A BIOSYSTEMATIC REVIEW GF THE BLOODSUCKING FSYCHODID FLIES OF COLOMBIA (DIPTERA: PHLEBOTOMIHAE AND SYCORACTNAE) By 'ID GRIER YOUNC A DISSERTATION PRESENTED TO THE GRADUATE COUNCIL OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY UNIVERSITY OF FLORIDA 1077 Dedicated to my father, Howard G. Young TABLE OF CONTENTS Page LIST OF FIGURES viii ABSTRACT xii INTRODUCTION 1 CLASSIFICATION AND TAXONOMIC TREATMENT A Keys to the Subfamilies of New World Psychodidae 6 GENERAL MORPHOLOGY AND TERMINOLOGY 8 METHODS AND MATERIALS 15 Field Collections 15 Laboratory Methods 17 DISTRIBUTION, COLLECTING LOCALITIES, AND CHARACTERISTICS OF THE FAUNA 19 Pacific Coast Region 20 Atlantic Coast Region 23 Andean Region 24 Oriental Region 25 MEDICAL IMPORTANCE 29 SUBFAMILY SYCORACINAE 31 Keys to the Sycorax Species 33 1. Sycorax coidieola n. sp 35 2. Sycorax colombiensis n. sp 36 3. Sycorax fairchildi n. sp 3rf 4. Sycorax trispinosa n. sp 30 SUBFAMILY PHLEBOTOMINAE 50 Key to the New World Genera 50 Genua Wariloya Hertig, 1948 52 Keys to Species 52 5. WariZeya (W.J nigrosacculus 53 6. Wan'.teya (W.) rotundipennis 5: Page Genus Brumptomyia Franga & Parrot, 1921 60 Key to Species 60 7. Brumptomyia av?.llavi 61 8. Brumptomyia beaupevtuyi 62 9. Brumptomyia galindoi 63 10. Brumptomyia hanata 65 11. Brumptomyia leopoldoi 66 Genus Lutzomyia Franca, 1924 76 Keys to Subgenera, Species Groups, and Ungrouped Species. . . 76 Subgenus Lutzomyia Franca, 1924 9.1 Keys to Species 91 12. Lutzomyia (L.) bifoliata 93 13. Lutzomyia (L.) lichyi 94 14. Lutzomyia (L.) longipalpis 96 15. Lutzomyia (L.) gomezi 97 16. Lutzomyia (L.) rnarinkellci n. sp 99 Migonei Group Theodor, l96i> Ii2 Keys to Species 112 17. Lutzomyia migonei 113 18. Lutzomyia marajoensis 114 19. Lutzomyia sp. de Baduel 116 20. Lutzomyia waVkevl 116 Saulsnvis Group Lewis et al., 1978 125 21. Lutzomyia saulensia 125 Vervucarum Group Theodor, 1965 130 Key to Species 130 22. Lutzomyia odax 133 23. Lutzomyia sevvona 134 24. Lutzomyia andina 135 25. Lutzomyia columbiana 136 26. Lutzomyia disiuncta 137 27. Lutzomyia evansi 137 28. Lutzorrpjia lonciflocosa 138 29. Lutzomyia movalesi n. sp 139 30. Lutzomyia nuneztovari 141 31. Lutzomyia ovaltesi 143 32. Lutzomyia quasitownsendi 14 5 33. Lutzomyia sauvoida 145 34. Lntzovvyia spinicrassa 146 Vbspertitionis Group Theodor, 1965 161 Key to Species 161 Page 35. Lutzomyia isovesperti.li.onts 161 35. Lutzomyia vespertilionis 162 Subgenus Damp forty ia Addis, 1945 167 37. Lutzomyia (D.) rosdbali 167 Subgenus Pintomyia Costa Lima, 1932 171 38. Lutzomyia (P.) spinosa 171 Subgenus Pressatia Mangabeira, 1942 176 Keys to Species 176 39. Lutzomyia (Pp.) oamposi 177 40. Lutzomyia (Pr.) dysponeta 178 41. Lutzomyia (Pr.) triaoantha 180 Baityi Group Theodor, 1965 186 42. Lutzomyia baityi 180 Subgenus Viannamyia. Mangabeira, 1941 190 Keys to Species 190 43. Lutzomyia (V.) caprir/x 19.1 44. Lutzomyia (V.) furcata 192 45. Lutzomyia (V.) tuberculata 194 Shannoni Group Theodor, 1965 200 Keys to Species 200 46. Lutzomyia abonnenci 202 47. Lutzomyia dasymera 203 48. Lutzomyia dendrophyla 204 49. Lutzomyia punctigenioulata 206 50. Lutzomyia shannoni 208 51. Lutzomyia. undulata 210 Longispina Group Theodor, 1965 224 Keys to Species 224 52. Lutzomyia longispina 225 53. Lutzomyia triramula 226 Aragaoi Group Theodor, 1965 232 Keys to Species 232 54. Lutzomyia aragaoi 234 55. Lutzomyia barret toi b arret boi 236 56. Lutzomyia barrettoi najuscula n. ssp 237 57. Lutzomyia eavpenteri 241 58. Lutzomyia. runoides 243 Page Drcisbachi Group Lewis et al . , 1978 255 Key to Species 25 5 5C2. Lutzomyia aolydifera 255 60. Lutzomyia drsisbaoki 257 Subgenus Triahopkororrryia Barretto, 1962 261 Key to Species 261 61. Lutzomyia (T.) auvaensis 262 62. Lutzomyia (T.) cellulana n. sp 264 63. Lutzomyia (T.) kouardi n. sp 266 64. Lutzomyia. (?.) rebur-va 267 65. Lutzomyia (T.) saltu^sa n. sp 269 66. IjUtzomyia (T.) ubiquitalis 270 Subgenus Nyssomyia Barretto, 1962 280 Keys to Species 280 67. Lutzomyia (II.) antunesi 283 68. Lutzomyia. ([.'. ) flavisscuLallata 285 69. Tjutzomyia (II.) olneoa bioolov 287 70. Lutzomyia (H.) brapidoi 289 71. Lutzomyia (!!.) urnbratilis 292 72. Lutzomyia (N.) ylephiletor 295 73. Lutzomyia (N.) yuilli 297 Subgenus Psycho dopy gus Mangabeira, 1941 311 Keys to Species 312 74. Lutzomyia (P.) bernalei 317 75. Lutzomyia (P.) chagasi 317 76. Lutzomyia (P.) fairtigi 318 77. Lutzomyia (P.) bispinosa 319 78. Lutzomyia (P.) ayrozzi 320 79. Lutzomyia. (P.) oar-rerai oewevai 323 80. Lutzomyia (P.) oarrerai thula n. ssp 326 81 . Lutzomyia. (P. ) davisi 331 82. Lutzomyia (P.) guyanensis 333 83. Lutzomyia (P.) hirsuta 335 84. Lutzomyia (P.) nooticola 338 85. Lutzomyia (P.) panarr.ensis 339 86. Lutzomyia (P.) vecuvva 341 87. Lutzomyia (P.) sp. of Tres Esquinas 342 VexatoT Group Theodor, 1965 366 Keys to Species 36 7 88. Lutzomyia cir-rita 369 89. Lutzomyia harbmar.ni 370 90. Lutzomyia. oso/noi 37.1 9.1. Lutzomyia sar.guinar-ia 372 Page 92. Lutzomyia scorzai 374 93. Lutzomyia strictivilla 378 94. Lutzonyia sp. of Pichinde. 380 Cayennensis Group Theodor, 1965 398 Keys to Species 398 95. Lutzomyia cayennensis 399 96. Lutzomyia micropyga 401 9 7. Lutzomyia atroolavata 405 98. Lutzomyia venezuelensis 407 Oswaldci Group Theodor, 1965 417 Keys to Species 417 99. Lutzomyia via 418 100. Lutzorwyia vovotaensis 420 101. Lutzomyia trinidadensis 422 LUL>u;>a utuu[j jii£uuCl, irOJ h.iZ 102. Lutzomyia pilosa 432 Ungrouped Species 4 37 103. Lutzomyia nordestina 4 37 104. Lutzomyia vangeliana 4 39 105. Lutzomyia sp. of Anchicaya , 440 SYNOPSIS OF NEW TAXA, SYNONYMS, FEMALE DESCRIPTIONS, AND NEW DISTRIBUTION RECORDS 448 REFERENCES 451 BIOGRAPHICAL SKETCH 481 LIST OF FIGURES Figure Page 1. Map of Colombia showing localities where phlebotomine and sycoracine flies have been collected 23 2. Syoo/'ox andicola 43 3. Syaorax colombiensis 45 4. Sycorax faivchildi 4 7 5. Syoorax trispinosa 49 6. V:av it °-ya nigvo^or-riylvs -- ,...,,<.-.,....,,., 57 7. Wavileya rotundipennis 59 8. Bmonptomyia avellai-i and Brumptomyia beaupertuyi 69 9. Brumptomyia galindoi 71 10. Erumptomyia hamata 73 11 . Bv.mptomyia leopoldoi and Brvrrotomyia guimavaesi 75 12. Lutzomyia (P.) hirsuta niearacuensis and Lutzomyia tounsendi 90 13. Lutzomyia (L.) bifoliaia 103 14. Lutzomyia (L.) liohyi 105 15. Lutzomyia (L.) longivalpis 107 16. Lutzomyia (L.) gomezi 109 17. Lutzomyia (L.) mavinkellei Ill 18. Lutzomyia migonei and Lutzomyia sp. de Baduel 120 19. Lutzomyia marajoensis 122 20. Lutzomyia walkeri 124 21. Lutzomyia saulensis 129 Fi gur e Page 22. Lutzomyia serrana 148 23. Lutzomyia andina, Lutzomyia disiuneta, and Lutzomyia nuneztovavi 150 24. Lutzomyia aolurribiana 152 25. Lutzomyia evansi 154 26. Lutzomyia moralesi 156 27. Lutzomyia ova'llesi 158 28. Lutzomyia sauvoida, Lutzomyia quasi towns endi > Lutzomyia. tongiflocosa} Lutzomyia spiniopassa, and Lutzonyia odax. . . 160 29. Lutzomyia vesper til ionis and Lutzomyia isovespsrtilionis . . 166 30. Lutzopvyia (D.i rosaboli 170 31. Lutzomy%a (Pi.) spinosa 175 32. Lutzomyia (Pp.) eamposi 183 33. Lutzomyia (Pp.) dysponeta 185 34. Lutzomyia (Pp.) tpicantha and Lutzomyia ha.ityi 189 35. Lutzomyia (V.) cappina and Lutzomyia (V. ) furaata 197 36. Lutzomyia (V.) tubepoulata 199 37. Lutzomyia abonnenei 213 38. Lutzomyia dasymeva 215 39. Lutzomyia dendpophyla 217 40. Lutzomyia punetigeniculata 219 41. Lutzomyia shannoni 221 42. Lutzomyia undulata 223 43. Lutzomyia longispina 229 44. Lutzomyia triramula 231 45. Lutzomyia avagaoi 246 46. Lutzomyia barvettoi majusoula 248 Figure Page. 47. Lutzormjia barrettoi maQusoula3 Lutzormjia barret toi barrettoi, and Lutzorrryia texana 250 43. Lutzormjia cavpentevi 252 49. Lutzormjia vunoides 254 50. Lutzormjia aelydifeva 260 51. Lutzormjia dreisbachi, Lutzormjia (T.) auraensisj and Lutzormjia ubiquitaiis 273 52. Lutzormjia (T.) cetlulana 275 53. Lutzormjia (T.) howardi and Lutzormjia (T.) sdltuosa 277 54. Lutzormjia (T.) reburra 279 55. Lutzormjia (f-1.) antunesi and Lutzormjia (N.) flaviscutellata. 300 56. Lutzormjia (N.) olmeca bioolor 302 57. Lutzormjia (N.) trapidoi 304 58. Lutzorrpjia (11.) umbratitis 306 59. Lutzormjia (N.) ylephiletor 308 60. Lutzormjia (N.) yuilli 310 61. Lutzonrjia (P.) bevnalei, Lutzormjia (P.) ch.aga.si, and Lutzormjia sp. of Tres Esquinas 345 62. Lutzormjia (P.) fairtigi 347 63. Lutzomjia (P.) bispinosa 349 64. Lutzormyia (P.) ayvozai 351 65. Lutzormjia, (P.) cawevai carrerai 353 66. Lutzormjia (P.) cavrerai thula 355 67. Lutzormjia (P.) davisi and Lutzormjia (P.) hirsuta hirsute/.. . 357 68. lutzormjia (P.) guyanensis 359 69. Lutzormjia (P.) nooticola 361 70. Lutzorrpjia (P.) panamensis 363 71. Lutzormjia (P.) vecurva 365 Figure Page 72. Lutzomyia air-vita 385 73. Lutzomyia kca>tmanni 387 74. Lutzomyia osornoi 389 75. Lutzomyia sanguinaria 391 76. Lutzomyia strictivilla 393 77. Lutzomyia soorzai 395 78. Lutzomyia sp . of Pichinde 39 7 79. Lutzomyia cayerm&nsis 410 80. Lutzomyia micropyga 412 81. Lutzomyia atroolavata 414 82. Lutzomyia venezuetensis 416 83. Lutzomyia pia 427 84. Lutzomyia r-orotaensis 429 85. Lutzomyia tvinidadensis 431 86. Lutzomyia, pilosa 4 36 87. Lutzomyia nordestina 443 88. Lutzomyia rangeliana 445 89. Lutzomyia sp . of Anchicaya 447 Abstract of Dissertation Presented to the Graduate Council if cite University of Florida in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy A BIOSYSTFi'tATIC REVIEW OF THE BLOODSUCKING PSYCHCDID FLIES OF COLOMBIA (DJ.PTERA: FHLFBOTOMINAE AND SYCORACINAR) By David Crier Young December, 1.977 Chairman: Graham B. Fairchild Major Departmenc: Entomology and Nernatology The psychodid subfamily Sycoracinae, previously unknown in Colombia, is represented by four new Sycovcx species described and illustrated in this review. Phlebotorainaa, a much larger subfamily containing vectors of human pathogens, is represented in the Republic by 101 species and subspecies — a number equivalent to one-third of the described New World taxa. Ten of these in the genus Lutzomyia. are described as new and are given formal or informal names. The previously urknown females of Lutzomyia pilosa, L. (P.) hi-vsuta ni ccwaguensis ?nd L. bifoliata are described. Nineteen phlebotcmin.e species are reported in Colombia for the first time. Other new distributional records include those from Ecuador (22 spp.), Panama (3 spp.), Honduras (2 spp.), Costa Rica, Cuba, Guatemala, and Paraguay (1 sp. each). In order to clarify the status of some forms, an attempt ■.-/as made to study specimens from as many Colombian and non-Colombian localities as possible. To this end, over 20,000 adults in both sub- families were examined from 1967 to the present. An extensive bibliography and synonomy is given for each previously described taxon. Actual and/or probable misidentif ications in the literature are discussed. identification keys, previously unavailable for the Colombian fauna, are provided along with numerous illustrations, most of ivhich are original. Although more information is needed, it is suggested that recent speciation in the neotropical phlebotominae was due to climate changes in the Pleistocene (refuge theory) . INTRODUCTION Prior to the last decade, we knew very little about the phlebotomine sand fly* fauna of Colombia. Osorno et al . (1967) in reviewing previous studies by others and adding records of their own, reported less than 25 species in the Republic. From 1967 to 1972, the late Dr. E. Osorno-Mesa and his colleagues at Instituto Nacional de Salud, Bogota added more records, bringing the total to 77 species and subspecies (Osorno et al., In this review 101 species and subspecies of Phlebotominae are re- ported 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 south- eastern region. The small subfamily Sycoracinae, whose members resemble those of Phlebotominae in being able to take vertebrate blood, is treated here for this reason. In order to clarify the status of some taxa and to study intraspeci- fic variation and distributional patterns, an effort was made to examine specimens from as many Colombian and non-Colombian localities as possible. To this end, I am vary grateful to the following persons for providing specimens, information or both: Dr. Jorge R. Arias, Instituto Nacional de Pesquisas da Amazonia (INPA) , Manaus ; Drs . Stephen C. Ayala and Pabln *I prefer to separate "sand" from "fly" in accordance with the suggestion of Borrer, Delong, and Triplehorn (1976) . Barretto, Universidad del Valle, Cali' Drs. Richard D. Ward and Habib Fraiha, Institute Evandro Chagas , Belem; Dr. David J. Lewis, British Museum (Nat. Hist.); Dr. Cornells J. Marinkelle, formerly Universidad de Los Andes, Bogota; Dr. AlberLo Morales-Alarcon and Che lace Dr. Ernesto Osorno-Mesa, Instituto Nacional de Salud, Bogota; Prof. Almilcar Vianna Martins, Universidade Federal de Minas Gerais; Dr. Nelson Papavero, Museu de Zoologia, Universidade de Sao Paulo; Drs. Charles H. Porter and Thomas M. Yuill, University of Wisconsin; Dr. Alan Stone, formerly Systematic Entomology Laboratory (ARS) ; Mr. Richard C. Wilkerson, University of Florida. Dr. Thomas H.G. Aitken, Yale University School of Medicine; Dr. Elisha S. Tikasingh, Caribbean Epidemiology Center (CAREC) , Trinidad; Dr. Harold Trapido, Louisiana State University Medical Center; Dr. Jorge E. Velasco, La Paz, Bolivia; Dr. Derek A. Duckhouse, University of Adelaide, South Australia; Maj . John F. Reinert, U.S. Army Medical Research and Development Command, Washington, D.C. Others who have supported this study and to whom I am indebted include members of the Atlantic-Pacific Interoceanic Canal Study Commis- sion, Canal Zone from 1967-1968. Col. Bruce F. Eldridge of this organi- zaCion was especially helpful in developing field programs and in providing data on Choco collections. Financial support from U.S. Army Medical Research and Development Command under contract no. DADA 17-72-C-2139 is gratefully acknowledged . Such support was provided also at various times by Drs. William G. Eden and Hugh Popenoe, University of Florida. Members of my graduate school committee at the University of Florida have been helpful in many ways, too numerous to mention here. For their help, T thank Drs. Franklin S. Blanton, Jerry F. Butler, Graham B. Fairchild, Dale H. Habeck, and Stephen G. Zam. Miss Stephanie Haney ably slide mounted thousands of specimens from 1972 to 1977. For typing the manuscript, I wish to thank Mrs. Adele Koehler. Dr. G.B. Fairchild deserves special recognition for his constant encouragement, stimulating discussions and expert assistance throughout the study, I ai., deeply indebted to him. Particular appreciation is expressed also to my wife Molly who was most understanding during the preparation of this review. Holotypes and allotypes of new taxa are to be deposited in the U.S. National Museum (Nat. Hist.). Paratypes, when available, will be held by the following institutions: Instituto Nacional de Salud, Bogota; Florida State Collection of Arthropods, Gainesville; and British Museum (Nat. Hist.), iondon. The remaining 20,000+ specimens, upon which most of this review is based, are stored in the author's personal collection. A synoptic collection will be given to the Instituto Nacional de Salud, Bogota. CLASSIFICATION AND TAXONOMIC TREATMENT Of the 6 recognized subfamilies of Psychodidae (Duckhouse, 1972, 1973), all but one, Horaiellinae from the Oriental Region, are represented in continental Colombia. The species of IloraieVla, Trtahomyia, and Suaorax and its allies are united by some in the subfamily Trichomyiinae but this appears to be artificial and does not "express the wide discon- tinuities and degrees of difference from other Psychodidae, especially in LIU- i'»»^^.v. .,._^.t,v,., v^^C OUS>C, A.JI*.;. The classification of Phlebotominae is interpreted differently according to author(s) . In a recent paper we reviewed most classifica- tory schemes and proposed one which is used here for the New World taxa (Lev/is et al . , 1978). It is essentially a modification of Theodor's 1965 classification. Three American genera are recognized — LutzonryLa Franca (ca. 290 spp.), Brumptomyia Franca and Parrot (ca. 23 spp.), Wariteya Hertig (5 spp.). The genus Hex'tigia Fairchild is considered to be a junior synonym of Waviteya by Lewis et al. (1978). The majority of the Lutzomyia spp. were described originally in the genus Phlebotomus (= Flebotomus) Rondani. Some authors continue to place them in this genus, ignoring the evidence to the contrary provided by Theodor (1948, 1965) and Christensen et al. (1971). The large genus Lutzomyia is divided into numerous subgenera, species groups, and series based upon adult morphology (Lewis et al., 197S) . The diversity of groups within the genus is attributed to "radiation or even 'explosive radiation' of some immigrating ancestral stocks in a newly colonized area having a wealth of ecological niches" (Theodor, 1965). This implies that these ancestral stocks originated in the Old World -- a hypothesis which may never be proven. It would appear that recent speciation in some groups (e.g. subgenus TlH-chophovornyia) may have been due to climatic changes in the past, especially during the Pleistocene, which served to isolate conspecific populations in moist refugia during dry periods (Haffer, 1974). Other remarks on the classification of New World phlebotomines are given by Lewis et al. (1978) and in this review under each supraspecif ic taxon. An extensive bibliography and synonomy is given for most taxa treated here. I usually omitted references which contain species lists but some of them including those by Fairchild (1955), Barretto (1955a, 1962), Theodor (1965) , and Martins & Morales (1972) are very important because of discussions related to classification and distribution. For each species, I cite the original description as well as most others published since Barretto' s 1947 catalog. In cases where a species was reviewed subsequent to 1946 (e.g. Forattini, 1973), I refer to that publication for full references. Abbreviations of words used in the text which may not be familiar to some readers are given below: Inst i tutions USNM — United States National Museum (Nat. Hist.), Washington, D.C. INPES — Instituto Nacional para Programas Especiales de Salud, Bogota INPA — Instituto Nactional de Pesquisas de Amazonia, Manaus UV -- Univcrsidad del Valle, Call GML -•- Corgas Memorial Laboratory, Panama UF — University of Florida Collectors C.H.P. •— Charles H. Porter C.J.M. — Cornelius J. Marinkelle R.C.lv. — Richard C. Wilkerson D.G.Y. — David G. Young Other Abbreviations ca. — about cf. — compared to classif. -- classification dsscr. — description (list. -- distribution morphol. — morphology pop. - — population Key to the Subfamilies of Psychodidae of the New World 1. Wing venation with 4 branched radius, P?,o fused into a single vein (Fig. 2B) 2 Wing venation with 5 branched radius (Fig. 12G) 3 2. Wing venation with short cubitus (cu) . Mandibles present Sycoracinae (Fig. 2) Wing venation with long cubitus. Mandibles absent. . . .Trichomyiinae -/- 3. Palpus of 4 segments, rarely 3. Eyes usually with eye bridge. Antennal f lagellomeres nodiform or barrel shaped Psychodinae Palpus of 5 segments. Eyes without eye bridge. Antennal f lagel- lomeres subcylindrical 4 4. Females with 1 spermatheca. Mandibles absent. Male aedeagus entire Bruchorayiinae Females with 2 spermathecae. Mandibles present. Males with bifid aedeagus. . . Phlebotominae (Fig. 12F) GENERAL MORPHOLOGY AND TERMINOLOGY This brief discussion focuses on the structures commonly used in taxonomic studies of phlebotomine sand flies. General accounts of adult anatomy include those by Fairchild and Hertig (1947a), Kirk & Lewis (1951), Abonnenc & Minter (1965), Davis (1967), Abonnenc (1972), Lewis (1973), and Forattini (1973). The terminology varies according to author; that used here applies also to the sycoracine flies except where otherwise stated. Although little studied, the immature stages are probably no less important in systematics; they are not considered here owing to lack of material. For information on the anatomy of these stages see Barretto (1941), Hanson (1968), Abonnenc (1972), Carnheiro & Sherlock (1964), Ward (1977), and Zimmerman et al. (1977). Descriptions of immatures of those species occurring in Colombia are cited in the appropriate species bibliography. The sand fly head and its appendages (Fig. 12A) are useful in associating conspecific males and females and for grouping taxa below the genus level. Head height, measured from the vertex to the tip of the clypeus, in relation to its greatest width is of value in distinguishing some of the Lutzomyia species. As a probable adaptive feature related to host finding, the size of the compound eyes varies according to species, especially those of Lutzomyia, but may be characteristic of subgenera and equivalent species groups. Contrary to Divis (1967), ocelli are lacking in the Phlebotomines and Sycoracines. Illustrations are by far the best way to indicate eye size, the relative terms such as "snail" or "large" being understood easily with associated figures. The interocular distance or eye separa- tion is the narrowest distance between the eyes and is measured directly or may be expressed by the number of facet diameters (or fractions there- of) needed to connect the eyes. The interocular suture (= post frontal suture of mosquitoes, Christophers, 1960) is complete in adults of BrumptonyLa (Fig. 8B) , Wavileya (Fig. 6A) , and Syaovax (Fig. 2A) but is incomplete in those of Lutzomyia (Fig. 12A) . In Phlebotominae, the maxillary palpus (Fig. 12A) consists of 5 segments, the basal one (palp 1) always the smallest and least important in taxonomy (Lev/is, 1973). Being partially fused with the second, it is difficult to accurately measure. The relative lengths of the segments are expressed often by a palp formula or by ratios with palp 1 being unity or 10 (Kirk &. Lewis, 1951). A palp formula of 1-4-2-3-5 indicates that palp 1 is the shortest, palp 4 the next shortest, etc. When two segments are equal in length they are enclosed in brackets, e.g. 1- (4-2) -3-5. The sycoracines have four, instead of five, palpal segments (Fig. 2E) . Species in both subfamilies have palpal sensilla (= Newstead's scales) which are small clubbed sensory organs attached to one or more segments in the phlebotomines and to palp 2 in the Colombian Syoovox spp. The cylindrical antennal f lagellomeres, unlike true segments, are not independently musculated (Imms, 1938). For this reason I join the increasing number of students who use the term "flagellomere" in the place of "antennal segment." The antenna consists of a scape, pedicel, -10- and 14 flagellomeres, each of which is designated by a Roman numeral (Fig. 12,\;. In Sycorax the terminal flagellomere is markedly reduced in size (Fig. 2D). The length of flagellomere 1 (= antennal segment 3), the most basal flagellomere, in relation to other flagellomeres, head height, or labrum length is useful as a diagnostic feature at the species or subspecies level. The paired antennal ascoids (fig. 12B) , often difficult to observe, are important owing to interspecific variation in their distribution, shape, and length. Some specimens may have but a single ascoid on a particular f lageilcmere. The raouthparts of New World phlebotomines in relation to feeding habits and taxonomy were studied in detail by Lewis (1975a). Following him, I use the term labrum for the labrum-epipharynx of various authors. Its length is measured from the apex of clypeus to tip of proboscis. The cibarium (= buccal cavity) lies within the clypeus and is armed with a variable number of teeth in the Lutzomyia and Brumptorrqiia females. Those in the latter genus are characteristically arranged in 4 longi- tudinal rows (Fig. 10M). Cibarial teeth are absent in both sexes of Warileya and Sycorax and are poorly developed or absent in the Lutzomyia and Bvurrp terry ia ma 1 e s . When viewed ventrally as in Fig. 12E, the horizontal teeth (= hind teeth) of most Lutzomyia females indeed look like teeth, their tips pointed, or not, depending on species. The vertical teeth (= fore teeth) usually appear as dark dots distad of the horizontal teeth at the same angle of view (Fig. 12E) . Lateral teeth may occur on both sides of cibarium near the horizontal teeth (Fig. 30M) . The number, position, shape, and size of these cibarial teeth are very important in species diagnosis and classification. -11- Two other features of the clbarium are noted in the descriptions and keys. The nhitinous arch (Fig. 12E), when complete, crosses the ventral wall of the cibarium. Its development varies according to species ana may be complete or not. The. pigment patch on the dorsum of cibarium (Fig. 12E) varies in size, shape, and degree of infuscation (= pigmenta- tion) in the Lutzomyia spp. The pharynx (Fig. 12C) , composed of 3 chitinous plates, is attached to the cibarium. Many species of Old World phlebotomines (genus Sevgentomyid) have spines on the posterior part of the pharynx but they arc relatively rare in the Lutzomyia spp., occurring mostly in some species in the eayannensis and oswatdoi groups. The majority of Lutzomyia females have transverse, unarmed ridges on the wider, posterior area of the pharynx. The length of the pharynx is given in descriptions although it is flexible and difficult to measure with accuracy (Lewis, 1967a). I adopt th.p terminology of Davis (195 7) as applied to the thorax. The mesonotum should correctly include the mesoscutum, mesoscutellum, and postnotum and should not be restricted to the. mesoscutum alone (Saether, 1971). Trie degree and distribution of pigmentation of the thorax and other body regions are important in associating sexes and for distinguishing species of Lutzomyia. Coloration has not been given much importance in systematic works but 1 find it generally dependable as a taxonomic character as applied to the New World phlebotomines . Pleural setae on the anepisterum (upper episternal setae) and katepisternum (lower episternal setae) are present in species of Lutzomyia. Brumptomyia, and Sycovax. The WariZeya spp. and the majority of Old World phlebotomines lack such setae (Abonnenc & Leger, 1975). Additional -12- 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. 2ii) . The 4 postspiracular setae in the phlebotomine species are reduced in sii'.e, barely visible (Fig. 12D) . The wing length is measured from the basal costal node (= rudiment of tegula; Snoigrass, 1935) to its apex; its width at widest part (Fig. 12G.i . Certain wing vein sections (Fig. 12G) are given useful, easily remembered names. Alpha (a) is the length of R from its junction with R to the costa. Beta (3) is the length or R from the junction of R to the junction of R„ + R, . Gamma (y) is the section of R from the r - m crossvein to the junction of R . Delia (5) is that part of R distad of the junction of R and R . It is negative when R ends before this junction . The legs are measured as in Fig. 12H. A few phlebotomine spp. have a row of short spines on the hind femur (Fig. 31F) . The length of the femur in relation to that of the tibia and/or basitarsus is sometimes used for distinguishing species of Lutzomyta. The setation of the abdominal tergites 2 and 6 is useful in the classification cf Old World phlebotomines but has been little studied for the New World species. lewis (1975b), however, discussed and figured the socket patterns for some species, noting that the patterns were of 5 types. Setae on the sides of tergite 8 may be present or not in New World females of Phlebotominae and Sycoracinae, The nature of the abdominal sternites, especially sternites 2 ^nd 3, is useful in associating male and female sand flies (Hertig & Fairchild, 1950) but the character state is variable among individuals of conspecific I ipulations (Forattini, 1954). The structures of the bilaterally symmetrical male genitalia, labelled in Fig. 2 and 12, sr-i extremely important ir systematics. After eclosion, the genitalia rotates 180° in the phlebotomine males but not in those of Sycor-ax. The style, attached to the apex of the coxite, bears i to 6 major spines* and 0 to many additional small setae. The presence of numerous strong spines is presumably a plesiomorphic feature (Tbeodor, 1965). The coxite is the dorsal basal appendage in the phlebotomine males and, of course, is the ventral basal one in the Sycovax males. It may or may not have a setal tuft or other nondeciduous setae on its inner sur- face. These setae are those which remain on the coxite after maceration and which usually differ in size or shape from other setae on the appendage. The fused, paired aedeagi, each of which is called the aedeagus is sub triangular and well sclerotized in most phlebotomine males. Some species (e.g. L. walkeri) have aedeagi with dorsal projections. The aedeagus of the Colombian Sycorax males is complex as in Fig. 2F. There is a basal, laterally flattened sperm pump within the abdomen, distal paired genital filaments and other structures mentioned on p. 31. The genital pump of the phlebotomine males is probably homologous with the sperm pump of the Sycovax spp. Leading from it are 2 genital filaments which pas.? through each aedeagus and which vary in length, width, and sclerotization according to species. The apices are simple, modified, inflated or not. *It is difficult to define major (= strong) spine, as it is a relative term referring here to size. One author's interpretation may differ from that of another. In doubtful cases, the illustrations should be studied to understand the meaning of statements in the text. -14- The phi ebo torn! ne parameres lie between the coxites and ventral lateral lobes when viewed laterally. Their shape and setation are im- portant in differentiating species. Simple parameres , i.e., I hose without arras or processes, are shown in Fig. 16F and ISA. Complex parameres are illustrated in Fig. 44H. The cercopods of the Syaorax males appear to correspond to the parameres of Phlebotominae and are very complex. The ventral, nonsegmented lateral lobes are more or less uniform in the phlebotomine males, although their length, width, and setation may offer good characters in some species. Corresponding structures are wanting in the eycoracines . The shape of the setose cerci may help associate conspecific sexes but in general these structures are of limited use in species diagnosis. The size and structure of the internal paired spermathecae and their ducts (Fig. 12F) are particularily important in the classification and identification of female sand flies. An idea of the variety of sperma- thecae can be gained by scanning the illustrations in this review. A well sclerotized, "Y" shaped genital fork or furca in the phlebotomines is associated with the spermathecae. METHODS AND MATERIALS Field Collections For sampling phlebotomine and sycoiacine populations, several methods should be used to determine species diversity and relative abundance. In Choc.o Department and elsewhere to a lesser extent, we used the following methods to capture these flies. Battery powered light traps (Sudia and Chamberlain, 1962) attract many psychodids, the numbers varying according to weather conditions, location, species composition, etc. We secured these, traps to tree branches 2 meters above ground level but did not use carbon dioxide as an adjunct attraction. At Curiche, where at least 1 trap per week was operated throughout the night (1800 hrs.