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VOLUME XIII 1977 11 CONTENTS Editorial — Comments About a Few Words in the Biological Sciences 1 Rempel, Heming and Church — The Embryology of Lytta viridana LeConte (Coleoptera: Meloidae). IX. The Central Nervous System, Stomatogastric Nervous System, and Endocrine System 5 Nimmo — The Adult Trichoptera (Insecta) of Alberta and Eastern British Columbia, And Their Post-Glacial Origins. I. The Families Rhyacophilidae and Limnephilidae. Supplement 1 25 Nimmo — The Adult Trichoptera (Insecta) of Alberta and Eastern British Columbia, And Their Post-Glacial Origins. II. The Families Glossosomatidae and Philopotamidae. Supplement 1 69 Belicek — Corrigenda on Coccinellidae of Western Canada and Alaska with Analyses of the Transmontane Zoogeographic Relationships Between the Fauna of British Columbia and Alberta (Insecta: Coleoptera: Coccinellidae) 73 Book Review — Matsuda, R. 1976. Morphology and Evolution of the Insect Abdomen 75 Evans — Geographic Variation, Distribution and Taxonomic Status of the Intertidal Insect Thalassotrechus barbarae (Horn) (Coleoptera: Carabidae) 83 Mendez — Mammalian-Siphonapteran Associations, The Environment, and Biogeography of Mammals of Southwestern Colombia 91 Sengupta — Changes in Acetycholinesterases and Cholinesterases During Development of Aedes aegypti (L.) (Diptera, Culicidae) 183 Book Review — Horn, D.J. 1976. Biology of Insects 191 Craig — Mouthparts and Feeding Behaviour of Tahitian Larval Simuliidae (Diptera: Nematocera) 195 Fredeen — A Review of the Economic Importance of Black Flies (Simuliidae) in Canada 219 Halffter — Evolution of Nidification in the Scarabaeinae (Coleoptera, Scarabaeidae) ... 231 Picchi — A Systematic Review of the Genus Aneurus of North and Middle America and the West Indies (Hemiptera: Aradidae) 255 Evans & Baldwin — Larval Exuviae of Attagenus bicolor von Harold (Coleoptera: Dermestidae) from an Archeological Site at Mesa Verde, Colorado 309 Book Review — Wiggins, G.B. 1977. Larvae of the North American Caddisfly genera (Trichoptera) 311 Steiner — Observations on Overnight Perch Constancy by a Female Digger Wasp, Ammophila azteca Cameron (Hymenoptera: Specidae), In Captivity 315 Fredeen — Black Fly Control and Environmental Quality with Reference to Chemical Larviciding in Western Canada 321 Griffiths — Studies on Boreal Agromyzidae (Diptera). XIII. Some Phytomyza and Chromatomyia Miners on Cichorieae (Compositae) 327 Reichardt — A Synopsis of the Genera of Neotropical Carabidae (Insecta: Coleoptera) 346 Editor’s Acknowledgements 495 Ill CORRIGENDA: Quaestiones Entomologicae, Volume 13 Mendez, E., Mammalian-Siphonapteran Associations, , pp. 91 — 182 page/line 112/ 28 112/ 35 158/ 35 166/ 30 166/ 52 change “ cincereus ” to “ cinereus ” change “Dyplomys” to “ Diplomys ” change “Aletes” to “ Ateles ” change “ gumnurus ” to “ gymnurus ” change “ pradoi ” to “ pardoi ” Craig, D.A. Mouthparts and Feeding Behaviour pp. 195-218 page/line 209/ 1 1 210/ 11 change “Corvert” to “Couvert” change “Aedes aegypti (1.) to “Aedes aegypti (L.)” Fredeen, F.J.H. A Review of the Economic Importance page/line 220/ 13 226/ 6 pp. 219-229 Change “Stone (1965)” to “Stone et al (1965)” Change “Onchocerea” to “ Onchocerca ” Halffter, G. Evolution of Nidification in the Scarabaeinae , pp. 231-253 page/line 247/41 249/ 1 change “(Croprini)” to “(Coprini)” change “(Eurysternina, Scarabaeini)” to “(Eurysternini, Scarabaeinae)” Picchi, V.D. A Systematic Review of the Genus Aneurus .............. pp. 255-308 page/line 255/ 1 259/ 31 272/ 34 272/ 40 281/41 295/ 1 change “ Curtis , 1 81 8” to “ Curtis , 1828 ” change “Stys (1974)” to “Stys (1974)” change “A tenuicornis ” to ‘A. leptocerus” change “Usingeri” to “Usinger” change “Stal 1873” to “Stal 1873” change “ vaurieri ” to “ vauriei ” 308/ 1 change “hispaniolensis” to “hispaniolensis’ Griffiths, G.C.D. Studies on Boreal Agromyzidae page/line 333/ 32 333/ 35 pp. 327-345 change “ Agoseris ” to “ Aposeris ” change “ Agoseris glauca” to “ Aposeris glauca” IV INDEX Abbott, D.P. ,84, 85,87 Acacia melanoceras, 104 Acarina, 182 Acer rubrum, 282 acety cholinesterases, 75, 183—189 A chat o carp us n igri ca ns, 102 Acrididae, 77 acropedes group, Rhyacophila, 27, 28 acropedes, Rhyacophila, 28 Adoratopsylla, 173 Adoratopsylla intermedia copha, 1 16, 117, 122, 166 Adoratopsylla (T.) intermedia, 166, 172, 174, 176 Adoratopsyllini, 117 aduncun, Piper, 104 Aedes aegypti, 75, 183-189, 210 aegypti, Aedes, 75, 183—189. 210 Aepopsis robinii, 84 Aepus marinus, 84 aequalis, Dolophilodes, 69 Agonini, 87 Agouti paca, 112, 142, 158, 166 Agouti paca quanta, 110 Agromyzidae, 327, 338, 339, 340, 341 aibonitensis, Aneurus, 255, 256, 263, 266, 268, 279, 289, 290, 291, 292, 293, 295 308 Akodon, 117, 140, 141, 176 Akodon pulcherrimus inambari, 141 Akodont, 132 alascense, Chilostigma, 46 alascensis, Chilostigma, 46 alascensis, Halesus, 46 alascensis, Platyphylax, 46 alascensis, Psychoglypha, 25, 65, 46, 47, 49 alba, Philo casca, 25, 45, 49, 50, 65, 67 albicans, Miconia, 105 albiceps, Phytomyza, 321 , 328, 329, 332, 334,337,338 albiforum, Hieracium, 333 albigularis, Oryzomys, 91, 110, 117, 124, 127, 131, 132, 139, 140, 141, 166 alfaroi, Oryzomys, 117, 132, 142, 166 allamandi, Galictus, 142 Allen, J.A., 113, 178 Allium cepa, 105 almifolia, Turnera, 102 Alouatta, 243 alpicola, Ixeris, 336 alpina, Cicerbita, 330, 331 alvatus, Limnephilus, 25, 27, 39, 40, 48, 50 51,63,67 Alysiinae, 339 americana, Mazama, 112, 113 americana, Periplaneta, 1 5 americana, Persea, 104 ammonites, 97 Ammophila azteca, 315, 316, 318, 320 breviceps, 319 pubescens, 318 amphibians, 1 1 1 amplexicaule, Hieracium, 331 Anabolia fusorius, 42 modesta, 42 Anabolia ( Asynarchus) fusorius, 42 Anacardium excelsum, 104 anatis, Leucocytozoon, 226, 229 Ancistropsyllidae, 180 Anderson, J.R., 226, 228 Anderson, N.H., 28, 34, 37, 53 Anderson, R.C., 226, 228 andinus, Eptesicus, 111 andinus, Sylvilagus, 112, Andosols, 100 Andres, V., Jr., (See Ellman, G.L.), 184, 186 Andropogon sp., 104 Anduaga, S. (see Halffter, V.), 238, 253 Aneuridae, Aneurinae, 288 Aneurosoma, 255, 270 Aneurus aibonitensis, 255, 256, 263, 266, 268, 277, 289, 290, 291, 292, 293, 295, 308 arizonensis n. sp., 255, 263, 266, 269, 275, 279. 280. 289. 290. 291. 292. 293. 295, 298,301,306 barberi, 263, 268, 289, 290, 291, 292, 298, 301, 308 borealis, n. sp., 255, 263, 266, 270, 282, 289, 290. 291. 292. 293. 295, 298, 301, 308 championi, 262, 266, 268, 278, 289, 290, 291, 292, 293, 296, 298, 301, 307 deborahae, n. sp., 255, 263, 266, 269, 274, 275, 289, 290, 291, 292, 298, 301 306 V Aneurus dissimilis, 255, 262, 263, 265, 266, 267, 270, 276, 289, 290, 291, 292, 293, 296,299,301,307 fiskei, 262, 266, 267, 269, 280, 281, 282, 289. 290. 291. 292, 293, 296, 299, 301. 306 froeschneri, 263, 265, 266, 269, 280, 289, 290. 291. 292, 293, 296, 299, 301, 307 gallicus, 288 haitiensis, 255, 256, 263, 266, 268, 273, 274, 289, 290, 291, 293, 296, 299, 302, 308 hispaniolensis n. sp., 255, 256, 263, 265, 266, 268, 273, 274, 289, 290, 291, 292, 299, 302, 308 hrdyi, 257, 263, 267, 288 inconstans, 259, 260, 263 266, 269, 271, 272, 280, 281, 282, 283, 289, 290, 292, 293, 296, 299, 302, 305 leptocerus, 262, 265, 266, 270, 272, 289, 290, 291, 292, 294, 296, 299, 302, 307 maryae, n. sp., 255, 263, 266, 270, 283, 289. 290, 291, 292, 295, 296, 299, 302. 307 minutus, 255, 263, 266, 269, 274, 275, 276, 277 , 278, 279, 280, 289, 290, 291, 292, 294, 296, 299, 302, 307 montanus, 263, 266, 270, 285, 289, 290, 291. 292. 294. 296. 299. 302, 308 nasutus, 263, 268, 286, 289, 290, 291, 292, 294, 296, 308 neojamaicensis n. sp., 255, 256, 263, 266, 268, 289, 291, 292, 294, 297, 300, 302. 308 patriciae n. sp. 254, 255, 263, 266, 267, 270, 274, 276, 287, 289, 290, 291, 292, 294, 297, 300, 302, 308 pisonae, 257, 268, 277, 308 politus, 263, 266, 267, 269, 271, 272, 278, 289, 290, 291, 292, 294, 297, 300. 302, 306 pusillus, 263, 266, 268, 277, 284, 292, 300, 302, 307 pygmaeus, 255, 263, 266, 269, 275, 276, 304, 307 roseae n. sp., 255, 263, 265, 266, 269, 279, 289, 290, 291, 292, 294, 297, 300. 303, 306 septentrionalis, 255, 278 Aneurus simplex, 255, 263, 266, 268, 278, 279, 289, 290, 291, 292, 294, 297, 300, 303, 305 slateri, n. sp., 255, 263, 266, 268, 285, 289, 290, 291, 292, 295, 297, 300, 303, 306 tenuis, 263, 266, 270, 272, 273, 289, 290, 291. 292. 295. 297. 300. 303. 308 usingeri, n. sp., 255, 263, 266, 269, 284, 285, 286, 289, 290, 291, 292, 295, 297, 300, 303. 308 vauriei, 256, 263, 266, 268, 273, 274, 275, 289, 290, 291, 292, 295, 298, 301, 303, 308 veracruzensis, n. sp., 255, 263, 265, 266, 269, 275, 285, 286, 289, 290, 291, 292, 295, 298, 307 wygodzinskyi, n. sp., 255, 263, 265, 266, 270, 271, 272, 289, 290, 291, 292, 295, 298, 301.303.308 angiosperm, 106 Anisogamus modes tus, 42 annato, 102 annectens, Lutra, 110, 112 Annelids, 14, 81 ant-eaters, 107 Anthomyidae, 83 Anthon, H. 197, 198,209 Aotus lemurinus, 1 1 1 trivirgatus, 1 1 1 Aphodiinae, 232 aposeridis, Phytomyza, 327 , 334 Aposeris foetida, 334 glauca, 333 apple tree, 282 aquilegiae, Phytomyza, 332 Aradidae, 255—308 Aradus sanquinosus, 281 Archaeopsyllini, 164 archhieracii, Phytomyza, 337 , 338 Archhieracium sp., 337 arcticum, Simulium, 219 — 224, 228, 229, 321-325 Arctopora pulchella 312 Arctopsychidae, 3 1 2 argenteus group, Limnephilus,, 41 argent eus, Limnephilus, 41 arizonensis, n. sp., Aneurus, 255, 263, 266, 269, 275, 279, 280, 289, 290, 291, 292, 293, 295, 298,301,306 armadillos, 107 VI armatum, Inophleum, 1 04 armiger, Cephalodesmius, 245 Arnason, A.P. 220, 228, 321, 324 Arnason, A.P. (See Fredeen, F.J.H.), 221, 228, 325 Arnason, A.P. (See Rempel, J.G.), 221, 222, 229 Artemisia, 339 Artibeus, cinereus, 112 jamaicensis, 1 1 0 lit ur at us, 110 arvensis, Sonchus, 330, 336 arvensis, uliginosus, Sonchus, 336, 337 aspen, 282 asper, Sonchus, 330, 334, 336 Astereae, 339 asteris, Chromatomyia, 335 atribarba, Crepis, 334 Asynarchus fusorius, 42 lapponicus, 25, 42, 48, 50, 51, 64, 57 modestus, 42 rhanidophorus, 42 Ateles fuscipes, 112, 158, 166 atricornis, Phytomyza, 339 atropurpureus tomentosus, Senecio, 334 Atta mexicana, 238 Attagenini, 309 Attagenus bicolor, 309, 310 audouini, Kenodactylus, 84, 86 Aulacopris maximus, 247 aurantiacum, Hieracium, 331 aureurn, Simulium, 228 aureus, Thomasomys, 110, 117, 121, 166 auripendulus, Eumops, 112 australis, Heteromys, 112, 166 A ustrosimulium ( A ustro simulium) tillyardianum, 209 autumnalis, Leontodon, 330 autumnalis n. sp., Rhyacophila, 25, 30* 34, 50, 51, 58, 60, 66 Avicennia marina, 104 avocado, 104 Axelrod, D.I. (See Raven, P.H.), 106, 107, 181 azarae, Didelphis, 111, 117, 121 azteca, Ammophila, 315, 316, 318, 320 Babock, E.B., 335, 338 Babers, F.H. and J.J. Pratt, Jr., 185, 186 Baccharis sp., 105 bacteria, 241 bairdii, Tapirus, 112 Baker, R.H., 108, 178 balachowskyi, Eurysternus, 249, 250 Baiba, M.H. (see Fredeen, F.J.H.), 325 balbisiana, Weinmannia, 105 Balduf, W.V., 34, 53 Baldwin, S.J., 309 Ball, G.E., 84, 87 Ball, G.E., and J. Negre, 86, 87 balsa, 102, 104 Banks, N., 34, 42, 44, 46,53 Baranov, N., 197, 198, 209 barbara, Eira, 112, 142, 166 barbarae, Thalassotrechus, 75, 83, 84, 85, 86, 87,88,89,90 Barber, H.G.,271,287 barberi, Aneurus, 263, 268, 286, 289, 290, 291, 292, 298, 301, 308 Barreropsylla, 176 Barrow, J.H. (see Herman, C.M.), 227 , 229 Basak, S.(see Sengupta, R.), 183 Bassaricyon gabbi medius, 112 bats (see Chiroptera) Beal, R.S., Jr., 309 bears, 107 Beckman, L., 184, 186 beebei, Polygenis roberti, 11.3, 116, 137, 140, 141, 154 beebi, Rhopalopsyllus, 140 Seek, K.J. and D.L. Bramao, 98, 178 beech tree, 282 beetles (see Coleoptera) beetles, dung, 252, 253 Befaria sp., 105 Beiger, M., 331 , 334, 338 Bembecinus godmani, 315 neglect us, 318 Bembicini, 318 Bendell, J.F., 227, 228 Bennett, G.F., 226, 228, 227 Bennett, G.F. (see Fallis, A.M.) 226, 227, 228 Bennett, G.F. (see Greiner, E.C.), 226, 229 Bennett, G.F. (see Laird, M.), 227, 229 Berck, B. (see Fredeen, F.J.H.)., 325 Bergroth, E., 256, 270, 271, 275, 278, 281, 287 Betten, C., 34, 42, 46, 53 biclavata, Twinnia, 204 bicolor, A ttagenus, 309, 310 Vll bicolor, Malagoniella, 247 biennis, Crepis, 331 bifila, Rhyacophila, 33 big belly tree, 104 bilobatum, Uroderma b., 112 birch 282 bispinus, Canthon, 247 Bixa orellana, 1 02 blackberries, 105 black rubber, 102 Blatchley, W.S., 271, 272, 275, 278, 281, 287 Blattella germanica, 77 Blepharoceridae, 209 Bocconia frutescens, 104 bogotensis, Sigmodon hispidus, 110, 112 bohlsi, Polygenis bohlsi, 116, 132, 142, 166, 172, 174 bohlsi, Pulex, 1 32 Bombycillidae, 226 Bombyx mori, 13, 15 bonariensis, Eumops, 1 1 0 Bonasa umbellus, 228 bonasae, Leucocytozoon, 228 borealis, n. sp., Aneurus, 225, 263, 266, 270, 282, 289, 290, 291, 292, 293, 295, 298, 301, 305 Bornemissza, G.F., 242, 251, 252 Borrero, J.I., 113, 178 boucardi, Copris, 243 bourgaei, Cicerbita, 331 brachinus, Polygenis, 138 brachiopods, 97 Braconidae, 339 Brady pus griseus, 1 1 2 brasiliensis, Eplesicus, 1 1 1 brasiliensis, Sylvilagus, 110, 111, 112 brasiliensis, Tadarida, 1 1 0 b/eviceps, Ammophila, 319 brevistyla, Rhizophora, 104 Briegal, H., 183, 186 Britton, W.E., 270, 278, 28 1 , 287 Brown, A.W.A. (see Arnason, A.P.), 220 228, 324 Brown, A.W.A. (see West, A.S.), 225, 229 bryozoans, 97 Brunellia sp., 105 brunneus, Zygodontomys brevicauda, 110 Buhr, H., 330, 331,332, 338 Burgl, H., 96, 97, 178 Burgl, H. (see Jacobs, C.H.), 97, 98, 180 buttonwood forest, 104 C. minus cuius, 40 Cabrera, A., 113, 178 Cabrera, A., and J. Yepes, 108, 113 cabuyas, 102 cacicus, Rhopalopsyllus saevus, 113, 115, 158, 162, 166, 172, 174 cactus, 275 Caenolestes, \16 fuliginosis, 124 obscurus, 113, 121, 124, 127, 131, 132, 166 Caenolestidae, 166 caespitosum, Hieracium, 331 Cafius 88 calabash, 102 Calamagrotis, 105 Calandra oryzae, 1 6 Calathus ruficollis, 86 Calhoun, E.H., 183, 186 Cali Virus Laboratory, 93, 178 calmnosus, Oryzomys, 111, 117, 121. 132, 137, 138, 140, 141, 142, 158, 164 caliginosus, Oryzomys (Melanomys), 140, 166 Caluromys derbianus derbianus, 108, 112 camels, 107 Cameron, A.E., 220, 221, 228 canadensis, Lactuca, 337 candezei, Megathoposoma, 247, 253 canaster, Galictis vittata, 1 1 2 cancrivora, Didelphis, 1 40 cancrivorus, Procyon, 1 12 candace, Nasua nasua, 1 1 0 canescens, Lantana, 102 caniceps, Diplomys, 1 1 2 Canidae, 107, 166 Canis familiaris, 142, 166 Canthidium, 241 Canthon bispinus, 247 cyanellus, 241, 243, 248, 252 cyanellus cyanellus, 247, 248 edentulus, 247 muticus, 247 virens, 247 Canthonina, 233, 235, 237, 242, 243, 245, 247 capers, 105 Vlll capillaris, Crepis, 331 Caprifoliaceae, 339 capucinus, Cebus, llz Carabidae, 75, 83, 84, 86, 87, 88 Carabidae, Neotropical-Indices, 479-493 caracasana, Euphorbia, 102 caracasana, Wigandia, 104 Carausius (=Dixippus) morosus, 6, 10, 12, 13, 15, 16 carbonero, 105 Carludovia palmata, 104 Carayon, J., 315, 318 carnivores, 107, 164, 166, 175 carolinus, Dichotomius, 238, 240, 242 Carollia castanea, 1 1 2 subrufa, 1 12 cascarillo, 105 cashew, 104 Casida, J.E., 185, 186 Cassia sp., 105 Cassiope mertensiana, 335 Castella elastica, 102, 104 Cathartidae, 226 cattle, 221, 222, 223, 224,238, 231, 242 catus, Felis, 164, 166, 175 caucae, Mormosa improvida, 110 caucensis, Polygenis n. sp., 91, 116, 137, 142, 144, 145, 166, 172, 174 caucenis, Scuirus, 1 1 1 Cavanillesia platani folia, 104 Cebidae, 166 Cebus capucinus, 112 Cecropia spp., 104 cedar, 279 Cedrela spp., 102 cedrillo, 105 Ceiba pentandra, 102 cepa, Allium, 105 Cephalocereus colombianus, 102 Cephalodesmius, 235, 243, 245, 247, 252 armiger, 245 laticollis, 245 quadridens, 245 cephalopods, 97 Ceratophyllidae, 131, 173, 180, 181 Ceratophyllinae, 131 Ceratophylloidea, 1 1 7 Ceratophyllus equatoris, 1 3 1 peripinnatus, 131 Cerdocyon thous, 113, 164, 166 Cespedesia macrophylla, 1 04 Cetraria sp. 104 Chabaud, A.G. 93, 178 Ceratopogonidae, 226 cervids, 241 cervipedis, Onchocerca, 226 Chalcidoidea, 333 Champion, G.C., 270, 271, 272, 275, 278, 285,288 championi, Aneurus, 263, 266, 268, 278, 289, 290, 291, 292, 293, 296, 298, 301, 307 Chance, M.M., 196, 197, 198, 201, 204, 205, 206, 209 cheesmanae, Simulium, 195, 196, 207 chickens, 222, 227 chayote, 104 cheopis, Pulex, 1 58 cheopis, Xenopsylla, 93, 1 14, 158, 164, 166, 167, 172, 174, 175 cherry tree, 105 cinctus, Oniticellus, 237, 251, 252, cinerascens, Liponeuria, 209 cinqulatus, Dysdercus, 1 3 Chilco, 105 chiloensis, Myotis, 1 1 1 Chilomys, 1 1 7 instans, 124 Chilo stigma alas cense, 46 alascensis, 46 Chimaeropsyllidae, 180 Chiroderma villosum, 1 12 Chironectes minimus, 110, 111, 112, 142, 166 Chironomidae, 322, 323 Chiroptera, 107, 108, 115, 164, 165, 175, 182 chocoensus, Dasyprocta, 1 12 cholinesterases, 75, 183—189 Choloepus hoffmani, 112 Chapman, F.M., 112, 178 Chromatomyia, 327 , 328, 324-337, Chromatomyia asteris, 335 kluanensis, 335 ixeridopsis n. sp., 327, 335, 336, 344, 345 lactuca, 327, 336, 342, 344, 345 senecionella, 327 , 334 syngenesiae, 327 , 328, 334, 335, 337 chryantha, Tabebuia, 104 Chylomys instans, 1 1 1 Chrysomelidae, 12 IX Cicerbita ( =Mulgedium), 331, 338 Cicerbita alpina, 330, 331 bourgaei, 331 prenanthoides, 331 Cichorieae, 327, 333, 334, 337, 341 Cichorium intybus, 337 Cicindelinae, 88 cinereiv enter, Thomasomys, 111, 113, 117, 121, 124, 127, 131, 132, 164, 166 cinereoargenteus, Urocyon, 110, 166 cinereus, Artibeus, 1 12 Citharexylum sp., 102 Clarke, C.H.D., 227, 228 Clay, T., 93, 178 Clayton, J.S. (see Mitchell, J.), 321, 325 Cleoptsylla, 173, 175, 176 monticola, 114, 124, 125, 166 Clifford, E.A. (see Smart, J.), 196, 198, 210 Clifford, H.F., 42, 53 Cnephia dacotensis, 200 Coelopidae, 83 Coleoptera, 3,5,6, 11, 12, 13, 14, 15, 16, 75, 78, 81, 83, 87, 88, 231, 232, 253, 309 Collembola, 77, 81, 86 Colombians, Metachiurus nudicaudatus, 111, 176 Colombians, Proechimys, 1 1 2 Colombians, Scolopsyllus, 115, 119, 141, 159, 160, 166, 172, 174, 181 coloradensis, Rhyacophila, 33 coloradensis group, Rhyacophila, 33 columbiana n. sp., Phytomyza, 327, 332, 333, 343, 345 comis, Tetrapsyllus, 91, 1 15, 132, 136, 166, 172, 174 communis, Lapsana (=Lampsana), 330, 331 Compositae, 327, 332, 334, 339, 341 compositus, Onthophagus, 352 concolor, Felis, 1 1 3 concolor, Oryzomys, 112 Condylarthra, 106, 107 Conepatus semistriatus, 1 10 conges tus var. palustris, Senecio, 334 conistylum, Prosimulium, 207, 209 Conley, D.L. (see Jacobs, C.H.), 97, 178 Cono carpus, 104 Coombs, R.F. (see Greiner, E.C.), 226, 229 Coprina, 235,243,245,247 Coprini, 233, 234, 238, 242, 243, 245, 247 Copris, 245 boucardi, 243 Coptopsyllidae, 180 copha, Adoratopsylla intermedia, 116, 117, 121, 166 copha, Stenopsylla intermedia, 1 1 7 Cora pavonia, 105 coralito, 104 Corbett, J.R., 183, 186 cordoncillo, 104, Coronapsylla, 175 Corvidae, 226 Costa Lima, A. da, 94, 178 Coulson, J.R. (see Stone, A.), 220, 229 Courtney, K.D. (see Ellman, G.L.), 184, 186 Couvert, L., 207, 209 Craig, D.A., 195, 196, 197, 198, 205, 206, 207, 209 Craneopsylla, \16 Craneopsyllinae, 124, 175, 176 Craneopsyllini, 124 Crepidinae, 327 , 330, 336, 337 Crepis atribarba, 334 biennis, 331 capillaris, 331 (Ixeridopsis) elegans, 335, 345 gracilis, 334 jacquini, 331 nana, 335 paludosa, 331 rubra, 331 runcinata, 334 sibirica, 331 tectorum, 334, 337 crepitans, Hura, 102 Crescentia cujete, 102 Cricetidae, 166, 175 Cricetinae, 107 crinoids, 97 Crosby, T.K., 198,209 Crosskey, R.W., 198, 201, 206, 209 Croton sp., 102, 104, 105 crustaceans, 83 cryptoctenes, R. lugubris, 158 Cryptotis medallinius, 1 1 1 thomasi, 1 1 1 Ctenidiosomus, 173, 175 croxtoni, Simulium, 226 croze tensis, Croze tia, 204, 209 X Crozetia crozetensis, 204, 209 Ctenocephalides, 173 felis, 114, 118, 164, 166, 168, 172, 175 Ctenidiosomus perplexus, 124 rex , 115, 121, 172, 174 traubi, 91, 94, 115, 121, 122, 124, 166, 172, 174 Ctenomys, 176 Ctenophtalminae, 117, 180 Cuculidae, 226 cuipo, 104 cujete, Crescentia, 102 Culicidae, 75, 198, 226 curculionidae, 1 1 Curtis, J. 225,256,288 Curtis, L.C., 220, 221, 228 cyanellus, Canthon, 241, 243, 248, 252 cyanellus cyanellus, Canthon, 247 , 248 Cyclopes didactylus, 1 1 2 Cyclorrhapha, 79 