (~, redescr., figs.). Lewis
& Garnham, 1959: 87 (9 descr., Belize). Fairchild & Hertig, 1959:122 (Central
American records). Scorza & Ortiz, 1960: 434 (ecology), Hanson, 1961: 317
(breeding sites), Johnson & Hertig, 1961: 765 (rearing data). Pifano et al., 1962:
370 et seq. (6, 9 keyed, redescr., figs,). Strangways-Dixon & Lainson, 1962:
297 (with flagellates, Belize). Johnson et al., 1963:110 (negative for flagellates).
McConnell & Correa, 1964; 527 (infected with fungus). Williams et al., 1965:
65, 70 (Belize). Rosabal, 1966:1 (Costa Rica). Biagi et al., 1966: 368 (Mexico,
9 figs.). Thatcher & Hertig, 1966: 50 (biting man and Potos, Panama). Scorza
et al., 1967: 191 (dist.). Hanson, 1968: 64 (larva, pupa, descr.). Osorno et al.,
1967: 29 (Colombia).
Sergentomyia ovallesi: Barretto, 1955: 185 (listed). Barretto et al., 1956: 51
(cf. tofirmatoi).
Lutzomyia ovallesi: Barretto, 1962:98 (listed). Aitken et al., 1968:264 (Trin-
idad, on man). Williams, 1970: 332 et seq. (summary of studies, refs., Belize).
Tesh et al., 1971a: 153 (blood meals). Chaniotis et al., 1971a: 100 (listed).
Osorno et al., 1972a: 28 (Colombian records). Shaw et al., 1972:719 (mention).
Chaniotis et al., 1972: 95 (collecting data). Christensen, 1972a: 88 (listed).
Christensen et al., 1972: 57 (light trap catch). Rutledge & Mosser, 1972: 366
(listed). Forattini, 1973: 122 et seq. (gen. review, refs., figs.). Lewis, 1975a:
500 et seq. (mouthpart morphol.). Rutledge et al., 1975: 181 (ecology). Rut-
ledge & Ellenwood, 1975a: 73 (ecology, refs.), 1975c: 87 (mention). Williams,
1976a: 603 (in caves, Belize); 1976b: 618 (mention).
Brumptomyia ovallesi: Lewis, 1965: 378 et seq. (internal morphol.).
Distribution: Mexico, Belize, Honduras, Nicaragua, Costa Rica, Panama,
Colombia (Antioquia, Boyaci, Caldas, Choc6, Cundinamarca, Magdalena,
Santander), Venezuela, Trinidad.
Material examined: Colombia. 34 6c, 19 99, Teresita (Choc6), in tree but-
tresses, March-Nov. 1967, D.G. Y. 1 c, near Rio Don Diego, E of Santa Marta
(Magdalena), 15 Aug. 1973, D.G.Y. & R.C.W. Panamd. 2 99, Almirante
(Bocas del Toro), Shannon trap, 13 July 1951, A. Quinones. Trinidad. Both
sexes from various localities, to be discussed in a forthcoming paper.
Discussion: The female of ovallesi, a man biter in several countries including
Colombia (Osorno et al., 1972a), is readily identified by the elongate constricted
spermathecae (Fig. 27J). Flagellates, apparently not Leishmania, have been
recovered from wild caught specimens in Panama and Belize (Williams, 1970).








B f


H- --
0\7; K-==, Z


FIGURE 27
Lutzomyiaovallesi A. Male head, B. Male flagellomere II, C, Female head, D. Female
flagellomere II, E. Female cibarium and pharynx, E Genital pump and filaments, same
scale as Fig 27G, G. Male genitalia, H. Female wing, I. Male wing, J. Spermathecae,
K. Female cibarium.
Male: Rio Sabana, Darien Prov., Panama
Female: Same locality as male


*u l,
j~ii


K






Evidence gathered by Thatcher and Hertig (1966) and Tesh et al. (1971a) indi-
cates that this species feeds primarily, if not exclusively, on mammals. In Pan-
ama, immatures were found in soil at the base of trees (Hanson, 1961), less
commonly on the open forest floor (Rutledge and Ellenwood, 1975a).