-0700 hrs.), we collected a total of 23 phlebotomine spp. (3786*6*. 789oo) from April to Dec, 1967 (39 trap nights). We recorded 36 sand fly species from Curiche based on all collection techniques . Shannon traps (Shannon, 1939) made from muslin bedshaets do not trap insects per se but are collecting devices similar to those used by lepi'dopterists in :,sheeting" for moths at night. A gasoline lantern pro- vider a light source enabling collectors to readily aspirate the psychodids which land on the illuminated cloth. They are attracted to the light, to the collectors, or to a combination of both. The species composition of light and Shannon trap captures is generally similar. -15- -16- Malalse ti'aps (Townes, 196?) and flight traps (similar to the design oi Gressitt and Oressitt, 1962) were placed in forest clearings, across trails or othei flyways at ground level and in the forest canopy. Insects which land on the trap baffles eventually die after making their way upwards to the killing jar(s) charged with potassium cyanide. Ordinarily, there trapp do not capture large numbers of psychodids but they are effective in sampling the species composition of a given area if operated over extended periods of time. For example, in the Curiche Forest at groend level, one Malaise trap captured 24 Phlebotomine species (3166*.:?, 482^o) from 3 April 1967, to 30 Nov. 1967 (109 trap days"). Whenever possible, we searched diurnal resting sites for sand flies. These included tree trunks, especially dark crevices between buttresses, animal burrows, and litter on the forest floor. Tree trunks are favored resting sites for several species, specimens of which are easily captured with a simple aspirator. During the survey in Choco Dept., especially at Curiche and Teresita, we collected large: numbers of phlebotomines on human bait. The majority of collections were made by 2 men sitting on the forest floor between 1830 and 2100 hours. Phlebotomines were captured with aspirators as they attempted to feed, the results computed on a man-hour basis, i.e., the total number captured in one hour divided by the number of collectors. Specimens were preserved dry in cardboard pill boxes rather than in alcohol to prevent hardening of muscle tissue. *0ne trap day equals 24 hours. -17- Laboratory Methods For routine. identification of phlebotomines and sycoracines, un- dissected specimens are first macerated (i.e., cleared) in 10-20% NaOH. The loss of pigmentation can be reduced by heating this solution (con- taining the files) to the boiling point rather than macerating them at room temperature for 12 hours or more. The flies should be left in the hot NaOH for 5 minutes, then placed directly into a drop of 80-95% liquid phenol (C,H OH) in a depression microslide. Within 5 minutes, the important internal structures such as the spermathecae and ducts are clearly visible and should be drawn if necessary, Once identified, the flies can be discarded, preserved in vials of 70% alcohol, or processed further for slide mounting. 1 follow the procedure of Fairchild and Hertig (1948c) except that Canada balsam or Euparal is substituted cor copaJ . Other slide-mounting procedures are discussed ny Osorno et al. (1966) Quate & Steffan (1966), and Lewis (1973) and others. The choice of which to use is a matter of personal preference but in all cases the best pre- parations are made from freshly killed flies. Sometimes it is necessary to remount specimens because of undesirable position and/or shrinkage. For those embedded in Canada balsam or other xylene—soluble media, it is advisable to submerge the entire slide in liquid phenol in a suitably closed container such as a petri dish. After i to 7 days, the specimens become soft and are easily handled. Xylene as a solvent should be avoided as the specimens tend to become brittle and easily damaged. -18- 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 present ecological factors, both biotic and physical, and by knowledge of past changes in climate, vegetation, and geology. Haffer (1967, 1974) discussed the probable speciation and distribution of some neotropical birds in relation to past changes, especially those occurring in the Pleistocene and post Pleistocene periods. During times of drought, popu- lations of forest birds and other organisms were restricted to isolated forest refugia which acted as core areas of speciation. As conditions became wetter, these forests enlarged in area, often merging with others to form zones of secondary contact. The presumed forest refugia in Colombia and elsewhere were discussed and mapped by Haffer (1967, 1974) snd Brown. (1975), the latter author studying speciation in forest butter- flies in relation to these presumed refugia. Although more data are needed, the refuge theory offers a plausible explanation for understanding recent speciation in the Phlebotominae and Sycoraciiiae. The majority of New World species inhabit forests, especially those in the lowland tropics which receive 2000+ mm of rain per year. For discussion purposes, it is convenient to divide Colombia into natural regions. D'Aliesandro & Barreto (19 71) delineate 7 such areas, their large "Oriental Region" consisting of both Amazonian forest and eastern natural savannah (llanos) . Within each region there are one or more life zones, each defined by a combination of biotemperature, annual -19- ■20- prec.Ipitation, humidity, and vegetation (Holdridge, i 96 /) . The life zones in Colombia were studied by Espinal and Montenegro (1963) . Their ''bosque humedo" translates to "tropical raoist forest" (2000-4000 mm of rain per year) and "bosque muy humedo" corresponds to "tropical wet forest" (4000-8000 mm of rain per year) . The "bosque pluvial" (= "tropipal rain forest"), tne wettest lowJand life zone, receives over 8000 runi of rain per year. Figure 1 depicts where phlebotomines and sycoracines have been col- lected in Colombia. Osorno et al. (1972a) provide specific data for most localities; others from which collections were made by me or my colleagues are discussed below in relation to faunal regions (D'Allesandro ?* Barreto, 1971) and distribution centers (Haffer, 1974). Pacific Coast Region Haffer (1967) attributes the "high concentration oi endemic species in the tropical lowland forests of western Colombia and Central America" tc a gradual accumulation of isolates in the trans-Andean forest region.* Tao dominant refugium on the Colombian Pacific coast is the Choco refugium extending from Lago Calima (Valle) to north of Quibdo (Choco) and across the Rio Atrato and San Juan river systems (Brown, 1975) . An analysis of the phlebotomine species and subspecies inhabiting the Pacific coast region indicates the following. The vast majority of the 41 known taxa, probably originated in forests east cf the Andes (cis-Audean region) . Most of these reached the *The tiVms-Andean region comprises "Middle America and the narrow Pacific lowlands along the western base of the Andes" (Haffer, 1967). Cis-Andean refers to lowland forests east of the Andes. -21- Pacific coast forests by advancing around, net across, the northern Andes. These species which now occur on both sides of the mountains include: Lutzomyta ayvozai, L. guyanensis, L. bispinosa> L. gomezi., L. ncrdestina, L. sewcena, L. pilosa, L. spinosa, L. trinidadensis , L. shannonij L. tvbeYculata.1 L. trn.rarr.ulCj and others. Nine of the 41 species and subspecies do not occur in other regions of the Republic. Of these, only Lutzomyia sp. of Anchicaya is precinc- tive, the others occurring also in the trans-Andean regions of Northwestern Ecuador and/or Central America. These species include: Brumptomyia hcj-.ata, Warileya nigrosaoaulus ., L. rebur-rc.., ."'. veauvva, L. isovesperti- lionis, L. ylephiletov , L. sangu.ina.via, and possibly L. od.ax. Two forms which probably evolved in Pleistocene forest refugia within the trans- Andean region include L. cavvevai thula n.spp. and L. barrettoi rnajusouta n.spp. , both of which now occur in trans-Andean regions of Ecuador, Colombia, and Central America and east to the northern foothills of the Central Cordillera of the Colombian Andes. There is a slight possibility that some phlebotomines were able to cross the Andes in southern Zcuador and northern Peru in times past when wet forests occupied the now dry valleys. Some Amazonian birds apparently colonized Pacific coast forests in such a manner (Haffer, 1967) . Phlebotomines, however, are rather weak fliers and it would have been necessary for lowland forms to survive at elevations above 2000 meters above sea level. The one possible example of successful colonization may be that of L. vehuri'a — a species whose closest allies occur entirely in the (•?•?'&— Andean region, especially in Amazonian forests. A few species such as L. vespertitionis and its sister species, L. isovespertitionis probably invaded the Pacific coast region of Colombia from Central America, the former species also occurring east of the Andes, at least as far south as Bolivar Department. Depart mant of Chocc Three collecting sites within this Department were described by Eidridge & Fairchild (1973) and Eldridge et a] . (19 73) . Maps accompany their descriptions. Cuptche. On narrow coastal plain between western slopes of Serrania del Baudo and Humboldt Bay on the Pacific coast; "tropical wet forest;'' elevation* less than 10 m. Sand flies were collected from April 1967, to :tec. 1967, in disturbed and undisturbed evergreen forests end near a mangrove swamp . Alto Cupi-dhe. About 3 km inland from Curiche on western ridge of Serrania del Baudo; "tropical wet forest;" elevation 302 m. The sand fly fauna is similar to that of nearby Curiche. Teresita. At eastern foothills of Serrania de Baudo, near Rio Truando; transitional life zone, "tropical moist/tropical wet forest"; elevation 35 m. Collections made in semi-disturbed forests on flat or gently rolling terrain from March-Dec. 1967. Department of Valle Ancth.iccvja Dam. About 35 km inland from Pacific Ocean, on the Rio Anchicaya; "tropical wet forest;" elevation ca . 560 m. This site is similar to that near the Rio Anori (Antioquia Dept.) in terrain, elevation, ^Elevation is expressed here and elsewhere in meters above sea level -23- and climate. Sycoracines were recorded from both localities and from one other in Cauca Department. 25 km E of Buenaventura. "Tropical rain forest" according to Espinal (1968); elevation ca. 50 m. Phlebotomines were captured in a patch of disturbed forest surrounded by cultivated fields. Atlantic Coast Region This area encompasses not only the Carribean lowlands but also the northern foothills of the western and central Cordilleras of the Andes. The presumed Nechi refugiura is located here, a center "heavily influenced by the Chor.o center to the west" (Haffer, 1967) . The present life zones are more diverse here than in the Pacific coast forests and except for the area near the Pvio Anori, the phlebotomine fauna has been little studied. Intensive collections of phlebotomines made by Dr. C.H. Porter and his colleagues near the Rio Anori (19 70-1971) yielded 37 species and sub- species. Thirty of these also occur in Pacific coast forests, a fact not surprising based on the distribution of other organisms (Haffer, 1967, 1974). Lutzorrvjia. stvictivitta may be endemic. Lutzomyia yuil-ti and L. noeticola are eis-Ardean taxa. The former reaches its northernmost limits in northern Colombia; whereas L. noeticola invaded extreme western Panama but populations between there and northern Colombia have since disappeared. Department of Antioquia Rio Anori (7° 19'N; 75° 04'W), about 24 km SW of Zaragoza; "tropical wet forest;" elevations from 500-700 m. Located in the foothills of the centra] Cordillera of the Andes, this site is mostly well forested with steep, well drained slopes. A detailed description is given by Porter & Defoliart (MS) . Pep artmeat of Hagdalena Three sites near Santa Marta yielded sand flies from tree trunks or under loose bark of standing trees in cultivated or semicleared areas (August, 1973). Rio Don Diego, E of Santa Marta within 1 km of Carribean Sea; "tropi- cal wet forest;" elevation ca . 3m. Fetuccn Santa McOfta and Mvnco.-, near a small stream; probably a transition zone between "tropical wet forest" and "tropical dry forest;" elevation ca. 80 m. Minca, SE of Santa Marta; life zone indeterminate but similar to preceding site; elevation 200 m. Andean Region Nearlv all records of phlebotoraine species occurring above 1000 m are given by Osorno et al. (1972a). Based on these and on my limited collec- tions from the Western Cordillera, it is apparent that the Lutzornyia species ill the Vewusaxtm and vexatov groups dominate the rather depauperate phiebotomine fauna. De partroent of Vail e Three sites W or SW of Cali — 1. Near Piohinde , 2. In forest at the television tower just E of Salidito, and 3. Near Rio Ranee, SW of Cali; -25- "lower montane very wet forest;" elevations from 3 570-1700 m. Limited collections from 31 July, 1973, to August, 1973. Department; ot Cauca iTasterr slope, near summit of Cerro Munchique, west of Popoyan; "lower montane rain forest;" elevation ca. 2500 in. Briefly surveyed on 5-6 August, 1973. One Si/eorax sp . taken but nc phlebotomines. Oriental Region Occupying a huge area east of the Andes, the Amazonian forest in Colombia is apparently rich in phlebotomine species but few collections have been made there or in the more northern Llanos which forms part of the Oriental Region (Haffer, 1967, fig. 1). Many of the species known from the Amazonian forests in Brazil, Peru, and Ecuador are expected to occur also in Colombia. C oreisana of Amazonas About 17 km W of Leticia; "tropical moist forest;" elevation 84 m. PhlebotoTtine flies were relatively rare in late July, 1973, the bulk of the material being taken in semi-disturbed forests near the Amazon River on tree trunks., less commonly in light traps. July and August are the two driest months at this locality. [n c eudcncia of Caquota Tres Esquinaj; transitional life zone, "tropical moist forest"/ 'tropical wet forest;" elevation ca. ?00 m. Using light traps, Dr. C.J larinkelle collected numerous phleboto.mines in November, .1971. Figure 1 Map of Colombia showing localities where phlebotomises and sycoracine flies have been collected (records based on Osorno et al., 1972a and this review)- . Shaded sections represent areas 1000 m+ above sea level, Legend » = Fhlebotomine record X = Sycoracine record 4 - Fhlebotomine and sycoracine record 1 = Atlantico Dept. 2 = Magdalena Dept. 3 = Norte de Santander Dept. 4 = Caldas Dept. b = Risaralda Dep1". 6 - Quindio Dept. 7 - Cundinaniarcs Dept. -12. V M V Choco ^f't^soM.K',, 3oyac< S ->-. "; v_.j_._i _.<_.-.._ t Lu O j-o.i_c.i_ uj.u meters. Antenna (including scape and pedicel) 1.61 long; flagellomere I (0.26 long), about 1.8xlength of flagellomere II; paired ascoids (Fig. 2C) , subequal in size, on f lagellomeres I-IV only; other antennal setae mostly deciduous, not arranged in a whorl-like pattern. Palpal length 0.17; ratio of segments: .1-1.02-0.72-0.67; second segment with 20+ strap- like sensory rods in a distinct patch. Pleura with 19-22 setae below wing base, without episternal setae. Wing venation as figured. Length of femora, tibiae, and basitarsi: Foreleg, 0.63, 0.73, 0.49; midleg, 0.63, 0.75, 0.47; hindieg 0.73, 0.82, 0.44. Visible sternites 2 and 3 with paired circular openings laterally. Genitalia complex: Coxite 0.31 long. Style 0.18 long wich 1 terminal spine, slightly beveled at tip and with a proximal row of 3 smaller spines. Paramere as shown with several setose lobes and projections, the most distal finger-like with rounded, upturned tips. Aede.agal complex of a laterally compressed sperm pump (0.20 long) within the abdomen; a median distal process (0.165 long) compressed dorso- ventrally, curved and slender in lateral view, upturned at tip, basal -36- two-thirds much wider than distal one-third when viewed dorsally; paired. sinuos, relatively long genital filaments. Cerci as figured. Material examirted: Colombia, c? holotype (no. 537), Cerro Munchique (Cauca) , 2450 m above sea level, light trap in cloud forest, 6 Aug. 1973, D.G.Y. and R.C.W. coll. c? paratype (no. 538), same data except collected by R.C.W., 8 Aug. 1975. Discussion: S. andicola, the largest Syoorax known from Colombia, closely resembles S. tvispinosa n. sp. in details of the male genitalia, setation of the antenna and in other character states. The two species, apparently allopatric, are separated by the characters given in the key. 2. Sycorax colombiensis n. sp, (Fig. 3) Male (holotype): Wing length 1.3 2; width 0.47. Head height 0.21; width 0.25; eyes separated by 0.126 mm or by distance = to 10 facet diameters. Flagellomere I (0.19 mm long), about 2.2xlength of flagello- mere II. Paired ascoids visible only on f lagellomeres I and II; other sensory hairs in a whorl-like pattern on f lagellomeres I-X, the remaining f lagellomeres missing. Palpal length 0.10, ratio of segments 1-0.91-0.80- 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. Parainere as figured, the lower distal projection blade-like, pointed at tip. Aedeagal complex: Sperm pump (0.10 long); median distal process (0.09 long), slender and subequal in width in dorsal view, with a slender, rounded tip in lateral view; a subtriangular plate, dorsoventrally flattened, with acute bifurcate tip, pointing to the rear, present below the base of the median process of aedeagus. Cerci as shown. Female (allotype): Wing length 1.37; width 0.59. Head height, 0.24; width, 0.29; eyes separated by 0.15 or by distance = to 10 facet diameters, Antenna (1.15 long), flagellomere I (0.22 long), about 2.4xlength of flagellomere II; paired ascoids on flagellomeres I- IV; other sensory hair- arranged in a whorl-like pattern on flagellomeres I-XV. Palpal length, 0.113; ratio of segments 1-1-0.75-0.58. Pleura with 16-18 setae below wing base and 0 or 1 upper episternal seta. Wing venation as shown. legs partly or wholly missing in all specimens, length of femora, tibiae, and basitarsi of foreleg: 0.43, 0.51, 0.27. Abdominal sternite 2 with paired circular openings at sides; other sternites apparently lacking these openings. Spernatheca and sperm duct length at least 0.15 from base to tip of button; furca spade-shaped at end. Material examined: Colombia, o* holotype (no. 529), Anchicaya Dam (Valle) , elev. ca. 400 m above sea level, light trap, 28 Jan. 1975, R.C.W. Q allotype (no. 530), same data as holotype except collected 10 June 1975. Paratypes (nos. 531-538), all from type locality in light traps; 1 c., 11 Aug. 1973, D.G.Y. and R.C.W. 1 -■ u ^V v^ -*/< V* / Figure 4 SycOTPax fatvohi'ldi — A. Male head, B. Female wing, C. Male wing, D, Female head, E. Male genitalia, dorsal view, F. Sperm pump, genital filaments, and median dorsal process of aedeagus, lateral view, G. Spermathecae, H. Female pedicel and flageilomeres I and II showing paired ascoids and other setae. Male: Anchicaya Dam, Valle Dept., Colombia Female: Same locality as male X'.: ^:Pmj j >' s^~ i I J H Ml •" ^ F ^^; c_^ Figure 5 Sycovax trispinosa — A. Male head, B. Female palpus, C. Female, terminal 4 f lagellomeres , D. Female pedicel and f lagellomeres I end TI showing ascoids, E. Female head, F. Female wing, G. Male wing, H. Male genitalia, dorsal view, I. Sperm pump, genital filaments, and median dorsal process of aedeagus, lateral view, J. Tip of furca, dorsal viev.'j K- Spenhathecae. Male: Anchicaya Dam, Valle Dept., Colombia Female: Same locality as male -49- / H . s r; 1 \ .1 B g ill j * 9 £> / If ml SUBFAMILY PHLEBOTOMINAE Previous studies on this subfamily in Colombia were reviewed by Osorno et al. (1972a) and will not be repeated here except for litez-ature citations in the species accounts. Nearly all of these studies involve systematica, especially species descriptions and distribution records. Owing to the paucity of bionomic studies on the Colombian phlebotomines , I felt that it would be desirable to provide references to such studies carried out in other countries. The results of these studies may not always apply to conspecific populations in Colombia. Information on the biting habits of some species in Choco Dept. is provided in the species accounts . With few exceptions I have not redescribed known taxa because of adequate original or subsequent descriptions. Hopefully, the figures and keys will serve to distinguish the taxa. Key to the New World Genera 1. Wing broad, rounded at tip; R„ + R„ + R, forks before, on same level as, or slightly beyond r-ni crossvein. Pleura without episternal setae. Female cibarium unarmed. Male genitalia with style longer than coxite Wcwileya (Fig. 6-7) Wing pointed at tip; R 4- R + R forks well beyond r-m crossvein. Pleura with episternal setae. Female cibarium armed with teeth. Male genitalia with style shorter than coxite 2 •50- -51- 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 Brumptonrjia (Fig. 8-11) Interocular suture incomplete. Female with 1 row of horizontal teeth; vertical and lateral teeth present or not. Male genitalia with style bearing 1-6 large spines, basal pair not borne on a common tubercle in those species having 5 spines . . .Lutzornyia (Fig. 12-89) -52- Genus Warn, ley a Hertig, 1948 This genus, recently enlarged to accomodate W. (H.) hertigi (Fchld.) by Lewis et al. (1978), contains 5 species, 2 of which in the subgenus Waviteya occur in Colombia. Little is known about the habits of any of the species although W. (W.) rotundipennis , one of the Colombian species, and W. (W.) phlebotomanica Hertig, the type species from Peru, are anthropophilic . Keys to Species Males 1. Palpal segment 5 longer than combined length of segments 2 + 3. Paired intraabdominal rods present. Genital filaments about 1.8X length of pump. Style lacking a basal bristle. Wing length less than 1.5 mm nigrosaccuhMS (Fig. 6) Palpal segment 5 shorter than combined length of segments 2+3. Intraabdominal rods absent. Genital filaments less than 1 . 3X length of pump. Style with a basal bristle. Wing length over 1.8 mm . . . rotundipennis (Fig. 7) Females 1. Spermathecae large and sac-like with smooth walls. Palpal segments 5 longer than combined length of segments 2+3. Wing length less than 2.0 mm nigrosaoculus (Fig. 6) Spermathecae cylindrical, somewhat worm-like, with transverse stria- tions. Palpal segment 5 shorter than combined length of segments 2 + 3. Wing length over 2.0 mm rotundipervnis (Fig. 7) -53- 5. Warileya (W. ) nigrosacaulus (Fig. 6) Warileya nigrosacaulus Fairchild & Hertig, 1951b: 428 (o. holotype, Cerro Campana, Panama Prov., Panama). Fairchild, 1953: 102 (cf. to Her tig ia hertig i) . Barretto, 1955a: 183 (listed). Forattini, 1971a: 107 (listed). Tesh et al., 1971a: 153 (blood meals, Panama). Martins & Morales, 1972: 366 (listed). Young & Chaniotis, 1972: 97 (cf, descr.). Chaniotis et al., 1972: 95-96 (in tree hollows, Panama). Christensen, 1972a: 88 (listed). Forattini, 1973: 536-533 (o, fig. ). Velasco & Trapido, 1974: (cf. to W. yungasi) . Lewis, 1975a: 500 et seq. (mouth- part morphol.). Lewis et al., 1978 (classif.). Distribution: Colombia (Choco, Valle) , Panama. Material examined: Colombia: 1 o", 1 o, Curiche (Choco), Malaise trap, 27 April 1977. 1 holotype, Cerro Campana (Panama Prov.), hollow log, 7 Jan. 1947, M. Hertig. 2 6cf, 5 op, near Gamboa, Canal zone, tree hollow, 10 Sept. 1969, B. Chaniotis. 4 6V, same data but taken 15 Jan. 1970. Discussion: A little known species, W. nigrosacaulus is easily distinguished from W. rotundipennis by the characters given in the key. Using a precipitin test for blood meal determinations, Tesh et al. (1971a) found that blood from 4 recently engorged nigrosacaulus females reacted with mammalian and reptile-amphibian antisera. Four other blood meals were nonreactive due to small volumes of blood and/or to the weakness of the reptile-amphibian antisera. 6. Warileya (W.) rotundipennis (Fig- 7) Warileya votundipennis Fairchild & Hertig, 1951b: 424 (c? hole type, o, Cerro Campana, Panama Prov. , Panama). Fairchild, 1953: 102 (cf. to Eevtigia hertigi) . Barretto, 1955a: 188 (listed). Fairchild & Hertig, 1959: 122 (Cosira Rica). Johnson & Hertig, 1961: 765, 775 (rearing). Hanson, 1968: 93 (larval fig.). Forattini, 1971a: 107 (listed). Young & Chaniotis, 1972: 366 (listed). Christensen, 1972a: 88 (listed). Forattini, 1973: 536-538 (cf, o. fig.). Velasco & Trapido, 1974: 436 (cf. to W. yungasi) . lewis, 1975a: 500 et seq. (mouthpart morphol.). Lewis, 1975b: 366 (hair sockets, fig.). Lewis et al. : 1978 (classlf.). Distribution: .Colombia (Antioquia, Choco, Valle), Costa Rica, Panama. Material examined: Colombia. 2 no, Teresita (Choco), tree buttress, 15 June 1967. 27 oo. Rio Anori (Antioquia), light traps, Sept. 1970. 5 w, same data but 29 May 1970, C.H.P. 8 6V, 2 no, Anchicaya Dam (Valle), light trap, 28 Jan. 1975, J.E. Browne. 6 def, 6 oo, same data but taken 10 June 1975, R.C.W. 5 <5b", 4 ot) , same data but taken 16 July 1975. Panama. 6* holotype (no. 2335), o allotype (no. 2312), Cerro Campana (Panama), Shannon trap, 24 Aug. 1950, M. Hertig & G. Fairchild. 2 c?6*, Rio Changena (Bocas del Toro) , Shannon trap. 8-11 Sept. 1961, R. Hartmann 6 P. Galindo. Costa Rica. 2 &<$, 1 o, Turrialba, Shannon trap, 20 Aug. 1961, G. Fairchild & M. Hertig. Discussion: Warileya rotundipennis , the only Warileya species lacking intra-abdominal rods (Lewis et al . , 1978), has been reported feeding on man in Panama (Fairchild & Hertig, 1951b) but its role, if any, in disease transmission is not known. Ur. C.H. Porter also has collected man-biting -55- feraales at the Rio Anori (Antioquia) locality, mostly in forest clearings (pers. comm. ) . The records of W. rotundipennis and W. nigvosaeovXus from Choco and Valle Depts. indicate that both taxa probably occur in other areas of the trans-Andean region perhaps as far south as Guayas Prov. , Ecuador. Figure 6 Wariloya. nigvosaocutus — A. Hale head, B. Male flageilomere II, C. Female head, D. Female flageilomere II, E. Female cibarium and pharynx, F. Male genitalia, dorsal, G. Tips of genital filaments, H. Female wing, I. Male v.dng, J. Spermathecae. Male: Curiche, Choco Dept., Colombia Female: Same locality as male -57- fife,, CM) Steffi V <3 A Figure 7 Warileyo. vctund.ipenn-Ls — A. Male head, B. Male f lagellomere II, C. Female head, D. Female f lagellomere II, E. Female cibarium and pharynx, F. Male genitalia, G. Female wing, H. Male wing, I. Sperma- checaa. Male: Cerro Campana, Panama Prov. , Panama Female: Rio Anori, Antioquia Dept., Colombia f / f ^ i l°% \\\ r ( / " ■ ■ ■ \ \i/ I l(l)/l Hi/A r7 V v -60- Genus Br-urnptornijia Franca and Parrot, 1921 Characterized by Tkecdor (1965), Lewis et al. (1978), and others, the genus Bvu^tomyi-a now contains over 21 species, 5 of which occur in Colombia. Except for the female of B. hatnata which has very short sperm ducts (Fig. 101) , the females of the other species in Colombia are virtually impossible to identify in the absence of males. Brwnptomyia avelXarn.-, one of the Colombian species, was reported feeding on armadillos in Brazil (Mangabeira, J 942b). This and other Brumptomyia spp. often rest in burrows dug by these mammals. They do not bite man and the preferred hosts of most species remain unknown. Key to Species Males 1. Genital filaments less than 4X length of pump . . . . hamata (Fig. 10) Genital filaments greater than 4X length of pump 2 2. Coxite tuft of mostly stout, spine-like setae 3 Coxite tuft of slender, hair-like setae 4 3. Coxite tuft of 20+ setae implanted on a distinct tubercle. Paramercs more slender as shown gdlindoi (Fig. 9) Coxite tuft of fewer than 20 setae, usually about 12, not implanted on a tubercle. Paramere broader as shown avell-ari- (Fig. 8) 4. Style with isolated spine well distad of proximal pair. Coxite tuft implanted on a distinct raised tubercle beaupertuyi (Fig. 8) -61- 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 (Fig." 8) Fhlebotomus avellari Costa Lima, 1932: 48 ((?, Lassance, Minas Gerais, Brazil). Mangabeita, 1942b: 225 et seq. (iir.matures , adults, descr., fig.). Barretto, 1947: 187-188 (full refs.). Fairchild & Hertig, 1947a: 615-616 (cf. to galindoi) . Barretto, 1951: 212 (dist.) . Floch & Abonnenc, 1952: 39, 45 (6*, o, keyed). Rodriguez, 1953b: 55 (mention). Forattini, 1954: 214-217 (second sternice, fig.). Forattini & dos Santos, 1955: 17 (Brazilian record). Ortiz, 1963: 320 (o, keyed). Brumptomyia avellari: Barretto, 1955a: 187 (listed). Martins et al., 1961b: 309 (mention). Martins et al., 1962a: 380 (Goias, Brazil). Sherlock, 1962: 332, 335 (mention). Carnheiro & Sherlock, 1964: 315 (pupa, keyed, fig.). Fraiha et al . , 1970a: 468 (6* keyed) . Christensen, 1972a: 88 (Panama). Osorno et al., 1972a: 14 (Boyaca, Colombia). Forattini, 1973: 122 et seq. (gen. review, fig.). Llanos et al., 1976: 480 (Peru). Martin et al . , 1976b: 496 (Peru). Ramirez et al . , 1976: 599 (Venezuela) . Distribution: Panama, Colombia (Boyaca), Peru, Venezuela, Brazil, Paraguay . Material examined: Colombia. 