dacotensis, Cnephia, 200 daphnis, Phanaeus, 233, 242, 244 Darlington, P.J., Jr., 84, 85, 87 Dasyprocta candelensis, 1 1 1 chocoensis, 1 1 2 fuliginosa, 1 1 1 punctata, 110, 112, 142, 166 Dasyproctidae, 166 Dasypsyllus, 173 Dasypsyllus gallinulae peripinnatuSj 91, 116, 131, 134, 166, 172, 174, 175 Dasypus novemcinctus, 110, 158 novemcinctus fenestratus, 1 5 8 Davies, D.M., 225, 228, Davies, D.M., (see Wood, D.M.), 206, 210 Davies, L., 195, 197, 204, 209, 224, 228 deborahae, n. sp., Aneurus, 225, 263, 266, 269, 274, 275, 289, 290, 291, 292, 298, 301,306 decemlineata, Leptinotarsa, 12, 16 decorum, Simulium, 224 deer, 107, 222 DeFoliart, G.R. (see Anderson, J.R.), 226 228 defoliarti, Simulium, 221 delpontei n. sp Polygenis, 91, 116, 138 146, 147, 148, 166, 172, 174 Dendragapus obscurus fuliginosus, 228 Denning, D.G., 27, 30, 33, 37, 39, 40, 46, 53, 54 derbianus, Caluromys derbianus, 108, 112 Dermaptera, 77, 79, 81 j Dermestidae, 309 Desmanthus virgatus, 102 Desmodontidae, 166 Desmodus rotundus, 112, 164, 166 j Dewhurst, S.A., 185, 186 Diaemus youngi, 112 j dichopticus, sp., near; Gymnopais, 204 Dichotomiina, 235, 238, 241, 243 Dichotomius carolinus, 238, 240, 242 tortulosus, 235 Dicosmoecinae, 34 Dicosmoecus, 34, 47 gilvipes, 25, 34, 47, 50, 51, 61, 66 j jucundus, 35 didactylus, Cyclopes, 1 1 2 Didelphidae, 166 Didelphis azarae, 111, 117, 121, 166 cancrivora, 140 marsupialis, 117, 121, 124, 137, 141, 142, 158, 158, 164, 166, 176 m. marsupialis, 110, 112 diomedes, Sphinctosylla, 91, 94, 1 14, 124, 166, 174 Diplura, 8 1 Diptera (flies), 11,75,77,78,79,83, 182, 195, 209, 210, 228, 229, 325, 327, 328, 339, 340, 341 discolor, Phyllostomus, 1 1 2 dissimilis, Aneurus, 225, 262, 263, 265, 266, 267, j 267, 270, 276, 289, 290, 291, 292, 293, 296, 299, 301, 307 distincta, Sternopsylla, 175 Dobzhansky, T., 86, 87 Dodds, G.S., 34, 54 Dolophilodes aequalis, 69 nora, 69, 71 novusamericanus, 69 Dolophilodes (D.) pallidipes , 69, 70 donaldi, n.sp Rhyacophila, 25, 29, 34, 50, 51, 58, 59, 66 Donvar-Zapolski, D.P., 338 Doratopsyllinae, 180 dorsalis, Vampyrops, 1 1 2 Douglas, J,, 256, 288 Drosophila melanogaster, 1 5 XI dry as, Marmosa, 113, 124 ducks (disease transmission to), 226, 227, 228, 229 dulce, Pithcellobium, 102 Dumbleton, L.J., 195, 197, 198, 204, 209 Dunn, L.H.,94, 178 dunni, Polygenis, 113, 116, 139, 141, 149, 172, 174 dunni, Rhopalopsyllus, 139 Dyplomys (=Diplomys) caniceps, 1 1 2 Dysdercus cingulatus, 13 ebria, Rhyacophila, 29 echidnophagoides, Juxtapulex, 158 Echimyidae, 140, 166 Echinoprocta rufescens, 1 1 1 echioides, Picris, 330, 334 edentates, 91 edentulus, Canthon, 247 Edmonds, G.F., 238, 252 edule, Sechium, 104 Edwards, F.W., 195, 197, 209 Eidt, R.C., 100, 178 Eira barbara, 1 12, 142, 166 elastica, Castella, 102, 104 elder, 282 elegans, Crepis (Ixeridopsis), 335, 345 elk, 226 Ellman, G.L., 184, 186 encenillo, 105 Encephalitis, Eastern Equine, 226, 228 encimo, 105 engelmanni, Picea, 279 Ephemeroptera, 77, 79, 322, 323 equatoris, Ceratophyllus, 131 equatoris, Pleochaetis equatoris, 117, 131, 172, 174 Equisetum, 39, 105 erigerophila, Phytomyza, 337 esmaralarum, Nectonys alfari, 111, 112 Escoto, J.A.V., 86, 88 espadero, 105 Espeletia, 105 Espinal, T., L.S., 94, 100, 102, 178 Essig, E.O.,34, 56, 54 Etnier, D.A., 43, 54 Eumops auripendulus, 1 1 2 Eumops bonariensis, 1 1 0 Euphorbia caracasana, 102 euryadminiculum, Simulium, 226 Eurysternini, 233, 237, 249 Eurysternus, 241, 249, 251, 252 balachowskyi, 249, 250 magnus, 249 mexicanus, 249, 251 Eusimulium, 227 Evans, H.E., 315, 316, 317, 318 Evans, W.G., 75, 83, 84, 88 Ewing, H.E.,93, 178 excelsum, Anacardium, 104 falcata, Kohlsia, 173 Fallen, C.F., 329, 339 Fallis, A.M., 226, 227, 228 fallisensis, Ornithofilaria, 226, 228 familiaris, Canis , 142, 166 farnesiana, Vachelia, 102 fasciatus, Oncopeltus, 6, 12, 13, 14, 15, 16 Featherston, F.M. (see Ellman, G.L.), 184, 186 Felidae 166 Felis catus, 1 64, 1 66 concolor, 1 1 3 pardalis, 1 1 2 tigrina pardionoides, 110, 112 yagouaroundi, 110, 112 felis, Ctenocephalides, 114, 118, 164, 166, 168, 172, 175 felis, Pulex, 1 64 femoralis, Limnephilus, 41 fenestratus, Dasypus novemcinctus, 158 ferns, 104 Fest, C., 183, 186 Festuca, 105 Fewkes, J.W., 309 Ficus (spp.), 102, 104 Fischer, F.C.J., 27, 30, 33, 34, 37, 39, 42, 43, 44, 46, 54 fish, 321, 322, 325 fiskei, Aneurus, 263, 266, 267, 269, 280, 281, 282 282, 289, 290, 291, 292, 293, 296, 299, 301,306 Fittkau, H.S., 107, 179 flavus, Pot os, 1 12 fleas, 91-94, 96-98, 102, 106-108, 110-117, 121, 122, 124, 127, 130, 131, 132, 134, 137 142, 149, 154, 158, 161-166, 168, 170-182 flea, bird, 165, 175 fleas, helmet Xll Flint, O.S., Jr., 34, 54 foetida, Aposeris, 334 Foote, R.H. (see Stone, A.), 220, 229 forests, tropical, evergreen, deciduous, mangrove, 98 Fortner, G., 196, 201, 206, 209 Fourcraea sp. 104 Fox, I. (see Tamsitt, J.R.), 94, 182 fox tail, 1 04 Fragaria sp., 105 frailijones, 105 frenata, Mustela, 110, 112, 166 Fredeen, F.J.H., 204, 209, 221, 222, 224, 228, 229, 321, 322, 323, Fredeen, F.J.H. (see Arnason, A.P.), 220, 228, 324 Frey, R., 331, 339 Freyvogel, T.A., 184, 186 Freyvogel, T.A. (see Briegel, H.), 183, 186 Freziera sericea, 1 05 Frick, K.E., 334, 336, 337, 339 frigidus, Petasites, 334 froeschneri, Aneurus, 263, 265, 266, 269, 280, 289, 290, 291, 292, 293, 296, 299, 301, 307 Frost, S., (see Kershaw, W.E.), 325 Frost, S.W., 336, 337, 339 frutescens, Bocconia, 104 Fugatera pterota, 102 fuliginosis, Caenolestes, 124 Fuller, H.S.,94, 179 fulvescens, Sylvilagus, 110, 111 fulvum, Prosimulium, 224 fungi, 241,255,257,259 Furcraea sp., 102 fusca, Phryganea, 42 fuscata, Marmosa, 1 24 fuscatus, Lasiurus ega, 1 1 0 fuscatus, Thomasomys, 110, 117, 121, 124 132, 140, 166 fuscipes, Ateles, 112, 158, 166 fusorius, Anabolia (Asynarchus), 42 fusorius, Asynarchus, 42 fusorius, Stenophylax, 42 fuscum, Prosimulium, 224 gabbi, Bassaricyon medius, 1 12 Galictis allamandi, 1 42 vittata canaster, 112 gallicus, Aneurus, 228 Gardner, J.C.M., 25 1 , 253 Garth, J.S., 85, 88 gasipals, Guilielma, 104 Gast Galvis, A., 94, 179 geese, 227, 228, 229 Geotrupinae, 232, 233 germanica, Blattella, 77 Ghosh, J.J. (see Sengupta, R.), 183, 187 gigantea, Trichantera, 104 Gillaspy, J.E. (see Evans, H.E.), 315, 316, 318 gilvipes, Dicosmoecus, 25, 34, 47, 50, 51, 61, 66 gilvipes, Stenophylax, 34 Glaphyrocanthon subhyalinus, 243 glauca, Aposeris, 333 glauca, Picea, 279 Gleicheniaceas, 104 Glossosomatidae, 52, 69 Glossophaga soricina, 110, 112, 164, 166 Glydenstolpe, N., 113, 179 Glynne-Williams, J. & J. Hobart., 84, 88 godmani, Bembecinus, 3 1 5 gossypiifolia, Jatropha, 1 02 goudotti, Odocoileus, 113 gracile, Hieracium, 333, 335 gracilis, Crepis, 334 Graham, S.A., 309 grandiflora, Lapsana, 331 grasses, 105 gregaria, Schistocerca, 13, 15 Greiner, E.C., 226, 229 Grenier, P., 197, 199, 109 Griffiths, G.C.D., 327, 328, 331, 332, 334, 335, 336, 339 griscescens, Philander, 110, 111, 112 griseipennis, Stenopsyche, 6, 11, 12, 13, 15 griseum, Simulium, 224 griseus, Brady pus, 1 1 2 Groschke, F., 334, 339 grouse, blue, 227, 228 grouse, ruffed, 227, 228 guamo, 104 guanta, Agouti paca, 110 guarumo, 102, 104, 105 guayacan, 104 Guilielma gasipals, 1 04 Guppy, R., (see Schmid, F.), 34, 37, 39, 56 Gust ania superb a, 104 guyannensis, Proechimys, 1 1 2 7\ xiii gymnurus, Hoplomys , 112, 140, 142, 166 Gymnopais sp. near dichopticus, 204 Gymnopleurus, 247 Gyorkos, H. (see Wood, D.M.), 206, 210 Haemoproteus, 227 Haffer, J., 108, 111, 179 Hagen, H.A., 34, 42, 54 haitiensis, Aneurus, 255, 256, 263, 266, 268, 273, 274, 289, 290, 291, 292, 293 296, 299, 302, 308 Halesus alascensis, 46 Halffter, G., 232, 233, 240, 244, 245, 246, 247, 253 Halffter, G., (see Edmonds, W.D.), 238, 252 Halffter, G., (see Halffter, V.), 238, 253 Halffter, V., 238, 244, 253 Hamelia patens, 104 Hammen, T.V.D., 98, 108, 179 Hanson, R.P., (see Anderson, J.R.), 226, 228 Hanssen, H. (see Mendez, E.), 94, 181 Harlan, T.P., (see Nichols, R.F.), 309, 310 Harper, P.P., (see Roy, D.), 38, 43, 44, 55 Hartig, R., 331, 339 has tat us, Phyllostomus, 1 1 1 Hathaway, C.R., (see Costa Lima, A. da), 94, 178 Hearle, E., 224, 229 Heideman, O, 278, 281, 288 Heliconia, 104 Heliotropium, sp. Ill helleri, Vampyrops, 112 hematozoa, 219, 220, 226, 227, 229 Heming, B.S., 81 Hemiptera, 11, 16, 81, 225, 284, 287, 288 Hendel, F., 329, 321, 332, 337, 339 Hennig, W., 79, 81 herbivore, 241 Herman, C.M., 227, 229 Hering, E.M., 329, 330, 331, 337, 340 Hering, E.M. (see Groschke, F.), 334, 339 Hering, M., 329, 330 331, 337, 339, 340 Hershkovitz, P.,97, 106, 107, 108, 111, 113, 173, 179 Heteromyidae, 166 Heteromys australis, 112, 166 Heteroptera, 6, 12, 13 15, 16, 287, 288 hieracina, Phytomyza, 329, 330 hieracioides, Picris, 330 Hieracium, 327 , 330, 335, 337, 338 albiflorum, 330 amplexicaule, 331 aurantiacum, 331 caespitosum, 331 gracile, 333, 335 japonicum, 334 lachenalii, 331 , 337 laevigatum, 331 laevigatum var. tridentatum, 331 murorum, 331 prenanthoides, 331 pulmonarioides, 331 sabaudum, 330, 331 schistosiphon, 33 1 silvaticum, 331 thapsoides, 331 transylvanicum, 331 triste, 333, 345 umbellatum, 331 villosum, 331 vulgatum, 330, 331 Hieracium sp., 331 hirtipes, Prosimulium, 224, 225 Hisaw, F.L. (see Dodds, G.S.), 34, 54 hispaniolensis n. sp Aneurus, 255, 256, 263, 265, 266, 268, 273, 274, 289, 290, 291, 292, 299, 302, 308 hispidus, Leontodon, 330 hispidius, Sigmodon, 1 1 2 Hodge, F.W., 309 hoffmani, Choloepus, 112 hogplum, 102 Holdrige, L.R., 102, 179 Holland, G.P., 165, 179 Hominidae, 166 Homoptera, 78 Homo sapiens, 164, 165, 166, 175, 220, 223, 224, 225,226, 227,242,247 Hooper, E.T., 179 Hopewell, W.W. (see Arnason, A.P.), 220, 228 324 Hopkins, G.H.E., 93, 165, 175, 179, 180 hopkinsi n. sp Polygenis, 91, 115, 116, 137, 150, 151, 166, 172, 174 Hoplomys gymnurus, 112, 140, 142, 166 Horn, G.H., 84, 88 horse, 107,221,222,226 horse tail (see Equisetum) XIV hrdyi, Aneurus, 257 , 263, 267 , 288 Hulten, E., 328, 340 Hunter, R.L. (see Freyvogel, T.A.), 184, 186 Hum crepitans, 1 02 Hussey, R.F., 272, 288 hyalinata, Rhyacophila, 33 hydrobat es, Ichthysomys, 1 1 1 Hydrochaeris hydrochaeris, 1 1 2 isthmius, 112 hydrochaeris, Hydrochaeris, 112 Hydrophilidae, 83, 88 Hydropsy chidae, 312 Hylepsyche plectrus, 44 hylophilus, Thomasomys, 124 Hymenoptera, 77, 315, 318 Hystrichopsyllidae, 117, 173, 180 Ichthyomys hydrobates, 1 1 1 nicefori, 1 1 1 Imania thomasi, 25, 35, 47, 50, 58, 62, 66 inambari, Akodon pulcherrimus, 141 incisus group, Limnephilus, 39 inconstans, Aneurus, 259, 260, 263, 266, 269, 271, 272, 280, 281, 282, 283, 289, 290, 291, 292, 293, 296, 299, 301, 305 indigo, 104 Indigobera sp. 104 inflatum, Prosimulium, 210 Inga spp. 1 04 Inophleum armatum, 104 Insecta, 14, 15, 16, 80 inseparata, Phytomyza, 329, 330, 332 instans, Chylomys, 111, 124 insularis, Limnephilus, 25, 39, 48, 50, 62, 67 intermedia, Adoratopsylla (T.), 166, 172, 174, 176 intybus, Cichorium, 337 invaria group, Rhyacophila, 33 iraca 104 irritans, Pulex, 93, 114, 164, 166, 169, 172, 174, 175 Isa, J.M. (see Savage, A.), 227, 229 Ischnopsyllidae, 127, 180 Ischnopsyllinae, 127 Isoderminae, 265 Isopoda, 86 isthmica, Marmosa, 112 isthmius, Hydrochaeris, 1 12 isthmius, Microsciurus falviventer, 1 12 Ixeridopsis, 321, 335, 336 ixeridopsis, n. sp., Chromatomyia, 327 , 335, 336. 344. 345 Ixeris alpicola, 336 Jacobs, C.H.,97, 98, 180 jacquini, Crepis, 331 jamaicensis, Artibeus, 1 10 janus, Limnephilus, 40, 63 japonica, Phytomyza, 334 japonicum, Hieracium, 334 Jatropha gossypiifolia, 102 Jeannel, R., 84, 88 johannseni, Simulium, 226 Johnson, F.M. (see Beckman, L.), 184, 186 Johnson, P.T., 94, 113, 140, 165, 175, 180 Johns, P.M., 84, 86, 88 Johnson, R.D. (see Kershaw, W.E.), 325 jucundus, Dicosmoecus, 35 Judulien, F., 247, 253 juliflora, Prosopis, 102 Juxtapulex echidnophagoides, 158 kangaroo, 241 Kaplan, W.D. (see Dewhurst, S.A.), 185, 186 Kapoor, I.P., 323, 325 kappleri, Peropteryx kappleri, 1 1 0 Karl, O., 331, 340 Keast, A., 106, 180 kelp, 83 Kenodactylus audouini, 84, 86 keratus, Lenar chus (Prolenarchus), 43 keratus, Limnephilus, 43 Kershaw, W.E., 323, 325 Kjellgren, B.L, (see Betten, C.), 34, 42, 46, 63 klagesi, Polygenis, 116, 140, 152, 166, 172, 174 klagesi, Pulex, 1 40 kluanensis, Chromatomyia, 335 Knight, K.L., 197, 198,209 Kohlsia 181 Kohlsia falcata 173 tiptoni, 173 kok-sghyz, Taraxacum, 334 Koble, H.J.,42, 54 Kormilev, N., 257, 270, 272, 273, 274, 275, 276, 279, 280, 281, 284, 285, 286, 288 Kristensen, N.P., 81 Kubska, J., 331, 340 Kurtak, D.C., 201, 209, Kurten, B., 107, 180 labialis, Noctilio, 110, 164, 166 ! lackberry, 276 || lachenalii, Hieracium, 331, 337 lactuca, Chromatomyia, 327 , 336, 342, 344, 345 lactuca, Phytomyza, 336 Lactuca canadensis, 337 scarida var. integrata, 337 serriola, 331, 336, 337, 345 spicata, 331 tatarica, 331 virosa, 331 Lactuca sp., 334 Ladenbergia, 105 laevigatum, Hieracium, 331 laevigatum, var. tridentatum, Hieracium, 331 lagopus, Ochroma, 102, 104 Laguncularia racemosa, 104 Laird, M., 227, 229 lampsanae, Phytomyza, 329, 330, 331 laniger, Thomasomys, 121 Laniidae, 226 Lantana canescens, 102 lapponicus, Asynarchus, 15, 42, 48, 50, 51, 64, 67 lapponicus, Limnephilus, 42 Lapsana (=Lampsana) communis, 330, 33 1 Lapsana grandiflora, 33 1 Lasiurus sp., 113 Lasiurus ega fuscatus, 1 1 0 Lateritics, Reddish-brown , 1 00 Latersol 100 laticollis, Cephalodesmicus, 245 latimanus, Rhipidomys, 110, 117, 131, 140, 141, 166 latipes, Simulium, 226, 228 Lee, V.H. (see Anderson, J.R.), 226, 228 Leech, H.B., 83, 88 Leg worms, 226 lemurinus, Aotus, 1 1 1 Lenarchus (Prolenarchus) keratus, 15, 43, 48, 50, 51, 64, 67 Leonard, F.A. (see Leonard, J.W.), 44, 54, 55 Leonard, J.W., 44, 54 55 Leontodon autumnalis, 330 hispidus, 330 Leontodontinae, 330 Lepidoptera, 11, 13, 88 Leptinotarsa decemlineata, 12, 16 leptocerus, Aneurus, 263, 265, 266, 270, 272, 289, 290, 291, 292, 294, 296, 299, 302, 307 Leptopsylla 173 segnis, 115, 131, 133, 166, 172, 174, 175 Leptopsyllidae, 180 Leptopsyllinae, 131 Lethierry, L., 270, 271, 275, 278, 281, 288 leucocytozoan infection, 219, 226, 227, 229 Leucocytozoon anatip, 226, 229 bonasae, 228 simondi, 227 , 228, 229 Lewis, R.E., 165, 180 Liatongus monstrosus, 238, 239, 241, 252, 253 lichens, 105 lilium, Sturnira, 1 12 Limnephilidae, 25, 26, 34, 43, 47, 50, 51, 312, Limnephilinae, 38 Limnephilus alvatus, 25, 27, 39, 40, 48, 50, 51 63, 67 argent eus, 41 femoralis, 41 insularis, 25, 39, 48, 50, 62, 67 janus, 40, 63 keratus, 43 lapponicus, 42 modestus, 42 nimmoi, 25, 27, 38, 48, 50, 52, 62, 66 rhanidophorus, 42 secludens, 39, 40 vernalis, 25, 41 , 48, 50, 5 1 , 63, 67 Lindner, E. (see Rubtzov, I.A.), 209 Lindroth, C.H., 84, 88 Ling, S.W., 37, 55 Linsley, E.G., 318 Linsley, E.G. (see Evans, H.E.), 315, 316, 317, 318 Liponeura cinerascens, 209 Listropsyllinae, 180 Litopterna, 106 lituratus, Artibeus, 1 10 lizards, 1 1 1 Locusta migratoria, 10, 12, 13, 14, 16 Loomis, F.G., 106, 180 Lopez, G. (see Halffter, G.), 232, 233, 243, 244, 253 luggeri, Simulium, 219, 223, 321, 323 lugubris, Rhopalopsyllus, 1 13, 115, 158, 163, 166, 172, 174, 176 XVI lulo, 105 Lundquist, A., 331, 340 Lutra annectens, 110, 112 Lygaeidae, 16 Lyneborg, L. (see Anthon, H.), 197, 209 Lytta viridana, 5, 6, 7, 9, 10, 11, 12, 13, 14, 15, 16 Macchiavello, A., 93, 94, 180, 181 macedero, 104 Machado-Allison, C.E. (see Tipton, V.J.), 113, 124, 139, 158, 182 Macropsyllidae, 180 macrophylla, Cespedesia, 104 Macropocopris, 241 magnus, Eurysternus, 249 Mahowald, A.P., 81 maize, 105 major, Molossus molossus, 110, 112, 164, 165, 166 Malachiidae 88 Malacopsyllidae 180 Malagoniella bicolor, 247 puncticollis tubericeps, 247 mammagua, 104 Mammalia, 75, 91, 92, 93, 97, 98, 102, 106, 107, 108, 110-117, 121, 124, 127, 131, 132,137-142, 158, 164, 165, 172-182, 220 man (see Homo sapiens ) mangrove, red 104 mangrove, black 1 04 mangrove, white 104 Manning, S.A., 331, 340 Mansingh, A. (see Smallman, B.N.), 183, 187 maple tree, 282 marginella, Phytomyza, 327 , 329, 332, 333, 337, 338, 342 marina, Avicennia, 104 marinus, Aepus, 84 Marmosa, 117 dry as, 113, 124 fuscata, 124 isthmica, 112 robinsoni, 112 improvida caucae, 1 10 Marsupiaha, 91, 106, 107, 127, 141, 175, 176, 241, 242, 247 marsupialis, Didelphis, 110, 112, 117, 121, 124, 137, 141, 142, 158, 164, 166, 176 mastodonts, 107 maryae, n. sp., Aneurus, 255, 263, 266, 270, 283, 289, 290, 291, 292, 294, 296, 299, 302, 307 Matchett, R.E. (see Kershaw, W.E.), 325 Matsuda, R. 75 Matsuda, R., (see Usinger, R.L.), 256, 265, 270, 271, 272, 275, 278, 281, 285, 288 Matthews, E.G., 233, 241, 243, 245, 247, 253 Matthews, E.G. (see Halffter, G.), 232, 233, 240, 243,245,246,247,253 Mayr, E., 86, 88 mays, Zea, 105 Mazama americana, 112, 113 americana zetti, 1 1 1 Mazur, J. 331, 340 maximus, Aulacopris, 247 Mecoptera, 77 medallinius, Cryptotis, 1 1 1 megalotus, Pot os flavus, 1 10 megastigmata, Rhynchopsyllus, 91, 94, 165 Megathoposoma candezei, 247, 253 Meijere, J.C.H. de, 329, 330, 331, 340 melanoceras, Acacia, 104 melanogaster, Drosophila, 15 melanurus, Philander, 1 12 Melinis minulti flora, 104 Melilotus, 315 Meloidae, 5, 14, 15, 16 Membrillo, 104 Mendez, E., 75,94, 181 Mendez, E (see Tipton, J.), 113, 121, 122, 130, 134, 139, 142, 149, 154, 158, 161, 162, 163, 165, 168, 170, 171, 173, 182 Menke, A.S., 318 mephistophiles, Pudu, 1 13 meridionale, Simulium, 226 Merkley, D.R., (see Ross, H.H.), 39, 42, 43, 55 mertensiana, Cassiope, 335 mesquite, 102, 276 Metachirus nudicaudates, 1 66 nudicaudates Colombians, 111, 176 Metcalf, R.L. (see Kapoor, I.P.), 325 mexicana, Atta, 238 mexicanus, Eurysternus, 249, 251 mexicanus, Reithrodontomys, 110, 138 micantha, Trema, 104 Michalska, Z., 331, 340 Michna, J., 331, 340 XVII Miconia albicans, 1 05 micranthum, Ocimum, 102 microfilaria, 226, 227 Microsciurus flaviventer isthmius, 1 1 2 migratoria, Locusta, 10, 12, 14, 16 Millar, J.L., 222, 229 milleri, Reithrodontomys mexicanus, 1 1 1 Milne, L.J., 27, 33, 34, 37, 40, 42, 46, 55 milnei, Rhyacophila, 25, 31, 34, 50, 60, 66 Mills, M.L. (see Kershaw, W.E.), 325 Mimosa, 104 minimus, Chironectes, 110, 111, 112, 142 minusculus, C., 40 minuscula, Phytomyza, 332 minutiflora, Melinis, 104 minutus, Aneurus, 255,263, 266, 269, 274, 275, 276, 277, 278, 280, 289, 290, 291, 292, 294, 296, 299, 302, 304, 307 minutus, Oryzomys, 124, 127, 132 mirae, Tylomys, 1 1 2 Miridae, 288 Mitchell, J., 321, 325 mixtum, Prosimulium, 224 modes ta, Anabolia, 42 modesta, Stenophylax, 42 modestus, Anisogamus, 42 modestus, Asynarchus, 42 modestus, Limnephilus, 42 Mokry, J.E., 207, 209 molitor, Tenebrio, 6,1, 11, 16 Moll, A. A., 94, 164, 181 Molossidae, 164, 166 Molossus molossus, 1 64 Molossus molossus major, 164, 165, 166 Mollusca, 97, 98 mombin, Spondias, 102 monkeys, 107, 243 montanus, Aneurus, 263, 266, 270, 285, 289, 290, 291, 292, 294, 296, 299, 302, 308 Montenegro, E. (see Espinal, T., L.S.), 102, 178 monteno, 105 monticola, Cleopsylla, 114, 124, 125, 166 monstrosus, Liatongus, 238, 239, 241, 252, 253 Moore, I., 85, 88 Moores, E. (see Valentine, J.W.), 106, 182 Moose, 226 mori, Bombyx, 13, 15 mosquito, 209, 210, 225 morosus, Carausius (=Dixippus), 6, 10, 12, 13, 15, 16 mortino, 105 mosquero, 104, 105 Moss, H.C. (see Mitchell, J.), 321, 325 mosses, 105 mouse, house, 131, 175,345 Muller, P., 107, 108, 112, 181 mulgedii, Phytomyza, 329, 330, 331 munchiquensis, Oryzomys, 110 muralis, Mycelis, 330, 331 Muridae, 158, 1 66, 173 murids, 158, 175 murorum, Hieracium, 331 Mus mus cuius, 110, 112, 131, 175, Mustela frenata, 110, 112, 166 Mustelidae, 166 muticus, Canthon, 247 Mycelis muralis, 330, 331 My otis sp., 113 Myotis chiloensis, 1 1 1 myxomatosis, 226 Maclnnes, C.D., (see Bennett, G.F.), 227, 228 McCaman, R.E. (see Dewhurst, S.A.), 185, 186 186 McLachlan, R., 34, 42, 55 nana, Crepis, 335 narvae, Rhyacophila, 33, 34 Nasua nasua, 1 1 2 nasua candace, 1 1 0 olivacea, 1 1 1 socialis, 140 nasua, Nasua, 1 1 2 nasutus, Aneurus, 263, 268, 286, 289, 290, 291, 292, 294, 296, 308 Navas, L., 33, 55 Neacomys tenuipes tenuipes, 112 Neave, F., 34, 35, 55 Nectomys alfari esmeraldarum, 111,. 