32. Lutzomyia quasitownsendi Osorno, Osorno & Morales
(Fig. 28)
Lutzomyia quasitownsendi Osorno, Osorno, & Morales, 1972b: 10 (6 hol-
otype, 9, Barbosa, Santander, Colombia). Osorno et al. 1972a. 29 (Colombian
records).
Distribution: Colombia (Santander).
Material examined: 1 6 (paratype no. 819), Barbosa (Santander), 30 Oct.
1968. 1 9 (paratype no. 2702), same data but taken on 23 June 1967. Both
specimens in collection at UF
Discussion: This little-known species was well described and illustrated by
Osorno et al. (1972b). I have nothing further to add.

33. Lutzomyia sauroida Osorno, Morales & Osorno
(Fig. 28)
Lutzomyia sauroida Osorno, Morales, & Osorno, 1972c: 433 (6 holotype,
vereda San Pedro, Municip. Santa Ana, Boyaca, Colombia; 9, same locality
except San Martin instead of San Pedro). Osorno et al., 1972a: 29 (Colombian
records).
Distribution: Colombia (Boyaca).
Material examined: Colombia. 1 3 (paratype no. 2690), 1 9 (paratype no.
2685, both in collection at INPES), type locality, 29 Oct. 1968.
Discussion: This species and L. townsendi (Ortiz) (Fig. 12) from Venezuela
are very similar. The males of sauroida have slightly stronger setae on the dor-
sum of the parameres but are otherwise indistinguishable. I can find no differences
between the females of the two species but hesitate to treat sauroida as a junior
synonym of townsendi until more material of the former species becomes
available.

34. Lutzomyia spinicrassa Morales, Osorno, Osorno & Hoyos
(Fig. 28)
Lutzomyia spinicrassa Morales, Osorno, Osorno, & Hoyos, 1969b: 383 (6C
holotype, vereda Umbabita, Municip. Almeida, Boyaca, Colombia; 9, vereda
Chitavita, Municip. Guayata, Boyach, Colombia). Osorno et al., 1972a: 30
(Colombian records). Forattini 1973: 213 et seq. (6 fig., keyed).
Distribution: Colombia (Boyaca).
Material examined: Colombia. 1 d (paratype no. 2654), 1 9 (paratype no.
2652), Guateque (Boyaca), 22 Oct. 1968, A. Morales & J. Ochoa. Both spec-
imens in collection at UF.




























-1//

-. .---, .






DE








FIGURE 28
Lutzomyia sauroida A. Male genitalia, B. Female cibarium.
Male: Municip. de Santa Ana, Boyaca Dept., Colombia
Female: Same locality as male

Lutzomyia quasitownsendi C. Male genitalia.
Male: Barbosa, Santander Dept., Colombia

Lutzomyia longiflocosa D. Male genitalia.
Male: Muncip. de Tello, Huila Dept., Colombia

Lutzomyia spinicrassa E. Male genitalia.
Male: Guateque, Boyaca Dept., Colombia

Lutzomyia odax F. Coxite tuft and paramere.
Male: Almirante, Bocas del Toro Prov., Panama






Discussion: The male of spinicrassa differs from other males in the verru-
carum group by the size and shape of the terminal spine of the style (Fig. 28D).
This species, known only from Boyaci Dept. at elevations ranging from 1430
to 1640 m above sea level, was found resting in tree holes and on walls inside of
houses.


VESPERTILIONIS GROUP THEODOR, 1965
With the center of distribution in Central America, the vespertilionis group
species are divided into 2 series following Theodor (1965)-the series vesper-
tilionis (5 spp.) and the series deleoni (3 spp.). Barretto (1962) created the
subgenus Coromyia to accommodate them and other species, mostly those in
the verrucarum group. Forattini (1971 a, 1973) divides the vespertilionis group
species by placing 1 in the subgenus Lutzomyia Franga and 2 in the subgenus
Dampfomyia Addis, while leaving the others in Coromyia. In my opinion, this
arrangement is unnecessary when the species are classified using a combination
of character states.
Two sibling species in the series vespertilionis are known to occur in Col-
ombia. The females feed on bats and are morphologically indistinguishable.