2 6b*, 1 o, Puerto Boyaca (Boyaca). light trap, 6 May 1973, C.J.M. Brazil. 1 6\ Coqueiros (Sao Paulo), armadillo burrow, Oct. 1953, Albertin. Panama. 1 o", Tocumen Airport (Panama), light trap, 23 March 1953, F. Blanton. Paraguay. 5 do\ Aca- Poi, San Pedro, burrow, 18 April 1950, M. Hertig. -62- Discussion: Brumptomyia avellari and E. brurnpti (Larousse) with sympatric populations in Brazil and Paraguay, closely resemble one another, the males differing chiefly by the shape of the parameres. I regard the Colombian specimens as being conspecific with the former species. 8. Brumptomyia beaupertuui (Fig. 8) Fhlebotomus beaupertuyi Ortiz, 1954: 235 (o* holotype, Duaca, Lara State, Venezuela). Scorza & Ortiz, 1960: 434 et seq. (ecology). Pifano et al., 1962: 383, 337 (o*, q, keyed), 411-412 (cf, o, descr., fig.). Ortiz, 1963: 322 (^ descr.). Ortiz & Scorza, 1963: 350 (listed), 354 {&> keyed) . Ortiz, 1965a: 208 (mention). Scorza et al., 1967: 193, 195 (d\ o keyed) . Leon, 1969: 30 (listed). Phlobotoinus galindoi (not galindoi Fairchild h Hertig, 1947) : Pifano & Ortiz, 1952: 138 (listed, Venezuela). Leon, 1969: 30 (listed). Brumptomyia beo-upertuyi: Barretto, 1955a: 187 (listed). Sherlock, 1962: 321 et seq. (6*. fig., Colombia). Fraiha et al., 1970a: 468 (6*, keyed). Forattini, 1971a: 98 (listed). Forattini, 1973: 522 et seq. (.gen. review, fig.). Distribution: Colorrbia (Santander), Venezuela. Material examined: Venezuela. 2 6$ (topotypes), Duaca (Lara), no date, T. Ortiz. Discussion- At present, this little-known species is represented in Colombia by a single male, skillfully illustrated by Dr. 0. Mangabeira (in Sherlock, 1962). Ortiz (1963) described the female based on Venezuelan specimens . •63- 9. Brumptomyia galindoi (Fig". 9) Phlebotomus galindoi Fairchild & Hertig, 1947a: 615 (cfholotype, near Roquete, Chiriqui Prov., Panama). Floch & Abonnenc, 1952: 40 (6* keyed). Ortiz, 1954: 238 (cf. to beaicpertuyi) . Rosabal, 1954: 30 et seq. (aesi is inserted at 0.59-0.61 of the segment whereas that of leopoldoi is located at 0.53. In addition to the leopoldoi specimens listed above, I examined 4 guimavciPsi males from southern Brazil: 2 <3a, Faz. Ribeirao de Baizo, Patos, Minas Gerais ; 1 ! C r D / cos ^- H /' V /* 8 . /> I v-" \ \ *& c ^ ^ Ml ^ Figure 11 Bx'imptomyia leopotdoi male — A. Head, B. Flagellomere II, C. Coxite tuft, paramere and aedeagus of male from near Santo Domingo de Los Colorados, Ecuador, D. Wing, E. Genital pump and filaments, F. Genitalia, G. Genital pump of male from Fig. 11C locality. MaJe: Rio Anori, Antioquia Dept-S Colombia (except for Figs, 11C and G) Bmenptomyia guimaraesi male — H. Genital pump, I. Coxite tuft, paramere and aedeagus, same scale as 11C. Male: Itaporanga, Sao Paulo State, Brazil •76- Germs _£ u tzorrcj ia Franga, 3 924 As mentioned earlier the classification of the Lutzomyia sand flies is based upon the scheme, of Lewis et al. (1978). It should be emphasized that this represents a flexible classification, subject to change as new species, new character states, or unknown sexes are discovered. It is hoped that the following keys and discussions will be helpful to those interested in sand flies and disease in Colombia and elsewhere in the neotrooics. Keys to Subgenera, Species Groups, and Un g ro upejd Species of Lutzonrjia In order to interpret some of the character states in these keys, the user is urged to closely examine the illustrations corresponding to the descriptive statements. Such relative terms as "strong," "small," "inflated," etc. are understood more easily with figures. This also holds true for the male and female genitalia, both of which may be complex and therefore difficult to describe by words alone. To a certain extent, the Lutzornyia subgenera and equivalent species groups are defined by the characters given here. It is important to note, however, that species in some groups, presently unknox^n in Colombia, may not exactly "fit" into a subgenus or species group as characterized by these keys. For example, L. nevesi (Damasceno and Arouck) belongs in the verpueavwri group although the male, unlike the others, lacks a coxite setal tuft. This species is very common at Limoncocha, Napo Prov. , Ecuador — a locality not far from Putumayo Intendencia, Colombia. As additional species are discovered in the Republic, these keys obviously will have to be revised. For the present, they are intended to be used with caution, applicable only to the Colombian fauna. Males 1. Palp 5 very short, less than half length of third and less than twice length of fourth subgenus Psyahodopygus (Fig. 61-71). . 2 Palp 5 subequal to or longer than half length of third and greater than twice length of fourth 4 2. (1) Style with 1 large terminal spine and 3 inconspicuous subapical setae. Coxite with dorsal margin indented near middle series sqi'.amiventr-is (Fig. 61) Style with 2-6 major spines. Coxite not indented near middle ... 3 3. (2) Style with 2 major spines, both terminal. Paramere simple. . . series ar-ihirei, L. bisp-inosa (Fig. 63) Style with 3-5 (sometimes 6) major spines. Paramere simple or complex series panamensis (Fig. 64-71) 4. (1) Lateral lobes markedly inflated . . vsspertilionis group (Fig. 29) Lateral lobes not inflated. .... 5 5. (4) Lateral lobes with spatulate setae at tips . . . subgenus Evandromyia, series infraspincsa* "Species of Evandvorrvji.a in this series have not been found as yet in Colombia but it is probable that at least one species occurs in the southeastern part of the Republic. The subgenus was reviewed by Young and Arias (19 7 7) . •78- Lateral lobes without spatulate setae 6. (5) Style deeply forked. Parameres with 2-4 modified apical setae as shown subgenus Viannamyia. (Fig. 35-35) Style simple, not forked. Parameres usually with simple setae but, if modified, then different from above 7 7. (6) Style with 2 major spines and 1 or 2 smaller accessory seta . . 8 Style with 3-6 major spines, smaller accessory setae present or not. 9 8. (7) Paramere with a dorsal setiferous arm. Coxite without nondeciduous setae •••.•J*.!., subgenus DoiTiV^oiwia L, vosciliciZi (Fig. 30) Paramere simple, without a dorsal arm. Coxite with a basal tuft or group of setae vevpv.cavwn group, series sewana (Fig. 22) 9. (7) Style with 3 major spines 10 Style with 4-6 major spines 11 10. (9) Coxite with a basal tuft of simple and modified setae. Style with a small subterminal and a small median seta in addition to the major spines subgenus Pressatia (Fig. 32-34) Coxite without a basal tuft but with a distal group of 154- simple setae. Style with only a small median seta and 3 major spines, no subterminal seta pilosa group, L. pilosa (Fig. 86) 11. (9) Style with 4 major spines 12 Style with 5, sometimes 6, major spines 32 *Males of L. vovotaensis usually have 5 major spines but those examined from ValJe Dept., have but 4 (see p. 421). -79- L2. (11) Antenna! ascoids with very long pointed or short bluut basal spurs. Style without a subterminal seta 13 Antermal ascoids with or without basal spurs, but if present they are very short and pointed. Style with or without a subterminal seta 14 13. (12) Coxite with 5-8 strong distal setae. Style with isolated basal spine clreisbacki group (Fig. 50-51) Coxite without distal setae. Style with basal spines more or less paired shannoni group (Fig. 37-42) 14. (12) Palp 5 shorter than palp 3. Style without a subterminal seta 15 Palp 5 as long as or longer than palp 3. Style with or without a subterminal seta 18 15. (14) Coxite with 2 or more basal and/or median setae 16 Coxite without nondeciduous setae 17 16. (15) Style with basal spine isolated. Antennal ascoids with very short pointed basal spurs. Flagellomere I shorter than f lagellomeres II + III subgenus Tviohophovorryia (Fig. 51-54) Style with paired basal spines. Antennal ascoids simple, no basal spurs visible. Flagellomere I longer than II + III . .subgenus NyssomyCa (in part), L. cmtunesi (Fig. 55) -80- 17. (15) Antennal ascoids with short, pointed basal suprs. Paramere with ? dorsobasal hump. Flagellomere I shorter than II + III avagaoi group (Fig. 45-49) Antennal ascoids simple. Paramere lacking a dorsobasal hump. Flagel- lomere I longer than II + III subgenus Nyssomyia (in part), (Fig. 55-60) 18. (14) Ccxite with 3+ basal and/or median setae in a tuft or group, d-istal setae may also be present 19 Coxite without a basal and/or median group of setae but long ventral ?et_ci£ llldy Dtr jJj.toCn>_ «.o 19. (18) Style with 4 short stubby spines, 2 of which are terminal; sub- terminal seta present. Paramere simple, undivided. Coxite with 4-8 strong distal setae baityi group, L. haityi (Fig. 34) Style with longer spines; if 2 are terminal then parameres are divided; subterminal seta present or absent. Coxite without distal nondeci.dtious setae 20 20. Hind femur with a row of 3-5 short spines subgenus Fintomyia, L. spinosa (Fig. 31) Hind femur without spines 21 21. (20) Paramere with 2 dorsobasal setae, much longer than others, hooked or fan-shaped subgenus Lutzomyia, series longipalpis (Fig. 13-15) Paramere without such setae 22 22. (21) Style with a subterminal seta 23 Style without a subterminal seta 26 23. (22) Genital filament tips enlarged, spoon-shaped, each with a dis- tinct inner "tooth" . . . migonei group, series walkevi (Fig. 19-20) Genital filament tips enlarged or not but lacking an inner tooth. 24 24. (23) Coxite with setae of basal tuft shorter than width of coxite migonei group, L. migonei (Fig. IS) Coxite with setae of basal tuft longer than width of coxite ... 25 25. Mesonotum & pleura pale . . saulensis group,* L. saulensis (Fig. 21) Mesouotum distinctly darker than pleura .... verrucarum group, series verrucar-ion (in part), (Fig. 23-24) 26. (22) Coxite with basal setae inserted on a raspberry-like, subcircular base subgenus Lutzomyia, series oruaiata (Fig. 16-17) Coxite with basal setae, if present, not implanted on suchabase. 27 27. (26) Style with paired basal spines. Coxite with basal tuft of slender, subequal setae . ver- rucarum group, series verrucavum (in part), L. nuneztovaTi (Fig. 23) Style with basal spine isolated. Coxite with either basal-median setae, 1 at least larger than others, or with a patch of slender distal sotae cayennencis group, series atvoclavata (Fig. 81-82) -VT am unable to separate the males on the group level using structural characters . 28. (18) Style with a small sub terminal seta 29 Style without a small sub terminal seta 31 29. (28) Paramere divided. Coxite with long ventral setae loagi spina group (Fig. 43-44) Paramere simple, undivided. Coxite without long ventral setae. . 30 30. (29) Palp 5 shorter than segments 3+4. Eyes large. Paramere slender, its width less than that of style ungrouped, L. sp . of Anchicaya (Fig. 89) Palp 5 longer than segments 3+4. Eyes small. Paramere broad, its width greater than that of style, ungrouped, L. ronyeliana (Fig. 88) 31. (28) Style with isolated, basal spine. Flagellomere I longer than head height; ascoids with very short basal spurs. Palp 5 shorter than segments 3 + 4 ungrouped, L. -aovdestina (Fig. 87) Style with paired basal spines. Flagellomere I shorter than head height; ascoids without visible basal spurs. Palp 5 longer than seg- ments 3 + 4 . . • cayennensis group, series ccrjennsnsi-s (Fig. 79-80) 32. (11) Style with a small subterminal seta oswaldoi group (in part) , L. pia (Fig. 83) Style without a small subterminal seta 33 33. (32) Coxite with 2 or more setae at base; others may extend to middle of structure forming a loose or compact tuft vexatov group, series per-uens'is (Fig. 72-78) Coxite without setae at base but with a few scattered setae near middle of coxite oswa.Zdoi group (in part), (Fig. 84-85) Females 1. Spermathecae associated with paired sclerotized stiuctures, intracel- lular ducts (= "hairs") covering most of spermathecae subgenus Viannar.tyia (Fig. 35-36) Spermathecae without associated sclerotized structures, intracellular ducts confined to small area, usually but not always, to terminal knob 2 2. (J) Pharynx with posterior spines 3 Pharynx without spines 6 3. (2) Cibariuir. with inner pair of horizontal teeth pointing inwards OSWdldoi group (in part), L. trinidadsnsis (Fig. 85) Cibarium with horizontal teeth pointing toward pharynx, not slanted 4 4. (3) Cibarium with 10+ horizontal teeth in a comb-like row . . cayen- nsis group, series cayennensis (in part),/,, cayennensis (Fig. 79) Cibarium with 4 horizontal teeth not in a comb-like row 5 5. (4) Foreleg with femur longer than tibia. Pharynx markedly enlarged with prominent, subequally spaced transverse ridges and conspicuous spines. Pleura pale cayennensic group, series atroolavata (Fig. 81-82) Foreleg with femur shorter than tibia. Pharynx nore slender, the transverse ridges and spines less defined. Pleura dark oayettnensi-s group, series oayennensis (in part), L. micvopyga (Fig. 80) 6. (2) Spermathecae with bubble-like expansions 7 Spermathecae without bubble-like expansions 8 7. (6) Cibariuni with 4 broad and blunt horizontal teeth and conspicuous lateral teeth. Mesonotum well pigmented, darker than pleura. Eyes very small subgenus Darrtpfomyiaf L. vosdbali (Fig. 30) Cibarium with 4 slender, sharp horizontal teeth, without obvious lateral teeth. Mesonotum and pleura pale. Eyes larger . . . . saulensis group, I. sav.lensis (Fig. 21) 8. (6) Antennal ascoids with long pointed or short blunt basal spurs . 9 Antennal ascoids simple or with short pointed basal spurs 10 9. (8) Cibarium with 10+ horizontal teeth. Spermathecae distinctly aimulated dreisbadki group (Fig. 50-51) Cibarium with 4-8 horizontal teeth. Spermathecae annulated or not . . . . shannoni- group (Fig. 37-42) 10. (8) Hind femur with row of 3-5 short spines .... .subgenus Pintomyia, L. spinosa (Fig. 31) Hind femur without spines 11 11. (10) Cibarium with 6 or more horizontal teeth 12 -85- Cibarium with 4 horizontal teeth 17 12. (13) Individual sperm ducts markedly convoluted, twisted . , . . . .avagaoi group, series, bras Hi ens is, L. runoides (Fig. 49) Individual sperm ducts not convoluted as above 13 13. (12) Palp 5 equal to or shorter than segments 3 + 4 14 Palp 5 longer than segments 3 + 4 16 14. (13) Spermathecae larger, sac-like, without annuli. Ascoids with short pointed basal spurs, avagaoi group, series aragaoi (Fig. 45-48) Spermathecae smaller, with incomplete or complete annuli. Ascoids with or without short basal spurs 15 15. (14) Flagellomere I longer than f lagellomeres II + III. Ascoids with short basal spurs subgenus THohophoromyia (Fig. 51-54) Flagellomere I shorter than II + III. Ascoids without visible basal spurs subgenus Nyssomyia (Fig. 55-60) 16. (13) Cibarium with 6 horizontal teeth. Spermatheca with terminal segment smaller than preceding segment. Common sperm duct longer than individual ducts . .OSWaldoi group (in part), L. pia* (Fig. 83) Cibarium with 8+ horizontal teeth. Spermatheca with terminal segment as large as others. Common sperm duct much shorter than individual kL. via is keyed twice because of intraspecif ic variation in the number of horizontal teeth (see p. 419). -86- ducts subge- nus Lutsomyia, series long ip alp is (in part), L. longipalpis (Fig. 15) 17. (11) Palp 5 ahorter than segments 3 + 4 18 Palp 5 longer than segments 3 + 4 21 18. (17) Spermathecae imbricated, annuli semi-telescoped. Individual sperm ducts heavily sclerotized, wrinkled, or rugose in part or whole subgenus Psychodopygus (Fig. 61-71) Spermathecae not imbricated, annuli, if present, not 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. nord.es tin 'a (Fig. 87) Cibarium with horizontal teeth pointing toward pharynx, not slanted inwards. Flagellomere I shorter than palpus. Ascoids without visible basal spurs -0 20. (19) Spermathecae as shown, longer than individual sperm ducts. Abdominal tergite 8 without lateral setae .... vespevtilionis group, series vespertilionis (Fig. 29) Spermathecae otherwise, shorter than individual sperm ducts. Ab- dominal tergite 8 with lateral setae 21 -87- 21. (20) Spermathecae less than twice as long as wide, terminal segment smaller than preceding segment. Common sperm duct longer than in- dividual ducts oswaldoi group (in part), L. pia (Fig. S3) Spermathecae cylindrical, its length at least 3 times width, terminal segment subequal to or smaller than others. Common sperm duct shorter: than individual ducts . . . vexatoP group, series perv.ensis (in part), (Fig. 72-74; 76-78) 22. (17) Individual sperm ducts heavily pigmented in part or whole. . . subgenus Prsssatia (Fig. 32-34) 23. (22) Cibarial arch absent or conspicuous only at sides 24 Cibarial arch complete or nearly so 26 24. (23) Cibarium with horizontal teeth like flattened plates viewed almost edgewise. Common sperm duct absent pilosx group, L. pilose. (Fig. 86) Cibarium with horizontal teeth otherwise. Common sperm duct present ?.j 25. (24) Wing venation with beta less than half alpha. Cibarium with horizontal teeth pointing tox^ards pharynx, not inwardly slanted . . vexator group, series per-uensis (in part), (Fig. 75) Wing venation with beta over half length of alpha. Cibarium with horizontal teeth pointing inwards oswaldoi group (in part), L. rofotaensis (Fig. 100) -88- 26. (23) Spermathecae completely smooth-walled 27 Spei.raathecae wrinkled, with transverse striations or indentations forming complete annul! or not 28 21, (26) Spermathecae tubular, much longer than wide and only slightly wider than individual ducts. Common sperm duct short, not exceeding 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 walkeri (Fig. 19-20) 28. (26) Spermathecae as shown, with some clearly defined annuli, ter- minal 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 stria- tions, complete or not longispina group (Fig. 43-44) Spermathecae otherwise, sac- like and wrinkled ..... 30 30. (29) Individual sperm ducts nearly absent, spermathecae seemingly joxned directly to common duct. Eyes very small. . . ungrouped, L. r-angetiana (Fig. 8S) Individual sperm ducts at least as long as half length of sperma- thecae. Eyes large vewucavion group (Fig. 24-27) Figure 12 Lutzomyia (P.) hirsuta nioaraguensis female — A. Head with palpal segments marked in Arabic numerals and flage Homer es designated by Roman numerals, B. Flagellomere II showing ascoids, C. Pharynx attached to cibarium, D. Metathoracic spiracle and postspiracular setae, E. Cibarium, F. Spe.rmathecae and associated structures, G. Wing, H. Femur, tibia, and basitarsus of front leg. Female: Rio Changena, Eocas del Toro Prov. , Panama Lutzomyia townsendi male — I. Male genitalia; small subterminal seta on sryle is present, J. Genital pump and filaments, sane scale as Fig. 121. Male: Rancho Grande, Aragua State, Venezuela -90- S: II H Jim .-« i \ , fl--J/-w its ?:■■:;' A Kv7A\. lnd;>:duol Sp-rm Ouct ss: '/7 >// .1 H U \\\ C-r..'al FilniKn! •91 Subgenus Lutzomyia _F r anca. , 1924 This subgenus, divided into 3 series (Lewis et al., 1978), contains 21 species including L. mafinkcllei n. sp. described in this review. Sever, of these are anthropophilic and others probably share the £ ime habit. Lutzomyia longipalpis , the principal vector of visceral leishmani- asis in the New World (Lewis, 1974), ranges from southern Mexico to central Argentina where it is locally common in dry, nonforested areas. Lutzomyia gomezi, also reported in Colombia, has been implicated as a vector of dermal leishmaniasis in Panama (Christensen et al., 1969). In Colombia the series longipalpis is represented by 3 species — L. longipalpis, L. liahyi, and L. bifoliata. Two species, L. gomezi and L. marinkellei are included in the series aruoiata. Except for longi- palpis, the females of these species are remarkably similar and are separated only by minor nonsexual characteristics. Keys to Species Males 1. Paramere with long dorsobasal arm bearing apical setae. Coxite with 2 very broad fan-like setae at inner bass 2 Paramere without dorsobasal arm. Coxite with 4 or more simple setae at inner base. . 3 2. Paramere with dorsobasal arm bearing 2 slender, recurved setae. Coxite with patch of long setae distad of fan-like setae bifoliata (Fig. 13) Pararaere with dorsobasal arm bearing 2 fan-like setae. Coxite without a distal patch of long setae lichyi (Fig. 14) 3. Paramere with 2 strongly developed, dorsobasal setae. Coxite tuft of 4 setae. Style with a subterrninal seta. . . longipalpis (Fig. 15) Paramere without, such setae. Coxite tuft of 10+ setae. Style without a subterminal seta 4 4. Pleura entirely pale. Style over half length of lateral lobe. Para- mere with dorsal setae on distal half of structure, .gomezi (Fig. 16) Pleura mostly dark. Style shorter than half length of lateral lobe. Paramere with short dorsal setae confined to distal third of structure marinkellei (Fig. 17) Females 1. Cibarium with at least 8 horizontal teeth. Spermathecae as shown, terminal annulation not markedly larger than others longipalpis (Fig. 15) Cibarium with 4 horizontal teeth. Spermathecae otherwise, terminal annulation spherical much larger than others 2 2. Pleura pale, contrasting with dark mesonotum. Flagellomere I shorter than or equal to length of labrura gomezi (Fig. 16) Pleura mostly dark. Flagellomere I longer than labrum 3 3. Flagellomere I less than 0.32 mm long marinkellei (Fig. 17) Flagellomere 1 greater than 0.32 mm long 4 4. Terminal flagellomere longer than preceding f lagellomere (XV) . . . lichyi (Fig. 14) Terminal flagellomere shorter than preceding flagellomere (XV) . . . bi.foZia.ta (Fig. 13) Series longipalpis 12. Lutzomyia (L.) bifoliata (Fig. 13) Lutzomyia bifoliata Osorno, Morales, Osorno, & Hoyos, 1970: 8 (c?, El Terminal, Municip. Puerto Boyaca, Boyaca, Colombia). Osorno et al., 1972a: 15 (listed). Forattini, 1973: 248 (gen. review, 6" fig.). Distribution: Colombia (Antioquia, Boyaca) . Material examined: Colombia. 13 <$d, 19 o, Rio Anori (Antioquia), tree buttresses, Sept. 1970, D.G.Y. 3 6*0", 1 cp , light traps, same locality May 1970, C.H.P. Discussion'. Lutzomyia bifoliata males from Rio Anori generally agree with the description and figures given by Osorno et al. (1970) but the strong erect seta on the dorsum of the paramere is lacking in our material. The female, described below, closely resembles L. lichyi but the terminal flagellomere is shorter, not longer than the preceding flagellomere. The longer labrum of bifoliata females (0.37-0.42 mm as opposed to 0.25-0.36 mm, n = 17, in lichyi) also may be useful in dis- tinguishing the species if it proves to be a consistent, nonoverlapping feature . The following description is based on 10 females captured near the Rio Anori . -94- Female: Wing length 2,35-2.57; width 0.73. Head, mesonotum, and abdominal tergites strongly pigmented: Rest of insect, including most of pleura, paler. Head height 0.46; width 0.44. Eyes separated by 0.14 or distance = to ca . 7.8 facet diameters. Flagellomere I (0.40-0.50 long), 1.2 x length of II + III; ascoids longer than those of 6", but not reaching ends of flagellcmeres , with almost indiscernible posterior spurs, present on all but last flagellomere. Length of palpal segments: 1 (0.06) 2 (0.15-0.18), 3 (0.19-0.21), 4 (0.09-0.12), 5 (0.30-0.37); palpal sen- si 11a on segments 2 and 3. Labrum length = 0.37-0.42. Cibarium with 4 nearly straight, sharp horizontal teeth and 20-30 vertical teeth dis- tributed as shown; chitinous arch complete, well defined; pigment patch sub triangular, darker, and broader posteriorally . Pharynx (0.2C long) unarmed. Pleura with 14-22 upper and 3-5 lower episternal setae. Length of wing vein sections: Alpha. (0.64-0.71), beta (0.27-0.34), delta (0.13-0.19), gamma (0.27-0.34). Length of femora, tibiae, and basitarsi of slide 618: Foreleg, 0.95, 1.22, 0.74; midleg, 0.91, 1.47, 0.81; hindleg, 0.98, 1.67, 1.27. Abdominal sternite 2 horseshoe-shaped, hollow in middle. Tergite 8 without setae. Spermathecae and sperm ducts as figured. 13. Lutzonyia (L.) lidhvi (Fig. 14) Phlebotomus lichyi Floch & Abonnenc, 1950a: 1 (o holotype, Rio Borburata, Carabobo, Venezuela). Morales et al., 1969a: 381 (Meta, Colombia) . Phlebotomus VexillaHus Fairchild & Hertig, 1952: 514 (6*, o, Panama) Plfano et al., 1960: 65 (Miranda State, Venezuela). Pifano et al., -95- 1962: 385 (keyed). McConnell & Correa, 1964: 527 (infected with fungi/. Flcch & Kramer, 1965: 1 (as synonym of lichyi) . Hanson, 1968: 90 (larva & pupa). Scorza et al., 1968: 35 (bionomics). Calderon, 1973: 87 (Merida State, Venezuela). PhZebotomus foliatus Mirsa & Ortiz, 1952: 249 (. Female cibarium. Male: Rio Anori, Antioquia Dept., Colombia Female: Same locality as male ■103- Figure 14 Lutzonvjia (L.) liohyi — A. Male head, B. Male flagellomere II, C. Female head, D. Female flagellomere II, E. Female cibarium and pharynx, F. Spermachecae, G. Female cibarium, H. Female wing, I. Male v/ing, J. Male genitalia. Male: Alto Curiche, Choco Dept., Colombia Female: Curiche, Choco Dept., Colombia -105- Figure 15 Lutzormjia (L.) longipalyis — A, Male head, B. Male flagellomere II, C. Feraale head, D. Female flagellomere II, E. Female cibarium and pharynx, F. Female wing, G. Male wing, H. Genital pump and filaments, I. Male genitalia, J. Body of spermatheca, K. Spermathecae, L. Genita] pump, same scale as Fig. 151, M. Paramere of male from Huila Dent., Colombia, same scale as Fig. 151, N. Female cibarium. Male: Calabozo, Guarico State, Venezuela Female: Same locality as male (except for Fig. 15M) -107- Figure 16 Lutzorwjia (L.) gomezi — A. Male head, B. Male flagellomere II, C. Female head, D. Female f lagellomere II, E. Female cibarium and pharynx, F. Male genitalia, G. Spermathecae, H. Female wing, I. Male wing, J. Female cibarium. Male : Trinidad Female: Trinidad Figure 17 Lutzomyia (L.) mapinkellei — A. Spermathecae, B. Female wing, C. Male. wing, D. Female flagellomere IT, E. Base of ascoid, F. Female head, G. Female cibarium, H. Male head, I. Male genitalia. Kale: Tres Esquinas, Caqueta Dept., Colombia Female; Rio At"ipuana, Mate Grosso -State, Brazil -111- V -112- Migonei Group Theodor, 1965 This group is divided into 3 series following Lewis et al. (1978). The series rrrCgonei , walkeri, and cGstalimai (= subgenus Boj.'rettomyia Martins and Silva, 1965) . Species in the latter series occur in Brazil but some of those in the other series are more widespread with 4 species occurring in Colombia. iSarretto (1562) and Forattini (1971a, 1973) in- clude some of the mi-gonei group taxa in the subgenus Coromyia Barretto, 1962. Based on present knowledge, the only man-biting species in this group is L. migonei, here reported in Colombia for the first time. Keys to Species Males 1. Genital filaments over 3 x length of pump, with simple tips. Coxite with basal tuft of 4-6 short setae migonei (Fig. 18) Genital filaments less than 3 x length of pump, their tips spherically enlarged with an inner "tooth." Coxite with basal tuft of 10+ longer setae and a median ventral group of 6+ setae 2 2. Aedeagus simple, without a dorsal projection. Genital filaments 0.30 mm or longer mava^oensis (Fig. 19) Aedeagus complex, with a dorsal cylindrical projection. Genital filaments less than 0.30 mm loalkeri (Fig. 20) ■113- Fema las 1. Spernathecae narrow and tubular, at least 3 x longer than wide. . . migonei (Fly. 18) Spermathecae capsule-shaped, less than 2 x as long as wide 2 2. Individual sperm ducts over 3 x length of common duct, wider and with fine transverse striations basally sp.de Baduel (Fxg. 18) Individual sperm ducts about 1/2 as wide as common duct r.arajoensis (Fig. 19) Individual sperm ducts more slender, about 1/3 as wide as common duct walkevi (Fig. 20) Series miaonei 1 7 . Lutzomyia migonei (Fig. 18) Phlebotomy^ migonei Franca, 1920: 230 (c? holotype, Assuncion, Paraguay). Barretto, 1947: 211-213 (full refs., synonyms). Forattini, 1954: 214 et seq. (sternites, figs.). Forattini, ]960: 479 (Amapa, Brazil). Pifano et al., 1962: 387, 388 (6*, o, keyed). Calderon, 1973: 87 (Merida State, Venezuela) . Lutzomyia migonei: Theodor, 1965: 182 (cf, o, figs.). Forattini, 1973: 122 et seq. (gen. review, refs., immatures, figs.). Lewis, 1975a: 500 et seq. (mouthpart morphol . ) . Distribution: Colorrbia (Magdalena) , Venezuela, Trinidad, Brazil, Paraguay, Argentina. -114- Material examined: Colombia. 2 SJ', Rio Don Diego, E of Santa Marta (Magdalena), tree trunks, 15 Aug. 1973, D.G.Y. & R.C.W. Brazil. 2 3$, Laphina (Ninas Gerais) , flight trap near cave, 28 Aug. 1974, D.G.Y. & P. Williams. Trinidad. 2 6*6', Bush Bush Forest, Nariva Swamp, chicken- baited trap, 17-18 May 1961, T.H.G. Aitken. 1 o_, same data but 22-23 Kay. Discussion: Ranging from Argentina to northern Colombia, L. nrigonei has been found naturally infected with flagellates of uncertain identity in Venezuela and Brazil (Johnson et al . , 1963; Forattini, 1973). Other information dealing with the biology of this anthropophilic species was reviewed by Forattini (1973) and need not be repeated here. Series walkeri 18. Lutzomyia mavajoensis (Fig. 19) Pklebotomus marajoensis Damasceno & Causey, 1944: 339 (6* holotype, Ilha do Marajo, Brazil). Fairchild & Hertig, 1961b: 250-254 (cf, o, redescr., figs., refs., distrib.). Pifano et al., 1962: 386 et seq. ( cM >*-^, ~\\ \ [\ il 7/ R| Figure 19 Lutzowyia marajoensis — A. Male head, B. Male f lagellomere II, C. Female head, D. Female f lagellomere II, E. Female cibarium and pharynx, F. Body of spermatheca, G. Male genitalia, H. Genital pump, I. Genital pump and filaments, J. Tips of genital filaments, X. Spermathecae, L. Female wing, M. Male wing, N. Female cibarium. Male: San Jose, Los Santos Prov. , Panama Female: Puerto Mensabe, Los Santos Prov., Panama Figure 20 Lutzomyia walkevi — A. Spermathecae, B. Female cibarium, C. Male genitalia, D. Tip of genital filament, greatly enlarged. Male: Trinidad Female r Trinidad -124- ( J \ \ 1® \ w i 4 i n A vs IV'X /v;;.m -125- So-'.lcnsis Group Lewis et. al., 1978 This group was created to accomodate two closely related species -- L. saulensis and L. wilsoni (Damascene) and Causey). Forattini (1971a, 19/3) places the former species in the subgenus Dampfomyia Addis but the spermathecae are structurally different (Theodor, 1965), the male para- meres lack a dorsal arm, and the style has no subterminal bristle. The females of saulensis and wilsoni are nearly identical (Vianna Martins, pers. comm.), their populations probably overlapping in northern Brazil. 