112 neglectus, Bembecinus, 3 1 8 Negre, J., 86, 87 Nematocera, 79, 195, 209 Neocanthidium, 241, 252 neojamaicensis, n. sp., Aneurus, 255, 256, 263, 266, 268, 277, 278, 289, 290, 291, 292, 294, 297, 300, 302, 308 Neophylacinae, 36, 47 XV111 Neophylax, 36, 47 pulchellus, 37 rickeri, 25, 37, 47, 50, 51, 58, 61, 66 Neotropical Carabidae, Indices, 479—493 Neotyphloceras, 173 rosenbergi, 116, 117, 120, 166, 172, 174, 176 Neotyphloceratini, 117 Neuroptera, 77 Newell, R.L., 28, 30, 33, 55 nice fori, Ichthyomys, 1 1 1 Nichols, R.F., 309, 310 nigricans, Achat ocarpus, 102 nigrifrons, Oxymycteris p., 141 nigripennis, Thalassotrechus, 84, 85 nigripennis, Thalassotrechus barbarae, 83, 86 Nimmo, A.P., 26, 27, 28, 30, 31, 32, 33, 34, 38, 40, 41, 42, 45, 46, 47, 48, 49, 50, 51, 52, 55, 57 nimmoi, Limnephilus, 25, 27, 38, 48, 50, 52, 62, 66 Noctilionidae, 166 Noctilio labialis, 110, 164, 166 Noel-Buxton, M.B., 323, 325 Nonapsylla, \16 nora, Dolophilodes, 69, 71 norvegicus, Rattus, 3, 11 0, 111, 1 12, 158, 166 Notungulata, 106 novemcinctus, Dasypus, 110, 158 novusamericans, Dolophilodes, 69 Nowak, Z., (see Michalska, Z.), 331, 340 Nowakowski, J.T., 330, 331, 337, 340 Nowicki, J., 331, 340 nudicaudates, Metachirus, 166 Nunberg, M., 331, 340 Nygren, W.E., 97, 111, 181 Nystrom, R.F. (see Kapoor, I.P.),'325 oak, 275 obliqua, Steniolia, 315 O’Brien, R.D., 183, 187 obscurus, Caenolestes, 113, 121, 124, 127, 131, 132, 166 obscurus fuliginosus, Dendragapus, 228 O chroma lagopus, 102, 104 Ocimum micranthum, 102 Odocoileus goudotti, 1 13 tropicalis, 112 Odocoileus virginianus, 110, 112, 113 Odonata 323 I officinale, Taraxacum, 331, 333, 337 oleraceus, Sonchus, 330, 334, 336 Oligophlebodes, 36 Oligoryzomys, 182 olivacea, Nasua, 1 1 1 O’Leary, S.B. (see Moll, A. A.), 94, 164, 181 Olsson, A.S., 97, 181 onca, Felis, 1 12 Onchocerca (=Wehrdikmansia) cervipedis, 226 Oncopeltus fasciatus, 6, 12, 13, 14, 15, 16 onion, 105 Oniticellini, 233, 235, 237, 238, 251 Oniticellus cinctus, 237 , 251, 252 Onitini, 233, 235, 238 Onitis, 243, 245 Onthophagini, 233, 235, 238, 253 Onthophagus, 238, 243 Onthophagus compositus, 252 parvus, 241 ophyrus, Rhyacophila, 27 opossum, Philander, 110, 111, 112, 117, 121, 166 Orcutt, A.W. (see Betten, C.), 34, 42, 46, 53 orellana, Bixa, 102 ornatus, Tremarctos, 113 Ornithofilaria fallisensis, 226, 228 O’Roke, E.C., 226, 229 Orthoptera, 11, 12, 13, 15, 16,77 79 Orthopteromorpha, 79 oryzae, Sitophilus (=Calendra), 11, 13, 17 Oryzomys albigularis, 91, 110, 117, 124, 127, 131, 129, 140, 141, 166 alfaroi, 117, 132, 166 alfaroi palmirae, 1 10 caliginosus, 91, 111, 112, 117, 121, 132, 137, 138, 140, 141, 142, 158, 164 capito, 1 1 2 concolor, 1 1 2 (Melanomys) caliginosus, 140, 166 minutus, 124, 127, 132 munchiquensis, 110 (Oligoryzomys), 117, 138, 166 physodes, 140 stolzmanni, 141 Osborne, D., 309 Osgood, W.H., 107, 113, 181 oviceps, Simulium, 194, 196, 197, 199, 200, 201, 203, 204, 205, 206, 207, 217, 218 XIX ovipennis, Trechus, 84 Oxymycteris p. nigrifrons, 141 paca, Agouti, 104, 112, 142, 158, 166 pallidipes, Dolophilodes (D.), 69, 70 palmata, Carludovica, 104 palmirae, Oryzomys alfaroi, 110 paludosa, Crepis, 331 paluster, Sonchus, 330 panamensis, Procyon, 112 paniculatum, Talinum, 102 Panicum barbinode, 96 maximum, 96 paramos, 96, 127 Paraphytomyza, 339 Paridae, 226 Parmenter, L., 331, 340 Parapsylline, 176 Parapsyllini, 132 pardalis, Felis, 1 1 2 pardionoides, Felis tigrina, 1 10 parnassum, Simulium, 224, 226 parvus, Onthophagus, 241 Pascual, R., (see Patterson, B.), 98, 106 107, 108, 176, 181 patens, Hamelia, 104 Patino-Camargo, L., 94, 181 patriciae, n. sp., Aneurus, 254, 255, 263, 266, 267, 270, 274, 276, 287, 289, 290, 291, 292, 294, 297, 300, 302, 308 Patrobini, 87 Patterson, B., 93, 98, 181 Patterson, B. & R. Pascual, 98, 106, 107, 108, 176 pavonia, Cora, 105, pecari, Tayassu, 112, 142 peccaries, 107 pejibaye, 104 penetrans, Tunga, 113, 165, 166, 171, 172, 174, 175 pentandra, Ceiba, 102 pentaphylla, Tabebuia, 102 peripinnatus, Ceratophyllus, 131 peripinnatus, Dasypsyllus gallinulae, 116, 131, 134, 166, 172, 174, 175 Periplaneta americana, 1 5 Peromyscini 107 Peromyscus, 173 Peropteryx kappleri kappleri, 1 1 0 perplexus, Ctenidiosomus, 124 Persea americana, 1 04 Petasites, 327 , 339 frigidus, 334 Petersen, B., 86, 88 Peterson, B.V., (see Davies, D.M.), 225, 228 Peterson, B.V., (see Wood, D.M.), 206, 210 Peterson, D.G., 225, 229 Peterson, D.G. (see West, A.S.), 225, 229 Peterson, D.G. (see Wolfe, L.S.), 225, 229 Phanaeina, 235, 242 Phanaeus, 243, 253 daphnis, 233, 242, 244 Phasmatodea, 6, 16, 79 Philander griscescens, 110 opossum, 110, 111, 112, 117, 121, 166 melanurus, 112 Philarctus quaeris, 313 Philocasca alba, 25, 45, 49, 50, 65, 67 thor, 45, 49 Phryganea fusca, 42 Phthiraptera 78 phyllisae, Plocopsylla, 91, 1 14, 127, 128, 166, 172, 174 Phyllostomus discolor, 112 hastatus, 1 1 2 Phyllotis, 176 Phytagromyza, 339 Phytomyza albiceps, 327 , 328, 329, 332, 334, 337, 338 aposeridis, 327 aquilegiae, 332 archhieracii, 337 , 338 atricornis, 339 columbiana n. sp., 327 , 332, 333, 343, 345, erigerophila, 337 hieracina, 329, 330 inseperata, 329, 330, 332 japonica, 334 lactuca, 336 lampsanae, 329, 330, 331 marginella, 327 , 329, 332, 333, 337, 338, 342 minuscula, 332 mulgedii, 329, 330, 331 prenanthidis, 329, 330, 331 robustella, 327 senecionella, 334 sonchi, 329,330,331,332,338 sonchi cicerbitae, 329 XX Phytomyza sonchi hieracina, 329, 338 sonchi lampsanae, 329 sonchi mulgedii, 329, 338 sonchi prenanthidis, 329 sonchina, 329, 330, 332 syngenesiae, 339 taraxaci, 331, 337, 338 Phytomyza sp., 329, 330, 331 Picea engelmanni, 279 glauca, 279 Picris, 330, 338 echioides, 330, 334 hieracioides, 330 pictipes, Simulium, 206 picturatus group, Limnephilus, 39 pigs fern, 104 Pilosa, Portulaca, 1 02 pinchaque, Tapirus, 1 1 3 Pinus ponder osa, 320 Pinus, sp., 320 phaeus, Phyllotis, 141 Phyllostomatidae, 166 Phyllotis. phaeus, 141 physodes, Oryzomys, 140 Piper aduncum, 104 pisonae, Aneurus, 257 , 268, 277 , 308 Pithcellobium dulce, 102 plantanifolia, Cavanillesia, 104 platanillos, 104 Platycentropus plectrus, 25, 44, 49, 50, 51, 65 Platyphylax alascensis, 46 Plecoptera, 322, 323 plectrus, Hylepsyche, 44 plectrus, Platycentropus, 25, 44, 49, 50, 51, 67 Pleochaetis, 94, 173, 180 equatoris equatoris, 117, 131, 172, 174 dolens quit anus, 131 smiti, 117, 131, 132, 134, 166, 172, 174 Pleridium aquilinum, 104 Philopotamidae, 52, 69 Plocopsylla, 173, 175 phyllisae, 91, 114, 127, 128, 166, 172, 174 scotinomi, 175 thor, 114, 127, 128, 166, 172, 174 Podalonia valida, 3 1 8 Pogonini, 84 politus, Aneurus, 263, 266, 267 , 269, 271, 272, 278, 289, 290, 291, 292, 294, 297, 300, 302, 306 Pollitzer, R., 94, 181 Polytricum sp., 105 Polygenis bohlsi bohlsi, 116, 132, 142, 166, 172 174 brachinus, 138 caucensis n. sp., 91 , 116, 137, 142, 144, 145, 166, 172, 174 delponti, 91, 116, 138, 146, 147, 148, 166, 172, 174 dunni, 113, 116, 139, 141, 149, 172, 174 hopkinsi n. sp., 91, 1 15, 1 16, 139, 150, 166, 172, 174 klagesi, 116, 140, 152, 166, 172, 174 klagesi samuelis, 91, 140 pradoi, 91, 116, 140, 153, 166, 172, 174 roberti, 172, 174 roberti beebei, 113, 116, 137, 140, 141, 154 thurmani, 91, 115, 141, 155, 166, 172, 174 trapidoi n. sp.,91, 115, 116, 141, 156, 166, 172, 174 popayanus, Thomasomys, 1 1 1 Portulaca pilosa, 1 02 potatoes, 105 Potos flavus, 1 12 flavus megalotus, 1 10 Potter, D.S. (see Newell, R.L.), 28, 30, 33, 55 potteri, Rhyacophila, 30 poultry, 226, 227 pradoi, Polygenis, 91, 116, 140, 153, 166, 172, 174 pradoi, Rhopalopsyllus, 140 Pratt, J.J., Jr. (see Babers, F.H.), 185, 186 Prenanthes, 331 purpurea, 330 prenanthidis, Phytomyza, 329, 330, 331 prenanthoides, Cicerbita, 331 primates, 107, 166 prinoides, Quercus, 272 prita, Psychoglypha, 15, 46, 49, 65 Procyon cancrivorus panamensis, 1 1 2 Proechimys, 140 colombianus, 112 guyannensis, 1 1 2 semispinosus, 112, 121, 140, 142, 166 Prolenarchus, 43, 48 prolixus, Rhodnius, 12 XXI Prosimulium, 228 conistylum, 207 , 209 fulvum, 224 fuscum, 224 hirtipes, 224, 225 inflation, 210 mixtum, 224 Prosopis juliflora, 1 02 Prosympiestinae, 265 Provancher, L., 281, 288 Prunus, 105 Psychodidae, 198, 209 Psychoglypha alascensis, 25, 46, 47, 49, 65 alaskensis, 46, 49 prita, 25 ? 49, 65 schmidi, 49 subborealis, 46, 49 ulla, 46, 47, 49 Psocoptera, 77 pterota, Fugatera, 102 Ptychopteridae, 198, 209 pubescens, Ammophila, 318 pucherani, Sciurus, 1 1 1 Pudu mephistophiles, 113 pulchella, Arctopora, 312 pulchellus, Neophylax, 37 Pulex, 1 73, 1 75 bohlsi, 132 cheopis, 158 felis, 164 klagesi, 140 irritans, 93, 164, 166, 169, 172, 174, 175 segnis, 131 simulans, 91, 114, 164, 166, 172, 174, 175 pulex, Rhynchopsyllus, 91, 94, 1 13, 164, 165, 170, 172, 174, 175 Pulicidae, 158 Pulicinae, 158 Pulicini, 164 Pulicoidea, 158 pulmonarioides, Hieracium, 331 punctata, Dasyprocta, 110, 112, 142, 166 puncticollis tubericeps, Malagoniella, 247 Puri, I.M., 198, 209 purpurea, Prenanthes, 330, 331 pusillus, Aneurus, 263, 266, 268, 277 , 284, 292, 300, 303, 307 Putnam, J.D., 34, 55 Pygiopsyllinae, 121, 180 Pygiopsyllidae, 121, 175 pygmaeus, Aneurus, 255, 263, 266, 269, 275, 276, 304, 307 quaeris, Philarctus, 3 1 3 quadridens, Cephalodesmius, 245 Quercus prinoides, 272 virginiana, 215, 276 Quassia sp., 105 quitanus, Ploechaetis dolens, 1 3 1 quitoense, Solanum, 105 rabbits, 107, 226, 247 racemosa, Langulallaria, 104 Rageau, J. (see Grenier, P.), 197, 199, 209 rainbow trout, 322 Rapnea sp. 105 rat, spiny, 140 Rattus, 3, 158, 175, 176 novegicus, 3, 110, 111, 112, 158, 166 rattus, Rattus, 3, 110, 111, 112, 131, 158, 166 Raven, P.H., 106, 107, 181 Ray, C., 86, 88 redbud tree, 176 Reed, E.B., 322, 323, 325 Reichardia, 330 Reithrodontomys mexicanus, 110, 138 mexicanus milleri, 1 1 1 Rempel, J.G., 221, 222, 229 Rempel, J.G., (see Fredeen, F.J.H.), 221, 228, 325 Rempel, J.G., (see Millar, J.L.), 222, 229 Rempel, J.G., (see Arnason, A.P.), 324 Renjifo-Salcido, S., 165, 181 re x, Ctenidiosomus, 115, 121, 172, 174 rhanidophorus, Asynarchus, 42 rhanidophorus, Limnophilus, 42 Rheomys, 1 17 Rhipidomys latimanus, 110, 117, 131, 140, 141, 166 similis, 117, 124, 166 Rhizophora brevistyla, 104 Rhodnius prolixus, 12 Rhopalopsyllidae, 132, 176, 180 Rhopsalopsyllinae, 132 Rhopalopsyllini, 132 Rhopalopsylloidea, 132 Rhopalopsyllus, 115, 164, 173, 176 australis tupinus, 115, 142, 161, 166, 172, 174, 176 XXII Rhopalopsyllus beebei, 140 cacicus saevus, 1 13, 115, 158, 162, 166, 172. 174. 176 dunni, 139 lugubris, 1 13, 1 15, 158, 162, 166, 172, 174. 176 lugubris cry ptocteres, 158 pradoi, 140 Rhyacophila acropedes, 28 autumnalis, n. sp., 25, 30, 34, 50, 51, 58, 60, 66 bifila, 33 coloradensis, 33 donaldi, n. sp., 25, 29, 34, 50, 51, 58, 59, 66 ebria, 29 hyalinata, 33 milnei, 25, 31, 34, 50, 60, 66 narvae, 33, 34 ophrys, 27 potteri, 30 rickeri, 30 vepulsa, 33 simplex, 25, 27, 34, 50, 58, 59, 66 unimaculata, 25, 30, 34, 50, 51, 60, 66 vobara, 33 vocala, 33 vao, 25, 27, 28, 34, 50, 51, 59, 66 Rhyacophilidae, 25, 26, 27, 50, 51, 31 1 Rhynchopsyllus megastigmata, 91, 94, 165 pulex, 91, 94, 113, 164, 165, 170, 172 174, 175 Rhypidae, 198, 209 rickeri group, Neophylax, 37 rickeri, Neophylax, 25, 37, 47, 50, 51,58, 61 , 66 rickeri, Rhyacophila, 30 robinii, Aepopsis, 84 roberti, Polygenis, 172, 174 Robineau-Desvoidy, J.-B., 329, 340 robinsoni, Marmosa, 1 1 2 roble, 102 robustella, Phytomyza, 327 Rodentia, 166 rodents, 92, 93, 102, 108, 124, 127, 131, 132, 141, 173, 175, 176, 241 rodents, caviomorph, 107, 175, 176 rodents, cricetine, 92, 106, 124, 127, 132, 141, 175, 181 rodents, myomorph, 107, 176 rodents, oryzomine, 132 rodents, sciuromorph, 107 roseae, n. sp., Aneurus, 255, 263, 265, 266, 269, 279, 289, 290, 291, 292, 294, 297, 300, 302, 306 rosenbergi, Neotyphloceras, 116, 117, 120, 172, 174, 176 rosenbergi, Typhlocerus, 117 Ross, H.H., 26, 27, 30, 33, 34, 37, 39, 40, 42, 43,44, 55 Rothmaler, W., 328, 241 Rothschild, M. (see Hopkins, G.H.E.), 165, 175, 180 rotunda group, Rhyacophila, 28 rotundus, Desmodus, 112, 164, 166 Roy, D. 38,43,44,55 Royer, L.M. (see Fredeen, F.J.H.), 325 rubber, 104 rubra, Crepis, 331 rubrum, Acer, 282 Rubtzov, I.A., 197, 198, 209 Rubus sp., 105 rufescens, Echinoprocta, 1 1 1 ruficollis, Calathus, 86 rugglesi, Simulium, 226 runcinata, Crepis, 334 Ryden, N.S., 329, 330, 331, 332, 341 sabaudum, Hieracium, 330, 331 Sabrosky, C.W. (see Stone, A.), 220, 229 saevus, Rhopalopsyllus cacicus, 158, 162, 166, 172, 174 Saha, J.G., (see Fredeen, F.J.H.), 325 samuelis, Polygenis klagesi, 140 Sanchez, E.H., 94, 181 Sangha, G.K. (see Kapoor, I.P.), 325 sanguinosus, Aradus, 281 sapiens, Homo, 164, 165, 166, 175 Sarkar, D. (see Sengupta, R.), 183, 187 Sasakawa, M., 334, 341 Sauer, C.O., 94, 181 Savage, A., 227, 229 Savage, J.M., 106, 181 Saxifragaceae, 339 Say, T., 271, 278, 288 Scarabaeidae, 231, 252, 253 Scarabaeinae, 231, 232, 233, 241, 242, 245, 249,251,253 Scarabaeini, 233, 235, 237, 243, 245, 247, 248, 249, 253 XX111 scariola var. integrata, Lactuca, 233 Schistocerca gregaria, 13, 15 schisto siphon, Hieracium, 33 1 Schmid, F., 26, 27, 28, 30, 33, 34, 36, 37 39,42,43,44, 55, 56 Schmid, F., (see Denning, D.G.), 30, 54 schmidi, Psychoglypha, 49 Schmidt, K.J., (see Fest, C.), 183, 186 Sciuridae, 166, 175 Sciurus granatensis valdiviae, 108, 166 pucherani caucenis, 1 1 1 Scolopsyllus, 173, 176 colombianus, 115, 119, 141, 159, 160, 166, 172, 174, 181 scotinomi, Plocopsylla, 175 Scott, J. (see Douglas, J.), 256, 288 Scott, W.B., 106, 107, 181 scrofa, Sus, 165, 175 Scudder, G.G.E., 77, 78 Sechium edule, 104 secludens, Limnephilus, 39, 40 segnis, Leptopsyllus, 91, 115, 131, 133, 166, 172, 174, 175 segnis, Pulex, 1 3 1 Sehgal, V.K., 334, 336, 337, 341 Seidel, J., 331, 341 semispinosus, Proechimys, 121, 140, 142, 166 semistriatus, Conepatus, 110 Senecio, 105, 327, 334, 339 atropurpureus tomentosus, 334 congestus var. palustris, 334 Senecioneae, 334, 339 senecionella, Chromatomyia, 327 , 334 senecionella, Phytomyza, 334 Sengupta, R., 75, 183, 187 sensitive-plant, 104 septentrionalis, Aneurus, 255, 278 sericea, Freziera, 105 Serra-Tosio, B., 207, 210 serriola, Lactuca, 331, 336, 337, 345 Severin, G. (see Lethierry, L.), 270, 271, 275,278,281,288 shale, 97 sheep, 222, 241 Sherman, F., 281, 288 Shewell, G.E., 226, 229 sibirica, Crepis, 331 sibirica group, Rhyacophila, 30 Sigmodon sp., 117, 132, 176 hispidus, 1 1 2 hispidus bogotensis, 110, 112 Sigmodontini, 107 silkworms, 16 silvaticum, Hieracium, 331 similis, Rhipidomys, 117, 124, 166 Simon, Jean-Pierre, 183, 187 simile, Simulium, 228 simondi, Leucocytozoon, 227 , 228, 229 simplex, Aneurus, 255, 263, 266, 268, 279, 289, 290, 291, 292, 294, 297, 300, 303, 305 simplex, Rhyacophila, 23, 27, 34, 50, 58, 59, 66 Simpson, G.G., 106, 107, 166, 181 simulans, Pulex, 91, 1 14, 164, 166, 172, 174, 175 Simuliidae, 195, 197, 198, 204, 206, 207, 209, 210, 219, 226, 228, 229, 322, 335 Simulium, 325 arcticum , 221, 222, 223, 224, 228, 229, 321, 322,323,324,325 aureum, 228 cheesmanae, 195, 196, 207 croxtoni, 226 decorum, 224 defoliarti, 221 euryadminiculum, 226 griseum, 224 johannseni, 22 6 latipes, 226, 228 luggeri, 219, 223, 321, 323, meridionale, 226 oviceps, 195, 196, 197, 199, 200, 201, 203, 204, 205, 206, 207, 217, 218, parnassum, 224, 226 pictipes, 206 rugglesi, 226 simile, 228 tahitiense, 195, 196, 197, 199, 200, 201, 202, 203, 204, 205, 206, 207, 21 1, 212, 214, 215,216,217 tuberosum, 224 venustum, 207, 219, 224, 225, 226 vittatum, 196, 200, 202, 206, 207, 224, 225, Siphonaptera, 77, 77, 81, 91, 94, 122, 130, 134 139, 140, 149, 154, 161, 162, 163, 165, 168, 170, 171, 173, 177, 180, 181 Sitophilus (=Calendra) oryzae, 11,13 Sittidae, 226 Skala, H., 331,341 XXIV slateri n. sp., Aneurus, 255, 263, 266, 268, 285, 289, 290, 291, 292, 295, 297, 300, 303, 306 Smallman, B.N., 183, 187 Smart, J., 196, 198, 210 Smith, E.L., 78 Smith, E.M. (see Freyvogel, T.A.), 184, 186 Smith, J., 184, 187 Smith, S.D., 27, 33, 56 Smithies, O., 184, 187 smiti, Pleochaetis, 117, 131, 132, 134, 166, 172, 174 socialis, Nasua, 140 Solanum quitoense, 105 tuberosum, 105 Solidago, 330, 332 sonchi, Phytomyza, 329, 330, 331, 332, 338 sonchi cicerbitae, Phytomyza, 329 sonchi hieracina, Phytomyza, 329, 338 sonchi lampsanae, Phytomyza, 329 sonchi mulgedii, Phytomyza, 329, 338 sonchi prenanthidis, Phytomyza, 329 sonchina, Phytomyza, 329, 330, 332 Sonchus arvensis, 330, 336 arvensis uliginosus, 336, 337 asper, 330, 334, 336 oleraceus, 330, 334, 336 paluster, 330 Sdnderup, H.P.S., 331, 341 Sortosa, 69, 70 soricina, Glossophaga, 110, 112, 164, 166 speciosa, Sternopsylla distincta, 115, 127, 130, 166, 172, 174 Spencer, G.J., (see Ross, H.H.), 27, 30, 33, 34, 37, 39, 55 Spencer, K.A., 329, 330, 331, 332, 336, 337, 341 Spencer, K.A. (see Hering, E.M.), 331, 339 Sphecidae Sphinctopsylla, 17 3, 175, 176 diomedes, 91,94, 114, 124, 126, 166, 174 tolmera, 114, 127, 166, 172, 174 spicata, Lactuca, 331 Spondias mombin, 102 sponges, 97 spurge, 102 squirrels, 107, 173 squirrel, red, 108 Stainer, J., 56 Stal, C., 271,278, 281,288 stalzmanni, Oryzomys, 141 Staphylinidae, 83, 88 Starke, H., 331,341 Stary, B., 331, 341 Stebbins, G.L., 330, 337, 341 Steiner, A.L., 315,318 Steniolia, 316, 318 obliqua, 315 Stenophylax fusorius, 42 gilvipes, 34 modesta, 42 Stenopsyche griseipennis, 6, 11, 12, 13, 15 Stenopsychidae, 15 Stephanocircidae, 124, 175, 180 Stephano circus, 175 Sternopsylla, 173 Sternopsylla distincta, 175 intermedia copha, 117 speciosa, 1 15, 127, 130, 166, 172, 174 Stirton, R.A., 98, 106, 182 Stone, A., 198,210, 220,229 strawberries, 105 Strepsiptera, 77 Sturnia lilium, 1 1 2 Stys, P., 257, 259, 262, 265, 271, 288 subborealis, Psychoglypha, 46, 49 subcentralis group, Limnephilus, 38 subhyalinus, Glaphyrocanthon, 243 subrufa, Carollia, 112 superba, Gustavia, 104 surrumbo, 104 Sus scro fa, 165, 175, 222 Sylvilagus andinus, 112 brasiliensis, 110, 111, 112 fulvescens, 110, 111 syngenesiae, Chromatomyia, 327, 328, 334, 335, 337 syngenesiae, Phytomyza, 339 Tabebuia chrysantha, 104 pentaphylla, 102 Tadarida sp., 127, 166 brasiliensis, 110, 127 tahitiense, Simulium, 195, 196, 197, 199, 200, 201,202, 203, 204, 205, 206, 207, 211, 212, 213,214,215,216,217 tajacu, Tayassu, 112, 142 Talinum paniculatum, 102 XXV Tamandua tetradactyla, 1 12 Tamsitt, J.R., 94, 182 tapirs, 107 Tapirus bairdii, 1 1 2 pinchaque, 1 1 3 taraxaci, Phytomyza, 331, 327, 338 Taraxacum, 331, 333, 337, 338 kok-sghyz, 334 officinale, 331, 333, 337 Tarshis, I.B. (see Herman, C.M.), 227, 229 tatarica, Lactuca, 331 Tate, G.H.H., 113, 182 Tayassu pecari, 112, 142 tajacu, 112, 142 tectorum, Crepis, 334, 337 Tenebrio molitor, 6, 7, 11, 16 Tenebrionidae, 6 tenuicornis, Aneurus, 272 tenuipes, Neacomys tenuipes, 1 1 2 tenuis, Aneurus, 263, 266, 270, 272, 273 289, 290, 291, 292, 295, 297, 300, 308 testaceus, Thalassobius, 84 Tetraonidae, 226 Tetrapsyllus, 173, 176 com/s, 91, 115, 132, 136, 166, 172, 174 tetradactyla, Tamandua, 1 1 2 Tettigoniidae, 77 Thalassotrechus barbarae, 75, 83, 84, 85, 86, 87,88,89,90 barbarae barbarae, 83, 86 barbarae nigripennis, 84, 85, 89 testaceus, 84 Thallomyscus, 182 thapsoides, Hieracium, 331 Thierry, A. (see Freyvogel, T.A.), 184, 186 thomasi, Crypto tis, 1 1 1 thomasi, Imania, 25, 35, 47, 50, 58, 62, 66 Thomasomys sp., 121, 127, 166, 173, 176 aureus, 110, 111, 117, 121, 166 cinereiv enter, 111, 113, 117, 121, 124, 127, 131, 132, 164, 166 fuscatus, 111, 117, 121, 124, 132, 140, 166 hylophilus, 124 laniger, 121, 124 po pay anus, 1 1 1 vestibus, 124 thor, Philocasca, 45, 49 thor, Plocopsylla, 114, 127, 128, 166, 172, 174 Thorpe, W.H.,317,318 thous, Cerdocyon, 112, 164, 166 Thraupidae, 226 thurmani, Polygenis, 91, 115, 141, 155, 166, 172. 174 Thysanoptera, 8 1 Thysanura, 77, 78 Tiamastus, \16 Tiarapsylla, 176 tillyardianum, Austro simulium ( Austro simulium), 209 Tipton, V.J. (see Wenzel, R.L.), 107, 165, 175, 176, 182 Tipton, V.J. and G.E. Machado-Allison, 1 13, 132, 139, 158, 182 Tipton, J&E. Mendez, 113, 121, 122, 130, 134, 139, 142, 149, 154, 158, 161, 162, 163, 165, 168, 170, 171, 173, 182 tiptoni, Kohlsia, 173 tolmera, Sphinctopsylla, 114, 127, 166, 172, 174 Torre-Bueno, J.R. de la, 281, 288 tortulosus, Dichotomius, 235 transylvanicum, Hieracium, 331 trapidoi, n. sp Polygenis, 91, 115, 116, 141, 156, 157, 166, 172, 174 traubi, Ctenidiosomus, 91, 94, 115, 121, 122, 124. 