Key to Species
Males
1. Paramere with dorsal setae on distal half of structure ...............
.......... ....................... L. vespertilionis (Fig. 29)
Paramere with dorsal setae restricted to distal third of structure .......
.................... ............ L. isovespertilionis (Fig. 29)


35. Lutzomyia isovespertilionis (Fairchild & Hertig)
(Fig. 29)
Phlebotomus isovespertilionis Fairchild & Hertig, 1958a: 516 (J holotype,
S, reared from eggs laid by Y from tributary of Rio Cocoli, Canal Zone). John-
son & Hertig, 1961: 765 et seq. (rearing data). Thatcher & Hertig, 1966: 46
(mention). Hanson, 1968: 88-90 (larva, pupa, descr.).
Phlebotomus vespertilionis: Fairchild & Hertig, 1947b: 617 et seq. (in part,
descr., fig.).
Lutzomyia isovespertilionis: Barretto, 1962: 97 (listed). Chaniotis et al.,
1971a: 344 et seq. (in part, pop. dynamics, Panama). Chaniotis et al., 1972: 94
et seq. (resting sites, Panama). Christensen et al., 1972: 57 et seq. (collecting
data, infected with trypanosomes, Panama). Christensen, 1972a: 88-89 (men-
tion). Forattini, 1973: 122 et seq. (gen. review, fig.). Christensen & Herrer,
1973: 579 et seq. (collecting data, Panama). Rutledge & Mosser, 1975; 411
(mention). Herrer & Christensen, 1969: 62 (collecting data, Panama).















L B D jd
A C E


\I.






I ^ > G '* '^ Wr^ ^- -j -~^ _






i K Y
I M







FIGURE 29
Lutzomyia vespertilonis A. Male head, B. Male flagellomere II, C. Female head, D.
Female flagellomere II, E. Female cibarium and pharynx, G. Paramere, H. Paramere of
male from Panama, I. Male genitalia, J. Spermathecae, K. Female wing, L. Male wing,
M. Female cibarium.
Male: Rio Anori, Antioquia Dept., Colombia (except Fig. 29H)
Female: same locality as male
Lutzomyia isovespertilionis E Paramere.
Male: Panama Canal Zone






Distribution: Panamd, Colombia (Choc6).
Material examined: Colombia. 1 S, Curiche (Choc6), tree buttress, 5 July
1967, D.G.Y. 4 dd, Teresita (Choc6), tree buttress, 18 May 1967, D.G.Y.
Panamd. 1 6, Cruces trail, Madden Forest, Canal Zone, hollow tree, 7 Sept.
1949, M. Hertig. 3 S Cerro Campana (Panama), tree hollows and buttresses,
2 Oct. 1949, M. Hertig & H. Trapido.
Discussion: See L. vespertilionis