21 . Lutzomyia saulensis (Fig. 21) Phlebotomus saulensis Floch & Abonnenc, 1944a: 11 (6* holotype, Saul, Haute Mana, French Guiana). Barretto, 1947: 222 (listed). Damasceno et al., 194'?: 831 (Brazilian records). Floch & Abonnenc, 1952: 35, 112- 114 (6* keyed, redescr., figs.). Fairchild & Hertig, 1958b: 204-205 (refs., taxonomic discussion). Fairchild & Hertig, 1959: 122 (distrib.). Sherlock, 1962: 321, 330 (Santander, Colombia). Pifano et al., 1962: 386, 389 (o*, o, keyed, Venezuela). Osorno et al . , 1967: 28 (mention). Phlebotomus pinealis Floch & Abonnenc, 1944a: 11 (cj> holotype, Crique Anguille, French Guiana). Fairchild & Hertig, 1958b: 204-205 (refs. as synonym of saulensis). Johnson &. Hertig, 1961: 765, 774 (rearing attempt). Hanson, 1968: 69 (1st instar larva, descr., fig.). Lutzonniia saulensis: Martins et al., 1963: 335 (Roraima, Brazil). Martins et al . , 1965: 3 (Rondonia, Brazil). Theodor, 1965: 196 (o*, g, figs.). Barreto, 1969: 466-467 (Valle, Colombia). Lewis et al., 1970: 215 (parous study). Fraiha et al . , 1970b: 215 (mention). -126- Christensen & Fairchild, 1971: 301 (Panama), Chaniotls et al., 1971a: 344 et seq. (pop. dynamics, Panama). Osorno et al., 1972a: 66 (Colombian records). Christensen, 1972a: 88 (listed). Chaniotls et al., 1972: 95 et seq. (resting sites). Shaw & Lainson, 1972: 710 et seq. (infected with nonlelshmanial flagellates, collecting data, Brazil). Llanos, 1973: 34 (6*, o, redescr., figs., Peru). Forattini, 1973: 327 et seq. (gen. review, figs.). Christensen & Herrer, 1973: 579 (listed). Ward et al., 1973: 178 (attracted to rodents and man, Brazil). Lainson et al . , 1973: 190 (negative for Leishmanial . Lewis, 1975a: 504 et seq. (mouthpart morphol.). Llanos et al., 1975b: 6 71 (Peru). Llanos et al., 19 76: 480 (Peru). Martins et al., 1976b: 496 (Peru). Distribution: Costa Rica, Panama, Colombia (Antioquia, Caqueta , Choco, Santander, Valle) , Peru, Brazil, French Guiana. Material examined: Colombia. 2 op, Rio Anori (Antioquia), light traps, 3 May 1970, C.H.P. 8 d'o", 11 oo, same data but some in tree hollows, Sept. 1970. D.G.Y. 14 6*d\ 55 ^o, Curiche (Choco), light, Shannon & Malaise traps, tree trunks, May-Dec. 1967, D.G.Y. 3 og, Alto Curiche (Choco), Malaise & light traps, Aug., Spet., Nov. .1967, D.G.Y. 1 g, Teresita (Choco), Malaise trap, 22 April 1967, D.G.Y. 4 ^o, Rio Atrato at Sautata (Choco), Malaise traps, Nov. -Dec. 1967, Jan. 1968, D.G.Y. 2 cfo", Anchicaya Dam (Valle), tree trunks, 9 Aug. 1973, D.G.Y. & R.C.W. 1 6", same data but light trap, 10 Aug. 1973. 2 o^ , 25 km E of Buenaventura (Valle), flight trap, 12 Aug. 1973, D.G.Y. & R.C.W. Brazil. 1 6", Belem (Para), 1 Feb. 1944, R. Damasceno & 0. Causey. Discussion: Although widely distributed in lowland forests, L. saulensis is a little known species. Ward et al . (1973) collected -127- specimens on rodent and human bait in Para, Brazil, but this sper-.ies can not be regarded as being anthropophilic. Both sexes are readily identi- fiable. Figure 21 Lutzomyia saulensis — A. Male head, B. Male flagellomere II, C. Female head, D. Female flagellomere II, E. Female cibarium and pharynx, F. Male genitalia, G. Genital pump and filaments, H. Spermathecae, I, Female wing, J. Male wing, K. Female cibarium. Male: Curiche, Choco DepL., Colombia Female: Same locality as male -130- Verruoarum Group Theodor, 1965 This group of 22 species is divided into 2 series based on male characteristics (Theodor, 1965) -- the series verruaarvm (14 spp.) and the series ssrrana (7 spp.). At present, L. buvsifortnis (Floch & Abonnenc) from French Guiana cannot be placed because the male remains undiscovered. Some authors include these species in the. subgenus Lutzomyia Franca (Forattini, 1971a, 1973) or in the subgenus Cororraia Barretto, 1962. Two species in the series sewana and 11 in the series vevruearum occur within or very near the borders of Colombia. The majority of species in the latter series are Andean, although L. ovallesi and L. evansi are widespread in lowland areas. The Andean taxa are mor- phologically 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 dis- tinguishing characteristics for some of the species. It may be appropri- ate to add that the presence of L. verrucarum (Townsend) in Venezuela (Anduze et al . , 1947; Floch and Abonnenc, 1950a; Forattini, 1973) can be confirmed only when the male is discovered in that country. Key to Species Males 1. Style of male genitalia with 2 or 3 strong spines (series s err ana) . 2 Style with 4 strong spines (series verruc-ai^uni) 3 -131- 2. Coxite tuft of 5-6 setae, upturned but not strongly recurved at tips, implanted in a straight row sen-ana (Fig. 22) Coxite tuft of 6-7 setae, markedly recurved apical ly, not implanted in a straight row odax (Fig. 28) 3. Basal spine of style isolated Basal spines of style paired, inserted more or less on the same level 7 4. Terminal spine of style very thick and crooked. Paramere as in Fig. 28£ spinier as sa (Fig. 28) Terminal spine of style relatively slender, not crooked. Paramere otherwise 5 5. Middle third of paramere with dorsal and ventral setae sauroida (Fig. 28) Middle third of paramere without dorsal or ventral setae 6 6. Paramere club-like, angular at tip and with dorsal subspical row of 13-16 short spine-like setae longiflocosa (Fig. 28) Paramere more or less rounded at end, less club-like, dorsal subapical setae longer, not spine-like quasitounsendi (Fig. 28) 7. Coxite with subapical group of long hairs. Basal spines of style in- serted on well marked rather long tubercles 8 Coxite without subapical group of long hairs. Basal spines of style not implanted on tubercles 9 •132- 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 lab rum ........ columbiana (Fig. 24) Paramere not clubbed. Coxite tuft of fewer than 7 setae. Flagellomere 1 short, subequal in length to labrum OVallesi (Fig. 27) 11. Basal coxite tuft of 18+ setae. Distal third of paramere wider than middle third disinnata (Fig. 23) Basal coxite tuft of 10 or fewer setae. Distal third of paramere not enlarged 12 12. Genital filaments over 4 x length of pump. Style with a subterminal seta evansi. (Fig. 25) Genital filaments less than 4 x length of pump. Style without a sub- terminal seta nuneztovari (Fig. 23) -133- U"ie3 8Cl'r\:-l:7 22. Lutzomyia odax (Fig. 28) Phlebotomus odax Faircbild & Hertig, 1961b: 239 (6* holotype, o, Almirante, Bocas del Toro, Panama). Lutzomyia odax: Barretto, 1962, 97 (listed). Martins et al . , 1965: 7 (cf. to dubia) . Christensen & Fairchild, 1971: 302 (Darien, Panama). Christensen, 1972a: 88 (listed). Christensen et al., 1972: 57 (Panama). Forattini, 1973: 213 et seq. (gen. review, figs.). Distribution: Guatemala, Honduras , Nicaragua, Panama. Material- examined'- Panama, cf holotype (no, 4168), Almrirante (Bocas del Toro), light trap, 15 Oct. 1952, A. Quinones. o allotype (no. 3327), same data, but human bait collection, 20 June 1951, A. Vivanco. 1 cf, same data but 17 June 1950, R. Hartmann. discussion: There is little doubt that this species occurs in Colombia, Christensen and Fairchild (1971) having reported its presence within 4 km of the Colombian border in Darien Province, Panama. Females closely resemble those of L. serrana but differ in the lengths of the individual sperm ducts (Fairchild & Hertig, 1961b) . Both species occasion- ally bite man but are more common in light trap and resting collections. Forattini (1973) may be correct in treating L. Cybia Martins, Falcao, & Silva as a junior synonym of L. odax but more evidence is needed to support this view. Martins et al . (1965) point out differences between the taxa, especially those of the male genitalia. •134 23. Lutzorrgia serrana (Fig. "22) Vhlebotomus serranus Damasceno & Arouck, 1949: 843 (6* holotype, Serra da Piriabas, Para, Brazil). Forattini, 1960: 479 (Amapa, Brazil). Blancas, 1959-1960: 125 (Peru). Fairchild & Hertig, 1961b: 237 (refs., descr., figs.). Fianson, 1961: 320 (breeding sites). Johnson & Hertig, 196.1: 765 (rearing data) . Sherlock, 1962: 330 (Santander, Colombia). Rosabal, 1966: 1 (Costa Rica). Hanson, 1968: 76 (larva, pupa, descr.). Phlebotomus guayasi Rodriguez, 1956: 76 (cf, o, Guayas, Ecuador). Fairchild & Hertig, 1961b: 237 (as synonym of serranus) . Lutzomyia serrana: Barretto, 1962: 97 (listed). Martins et al . , 1965: 3 (Rondonia, Brazil), 6 (cf. to dvbid) . Lewis, 1967a: 74 (listed) Forattini, 1971a: 100 (listed). Tesh et al . , 1971a: 152 (blood meals). Chaniotis et al., 1971a: 345 (pop. dynamics, Panama). Osorno et al., 1972a: 30 (Colombian records). Tesh et al., 1972: 90 (blood meals). Chaniotis et al., 1972: 95 (listed). Christensen, 1972a: 88 (listed). Forattini, 1973: 123 et seq. (gen. review, figs.). Llanos, 1973: 32 (Peru, figs.). Velasco, 1973: 77 (Bolivia). Llanos et al . , 1975b: 668 (unusual spermathecae, fig.), 67]. (Peru). Herrer & Christensen, 1976a: 62 (Panama). Llanos et al . , 1976: 480 (Peru). Martins et al . , 1976a: 488 (Peru) . Martins et al., 1976b: 496 (Peru). Distribution: Mexico, Honduras , Nicaragua, Costa Rico,, Panama, Colombia (Antioquia, Bolivar, Boyaca, Caldas, Chocc, Cundinamarca, Sant- ander), Ecuador, Peru, Bolivia, Brazil. Material examined: Colombia. 1 c?, Alto Curiche (Choco) , tree, but- tress, 21 May 1967, D.C,.^. 1 cf, Rio Anori (Antioquia), tree buttress, 6 Feb. 1970, C.H.P. 1 0, same data but at light, 21 Sept. 1970, D.G.Y. -135- Panama. 1 ] <2, Rio Platanar (Panama), hollow tree, 20 July 1950, R. Hartmaan. Discussion: In Panama, Tesh et al .. (1971a, 1972) identified the blood meals of 14 females using the precipitin test. Ten had fed upon rodents, 2 on edentates and 2 on marsupials. Dr. C.H. Porter (in litt.) collected less than 10 specimens on human bait over a one year period at Ric Anori (Antioquia) , Colombia. Series verrucarum 24 . Lutzomyia andina (Fig. 23) Lutzomyia andina Osorno, Osorno, & Morales, 1972b: 2 (c? holotype, p, El Chariquito, Cundinamarca, Colombia). Osorno et al., 1972a: 24 (Colombian records) . Fh'lebotomus verruaaricn: Osorno et al., 1967: 29 (not verrueanm Townsend) . Distribution: Colombia (Cundinamarca). Material examined: 1 6", paratype, type locality, 2550 m above sea level, Aug. 1966. 1 o, paratype, same data but Sept. 1966. Discussion: Specimens of this recently described species, known only from the type locality, were taken on human bait and in tree holes. Females bite man during the day and night. The shorter of the paired basal spines of the male sytle is curiously modified, the apical part widened and with a flange (Fig. 2 3A) . This characteristic, if shared by other andina males, will serve to separate it from other males in the verrucar-jm group. -136- 25 . Lu tzomyia co I vjrb i an a (Fig. 24) Phlebotomus columbianus Ristorcel]i & Van Ty, 194.1: 263 (o holotype, Valle He Capuli, Narino, Colombia). Rozeboom, 1947b: 705 (6*, figs.). Sherlock, 1962: 322 (cf, o, figs.). Osorno et a.l . , 1967: 28 (mention). Phlebotomus montieolus var. incarwn Ristorcelli & Van Ty, 1941: 266 (o, Valle de Capuli, Narj.no, Colombia). Rozeboom, 1947b: 705 (as synonym of colurrbiand) . Lutzomyia colvmbiana: Rarretto, 1962: 97 (listed). Osorno et al., 1972a: 24-26 (Colombian records). Forattini, 1973: 206 et seq . (gen. review, figs . ) . Distribution: Colorrbia (Cauca, Guajira, Narino, Valle). Material examined: Colombia. 5 6*6", 3 oo, Bolivar (Cauca), Jan. 1944, J. Burbano. 1 (J1, 3 oo, San Pedro (Narino), Jan. 1944, J. Burbano . 2 6*6*, ca. 30 km NW of Cali, near El Carmen (Valle), tree trunks, 12 Aug. 1973, D.G.Y. & R.C.W. Discussion: This precinctive Colombian species was the suspected vector of bartonellosis during the outbreak in southeastern Colombia over 40 years ago (Rozeboom, 1947b) . The record of colvmbiana. from Montes de Oca and Serra Kacuira in Guajira (Osorno et al . , 19 72a) is very interesting, these localities being widely separated from southeastern Colombia where colurrbiana, was first discovered. The Andes, especially the eastern chain, may have acted as a corridor for the dispersal of this species. 1 have seen other specimens of colunfoiana in the collections of the California Academy of Sciences and University of Florida. All were cap- tured in southeastern Colombia and were identified by 0. Mangabeira and P. Galindo. ■137- 26. Lutzorr^ia disiunata (Fig. 23) Lutzomjia disiunata Morales, Osorno, & Osorno, 1974: 446 (cfholo- type, o, Hacienda La Conejera, Municip. de Suba, Bogota, Cundinamarca, Colombia) . Distribution: Col-orrhia (Cesar, Cundinamarca). Material examined: Colombia. 8 3d1, 21 oo, Municip. Agustin Cadazzi (Cesar), Sept. 1969 (INPES) . 3 do" (including holotype) , 9 o^, type locality, Aug. -Sept. 1969 (INPES). Discussion: This is another Andean species which is anthropophilic , specimens having been taken on human bait during the day and night (Morales et al., 1974). Other specimens were captured in tree holes in Cesar Department. 27. Lutzcyia evansi (Fig.' 25) Phlebotomus evansi: Nunez-Tovar, 1924: 44 (cf, Mariara, Carabobo, Venezuela). Barretto, 1947: 198 (full refs.). Mirsa, 1953: (rearing data). Fairchild & Kertig, 1959: 122 (distrib.). Pifano et al . , 1962: 412 (refs., redescr., figs.). Rosabal & Trejos, 1964: 167 (El Salvador) Rosabal & Trejos, 1965: 222 (El Salvador). Osorno et al . , 1967: 28 (Narino, Colombia) . Lutzomyia evansi: Theodor, 1955: 183 (listed). Morales et al., 1969b: 385 (mention). Osorno et al . , 1972a: 27 (Colombian records). Llanos, 1973: 34 (d*, o. redescr., figs., Peru). Forattini, 1973: 122 et seq. (gen. review, figs.). Llanos et al . , 19 75b: 670 (Peru). -138- Distribution: Guatemala,, El Salvador, Honduras, Nicaragua, Costa Rica, Colombia (Bolivar, Guajira, MagJalena, Narino, Santander) , Peru, Venezuela. Material examined: Colombia. 1 q, Arjona (Bolivar), 27 May 1940. 1 o, 10 km SE of Santa Marta (Magdalena) , tree trunk, 17 Aug. 1973, D.G.Y, & K.C.W. Honduras: 30+ d*cf, 100+ oo, Chumbagua, Santa Barbara, light traps, July 1966, J. Matta. Other specimens from Central America and Venezuela in collection at UF. Discussion: The female of evansi has very long sperm ducts and relatively small spermathecae (Fig. 25J) , features which separate it from other verrucavum group females in Colombia. This species has been re- ported biting man in several countries including Colombia (Osorno et al., 1972a). It and L. longipalpis often, but not always, occur together in Central America, Colombia, and Venezuela. The apparent absence of this species from Panama is most peculiar. 28. Lutzomyia longifloeosa (Fig. 28) Lutzomyia longifloeosa Osorno, Morales, Osorno, & Hoyos, 1970: 1 (c? holotype, Municip. de Tello , Huila, Colombia. Osorno et al., 1972a: 28 (Colombian records). Osorno et al . , 1972b: 18 (cf. to andina and quasitownsendi) . Distribution: Colombia (Huila) . Material examined: Colombia. 1 d" (paratype no. 1641), type local- ity, 9 Feb. 3 968, E. Osorno et al. Discussion: Other than resting in tree holes at the. type locality, nothing is known about the habits of this species. The female is unknown. •139- 29. LutzomyLa moralesi n. sp. (Fig. 26) Male: Wing length 1.96; width 0.58. Head, mesonotum, and genitalia strongly pigmented, rest of insect pale. Head height 0.35; width 0.32. Eyes separated by 0.11 or by distance = to 5.9 facet diameters. Flagel- lomere I (0.33 long), 1.2 x length of II + III; ascoids simple, those on flagellomere IT ending near middle of structure, on all f lagellomeres except last. Length of palpal segments: 1 (0.04), 2 (0.15), 3 (0.16), 4 (0.1 1), 5 (0.32); palpal sensilla (ca. 10) grouped loosely at middle of segment 3. Labrum length 0.21. Cibarium with 30+ vestigial, dot-like teeth; chitinous arch complete, diffuse in middle; pigment patch in- visible. Pharynx 0.15 long, unarmed. Pleura with 8-9 upper and 4 lower episternal setae. Length of wing vein sections: Alpha (0.53), beta (0.18) delta (0.17), garmr.a (0.32). Length of femora, tibiae, and basitarsi: Foreleg, 0.78, 0.90, 0.54; midleg, 0.74, 1.07, 0.61; hindleg 0.83, 1.22, 0.69. Abdominal sternites hardly visible some with lateral, circular openings. Genitalia: Style 0.13 long, shaped as shown, the proximal 2 spines inserted at the same level, the longer on a long process, no sub- terminal seta. Coxite (0.32 long x 0.12 wide) with a distal patch of 30+ straight or nearly straight setae, a dense tuft of ca . 30 thick sinuous setae at base. Paramere as shown but variable according to angle of view; the apical part with an incipient lateral lobe and a ventral heel. Aedeagus subtriangular, unmodified. Lateral lobe relatively long (0.42). Genital pump (0.155 long), each filament 0.63 long or 4 x length of pump, tip enlarged. Cercus as shown. Female: Wing length 2.45; width 0.76. Coloration as in d1. Head height 0.43; width 0.39. Eyes separated by 0.13 or distance = to ca. 6.6 -140- facet diameters. Flagellomere I (0.31 long) nearly 1.3 x length of II + III; .ascoids longer than those of o but not reaching end of flagellomere II; on all flagellomeres except last. Length of palpal segments: 1 (0.014), 2 (0.20), 3 (0.19), 4 (0.12), 5 (0.39); palp 2 with ca. 5 apical sent, ilia, palp 3 with 25+ sensilla on apical half. Lab rum 0.36 long, Cibarium with 4 sharp horizontal teeth evenly spaced, 2 transverse rows of subequal vertical teeth, ca . 13 teeth in each row; chitinous arch complete; pigment patch subtriangular , well defined. Pharynx 0.20 long, unarmed but with distinct ridges. Pleura with 9-10 upper and 2 lower episternal setae. Length of wing vein sections: Alpha (0.67), beta (0.20), delta. (0.26), gemma. (0.41). Length of femora, tibiae, and basi- tarsi: 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. Cerci unremarkable . Distribution: Colombia. (Valle) . Material examined: Colombia. 6* holotype (no. 512), ca. 15 km SW of Cali (1600 m above sea level) near Rio Pance (Valle), on tree trunk, 8 Aug. 1973, D.G.Y. & S. Ayala. ^ allotype (no. 513), ca . 10 km W of Cali (1800 m above sea level), near Pichinde (Valle), rock crevice, 2 Aug. 1973, D.C.Y. & R.C.W. Paratypes (nos. 514-526), 1 v ^> ■:'';>:': _j,- \\ *5bLJ> -161- Vespertilionis Group Theodor, 1965 With the center of distribution in Central America, the vespertili.- onis group species are divided into 2 series following Theodor (1965) -- the series vespertitinois (5 spp.) and the series deleoni (3 spp.). Barretto (1962) created the subgenus CoTomyia to accomodate them and other species, mostly those in the verrucamm group. Forattini (1971a> 1973) divides the vesper tiliorvis group species by placing 1 in the sub- genus Lutzomyia Franga, 2 in the subgenus Dampfomyia Addis, while leaving the others in Coromyia. In my opinion, this arrangement is unnecessary when the species are classified using a combination of character states. Two sibling species in the series vespertili-onis are known to occur in Colombia. The females feed on bats and are morphologically indis- tinguishable . Key to Species Males 1. Paramere with dorsal setae on distal half of structure L. vespertilionis (Fig. 29) Paramere with dorsal setae restricted to distal third of structure L. isovespertilionis (Fig. 29) 35. Lutzomyla isovespei-tili-on-ls (Fig. 29) Phlebotomies ■isovespertilioni-s Fairchild & Hertig, 1958a: 516 (6* holotype, p, reared from eggs laid by p from tributary of Rio Cocoli, Canal Zone). Johnson & Hertig, 1961: 765 et seq. (rearing data). -162- Thatcher & Hertig, 1966: 46 (mention). Hanson, 1968: 88-90 (larva, pupa, descr.) . Phleho tonus vespevtitionis : Fairchilci b. Hertig, 1947b: 617 et seq. (in part, descr., fig.). Luizcmuia isovespevtilionis: Barretto, 1962: 97 (listed). Chaniotis et al . , 1971a: 344 et seq. (in part, pop. dynamics, Panama). Chaniotis et al., 1972: 94 et seq. (resting sites, Panama). Christensen et al . , 1972: 57 et seq. (collecting data, infected with trypanosomes , Panama). Christensen, 1972a: 88-89 (mention). Forattini, 1973: 122 et seq. (gen. review, fig.). Christensen & Herrer, 1973: 579 et seq. (collecting data, Panama). Rutledge & Mosser, 1975: 411 (mention). Herrer & Christensen, 1969: 62 (collecting data, Panama) . Pistx'ibution: Panama, Colombia (Choco). Material examined'. Colorrbia. 1 o", Curiche (Choco), tree buttress, 5 July 1967, D.G.Y. 4 dcf, Teresita (Choco), tree buttress, 18 May 1967, D.G.Y. Panama. 1 6", Cruces trail, Madden Forest, Canal Zone, hollow tree, 7 Sept. 1949, M. Hertig. 3 6*cf, Cerro Campana (Panama Prov.), tree hollows and buttresses, 2 Oct. 1949, M. Hertig & H. Trapido. Discussion: See L. vespertilicnis 36. Lutzomyia vespertilioni (Fig. 29) Phlebotomies vesperhitionis Fairchild & Hertig, 1947b: 617 (6" holo- type, Cerro Campana, Panama Prov., Panama). Hertig & Fairchild, 1950: 91 (abdor.iin.-il steraites). Barretto, 1951: 225 (distrib.). Vargas & Difiz-Najera, 1951b: 101 (cf. to beltrani) . Floch & Abonnenc, 1952: 30-48 (d\ o, keyed). Rosabal, 1954: 10, 28-29 (measurements, figs., Costa Rica, sp. no. 12 = Vesper tilionis) . Rodriguez, 1956; 76 et seq. (Ecuador). Fairchild £< Hertig, 1958a: 510 (keyed). 514 (o, o, redescr., figs.). Fairchild & Hertig, 1959: 122 (Central American records). Hanson, 1961: 320 (breeding site, Panama). Johnson & Hertig, 1961: 765 et seq. (rearing). McCormell & Correa, 1964: 523 et seq. (infected with trypanosomes & fungi, Panama). Thatcher & Hertig, 1966: 46 et seq. (mention). Hanson, 1968: 88-90 (larva, pupa, descr., figs.). Lutzomyia vespertilionis: Barretto, 1962: 97 (listed). Theodor, 1965: 184 (o*, o, figs.). Barreto, 1969: 468 (Valle, Colombia). Zele- T don & Rosabai, 1969: 221-227 (as vector of bat trypanosome) . Tesh et ai., 1971a: 152-153 (in part, blood meals). Chaniotis et al . , 1971a: 344 et seq. (in part, pop. dynamics, Panama). Christensen et al., 1971: 116 et seq. (pyloric armature, figs.). Tesh et al . , 1972: 90 et seq. (in part, blood meals). Chaniotis et al., 1972: 94 et seq. (resting sites, Panama). Christensen, 1972a: 88-89 (mention). Christensen, 1972b: 683 et seq. (rearing). Osorno et al . , 1972a: 32-33 (Colombian records). Rutledge & Mosser, 1972: 307 (mention). Christensen & Herrer, 1973: 579 et seq. (collecting data, infected with trypano- somes, Panama). Forattini, 1973: 123 et seq. (gen. review, figs.). Rutledge & Mosser, 1975: 411 (mention). Distribution: Costa Rica, Panama, Colombia (Antioquia, Bolivar, Boyaca, Choco, Valle), Ecuador. Material examined: Colombia. 3 66, Rio Anori (Antioquia), tree trunks, 6 Feb. 1970, C.H.P. 2 oV, 1 o, same data but Sept. 1970, D.G.Y. 4 dc/, 2 co, Anchicava Dam (Valle), tree trunks, 9 Aug. 1974, D.G.Y. & t i R.C.W. 7 dV, 14 oo, same data but light trap at entrance of tunnel, -164- 10-11 Aug. 1974. 1 6*, 1 o, 25 km E of Buenaventura (Valle) , tree trunks, 11 Aug. 1974, D.G.Y. & R.C.W. Ecuador. 15 c?tf, 1 o, 17 kn E of Santo Domingo do los Colorados (Pichincha) , tree trunks, 4-6 May 1976, D.G.Y. & 1. Rogers. Panama. 1 o* (holotype no. 761), Cerro Campana (Panama), shallow cave, 17 Jan. 1947. Discussion: In structure and habits, L. vesper til-ionis and L. isovespertilionis are remarkably similar, both species often being found in tree hollows associated with bats. At least one species of Trypanosoma is transmitted to these mammals by one or both of these flies (Zeledon and Rosabal, 1969) . The preferred breeding site(s) of vespertilionis and -isovespertilionii has not been determined (Rutledge and Mosser, 1972), the only wiJd caught larvae being reported by Hanson (1961, 1968) who recovered this stage on two occasions in soil and bat guano inside of hollow trees. Apparently the larvae and pupae of these species are indistinguishable (Hanson, 1968). Lubzomyia vespertiZionis ranges as far south as Guayas Province, Ecuador. Luty.omijia isovespertilionis probably has a more limited dis- tribution as noted by Fairchild and Hertig (1958a) , its southernmost locality at present being Choco Department, Colombia. These are the only vespertilionis group species which occur in South America. Fairchild and Hertig (1958a) provide further information on collecting data, distri- bution, and taxonomy. Figure 29 Lutzonyia Vespertilonrs — A. Male head, R. Male flagellomere II, C. Female head, D. Female flagellomere II, E. Female ciharium and pharynx, G. Paramere, H. Paramere of male from Panama, I. Male geni- talia, J. Spermathecae, K. Female wing, L. Male wing, M. Female cibarium. Male: Rio Anori, Antioquia Dept. , Colombia (except Fig. 29H) Female: Same locality as male Lutzomyia isovespeptilionis — F. Paramere. Male: Panama Canal Zone -166- Cf :<:■?- f\ 0 D ~^^2rf> H t\it-'^7 ^ x^ V \. \ M w Ml. ;% /e =Z7 w \ ! \ -167- S ubgenus 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 vespsrti.li.onis group and mem- bers of the delpozoi group Lewis et al. (1973). The spermathecae of the Dampfomyia species are large with bubble-like expansions and could have been derived from the d&leoni-type spermathecae of the vespertilionis group. Like species in these two groups, ths Dampfomyia spp. do not ordinarily bite man and there are more species in Mexico and Central America than elsewhere. The preferred host(s) of L. vosabali, the only Dampfomyia sp . in Colombia or South America, is unknown. Related species such as L. anthophora (Addis) and L. permiva (Fairchild and Hertig) feed on small mammals, the former species probably transmitting Rio Grande virus to Neotoma woodrats in Texas (Calisher et al., 1977). 37. Lutzomyia (D.) rosabali (Fig. 30) Phlebotomus vosabali Fairchild & Hertig, 1956: 310 (6* holotype, Puerto Armuelles, Chiriqui Prov. , Panama; o, Guarare, Los Santos Prov., Panama). Rosabal, 1954: 10 et seq. (as sp . no. 9, Costa Rica). Fairchild & Hertig, 1959: 122 (Central American records). Rosabal & Trejos, 1964: 168 (cf . to dodgei) . Lutzomyia rosabali: Barretto, 1962: 95 (listed). Theodor, 1965: 193 (listed, spermatheca fig.). Forattini, 1971a: 100 (listed). Osorno et al., 1972a: 61 (Narino, Colombia). Christensen, 1972a: 88 (listed). Forattini, 1973: 208 (spermatheca fig.), 329 (as synonym of dodgei). •168- Distribution: Costa Rica, Panama, Colombia (Cauca, Narino). Material examined'. Colombia. 6 c5c?, 8 oo, Capuli (Narino), rock crevices, March 1945, M. Hertig. 3 go, Bolivar (Cauca), Jan. -Feb. 1944, J. Burbano. Panama. 1 o" (holotype no. 3209), type locality, tree but- tress, 9 June 1951, M. Hertig. 1 o (allotype), Guarare (Lcs Santos), light trap, 23 Oct 1952, F. Blanton. Discussion: Forattini (1973) treats L. rosabali as a junior synonym of L. dodgei (Vargas and Najera), believing, perhaps correctly, that the differences between the males are due to intraspecif ic geographic varia- tion. Without a larger sample of specimens from various localities, however, I prefer to consider them a distinct based on the. discussion and figures of Fairchild and Hertig (1956) . The Colombian males (Fig. 30) agree more closely with L. rosabali than with L. dodgei in the shape and setation of the dorsal arm of the paramere but the basal spine of the style is longer and more slender. The small bristle near the basal spine is absent or: one style of the rosabali hoiotype and from the styles of the Colombian males. Figure 30 Lutzomyia (D.j rosabali — A. Male head, B. Male flagellomere II, C. Female head, D. Female flagellomere II, E. Female cibarium and pharynx, Panamanian specimen, F. Male genitalia, G. Dorsal arm of puramere, H. Different views of paramere, I. Genital pump and filaments, J. Spermathecae, K. Female wing, L. Male wing, M. Female cibarium, Panamanian specimen. Male: Capuli, Narino Dept., Colombia Female: Same locality as male (except Fig. 30E and 30M) ■170- -171- Sub genus Pintomyia Costa Lima, 1932 The 5 described species in this subgenus are characterized by a row of short spines on the hind femur. Forattlni (1971a, 197 3) treats Pintomyia as a genus, placing, I believe, undue importance on these spines which, although not exactly the same, are shared by 2 species of Serge atomy ia Franca and 1 of Bnanptomyia. The Pintomyia females, like those of Pressatia, have well sclerotized individual sperm ducts but differ from them in other aspects (Theodor, 1955). At present, only L. spinosa is known to occur in Colombia. Lutzomyia damaseetioi (Mar.g.), a clos^ ally, was repoi'ted in Meta Department by Morales et al. (3 969a) but was excluded in a later work on the Colombian fauna (Osorno et al., 1972a) probably because of the uncertain identity of the specimens (2 oo) . Except for L. pessoai (Coutinho and Barretto) and L. fisokeri (Pinto) . the Pinto.nyia spp . have not been reported feeding on man. Lutzomyia gihsoni (Pifano and Ortiz), recently described from Venezuela from a single female, definitely belongs in the subgenus but its identify re- mains uncertain based upon the original description. 