174 Trechini, 84 Trechus ovipennis, 84 Trema micantha, 104 Tremarctos ornatus, 113 Trichant era gigantea, 1 04 Trichoceridae, 198,209 Trichoptera, 6, 15, 25, 26, 27, 30, 60, 311, 312, 313,322,323, trilobites, 97 tripunctata group, Imania, 35 triste, Hieracium, 333, 345 Triticum, 105 trivirgatus, Aotus, 1 1 1 trompet, 104 tropicalis, Odocoileus, 1 1 2 Trypanosoma, 226, 227, 228 Tschirnhaus, M. von, 328, 341 tuberosum, Simulium, 224 tuberosum, Solanum, 105 Tunga, 173, 175 Tunga penetrans, 114, 165, 166, 171, 172, 174, 175 XXVI Tunginae, 164 tupinus, Rhopalopsyllus australis, 115, 142, 161, 166, 172, 174, 176 Turdidae, 226 turkey, 227, 229 Turnera almifolia, 102 turtles, 181 Tussilago, 339 Twinnia biclavata, 204 Tylomys mirae, 112 typhus, murine, 93 Typhloceras rosenbergi, 1 1 7 Tyrannidae, 226 Udvardy, M.D.F., 86, 88 Uhler, P.R., 225, 271, 278, 281, 288 ulla, Psychoglypha, 46, 47, 49 Ulmer, G., 34, 42, 46, 56 umbellatum, Hieracium, 331 Umbelliferae, 332 umbellus, Bonasa, 228 ungulates, 91, 106, 164 unimaculata, Rhyacophila, 25, 30, 34, 50, 51, 60, 66 Urocyon cinereoargenteus, 110, 166 Uroderma b. bilobatum, 1 12 Usinger, R.L., 256, 265, 270, 271, 272, 275,278,281,285,288 usingeri n. sp., Aneurus, 255, 263, 266, 269, 284, 285, 286, 289, 290, 291, 292, 295, 297, 300, 303, 308 Vaccinium sp., 105 Vachelia farnesiana, 102 Vadlamudi, S. (see Anderson, J.R.), 226, 228 vagrita group, Rhyacophila, 31 valdiviae, Sciurus granatensis, 108, 166 Valentine, J.W. and E. Moores, 106 Valeriana, 339 valida, Podalonia, 318 Vampyrops dorsalis, 112 helleri, 1 1 2 Van Duzee, E.P., 272, 275, 278, 281, .288 Van Dyke, E.C., 83, 84, 88 Vanzolini, P.E., 108, 182 vao, Rhyacophila, 25, 27, 28, 34, 50, 51, 59, 60 vaueri, Aneurus, 256, 263, 266, 268, 273, 274, 275, 289, 290, 291, 292, 295, 298, 301,303,308 venestus, Rhipidomys, 124 venestum, Simulium, 207, 219, 224, 225, 226 vepulsa, Rhyacophila, 33 veracruzensis n. sp., Aneurus, 225, 263, 265, 266, 269, 275, 285, 286, 289, 290, 292, 295, 298, 307 Vermipsyllidae, 180 vernalis, Limnephilus, 25, 41, 48, 50, 51, 63, 67 verrula group, Rhyacophila, 30 vestibus, Thomasomys, 124 villosum, Chiroderma, 112 villosum, Hieracium, 331 Vieronidae, 226 virens, Canthon, 247 virgatus, Desmanthus, 102 viridana, Lytta, 5, 6, 7, 9, 10, 11, 12, 13, 14, 15 16 virginiana, Quercus, 275, 276 virginianus, Odocoileus, 110, 112 virosa, Lactuca, 331 vittatum, Simulium, 196, 200, 202, 206, 207, 224, 225 vobara, Rhyacophila, 33 vocala, Rhyacophila, 33 vofixa group, Rhyacophila, 27, 33 Voigt, G., 331, 341 Vuilleumier, B.S., 98, 182 vulgatum, Hieracium, 330, 331 wasps, 315, 316, 317 Waite, E.R.,247,253 Walker, F., 255, 278, 288 Wallengren, H.D.J., 42, 56 Waters, T.F., 323, 325 Weeks, L.G., 98, 182 Wenzel, R.L., 107, 165, 175, 176, 182 West, A.S.,-225, 229 Weinmannia sp., 105 Weinmannia balbisiana, 105 West, R.C., 182 wheat, 105 White, E.M. (see Greiner, E.C.), 226, 229 Wickware, A.B., 226, 229 Wigania caracasana, 1 04 Wiggins, G.B.,311 Wille, A., 247, 253 Williams, T.R., (see Kershaw, W.E.), 325 Wirth, W.W. (see Stone, A.), 220, 229 Wobeser, G., 226, 229 XXV11 Wold, J.L. (see Anderson, N.H.), 28, 37, 53 Wolfe, L.S. (see Peterson, B.V.), 225, 229 Wood, D.M., 206, 210 Wood, D.M. (see Davies, D.M.), 225, 228 worms, leg, 226 wrack, 83 wygodzinskyin. sp., Aneurus, 255, 263, 265 266, 270, 271, 272, 289, 290, 291, 292, 295, 298, 301, 303, 308 Xenopsylla, 173 cheopis, 93, 114, 158, 164, 166, 167, 172, 174, 175 Xenopsyllini, 158 yagouaroundi, Felis, 1 1 0 yaragua, 104 Yepes, J. (see Cabrera, A.), 108, 113 Yin, R-S, L., 197, 198, 210 youngi, Diaemus, 1 1 2 Zavrel, H., 331, 341 Zavrel, H. (see Skala, H.), 331, 341 Zea mays, 105 Zetterstedt, J.W., 42, 56 zetti, Mazama americana, 1 10 Xiphiopsyllidae, 180 Zoraptera, 79 Zygodontomys brevicauda brunneus, 110, 112 ffl-A i -f£/ 6!/3 ; 8vcr< Quaestiones Entomologicae A periodical record of entomological investigations, published at the Department of Entomology, University of Alberta, Edmonton, Canada. VOLUME 13 NUMBER 1 JANUARY 1977 QUAESTIONES ENTOMOLOGICAE ISSN 0033-5037 A periodical record of entomological investigation published at the Department of Entomology, University of Alberta, Edmonton, Alberta. Volume 13 Number 1 January 1977 CONTENTS Editorial — Comments About a Few Words in the Biological Sciences 1 Rempel, Heming and Church — The Embryology of Lytta Viridana LeConte (Coleoptera: Meloidae). IX. The Central Nervous System, Stomatogastric Nervous System, and Endocrine System 5 Nimmo — The Adult Trichoptera (Insecta) of Alberta and Eastern British Columbia, And Their Post-Glacial Origins. I. The Families Rhyacophilidae and Limnephilidae. Supplement 1 25 Nimmo — The Adult Trichoptera (Insecta) of Alberta and Eastern British Columbia, And Their Post-Glacial Origins. II. The Families Glossosomatidae and Philopotamidae. Supplement 1 69 Belicek — Corrigenda on Coccinellidae of Western Canada and Alaska with Analyses of the Transmontane Zoogeographic Relationships Between the Fauna of British Columbia and Alberta (Insecta: Coleoptera: Coccinellidae) 73 Editorial — Comments About a Few Words in the Biological Sciences Editors, perhaps to a greater extent than most individuals who are interested in literature, find word usage a subject for serious consideration, from which they derive both pleasure and pain. An editor may develop concepts of the meanings of certain Words, or about how certain words should be used, with his notions differing from those that seem generally accepted. These concepts are expressed in modifications to those manuscripts that an editor is invited to consi- der for publication in his journal. Of course, each editor is convinced that his concepts are correct and at least worthy of at- tention, if not emulation, by others. This is the thought that has impelled me to list in alpha- betical order and discuss some of the words that seem to me to be commonly mis-used in bio- logical literature generally, and especially in the literature of systematic biology. Because I am a systematist, I have also listed some commonly used words that I think are inappropriate in writing about systematics. ALWAYS (antonym - never). — I think it’s best to avoid this word and its antonym in wri- ting about fields that rely mainly on observation for information about nature, and hence are mainly inductive. Further, does the statement “prolegs are always absent” mean anything dif- ferent from “prolegs are absent”? Unqualified, “absent” means just that. I suppose “always” gives emphasis to the adjective, but this seems unnecessary. ANATOMY. — This word refers, literally, to dissection. It is not a synonym of morpholo- gy (study of structure and form) except by virtue of careless usage. Neither should it be used as botanists and some entomologists do: that is, to refer to internal structures and organs. I suppose this unfortunate usage developed because one must dissect (anatomize) to examine internal organs. The noted arachnologist, T.H. Savory wrote (Browsing Among Words of Sci- ence, 195 1, page 67): “we read of . . . external anatomy, which if it means anything can only mean shaving; and of comparative anatomy, which ought to be a contrast between the use of scissors, scalpels and saws . . . .” Why not use the simple terms “external structure” and “in- ternal structure” when referring to these aspects of morphology? See also MORPEIOLOGY, below. 2 BIOLOGY. — This is a term that refers to a field of human endeavor. It means “study of life”. It seems to me utterly incorrect to use “biology” for the non-morphological, non-class- ificatory portions of the general field. By implication then, morphology and systematics are excluded from biology, and in turn, morphologists and systematists are not biologists! An organism or group of organisms cannot have a biology. An organism has a life, and study j of its life (in the broadest sense) is the domain of biology. j Rather than entitling that section of a systematic treatment “Notes on Biology” that deals j with some special aspects of the living members of a particular taxon, it’s best to be more spe- ! cific, and more accurate. If the notes deal with host associations, then use these words for the title; if they deal with habitat and life history, then use these words for the title. j CASE (synonym - situation), as used in the expression “in this case”. — I doubt that this phrase need be used outside the literature of the legal profession or the brewing or cartage in- dustries. More specific expressions are available for biological literature. CLASSIFICATION. - See TAXONOMY. DIVERGENCE. — This term should be used to express relative amount of difference among taxa. One group, whose members are removed by appreciable structural differences from mem- bers of other taxa, is said to exhibit marked divergence, or to be highly divergent. A higher taxon whose member taxa exhibit considerable difference from one another is said to be high- ly divergent in comparison with another taxon whose member taxa show slight differences. See also DIVERSITY. DIVERSITY. — This term is best restricted to reference to numbers of members of groups (either individuals or taxa, or both). A higher-ranking taxon including many lower-ranking taxa is highly diverse, but some exhibit slight divergence. On the other hand, some highly divergent taxa exhibit low diversity. In any event, it’s best to refer to inclusiveness of a taxon in terms of amount of diversity or words implying this (that is, “this genus includes few (or many) species”), rather than as “large” or “small”. These latter terms deal with size, too, but the con- text is different. Also, as these words are used, a “large genus” can be either one including many species, or one that includes species whose members are of large size. DIVISION. — This is not an appropriate word to describe the actions of systematists with reference to taxa. Activities include grouping, re-grouping, and arranging. Re-grouping of spe- cies that results in more genera than were previously recognized seems to be described appro- priately by the word division, but it is not the same thing as, say, cutting a pie, which has a high degree of physical continuity, or cutting a piece of string. Of course, dissection is a form 1 of division, and when a biologist makes a dissection he is indeed dividing. LARGE (as applied to taxa). - See DIVERSITY. MAY. — As used with reference to occurrence or non-occurrence of characters among mem- bers of a taxon, this word is inappropriate. In fact, in each instance, a given character state is or is not exhibited. So, rather than writing “structure X may be present”, it’s better to write “structure X is present in a few individuals; it is absent from most”. MORPHOLOGY. — Means study of form and structure. An individual (not a taxon) has a particular structure, or a particular combination of structures. It does not have a morphology. As with the term “anatomy”, what can “external morphology” possibly mean: “the external study of form and structure”? When referring to parts of a particular organism, the word struc- ture is adequate, and does not need the embellishment of the word morphology. NEVER. - See ALWAYS. OCCASIONALLY (related words - rarely, sometimes, usually). - This word is not appropr- iate in description of distribution of static characters states among taxa. What is implied by most authors is that a particular character is represented by more than one state, and expression of the less frequent one is described as occasional. It is best to indicate the matter more precisely. 3 without reference to time, as follows: “members of most species with red antennae, those of a few species with black antennae”; in preference to “antennae usually red, occasionally (or sometimes, or rarely) black”. On the other hand, if antennal color of an individual changes periodically and is usually red, then it would be occasionally (or sometimes, or rarely) black. RARELY. - See OCCASIONALLY. SMALL (with reference to taxa). — See DIVERSITY. SOMETIMES. - See OCCASIONALLY. SPECIES (and other taxa). — These have members, which are represented by eggs, immatur- res, or adult males and females. These members have characters — structural, behavioural eco- logical, genetical, physiological, and so on. Species and other taxa do not have characters. Thus it is incorrect to write that “A few species deposit their eggs in the nests of other bees”, or that “the aedeagus of this species has a knob on the distal end”. Rather, one should write that “Lemales of a few species deposit their eggs . . . ” et cetera', or that “aedeagi of males of this species” . . . et cetera. SYSTEMATICS. - See TAXONOMY. TAXONOMY. — This is the general term for study of the principles of classification, appli- cation of the principles to formation and ranking of formal groups (taxa), and naming the taxa. Systematics is the field of organic diversity, and taxonomy is a portion of this field. Classification is used in two senses: 1. for the process of establishing taxa; 2. for the resul- ting formal system of arrangement. Thus, one classifies as one constructs a system. The sys- tem is a classification. I doubt that it is correct to write, as the title of a paper: “The Taxonomy of the Genus X-us ”. Genus X-us does not have a taxonomy. Rather, its species are arranged, or classified in a particular way, according to taxonomic principles. The word taxonomy also seems to me to be mis-used with reference to study of an existing classification for the purpose of learning to identify specimens. This type of activity is not tax- onomy. TECHNOLOGY. — This word must mean “study of techniques”. It is used by many writers and speakers in place of technique, so that even the most mundane and inconsequential of pro- cedures is given some aura when it is referred to as a technology! THE. — Categorical expressions (as implied by use of “the”) are to be avoided with referen- ce to selected individuals. For instance, vertebrate physiologists commonly use the expression “the rat” in published work, when what is meant is groups of rats of the species Rattus norve- gicus on which some experiments have been conducted. “Rats”, if a more modest expression than the categorical “the rat”, is also a more accurate expression. UNIQUE. — Because members of each taxon have some unique features, it is inappropriate to state that a given taxon is unique without specifying in what way this is so. USUALLY. - See OCCASIONALLY. VARIABLE. — This means that a feature of an individual has the capacity to exhibit varia- tion. However, in descriptive writing, what is meant by most authors is that a particular char- acter is expressed in one or more states. Each state is either expressed or not expressed in a given individual, and is non-varying therein. Hence, it seems preferable to use “varied”, or “var- ious”. For example, I prefer: “The general form of beetle larvae is widely varied” rather than “ . . . form of beetle larvae is variable”. To me, the second expression implies that a given indi- vidual at different times might be compodeiform, scarabaeiform, and eruciform. Of course, when one writes about larvae of meloid beetles (and larvae of some other groups, too), “var- iable” is indeed the word of choice, because form of an individual larva changes markedly dur- ing development. 4 Examples of uses of many of these words that are counter to my present views are in my earlier publications. Hopefully, these past actions will not be held against me, or used to ne- gate my arguments! I invite the readers of Quaestiones Entomologicae to consider not only my views on the use of these words, but also the more general question of economy and ac- curacy of written expression of observations and ideas. I’ll be pleased to receive comments about these matters. George E. Ball THE EMBRYOLOGY OF L YTTA VIRIDANA LE CONTE (COLEOPTERA: MELOIDAE). IX. THE CENTRAL NERVOUS SYSTEM, STOMATOGASTRIC NERVOUS SYSTEM, AND ENDOCRINE SYSTEM 1 2 3 o J.G. Rempel Department of Biology University of Saskatchewan Saskatoon, Saskatchewan Canada B.S. Heming ^ Department of Entomology University of Alberta Edmonton, Alberta Canada T6G 2E3 N.S. Churchy- Research Station Agriculture Canada Saskatoon, Saskatchewan Quaestiones Entomologicae Canada 13: 5-23 1977 The central nervous system of Lytta viridana arises in the usual way from ventral, longitu- dinal files of neuroblasts in the head, thorax, and first 10 abdominal segments of the embryo. Median nerve strand cells have two fates: clumped, inter segmental cells shift cephalad and differentiate into ganglion cells ; intrasegmental cells probably develop into glial elements of the lateral nerve cords. The perilemma appears to originate from modified outer ganglion cells. The stomatogastric nervous system develops from three evaginations in the roof of the stomodaeum: the frontal ganglion from the first, the hypo cerebral ganglion and corpora car- diaca from the second, and the ventricular ganglion from the third. It is suggested that these three ganglia are not serially homologous with the intersegmental clumps of the post-oral me- dian nerve strand. A corpus allatum invaginates inward from the anterior face of each maxillary base and eventually fuses with a corpus cardiacum below the brain. Probable prothoracic glands pro- liferate inward between the inter ganglionic connectives from the ventral, labial-pro thoracic intersegmental ectoderm. Their cells eventually spread over the surface of a transverse trach- eal commissure emanating from branch 2 of the mesothoracic spiracular tracheal system. Observations are discussed in relation to findings on the origin and function of compara- ble structures in other insects. 1. Previous papers in this series were published in the Canadian Journal of Zoology. Dr. Rempel stipulated that he wished all subsequent ones to appear in Quaestiones Entomologicae. 2. Deceased. 3. Requests for reprints should be sent to this address. 