36. Lutzomyia vespertilionis (Fairchild & Hertig)
(Fig. 29)
Phlebotomus vespertilionis Fairchild & Hertig, 1947b: 617 (d holotype,
Cerro Campana, Panama Prov., Panama). Hertig & Fairchild, 1950: 91 (ab-
dominal sternites). Barretto, 1951: 225 (dist.). Vargas & Diaz-Najera, 1951b:
101 (cf. to beltrani). Floch & Abonnenc, 1952: 30-48 (d, ?, keyed). Rosa-
bal, 1954: 10, 28-29 (measurements, figs., Costa Rica, sp. no. 12 = vesperti-
lionis).Rodriguez, 1956: 76 et seq. (Ecuador). Fairchild & Hertig, 1958a: 510
(keyed), 514 (d, 9, redescr., figs.). Fairchild & Hertig, 1959: 122 (Central
American records). Hanson, 1961: 320 (breeding site, Panama). Johnson &
Hertig, 1961: 765 et seq. (rearing). McConnell & Correa, 1964: 523 et seq.
(infected with trypanosomes & fungi, Panami). Thatcher & Hertig, 1966: 46
et seq. (mention). Hanson, 1968: 88-90 (larva, pupa, descr., figs.).
Lutzomyia vespertilionis: Barretto, 1962: 97 (listed). Theodor, 1965: 184
(d, 9, figs.). Barreto, 1969:468 (Valle, Colombia). Zeledon & Rosabal, 1969:
221- 227 (as vector of bat trypanosome). Tesh et al., 1971 a: 152- 153 (in part,
blood meals). Chaniotis et al., 1971a: 344 et seq. (in part, pop. dynamics. Pan-
ama). Christensen et al., 1971: 116 et seq. pyloricc armature, figs.). Tesh et al.,
1972: 90 et seq. (in part, blood meals). Chaniotis et al., 1972: 94 et seq. (resting
sites, Panami). Christensen, 1972a: 88-89 (mention). Christensen, 1972b:
683 et seq. (rearing). Osorno et al., 1972a: 32- 33 Colombian records). Ru-
tledge & Mosser, 1972: 307 (mention). Christensen & Herrer, 1973: 579 et seq.
(collecting data, infected with trypanosomes, Panama). Forattini, 1973: 123 et
seq. (gen. review, figs.). Christensen & Herrer, 1975: 477-478 (as host for bat
trypanosomes). Rutledge & Mosser, 1975: 411 (mention).
Distribution: Costa Rica, Panamd, Colombia (Antioquia, Bolivar, Boyaci,
Choc6, Valle), Ecuador.
Material examined: Colombia. 3 d, Rio Anori (Antioquia), tree trunks, 6
Feb. 1970, C.H.P. 2 dS, 1 9, same data but Sept. 1970, D.G.Y. 4 Sd, 2
99, Anchicaya Dam (Valle), tree trunks, 9 Aug. 1974, D.G.Y. & R.C.W 7
SS, 14 99, same data but light trap at entrance of tunnel, 10- 11 Aug. 1974.
1 c, 1 25 km E of Buenaventura (Valle), tree trunks, 11 Aug. 1974, D.G.Y,
& R.C.W Ecuador. 15 SS 1 9, 17 km E of Santo Domingo de los Colorados
(Pichincha), tree trunks, 4-6 May 1976, D.G.Y. & T. Rogers. Panamd. 1 I
holotypee no. 761), Cerro Campana (Panami), shallow cave, 17 Jan. 1947.






Discussion: In structure and habits, L. vespertilionis and L. isovespertilionis
are remarkably similar, both species often being found in tree hollows associ-
ated with bats. At least one species of Trypanosoma is transmitted to these
mammals by one or both of these flies (Zeledon and Rosabal, 1969; Christensen
and Herrer, 1975).
The preferred breeding site(s) of vespertilionis and isovespertilionis has not
been determined (Rutledge and Mosser, 1972), the only wild caught larvae
being reported by Hanson (1961, 1968), who recovered this stage on two oc-
casions in soil and bat guano inside of hollow trees. Apparently the larvae and
pupae of these species are indistinguishable (Hanson, 1968).
Lutzomyia vespertilionis ranges as far south as Guayas Province, Ecuador.
Lutzomyia isovespertilionis probably has a more limited distribution as noted
by Fairchild and Hertig (1958a), its southernmost locality at present being
Choc6 Dept., Colombia. These are the only vespertilionis group species which
occur in South America. Fairchild and Hertig (1958a) provide further infor-
mation on collecting data, distribution, and taxonomy.



SUBGENUS DAMPFOMYIA ADDIS, 1945
The 5 species in this subgenus are characterized by several features (Fairchild
and Hertig, 1956; Theodor, 1965), some of which are shared by L. aquilonia
(Fairchild and Harwood) of the vespertilionis group and members of the del-
pozoi group Lewis et al. (1977). The spermathecae of the Dampfomyia species
are large with bubble-like expansions and could have been derived from the
deleoni-type spermathecae of the vespertilionis group. Like species in these
two groups, the Dampfomyia spp. do not ordinarily bite man, and there are
more species in Mexico and Central America than elsewhere.
The preferred host(s) of L. rosabali, the only Dampfomyia sp. in Colombia
or South America, is unknown. Related species such as L. anthophora (Addis)
and L. permira (Fairchild and Hertig) feed on small mammals, the former spe-
cies probably transmitting Rio Grande virus to Neotoma woodrats in Texas
(Calisher et al., 1977).