38. Lutzomyia (Pi.) spinosa (Fig. 31) Pklebntomv3 spinosus Floch & Abonnenc, 1942b: 1 (o* holotype, Gallion, near Cayenne, French Guiana). Barretto, 1947: 224 (full refs.). Hertig L Fairchild, 1950: 92, 93 (abdominal sternites 1 & 2, figs., Panama). Barretto, 1951: 224 (aistrib.). Floch & Abonneuc, 1952: 20 et seq . (a*, o, keyed, redescr. , figs.). -172- Blancas, 1959-1960: 129-130 (cf. to gorbitzi) . Johnson & Hertig, 1961: 765 et seq. (rearing data). Hanson, 1968: 80 (larva, pupa, descr., figs.). Morales et al . , 1969a: 381 (Meta, Colombia). Pifano & Ortiz, 1972: 31 (cf. to gibsoni) . Pifano et al., 1973: 153-154 (cf. to gibsoni) . Lutzonvjia spinosa: Barretto, 1962: 93 (listed). Martins et al., 1962a: 381 (Ooias, Brazil). Martins et al . , 1963: 335 (Brazilian records). Christensen & Fairchild, 1971: 302 (Darien Prov., Panama). Christensen, 1972a: 88 (listed). Osorno et al., 1972a: 60-61 (Colombian records). Ward et al., 1973: 178-181 (collecting data, Para, Brazil). Christensen & Herrer, 1973: 579 (Collecting data, Panama). Lewis, 1975a: 503 et seq. (mouthpart morphol.). Pintomyia spinosa: Forattini, 1971a: 104 (listed). Forattini, 1973: 123 et seq. (gen. review, figs.). Distribution: Panama, Colombia (Amazonas, Antioquia, Boyaca, Caqueta, Choco, Meta), French Guiana, Brazil. Material examined: Colombia. 70 6*0*, 12 09, 17 km W of Leticia (Amazonas), tree trunks, 24-26 July 1973, D.G.Y. & R.C.W. 1 6", Rio Anori (Antioquia) , light trap, 14 May 1970, C.H.P. 38 $3, 20 ^, same data but tree trunks, Sept. 1970, D.G.Y. 1 o, Curiche (Choco), Malaise trap, 7 July 1967, D.C.Y. Brazil. 11 6*0*, 1 o, Labrea (Amazonas), tree trunks, 9 Oct. 1972, D.G.Y. 2 dV, near Altamira (Para), tree trunks, 1 Oct. 1972, D.G.Y. 1 ^, Itiatuba (Para), tree trunk, 4 Oct. 1972, D.G.Y. Panama. 2 33 , 1 o, Cerro Campana (Panama) , tree buttresses. Discussion: The male parameres of spinosa are rather slender and the dorsal setae are distributed over the distal half of the structure. The coxite tuft consists of 6-8 setae. In L. domascenoi the parameres -173- are broader, slightly clubbed apically, and the dorsal setae are less numerous, being restricted to the distal third of the structure. The coxite tuft consists of 12+ setae. The females of the two species are very similar, but differ in the relative lengths of the sperm ducts and in the distribution of their sclerotization. The common sperm duct of spinosa is longer than 2 x the length of the individual ducts and the basal one-fourth of the latter is nonsclerotized, unlike damascenoi in which the individual ducts are almost completely sclerotized and the common duct is shorter, less than 2 x the length of the individual ducts. The female cibarium of L. gihsoni has well defined, long horizontal teeth as shown by Pifano and Ortiz (1972) . Those of spinosa and damascenoi are short and irregularly spaced. Adults of L. spinosa are commonly collected on tree trunks. Near Leticia (24-26 July 1973), this species was by far the most abundant sand fly taken in such resting sites. Figure 31 LutzomyLa spinosa — A. Male head, B. Male f lagellomere II, C. Female head, D. Female flagellomere II, E. Female cibarium and pharynx, F. Female hind femur showing spines, G. Male genitalia, H. Sperraathecae. I. Female wing, J. Male wing, K. Female cibarium. Male : Panama Canal Zone Female: Same locality as male -175- X ■n o / A f to I VO*Ot 1! W c \ ,:;-.i 0 C / w iV>/7 u 'I fe,y?4$0Z~\ ^V^l. xx: 7 4 / ■~vk / VI / A n o w. !s -176- Subgenus Pvessatic: Mangabeira, 1942 This snail distinctive group is represented in Colombia by 3 of the 6 known species Forattini (1971a, 1973) elevates the category to the genus level and adds several other species, which, in my opinion, belong in other groups (Young and Arias, 197 7; Lewis et al . , 1978). Except for L. oamposi on rare occasions, the Pressatia species do not feed on human blood. There are but slight differences among the 5 described females. Lutsomyia tviacantha , known from a single female in Colombia, may or may not be that species and confirmation of its presence there must await the discovery of the male. Keys to Species Males 1. Paramere as shown, with ca. 10 dorsally directed, blade-like setae at base oamposi (Fig. 32) Paramere otherwise, without blade-like setae 2 2. Coxite with patch of fine hairs above basal tuft. Paramere broader, its width at level of ventral acute process greater than width of lateral lobe tviacantha (Fig. 34) Coxite without fine hairs above basal tuft. Paramere more slender, its width at level of ventral acute process less than or = to width of lateral lobe dysponeta (Fig. 33) •17 7- 1. Individual sperm duct as long as or longer tnan common duct; basal 1/2 thin vailed, not well sclerotized camposi (Fig. 32) Individual sperm duct much shorter than common duct, thick walled, well sclerotized throughout most of its length . .dysponeta (Fig. 33) triacantha 39. Lutzorryia (Pp.) camposi (Fig. 32) Vh.lebotorn.us camposi- Rodriguez, 1952 (1950): 7 (o* holotype, Quevedo, Los Rios Prov., Ecuador). Fairchild & Hertig, 1958b: 205 (refs.). Fairchild & Hertig, 1959: 121 (Central American records). Arzube, 1960: 157 (Ecuadorian records). Hanson, 1961: 320 et seq. (breeding sites, Panama). Johnson & Hertig, 1961: 765, 773 (rearing data). Sherlock, 1962: 322 (Santander, Colombia). McConnell & Correa, 1964: 526 (in- fected with gregarines & fungi). Hanson, 1968: 52 (larva, descr., figs.) Phlebotomies acanthobasis Fairchild & Hertig, 1952: 508 (cf, o, Panama, figs.). Fairchild & Hertig, 1958b: 205 (as syonoym of camposi). Lutzomyia. camposi: Barretto, 1962: 95 (listed). Barreto, 1969: 463 (Valle, Colombia). Chaniotis et al . , 1971a: 345 et seq. (pop. dyna- mics, Panama). Osorno et al . , 1972a: 59 (Colombian records). Chaniotis et al., 1972: 95 (resting sites). Christensen, 1972a: 88 (listed). Christensen et al . , 1972: 57 et seq. (collecting data, Panama). Christensen & Herrer, 1973: 579 et seq. (collecting data). Rutledge & Mosser, 1975: 410 (biting man, Panama). Lewis, 1975a: 502 et seq. (mouthpart morphol.). Herrer & Christensen, 1976a: 62 (collecting data). -178- Pvessatia oamposi: Forattini. 1971a: 106 (listed). Forattlni, 1973: 123 et sec. (gen. review, figs.). Gomes, 1975: 8 (listed). Distribution: Costa Rica, Panama, Colombia. (Antioquia, Boyaca , Caldas, Choco, Santander, Valle), Ecuador. Material examined: Colombia. 26 (?, same locality, Shannon trap, 20 Aug. 1967, D.G.Y. 6 do*, 17 o<^, Teresita (Choco), light, Malaise, Shannon traps, tree trunks, March-Dec. 1967, D.G.Y. Panama. 6* holotype (no. 2012), type locality, 13 Dec. 1949, light trap, M. Hertig. p allotype (no. 2012), type local- ity, 15 Dec. 1949, M. Hertig. -180- Disaussion: In Panama, Thatcher (1968b) recovered dysponeta larvae from dead leaves and detritus 6+ meters above the ground in trees. Rutledge and Ellenwood (1975a), using soil emergence traps, collected 1 adult which had developed on the open forest floor. The immature stages have not been described. Observations on hosts include pigs (Fairchild and Hertig, 1952) , rodents (Christensen and Hertig, 1973), and horses (Rutledge and Mosser, 1975) . 41. Lutzomyia (Pi'.) triaoantha (Fig. 34) PklebotonvAS triacanthus Mangabeira, 1942a: 119 (, Rio Napo at Limoncocha (Napo) 21-24 May 1976, light and flight traps, D.G.Y. & T. Rogers. Discussion: The record of L. triaoantha from Venezuela is based on Pifano et al. (1962) who included the male in a key to the phlebotomines of chat country. No additional data were given. Without males, I am unable to separate the females of triacantka from those of dysponeta, although the species probably do not occur together in Colombia or elsewhere. Figure 32 Lutzomyia (Pr.) carrtposi — A. Male head, R. Male f lagellonere II, C. Female head, D. Female flagellomere II, E. Female cibarium and pharynx, F. Male genitalia, G. Genital pump and filaments, H. Sperma- thecae, slightly collpased, normal shape similar to those of Fig. 33H, I. Female wing, J. Male wing, K. Female cibarium. Male: Curiche, Choco Dept., Colombia Female: Same locality as male -183- vfely fl Figure 33 Lutzornyia (Pr. ) dysponeta — A. Male head, B. Male f lagellomere II, C. Female head, D. Female f lagellomere II, E. Female cibarium and pharynx, F. Male genitalia, G. Genital pump and filaments, H. Sperma- thecae, I. Female wing, J. Male wing, K. Female cihariurn. Male: Teresita, Choco Dept., Colombia Female: Same locality as male -185- •18( Baityi Group Theoclor, 1965 The two species comprising this group do not" show obvious affinities to other Lutzorrryia species and I prefer to separate them, unlike Forattini (1973) who places them in the subgenus Cororyia Barretto. The female of L. baityi remains unknown but it probably resembles that of L. gorbitzi (Blancas) figured by Blancas (1959-1960) , Fairchild & Hertig (1961b, as hansoni) , and Theodor (1965). At present, only L. baityi is known to occur in Colombia but populations of gorbitzi, discovered in Darien Prov., Panama, may extend into Choco Department. 42. Lutzorrryia baityi (Fig. 34) Phlebotomies baityi Damasceno, Causy, & Arouck, 1945: 22 (o^holotype, Cuiteua, Irituia, Para, Brazil). Barretto, 1947: 188-189 (listed). Damasceno et al . , 1949: 819 (Brazilian records). Floch & Abonnenc , 1952: 34 (o* keyed) . Fairchild & Hertig, 1961b: 2^6 (cf. to hansoni). Pifano et al . , 1962: 386 (d* in key to Venezuelan spp.). Leon, 1968: 31-32 (6" redescr., figs., Venezuela). Lutzomyia baityi: Martins et al., 1963: 334 (Roraima, Brazil). Theodor, 1965: 194 (listed, fig.). Forattini, 1971a: 101 (listed). Forattini, 1973: 274 et seq. (gen. review, fig.). Distributional Colombia (Boyaca) , Ecuador, Venezuela, Brazil. Material examined: Colombia . 1 cf [UV], San Pablo de Borbur (Boyaca), tree hole, 10 March 1945, A. Cast. Brazil. 1 cf, Boa Vista (Roraima), 16 Dec. 1960, J. Silva & L. Ferreira. Ecuador. 3 $$ \ Rio Napo at Limoncocha (Napo) , tree trunks, 18-22 May 1976, D.G.Y. & T. Rogers . -187- Disoussion: The single Colombian male of baityi, examined by me through the kindness of Dr. Pablo Earreto, is housed in the collection at the University of Valle, Faculty of Medicine, Cali, Colombia. Future search will probably reveal its presence in other localities within the Republic, especially those in Amazonas. Figure 34 Lutzormjia (~Pr.) triacantha — A. Male genitalia, B. Spermathecae, C. Female cibarium. Male: Limoncocha, Napo Prov., Ecuador Female: Puerto Lleras, Meta Dept., Colombia Lutzomyia baityi male — D. Genitalia. Male: Limoncocha, Napo Prov., Ecuador 189 -190- Subgenus Viar.no/rpjig Hangab eir a , 1941 This small distinctive group of four species is treated as a genus by Fcrattini (1971a, 1973). Three species occur in Colombia; the fourth, L. faviasi (Damasceno, Causey, and Arouck) , is known from Brazil and French Guiana. The preferred hosts of these species remain unknown. Specimens are observed usually on tree trunks, sometimes in burrows and at light. Keys to Species Males 1. Style with both proximal spines inserted on long arm or process. Parameres relatively slender 2 Style with proximal spines inserted on separate processes or tuber- cles. Parameres broader tvberculata (Fig. 36) 2. Parameres slender and downwardly turned apically. . fuvoata (Fig. 35) Parameres enlarged distally, shaped like a goat's head caprina (Fig. 35) Females 1. Spermathecae cylindrical, associated sclerotized processes funnel- shaped tuberoutata (Fig. 36) Spermathecae oval, sclerotized processes subequal in width throughout eaprina (Fig. 35) furcata -191- 43. Lutzomyic, (V.) oapvina (Fig. 35) LutsomyCa oapvina Osorno, Morales, & Osornn, 1972c: 437 (6* holotype, veroda El Llano, Municip. Victoria, Caldas, Colombia; p, El Terminal, Municip. Puerto Boyaca, Boyaca, Colombia). Osorno et al . , 1972a: 58 (Colombian records). Gomes, 1975: 8 (listed). Martins et al., 1976b: 496 (Peru) . Phlebotomus furcatus (not furcatus Mangabeira, 1941) : Fairchild, 1943: 571 (Panama records) . Phlebotomus arborealis (not arborealis Floch & Abonnenc, 1944) : Fairchild & Hertig, 1959: 121 (in part, Honduras & Panama records only). Johnson & Hertig, 1961: 765 et seq. (rearing data). Hanson, 1968: 48- 50 (larva, pupa, descr., figs.). Thatcher, 1968a: 296 (6*, in Disney trap) . Lutzomyia furcata (not furcatus Mangabeira, 1941): Osorno et al., 1970: 11 (Boyaca, Colombia). Christensen 5. Fairchild, 1971: 302 (Darien, Panama). Christensen, 1972a: 88 (listed). Christensen & Herrer, 1973: 579 (in light traps, Panama). Lutzomyia arborealis (not arborealis Floch & Abonnenc, 1944): Chaniotiset al., 1971a: 344 et seq. (pop. dynamics, Panama). Chaniotis et al., 19/2: 95 (resting sites). Distribution: Honduras _, Panama, Colombia (Antioquia, Boyaca, Caldas, Choco) , Peru. Material examined: Colombia. 1 o, Rio Anori (Antioquia), tree buttress, 6 Feb. 1970, C.H.P. 1 o, 1 g, same data but Sept. 1970, D.G.Y. 1 g, Curiche (Choco), 27 July 1967, D.G.Y. 1 g, Alto Curiche (Choco), tree buttress, 31 May 1967, D.G.Y. 1 o. same data but light trap 27 Aug. - 192 1967. 3 op, Teresita (Choco) , light trap & tree buttresses, May & Sept. 1967, D.G.Y. Honduras. 1 <$ , 4 oo, Tela, Lancetilla Valley, light traps 6 tree buttresses, Dec. 1953 & Jan. -Feb. 1954, V.'. Hils. Panama. 10 do', 20 go from various localities in the Canal Zone, Bocas del Toro Prov. , Color) Prov., Darien Prov., & Panama Prov., tree buttresses & light or Shannon traps. Discussion: See L. furcata. 44. Lutzomyia (V.) furcata (Fig. 35) Phlebotomus furcatus Mangabeira, 1941b: 256 (cf holotype, Aura, Belem, Para, Brazil). Barretto, 1947: 201 (refs.) . Barretto, 1950a: 104 (keyed). Barretto, 1951: 217 (Brazilian records). Floch & Abonnenc, 1952: 31 (keyed), 78-80 (c?, redescr., figs., French Guiana). Fauran, 1960: 8, 9 (mention. Forattini, 1960: 479 (Amapa, Brazil). Pifano et al., 1962: 38 3 (c? keyed) . Phlebotomus arborealis Floch & Abonnenc, 1944a: 8 (o, Baduel, French Guiana). Barretto, 1947: 186 (refs.). Floch & Abonnenc, 1952: 43 (keyed), 179-181 (redescr.. figs.). Fairchild, 1955: 190 (as probable o of furcatus). Fauran, 1960: 9 (mention). Lutzomyia fur cata: Barretto, 1962: 94 (listed). Martins et al., 1963: 334 (Roraima, Brazil). Martins et al., 1965: 3 (Rondonia, Brazil). Wijers & Huisenga, 1967: 397 (cf. to tuberculata) . Osorno et al., 1972a: 58-59 (Colombian records). Osorno et al . , 19 72c: 441 (cf. to caprina) . Shaw & Lainson, 1972: 713 (mention). Lewis, 1975a: 502 et seq . (mouthpart morphol.). Llanos et al., 1976: 480 (Peru). Lainson et al., 1977 (Mato Grosso, Brazil). -193- butzomyia arbovealis: Barretto, 1962: 94 (as synonym of furcata) . Viannamyia furcata: Forattini, 1971a: 106 (listed). Forattini, 19 73: 123 et seq. (in part, refs. based on material from Panama not furcata Mang., gen. review, figs.). Distribution: Brazil, French Guiana, Venezuela, Peru, Colombia (Caqueta, Santander) . Material examined: Colorrbia. 1 6* (INPES no. 3825), Vereda La Nevera, Solano (Caqueta), tree hole, 25 Aug. 1969. Brazil. 1 d1, 1 o, Serra do Navio (Araapa), tree trunks, 21 Sept. 1972, D.G.Y. & H. Fraiha. 1 if, 2 oo., Labrea (Amazonas) , tree trunk, 9 Oct. 1972, D.G.Y. 2 o'o", 2 cj>o , Rio Aripuana at Humboldt (Mato Crosso) , tree trunks, 17-20 Aug. 1974, D.G.Y., J. Shaw, & II. Fraiha. 1 6*, Belem (Para), 3 Aug. 1964, F. Guimaraes. 1 o, same locality, light trap, 18 Aug. 1972, D.G.Y. 2 o^, same locality, flight traps, 29 July-6 Aug. 1974, D.G.Y. 1 ^, Itiatuba (Para), light trap, 4 Oct. 1972, D.G.Y. 1 o. , 27 km E of Maraba (Para), light trap, 27 Setp. 1972, D.G.Y. & H. Fraiha. 1 d", 1 £, W of Altamira, km 164 (Para), tree trunk, 9 Nov. 1974, J. Reinert. Discussion: The distal part of the parameres of L. furcata (de- scription of Mangabeira, 1941b, and males from Brazil, French Guiana, and Caqueta, Colombia) is long and slender, downwardly curved with a short, stout apical spine (Fig. 35G) . In contrast, the parameres of L. caprina (description of Osorno et al . , 1972c, and male from Antioquia, Colombia) are broader apically and lack a stout terminal spine (Fig. 35H) . The occurrence of caprina in Loreto Prov. , Peru (Martins et al . , 1976b) suggests that it and furcata are distinct taxa. The females, however, from these and all other localities are morphologically inseparable, the record of fui'cata (o only) in Antioquia Dept . (Osorno et al., 1972a) probably representing not that species but L. caprina. -194- Without additional material from Central America, the identity of the specimens from Panama and Honduras remains uncertain. Panamanian males generally resemble capr-ina in the shape of the parameres but each terminates in an apical spine, longer than that of fuTCata. The male from Honduras, on the other hand, more closely resembles furcata in paramere shape but again the apical spine is much longer. It is dif- ficult to determine whether or not these character states reflect inter or intraspecific variation but for the present I arbitrarily treat specimens from Panama and Honduras as variants of caprina, primarily on the basis of distribution. Lutzomyia furcata was included in a key to the phiebotomines of Venezuela (Pifano et al . , 1962) but no additional information was given. 45. Lutzomyia (V.) tuberoulata (Fig. 36) P'hlebotomus tuberculo.tus Mangabeira, 1941b: 251 (6* holotype, Aura, Belem, Para, Brazil). Barretto, 1947: 227 (refs.). Damasceno et al., 1948: 699 (cf. to f arias i) . Damasceno et al . , 1949: 833 (Brazilian records). Barretto, 1950a: 104 (keyed). Barretto, 1951: 225 (distrib.) Floch & Abonnenc, 1952: 31 (keyed). Fauran, 1960: 8-9 (6", redescr., figs.). Phlebotomus sp. x Floch & Abonnenc, 1944a: 10-12 (o, French Guiana). Floch & Abonnenc, 1952: 43 (keyed), 181 (redescr., figs.). Barretto, 1962: 94 (as female of tubex'cu'lata) . Lutzomyia. tuberoulata: Barretto, 1962: 94 (listed). Martins et al., 1963: 335 (Roraima, Brazil). Theodor, 1965: 190 (6*, £, figs.). Barreto, 1969: 467 (Valle, Colombia). Lewis et al . , 1970: 215 (ovaries -195- & accessory glands). Christensen 1972a: 38 (Panama). Osorno et al., 1972a: 59 (Colombian records). Lewis, 1975a: 502 et seq . (mouthpart morphol.). Lainson et al., 1977 (Mato Grosso, Brazil). Lutzomyia munangai Wijers & Huisenga, 1967: 395 (d1 holotype, near Moengo, Surinam), Lewis, 1975a: 511 (as synonym of tubsroulata) . Viannamyia munangai: Forattini, 1971a: 106 (listed). Forattini, 1973: 516 (cf. to tuberculata.) . Gomes, 1975: 8 (listed). Viannamyia tuberculata: Forattini, 1971a: 106 (listed). Forattini, 1973: 507 et seq. (gen. review, figs.). Gomes, 1975: 8 (listed). Distribution:. Panama, Colombia (Antioquia, Caqueta, Choco, Valle), French Guiana, Surinam, Brazil. Material examined: Colombia. 1 o, Rio Anori (Antioquia), tree trunk, 4 March 1971, C.II.P. 1 £, Curiche (Choco), Malaise trap, 25 July 1967, D.G.Y. 2 00, Alto Curiche (Choco), 7 July 1967, D.G.Y. 1 o, Anchicaya + T "f Dam (Valle), tree trunk, 9 Aug. 1973, D.G.Y. & R.C.W. 6 JcT, 8 00, 25 km E of Buenaventura (Valle), tree trunks & light trap, 11-12 Aug. 1973, D.G.Y. & R.C.W. Brazil. 2 op, Serra do Navio (Amapa) , tree trunks, 21 Sept. 1972, D.G.Y.& H. Fraiha. 1 0, W of Altamira (Para), light trap, 1 Oct. 1972, D.G.Y. 2 00, Belem (Para), light trap, 18 Oct. 1972, D.G.Y. 2 061. 5 00, Belem (Para), tree trunks, flight trap, 29-31 July, 1974, D.G.Y. 3 0*0*, 2 00, Rio Aripuana at Humboldt (Mato Grosso), tree trunks, 16-19 Aug. 1974, D.G.Y., J. Shaw, & H. Fraiha. Panama. 2 00, Almirante (Bocas del Toro) , Aug. -Sept. 1951, A. Quinones. Discussion: Lutzomyia tuberculata, a little known species, is not considered to be anthropophilic although Barreto (1969) reported the capture of a single female on man in Valle Dept., Colombia. Both sexes are easily recognized. Figure 35 Lutzomyia (V.) oaprina — A. Male head, B. Male flagellomere II, C. Female head, D. Female flagellomere II, E. Female cibarium and pharynx, F. Male genitalia, H. Tip of paramere of male from Rio Anori, Antioquia Dept., Colombia, I. Genital pump and filaments, J. Sperma- thecae, K. Female wing, L. Male wing, M. Female cibarium. Male: Panama Canal Zone (except Fig. 35H) Female: Curiche, Choco Dept., Colombia Lutzomyia (V. ) furcata — G. Tip of paramere. Male: Para State, Brazil •197 "X Figure 36 Lutsonyia (V.) tubevoulata — A. Male head, B. Male flagellomere II, C. Female head, D. Female flagellomere II, E. Female cibarium and pharynx, F. Male genitalia, G. Spermathecae, H. Female wing, I. Male wing, J. Female cibarium. Male: Valle Dept., Colombia Female: Curiche, Choco Dept., Colombia -199- -200- Shannoni Group Theodor, 1965 Barret to (1962) created the subgenus Psathyromyia to accomodate L. shanncmi and rits allies, some of which (the aragaoi group species) were separated later by Theodor (1965) . Rased on both sexual and non- sexual characters, the shannon!, and aragaoi species are very similar as noted by Fairchild (1955) and Barretto (1962). Forattini (1971a, 1973), for reasons not fully understood, places the species in the aragaoi group in the genus Psychodopygus but retains L. channoni and its relatives in the genus Lutzorryia in the expanded subgenus Trickopygomyia Barretto. This seems to be unnecessarily arti- ficial ana I prefer to follow the arrangement of Theodor (1965) and Lewis et al. (1978). Six of the 14 described shannoni group species occur in Colombia but others such as L. lutziana (Costa Lima), L. campballi (Damascene, Causey & Arouck) , and possibly I. soaffi (Damascene & Arouck) may eventually be discovered in the Amazonian region of the Republic. Keys to Species Males 1. Paramere with a basal tuft of long setae implanted on a short dorsal process dasyrrera (Fig. 33) Paramere simple, lacking a setiferous dorsal process 2 2. Antennal ascoids with short, blunt posterior spurs punetigenieulata (Fig. 40) Antennal ascoids with long pointed posterior spurs 3 -201- 3, Paramere with sinuous dorsal setae undulata (Fig. 42) Paramere without such dorsal setae 4 4. Paramere with dorsal setae restricted to apical third of structure . , abonnenci (Fig. 37) Paramere with dorsal setae covering at least the apical half of structi i 5. Paramere us in Fig. 39A. The apical dorsal setae slightly foliaceous and recurved unlike the shorter proximal setae, dendrophyla (Fig. 39) Paramere otherwise (Fig. 41G) , the dorsal setae unmodified, more or less erect shannoni (Fig. 41) Females 1. Cibarium with 6+ horizontal teeth . 2 Cibarium with 4 horizontal teeth* shannoni (Fig. 41) dendrophyla (Fig. 39) abonnenci 2. Spermathecae annulated undulata (Fig. 42) Spermathecae smooth-walled, without annulations 3 *Morales et al. (1974) noted that the common sperm duct of dendrophyla is shorter than that of shannoni (0.049-0.099 mm as opposed to 0.145- 0.198 mm). This may be helpful in separating the taxa but without associated males I nesitate to identify females to species. -202- 3. Antennal ascoid wich short blunt posterior spur. Individual sperm duct, about half length of common duct and less than 2 >: length of spermatheca punctigenioulata (Fig. 40) Antennal ascoid with long pointed posterior spur. Individual sperm duct much longer than common duct and over 3 x length of spermatheca dasymsra (Fig. 38) 46. Lutzomyia abonn&noi (Fig. 37) Fhlebotomus abonnenoi Floch & Chassignet, 1947c: 1 (, 17 km W of Leticia (Amazonas), tree trunks, 24-26 July 1973, D.C.Y. & R.C.W. Bolivia. 1 6\ 1 o, 5 km N of Tajlewi (Larecaja) , tree trunks, 13 Aug. 1971, J. Velasco. 1 d", 1 o, Misquirnayo (Sud Yungas) , chicken coop, 1700 m above sea level, 3 Sept. 1971, J. Velasco. Brazil. (all on tree trunks). 5 c?cf, Serra do Kavio (Amapa), 21 Sept. 1972, D.G.Y. & H. Fraiha. 12 6"d\ Labrea (Amazonas), 9-10 Oct. 1972, D.C.Y. 72 o'c?, Rio Aripuana at Humboldt (Mato Grosso), 13-20 Aug. 1974, D.G.Y. , J. Shaw, & H. Fraiha. 3 S3, 27 km -206- SE of Maraba (Para), 26-28 Sept. 1972, D.G.Y. & H. Fraiha. 1 6*, W of Altamira (Para), 1 Oct. 1972, D.G.Y. 5 6*0*, Belern (Para), 29 July-1 Aug. 1974, D.G.Y. Ecuador. 90 cf6*, Rio Napo at Limoncocha (Napo) , tree trunks, 19-24 Kay 1976, D.G.Y. & T. Rogers. French Guiana. 1 6*, Baduel, Dec. 1947, no other data. Peru. 2 6*6*, Cachicote (Huanuco) , 12 Sept. 1965, J. Hitchcock. Discussion: Lutzormjia dendrophyla is an Amazonian species which occurs often but not always with L. shannoni where their ranges overlap. We collected only dendrophyla at Leticia, the spermathecae (Fig. 39B) drawn from a female taken from there. The females of both species occasionally bite man (Lainson et al . , 1977; Forattini, 1973) and are very difficult to separate in the absence of associated males. 49. Lutzomyia puna tig eniculata (Fig. 40) Phlebotomus punctigeniculatus Floch & Abonnenc, 1944b: 5 (6* holo- type, near Cayenne, French Guiana). Barretto, 1947: 220 (refs.). Floch & Abonnenc, 1948b: 1, 12, 24 (Venezuela, 0*, figs.). Damasceno et al . , 1949: 831 (Brazilian records). Fairchild & Hertig, 1950: 524 et seq. (measurements, taxonomy, figs.). Barretto, 1951: 222 (distrib.). Pifano & Ortiz, 1952: 144, 146 (keyed). Floch & Abonnenc, 1952: 37, 46 (keyed), 153 (6*, o, redescr.). Barretto & Duret, 1953: 344 (mention) Pifano et al., 1962: 385, 388 (6", o, keyed), 397 (6*, o, redescr., refs., figs., Venezuela). Ortiz, 1965c: 416 (mention). Leon, 1968: 30 (listed). Ortiz, 1972a: 23 (mention). Pifano et al., 1973: 158 (mention) . 207- Phlebotomus chrisiophersoni Damasceno & Causey, 1944: 347 (25- 52. Lutzomyia longispina (Fig. 43) Phlebotomus longispinus Mangabeira, 1942a: 186 (c? holotype, Aura, Belem, Para, Brazil). Mangabeira, 1942d: 251 (iramatures descr.). Barretto, 1947: 209 (catalog). Causey & Damasceno, 1948: 628 et seq . (cf. to wagleyi, 6* fig.). Damasceno et al . , 1949: 827 (Brazil). Vargas & Diaz-Najera, 1951a: 22 (cf. to pratti) . Pifano & Ortiz, 1952: 142 (Bolivar, Venezuela). Floch & Abonnenc, 1952: 33, 48 (cf, y keyed). Fairchild & Her tig, 1952: 518 (cf. to triramulus) . Pifano et al . , 1962: 385, 389 (cf, o keyed). Sherlock & Pessoa, 1964: 333 (Bahia, Brazil). Hanson, 1968: 88 (larva, cf. to triranulus) . Lutzomyia longispina: Barretto, 1962: 98 (listed). Theodor, 1965: 189 (d", figs.). Martins et al . , 1965: 14 (cf. to rondoniensis) . Osorno et al . , 1972a: 57 (Caqueta, Colombia). Forattini, 1973: 172 et seq. (gen. review, figs.). Lewis, 1975a: 502 et seq. (mouthpart morphol.). Ramirez et al., 1976: 602 (mention). Martins et al . , 1976a: 491 (cf. to elegans) . Distribution: Brazil, Venezuela, Colombia (Caqueta) . Material examined'. Colombia. 1 o (INPES no. 7068), San Miguel, Municip. Florencia (Caqueta), tree hole, 2j June 1971, E. Osorno & A. Morales. Brazil. 1 6*, Cacheira (Bahia), armadillo burrow, 7 July 1959, Afonso. 1 29- Figure 44 Lutzomyia trtvamula — A. Male head, B. Male flagellomere II, C. Female head, D. Female flagellomere II, E. Female cibarium and pharynx, F, Female cercus, G. Genital pump and filaments, same scale as Fig. 44H, H. Male genitalia, I. Spermathecae, J. Female wing, K. Male wing, L. Female cercus, specimen from Valle Dept., Colombia, M. Femalei cibarium. Male: Curiche, Choco Dept., Colombia Female: Same locality as male (except Fig. 44L) -23J V \ XSA -232- Avagaoi Croup Theodor, 1965 Theodor (1965) divides these species into 2 categories, the series avagaoi (7 spp.) and series bvasitiensis (4 spp.), based upon the sperma- thecae which are sac-like in the series avagzoi females, tubular in those of the bvasiliensis series. Fairchild (1955), Barretto (1962), Ortiz (1972a) , and others combine the avagaoi group species with those in the sharmoni group on the basis of several characteristics, the latter two authors placing them in the subgenus Psatkyvonyla Barretto, 1962. The status of L. pifa.noi (Ortiz), a Venezuelan species is not yet clear. It certainly belongs in the aragaoi group but is known from a single male which somewhat resembles L. vv.noid.es . Lutzomyia abunaensis Martins, Falcao and da Silva (1965), from Brazil and Ecuador, also described from the male, is similar to that of L. bvasiZiensis (Costa Lima, 1932) . I examined a male and several females of abunaensis , to be described in another paper, from Limoncocha, Napo Prov., Ecuador. The females differ from those of bvasiliensis in having 8+ horizontal cibarial teeth instead of 4 and by the shorter sperm ducts. The spermathecae of both females are similar in shape and size. The presence of one or both of these species in Colombia would not be surprising. Keys to Species Males 1. Genital filament tips hooked or twisted 2 Genital filament tips simple, pointed, or spear shaped .3 -233- 2. Style with isolated basal spine. Genital filament tips shaped like a burton hook. Aedeagus rao.re slender. Paramere as shown . . carpenteri (Fig. 48) Style with paired or nearly paired basal spines. Genital filament tips twisted like a corkscrew. Paramere variable but never as above aragaoi (Fig. 45) 3. Genital filament tips markedly inflated, spear-shaped. Coxite without scattered setae on inner surface runoides (Fig. 49) Genital filament tips simple, not inflated. Coxite with scattered setae on inner surface 4 4. Lateral lobes greater than 0.56 mm. Parameres as shown, the ventral "heel" better developed barrettoi majuscula (Fig. 4 6) Lateral lobes less than 0.50 mm. Parameres otherwise, more slender, lacking a ventral heel bavrettoi barrettoi (Fig. 47) females 1. Spermathecae tubular with convoluted individual ducts runoides (Fig. 49) Spermathecae sac-like, much wider than nonconvoluted ducts, .... 2 2. Spermathecae pear shaped with an expanded neck, individual ducts with fine transverse striations carpenteri (Fig. 48) Spermathecae otherwise, ovoid or spherical, individual ducts saiooth walled 3 -234- 3. Spermathecae smaller, subspherical with apical "hairs" not arising from a differentiated area. . . , avagaoi (Fig. 45) Spermathecae larger, ovoid with apical "hairs" arising from a raised circular area 4 4. Spermathecae larger, their ducts expanded where they enter sperma- thecae bawettoi bavvettoi (Fig. 47) Spermathecae smaller, their ducts more slender, subequal in width throughout barrettoi majuscula (Fig. 46, 4 7) Series avagaoi 54. Lutzomgia avagaoi, (Fig". 