6 Rempel, Heming & Church Le systdme nerveu central de la Lytta viridana se forme d’une facon normale a partir de files ventrales et longitudinales de neuroblastes dans la tdte, le thorax, et les dix premiers segments abdominaux de I’embryon. Les cellules nerveuses et dtroites apparaissent en deux groups: les cellules intersegmentaires groupdes s’assemblent antdrieurement et se diffdrencient en cellules ganglionaires et les cellules intrasegmentaires probablement se develop pent en dldments gliaux de la chaine ner- veuse la t dr ale. Le perilemme semblent dtre formd de cellules ganglionaires externes et modifides. Le systdme nerveux stomatogastrique se ddveloppe a partir de trois evaginations dans la partie supdrieure du stomodeum: du premier vient le ganglion frontal, du second le ganglion hypocdrdbral et les corpora cardiaca, et du troisieme le ganglion ventriculaire. Nous suggdrons que ces trois ganglions ne sont pas des homologues en sdrie avec les groupes de cellules inter- segmentaires du ruban nerveux median et post-oral. Un corpus allatum pdndtre a Vinterieur a partir de la face antdrieure de la base de chaque maxillaire et dventuellement se fusionne avec un corpus cardiacum sous le cerveau. Les glandes prothoraciques probables proliferent vers Vintdrieur entre les connectifs rdunissant les ganglions subesophageaux et prothoraciques Vectoderme intersegmentaire labial-prothoracique du cdtd ventral. Leurs cellules dventuellement se repandent au-dessus de la surface d’une commissure trachdaire transverse dmanant de la branche 2 du systdme trachdaire et spiraculaire de mdsothorax. Les observations sont discutdes en relation aux ddcouvertes sur Vorigine et la fonction de structures comparables chez d’autres insectes. INTRODUCTION Early stages in the embryonic development of blister beetles of the species Lytta viridana LeConte, have been described in previous papers in this series (Rempel and Church 1 965, 1969 a, b, 1971; Church and Rempel 1971). Organogenesis and differentiation of individual organ systems are being treated in separate papers and the first of these, on the respiratory system, has already appeared (Rempel and Church 1972). This paper is devoted to the ner- vous and endocrine systems, to which brief reference has already been made (Rempel and Church 1969b; Church and Rempel 1971). Since Ullmann (1967) described development of the nervous system in Tenebrio molitor L. (Coleoptera, Tenebrionidae) in considerable detail, and because events in L. viridana are very similar, we here concentrate on aspects that supplement her publication. Embry ogenesis of the brain and lateral nerve cords of insects is well known (Edwards 1 969; Anderson 1972 a, b, 1973) and quantitative analysis of the events involved has begun (Bate 1976; Kankel and Hall 1976). However, there is still controversy regarding the development and ultimate fate of the median cord or nerve strand. Although authors agree that the median cord arises from a continuous strip of median ectoderm between the lateral cords, few studies describe its development throughout embryogenesis. Ontogeny of the stomatogastric nervous system has been thoroughly described recently for T. molitor by Ullmann (1967), for Carausius (= Dixippus) morosus Br. (Phasmatodea) by Scholl (1969), for Oncopeltus fasciatus Dallas (Heteroptera) by Dorn (1972, 1975 a, b) and for Stenopsyche griseipennis MacLachlan (Trichoptera) by Miyakawa (1974). Like ear- lier workers, these authors believed the system to develop from three evaginations in the roof of the stomodaeum. According to Ullmann ( 1967), the most anterior evagination gives rise to the frontal ganglion, the second to the hypocerebral ganglion, and the third to the ventricular ganglion. Scholl (1969) also followed development of the frontal ganglion back to the first evagination, but he considered the hypocerebral and ventricular ganglia to come from the third evagination and the corpora cardiaca from the second. Miyakawa ( 1974) clai- med that the recurrent nerve arose from the second evagination. All three interpretations differ slightly from the traditional one of Roonwal (1937) where evagination 2 gives rise to the hypocerebral (occipital) ganglion and corpora cardiaca (pharyngeal ganglia). Although many reviews are available about the structure and function of insect endocrine organs (e.g. Cazal 1 948; Pflugfelder 1958; Herman 1967; Dorn 1972; Gilbert and King 1973; Slama, et al. 1974; Novak 1975), their embryonic development has received little attention. In this paper, we describe the embryogenesis of the central nervous system, stomatogastric Embryology of Lytta viridana IX 7 nervous system and endocrine system of L. viridana. Limited comment is made concerning the neuropile and neurosecretory cells since stains appropriate for these structures were not used. We have described all our methods in previous papers (Rempel and Church 1965, 1969b). OBSERVATIONS Central Nervous System In Lytta viridana , segmentation precedes neurulation (Rempel and Church 1969b). The former process is initiated at approximately 36 h, with coelom formation being well advan- ced by 50 h. At this time, the ectoderm (ect), as seen in parasagittal section (Fig. 1), consists of a single layer of columnar cells with large nuclei and conspicuous vacuoles at their inner ends. In median sagittal section (Fig. 2), the cells appear to form an irregular double layer. Neurulation begins at 56 h. The sequence of the two processes involved is similar to events in T. molitor embryos (Ullmann 1964, 1967) and is probably general in insect development. Many ectodermal cells (Fig. 3. nbl) enlarge in both their nuclei and cytoplasm and begin to stain more strongly than neighbouring cells. Gradually, they withdraw from the surface and move inward. As this process continues, the germ band soon separates into an outer dermato- gene layer (dg. 1) and an inner neurogene layer (ng: 1). The latter consists of neuroblasts (nbl) which, by successive but unequal, vertical, teloblastic division, give rise to small pre-ganglion and ganglion cells (Fig. 4, ggl.c; see also Fig. 3 and 16 in Church and Rempel 1971). Pre-gang- lion cells divide at least once, equally, and often at right angles to the neuroblasts, and gen- erate ganglion cells. This process occurs in continuous, longitudinal files of cells extending on either side of the midline from the region of the stomodaeum to the proctodaeum. Simultaneously, along the midline, cells in intrasegmental regions continue, by equal divi- sion, to form an apparent multi-layered strand of cells (m.n.c), while those in intersegmental regions develop into clumps of large, neuroblast-like, darkly-staining cells (Fig. 5, m.c.c; see also Fig. 17 and 18 in Church and Rempel 1971). At first, the dermatogene layer does not cover these clumps ventrally. These two classes of cells comprise the median nerve strand. This strand begins anteriorly in the intercalary segment (intc. seg) and extends posteriorly to a region behind the tenth lateral cord ganglia (Fig. 7, m.n.c). The first clump of cells is located in the intersegmental region between the mandibular and maxillary segments (Fig. 7, 11, 47), not, as was incorrectly stated earlier (Church and Rempel 1971, but see their Fig. 2), between the intercalary and mandibular segments. Cephalic ganglia develop in the same way as do the lateral nerve cords. Large neuroblasts separate from the dermatogene layer, and, by repeated, unequal, teloblastic division, give rise to pre-ganglion and ganglion cells. The first pair of ganglia arise pre-orally and ultimately form the protocerebrum (Fig. 8, protc; as in T. molitor (Ullmann 1967) these ganglia are tri-lobed, the optic ganglia (op. 1) arising as separate, ectodermal invaginations — (Fig. 42)); the second pair form paraorally and develop into the deutocerebrum (deutc) (Fig. 8); and the third pair arise postorally and later move into a pre-oral position to form the tritocere- brum (Fig. 9, 18, trite). Simultaneously, the stomatogastric nervous system (stmg. n.s) arises from three evaginations in the roof of the stomodaeum (Fig. 10; in 1969, Rempel and Church, incorrectly referred to these as invaginations, as did Ullmann (1967) . Although they are mvaginations of the body wall, they are evaginations of the stomodaeum). The cells surround- ing the evaginations have large, light-staining nuclei and resemble neuroblasts of the central nervous system (Fig. 46). However, as Ullmann ( 1967) pointed out, they divide equally not teloblastically. By 64 h, the lateral nerve cord ganglia have enlarged and have moved mesad. Meanwhile, the clumps of median nerve strand cells (m.c.c) have shifted cephalad from a strictly inter- Quaest. Ent., 1977 13 (1) 8 Rempel, Heming & Church segmental position into an intrasegmental one in the preceding segment (Fig. 6, 39). Here, they later contribute to the posterior gangliomere of each ganglion. By 88 h (see Fig. 1 in Rempel and Church 1971), neuroblasts are prominent in the proto- cerebrum which, by active division, produce columns of pre-ganglion and ganglion cells. The innermost of these, beginning at 72 h, have begun to grow out as axons (Fig. 12). Thus, by this time, a neuropile (npl; often called neuropil) is evident, both here and in all central nerve cord ganglia and connectives (Fig. 40). In each antenna, an ectodermal invagination has arisen which will eventually differentiate into an antennal sense organ (Fig. 12, ant. s.o). Its ontogeny will be the subject of a future contribution. At 88 h, the tritocerebrum (tr. com) is still post oral and is still attached to the intercalary ectoderm (Fig. 11, 13, intc. seg). The mandibular (md. ggl), maxillary (mx. ggl) and labial ganglia (lb. ggl) have moved closer to each other, foreshadowing formation of the subesoph- ageal ganglion (Fig. 1 1) but the 10 abdominal ganglia are still separate at this time. Cells of the median nerve strand clump (m.c.c) are still larger than their neighbours and their cyto- plasm still stains more darkly (Fig. 1 1). They have become oval and have developed clearly discernible axons, which join in a bundle and extend forward and upward, reaching the dor- sal region of the ganglion midway between the anterior and posterior cross commissures (Fig. 1 1, 47, com). These cells maintain this appearance until the end of embryogenesis, although not as obviously. In cross sections through the tritocerebrum (Fig. 13), cells of the median strand differ little from those of ordinary, body wall ectoderm. In the mandibular region (Fig. 14), the strand appears as a cluster of cells having faintly stained nuclei. Sections through the poster- ior cross commissure (com), of the maxillary segment (Fig. 1 5) show numerous axonal ex- tensions into the neuropile. Here, the median strand cells are small, rectangular and more lightly staining than neighbouring ganglion cells. Sections through the median nerve strand clump (m.c.c) of the labial ganglion (Fig. 16, 41) show its cells to be distinctly separate from those of the lateral cord ganglia. Finally, in the prothoracic-mesothoracic interganglionic re- gion (Fig. 17), median strand cells are indistinguishable from those of ordinary, body wall ectoderm, suggesting that, at this time, the median strand has become discontinuous in inter- segmental regions. By 120 h(see Fig. 18 and 19 in Rempel and Church 1971), cephalization of the embryo has become more pronounced, and the head has become clearly set apart from the rest of the body. The gnathal ganglia have fused to form the subesophageal ganglion (Fig. 19, sb. ggl) and abdominal ganglia 9 and 10 have amalgamated (see Fig. 6 in Rempel and Church 1972). The lateral nerve cord ganglia have moved to the midline and have fused to form a single ganglion- ic mass in each segment. Ganglion cells have encroached ventrally upon the median strand, restricting it to the dorsal region of each ganglion (Fig. 20, 43, m.n.c). The strand seemingly has disappeared from intersegmental regions (Fig. 21) of the ventral nerve cord but it and its clumps (m.c.c) are retained in intrasegmental regions. For example, three clumps are clearly visible in the subesophageal ganglion. (Fig. 19). A characteristic feature of this stage in development, is the presence of long, cytoplasmic strands (cyt. std) in the intersegmental regions, extending from the body wall (bd. w) to the interganglionic connectives (int. cn) and from there to the developing midgut (Fig. 21, mdgt). We do not know what significance they have and in embryos older than 132 h, they are no longer present. By 132 h, a perilemma has appeared. It is best developed around the interganglionic con- nectives (Fig. 22) and consists of a layer of cells, the perineurium (prn), and its secretion pro- duct, the neural lamella (nr. 1ml; often termed neurilemma or neural lemma). Here and there, the interface between ganglion cells and neuropile (npl) is occupied by cells (gll. c) that have Embryology of Lytta viridana IX 9 small, light-colored nuclei (Fig. 20, 43). These cells first appear at about 104 h, and apparently originate from the median strand. We believe them to be glial. Springer (1967) and Springer and Rutschky (1969) referred to them as the inner sheath. By 180 h, the entire central nervous system is enclosed by a well-developed perilemma. It is especially prominent about the interganglionic connectives (Fig. 25, 28, 56) and dorsally in each ganglion (Fig. 44). By this time, most neuroblasts have disappeared, although a few dividing in the brain remain active until hatching. The sheath of glial cells (gll.c) separating neuropile from ganglion cells within each ganglion is much more pronounced (Fig. 44) and a few small, dark-staining glial cells are scattered among ganglion cells throughout the central nervous system. Although inadequate staining makes their details impossible to sort out, fibre tracts and glomeruli are now clearly visible within the brain and ventral nerve cord ganglia. These first begin to take shape at 120 h (Fig. 43) and become steadily more complex until hatching at 250-264 h. (Fig. 44, 48). Between 1 80 and 200 h, most changes affect the distribution of glial elements enclosing the neuropile (Fig. 23). The cells of the median strand clump (m.c.c) are still recognizable but can be confused easily with other ganglion cells. In parasagittal sections of the nerve cord (Fig. 24), the glial cells (gll. c) appear to be stacked up vertically at each end of each intergang- lionic connective, but transverse sections through these regions (Fig. 45) show they are not. In cross sections (Fig. 25-28) made at the points indicated by lines in Fig. 23 and 24, differ- ences in distribution of glial cells can also be seen. In interganglionic connectives, the perineu- rium (prn) is thick dorsally and very thin ventrally, whereas the opposite is true of the neural lamella (nr. 1ml) (Fig. 24, 25, 28, 56). Except for continued differentiation of neuropile cen- tres, no other significant changes were observed in nervous system development up to the time of hatching (250 to 264 h). Stomatogastric Nervous System (stmg. n.s.) In Lytta viridana embryos, when the stomodaeum begins to invaginate at 56 h, the three evaginations in its roof — the anlagen of the stomatogastric nervous system (stmg. n.s) - are already evident (see Fig. 50-53 in Rempel and Church 1969b). At 64 h (Fig. 10), nuclei of cells surrounding the evaginations become enlarged and, from 72 to 80 h (Fig. 1 1, 46), strands of cells arise behind each evagination and begin to stream forward over the roof of the stomo- daeum. This movement is accompanied by considerable mitotic activity. By 96 h, the frontal ganglion (frt. ggl) has begun to enlarge and neuropile to differentiate within it (Fig. 47) (the latter actually commences at about 88 h). Simultaneously, fibres of the recurrent nerve (ret. nv) become apparent just behind the frontal ganglion. At 104 h (Fig. 29), forward streaming of nerve cells from the three stomodaeal evaginations is still evident but, by 1 20 h (Fig. 19), evaginations 1 and 2 have begun to disappear, and the three cell streams are no longer separ- able. By 120 h, the frontal ganglion (frt. ggl) is well-developed (Fig. 19) and by 132 h, the hypo- cerebral ganglion (hyp. ggl) has appeared as a slight swelling in the recurrent nerve (ret. nv) below the pars intercerebralis (prs. intr) of the brain (Fig. 33). Evaginations 1 and 2 have dis- appeared and all cell proliferation now seems to arise from evagination 3 — an indication that this is a source of most cells comprising the recurrent nerve (ret. nv). The tritocerebral (= fron- tal) connectives of the frontal ganglion form at 1 20 h, (see Fig. 31, C, D in Rempel and Church 1971). By 156 h, the system is essentially complete, and from here on most changes involve ganglionic enlargement and an increase in length of the recurrent nerve to match continued growth of the stomodaeum. Fig. 38 shows the system as it appears at 216 h and Fig. 48 at 264 h. Quaest. Ent., 1977 13 (1) 10 Rempel, Heming & Church Endocrine System The endocrine system of L. viridana consists of neurosecretory cells, which we will not consider, and paired corpora cardiaca, corpora allata, and prothoracic glands. The corpora cardiaca (crp. crd) first become recognizable at 96 h. They originate from cells emanating from evagination 2 in the roof of the stomodaeum, also the source of the hypocere- 1 bral ganglion (Fig. 29, 47). These cells move laterally around the stomodaeum (Fig. 31 c, 49) and, by 1 1 2 h, have been carried forward by the cell streams to the region of the pars inter- cerebralis (prs. intr). As this movement occurs, the cells comprising the cardiaca, become more and more loosely arranged (Fig. 50). Later, each gland rudiment shifts laterally and attaches to the posterior surface of the brain. By 132 h, the glands have established contact posterior- ly with the corpora allata (crp. all) (Fig. 37). Differentiation of the glands’ cells first becomes evident at about 1 20 h, both secretory(s) and glial elements being present (Fig. 50). The former j resemble ganglion cells in their dark-staining, homogeneous, cytoplasm; the latter form a loose, j parenchymatous network. By 250 h (Fig. 37, 51, 54), this network has contracted, so that the fully-formed glands are not much larger than the allata. The corpora allata (crp. all) first become apparent at 56 h, as small, ectodermal invagina- tions of the anterior base of each maxilla (Fig. 34, 52). Each invagination grows inward and dorsad until, at 96 h (Fig. 53), it reaches a position below the caudal extension of the anter- ior tentorial arm (a. tent). It now shifts slightly laterally and then dorsally over the tentorial arm, maintaining, throughout this movement its connection with the body wall (Fig. 35, 36). At 1 1 2 h, this connection breaks. In the meantime, the ventral tracheal trunk of each side (2 in Fig. 6 in Rempel and Church 1972) has extended forward from the mesothoracic spiracle over the posterior tentorial arm (p. tent). At 120 h, this trunk sends one branch (2b) to the gnathal appendages and another (2a) to the brain. The second branch becomes closely applied to the corpus cardiacum-corpus allatum complex (Fig. 13 and 17 in Rempel and Church 1972). At 120 h, each corpus allatum reaches its final position and, at 132 h, re-establishes its connection with the body wall via a clear, tendon-like strand (Fig. 37, 54). The connective tissue sheath of the corpus allatum, described by Weismann ( 1 926) in C. morosus and by Roonwal (1937) in L. migratoria , was not evident in our preparations, even under oil immersion (Fig. 54). At 96 h, two, small protuberances proliferate inward from the ventral, labial-prothoracic intersegmental ectoderm (Fig. 30, 55, pthr. gl). They originate close together and grow dor- sad between the developing interganglionic connectives, (int. cn). At the same time, tracheae 2 (Fig. 6, 9 and 17 in Rempel and Church 1972) each give off a medially-directed branch (tracheae 9) to form a commissure above the ventral nerve cord between the subesophageal and prothoracic ganglia (Fig. 55, 56, trch). The proliferated cells become associated with the tips of these branches as they advance, but maintain their connections with the body wall until 156 h (Fig. 32, 33). Gradually, the cells from the proliferations spread over the surface of the tracheal commissure (Fig. 25, 38, 56). At later stages, they are very difficult to see be- cause they so closely resemble cells of the tracheal epithelium (Fig. 25, 56). We believe these cellular proliferations to constitute the prothoracic glands. We have no idea how far they spread over tracheae 1, 2 and 9 (Rempel and Church 1972, Fig. 6, 9, 17), since, even in mature larvae of other beetles, such glands are difficult to trace (Svrivastava 1959). Additionally, the area between the subesophageal-prothoracic interganglionic connec- tives is eventually occupied by a complex array of tracheal branches (Fig. 56). This also causes difficulties in distinguishing between glandular and tracheal tissues. Embryology of Lytta viridana IX 11 DISCUSSION Central and Stomatogastric Nervous Systems Embryogenesis of the brain, lateral nerve cords and stomatogastric nervous system in Lytta viridana has been described very briefly because the sequence of events is so similar to that occurring in Tenebrio molitor L. (Ullmann, 1967) and Sitophilus [= Calendra) oryzae (L.) (Coleoptera, Curculionidae) (Tiegs and Murray 1938). The median strand - Authors agree generally about the origin of this system, and the des- cription we have presented outlines the usual pattern. The median strand arises from ectoder- mal cells along the mid-ventral line of an embryo in a region extending from behind the in- tercalary segment to the end of the tenth abdominal segment. Early in its development, two types of cells are recognized. In intrasegmental regions, cells of the median strand resemble those of the body wall ectoderm; in intersegmental regions, they assume the appearance of neuroblasts. These latter cells divide teloblastically, like typical neuroblasts, and in fact, seem to be neuroblasts. Nonetheless, we prefer to use the word “clump” for groups of these cells, as did Springer (1967) and Springer and Rutschky ( 1 969). The forward shift of each clump of the median strand from an intersegmental to an intrasegmental position is in agreement with descriptions of this process for other insects (Springer 1 967; Springer and Rutschky 1969). General agreement about development of the median strand gives way to disagreement and controversy about fate and role of the components of this system (Edwards 1969; An- derson 1972a and b and 1973). Details are provided below. Fate and role of the clumps of the median strand — Ullmann ( 1 967) and Springer and Rutschky ( 1 969) summarized information presented by earlier workers about this topic. It was claimed by Ullmann and others that clumps in embryos of S. oryzae and T. molitor contribute to development of the definitive ganglia, whereas in the insects studied by Sprin- ger and Rutschky (various hemipterans, orthopterans, beetles, moths and dipterans), the clumps disappear after katatrepsis. In embryos of L. viridana , the clumps are present from their first appearance (56 h) until after hatching (±250 h). We agree with Ullmann (1967) that these cells probably act as ganglion cells because they develop axons (Fig. 11, 19, 47). Although, for most insects studied, the clump cells seem to be involved with development of the nervous system, Miyakawa ( 1 974) reports that in embryos of S. griseipennist the inter- ganglionic portions of the median cord of the thoracic segments develop into furcae, and thus do not have a nervous function. Clumps of the median strand and segmentation of the insect head.