37. Lutzomyia (D.) rosabali (Fairchild & Hertig)
(Fig. 30)
Phlebotomus rosabali Fairchild & Hertig, 1956: 310 (6 holotype, Puerto
Armuelles, Chiriqui Prov., Panama; 9, Guarare, Los Santos Prov., Panama).
Rosabal, 1954: 10 et seq. (as sp. no. 9, Costa Rica). Fairchild & Hertig, 1959:
122 (Central American records). Rosabal & Trejos, 1964: 168 (cf. to dodgei).
Lutzomyia rosabali: Barretto, 1962: 95 (listed). Theodor, 1965: 193 (listed,
spermatheca fig.). Forattini, 1971a: 100 (listed). Osorno et al., 1972a: 61 (Na-
riiio, Colombia). Christensen, 1972a: 88 (listed). Forattini, 1973: 208 (sper-
matheca fig.), 329 (as synonym of dodgei).










































..0

I' .. ci.








S/M


FIGURE 30
Lutzomyia (D.) rosabali A. Male head, B. Male flagellomere II, C. Female head, D.
Female flagellomere II, E. Female cibarium and pharynx, Panamanian specimen, E Male
genitalia, G. Dorsal arm of paramere, H. Different views of paramere, I. Genital pump
and filaments, J. Spermathecae, K. Female wing, L. Male wing, M. Female cibarium,
Panamanian specimen.
Male: Capuli, Narifio Dept., Colombia
Female: Same locality as male (except Fig. 30E and 30M)






Distribution: Costa Rica, Panamd, Colombia (Cauca, Narifio).
Material examined: Colombia. 6 6d, 8 9, Capuli (Narifio), rock crevices,
March 1945, M. Hertig. 3 99, Bolivar (Cauca), Jan.-Feb. 1944, J. Burbano.
Panamd. 1 & holotypee no. 3209), type locality, tree buttress, 9 June 1951,
M. Hertig. 1 9 (allotype), Guarare (Los Santos), light trap, 23 Oct. 1952,
E Blanton.
Discussion: Forattini (1973) treats L. rosabali as a junior synonym of L.
dodge (Vargas and Diaz-Najera), believing, perhaps correctly, that the
differences between the males are due to intraspecific geographic variation.
Without a larger sample of specimens from various localities, however, I prefer
to consider them a distinct based on the discussion and figures of Fairchild and
Hertig (1956).
The Colombian males (Fig. 30) agree more closely with L. rosabali than
with L. dodgei in the shape and station of the dorsal arm of the paramere, but
the basal spine of the style is longer and more slender. The small bristle near
the basal spine is absent on one style of the rosabali holotype and from the
styles of the Colombian males.


SUBGENUS PINTOMYIA COSTA LIMA, 1932
The 5 described species in this subgenus are characterized by a row of short
spines on the hind femur. Forattini (1971a, 1973) treats Pintomyia as a genus,
placing, I believe, undue importance on these spines, which, although not ex-
actly the same, are shared by 2 species of Sergentomyia Franga and 1 of Brump-
tomyia.
The Pintomyia females, like those of Pressatia, have well sclerotized indi-
vidual sperm ducts but differ from them in other aspects (Theodor, 1965).
At present, only L. spinosa is known to occur in Colombia. Lutzomyia da-
mascenoi (Mang.), a close ally, was reported in Meta Dept. by Morales et al.
(1969a), but was excluded in a later work on the Colombian fauna (Osorno
et al., 1972a), probably because of the uncertain identity of the specimens
(2 $9).
Except for L. pessoai (Coutinho and Barretto) and L. fischeri (Pinto), the
Pintomyia spp. have not been reported feeding on man. Lutzomyia gibsoni (Pi-
fano and Ortiz), recently described from Venezuela from a single female, defi-
nitely belongs in the subgenus, but its identity remains uncertain on the basis
of the original description.

38. Lutzomyia (Pi.) spinosa (Floch & Abonnenc)
(Fig. 31)
Phlebotomus spinosus Floch & Abonnenc, 1942b: 1 (6 holotype, Gallion,
near Cayenne, French Guiana). Barretto, 1947: 224 (full refs.). Hertig & Fair-
child, 1950: 92, 93 (abdominal sternites 1 & 2, figs., Panama). Barretto, 1951:
224 (dist.). Floch & Abonnenc, 1952: 20 et seq. (3, 9, keyed, redescr., figs.).




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