45) Phlebotomus avagaoi Costa Lima, 1932: 48 (6" Lassance, Minas Gerais, Brazil). Barretto, 194 7: 185-186 (full refs.). Floch & Abonnenc, 1952: 131-134 (in part, c? only) . Fairchild & Hertig, 1953a: 21 et seq. (o", o, redescr., figs., keyed, refs., Panama). Vargas & Diaz-Najera, 1959: 146 (cf. to humboldti). Fairchild & Hertig, 1961b: 250 (o. cf. to botella) ■ Lucena & Almeida, 1964: 187 et seq. (cf, o redescr., figs.). Sherlock & Pessoa, 1964: 333 (resting sites, Brazil). Lucena, 1967: 271 (mention). Ortiz, 1972a: 23-25 (listed, cf. to pifanoi) . Phlebotomus keckenvothi Floch u AbonneiiCj 1942b: 8 (o, French Guiana) Barretto, 1946c: 430 (listed). Floch & Abonnenc, 1952: 47, 176 (c. keyed, redescr., figs.). Fairchild & Hertig, 1953a: 21 et seq. (o keyed, cf. to avagaoi, refs.). Ortiz, 1972a: 23 (listed). Pifano et al . , 1973: 158 (listed). Forattini, 1973: 456 (as synonym of avagaoi)- -235- Lutzomyia aragaoi: Barretto, 1962: 99 (listed). Theodor, 1965: 186 ((?, o, figs). Martins et al., 1965: 2 (Rondonia, Brazil). Barreto, 1969: 462-463 (Valle, Colombia). Lev/is et al., 1970: 215 (parous rates). Chanictis et al., 1971a: 344 et seq. (collecting data, Panama). Christensen, 1572a: 88 (listed). Osorno et al., 1972a: 37 (Vichada, Colombia). Chcmiotis et al., 1972: 95 (resting sites). Lewis, 1975a: 501 (inouthpart morphol., cibarium fig.). Martins et al . , 1976a: 488 (Peru) . Lutzomyia heckenrothi: Barretto, 1962: 99 (listed) . Martins & Morales, 1972: 367 (listed). Psychodopygus aragaoi: Forattini, 1971a: 105 (listed). Forattini, 1973: 416 et seq. (gen. review, figs.). Distribution: Panama, Colombia (Amazonas, Antioquia, Choco, Valle, Vichada), Peru, Brazil, Paraguay, French Guiana, Trinidad. Material examined: Colombia. 1 6", 17 km W of Leticia (Amazonas) , light trap, 26 July 1973, D.G.Y. & R.C.W. 2 66, 1 g, Rio Anori (Antioquia) 2-5 May 1970, C.H.P. 7 ^o., same data but 20-22 Sept. 1970, D.G.Y. 1 6, same data but 14 April 1971, C.H.P. Id1, Tres Esquinas (Caqueta) , light trap, 10 Nov. 1971, C.J.M. 52 do', 70 ^>§, Curiche (Choco), light h Malaise traps, April-Nov. 1967, D.G.Y. 2 oo., Alto Curiche (Choco), light trap, 2 July 1967, D.G.Y. 2 do", 2 ^o, Anchicaya Dam (Valle), light traps, 10-11 Aug. 1973, D.G.Y. & R.C.W. 1 o., 25 km E of Buenaventura (Valle), light trap, 12 Aug. 1973, D.G.Y. & R.C.W. Brazil, French Guiana, Panama, and Paraguay. Those specimens listed by Fairchild & Hertig (1953a). Trinidad. 50+ 66, 50+ oo, from various localities to be treated in a forthcoming paper. •236- Discussion: According to locality, the Tiales of L. aragaoi vary in the shape of the parameres. Those seen by me or figured by "•(hers from Brazil (Costa Lima, 1932), Colombia (Rio Anori, near Leticia, and Anchicaya Dam), Panama (Fairchild & Hertig, 1953a), and Peru (Lucena & Almeida, 1964) have relatively slender parameres compared to those from French Guiana (Floch & Abonnenc, 1952), Trinidad, and Colombia (Choco Dept. and Tres Esquinas in Caqueta Dept.)- Whether clinal or not, I believe that this variation is intraspecif ic, the other character states of both sexes being remarkably similar. Like Forattini (1973), I consider L. heskenrothi (Floch & Abon- nenc, to be the female of L. aragaoi. Floch & Abonnenc' s subsequent figure (1945c) shows long sperm ducts with warty protuberances, as noted by Fairchild & Hertig (1953a) , but this probably represents an abnormal specimen, a poor slide mount or, less ]ikely, another species. 55. Lutzomyia bavrettoi bavvettoi New Status (Fig. 47) Phlebotorrrus bavrettoi Mangabeira, 1942a: 148 (o* holotype, Belem, Aura, Para, Brazil). Barretto, 1947: 189 (refs.). Floch & Abonnenc, 1952: 38 (c? keyed), 144 (6* redescr . , figs.). Fairchild & Hertig, 1953a: 26 (refs.). Fairchild & Hertig, 1959: 121 (in part, Brazil & French Guiana). Vargas & Diaz-Najera, 1959: 146 (cf. to humboldti) . Sherlock & Carnheiro, 1964: 206, 208 (internal morphol., (7, fig.). Rosabal, 1966: 1-2 (in part, distrib.). Ortiz, 1972a: 23-24 (listed). Phtebotomus sp. de Haripa Floch & Abonnenc, 1946: 3 (o, French Guiana). Floch & Abonnenc, 1952: 46, 191 (o keyed, redescr., figs.). -237- Lutzomyia barrettoi: Barretto, 1962: 99 (listed). Martins et al., 1976c-: 483 (Peru). Psychodopygus barrettoi: Forattini, 1971a: 105 (listed). Forattini, 1973: 416 et seq. (in part). Distribution: Colombia (Caqueta), Peru, Brazil, French Guiana, Trinidad. Material examined: Colombia. 2 , Tres Esquinas (Caqueta), light trap. 10-11 Nov. 1971, C.J.M. Brazil. 1 d", 0 , Mangabeira, no. 1075, no other data. Trinidad. 1 o, Macqueripe U.S. Naval Station, light trap, 19 Nov. 1956. T.H.G. Aitken. 1 6", Tucker Valley U.S. Naval Station, light trap, 14 Nov. 1957, T.H.G. Aitken. 1 tf, San Jose Point U.S. Naval Station, light trap, 18 Nov. 1957, T.H.G. Aitken. Discussion: Specimens of L. barrettoi from east of the Andes in Colombia (Caqueta), Brazil, French Guiana, Trinidad, and probably Peru agree closely with the descriptions of Mangabeira (1942a) and Floch & Abonnenc (1946, 1952, as sp. de Maripa) . I am convinced that Lutzomyia sp. de Maripa is the female of L. b. barrettoi based upon morphology and collecting data in Trinidad and Caqueta Dept., Colombia. The spermathecae are large and oval, their ducts subequal in width throughout (Fig. 47B) . Other features of diagnostic importance are discussed under L. barrettoi majuscula n. ssp. 56. Lutzomyia barrettoi majuscula. n. ssp. (Fig. 46, 47) Phlebotorrus barrettoi: Fairchild 6 Hertig, 1953a: 21 et seq. (d\ p, descr., keyed, figs.). Rodriguez, 1956: 79 (Ecuador). Fairchild & Hertig, 1959: 121 et seq. (in part, Central American records). Fairchild -238- & Hertig, 1961b: 250 (cf. to botella) . Johnson & Hertig, 1961: 765 (rearing data). Rosahal & Trejos, 1964: 167 (El Salvador). Rosabal, 1966: 1--2 (Costa Rica). Hanson, 1968: 51 (first instar larva, descr., tig.). Fklebotomus sp. no. 6 Rosabal, 1954: 6, 46 (Costa Rica). Lutzomyia barrettoi: Barreto, 1969: 463 (Valle, Colombia). Chaniotis et si., 1971a: 344 et seq. (collecting data, Panama). Christensen, 1972a: 88 (listed). Christensen et al., 1972: 57 (in animal burrow, Panama). Chaniotis et al., 1972: 95 (resting sites). Osorno et al., 3 972a: 38 (mention). Psyehodopygus barrettoi Forattini, 1973: 416 et seq. (in part). Distribution El Salvador, Nicaragua, Costa Rica, Panama, Colombia (Antioquia, Choco, Valle) , Ecuador. Material examined: Panama. 1 6* (holotype no. 6320), Cruces Trail, Madden Forest Reserve, Canal Zone, animal burrow, 6 Jan. 1957, M. Hertig. 1 c (allotype no. 4732), Camp Pina, Canal Zone, light trap, 24 May 1954, F. Blanton. 1 6" (paratype no. 4200), San Lorenzo, Canal Zone, light trap, 15 Aug. 195.2, F. Blanton. 1 o (paratype no. 3914), Mojinga Swamp, Canal Zone, light trap, 31 Jan. 1952, F. Blanton. 1 o (paratype no. 4735), Canal Zone, light trap, 29 June 1954, G. Field. 3 $S , 4 jo. (paratypes nos. 270-276), Cerro Azul (Panama), Burrow, 24 March 1968, D.G.Y. Colombia. 1 o (paratype no. 277), Rio Anori (Antioquia), light trap, 22 Sept. 1970, D.G.Y. 1 cf (paratype no. 278) Alto Curiche (Choco), light trap, 7 July 1967, D.G.Y. 1 o (paratype no. 279), Curiche (Choco), Malaise trap, 29 June 1967, D.G.Y. Costa Rica. 1 6* (paratype no. 3447), Suerre de Guapiles, tree buttress, 29 April 1951, R. Rosabal. Ecuador. 1 6* (paratype no. 280), 17 km E of Santo Domingo de Los Colorados -239- (Pichincha), light trap, 27 May 1976, D.G.Y., T. Rogers, & G. Fairchild. 1 o (paratype no. 281), same data but May 1976. Nicaragua. 1 cf (para- type no. 4457), Guapilonar, Carrazo, light trap, 21 Oct. 1953, A. Adames. Other material examined: Colombia. 1 d", 5 09, Curiche (Choco) , light & Malaise traps, 5 July-28 Aug. 1967, D.G.Y. 1 o., 25 km E of Buenaventura (Valle), light trap, 12 Aug. 1973, D.G.Y. & R.C.W. Discussion: Male: As described and illustrated by Fairchild & Hertig (1953a, as L. baivettoi) with the following additions based upon the holotype of L. b. majuscula n. ssp. Wing length 2.47; width 0.78. Head height 0.38; width 0.44. Eyes separated by 0.10 or by distance = to 4.6 facet diameters. Flagellomere I (0.30 long); combined length of f lagellomeres II + III = 0.38. Length of palpal segments: 1 (0.03), 2 (0.08), 3 (0.12), 4 (0.07), 5 (0.16). Labrum 0.20 long. Cibarial arch nearly complete; pigment patch distinct, subtriangular. Pharynx 0.20 long. Pleura with 13 upper and 4-5 lower episternal setae. Length of wing vein sections: Alpha (0.80), beta (0.22), delta (0.42), gamma (0.19). Length of femora, tibiae, and basitarsi: Foreleg, 1.06, 1.56, 0.95; midleg, 0.98, 1.66, 0.98. Hindleg missing. Genitalia: Length of style 0.28; coxite 0.49; lateral lobe 0.60; genital pump 0.21, filaments 0.78 (the latter 3.7X length of pump) . Female: As described and illustrated by Fairchild & Hertig (1953) with the following additions based upon the allotype of L. b. majuscula n. ssp. Wing length 2.60; width 0.88. Head height 0.42; width 0.46. Eyes separated by 0.45 or by distance = to 5 facet diameters. Flagello- mere I 90.29 long); combined length of f lagellomeres II + III = 0.37. Length of palpal segments: 1 (0.03), 2 (0.09), 3 (0.14), 4 (0.07), 5 -240- (0.17). Cibarium with 15 horizontal teeth, about 20 vertical teeth in 2-3 transverse rows, the 7 closest to horizontal teeth markedly enlarged; cibarial arch nearly complete, pigment patch prominent as shown by Fairchild & Hertig (1953a, fig. 21). Labrum 0.19 long. Pharynx 0.20 long. Pleura with 15-16 upper and 4-5 lower episternal setae. Length of wing vein sections: Alpha (0.88), beta (0.22), delta (0.51), gamma (0.19). Legs missing. Spermathecae and ducts as figured. Although larger than L. b. barvettoi , the female of L. b. ma jus aula has smaller spermathecae and the ducts are noticeably swollen where they enter the spermathecae (Fig. 47A) . The tip of the genital fork stem, sha.llowly bifid in majusQula, is variable in b. bavvettoi (Fig. 47C-E). Without females or locality data, the males are difficult to distinguish but the parameres of majusaula are slightly broader with a better developed ventral "heel." The lateral lobes of this taxon range between 0.57-0.61 mm in length; those of b. bavrettoi are shorter, not exceeding 0.50 mm. The wing length of majuscula 8<3 ranges from 2.3 to 2.7 mm; go from 2.5 to 2.8 mm. The wing length of b. bavrettoi 6o ranges from 1.8 to 2.2; go from 2.1 to 2.4. The subspecific name refers to the larger size of the majuseula specimens. This subspecies like L. eavvevai thula n. ssp., occurs west of the Andes in Ecuador and Colombia, then northward into Central America to El Salvador. Sherlock & Carnheiro (1964) suggest that L. texana (Dampf) from Texas (U.S.A.), Mexico, and Honduras (unpub . data) and L. barvettoi may be conspeeific, basing their interpretation on the description of texana by Dampf (1938) . While examining numerous specimens of the former species (Young, 1972b), 1 noted that the males of the two species can not -241- be separated by the key given by Fairchild & Hertig (1953a)., the styles of both species with the isolated spine set upon a well marked tubercle doner to the basal spines than to the terminal spine. They differ, however, in the relative lengths of the genital filaments (less than 2.6 r. length of the pump for texana.; 2.8 x or greater for the barrettoi subspecies) . The scattered hairs on the inner surface of the coxites are long in texana but rather short in barrettoi. The females are more easily distinguished, as was done by Fairchild & Hertig (op. cit., in key) and by the nearly spherical, smaller spermathecae of texana. (Fig. 471). 57. Lutzomyia earpenteri (Fig. 43) Phlebotomus earpenteri Fairchild & Hertig, 1953a: 28 (6* holotype, Chiva Chiva, Canal Zone; o, near Pacora, Panama Prov. , Panama). Fairchild & Hertig, 1959: 121-122 (Central American records). Fairchild & Hertig, 1961b: 2.50 (cf. to botella) . Johnson & Hertig, .1961: 765, 773 (rearing data). Biagi et al . , 1966: 141 et seq. (d", p, figs., Mexico). Hanson, 1968: 53 (first instar larva, descr., fig.). Ortiz, 1972a: 23-24 (listed) . Lutzomyia aarpenteri.: Barretto, 1962: 99 (listed). Williams, 1970: 332 et seq. (summary of collecting data, Belize). Tesh et al., 1971a: 153 (blood meal from bird). Chaniotis et al., 1971a: 344 et seq. (col- lecting data, Panama) . Christensen, 1972a: 88 (listed) . Chaniotis et al., 1972: 95 et seq. (resting sites). Shaw & Lainson, 1972: 709 (mention). Christensen et al . , 1972: 57 et seq. (collecting data, nega- tive for flagellates, Panama). Christensen & Herrer, 1973: 579 et seq. (collecting data, negative for trypanosomatids) . Chaniotis & Correa, -2A2- 1974: 115 (light trap data). Lewis, 1975a: 501 et seq. (mouthpart morphol.)- Rutledge & Ellenwood, l?75a: 73 (breeding site, open forest floor, Panama). Rutledge et al . , 1975: 179 et seq. (light trap data). Williams, 1976a: 604 (in caves, Belize). Psyehodopygus carpentevi: Forattini, 1971a: 105 (listed). Forat- tini, 1973: 390 et seq. (gen. review, figs.). Distribution: Mexico, Belize, Costa. Rica, Panama, Colombia (Antioquia, Choco) . Material examined'. Colombia. 72 6*cf, 10 on, Rio Anori (Antioquia), light traps, May 1970, C.H.P. 12 cfc?, 25 o^, same data but Sept. 1970, D.C.Y. 29 eft?, 33 go, Curiche (Choco), light, Shannon, & Malaise traps, April-Dec. 1967, D.G.Y. 1 o, Teresdta (Choco), Malaise trap, 22 April 1967, D.G.Y. 1 o", same data but light trap, 29 Sept. 1967. 1 o, same data but 28 Oct. 1967. Panama. 1 6* (holotype no. 1345), Chiva Chiva, Canal Zone, light trap, Oct. 1948, S. Carpenter. 1 c- (allotype no. 2301) near Pacora (Panama Prov.), burrow, 13 Aug. 1950, M. Hertig. 9 <$48- i l§ "V =^-" ft \ i Figure 4 7 Lutzomyia, barpettoi majusaula — A. Sparmatheca. Female: Cerro kzu.1, Panama Prov. , Panama Lutzomyia bajrrettoi barvettoi — B. Spermathecae, same scale as Fig. 46A, C. Tip of genital fork stem, D. Tip of genital fork stem, E. Tip of genital fork stem of female from Caqueta, Colombia, F. Female cibarium, G. Female f lagellomere II, H. Male genitalia. Kale: Trinidad Female: Trinidad (except Fig. 46E) Lutzomyia texana — I. Spermatheca, same scale as Fig. 46A. Female: Sinton, Texas, U.S.A. -250- Figure 48 Lutzonyia oavpentevi — A. Male head, B. Male flagellomere II, C. Fe- male head, D. Female flagellomere II, E. Female cibarium and pharynx, F. Body of spermatheca, G. Genital pump and filaments, same scale as Fig. 48H, H. Mai e genitalia, I. Tips of genital fiimanets, J. Sperma- thecae, K. Female wing, L. Male wing, M. Female cibarium. Male: Curiche, Choco Dept. , Colombia Female: Same locality as male Figure 49 Lutzomyia vunoides — A. Male head, B. Male flagellomere II, C. Female head, D. Female flagellomere II, E. Female cibarium and Pharynx, F. Genital pump and filaments, G. Male genitalia, H. Spermathecae, I. Female wing, J. Male wing, K. Female cibarium. Male: Rio Anori, Antioquia Dept., Colombia Female: Same locality as male -254- -255- Dreisbachi. Group Lewis et a 1. , 1978 Ry virtue of the annulated spermathecae and cibarial armature, the females of this group resemble those in the subgenera Nyssomyia Barretto and Tviakopkovomyia Barretto but the ascoids have long posterior spurs and the males have 2 or more strong distal setae on each coxite. Four species comprise the dveisbaahi group, two of which occur allopatrically in Colombia. The females ordinarily do not attack man and are very similar, if not indistinguishable, in structural details. With- out associated males or knowledge of locality, I am unable to identify them beyond the group level. Key to Species Males 1. Paramere simple but with modified dorsal seta at apex and tuft of apically recurved setae near middle. Coxite with 2 large modified setae at ventral median surface aalydifeva (Fig. 50) Paramere bifurcate; it & coxite without modified setae dveisbaohi (Fig. 51) 59 . Lutzomyia aalydifeva (Fig". 50) Phlebotomus aclydifevus Fairchild & Hertig, 1952: 511 (c? holotype, Mcjinga Swamp, near Catun, Canal Zone; (j>, Juan Mina, Canal Zone) . Fair- child & Hertig, 1959: 121 (Mexican & Central American records). Johnson & Hertig, 1961: 765, 773 (rearing data). Hanson, 1968: 44 (larva, pupa. descr. , figs.) . -256- Lutzomyia aclydifera: Theodor, 1965: 193, 194 (6*, n figs., listed). Barreto, 1969: 46?. (Valle, Colombia). Sherlock & Guitton, 1970: 140 (classif.). Martins et al . , 1970a: 550 (cf. to 1 ienna.nl en bi) . Chaniotis et al . , 1971a: 344 et seq. (pop. dynamics, Panama). Young, 1972a: 312 (mention). Martins & Morales, 1972: 365 (listed). Osorno et al . , 1972a: 61 (mention). Chaniotis et al., 1972: 95 (resting sites, mostly burrows) Christcnsen, 1972a: 88 (listed). Christensen et al . , 1972: 57 et. seq. (collecting data). Chaniotis & Correa, 1974: 115 (light trap data). Rutledge et al., 1975: 181 (light trap data). Rutledge & Ellenwood, 1975a: 72 (breeding on open forest floor). Lewis , 1975a: 502 et seq. (rnouthpaxt morphol.). Martins et al . , 1976a: 493 (cf. to ruparupa) . Herrer & Christensen, 1976a: 62 (collecting data). Psychodopygus aclydifevus: Forattini, 1971a: 105 (listed), Forat- tini, 1973: 189 et seq. (gen. review, figs.). Gomes, 1975: 9 (listed). Distribution: Mexico, Honduras, Nicaragua., Costa. Rica, Panama, Colombia (Antioquia, Choco, Valle), Ecuador. Material examined: Colombia. 58 dcT1, 16 oo, Rio Anori (Antioquia), light traps, May 1970, C.H.P. 6 6*o*, 6 (ft, same data but Sept. 1970, D.G.Y. 190 6*6*, 296 o^, Curiche (Choco), light, Shannon & Malaise traps, tree trunks, April-Dec. 1967, D.G.Y. 26 do*, 28 oo, Alto Curiche (Choco), light & Shannon traps, Aug. -Nov. 1967, D.G.Y. 28 do1, 59 o^, Teresita (Choco), light & Shannon traps, June-Nov. 1967, D.G.Y. 1 o, Anchicaya Dam (Valle), light trap, 11 Aug. 1973, D.G.Y. & R.C.W. Ecuador. 15 dcf, 10 op , 17 km E of Santa Domingo de los Colorados (Pichincha) , light & Malaise traps, May 1976, D.G.Y., T. Rogers, & G. Fairchild. Panama. d'holotype, o allotype and 50+ specimens from various localities, most of which were listed by Fairchild & Hertig (1952) . Discussion: At Curiche, L. aolydifeva was the most abundant phle- bototoine species taken in light traps from April to Dec, 1967 (39 trap nights). Of a total catch of 1167 specimens (23 species), accounted for 30% or 350 specimens (155 6*o", 195 oo) . The feeding habits of this species remain unknown. 60 . Lutzomyia dreisbachi (Fig. 51) Phlebotomus dreisbachi Causey & Damascene, 1945: 649 (6* holotype, Coarij Rio Solimoes, Amazonas, Brazil). Barretto, 1947: 197 (refs.). Damasceno et al., 1949: 824 (Brazilian records). Barretto, 1950a: 106 (c? keyed). Barretto, 1951: 216 (distrib.). Floch h Abonnenc, 1952: 20 et seq. (cf, o, keyed, redescr., figs., French Guiana). Vargas h Diaz- Najera, 1953a: 49 (listed). Lutzomyia dreisbachi: Theodor, 1965: 185 (listed). Martins et al., 1965: 2 (Rondonia, Brazil). Sherlock & Guitton, 1970: 140 (classif.) . Martins et al., 1970: 550 (cf. to hermanlenti) . Young, 1972a: 312 (mention). Martins & Morales, 1972: 367 (listed). Martins et al., 1976a: 493 (cf. to ruparupa) . Martins et al . , 1976b: 498 (mention). Psychodopygus dreisbachi: Forattini, 1971a: 105 (listed) . Forat- tini, 1973, 390 et seq. (gen. review, figs.). Gomes, 1975: 9 (listed). Distribution: Brazil, French Guiana, Colonbia (Amazonas) . Material examined: Colombia. 1 d*, 17 km W of Leticia (Amazonas), light trap, 26 July 1973, D.G.Y. & R.C.W. Discussion: A little known Amazonian species, L. dreisbachi was discovered first in an armadillo burrow in Amazonas, Brazil, where over 2,000 males were captured (Causey and Damasceno, 1945). Future collections -258- should reveal its presence in other localities within the Amazon Basin. I have not examined females of this species but a description and good figures are provided by Floch and Abonnenc (1952) . Figure 50 Lutzomyia aclydifevo. — A. Male head, B. Male flagellomere II, C. Fe- male head, D. Female flagellomere II, E. Female cibarium and pharynx, F. Genital pump and filaments, G. Spermathecae, H. Male genitalia, I. Body of spermatheca, J. Female wing, K. Male wing, L. Female cibarium. Male: Curiehe, Choco Dept., Colombia Female: Same locality as male -260- -261- Subgenus Tr-ichopk o vonvjio. Barretto, 1962 This is a distinct group as treated by Barretto (1962) and Sherlock and Guitton (1970), the species of which occur in or near the Amazon Basin except for L. reburra. The 20+ Triahophoromjia species, especially females, resemble those in the subgenus Nyssomyia Barretto, 1962, as noted by several authors. Forattini (197] a, 1973) in fact enlarges TvLchophor-omyia to accomodate these and other species, some of which such as L. har>trnanni (Fairchild and Hertig), L. sangv.-inaria (Fairchild and Hertig) , L. fluvCatilis (Floch and Abonnenc) , andL. nordest-ina (Mang.) do not appear to be closely related to the others based on a combination of characters. Sherlock and Guitton (19 70) provide a useful key to the Trichophoro- myia males. The few described females, being similar in structure, are difficult to identify in the absence of males. At present, 6 Tvichophovonvjia species are known to occur in Colombia but others certainly exist in the Amazonian region. The species are attracted to lights but their preferred resting sites and hosts remain virtually unknown. None of the species, unlike those of Nyssomyia, is known to be anthropophilic. Key to Species Males 1. Genital filaments less than 4 times length of pump 2 Genital filaments greater than 4 times length of pump •262- 2. Paramere broader, less clubbed, most of dorsal setae sinuous. Eyes large, separated by less than 4 facet diameters . . vphurra (Fig. 54) Paremere otherwise, dorsal setae straight or nearly so. Eyes smaller. separated by more than 6 facet diameters. . . . vbiquitdlis (Fig. 51) 3. Coxite with only a distal group of 4-6 straight setae. Paramere as shown saltuosa (Fig. 53) Coxite with basal and/or median tufts, the total number of setae greater than 10. Paramere otherwise 4 4. Coxite with 3 groups of setae, a basal patch of about 7 short setae, a row of 4 longer hairs on median ventral surface & a distal patch of about 8 slender setae. Paramere as shown. . . eelluZana (Fig. 52) Coxite wir.h 1 or 2 groups of setae. Paramere otherwise 5 5 Coxite tuft circular and compact, the tips of nearly all setae down- wardly curved howardi (Fig. 53) Coxite tuft otherwise, the setae loosely arranged, the tips of most not downwardly curved avJ'O.ensis (Fig. 51) 61. Lutzomyia (T.) aw?o.ensis (Fig. 51) Phlehotomu.s auvaensis Mangabeira, 1942a: 161 (d1 holotype, Aura, Belem, Para, Brazil). Barretto & Coutinho, 1943: 188 (mention). Damasceno et al . , 1949: 819 (Brazilian records). Barretto, 1950a: 108 (o keyed). Vargas & Diaz-Najera, 1951a: 22 (mention). Barretto, 1951: 212 (Brazilian records). Floch & Abonnenc, 1952: 36 (c? keyed) . Llanos. 1964: 373-375 (V auvaensis, o^redescr., figs., Peru). -263- Lutzomyia auraensis: Barrettc, 1962: 97 (listed). Martins et al., 1965: 2 (Rondonia, Brazil). Theodor, 1965: 185 (listed, 6*, o. figs.). Sherlock & Guitton, 1970: 143 et seq. ( 0.40 mm long), longer than II + III olmeca bicolor' (Fig. 56) 4. Style with subterrninal and terminal spines nearly paired, separated by distance = to greatest width of terminal spine ylephiletor> (Fig. 59) Style with distal pair of spines more widely separated 5 5. Genital filament tips slightly inflated, simple. Coxite longer than lateral lobe trapidoi (Fig. 57) Genital filament tips modified. Coxite and lateral lobe subequal in length 6 6. Genital filament tips forked, claw-like. Style over half length of coxite or lateral lobe. Paramere somewhat acute at ventral basal margin. Mesonotum partly pigmented vjnbvatilts (Fig. 58) Genital filament tips otherwise. Style shorter than or = to half length of coxite or lateral lobe. Paramere rounded at lower basal margin. Mesonotum pale yuilli (Fig. 60) -282- Females 1. Common sperm duct as long as or longer than individual ducts. ... 2 Common sperm duct, if present, shorter than individual ducts. ... 4 2. Head height subequal to width. Flagellomere I shorter than labrum. Spermatheca with fewer than 8 distinct segments, about 2 x length of individual sperm ducts antunesi (Fig. 55) Head much longer than wide. Flagellomere I = to or longer than labrum. Spermatheca with 9-12 distinct segments, = to or longer than individual ducts 3 3. Flagellomere I = to or longer than II + III. Genital fork stem broad and blade-like. Cibarium with 8-10 (rarely 7) horizontal teeth . . olmeoa bioolor (Fig. 56) Flagellomere I shorter than II + III. Genital fork stem slender. Cibarium with 6-7 horizontal teeth flaviscutellata (Fig. 55) 4. Individual sperm ducts mostly striated and/or with refringent ex- crescences (papules) 5 Individual sperm ducts smooth-walled 6 5. Mesonotum pale. Individual sperm ducts" with numerous excrescences forming a feather-like pattern yuilti (Fig. 60) Mesonotum pigmented in part. Individual sperm ducts transversely striated but lacking excrescences imbratilis (Fig. 5S) -233- 6. Common sperm duct short but discernible. Spermathecae with segments increasing in size from base to apex ylephiletoi' (Fig. 59) Common sperm duct apparently absent. Spermathecae with segments subequal in size trapidoi (Fig. 57) 67. Lutzomyict (N.) aniunesi (Fig. 55) Phlebotomus centimes £ Coutinho, 1939: 181 (d7, Manacapuru, Amazonas, Brazil). Barretto, 1946b: 529-532 (refs., synonyms). Barretto, 194 7: 185 (refs., synonyms). Damasceno et al . , 1949: 818 (distrlb., Brazil). Barretto, 1950a: 108 (keyed). Barretto, 1951: 212 (distrib.). Vargas & Diaz-Najera, 1951a: 22 (mention). Floch & Abonnenc, 1952: 36 (o, Alto Curiche (Choco), light Shannon traps, biting man, tree trunks, June-Dec. 1967, D.G.Y. 12 6*0*, 6.1 oo, Teresita (Choco), light, Malaise & Shannon traps, tree trunks, biting man, March-Dec. 1967, D.G.Y. 3 53, Rio Atrato at Sautata (Choco), Malaise trap, Jan. 1968, D.G.Y. 1 6*, 8 oo, Anchicaya Dam (Valle), light flight traps, tree trunks, 9-11 Aug. 1973, D.G.Y. & R.C.W. 52 &?, 39 oo, 25 km £ of Buenaventura (Valle), light & flight traps, tree trunks, 11-12 Aug. 1973, D.G.Y. h R.C.W. Costa Rioa. 4 oo , Moravia, Chirripo (Cartage), Shannon trap, 4 March 1966, R. Zeledon. Ecuador. 84 6*0*,, 33 00. 17 km E of Santo Domingo de los Colorados (Pichincha), light & flight traps, tree trunks, May 1976, D.G.Y., T. Rogers, & G. Fairchild . Honduras. 3 op, Tela, Lancetilla Valley, light trap, 24 Feb. 1954, W. Mis. Panama. 1 o* (holotype no. 3291), type locality, 19 June 1951, A. Quinones & R. Hartmann. 1 (j> (allotype no. 2474), Finca Nievecita, Almirante (Bocas del Toro) , tree buttress, 21 June 1950, R. Hartmann. 100+ 6*0", 100+ op, various localities, most of which were listed by Fairchild and Hertig (1952). Discussion: Specimens of L. trapidoi from Valle Dept., Colombia and Pichincha Prov., Ecuador are darker than those from other localities, the heads and mesouota of both sexes being moderately infuscated. Structurally, however, they agree with the holotype and allotype of trapidoi. 1 have cited only recent references to this species, Forattini (19/3) having revisited the important papers published prior to 1971. At Curiche from April to Dec, 1967, we secured 52 00 of trapidoi in routine human bait collections. This represents 1.4% of the total catch -292- of J5 LutRomyia spp. (3763 go, 141 nan hours). At Teresita less than 1% of i'.he biting sand flies captured during an eight month period repre- sented this species. We did not attempt, however, to collect sand flies in the forest canopy where trapidoi is likely to be more abundant (Fairchild & Hertig, 1952; Chaniotis et al., 1971b). During the day, trapidoi adults rest commonly on the forest floor, less frequently on living plants or on tree trunks (Chaniotis et al . , 1972). The larvae live on the open forest floor (Rutledge & Ellenwood, 1975a, b,c) where they tend to be surface feeders (Hanson, 1968). Flagellates, believed to be leishmanial promastigotes, were recovered from wild caught trapidoi females in Panama (Johnson et al. , 1952; McConnell, 1963) . This species may also play an important role in the epidemiology of certain arboviruses in that country, Tesh et al. (1974) having isolated a number of strains from males and females. There is presumptive evidence that vertical (transovarial) transmission of some viruses occurs in L. trapidoi and L. ylephiletor (Tesh & Chaniotis, 1975). 71. Lutzomyia (N.) umbratilis (Fig. 58) Lutzomyia wrbvatilis Ward & Fraiha, 1977 (o ho lo type, Monte Dourado, Rio Jari, Para, Brazil). Phlebotomus intermedia (not intermedia, Lutz & Neiva, 1912) : Floch & Abonnenc, 1941a: 2 (cf, descr. , figs.). Floch & Abonnenc, 1942a: 9 (o, descr. , figs .) . Phlebotomus anduzei (not andazei Rozeboom, 1942): Floch & Abonnenc, 1944c: 11 (o", 0, taxonomy, figs.). Floch & Abonnenc, 1945c: 4 (o , -293- keyed, figs.). Floch & Abonnenc, 1947a: 18 (d1, keyed). Fairchild & Hertig, 1952: 523 et seq. (cf. to ylephiletor & bra.pid.oi). Floch & Abonnenc, 1952: 37, 43 (6", o, keyed), 121 (cf, o, redescr., figs.). Fairchild & Hertig, 1959: 121 (distrib. , in part). Forattini, 1960: 476 (Amapa, Brazil). Wijers & Linger, 1966: 501 et seq. (Surinam, as probable vector of Leishmanial . Lutzomyia anduzei (not anduzei Rozeboom, 1942): Martins et al . , 1963: 334 (Roraima, Brazil). Martins et al., 1965 (Rondonia & other Brazilian localities). Almeida, 1970a: 2 et seq. (anomalous , redescr., figs., Peru). Lewis, 1975a: 501 et seq. (mouthpart morphol.). Llanos et al., 1975b: 669 (Peru). Martins et al., 1976a: 487 (Peru). Lainson et al., 1976: 171-172 (as vector of Leishmania in northern Brazil). Ward, 1977: 228 et seq. (larva descr., fig., keyed). Lainson et al., 1977 (Mato Grosso, Brazil). Lutzomyia sp. no. 260.31 Ward et al . , 1973: 178 (collecting data, Para, Brazil). Ward & Killick-Kendrick, 1974: 216, 219 (Para, Brazil). Psyohodopygus anduzei: Forattini, 1973: 388 et seq. (in part, gen. review, figs., refs.). Distribution: Colombia (Amazonas, ?Boyaca, ?Caqueta) , Peru,, Brazil, Surinam, French Guiana. Mati-rrial examined: Colombia. 6 ob*, 1 o, 17 km W of Leticia (Amazonas), tree trunks, 24-26 July 1973, D.G.Y. &■ R.C.W. Brazil. 15 do", 10 op, Serra do Navio (Amapa), tree trunks, 21 Sept. 1972, D.G.Y. & H. Fraiha. 1 cf, Rio Aripuana at Humboldt (Mato Grosso), tree trunk, 16 Aug. 1974, D.G.Y. & J. Shaw. 1 o, 27 km SE of Maraba (Para), light trap, -294- 26-28 Sept. 1972, D.G.Y. & H. Fraiha. 