— Because the clumps are intersegmental in all post-oral segments, we assume that in ancestral hexapods they were present also in those segments that arise post-orally but which, in more highly evolved stocks, become pre-oral. If one could recognize the clumps in heads of extant insects, one would have additional evidence for deducing the number of segments involved therein. (See Malzacher 1968; Scholl 1969; Rempel and Church 1971 ; and Rempel 1975 for comprehensive discus- sions of head segmentation). Although it is tempting to suggest that the stomatogastric ner- vous system with its three ganglia (frontal, hypocerebral, and ventricular) is the forward con- tinuation of the median strand, and that the ganglia represent respectively the clumps of the preantennal (labral), antennal, and tritocerebral segments, this is probably not so, for the fol- lowing reasons: As cephalization occurred, the floors of these segments supposedly contribu- ted to the floor of the stomodaeum. However, each post-oral clump is situated behind the pos- terior cross commissure of its ganglion. How then could the clump of, for instance, the tri- tocerebral segment, get onto the roof of the stomodaeum to form the ventricular ganglion while its commissure remained post-oral? It seems developmental^ impossible. Thus, it also Quaest. Ent., 1977 13 (1) 12 Rempel, Heming & Church seems impossible that these ganglia are the homologues of clumps of the median strand. Pro- bably in extant insects, the segments in question are without a median strand and without clumps, and this is probably the result of atrophy occurring in the extinct ancestral stock. Role of the intraganglionic portions of the median strand. - Springer (1967) and Springer and Rutschky (1969) showed that cells in this region do not become functional ganglion cells, and this seems to be generally accepted. However, authors disagree about what these cells do. Ullmann (1967) claimed that those near the periphery contribute to formation of the perilem- ma in dorsal portions of the ganglia. On the other hand, Miyakawa ( 1 974) reported that, in S. griseipennis embryos, the perilemma over most of each ganglion seemed to arise from mod- ified outer ganglion cells. Our observations for L. viridana embryos support the conclusion of Miyakawa. Cells of the intraganglionic portions of the median strand are involved in production of glial cells. We believe that glial elements associated with axons of the interganglionic connec- tives also originate from the median strand. We conclude, therefore, that the principal role of the median cord is to form glial cells associated with nerve fibres of the lateral nerve cords. More specifically, in embryos of L. viridana at 104 h, some median strand cells move lat- erally between neuropile and innermost ganglion cells to form an inner sheath of glial cells (Fig. 20, 43, gll. c), a process very similar to that occurring in embryos of S. griseipennis (Miy- akawa 1974). We disagree with Springer and Rutschky (1969) who claimed that the inner sheath develops from the innermost ganglion cells. Glial cells. - Three of the four types of glial cells described by Wigglesworth (1972) from specimens of Rhodnius prolixus Stal (Heteroptera) are evident in larvae of L. viridana that are ready to hatch (prolarvae). These cells are illustrated in Fig. 44: type i (perineurium); type ii (cells scattered among ganglion cells); and type iv (neuropile sheath). Type iii cells having giant nuclei are not present, although some type iv cells have quite large nuclei. All three types of glial cells are also evident in ganglia of the stomatogastric nervous system. Endocrine System This includes the corpora cardiaca, corpora allata, and prothoracic glands. Each of these paired glands is discussed below. Corpora cardiaca. — Embryogenesis of these glands has been studied by a number of work- ers, notably Weismann ( 1 926, in C. morosus), Roonwal (1937, in Locusta migratoria (L.) (Orthoptera)), Pflugfelder ( 1 937, in C. morosus ), and Dorn (1972, and 1975a, in O. fasciatus). All agree that ontogenetically, these glands originate with the hypocerebral ganglion, by cell migration from the roof of the stomodaeum. Dorn (1975a) followed embryogenesis of the corpora cardiaca in eggs of O. fasciatus, using the transmission electron microscope. In these insects, the glands appear at 56 h, and begin to differentiate into glandular and glial components at 96 h. The glandular cells assume a spherical distribution about a lumen (his Fig. 4 and 8) into which grow cell projections, pro- bably axons of the nervi corporis cardiaci. This occurs when the glands attach to the aorta. Evidence of protein synthesis appears at 96 h, and by 1 1 1 h, neurosecretory granules are evident. During hatching (at 124 h) the cells of the glands appear to be secretory. Secretory cells in the cardiaca of embryos of L. viridana are probably homologous with the “instrinsic secretory cells” described by Schoonveld (1970) in the cardiaca of adult speci- mens of Leptinotarsa decemlineata Say (Coleoptera, Chrysomelidae). Phylogenetically, Hanstrom ( 1 942) assumed the corpora cardiaca to have evolved from a stomodaeal ganglion. Because these glands occur in all apterygote insects which have been examined for them, and because the corpora allata do not, Novak (1975) assumed that the former glands are evolutionarily older than the latter glands. Embryology of Lytta viridana IX 13 Corpora allata. — In embryos of different taxa of insects, these glands seem to originate from different germ layers, and in different positions. For example, in embryos of L. virid- ana, O. fasciatus (Dorn 1972), and S. griseipennis (Miyakawa 1974), they arise as ectodermal invaginations at the anterior base of each maxilla. On the other hand, in L. migratoria (Roon- wal 1937) and C. morosus embryos (Pflugfelder 1937), the glands arise as paired, ectodermal invaginations between the mandibular and maxillary segments. And, in embryos of S. oryzae, they originate from mesoderm of the antennal coelomic sacs (Tiegs and Murray 1938). Accor- ding to Pflugfelder (1937), some authors have even reported corpora allata as being of endo- dermal origin. Probably some of these observations are incorrect. Certainly the question of ontogenetic origin of these glands should be investigated using a wide taxonomic spectrum of pterygote insects and a diversity of approaches. The ultrastructure and function of the developing corpora allata of O. fasciatus embryos have been studied by Dorn (1975 b) who showed that high titers of juvenile hormone present just before hatching are probably the result of activity of these glands. Pro thoracic glands. - For Coleoptera, Srivastava (1959) described the prothoracic glands of larvae of 1 5 species (none were meloids). These glands occur in the head, cervical region and prothorax as thin cords or sheets of cells closely associated with one or both of two large tracheae extending from the prothoracic spiracles into the head. Embryogenesis of the pro- thoracic glands was previously unknown for Coleoptera, and has been little studied in any insect species. Differences of opinion among authors suggest i) that the prothoracic glands of different insects are not homologous, or ii) that some accounts of their origin are in error, or iii) that some tissues referred to as prothoracic glands are some other structure. We cannot resolve the problem, but we review the different viewpoints, below. According to Gilbert and King ( 1 973), Toyama ( 1 902) identified the prothoracic glands in embryos of Bombyx mori F. (Fepidoptera) as epithelial invaginations of the labial segment of the head. A similar origin was postulated for the glands of embryos of Dysdercus cingula- tus (Fab.) (Heteroptera) by Wells ( 1 954), and for those of O. fasciatus embryos by Dorn ( 1 972). In embryos of Schistocerca gregaria (Forskal) (Orthoptera), the prothoracic glands invaginate before katatrepsis from lateral ectodermal regions between the maxillary and labial segments (Micciarelli and Sbrenna 1972). Novdk(1975) suggested that the prothoracic glands of most pterygote insects originate from the ventral margin of the prothoracic segment, basing his con- clusion on their innervation from the prothorax, and on their supposed homology with the cephalic nephridia of apterygote insects. The proposed labial-prothoracic origin for these glands in L. viridana embryos, if proved, would support Novak’s conclusion. We at first thought that the labial diverticula (Fig. 19, lb. div), not the intersegmental pro- liferations suggested here, might be the progenitors of the prothoracic glands, because these diverticula originate from the labial segment, as do the glands of some of the insects named above. The diverticula grow caudad under the suboesophageal ganglion where they bend ab- ruptly upward to end in the labial-prothoracic region. Ullmann (1967) detailed evidence to suggest that these diverticula develop into “maxillary glands”. We have followed developmen- tally the “glands” of L. viridana embryos to similar but very indistinctly developed structures in prolarvae of this species. Publications also present conflicting evidence concerning the function of embryonic pro- thoracic glands (see discussions and refs, in Dorn 1972; and Micciarelli and Sbrenna (1972)). Dorn (1 972) noted that the glands of O. fasciatus showed three cycles of activity (as did the neurosecretory cells and corpora cardiaca) that correlated well with secretion of three pre- hatch cuticles. A similar correlation was noted by Micciarelli and Sbrenna (1972) in the two moults of S. gregaria embryos. However, these authors showed that isolated embryonic abdo- mens, lacking these glands, were also capable of secreting two cuticles and concluded that Quaest. Ent., 1977 13 (1) 14 Rempel, Heming & Church ! embryonic apolyses are not under control of the prothoracic glands. Since much recent evi- dence suggests that ecdysone can be synthesized in organs other than the prothoracic glands (Nakanishi, et al. 1 972; Romer, et al. 1 974; Hsiao, et al. 1 975), action of this hormone in em- bryonic moults has still not been ruled out. Embryos of L. viridana produce a single, delicate embryonic cuticle between 1 20 and 132 h, shortly after katatrepsis, about the time that secretory cells become evident in the corpora cardiaca. However, there is no obvious change at this time in cells of the corpora allata or pro- ' thoracic glands. Neither is any change noted in these cells at 168 to 1 80 h, when deposition of larvae cuticle begins. Thus, we have no positive evidence of endocrine function in embryos of L. viridana. Writing the above discussion on the origin and function of embryonic insect endocrines was frustrating because of the conflicting results presented in the literature. We do not believe that structures as fundamental to insect development as the corpora allata and prothoracic glands can have so many different ontogenetic origins. Diversity in site of origin implies multiple in- dependent origin during insect evolution. This conflicts with the proven similarity in biochem- ical function of these glands in representative insects of most orders (Gilbert and King 1 973; Slama, et al. 1 974; Novak 1975). What is required to resolve the conflict are detailed compara- tive embryological studies of the most critical kind using all methods available. Dorn’s (1972, 1975 a, b) studies in which conventional, ultrastructural and experimental methods are com- bined, provide a start in the right direction. It would also help if embryological studies the quality of Dorn’s were carried out on individuals of species for which experimental proof of gland function exists. ACKNOWLEDGEMENTS An early draft of this manuscript was read and constructively criticized by G.E. Ball, D.A. Craig and B.K. Mitchell. Dr. Craig also offered suggestions to B.S. Heming on photomicrogra- phy. J. Scott prepared the photographic plates, Mrs. Patricia Cookson typed the manuscript and H. Goulet translated the abstract into French. We thank all these individuals for their generous assistance. Publication costs were met by grants from the Strickland Memorial Trust Fund and the National Research Council of Canada (A5756 to B.S. Heming). REFERENCES Anderson, D.T. 1972a. The development of hemimetabolous insects. In Counce, S.J. and C.H. Waddington (eds). Developmental Systems: Insects. Vol. I. Academic Press, London and N.Y. pp. 95-163. Anderson, D.T. 1972b. The development of holometabolous insects. Ibid. pp. 165-242. Anderson, D.T. 1973. Embryology and Phylogeny in Annelids and Arthropods. Pergamon Press, N.Y. and Oxford 495 p. Bate, C.M. 1976. Embryogenesis of an insect nervous system. I. A map of the thoracic and abdominal neuroblasts in Locusta migratoria. Journal of Embryology and Experimental Morphology 35; 107-123. Cazal, P. 1 948. Les glandes endocrines retrocerebrales des insectes. Etude morphologique. Bulletin biologique de la France et de la Belgique. Supplement 32: 1-228. Church, N.S. & J.G. Rempel 1971. The embryology of Lytta viridana LeConte (Coleoptera: Meloidae). VI. The appendiculate 72-h embryo. Canadian Journal of Zoology 49: 1 563-1570. Dorn, A. 1972. Die endokrinen Drusen im Embryo von Oncopeltus fasciatus Dallas (Insecta, Embryology of Lytta viridana IX 15 Heteroptera). Morphogenese, Funktionsaufnahme, Beeinflussung des Gewebewachstums und Beziehungen zu den embryonalen H&utungen. Zeitschrift fur Morphologie und 6kologie der Tiere 71 : 52-104. Dorn, A. 1 975a. Electronenmikroskopische Studien iaber Differenzierung und Funktionsauf- nahme der Corpora cardiaca im Embryo von Oncopeltus fasciatus Dallas (Insecta, Heteroptera). Cytobiologie 10: 235-248. Dorn, A. 1975b. Struktur und Funktion des embryonalen Corpus allatum von Oncopeltus fas- ciatus Dallas (Insecta, Heteroptera). Verhandlungen der Deutschen zoologischen Gesellsch- aft 67: 85-89. Edwards, J.S. 1969. Postembryonic development and regeneration of the insect nervous system. Advances in Insect Physiology 6: 97-137. Gabe, M. 1966. Neurosecretion. Pergamon Press, N.Y. 872 p. Gilbert, L.I. & D.S. King 1973. Physiology of growth and development: endocrine aspects. In Rockstein, M. (ed.) The Physiology of Insecta. 2nd Ed. Vol. 2. Academic Press. N.Y. pp. 249-370. Hanstrom, B. 1942. Die Corpora cardiaca und Corpora allata der Insekten. Biologia generalis 15: 485-531. Herman, W.S. 1967. The ecdysial glands of arthropods. International Review of Cytology 22: 269-347. Hsiao, T.H., C. Hsiao & J. de Wilde 1 975. Moulting hormone production in the isolated abdo- men of the Colorado beetle. Nature 255: 727-728. Kankel, D.R. & J.C. Hall 1 976. Fate mapping of nervous system and other internal tissues in genetic mosaics of Drosophila melanogaster. Developmental Biology 48: 1-24. Malzacher, P. 1968. Die Embryogenese des Gehirns paurometaboler Insekten. Untersuchungen an Carausius morosus und Periplaneta americana. Zeitschrift fhr Morphologie und Okologie der Tiere 62: 103-161. Micciarelli, A.B. & G. Sbrenna 1972. The embryonic apolyses of Schistocerca gregaria (Orth- optera). Journal of Insect Physiology 18: 1027-1037. Miyakawa, K. 1 974. The embryology of the caddisfly Stenopsyche griseipennis MacLachlan (Trichoptera, Stenopsychidae). III. Organogenesis: Ectodermal derivatives. Kontyu 42: 305-324. Nakanishi, K., H. Moriyama, T. Okauchi, S. Fujioka & M. Koreeda 1972. Biosynthesis of a and (5 ecdysones from cholesterol outside the prothoracic gland in Bombyx mori. Science 176: 51-52. Novak, V.J.A. 1975. Insect Hormones. 2nd English Ed. Chapman & Hall, London. 600 p. Pflugfelder, O. 1937. Bau, Entwicklung, und Funktion der Corpora allata und cardiaca von Dixippus morosus Br. Zeitschrift fur wissenschaftliche Zoologie 149: 477-512. Pflugfelder, O. 1958. Entwicklungsphysiologie der Insekten. 2 Auflage. Akad. Verlag. Geest & Portig. K - G., Leipzig. 490 p. Rempel, J.G. 1975. The evolution of the insect head: the endless dispute. Quaestiones Ento- mologicae 11: 7-25. Rempel, J.G. & N.S. Church 1965. The embryology of Lytta viridana Le Conte (Coleoptera: Meloidae). I. Maturation, fertilization and cleavage. Canadian Journal of Zoology 43: 915- 925. Rempel, J.G. & N.S. Church 1969a. The embryology of Lytta viridana LeConte (Coleoptera: Meloidae). IV. Chromatin elimination. Ibid. 47: 351-353. Rempel, J.G. & N.S. Church 1969b. The embryology of Lytta viridana LeConte (Coleoptera: Meloidae). V. The blastoderm, germ layers, and body segments. Ibid. 47: 1 1 57-1 1 71 . Rempel, J.G. & N.S. Church 1971. The embryology of Lytta viridana LeConte (Coleoptera: Quaest. Ent., 1977 13 (1) 16 Rempel, Iieming & Church Meloidae). VII. Eighty-eight to 132 h: the appendages, the cephalic apodemes, and head segmentation. Ibid. 49: 1571-1581. Rempel, J.G. & N.S. Church 1972. The embryology of Lytta viridana LeConte (Coleoptera: Meloidae). VIII. The respiratory system. Ibid. 50: 1547-1554. Romer, F., H. Emmerich & J. Nowock. 1974. Biosynthesis of ecdysones in isolated prothora- cic glands and oenocytes of Tenebrio molitor in vitro. Journal of Insect Physiology 20: 1975-1987. I Roonwal, M. L. 1 937. Studies on the embryology of the African migratory locust Locusta migratoria migratorioides, R & F. II. Organogeny. Philosophical Transaction of the Royal Society. Series B 227: 175-244. j Scholl, G. 1 969. Die Embryonalentwicklung des Kopfes und Prothorax von Carausius morosus \ Br. (Insecta, Phasmida). Zeitschrift fur Morphologie und Okologie der Tiere 65: 1-142. Schoonveld, H. 1970. Structural aspects of neurosecretory and corpus allatum activity in the adult Colorado beetle, Leptinotarsa decemlineata Say, as a function of day length. Nether- lands Journal of Zoology 20: 1 51-237. Slama, K., M. Romanuk & F. Sorm. 1974. Insect Hormones and Bioanalogues. Springer Verlag N.Y. and Wien. 477 p. j Springer, C.A. 1 967. Embryology of the thoracic and abdominal ganglia of the large milkweed bug, Oncopeltus fasciatus (Dallas) (Hemiptera, Lygaeidae). Journal of Morphology 122: 1-1 8. j Springer, C.A. & C.W. Rutschky, III. 1969. A comparative study of the embryological develop- ment of the median cord in Hemiptera. Ibid. 1 29: 375-400. Svrivastava, U.S. 1959. The prothoracic glands of some coleopteran larvae. Quarterly Journal of Microscopical Science 100: 51-64. Tiegs, O.W., & F.V. Murray 1938. The embryonic development of Calandra oryzae. Ibid. 80: j 159-271. Toyama, K. 1902. Contributions to the study of silkworms. I. On the embryology of the silk- worm. Bulletin of the College of Agriculture. Tokyo Imperial University 5: 73-1 18. Ullmann, S.L. 1964. The origin and structure of the mesoderm and the formation of the coelomic sacs in Tenebrio molitor L. (Insecta, Coleoptera). Philosophical Transactions of the Royal Society. Series B 248: 245-277. Ullmann, S.L. 1967. The development of the nervous system and other ectodermal derivatives j in Tenebrio molitor L. (Insecta Coleoptera). Ibid. 252: 1-25. Wells, M. J. 1 954. The thoracic glands of Hemiptera-Heteroptera. Quarterly Journal of Micro- scopical Science 95: 231-244. Weismann, R. 1926. Entwicklung und Organogenese der Colomblasen. pp. In Leuzinger, H., R. Weismann und F.E. Lehmann (eds.). Zur Kenntnis der Anatomie und Entwicklungeschich- te der Stabheuschrecke Carausius morosus Br. G. Fisher Verlag, Jena. Wigglesworth, V.B. 1972. The Principles of Insect Physiology. 7th Ed. Chapman and Hall. London. 827 p. Embryology of Lytta viridana IX 17 ABBREVIATIONS abd. ggl abdominal ganglia mdgt midgut ant antenna mes mesoderm ant s. o antennal sense organ m. n. c median nerve cord a. tent anterior tentorial arm mx maxilla bd. w body wall mx. ggl maxillary ganglion brn brain nbl neuroblast com commissure ng. 1 neurogene layer crp. all corpus allatum npl neuropile crp. crd corpus cardiacum nr. 1ml neural lamella cyt. std cytoplasmic strand nv nerve deutc deutocerebrum op. c optic cup dg. 1 dermatogene layer op. 1 optic lobe ect ectoderm prn perineurium ent r enteron rudiment protc protocerebrum frt. ggl frontal ganglion prs. intr pars intercerebralis ggl- c ganglion cell p. tent posterior tentorial arm gll. c glial cell ret. nv recurrent nerve haem haemocytes s secretory cells of corpora cardiaca hyp. ggl hypocerebral ganglion sb. ggl subesophageal ganglion int. cn interganglionic connectives spl. m splanchnic mesoderm intc. ect intercalary ectoderm stmg. n. s stomatogastric nervous system intc. seg intercalary segment stom stomodaeum lb labium stom. rf roof of stomodaeum lb. div labial diverticulum sub. b subesophageal body lb. ggl labial ganglion tr. com tritocerebral commissure lm labrum tent tentorium 1. n. c lateral nerve cord thr. ggl thoracic ganglia m. c. c median strand clump trch trachea md mandible trite tritocerebrum md. fix. ap mandibular flexor apodeme vnt. ggl ventricular ganglion md. ggl mandibular ganglion yk yolk Quaest. Ent., 1977 13 (1) 18 Rempel, Heming & Church Fig. 1. Body wall and coelomic sacs at 50 h, parasagittal section. Fig. 2. Body wall at 50 h, median sagittal section. Fig. 3. Same, at 56 h, parasagittal section, showing dermatogene (dg. 1) and neurogene (ng. 1) layers, the latter with neuroblasts (nbl). Fig. 4. Same, at 60 h, showing formation of pre ganglion and ganglion cells (ggl. c). Fig. 5. Same, median sagittal section, show- ing “clumps” of median strand cells (m.c.c). Fig. 6. Same, at 64 h, showing forward shift of median strand clump. Fig. 7. Diagram of central nervous system, showing cephalic ganglia, lateral nerve cords (l.n.c) and median nerve strand (m.n.c). Fig. 8. Parasagit- tal section through protocephalic lobe and antenna (ant) at 56 h, showing formation of protocerebral (protc) and deutocerebral (deutc) neuroblasts. Fig. 9. Same, through stomodaeum, showing formation of tritocerebral (trite) neuroblasts from intercalary ectoderm (intc. seg). Fig. 10. Median sagittal section through stomodaeum at 64 h, showing formation of stomatogastric nervous system (stmg. n.s). Fig. 11. Same, at 88 h, showing median strand clumps (m.c.c). Fig. 12-17. Transverse sections through dev- eloping nerve cord at 88 h, taken at points indicated by lines in Fig. 11: 12, Through antenna and stomodaeum, showing devel- oping ganglion cells, stomatogastric nervous system, (stmg. n.s) and antennal sense organ (ant. s.o); 13, Through tritocerebral ganglion; 14, Through mandibular ganglion; 15, Through posterior commissure of maxillary ganglion; 16, Through median strand “clump” (m.c.c) of labial ganglion; 17, Through prothoracic-mesothoracic intersegmental region and inter ganglionic connectives (int. cn). Fig. 18. Parasagittal section through tritocerebral ganglion (trite) and anterior tentorial arm (a. tent). Quaest. Ent., 1977 13 (1) 20 Rempel, Heming & Church 'sp * 5 o S o — 5 g -r -a o -2 >> .tJ .O bfi g 13 s § PU CU •S if 2 =3 3 S O O- V >. M S £ -S a E £ £ O - T) x £ e S -o 6 i— I &h r~| <« £ M ;i § H £ P o 1/3 ^ H .O O S £ “ M z UJ o 0 -J > 1 Cl Fig. 1. Distribution of ovariole type in the Insecta. 81 (knowledge of whose relationships is still inadequate) accepted sister group relations of the insect orders are becoming stabilized (see Hennig, W. 1969. “Die Stammesgeschichte der In- sekten” and Kristensen, N.P. 1975. Zeitschrift fur zoologische Systematik und Evolutions- forschung 13: 1-44). If details of particular abdominal structures or developmental sequences are plotted on a phylogenetic dendrogram of the insect orders, one sees quickly whether they arose independently several times or only once in the common ancestor of monophyletic ass- emblages of orders. For example, if one does this for ovariole type (Fig. 1) one sees at a glance that the panoistic type is plesiomorphic but has been lost and secondarily regained either once (Siphonaptera) or twice (Thysanoptera and Siphonaptera), that the polytrophic type has in- dependently arisen four times (in Collembola, Diplura, Dermaptera, and in the common an- cestor of all remaining orders), and that the acrotrophic type has arisen independently twice (Hemiptera and Coleoptera) from ancestors having the polytrophic type. If such a plot is then superimposed on an identical diagram in which species diversity per order is indicated, one sees that the evolution of trophocytes (nurse cells) has probably contributed directly to the evolu- tionary success of those orders whose members’ ovarioles are characterized by their presence. The selective advantage of meroistic (nurse cell-containing) ovarioles is that their oocytes grow faster and are produced in greater numbers than those of panoistic ovarioles because of the large amount of template DNA (up to 1024C) available for oogenesis in the polyploid nuclei of their nurse cells (only 4C amounts of DNA are available for RNA synthesis in the germinal vesicles of panoistic oocytes - see Mahowald, A.P. 1972 in Vol. Developmental Systems: In- sects). Use of Hennig’s methods would add rigor to Matsuda’s analysis and would make more ob- vious structural and developmental dines. This would result in Matsuda’s book being of use to a larger number of workers than it is now. It would also enable readers to make conpari- sons more easily and to use the book in making predictions about structure of the abdominal appendages in unstudied taxa. The volume is well-produced but there are many typographical errors and some of the print of my copy ended up on my fingers and on the pages of the first draft of this review. Some fig- ures, though fully-labeled, are difficult to interpret because they were sloppily done or repro- duced with too much reduction. They compare unfavourably with those of Anderson’s recent treatise (1973. “Embryology and Phylogeny in Annelids and Arthropods”) in the same series. In spite of these criticisms, I consider Matsuda’s book to be a major contribution because of its comprehensive bibliography and because of the vast amount of information it contains. ACKNOWLEDGEMENT I thank G.E. Ball for constructive criticism and discussion during the writing of this review. B.S. Heming Department of Entomology University of Alberta Edmonton, Alberta T6G 2E3 GEOGRAPHIC VARIATION, DISTRIBUTION AND TAXONOMIC STATUS OF THE INTERTIDAL INSECT THALASSOTRECHUS BARBARAE (HORN) (COLEOPTERA: CARABIDAE) WILLIAM G. EVANS Department of Entomology University of Alberta Edmonton, Alberta T6G 2E3 Quaestiones Entomologicae 13: 83-90 1977 Populations of Thalassotrechus barbarae (Horn), a nocturnal, flightless, intertidal carabid beetle species, are distributed linearly from Point St. George ( Crescent City), California to Bah- ia Magdalena, Baja California, Mexico. Mean elytral lengths vary clinally from 2.68 mm in the extreme southern part of the range to 3.54 mm in the northern part. Elytral color also varies clinally with northern populations being darker. These data are not consistent with recogni- tion of subspecies. Consequently, T. b. barbarae (Horn, 1892; type locality - Santa Barbara, California), and T. b. nigripennis (Van Dyke, 1918; type locality - Moss Beach, San Mateo County, California) are consubspecific, and their names are synonyms. North-South temper- ature and rainfall gradients may be implicated in selection of local variants along the linear dine of elytral length and color. Les populations de Thalassotrechus barbarae (Horn), une especes de carabique nocturne, et apte're se trouvent dans la zone des maries et se rencontrent de la pointe St. George (Crescent City), Californie jusqu’a Bahia Magdalena, Basse Calif ornie, Mex- ique. La longeur moyenne des ilytres varie graduellement a partir de 2.68 mm dans V extreme sud de leur distribution jusqu’a 3.54 mm dans le nord. La couleur des ilytres varie de la mime facon avec les populations les plus foncies dans le nord. Ces donnies ne sont pas compatibles avec notre concept de la sous-espece. Done T.b. barbarae (Horn, 1892; localite type - Santa Barbara, Californie) et T. b. nigripennis (Van Dyke, 1918; localiti type - Moss Beach, San Mateo County, Californie) ne repri- sentent qu’une seule sous-espece, et leurs noms deviennent synonymes. Le changement graduel de la tempirature et de la pri- cipitation entre le nord et le sud pourrait expliquer la silection de variations locales en fuonction de changement liniaire de la couleur et de la longeur des elytres. On most coastlines of the world certain insects are conspicuous on rocky shores and sandy beaches, particularly in those areas characterized by extensive beds of offshore kelp and by diverse intertidal algal communities. During daytime low tide in such places, flies are very evi- dent among cast-up algal masses (wrack) and occasionally swarms of beetles (staphylinids and hydrophilids) are encountered (Leech, 1971); during periods of low tide at night predatory and scavenging beetles are active on sand or rock surfaces (Evans, 1977). However, the major- ity of intertidal insects, particularly their larval stages, are not easily observed since they inha- bit subsurface or rock crevice habitats or algal vegetation. In general, similar kinds of intertidal insects occur in similar habitats all over the world. Kelp flies of the families Anthomyidae and Coelopidae, for instance, are found wherever wrack ac- cumulates and an extensive beetle fauna is also associated with this food base either as preda- tors or scavengers. Some beetles eat other insects or small crustaceans stranded by receding tides, and members of Thalassotrechus barbarae (Horn) are such predators. Adults and larvae of this species live in cracks in rocks in the intertidal zone; adults are brachypterous, and there- fore flightless. Larvae are restricted to crack habitats but adults emerge at night and walk over the rock surfaces; feeding and mating takes place at this time. INTRODUCTION 84 Evans Ecological equivalents of this species are undoubtedly found in other parts of the world and probably include carabid beetles of the tribe Trechini such as Kenodactylus audouini (Guerin) I occurring in the subantarctic islands of New Zealand (the Antipodes Islands, Auckland Islands, j Campbell Island, Snares Island, Stewart Island), the Falkland Islands and Patagonia (Darlington, ! 1964; Johns, 1974), the European Aepus marinus (Strom) and Aepopsis robinii (Laboulbene) (Glynne-Williams and Hobart, 1952) and Thalassobius testaceus Sober from Chile (Jeannel, 1926). S GEOGRAPHICAL DISTRIBUTION OF THALASSOTRECHUS BARBARAE (HORN) The geographic range of Thalassotrechus barbarae extends from Point St. George, (near Cre- scent City) Del Norte County, California to Punta Belcher, Bahia Magdalena, Baja California (Fig. 1), a straight-line distance of approximately 2400 km that spans 17° 10' of latitude and 12° 03' of longitude. Populations of this species occur on rocky shores or on rocky outcrops of sandy beaches along this range and conceivably south of Bahia Magdalena as far as Cabo San Lucas, some 185 km away at the tip of Baja California. A part of this range is in the mar- ine biogeographic zone (based on sea water surface temperature) called Cold-Temperate which , extends from Point Conception, California to Alaska (Abbott, 1966) while the remaining part coincides with the Warm-Temperate zone between Point Conception and Bahia Magdalena. So j the latter locality is most likely to be very close to the southern limit of the range but as far as I know no collections have been made between Bahia Magdalena and the southern tip of the peninsula. The northern limit of the range is more exactly defined because north of Crescent City, California, where collecting sites are more accessible than in Mexico, I (and presumably other collectors) have failed to find any specimens. TAXONOMIC CONSIDERATIONS AND GEOGRAPHICAL VARIATION Taxonomic history of Thalassotrechus barbarae T. barbarae , though trechine in general form and habits, has been placed in the Poginini on the basis of general structure and particularly on characters of the mouthparts (Van Dyke, 1918) 1 and genitalia (Darlington, 1938). Pogonines are halobiontic (Ball, 1968), that is, they inhabit alkali soils with high concentrations of sodium chloride, but T. barbarae is the only member of this predominantly Palaearctic group that is found in the rocky intertidal zone, whereas mem- bers of the Trechini are commonly found in this habitat. Adults and larvae of Trechus ovipen- nis Motschulsky, for instance, occur in crevices and under stones of the high intertidal zone on rocky shores from central California to southern Alaska (Evans, 1977; Lindroth, 1961) and the trechine species mentioned above have similar habits. Thus, T. barbarae is probably a relict gen- us derived from a stock that gave rise to the present day halobiontic pogonines (Ball, pers. comm.) but because it has become adapted to living in the rocky marine littoral it has convergently be- come similar in color and form to marine trechine beetles (Darlington, 1938). Horn (1892) described Trechus barbarae from specimens collected at Santa Barbara, Califor- na, the type locality. 1 Van Dyke (1918) transferred this species to the Pogonini, based on it the new genus Thalassotrechus, the name indicating resemblance of its members to members of Trechus. He also described a second species, T. nigripennis, from northern California (type locality - Moss Beach, San Mateo County; see Fig. 1), adults of which were larger and darker, and with proportionately broader prothoraces (Van Dyke probably judged this last feature, as taxonomists generally did at that time, rather than taking measurements). Apparently, however, 1. During 1969, specimens could not be found anywhere near Santa Barbara presumably because of the massive oil spill that occurred early in the year (Evans, 1970). Thalassotrechus barbarae (Horn) 85 he later came to regard T. barbarae and T. nigripennis as subspecies, a conclusion amplified by Darlington (1938) on the basis of comparison of adults from Santa Barbara with adults from the San Francisco area. The elytra of adults of the southern population examined by Darlington were rufotestaceous or castaneous in contrast to the black and dull-textured ely- tra of adults of the northern group. Also, the anterior prothoracic angles of adults of barbar- ae were less flattened than those of the northern population. Moore (1956) also agreed with this arrangement even though he found no consistent structural difference between series of both populations. Geographic variation of T. barbarae Since clinal variation is expected to occur in Thalassotrechus over the predominantly NW- SE trend of the coastline on which it is found I examined adult specimens loaned by the Cali- fornia Academy of Sciences as well as collected by myself. With the use of an ocular microme- ter, measurements of elytral length (base to apex along the suture) and maximum prothoracic width were made of individuals obtained from localities shown in Table 1 . These localities re- present collecting sites that vary in size from individual rocks (at Bahia Magdalena) to indivi- dual beaches (Pismo Beach) to long stretches of coastline (Del Norte, Humboldt and Mendoc- ino Counties). The data, shown in Fig. 1, clearly indicate that the mean elytral length of Thalassotrechus populations increases progressively in a north-westerly direction agreeing with Bergman’s Rule (Mayr, 1942) which states that the smallest sized individuals of a species are found in the sou- thern part of the range and the largest in the northern part. This clinal progression consists of consecutive, overlapping ranges of elytral length, and of standard deviations of elytral length (except for San Diego County and Bahia Magdalena) with the largest step occurring between Bahia Magdalena, Mexico and San Diego, California, a straight-line distance of approximately 1060 km. Because of this distance I would expect that further sampling between these two localities would give populations that would reduce this large step to a series of overlapping smaller ones. A population, then, from any locality would not differ significantly in size from an adjacent population. Prothoracic width/elytral length ratios do not differ significantly be- tween adjacent localities or between the most northerly and the most southerly localities (Table 1) indicating that the samples were drawn from a population of individuals of different sizes but not of different form. As shown in Fig. 1 , the rate of change of mean elytral length appears to be greater in the middle of the range than at the ends. Theoretically, this may be expected because there would be a decreasing tendency for selection of variants that reflect submarginal environmental con- ditions that presumably exist towards the ends of the range. Alternatively, the greater rate of change may be due to an equally greater rate of change of some environmental variable such as sea water temperature. An abrupt change such as this probably occurs in the middle of the range near Point Conception, California (see Fig. 1) a well-known locus of marine inshore fau- nal discontinuity (Abbott, 1966; Garth, 1955). Adult specimens were also examined for variation in elytral color. Five color categories could be distinguished and individuals were assigned to a category as accurately as possible by this subjective method. The proportions of the populations from each locality (not identical to the above localities because all the specimens were not available at the time of this analy- sis) in each of these color categories are shown in Fig. 2 where the suspected color cline is sat- isfactorily demonstrated. Elytral color progresses from pale testaceous in the extreme south to black in the northern part of the range. It is interesting to note, however, that color varies even at the ends of the range so that the elytra of individuals from Bahia Magdalena are not Quaest. Ent., 1977 13 (2) all pale testaceous and individuals from Point St. George do not all have black elytra. Fin- ally, color variation is apparently greater in the middle part where the elytral color of indi- viduals