1 cf, near Altamira (Para), tree trunk, 1 Oct. 1972, D.G.Y. Discvjssioni Lutzomyia urribratilis , confused with L. anduzei (Roze- booui) for many years, differs from it by the darker mesonotum, by the striated sperm ducts (smooth walled in anduzei), by the relative lengths of palpal segments 2 and 5, and by other features discussed by Ward & Fraiha (1977). The male of anduzei has not been described. Fairchild & Hertig (1959) reported the presence of L. anduzei in Costa Rica and Panama. I have not examined Costa Rican specimens but 3 op from Almirante (Bocas del Toro Prov., 19-21 June 1950, R. Hartmann) seem to be conspecific with L. anduzei (Rozeboom) . The fifth palpal segment is shorter than the third, post of the insect is pale and the spermathecae and ducts are similar to those of L. anduzei Rozeboom ex- cept for some transverse striations on the individual ducts. There is no doubt that the specimens from near Leticia, Colombia, are conspecific with L. vrrbratilis as described by Floch & Abornienc (1944, 1952, as anduzei) and by Ward & Fraiha (1977). I am assuming, perhaps incorrectly, that the records of anduzei by Osorno et al. (1972a) from Boyaca and Caqueta Departments represent L. wrbratilis , not anduzei P.ozeboom. Lainson et al . (1976) found promastigotes in 4 out of 55 wrbratilis females examined in northern Brazil. These subsequently proved to belong in the genus Leishmania and may be identical to the type discussed by Wijers & Linger (1966) in Surinam (Lainson et al., op. cit.). ■295- 72. Lutzomyia (N.) ylephiletov (Fig. 59) Ffolebotomus ylephiletor Fairchild 6 Hertig, 1952: 518 (6* holotype, Almirante, Bocas del Toro Prov. , Panama; o, Cacique, Colon Prov. , Panama). Lewis & Garnham, 1959: 82 (Belize). Fairchild & Hertig, 1959: 122 (distrib.) • Forattini, 1960: 474 (not ylephiletor Fairchild & Hertig, 1952). Hanson, 1961: 320 et seq. (larvae on forest floor). Johnson & Hertig, 196i: 765 (rearing data). Hanson, 1968: 95 (larva, pupa, figs.) Lutzomyia ylephiletor (or ylephiletrix) : Barretto, 1962: 98 (listed). Barreto, 1969: 468 (Valle, Colombia). Williams, 1970: 332 et seq. (summary of collecting data, Belize). Tesh et al . , 1971a: 152 et seq. (blood meals). Chaniotis et al., 1971a: 344 et seq. (pop. dynamics, Panama). Chaniotis et al . , 1971b: 415 et seq. (man-biting study). Osorno et al., 1972a: 36 (listed). Tesh et al . , 1972: 88 et seq. (blood meals). Chaniotis et al., 1972: 91 et seq. (resting sites). Christensen, 1972a: 88 (listed). Christensen et al., 1972: 55 et seq. (collecting data). Zeledon & Alfaro, 1973: 416 (naturally infected with Leishmania, Costa Rica). Christensen & Herrer, 1973: 579 et seq. (col- lecting data, Panama). Chaniotis, 1974a: 73 et seq. (sugar feeding tests). Chaniotis & Correa, 1974: 115 (collecting data). Chaniotis, 1974b: 501 (keyed). Tesh et al., 1974: 258 et seq. (summary of virus studies). Chaniotis et al . , 19 74: 372 et seq. (dispersal study). Lewis, 1975a: 501 et seq. (mouthpart morphol.) ■ Lewis, 1975b: 366 (hair sockets, figs.). Tesh & Chaniotis, 1975: 125 et seq. (virus transmission studies). Herrer & Christensen, 1976a: 62 (collecting data), 1976b: 57 (collecting data). Williams, 1976a: 604 (in caves, Belize). Zimmerman et al., 1977: 574 et seq. (egg, descr., figs.). -296- Psychodopygus ylephiletor: Forattini, 1971a: 1G6 (listed). Forattini, 1973: 123 et seq. (gen. review, refs., figs.). Gomes, 1975: 9 et seq. (gen. review, distrib.). Distribution: Mexico, Guatemala.;, Belize, Honduras, Nicaragua, Costa Rica, Panama, Colombia (Choco, Valle) . Material examined'. Colombia. 17 oo, Curiche (Choco), Shannon & Malaise traps, biting man, tree trunks, 26 June-25 Nov. 1967, D.G.Y. Mexico. 1 o, Palenque (Chiapas), tree buttress, 1 April 1951, C. Fair- child & R. Hartmann. 2 oo, Teapa (Tabasco), light traps, 19 Aug. & 10 Sevt. 1953. Guatemala. 8 oo, Montana del Mico, Rio Blanco, human bait, no data (but probably March 1956), J- Boshell. Honduras. 19 oo, Tela, Lancetilla Valley, human bait, March-April, 1954, W. Hils. Nicaragua. 1 6*, 1 o, Villa Soraoza, 15 June 1953. Costa Rica. 2 oo. Esquinas (Puntarenas) , tree buttresses, 28 Nov. 1949, H. Trapido. Panama. 1 ermathecae. Male: Trinidad Female: Trinidad jUtzomy-ia (N.) flaviseutellata — D. Spermathecae, £. Genital pump and genitalia, F. Female cibarium. Male: Leticia, Amazonas Comisaria, Colombia Female: Trinidad -300- P=*> :0^ A W|\\-^%^ v ■-•>--5A> ^iMS^f F\\\ j I Figure 56 Lutzonvjia (N.) olrneoa bicol-ov — A. Male head, B. Male flagellomere II, C. Female head, D. Female flagellomere II, E. Female cibarium, F. Male genitalia, G. Genital pump and filaments, H. Spermathecae, I. Female wing, J. Male wing, K. Female wing. Male: Curiehe, Choco Dept,, Colombia Female: Same locality as male -302- Figure 57 Lutzormjia (N.) tvapidoi — A. Male head, B. Male flagellomere II, C. Female head, D. Female flagellomere II, E. Female cibarium and pharynx, F. Male genitalia, G. Cenital pump and filaments, H. Sperma- thecae, I. Female wing, J. Male wing, K. Female cibarium. Male: Curiche, Choco Dept., Colombia Female: Same locality as male -304- c^fr% Figure 58 Lutzomyia (a.) umbrat-ilis -- A. Male head, B. Female head, C. Female cibarium, D. Female wing, E. Male genitalia including genital pump, F. Tip of genital filament, G. Spermathecae. Male: Leticia, Amazonas Comisaria, Colombia Female: Para State, Brazil ■306- Figure 59 Lutzomyia (N. .' yZephiletov — A. Male head, B. Male flagellomere II, C. Female head. D. Female flagellomere II, E. Female cibarium and pharynx, F. Male genitalia, G. Spermathecae of female from Costa Rica. H. Female wing, i. Male wing, J. Female cibarium. Male: Aimirante, Bocas del Toro Prov., Panama Female: Panama Canal Zone (except Fig. 59G) ■308- |LjI D Figure 60 Lutzomyia (N.) yv.illi — A. Male head, B. Male flagellomere II, C. Female head, D. Female flagellomere II, E. Female cibarium and pharynx, F. Different views of paramere, G. Male genitalia, H. Tip of genital filament, I. Spermathecae, J. Female wing, K. Male wing, L. Female cibarium (all figures from Young & Porter, 1972). Male: Rio Anori, Antioquia Dept., Colombia Female: Same locality as male ■31! -311- S ub genus Psyc iho dopygus Mangabeira, 1941 • Some of the most no torious man-biters among the Lutzomyia sand flies be loi •g in Lh is subgenus, a well defined group of about 25 taxa which is some? imes tr eated as a genus (F arattini, 1971a, j 973; . Members of Ps ychodopygu 8 range from Mexico to southern Brazil and are commonly ob- served in forests attacking man. Several species have been incriminated as vectors of dermal leishmaniasis caused by Leishmania bvaziliensis (Lamson et al., 1973). Many of the species were described originally from a single sex from a single locality (nondimensional species) and this, along with the dif- ficulty of identifying females alone, has led to numerous errors in the. taxonomic and medical literature. Some corrections and name changes, discussed in mere detail under the species, are summarized as follows. Lutzomyia tintinnabula Chris tensen and Fairchild, is a junior synonym of L. ayvozai (Barretto and Coutinho) (NEW SYNONYM). Lutzomyia pessoana (Barretto) from southern Brazil is a junior synonym of L. cavvevai (Barretto) described from Meta Dept., Colombia. Lutzomyia 'pessoana of various authors, treated as Pklebotomus paraensis by Fairchild and Hertig, 1951a, differs subspecifically from L. cavvevai and is given a new name, L. cavvevai thula n. ssp. The female of L. c. cavrevai (NEW STATUS) or possibly that of L. faivchiidi Barretto was incorrectly described as L. davisi by Root (1934). Martins et al., (1973b) and Llanos (1973) correctly associated the sexes of L. davisi. Forattini's descriptions (1960, 1973) of the female of L. paraensis (Costa Lima) represent not that species but L. davisi. Lutzomyia hirsuta (Mang.) and L. nicavaguensis (Fairchild and Hertig) differ subspecifically (NEW STATUS) . The female of L. h. hivsuta, was -312- incorrectly described as L. davisi (not davisi Root, 1934), by Forattini (1960, 1973) . Only the nominate subspecies occurs in Colombia where it was mis identified as L. paraensis by Osorno et al . (19 72a). The other records of paraensis in Colorbia (Barreto, 1969) represent not that species but L. earrerai thula and/or L. ayvozai. Antunes (1937) reported the presence of L. sqiianrii'entr'is (Lutz and Neiva) in Meta Dept., Colombia but this record is based on females alone and I consider it to be doubtful, believing that the specimens represented another species in the series squamiventris , probably L. fairtigi Martins. Based on male genitalia, the Psychodopygus species are divided into 3 series following Theodor (1965) and Lewis et al. (1978). This agree- ment, admittedly artificial, is useful nevertheless for discussion and identification purposes. The short fifth palpal segment and imbricated spermathecae, both apomorphic features, are characteristic of all the Psychodopygus species. Studies on feeding habits indicate that most, if not all, females feed on mammals. The male genitalia may be simple. (series arthuri) or very complex (series sqi'jxmiventris and panamensis) . At present, 14 species and subspecies in the subgenus are known to occur in Colombia. Keys to Species Males Series squamiventris 1. Paramere as shown, with dense apical patch of 304- long erect setae and with a slender upwardly directed process at apex. . chagasi (Fig. 61) Paramere otherwise, without dense patch of dorsal setae but with a group of 5-20 setae near base of subterminal dorsal arm 2 2. Main lobe of paramere slender and upturned apically, with a group of about 15 long setae near base of dorsal arm . . . .bernalei (Fig. 61) Main lobe of paramere broad, not upturned apically, with about 5 short stout setae near base of dorsal arm fo.ivtigi (Fig. 62) Series arthuri. The male of L. bispinosa, the sole representative of this series in Colombia, differs from other Psycho dopy gus males by having only 2 spines on the style, both of which are terminal. Series pancavens-is 1. Style with 5 (rarely 6) strong spines. Paramere as figured .... ■ davisi (Fig. 67) Style with 3 strong spines and 1 smaller bristle. Paramere other- wise 2 2. Paramere simple, slender distally, without arms or other extensions ..... guyanensis (Fig. 68) Paramere complex, with ventral and/or dorsal arm 3 3. Paramere with recurved apical setae on main lobe and ventral arm; dorsal arm present. Aedeagus long and slender, reaching to ends of longest recurved setae on main lobe of paramere . . Tsaia-va (Fig. 71) Paramere otherwise, without dorsal arm or recurved setae. Aedeagus rather short and stout 4 -31 4. Main lobe of paramere with 2 tufts of blade-like setae; ventral arm wirh 2 enlarged setae, one terminal and one sub terminal , panamensis (Fig. 70) Main lobe of paramere with 1 group of setae, blade-like or not; ven- tral arm with smaller, more numerous setae 5 5. Paramere as shown, ventral arm inserted near base of main lobe. . . hirsuta hivsuta (Fig. 67) Paramere otherwise, ventral arm shorter, inserted at or near apex of main lobe 6 6. Mesonotum and pleura pale ........ car-rerai oawevai (Fig. 65) carrevai thula (Fig. 66) Mesonotum partly or entirely pigmented, contrasting with pale pleura* 7 7. Paramere with a short ventral arm (length subequal to width of coxite) apical setae (20+) shorter, not blade-like. Mesonotum, katepisternum, procoxae, and mesocoxae pigmented ayrozai (Fig. 64) Paramere with a longer ventral arm, its length definitely greater than width of coxite; apical setae (less than 20) blade-like and long. Only mesonotum pigmented noat-icota (Fig. 69) *The katepisternum of L. ayrozoi is faintly pigmented but the rest of the pleura is decidedly paler than the mesonotum. -315- Females 1. Common sperw duct completely smooth-walled, not rugose* bevaalei ehagasi fairtigi (Fig. 62) Common sperm duct rugose in part and/or with transverse striations. 2 2. Cibarium with small subequal vertical teeth in 1-3 transverse rows, without longitudinal rows reourva (Fig. 71) Cibarium with vertical teeth of varying size, those in the middle enlarged and forming 2 or more longitudinal rows, irregular or not. 3 3. Sparmatheca shorter than individual duct 4 Spermatheca longer than individual duct .... 5 4. Common sperm duct with complete transverse striations guyanensis (Fig. 68) Common sperm duct with incomplete transverse striations appearing as short, thick lines sp . of Tres Esquinas (Fig. 61) 5. Mesonotum and pleura pale 6 Mesonotum partly or entirely pigmented, contrasting with pale pleura 7 *I am not able to separate these females without associated males. The female of L. bermalei, presently unknown, probably resembles the other 2 species, all of which belong in the series sqi'amiventris . -316- • 6. Lai-rum shorter than 0.30 rum or shorter than combined length of scape and f Lagellomere I carrevai. aavrerai (Fig. 65) Labrum longer than 0.30 mm or greater than combined length of scape and f lagellomere I cavvevai thula (Fig. 66) 7. Cibarinm with short inwardly directed horizontal teeth. Katepisternum, procoxae, and mesocoxae pigmented cyrosai (Fig. 64) Cibarium with longer erect horizontal teeth. Katepisternum, procoxae, and mesocoxae pale 8 8. Common sperm duct completely smooth-walled below rugose portion . . 9 Common sperm duct with transverse striations or dot-like thickenings below rugose portion 10 y. Terminal annulation of spermatheca asymmetrical. Mesoscutellum pig- mented panarnensis (Fig. 70) Terminal annulation of spermatheca symmetrical. Mesoscutellum pale noaticola (Fig. 69) 10. Common sperm duct with complete or nearly complete transverse stria- tions. Stem of genital fork broad, blade-like . . bispinosa (Fig. 63) Common sperm duct heavily rugose below junction of individual ducts; rest of duct with dot-like thickenings, not transversely striated. Stem of genital fork more slender, subacute at tip 11 11. Individual sperm duct less than half the length of spermathcea; rugose section of common duct swollen near junction of individual ducts. . hirsute hivsuta (Fig. 67) -317- Individual sperm duct over half the length of sperniatheca; rugose section of common duct not swollen, more or less equal in width throughout davisi (Fig. 67) Series squamiventris 74. Lutzomyia (P.) bernalei (Fig. 61) Phlebotomus bernalei Osorno, Morales, & Osorno, 1967: 30 (6* holo- cype, Araracuara, Caqueta, Colombia). Lutzomyia bernalei, '. Martins et al., 1968: 22 (mention). Osorno el: al., 1972a: 45 (listed). Psyehodopygus bernalei: Forattini, 1971a: 104 (listed). Forattini, 1973: 392 et seq. (review, keyed, fig.). Distribution: Colombia (Caqueta) . Material examined: , redescr., figs., refs.) . Psychodopygus unisetosus: Forattini, 1971a: 104-105 (listed). Psychodopygus chagasi: Forattini, 1973: 392 et seq. (review, keyed) Fraiha et al., 1974: 89 (cf, figs., biting man). Distribution: Colombia. (Caqueta) , Material examined: Colombia. 4 tfd1, Tres Esquinas (Caqueta), light trap, 10 Nov. 1971, C.J.M. Brazil. 1 o. , km 13 Macajai-Caracarai (Roraima) , human bait, 16 July 1973, H. Fraiha. Discussion: This man-biting species, precinctive in the. Amazon Basin, is easily recognized but I am unable to separate the females from those of L. fairtigi, L. squamiventris , and related species in the series squamiventris without associated males. 76. Lutsomyia (P.) fairtigi (Fig. 62) Lutsomyia fairtigi Martins, 1970: 279 (c? holotype, Villavicencio, Meta Dept., Colombia). Martins & Morales, 1972: 366 (listed). Phlebotomus squamiventris: Fairchild & Hertig, 1951a: 414 (6", £, redescr., figs., not squamiventris Lutz & Neiva, 1912). Psychodopygus fairtigi: Fraiha et al., 1971: ^95-496 (cf. to wellccmei) . Forattini, 1971b: 152-154 (Mato Grosso, Brazil). Forattini. 1973: 392 et seq. (distrib., figs., keyed). -319 - Distribution: Colorrbia (Meta, Caqueta) , ?Brazil (Mato Grosso). Material examined: Colombia, c? holotype, o allotype, 6 o_g paratypes, Vil lavicancia (Meta), stable trap, 10 June 1948, C.Y. Chow. 3 d*d"s Tres Esquinas (Caqueta), light trap, 10 Nov. 1971, C.J.M. Discussion: A little known species^ L. fairtigi was first collected in a donkey-baited stable trap at Villavicencio and later in a light trap at 'Ires Esquinas. Forattini (1971b, 1973) lists this species (females only) from Salto Augusto (Mato Grosso), Brazil, but its presence there can only be confirmed when the male is discovered. Series avthuri 77. Lutzomyia (P.) bispinosa (Fig. 63) Phlebotomus bispinosus Fairchild & Hertig, 1951a: 410 (6* holotype, La Victoria, Cerro Jefe, Panama). Fairchild, 1955: 194 (listed). Lewis & Garnham, 1959: 80-81 (o. figs., Belize). Fairchild h Hertig, 1959: 121, 123 (distrib., Central America). Forattini, 1960: 479 (Amapa, Brazil). Williams et al . , 1965: 65 et seq. (Belize). Lutzomyia bispinosa: Barretto, 1962: 94 (listed). Theodor, 1965: 188 (listed). Barretto, 1966: 141, 144 (keyed). Williams, 1970: 332 et seq. (Belize). Chaniotis et al., 1971a: 344-345 (collecting data, Panama). Christensen, 1972a: 88 (listed). Lewis, 1975a: 500 et seq. (mouthpart morphol.). Lainson et al., 1977 (Mato Grosso, Brazil). Psyc-liod.opygus bispinosus: Forattini, 1971a: 104 (listed). Forat- tini, 1973: 170 et seq. (distrib., cf, o, keyed, figs.). Distribution: Belize, Honduras, Nicaragua., Panama., Colombia (Choco)., Ecuador, Northern Brazil. -320- tiatevial examined: Colombia. 24 oo, Curiche (Choco) , biting man, tree trunk, Malaise, light, & Shannon traps, May-Oct. 196 7, D.G.Y. 1 c. , Alto Curiche (Choco), light trap, 8 Sept. 1967, D.C.Y. 10 6"cT, 66 05, Teresita (Choco), biting man, tree trunks, in Malaise, light, & Shannon traps, May-Dec 1967, D.G.Y. Ecuador. 3 op, Rio Napo at Limoncocha (Napo), flight trap, 22-24 May 1976, D.G.Y. & T.E. Rogers. Panama. , sane data but flight trap, 3 May 1970. 2 o^, same data but biting man, 25 June 1970. 107 eft?, 204 00, same data but light traps, Sept. 1970, D.C.Y. 1 q, same data but tree crunk, 21 Sept. 1970. 2 88, 4 go., Puerto Boyaca (Boyaca) , light trap, 10 Aug. 1971, C.J.M. 4 38, 8 00, Trcs Esquinas (Caqueta) , light trap, 10 Nov. 1971, C.J.M. 2 68, 177 00, Curiche (Choco) , biting man, Malaise, Shannon, & light traps, April-Sept. 1967, D.G.Y. 3 68, 5 5$, Alto Curiche (Choco), light & Shannon traps, July 1967, D.C.Y. 2 00 , Anchicaya Dam (Valle), flight trap, 10 Aug. 1973, D.G.Y. & R.C.W. 1 g, same data but 10 June 1975, R.C.W. 1 o, 25 km E of Buenaventura (Valle), light tr^p, 12 Aug. 1973, D.G.Y. & R.C.W. Brazil. 1 £, Manacapura (Amazonas) , 1936. 1 p, near Manaus (Amazonas), 29 Oct. 1974, J. Arias. 2 68, Mata Prefeitura, Oliveira (Minas Gerais) , 22 March 1960, Silva. 1 d\ 1 o, M.D. dc; Rio Santa Antonio (Minas Gerais), 3 Jan. 1970, Silva. 5 6*0*, 2 00 , Belem (Para), flight trap, 2-5 Aug. 1974, D.G.Y. 1 6 (cotype) , Horto Florestal, Sao Paulo (Sao Paulo), 1941, M. Barretto & J. Coutinho. 1 8, S. Jose Campos (Sao Paulo), at light, 11 Nov. 1940, M. Barretto & J. Coutinho. Panama. 1 6", 1 o (paratypes of tintinnabula) , Cerro Quia, Finogana (Darien Prov.), 14 March 1970, F. Beam. Trinidad. 1 (j>, Bush Bush Forest, Nariva Swamp, 20 Nov. 1962, T.H.G. Aitken. Other specimens, both males & females, from Trinidad, various dates and localities, to be treated in another paper on the phlebotomine fauna of Trinidad. Discussion: My treatment of L. tintinnabula as a junior synonym of L. aypozai is based on the following considerations. The female of L. aupozai, correctly associated with the male and described by Forattini and Galati (1977) , was believed earlier to be that -323- described by Barretto and Coutinho (1943) but, their 1943 description represented that of L. guyanensis (Floch and Abonnenc) , not L. auvozai (Barretto and Coutinho) . This misled others including Christensen and Fairchild (1971) who described L. iintinnabula as a distinct species primarily on the basis of female characters, notably the different sperma- thecae and ducts. The true female of ayvozai and that of L. tintinnabu'la are indistinguishable, however. The character states used in separating the males of tintinnahula and ayvozai-, discussed by Christensen and Fairchild (1971), are variable and reflect intraspecif ic , not interspecific, variation. Delta, of wing vena- tion is shorter than, equal to or longer than gamma depending on the individual specimen. The shape of the parameres varies slightly according to geographic locality but not to the extent that it repi-esents a dif- ference at the species level. One feature shared by all the specimens examined is that the procoxae, mesocoxae, and areas just above them on the pleura are moderately infuscated. Lutzomyia ayvozai is a widespread, anthropophilic species which probably occurs in other parts of Colombia, Brazil, and possibly Venezuela, Peru, and Ecuador. At Curiche (Choco) from April to Dec, 1967, we col- lected 144 females on human bait, the majority of which were taken in June (122 specimens) . The total catch represented 3.8% of all biting specimens taken during this 8 month period (141 man hours) . 79. Lutzorrryia (P.) cavrovai car'vevai (NEW STATUS) (Fig. 65) Phlebotornus cavrevci Barretto, 1946d: 286 (c? holotype, o, Vega Grande, Restrepo, Meta Dept., Colombia). Fairchild & Hertig, 1951a: 401 ■324- et seq. (listed, keyed). Ortiz & Alvarez, 1972: 140 et seq. (listed, cf. to parimaensis) . Ortiz, 1972b: 222 et seq. (taxonomic discussion, figs. , keyed) . ?Phlebotomus davisi: Root, 19 34: 233 (in part, c. only). Sergentomyia carrerai: Barretto, 1955a: 82 (listed). Barretto, 1955b: 249 (cf. to pessoana) . Sergentomyia pessoana Barretto, 1955b: 247 (, Caravelas (Bahia) , • ran .1931, N. Davis. 1 cf, ] o, near Bacuri, N of Maraba (Para), flight trap, 2C' Oct. 1974, J.F. Reinert. 1 o, same data but 1 Nov. 1974. 6 3d, 40 oj, near Altamira (km 165) (Para), biting man & Shannon trap, 6 Nov. 1974, J.F. Reinert. 1 o, same data but flight trap, 13 Nov. 1974. 1 d*, 1 o, Itatiaia (Rio de Janeiro), at light, Feb. 1955, M. Barretto. Ecuador. 2 no, Limoncocha, near Rio Napo (Napo) , flight & light traps, 20-24 May 1976, D.G.Y. & T. Rogers. Peru. 1 cf, 2 , descr., figs., Panama). Bruijning, 1957: 229 (Surinam). Lewis & Garnham, 1959: 80-81 (o, figs., Belize). Johnson & Hertig, 1961: 765 (rearing). Hanson, 1968: 58 (larva descr., figs.). Phlebotomus ayvozai: Barretto & Coutinho, 1943: 183 (o not Barretto & Coutinho, 1943) . Lutzomyia geniculate: Barretto, 1962: 94 (listed). Barretto, 1966: 141, 145 (keyed). Williams, 1970: 332 (Belize). Osorno et al . , 1972a: 46 (Caquata, Colombia). Lewis, 1975a: 502 et seq. (mouthpart morphol.) . Llanos et al . , 1975b: 670 (Peru). Lutzomyia guyanensis: Barretto, 1962: 94 (listed). Barretto, 1966: 145 (keyed). Lewis, 1975a: 502 et seq. (mouthpart morphol.). -334- Psyohodopygus geniculatus: Forattini, 1971a: 105 (listed). Forattini, 19 73: 124 et seq. (as synonym of guyanensis). Shaw et al., 1972: 720 (biting man). Ward et al., 1973: 178 (biting man. Para, Brazil). Lainson et al . , 1973: 190 (Para, Brazil). Ward, 1976: 239 (larva keyed) . Psyohodopygus guyanensis: Forattini, 1971a: 105 (listed). Forattini, 1973: 124 et seq. (gen. review). Forattini & Galati, 1977: 25 (o, of. to ayrozai) . Distribution: Belize, Panama, Colombia. (Caqueta, Choco) , Ecuador, Peru, SuY-lnam, Brazil. Material examined'. Colombia. Id1, 2 ra, Tres Esquinas (Caqueta), light trap, 10 Nov. 1971, C.J.M. 117 op, Curiche (Choco), biting man, light & Shannon traps, June-Sept. 1967. 16 oo, Alto Curiche (Choco), biting man, light traps, June-Sept. 1967. Brazil. 6 oo, Belem (Para), light trap, 18 Oct. 1972, D.G.Y. 1 o, Cabeca da Anta (Sao Paulo), at light, 1941, M. Barretto & J. Coutinho. Ecuador. 6 S3, 208 o^> , Rio Napo at Limoncocha (Napo), light traps, 19-23 May 1976, D.G.Y. & T. Rogers. 5 qq , same data but flight traps. Panama. 8 op, Almirante (Bocas del Toro), Shannon trap, 22 June 1951, W. Hils . 1 o*, same data but light trap, 30 March 1953, A. Quinones. Discussion: Although Forattini (1973) is probably correct in treat- ing L. genicula.ta as a junior synonym of L. guyanensis, there remains the possibility that they are distinct taxa. Additional material, particu- larly males, from or near the type locality of guyanensis , is needed to confirm the synonomy. At Curiche in 1967, we collected 102 guyanensis females biting man, the majority of which were taken in June (98 specimens, 24 man hours) . -335- The remaining 4 man-biting females were captured in July and August (43 nan hours) . Williams (1970) in Belize dissected 19 females but. found no flagellates. He captured specimens on dead leaves on the forest floor, on man, in light traps, and in a rodent-baited Disney trap. 83. Lutzomyia (P.) hirsute hirsute NEW STATUS (Fig. 67) Phlebotomus hir-sutus Mangabeira, 1942a: 116 (, Cano Saddle, near Gatun, Canal Zone). Barretto, 1947: 216 (complete refs.). Fairchild & Hertig, 1951a: 399 et seq. (o*, o_, descr., discussed, figs.). Osorno et al., 1967: 28 (Meta, Colombia). Morales et al . , 1969a: 381 (Meta, Colombia). Ortiz, 1972b: 22 et seq. (cf, o_ , keyed, figs.). Ortiz & Alvarez, 1972: 140 (listed). Lutzomyia panamensis : Barretto, 1962: 94 (listed). Theodor, 1965: 188 (listed, figs.). Barretto, 1966: 142, 144 (keyed). Barreto, 1969: 465 et seq. (Valle, Colombia). Tesh et al., 1971a: 153 (blood meals). Osorno et al., 1972a: 47-48 (Colombian records). Tesh et al., 1972: 90 (blood meals). Young, 1973: 108 (mention). Chaniotis & Correa, 1974: 115 (biting habits, Panama). Chaniotis, 1974b: 501 (keyed). Lewis, 1975a: 502 et seq. (mouthpart morphol.). Rutledge & Ellenwood, 1975a: -340- 12 et seq. (breeding sites, refs. on bionomics). Rutledge et al. , 1975: 179 (ecology, Panama), Williams, 1976a: 604 (in caves, Belize). Phlebotomus peasoanus: Strangways-Dixon & Lainson, 1966: 192 (not pessoana Barretto, 1955, Belize). Psychodopygus panamensis : Lewis, 1965: 376 et seq. (internal morphol.). Forattini, 1973a: 89 et seq. (gen. review, figs., cf, p) . Ward, 1976: 239 (larva keyed) . Distribution: Mexico, Central America, Colombia (Antioquia, Boyaca, Caldas, Caqueta, Choco, Guajira, Meta, Norte de Santander, Santander, Valle) , Ecuador, Peru, Brazil, Venezuela. Material examined: Colombia. 40 do", 45 op, Rio Anori (Antioquia), light traps, May 1970, C.H.P. 5 do*, 8 o^, same data but Sept. 1970, D.G.Y. 4 on, Puerto Boyaca (Boyaca), light trap, 10 Aug. 1971, C.J.M. 1 d", 4 op, same data but 6 May 1973. 5 do", 6 go, Tres Esquinas (Caqueta), light trap, 10 Nov. 1971, C.J.M. 65 do*, 2386 o^ , Curiche (Choco), biting man, light, Shannon, & Malaise traps, April-Dec. 1967, D.G.Y. 12 do*, 474 op, same data but Alto Curiche, June-Dec, 1967. 72 dcf, 1528 (jig, same data but Teresita, March-Dec. 1967. Ecuador. 1 o, 16 km E of Santo Domingo de Los Colorados (Pichincha) , flight trap, 28 May 1976, D.G.Y., T. Rogers, & G. Fairchild. Panama. 3 dcf, Almirante (Bocas del Toro) , tree buttress, 21 June 1950, R. Hartmann. 5 oo, Juan Mina (Canal Zone) biting man in daytime, Sept. 1949, H. Trapido. Numerous other specimens from Panama, Mexico, Nicaragua, and Venezuela in collection at UF. Discussion: Lutzomyia panamensis , a widely distributed, strongly anthropophilic species, was the dominant man biter at Curiche, Alto Curiche, and Teresita during medical ecology studies in the Choco in 1967, At Curiche, we captured a total of 1751 panamensis females on human bait 341- from April Lo December (141 man hours), over half of which were taken in June (624 oo, 24 man hours) . One extraordinary biting collection on 8 June from 2000 to 2200 hours (4 man-hours) yielded 1151 Lutzomyia females of 11 species. Lutzomyia panamensis accounted for 250 of these. Most of the information on the bionomics and disease relationships of this important species is summarized by Forattlni (1973) . A few additional papers, published since then, are cited here in the species bibliography . 86 . Lutzomyia (P. ) reourva (Fig. 71) Lutzomyia reourva Young, 1973: 106 (6* holotype, o, Curiche, Choco, Colombia). Lewis, 1975a: 505 (listed). Distribution: Panama (Darien), Colombia (Choco). Material examined: Colombia. 11 3>3, 480 oo, Curiche (Choco), biting man & light, Shannon, & Malaise traps, April-Dec. 1967, D.G.Y. Panama. 1 6*, 2 op, Altos de Quia (Darien), 1970, F. Beam (GML collection). Discussion: Lutzomyia reourva and L. amazonensis (Root, 1934) , both anthropophilic, apparently are closely related. The male of the latter species, unknown for many years, was recently described by Llanos et al . (1975b). It and reourva, unlike the other series panamensis males, possess a slender, arched dorsobasal arm on each paramere but differ from each other in the shape and setation of the paramere. Lutzomyia reourva, known only from Choco Dept., Colombia, and Darien Prov., Panama, occurs well outside the range of amazonensis. Specific data on human biting collections of reourva were given in an earlier paper (Young, 1973) . 17. Lutzomyia (P.) sp. of Tres Esquinas (Fig. 61) Female (description based on 3 09) : Wing length 2.0-2.3; width 0.6. Mesonotum and abdominal tergites well pigmented, head less so, rest of insect pale. Head height 0.41-0.45; width 0.38-0.40. Eyes separated by 0.10-0.11 or by distance = to ca . 5.2 facet diameters. Flageliomare I (0.23-0.25 long), 1.2 x length of II + III; ascoids simple, reaching to or beyond ends of their respective segments, on all but last 3 flagel- lomeres. Length cf palpal segments: 1 (0.05), 2 (0.13-0.14), 3 (0.15- 0.16), 4 (0.05), 5 (0.08); palpal sensilla (8-12) on distal 1/3 or segment 3. Labrum 0.30-0.33 long. Cibarium with 4 horizontal teeth arranged as shown (1 o has 5) and 15-20 vertical teeth, those in middle enlarged; chitinous arch not visible; pigment patch well defined, moderately pig- mented. Pharynx 0.20 long, unarmed. Pleura with 10-19 upper and 6-7 lower episternal setae. Length of wing vein sections: Alpha (0.45-0.61), beta (0.21-0.31), delta (0.05-0.16), gamma (0.18-0.20). Length of femora, tibia, and basitarsi of slide 658: Foreleg (missing), midleg, 0.83, 1.4, 0.88; hindleg, 0.93, 1.6, 0.98. Abdominal sternites entire, lacking clear areas. Tergite 8 with 10+ setae on each side. Spermatheca with 6-8 well formed annuli, the terminal one symmetrical; individual sperm duct over 2 x length of spermatheca; common duct as shown, the apical 1/5 rugose and with transverse striations appearing as complete lines, the remainder with incomplete striations more pronounced and numerous on the basal half of duct. Cerci unremarkable. D-is tpibution: Colombia (Caqueta) . Material examined: Colombia. 3 09, Tres Esquinas (Caqueta), light trap, 10 Nov. 1971, C.J.M. -343- Discussion: I hesitate to formally name this species because the male is unknown and because I am not entirely convinced that it and L. guyanensis are distinct taxa. Except for differences in the sperm ducts and slightly longer individual ducts of the Tres Esquinas specimens, the females appear to be indistinguishable. Figure 61 Lutzomyi-a (P.) hevaalei — A. Male genitalia (after Osorno et al . , 1967). Male: Araracuara, Caqueta Intendencia, Colombia Lutzomyia (P.) ohagasi — B. Male genitalia. Ma.le: Tres Esquinas, Caqueta Intendencia, Colombia Lutzomyi.a sp. of Tres Esquinas, female — C. Cibarium, D. Head, E. Flagel- iomere II, F. Wing, G. Spermathecae . •34b- Figure 62 Lutzomyia (P.) failrtigi — A. Male head, B. Male f lagelloinere II, C. Female head, D. Male flagellomere II, E. Sperrnathecae, F. Paramere, G. Male genitalia, H. Female cibarium, I. Female cibarium and pharynx, J. Female wing, K. Male wing. Male: Villavicencio, Meta Dept., Colombia Female: Same locality as male 347- Figure 63 Lutzomyia (P.) bispinosa -- A. Male head, B. Male flagellomere II, C. Female head, D. Female flagellomere II, E. Female cibarium and pharynx, F. Male genitalia, G. Genital pump and filaments, same scale as ?'ig. 63F, H. Body of spermatheca, I. Spermathecae, J. Female wing, K. Male wing, L. Female cibarium. Male: Cerro Jefe, Panama Prov. , Panama Female: Curiche, Choco Dept., Colombia ■349- ra J i/imk 4 IU .-;'^ > //I V ^. ft /ft ^ Figure 64 Lutzomy-ia (P.J ayvozai — A. Male head, B. Male flagellomere II, C. Female head, D. Female flagellomere II, E. Female cibarium and pharynx, F. Genital pump and filaments, G. Male genitalia, H. Body of spermatheca, I. Spermathecae, J. Female wing, K. Male wing, L. Female cibarium. Male: Curiche, Choco Dept., Colombia Female: vSame locality as male -351- Figure 65 Lubzormjia (P.) eavvevax carrerai — A. Male genitalia, B. Spermathecae, C. Female head. Male: Tres Esquinas, Caqueta Intendencia, Colombia Female: Same locality as male 353- Figure 66 Lutzomyia (P.) carvevai tkula — A. Male genitalia, B. Female head, Panamanian specimen, C. Spermathecae, D. Female cibarium. Male: Curiche, Choco Dept., Colombia Female: Same locality as male (except Fig. 66B) -355- Figure 67 Lutzonryia (P.) davisi — A. Spermathecae of female from Para State, Brazil. B. Male genitalia, C. Female cibarium. Male: Tres Esquinas, Caqueta Intendencia, Colombia. Female: Same locality as male (except Fig. 67A) Lutzomyia (P.) kirsuta hirsuta — D. Spermathecae, E. Male genitalia, F, Female cibarium. Male: Tres Esquinas, Caqueta Intendencia, Colombia Female: Same locality as male -357- rj [V3 M J tx .\ i w Figure 68 Lutzomyia (P.) guyanensis — A. Male head, B. Male flagellomere II, C. Female head, D. Male flagellomere II, E. Female cibarium and pharynx, F. Male genitalia, G. Body of spermatheca, H. Spermathecae, I. Female wing, J. Male wing, K. Female cibarium. Male: Almirante, Bocas del Toro Prov. , Panana Female: Curiche, Choco Dept., Colombia r- ft«S\ f\ Gj 0 Figure 69 Lutzomyia (P.) noctieola — A. Male head, B. Male flagellomere II, C. Female head, D. Female flagellomere II, E. Female cibarium and pharynx, F. Aedeagus and paramere, G. Male genitalia, H. Spermathecae, I, Proximal end of genital pump, J. Genital pump and filaments, K. Female wing, L. Male wing, M. Female cibarium (all figures from Young, 1973). Male: Rio Anori, Antioquia Dept., Colombia Female: Same locality as male -361- -^ it yftvs 7> ii'l B W » u Figure 70 Lutzormjia (P.) panamensis — A. Male head, B. Male flagellomere II. C. Female head, D. Female flagellomere II, E. Female clbarium and pharynx, F. Genital filaments, G. Male Genitalia, H. Spermathecae, I. Female wing, J. Male wing, K. Female cibarium. Male: Curiche, Choco Dept., Colombia Female: Teresita, Choco Dept., Colombia -363- Figure 71 Lutzowjia (P.) recurva — A. Male head, B. Male flagellomere II, C. Female head, D. Spermathecae, E. Genital pump and filaments, F. Female cibarium, G. Aedeagus and paramere, H. Female wing, I. Male ■•ring, J, Male genitalia. Male: Curiche, Choco Dept., Colombia Female: Same locality as male -365- -366- Vexator Group The odor 1965 Defined and discussed by Fairchild and Hertig (1957) and Theodor (.1965), this group is divided into 2 series — the series vexa£or(/i+ species, Central America and Nearctic Region) and the periiensis (14+ species, Central and South America). Most of the described species in the latter series occur in mountainous areas but a few including L. hart- manni, L. sanguinaria, and an undescribed Amazonian species from Ecuador inhabit lowland forests where they are strongly anthropophilic. Forattini (1971a, 1973) places hcrtmanni and sanguinaria in the sub- genus Triahophoromyia Barretto, gsnus Psychodopygus Mang., but this does not seem to reflect a natural arrangement. The females, unlike those of the Triahophoromyia spp. (in the sense of Sherlock and Guitton, 1970) have only 4 horizontal teeth in the cibarium and the males have 5 major spines on each style. The length of palp 5, variable in the vexator group spp., is relatively short in the Triahophoromyia spp. The single known fossil phlebotomine in the New World, L. paterna (Quate, 1963), was discovered in Mexican amber (late Oligocene or early Miocene). It, a male, differs very little from some extant species in the vexator' group (Quate, op. cit.). For the most part, the females in this group are keyed on the basis of external characters owing to similar cibaria and spermathecae, the latter being difficult to observe in per- manent mounts. Not being completely satisfied with this key, I recommend that associated males be on hand before confirming specific identifica- ■367- Keys to Species Kales 1. Coxite with 20+ setae forming a tuft or loosely arranged patch. . . 2 Coxite with fewer than 5 setae at base .4 2. Coxite setae loosely arranged. Genital filaments ca. 5X length of pump. Palpal segment 5 shorter than third. Flagellomere I shorter than or = to head height airrita (Fig. 72) Coxite setae forming a compact tuft on a subcircular base. Genital filaments shorter than 4X length of pump. Palpal segment 5 longer than third. Flagellomere I longer than head height 3 3. Palpal segment 5 shorter than segments 2-1-3. Paramere slender. Coxite tuft on a smaller subcircular base, its diameter less than greatest width of cercus striativilla (Fig. 76) Palpal segment 5 longer than segments 2+3. Paramere broader. Coxite tuft on a larger subcircular base, its diameter = to greatest width of cercus osomoi (Fig. 74) 4. Aedeagus and genital filaments with blunt tips . sanguinaria (Fig. 75) Aedeagus and genital filaments with pointed tips 5 5. Coxite tuft of 2-3 setae, one markedly thickened, over twice length of others and much wider at base. Style with all spines on apical half of structure soorzai (Fig. 77) Coxite tuft of 3-5 setae, none markedly thickened as above. Style with basal spine usually on proximal half of structure 6 -368- 6. Coxite tuft of 3-5 setae, one or more distinctly longer than others. Eyes separated by distance = to 5 or more facet diameters. Lab rum or palp 5 shorter than 0.28 mm hartmanni (Fig. 73) Coxite tuft of 3 setae subequal in length. Eyes closer together, separated by less than 4.5 facet diameters. Labrum or palp 5 longer than 0.30 mm sp. of Pichinde (Fig. 78) Females 1. Spermathecae carrot-shaped and large, each with a relatively broad terminal knob. Basitarsus of hind leg shorter than or = to length of hind femur • sanguinavia (Fig. 75) Spermathecae slender and smaller, each with a narrow, more protruding terminal knob. Basitarsus of hind leg longer than hind femur. . . 2 2. Labrum (< 0.43 mm long) shorter than head width. Combined length of palpal segments 1+2+3 shorter than 0.50 mm 3 Labrum (> 0.43 mm long) longer than head width. Combined length of palpal segments 1+2+3 over 0.50 mm 4 3. Flagellomere I (> 0.40 mm long) much longer than labrum. Palpal seg- ment 5 (> 0.25 mm long) = to or longer than segments 3+4 stvictivi-lla (Fig. 76) Flagellomere I (< 0.40 mm long) shorter, = to or but slightly longer than labrum. Palpal segment 5 (< 0.25 mm long) shorter than segments 3 + 4 havtmanni (Fig. 73) -369- 4. Flagellomere I longer than labrum osornci (Fig. 74) Flageliomere I shorter than labrum 5 5. Genital fork stem markedly enlarged at tip .... cirrita (Fig. 72) Genital fork stem subequal in width throughout sp. of Pichinde (Fig. 78) scorzai* (Fig. 77) 88. Lutzonvjia cirrita. (Fig/ 72) Lutzomyia cirrita Young & Porter, 1974, 321 (cf holotype, o, Rio Anori, Antioquia Dept,, Colombia). Distribution: Colombia (Antioquia, Valle) . Mats-rial examined: Colombia. 2 o*c?, 9 o (including holotype and allotype), type locality, other specifics by Young & Porter (1974). 1 <7, Anchicaya Dam (Valle), flight trap, 10 June 1975, R.C.W. Discussion: Other than biting man and being attracted to light traps (Young and Porter, op. cit.), this species is little known. The male, with the scattered coxite setae, is readily distinguished from other Colombian males in the vexator group. ''According to Ortiz (1965b), the length of palp 5 in Venezuelan scorzai females ranges from 0.28-0.33 mm. In the Colombian females it is gen- erally shorter, ranging from 0.22-0.29 mm, average length 0.26 mm, n = 9. The length of this segment in Lutzomyia sp. of Pichinde ranges from 0.31- 0.36 mm, average length 0.35 m, n = 9 so that in Colombia, at least, this difference may be helpful in separating the females of scorzai and Lutzomyia sp. of Pichinde, both of which occur together in Valle Dept., near Cali. -370- 89. Lutzomyia havtmanni (Fig. 73) Fhlehotomus havtmanni Fairchild & Hertig, 1957: 328 (6* holotype, o., Cerro Campaaa, Panama Prov., Panama). Johnson & Hertig, 1961: 765 et seq. (rearing data). Ortiz & Alvarez, 1963b: 321 (listed). McCcnnell &. Correa, 1964: 526 (infected with gregarines) . Ortiz, 1965b: 25 et seq. (listed, cf. to soovzai , measurements). Hanson, 1968: 63 (larva cf. to sariguinaria , figs.). Lutzomyia hartmanni: Barretto, 1962: 96 (listed). Barreto, 1969: 464 (Valle, Colombia). Christensen & Fairchild, 1971: 302 (Darien, Panama). Osorno et al., 1972a: 31 (Colombian records). Christensen, 1972a: 88 (listed). Young, 1973: 109 (Choco, Colombia). Young & Porter, 1974: 323 (cf. to civvita) . Psychodopygus havtmanni: . Forattini, 1971a: 105 (listed). Forattini, 1973: 123 et seq. (gen. review, figs.). Distribution: Panama, Colombia (Antioquia, Choco, Valle) , Ecuador. Material examined: Colombia. 16 6*6", 8 oo, Rio Anori (Antioquia), light traps, May 1970, C.H.P. 1 cf, 8 oo , same data but Sept. 1970, D.G.Y. 8 cW, 1,066 yy, Curiche (Choco), biting man, tree trunks, light, Malaise & Shannon traps, March-Dec. 1967, D.G.Y. 1 cf, 114 c^o. , same data but Alto Curiche (Choco), June-Nov. 1967. 2 cfcf, 13 oo., Teresita (Choco), biting man, tree trunks, Shannon traps, May-Oct. 1967, D.G.Y. 1 cf, Rio Raposo (Valle), light trap.y 4 May 1964, P. Barreto. 3 oV, 6 o^, Anchicaya Dam (Valle), light, flight, traps, tree trunks, 9-11 Aug. 1973, D.G.Y. & R.C.W. 8 op, 25 km E of Buenaventura (Valle), light traps, 12 Aug. 1973, D.G.Y. & R.C.W. Ecuador. 2 6'cf, 3 oo , 17 km E of Santa Domingo de los Colorados (Pichincha) , light and flight traps, May 1976, D.G.Y., T. Rogers, -371- (t. Falrchild. Panama. 1 6" (holotype no. 3997), type locality, light trap, 24 April 1952, F. Blanton. 1 p (allotype no. 2904), type locality, Shannon trap, 10 Feb. 1951, M. Hertig et al. 2 S3, 8 oc_, typa locality; Almiraute (Bocas del Toro) ; Canal Zone; Cerro Azul (Panama), various dates and collectors. Discussion: At Curiche (Choco) , this species represented nearly 20% (733 po) of all man-biting sand flies taken from April to November, 1967 (141 man hours). Only L. panamensis was more abundant in human bait collections, accounting for 46.5% (1751 ) of 3707 total females. According to Hanson (1968) , the larvae of hartmanni and sanguinaria are very close, differing only in the length of certain setae on the thorax and abdomen. The breeding sites of the havtmanni immatures and the diurnal resting sites of the adults remain virtually unknown. At Curiche, we collected 1 &* 2 op in tree buttresses but other resting sites were not sampled thoroughly. 90. Lutzomyia osornoi (Fig. 74) Phlebotomus osornoi Ristorcelli & Van Ty, 1941: 260 (o holotype, Valle de Capuli, Narino Dept., Colombia). Rozeboom, 1947a: 177 (6", dascr . ) - Phlebotomus montoyai Sherlock, 1962: 328 (ictivilla n. sp. (Fig. 76) Male (holotype) : Wing length 2.10; width 0.67. Whole insect dusky except for darker head, part of mesonotum, procoxae, and abdominal ter- gites 1-3. Head height 0.42; width 0.37. Eyes separated by 0.12 or by distance = to 6 facet diameters. Flageliomere I (0.50 long), slightly longer than II + III; ascoids simple, short, those on II not extending beyond middle of structure, on all f lagellomeres except last. Length of palpal segments: 1 (0.04), 2 (0.13), 3 (0.17), 4 (0.09), 5 (0.25); palpal sensilla (ca. 20), on distal two-thirds of segment 3. Labrum 0.25 long. Cibarium with ca. 20 vestigial teeth; pigment patch slender, it and chitinous arch nearly invisible, the latter seemingly complete. Pharynx 0.23 long, unarmed. Pleura with 11-12 upper and 5-6 lower episternal setae. Length of wing vein sections: Alpha (0.67), beta (0.17), delta (0.22), gamma (0.40). Length of femora, tibiae, and basitarsi: Foreleg, 1.12, 1.71, 1.20; midleg, 1.0, 1.94, 1.32; hindleg, 1.23, 2.33, 1.47. Abdominal sternites entire, no openings visible. Genitalia: Style 0.24 long, with 5 major spines, basal pair on different levels, not quite paired, no subterminal bristle. Coxite (0.41 long x 0.08 wide), with basal tuft of ca. 25 slender setae inserted on a cir- cular, raspberry-like base. Paramere simple, rather slender as shown. Aedeagus sub triangular, pointed at tip. Lateral lobe 0.33 long. Genital pump 0.12 long, each filament very thin, 0.43 long or nearly 3.6X length of pump. Cerci unremarkable. Female (allotype): Wing length 2.30; width 0.78. Coloration as in male. Head height 0.47; width 0.40. Eyes separated by 0.13 or by distance = to 6.3 facet diameters. Flageliomere I (0.4.1 long), slightly -379- longer than II + III; ascoids simple, longer than in 6* but not reaching end of flagellomere II, on all but last flagellomere. Length of palpal segments: 1 (0.05), 2 (0.15), 3 (0.19), 4 (0.09), 5 (0.27); palpal sensilla as in d1. Labrum 0.32 long. Cibarium with 4 straight, equi- distant horizontal teeth, ca, 15 small vertical teeth in an irregular transverse row; pigment patch slender, wider near vertical teeth; chitinous arch complete but hardly visible in middle. Pharynx 0.21 long. Pleura. with .10-11 upper and 3-4 lower episternal setae. Length of wing vein sections: Alpha (0.77), beta (0.16), delta (0.33), gamma (0.37). Length of femora, tibiae, and basitarsi: Foreleg, 0.98, 1.40, 0.95; midleg, 0.88, 1.50, 1.05; hindleg missing. Abdominal sternites as in d*. Tergite 8 with 2-5 setae on each side. Spermathecae and ducts as shown. Cerci unremarkable . Distribution: Colombia (Antioquia) . Material examined: Colombia. females, not exceeding 0.37 mm in 18 oo measured; whereas in stria Hvilla females it ranges from 0.41-0.48 mm in length (n = 7) . Of the vexator group species, L. otrietiviVla most closely resembles L. osowoi but that species is very large, the wing being 3.0 mm or longer in both sexes. In contrast to the osovnoi male, L. stvictivilla has relatively slender parameres, the coxite tuft is smaller and consists of fewer .setae. Other than size differences, I am unable to separate the females of stvictivilla and osornoi . The labrum of the latter exceeds 0.47 mm in length; whereas in stvictivilla females, the labrum is less than 0.39 mm long (0.32-0.38 mm, n = 7) . Future studies based on material from other localities may reveal that these taxa are distinct as subspecies or geographic variants but until, or unless, evidence to support this becomes available, 1 will con- tinue to treat them as distinct species. The specific name means "vile woman" and refers to the man-biting habit of the females, Dr. C.H. Porter having collected over 20 females attacking man at Rio Anori in 1970 and 1971 (pers. comra.) . 94. Lutzomyia sp. of P ichinde (Fig. 78) Distribution: Colombia (Valle) . Material examined: Colombia. 6 6V, 2 op, ca. 10 km W of Cali (1750 m) (Valle), tree trunks, rock crevice, 31 July 1973, D.G.Y. & R.C.W. 3 &f, 1 q, same data but 2 Aug. 1973. 1 d", 7 on, same locality ■381- but light trap, 2 Aug. 1973. 2 6*6*, ca. 15 km SW of Cali near Rio Pance (1600 m) (Valle), tree trunk, 7 Aug. 1973, D.G.Y. & S. Ayala. Discussion: Owing to the uncertain status of this taxon, I prefer to give it an informal name, hoping that future studies will firmly establish its position as a distinct species or as a geographic variant of L. aavtmanni . Lutzomyia sp. of Pichinde is larger than hartnanni, possibly re- flecting altitudinal differences as noted for B. galindoi and L. soovzai. Thus far, specimens of sp. of Pichinde have been taken only near Cali (Valle) above 1200 in. It has not been determined whether or not the females are anthropophilic. The coxite tuft of the male consists of 3 setae, all subequal in size, and palp 5 is always longer than palp 3. Both sexes were collected together in forests on tree trunks, in a rock crevice and once in a light trap. Lutzomyia hartmanni, an anthropophilic species most common in the lowlands of Colombia, has been taken as high as 950 m at Cerro Campana, Panama Prov. , Panama --- the type locality. Specimens from there, including the holotype and allotype, are larger than those from the lowlands but do not differ in structure. One seta of the coxite tuft is larger than the others. Palp 5 is variable, being less than, equal to, or but slightly longer than palp 3. This species is rarely found resting on tree trunks. A description of Lutzomyia sp. of Pichinde is given below. Both the male and female were taken together on a tree trunk near Pichinde on 31 July 1973. Male (no. 593): Wing length 3.09; width 0.98. Head, mesonotum and genitalia well pigmented, contrasting with rest of insect which is faintly pigmented. Head height 0.49; width 0.44. Eyes narrowly separated, -382- interocuiar distance = 0.76 or 3.9 facet diameters. Flagellomere I (0.50 long), 1.35X length of IT + III; ascoids simple, tips extending beyond middle of II, on all f lagelloraeres except last. Length of palpal segments; 1 (0.06), 2 (0.18), 3 (0.22), 4 (0.11), 5 (0.35); palpal sensilla (ca. 15) on distal two-thirds of segment 3. Labrum length 0.35. Cibarium with ca. 20 small remnants of teeth. Pigment patch slender, hardly visible; chitinous arch indefinite. Pharynx 0.24 long, unarmed. Pleura with 15 upper and 6 lower episterna] setae. Length of wing vein sections: Alpha (0.84), beta (0.35), delta (0.26), gamma (0.38). Length of femora, tibia, and basitarsi of slide no. 600: Foreleg, 1.07, 1.83, 1.24: midleg, 1.02, 1.93, 1.34; hindleg, 1.20, 2.30, 1.57. Abdominal sternite 2 not visible (other cfo* have paired lateral openings). Genitalia: Style 0.20 long, with 5 major spines, the most basal on proximal one-half of structure, no subterminal seta. Coxite (0.36 x 0.08 wide) with 3 monomorphic, thin setae at base, none longer than greatest width of coxite. Aedeagus subtriangular, pointed at tip. Lateral lobe 0.31 long. Genital pump 0.15 long, each filament 0.51 long or 3.4X length of pump. Cercus as shown. Female (no. 599): Wing length 3.29; width 1.1. Coloration as in 3. Head height 0.60; width 0.51. Eyes separated by 0.13 or by distance = to 6.2 facet diameters. Flagellomere I (0.53 long), 1.32X length of II + III; ascoids as in o but slightly longer as shown. Length of palpal segments: 1 (0.07), 2 (0.28), 3 (0.31), 4 (0.13), 5 (0.35); palpal sen- silla as in Para States. Brazil). Barretto, 1946b: 527 (as synonym of baduelensis, refs.). Fairchild & Hertig, 1948a: 255 (as synonym of trinidadensis) ■ PhZebotomus baduelensis Floch & Abonnenc, 1944d: 1 (, Almirante (Bocas del Toro) , light traps, Nov. 1952-Jan. 1953, W. Hils. coll. 1 6", Rio Platanar (Panama), hollow tree with bats, 20 July 1950, R. Harmann. Trinidad. 5+ 6V, 1 Q_, Nariva Swamp, Bush Bush Forest, various dates, T.H.G. Aitken. Discussion: Lutzomyia pilosa and L. chassigneti are very similar, the male of the former species having a distal patch of coxite setae, lacking in chassigneti. The pilosa female, known but undescribed, is described here based on 2 specimens from Alto Curiche (Choco) Colombia and 1 from Almirante (Bocas de Toro), Panama. As Theodor (1965) mentioned, the yilosa and chassigneti females are indistinguishable. Female: Wing length 1.64-1.80; width 0.49-0.57. Head, mesonotum, abdominal tergites moderately pigmented, rest of insect dusky, pleura slightly paler than mesonotum. Head height 0.32-0.35; width 0.30-0.32. Eyes separated by 0.11 or by distance = to ca. 5.5 facet diameters. Rlagellomere I (0.25-0.27 long), nearly 1.2 x length of II + III; ascoids sinple, those on flagellomere II reaching to or beyond end, on all flagellomeres except last. Length of palpal segments: 1 (0.03), 2 (0.09-0.11), 3 (0.14-0.16), 4 (0.10), 5 (0.34-0.37); palpal sensilla at -434- apex of segment 3. Labrum 0.15-0.16 long. Cibarium as figured. Pharynx 0.13-0.14 long, unarmed. Pleura with 5-10 upper and 3-6 lower episternal setae. Length of wing vein sections: Alpha (0.34-0.47), beta (0.29-0.31) delta (0.15-0.25), g aroma (0.18-0.20). Length of femora, tibiae, and basitarsi of slide 4276 (Almirante) : Foreleg, 0.71, 0.71, 0.39; midleg, 0,67, 0.83, 0.46; hindleg, 0.73, 0.98, 0.54. Abdominal sternite 2 entire, without openings, sternite 2, at least, with paired circular openings at sides. Tergite 8 with 5-10 setae on each side. Spermathecae, ducts, and cerci as shown. Figure 86 LubzomyLa pilosa — A. Male head, B. Male f lagellomere II, C. Sperma- thecae, D. Female head, E. Female flagellomere II, F. Male genitalia, G. Genital pump and filaments, H. Tip of genital filament, I. Female cibarium and pharynx, J. Female wing, K. Male wing, L. Female cibarium. Male: Alto Curiche, Choco Dept., Colombia Female: Same locality as male -436- -437- Ungrouped Species 103. Lutzomyia nordestina (Fig. 87) Phlebotomus novdestinus Mangabeira, 1942e: 327 (o holotype, Nova Olinda, Ceara, Brazil). Barretto, 1947: 215 (refs.). Damascene et al . , 1949: 830 (Brazilian records). Barretto, 1951: 221 (distrib.). Floch & Abonnenc, 1952: 37, 45 (o*, o keyed), 145 (d*, 9, redescr., figs.). Rodriguez, 1956: 76 et seq. (Ecuador). Hanson, 1961: 321 (breeding site, Panama). Johnson & Hertig, 1961: 765 et seq. (rearing data. Thatcher & Rertig, 1966: 52 (in burrows, Panama). Pklebotonras longieovnutus Floch & Abonnenc, 1943: 6 (d holotype, p, Montabo, French Guiana). Barretto, 1946b: 534 (as synonym of nordestina). Forattini, 1960: 478 (Amapa, Brazil). Lutzomyia nordestina: Barretto, 1962: 96 (listed). Martins et al. , 1962a: 381 (Goias, Brazil). Theodor, 1965: 195 (characterized). Martins et al., 1965: 3 (Rondonia, Brazil). Martins et al., 1970: 553 (mention). Tesh et al., 1971a: 152 (blood meals). Chaniotis et al., 1971a: 344 et seq. (pop. dynamics, Panama). Chris tensen & Fair child, 1971: 301 (Darien, Panama). Osorno et al., 1972a: 65-66 (Colombian records). Christensen, 1972a: 83 (listed). Christensen et al . , 1972: 57 (collecting data). Chaniotis et al . , 1972: 95 (resting sites). Christensen & Herrer, 1973: 579 (collecting data, Panama). Lewis, 1975a: 504 et seq. (mouthpart morphol . ) . Llanos et al . , 1976: 480 (Peru). Martins et al., 1976b: 496 (Peru). -438- Psychodopygus noraestina: Forattini, 1971a: 105 (listed). Forattini, 1973: 140 et seq. (gen. review, figs.). Distribution: Panama, Colombia (Amazonas, Antioquia, Boyaca, Caqueta, Choco, Tolima, Valle) , Ecuador, Panama, Peru, French Guiana, Trinidad, Brazil. Material examined: Colombia. 1 o, 17 km W of Leticia (Amazonas), burrow, 26 July 1973, D.G.Y. & R.C.W. 1 o_, Rio Anori (Antioquia), light trap, 3 May 1970, C.H.P. 4 do*, same locality, tree trunks, Sept. 1970, D.G.Y. 1 0, Puerto Boyaca (Boyaca), light trap, 10 Aug. 1971, C.J.M. 6 $6\ 16 00, Curiche (Choco), light, Malaise & Shannon traps, May-Oct. 1967, D.G.Y. 1 6*, 7 OO, Alto Curiche (Choco), light & Shannon traps, tree trunk, June-Sept. 1967, D.G.Y. 1 6*, Anchicaya Dam (Valle), tree trunk, 9 Aug. 1973, D.G.Y. & R.C.W. 2 6*0*, 1 o., 25 km E of Buenaventura (Valle), light & flight traps, 12 Aug. 1973, D.G.Y. & R.C.W. Brazil. 2 do, 5 00, 27 km SE of Maraba (Para), light traps, 26-28 Sept., 1972, D.G.Y. & H. Fraiha. 5 cfcf, 1 o, same locality, burrow in leaf-cutting ant nest, 26 Sept. 1972, D.G.Y. 1 6*, near Altamira (Para), tree trunk, 1 Oct. 1972, D.G.Y. 1 o_, Belem (Para), flight trap, 2 Aug. 1974, D.G.Y. 1 o, near Bacuri, N of Maraba (Para), light trap, 28 Oct. 1974, J. Reinert. 2 6*6*, 5 00, Rio Napo at Limoncocha (Napo) , light & Flight traps, 19-24 May 1976, D.G.Y. & T. Rogers. 2 00., 17 km E of Santa Domingo de los Colorados (Pichincha) , burrow, 4 May 1976, D.G.Y. & T. Rogers. Panama. 12 c?cf, 24 00, various dates and localities in the Canal Zone, Colon, 4 Panama Provinces. Trinidad. 1 o_, Nariva Swamp . Bush Bush Forest, light trap, 2-3 Nov. 1965, T.H.G. Aitken. Discussion: Lutzomyia nordestina, a widespread species occurring on both sides of the Andes in Colombia and Ecuador, is often found resting -439- in animal burrows. There is some evidence indicating that females feed on cold-blooded vertebrates (Tesh et al., 1971a). 1 have examined females of an undescribed Lutzomyia sp. from Aitamira (Para), Brazil (km 164), which resembles that of nordestina in most nonsexual and sexual characters but the cibarial armature is quite different. Other than this species, L. nordestina does not seen to be closely allied to any other described taxon. 104. Lutzomyia rangeiiana (Fig. 88) Phlebotomus rangelianus Ortiz, 1952: 153 (6* holotype, Duaca, Lara State, Venezuela). Pifano et al., 1962: 384, 389 (6\ o_, keyed), 406- 407 (M V V- Figure 89 Lutzomyia sp. of Anchicaya male — A. Head, B. Wing, C. Genital pump and filaments, D. Flagc-;] ioroere. II, E. Genitalia. Kale: r>:ich i.caya Darn, Val le Dept., Colonbia -film # » \ \ i . .» ! U 1 t — ? M i i / : _y" // v\- SYNOPSIS OF NEW TAXA, SYNONYMS, FEMALE DESCRIPTIONS, AND NEW DISTRIBUTION RECORDS Taxa Sycorax andicola Sycorax aolombiensis Sycorax faivchi Idi Sycorax trispinosa Luizomyia (L.J marinkellei Lutzomyia moralesi Lutzomyia, barrettoi majuscula Lutzomyia (T. ) howardi Lutzomyia (T.) cellulana Lutzomyia (T. ) saltuosa Lutzomyia (P. ) carrerai thul-a Lutzomyia. (P. ) of Tres Esquinas Lutzomyia sp. of Pichinde Lutzomyia sp. of Anchicaya Lutzomyia tintinnabula = L. ayrozai Lutzomyia gasti - L. waVkeri ■male Descriptions Lutzomyia (L.) bifoliata Lutzomyia (P. ) hirsuta r.icaraguensis Lutzomyia pilosa New Distribution Records COLOMBIA Brumptomyia hamata Brumptomyia leopo~ldx)i Lutzomyia baityi Lutzomyia (P.) bispinosa Lutzomyia carpenteri Lutzomyia dreisbachi Lutzomyia (Pr.J dysponeta Lutzomyia isovesp>e.rti lionis Lutzomyia rnigonei Lutzomyia nunsztovari Lutzomyia scorzai -449- COSTA RICA Lutzomia piu CUBA Lutzomyia oayennensis ECUADOR Bvumptomyia galindoi Lutzomyia abunaensis Lutzomyia aalydifeva Lutzomyia baity i Lutzomyia (P. ) bispinosa Lutzomyia (P. ) cavrerai oarrerai Lutzomyia (P. ) cavrevai thula Lutzomyia (P, ) davisi Lutzomyia dendrophy'uz Lutzomyia (II.) flavisoutellata Lutzomyia guyanensis Lutzomyia hartmanni Lutzomyia (P.) hirsuta kirsuta Lutzcrryia micropyga Lutzomyia nevesi Lutzomyia (P.) nocticola Lutzomyia (S. ) olmeaa bicolor Lutzomyia (P. ) pananensis Lutzomyia (T. ) rebwra Lutzomyia ur.dulata Lutzomyia walker I Lutzomyia (N.) yuilli GUATEMALA Lutzomyia (II.) ylepkiletov HONDURAS T'Utzomyia evansi Lutzomyia texawa -450- PANAtlA Lutzomyla (P. ) noa Lutzomyia vangeVLa. 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BIOGRAPHICAL SKETCH David Grier Young was born May 9, 1940, at Dayton, Ohio. In June, J.953, he was graduated from Oakwood High School, Dayton, Ohio. In Decem- ber, 1962, he received the degree of Bachelor of Science with a major in entomology from the University of Florida. From i.963 to 1968, he served as a commissioned officer in the Medical Service Corps, U.S. Army at Ft. Sam Houston, Texas, and the Panama Canal Zone. Following his dis- charge, he worked on a research project involving siudli mammals and ectoparasites in Ohio (Dayton Museum of Natural History) In 1968, be enrolled in the Graduate School of the University of Florida, receiving the degree of Master of Science in June, 1971, with a major in entomology, From 1972 to December, 1977, he Las been co-investigator and principal investigator of a U.S. Army Medical Research and Development contract dealing with the systematics of phlebotomine sand flies. The author is a member of the American Association for the Advance- ment of Science, the Entomological Society of America, and the Florida Entomological Society. He is a research associate of the Florida State Collection of Arthropods. David Young is married to the former Marianna Brown. They have twin sons, Alexander and Mark. -48.1 1 certify that I have read this study and that in my opinion it coniorns Co acceptable standards of scholarly presentation and is fully adequate, i;i scope and quality, as a dissertation for the degree of Doctor of Philosophy. ^7 ,/X Associate Professor of Microbiology This dissertation was submitted to the Graduate Faculty of the College of Agriculture and to the Graduate Council and was accepted as partial fulfillment of the requirements lor the degree of Doctor of Philosophy. December 1977 vok dC.