may vary from the lightest to the darkest. Because variation in supposed diagnostic features is either non-existent (ratio to prothora- cic width/elytral length) or clinal (elytral length and color), I conclude that recognition of subspecies would be wholly arbitrary, and would serve no useful purpose. Consequently, T. j b. barbarae and T. b. nigripennis are consubspecific, and their names are synonymous. The j species is thus monobasic, and a binomial name is sufficient. Discussion of Pattern of Variation of T. barbarae Clines result from gene flow between populations and from selection of individuals that are adjusted to the local environment (Mayr, 1963). Since T. barbarae is distributed along a northwest-southeast trending coastline, that is, linearly (or unidimensionally, Udvardy, 1969), environmental factors that show a north-south gradient are more likely to be implicated in the selection of local varients. One such factor, temperature, is an obvious example, and its role in influencing the body size or body proportions of poikilotherms has been demonstrated by Ray (1960). A north-south gradient for the California coastline is well-documented for air temper- ature (U.S. Department of Commerce, 1968) and sea water temperature (U.S. Department of Commerce, 1967) both of which must influence metabolism of T. barbarae but other factors such as salinity, rainfall, humidity, wave action, and availability and kind of food, are proba- bly also involved. Johns (1974) suggests that body size of marine carabids of the species Keno- dactylus audouini may be related to the amount of exposure to sea water since the largest spe- cimens of this species are in supralittoral habitats where sea water is much diluted and where prey such as Collembola and Isopoda, and larvae and eggs of various arthropods are more avai- lable. A rainfall gradient exists between the ends of the range of T. barbarae with Point St. Geo- rge (Crescent City), California having the highest average annual rainfall, about 178 cm (U.S. Department of Commerce, 1968) and Bahia Magdalena, Mexico receiving less than 25 cm a year (Escoto, 1964). The increasing dilution of sea water in a northerly direction may very well be implicated in the dine of body size in T. barbarae but perhaps the rainfall gradient, expressed as a humidity gradient, may be implicated in the clinal variation in elytral color. Discussing Gloger’s Rule, Dobzhansky (1937), concludes that pigmentation in insects increa- ses in humid and cool and decreases in dry and hot climates with humidity being more effect- ive than temperature. If, indeed, this is correct, the correlation between elytral color and hum- 1 idity.in T. barbarae could be explained but the underlying mechanism for this phenomenon is unknown. In all likelihood, several factors are involved in geographic variation of T. barbar- ae making it difficult to determine the actual cause of this variation even though the distribu- tion of this insect is linear, therefore, seemingly much more simple to analyze than two-or three-dimensional distributions (Udvardy, 1969). In the latter case, for example, a plot of a single character gradient such as color on a distribution map results in, at best, a series of non- parallel, crude isophenes (see Petersen, 1947). Or, when geographic variation in body length of a nonlinearly distributed insect is plotted on a Hubbs-Hubbs diagram, as Ball and Negre (1972) did for a xerocolous carabid, Calathus ruficollis Dejean, the resulting pattern can be very complex indeed. But this pattern merely reflects complex gradients of environmental var- iables. In order to elucidate relationships between biological and environmental variation pre- cise measurements of such parameters as temperature and rainfall or humidity are needed in addition to data on several character states of populations of linearly distributed species. This study suggests that such an approach is possible. Thalassotrechus barbarae (Horn) 87 ACKNOWLEDGEMENTS I am grateful to H.B. Leech and D.H. Kavanaugh for the loan of specimens. Bahia Magdal- ena specimens were collected by the author during Cruise 1 8 of Stanford Oceanographic Ex- peditions which was supported by NSF grants GB 6870 and GB 6871 to Stanford University. My thanks are extended to G.E. Ball for his encouragement and for his critical review of the manuscript and to J.S. Scott for drafting the figures. Table 1. Prothoracic width/elytral length ratios of populations of Thalassotrechus barbarae collected at various localities. Locality Code (see Fig. 1) Mean prothoracic width x 100/elytral length ratio S.D. CALIFORNIA Del Norte, Humboldt and Mendocino Co’s. 1 36.04 3.73 Marin Co. 2 36.15 4.04 San Francisco and Contra Costa Co’s. 3 35.12 2.27 Pacific Grove 4 35.44 1.42 Pismo Beach 5 34.4 2.7 Santa Barbara Co. 6 37.59 3.3 Los Angeles and Orange Co. 7 36.95 2.4 San Diego 8 37.8 1.19 MEXICO Punta Belcher, Bahia Magdalena 9 35.4 2.11 LITERATURE CITED Abbott, D.P. 1966. Factors influencing the zoogeographic affinities of the Galapagos inshore marine fauna, pp. 108-122. In Bowman, R.I. (editor), The Galapagos. University of Califor- nia Press. Berkeley, California xvii + 318 p. Ball, D.E. 1968. Carabidae, Fascicle 4, pp. 55-174. In Arnett, R.H. The beetles of the United States (a manual for identification). Catholic University of America Press, Washington, D.C. xi + 1 1 12 p. Ball, G.E. and J. Negre. 1972. The taxonomy of the nearctic species of the genus Calathus Bonelli (Coleoptera: Carabidae: Agonini). Transactions of the American Entomological Society, 98: 412-533. Darlington, P.J., Jr. 1938. The American Patrobini (Coleoptera, Carabidae). Entomologica Americana, 18: 135-187. Darlington, P.J., Jr. 1964. Insects of Campbell Island. Coleoptera: Carabidae. Pacific Insects Monograph 7: 335-339. Dobzhansky, T. 1937. Genetics and the origin of species. Columbia University Press, New York, xvi + 364 p. Quaest. Ent., 1977 13 (2) 88 Evans Escoto, J.A.V. 1964. Weather and climate of Mexico, pp. 187-215. In Wauchope, R. and R.C. West (editors). Handbook of Middle American Indians, Volume 1. Natural environment and early cultures, University of Texas Press, Austin, Texas. 570 p. Evans, W.G. 1970. Thalassotrechus barbarae (Horn) and the Santa Barbara oil spill. The Pan- Pacific Entomologist, 46: 233-237. Evans, W.G. 1977. Insects and relatives. In Morris, R.H. and D.P. Abbott (editors). Marine invertebrates of the California shore. Stanford University Press, Palto Alto, California (In Press). Garth, J.S. 1955. The case for a warm-temperate marine fauna on the west coast of North America, pp. 19-27. In Essays in the natural sciences in honor of Captain Allan Hancock. j On the occasion of his birthday, July 26, 1955. University of Southern California Press, Los Angeles, California xii + 345 p. Glynne- Williams, J. and J. Hobart. 1952. Studies on the crevice fauna of a selected shore in Anglesey. Proceedings of the Zoological Society of London, 122: 797-825. Horn, G.H. 1892. Random studies in North American Coleoptera. Transactions of the Ameri- can Entomological Society, 29: 40-48. Jeannel, R. 1926. Monographic des Trechinae. Morphologie comparee et distribution geogra- phique d’un groupe de Coleopteres. Part I. L’Abeille, Tome 32: 221-550. Johns, P.M. 1974. Southern New Zealand, Patagonian and Falkland Island Carabidae. Arth- ropoda of the Subantarctic Islands of New Zealand (I) Coleoptera: Carabidae. Journal of the Royal Society of New Zealand, 4: 283-302. Leech, H.B. 1971. Nearctic records of flights of Cafius and some related beetles at the sea- shore. (Coleoptera: Staphilinidae and Hydrophilidae). Wasman Journal of Biology, 29: 65- 70. Lindroth, C.H. 1961. The ground beetles (Carabidae, excl. Cicindelinae) of Canada and Alas- ka. Part 2. Opuscula Entomologica, Supplement 20: 1-200. Mayr, E. 1942. Systematics and the origin of species. Columbia University Press, New York, 334 p. Mayr, E. 1963. Animal species and evolution. The Belknap Press of Harvard University Press, Boston, Mass, vii + 797 p. Moore, I. 1956. Notes on some intertidal Coleoptera with descriptions of the early stages (Carabidae, Staphylinidae, Malachiidae). Transactions of the San Diego Society of Natural History, 12(11): 207-230. Petersen, B. 1947. Die geographische variation einiger Fennoskandischer Lepidopteren. Zoo- logiska bidrag fran Uppsala, 26: 330-531. Ray, C. 1960. The application of Bergmann’s and Allen’s rules to the poikilotherms. Journal of Morphology, 106: 85-108. Udvardy, M.D.F. 1969. Dynamic zoogeography. With special reference to land animals. Van Nostrand Reinhold Publishing Co. New York, xviii + 445 p. United States Department of Commerce. Environmental Science Services Administration, Coast and Geodetic Survey. 1967. Surface water temperature and density. Pacific Coast, North and South America and Pacific Ocean Islands. Publication 31-3, 2nd edition. United States Department of Commerce. Environmental Science Services Administration. Environmental Data Service. 1968. Climatological Data, California, Volume 72, No. 13. Vay Dyke, E.C. 1918. New inter-tidal rock-dwelling Coleoptera from California. Entomolo- gical News, 29: 303-308. Thalassotrechus barbarae (Horn) 89 Fig. 1. Mean (vertical line), standard deviation (black bar) and range (horizontal line) of elytral lengths of populations of Thalassotrechus barbarae collected from localities given in Table 1. A- Type locality for T. nigripennis. •- Type locality for T. barbarae. Quaest. Ent., 1977 13 (2) 90 Evans 68 55 12 ®00 0 0 0 0000'' / 125° / 120° / 115° Fig. 2. Proportions of populations of Thalassotrechus barbarae collected from various localities in five elytral color categor- ies. Black portion of circle represents the proportion of a population in that particular color category. Locations: 10. Men- docino, Humboldt and Del Norte Counties, Calif.; 11. Marin, Sonoma and Contra Costa Counties, Calif.; 12. Pacific Grove, Calif.; 13. Pismo Beach, Calif.; 14. Santa Barbara and Ventura Counties, Calif.; 15. Punta Belcher, Bahia Magdalena, Baja California, Mexico. MAMMALIAN-SIPHONAPTERAN ASSOCIATIONS, THE ENVIRONMENT, AND BIOGEOGRAPHY OF MAMMALS OF SOUTHWESTERN COLOMBIA EUSTORGIO MENDEZ Gorgas Memorial Laboratory P.O. Box 2016 Balboa Heights Panama Canal Zone Quaestiones Entomologicae 13: 91-182 1977 A synopsis of the fleas and their mammalian hosts in southwestern Colombia, with parti- cular reference to the Departamento del Valle is presented and information about the Siph- onaptera of Colombia is reviewed. Descriptions and illustrations of diagnostic features characterize the following new species o/Polygenis Jordan: P. caucensis (Type locality : Alto Anchicay a, Depto. del Valle, COLOM- BIA; type host: Oryzomys caliginosusj, P. delpontei (Type locality: Quebrada Honda near Pichinde, Municipio de Cali, Depto. del Valle, COLOMBIA; type host: Oryzomys caliginosus/ P. hopkinsi (Type locality: Pena del Cerro, Cerro Munchique, Depto. del Cauca, COLOMBIA; type host: Oryzomys albigularisA P. trapidoi (Type locality: Valle del Rio Pichinde, Munici- pio de Cali, Depto. del Valle, COLOMBIA; type host: Oryzomys caliginosusj. In addition, des- criptions are given of the male of Ctenidiosomus traubi Johnson, and the female o/Sphincto- sylla diomedes Johnson. The former species was previously known from the female, and the latter species was originally described from the male. The following taxa are reported for the Republic of Colombia for the first time: Plocopsylla phyllisae Johnson, Leptopsylla segnis (Schonherr), Dasypsyllus gallinulae peripinnatus (Baker), Tetrapsyllus comis Jordan, Polygen- is pradoi (Wagner), P. thurmani Johnson, P. klagesi samuelis (Jordan and Rothschild), and Pulex simulans ( Baker). The taxon Rhynchopsyllus megastigmata Traub and Gammons, 1958, is considered to be conspecific with Rhynchopsyllus pulex Haller, 1895. Records contained herein bring the total number of known Colombian Siphonaptera to at least 44 species and subspecies. Of this number, more than two-thirds are reported from the southwestern part of the country. A key to the fleas of southwestern Colombia, and illustrations of diagnostic features of most of the taxa are included. An account is given of the history and zoogeography of the mammalian fauna of south- western Colombia, as well as comments on host-parasite relationships. The history of the mam- malian fauna of South America was probably initiated at the beginning of the Tertiary. This early fauna included marsupials, certain edentates and primtive ungulates that came from North America. Subsequent isolation of North America from South America interrupted, but not completely stopped, an interchange of animals by island-hopping. The faunal interchange was re-established when the Isthmian land bridge appeared at the end of the Tertiary. It is also possible that during the Cretaceous and early Tertiary, South America received faunal elements from Africa and from Australia. The majority of families and genera of South American mammals originated during periods of total or incomplete isolation. Presently, in the Pacific Coastal Lowlands of Colombia ubi- quituous mammals outnumber endemic forms. This fauna has strong affinities with those of the Amazon Region and the coastal lands of Panama and Ecuador. The remaining taxa of the southwest portion of Colombia are more concentrated in the Andes and include some forms that probably originated in the lowlands. Evolution and radiation of the fleas of this territory probably was correlated to evolution 92 Mendez of their hosts. This phenomenon is more apparent in fleas parasitizing small mammals, such as cricetine rodents. The lowland flea fauna of the Pacific sector is particularly poor, both in numbers of taxa and endemism, while that of the Andean mountains displays more local ele- ments in addition to being significantly diversified. The Southwestern Colombia flea fauna ex- hibits little specificity and various species may parasitize the same host. On the other hand, there are related and unrelated host species that harbor the same flea taxon. It is evident that the flea fauna of the Colombian Pacific lowlands is related to those of Panama and the Amazon and Orinoco basins. The strong mammalian relationship existing throughout the Andean Cor- dillera in Colombia and Ecuador is reflected in the affinity observed in the flea fauna. Geographic aspects, such as topography, geology, soil, climate and vegetation concerned with the pertinent biota are discussed. Southwestern Colombia is characterized by a diversity of ecological situations resulting from a complex topography and the concomitant climatic regimes. The Pacific Coastal lands, and the Andean highlands, in addition to the Cauca Valley, represent the major geographic areas of this territory. Se presenta un sumario de la fauna de sifondpteros y sus mamiferos hospederos en la regidn suroccidental de Colombia, principalmente en el Departamento del Valle; al mismo tiempo, se revisa la informacidn existente sobre las pulgas de Colom- bia. Se incluyen descripciones e ilustraciones de aspectos diagndsticos que caraceterizan las siguientes especies nuevas del gdn- ero Polygenis Jordan: P. caucensis (Localidad tipo: Alto Anchicaya, Depto. del Valle, COLOMBIA; hospedero tipo: Oryzo- mys caliginosus^, P. delpontei (Localidad tipo: Quebrada Honda, cerca de Pichindd, Municipio de Cali, Depto. del Valle, COLOMBIA; hospedero tipo: Oryzomys caliginosus^, P. hopkinsi (Localidad tipo: Pena del Cerro, Cerro Munchique, Depto. del Cauca, COLOMBIA; hospedero tipo: Oryzomys albigularisy, P. trapidoi (Localidad tipo: Valle del Rio Pichindd, Munici- pio de Cali, Depto. del Valle, COLOMBIA: hospedero tipo: Oryzomys caliginosus/ Adema's, se describen en este trabajo el macho de Ctenidiosomus traubi Johnson y la hembra de Sphinctopsylla diomedes Johnson. De la primera especie unicamen- te se conicia la hembra, mientras que la segunda especie se describid originalmente del macho. Los siguientes taxa son citados por vez primera para la Repiiblica de Colombia: Plocopsylla phyllisae Johnson, Leptopsylla segnis (Schonherr), Dasypsyllus gallinulae perpinnatus (Baker), Tetrapsyllus comis Jordan, Polygenis pradoi (Wagner), P. thurmani Johnson, P. klagesi samue- lis (Jordan and Rothschild), and Pulex simulans (Baker). El taxon Rhynchopsyllus megastigmata Traub & Gammons, 1958, es considerado sindnimo de Rhynchopsyllus pulex Haller, 1895. Los registros contenidos en este trabajo permiten estimar que cerca de cuarenta y cuatro especies y subespecies de pulgas son conocidas de Colombia y que de este numero mas de dos tercios esta'n citadas para la parte suroccidental del pai's. Se incluye una clave para separar las pulgas del soroeste de Colombia y se ilustran aspectos diagndsticos de la mayoria de los taxa. Se expone un bosquejo de la historia y zoogeografi'a de la fauna de mamiferos de la parte suroccidental de Colombia, asi' como tambien comentarios sobre las relaciones entre pardsitos y hospederos. La historia de la fauna de mamiferos de Sur Amdrica probablemente se inicid al comienzo del peri'odo Terciario. Esta fauna primitiva consistid de marsupiales, cier- tos edentados y ungulados primitivos que vinieron de Norte Amdrica. El posterior aislamiento entre Norte Amdrica y Sur Amdrica logrd interrumpir pero no detener completamente un intercambio de animates mediante su traslado de isla a isla. El intercambio fauni'stico fue restablecido cuando aparecid el puente Istmeno al final del Terciario. Tambidn es posible que durante el peri'odo Cretacico y el comienzo del Terciario, Sur Amdrica recibid elementos fauni'sticos de Africa y de Australia. La mayoria de las familias y gdneros de mamiferos suramericanos se originaron durante periodos de parcial o total aisla- miento. En los tiempos actuates, en las tierras bajas de las costas colombianas del Pacifico aquellos mamiferos de amplia dis- tribucidn sobrepasan las formas enddmicas. Estas faunas tienen una fuerte afinidad con aquellas de la regidn amazdnica y de las tierras costeras de Panamd y Ecuador. Las restantes formas de la porcidn suroeste de Colombia estdn mds concentradas en los Andes y contienen algunos elementos que probablemente se originaron en las tierras bajas. La evolucidn y radiacidn de las pulgas de este territorio probablemente estaban correlacionadas con aquellas de sus hos- pederos. Este fendmeno parece ser mds aparente en las pulgas que parasitan mamiferos pequenos, tales como los roedores cricetinos. La fauna de pulgas del sector Pacifico es particularmente pobre, tanto en composicidn como en endemismo; mien- tras que aquella de las montanas Andinas muestra mds elementos locales ademds de estar apreciablemente diver sificada. La fauna de pulgas del suroeste de Colombia exhibe poca especificidad y varias especies pueden parasitar el mismo hospedero. Por otra parte, existen especies de hospederos relacionados y no relacionados que comparten el mismo taxon de pulga. Es evidente que la fauna de pulgas de las tierras bajas del Pacifico colombiano estdn relacionados con las de Panamd y las cuencas de los rios Amazona y Orinoco. La fuerte relacidn de los mamiferos exist entes a traves de las cordilleras Andinas en Colombia y Ecuador, se refleja en la afinidad que se observa en la fauna de pulgas. Se discuten tambidn en este trabajo aspectos geograficos tales como topografia, geologia, suelo, clima y vegetacion que conciernen al pertinente biota. El suroeste de Colombia se caracteriza por una diversidad de situaciones ecologicas que Mammalian-Siphonapteran Associations 93 resultan de una topografih compleja y los regi'menes climaticos concomitantes. Las tierras costeras del Pacifico y las elevadas tierras Andinas, adema's del Valle del Cauca, representan las principales areas geogrdficas de este territorio. CONTENTS Introduction 93 Physical Environment of the Departamento del Valle 94 Vegetation Formations of the Departamento del Valle 102 Historic and Zoogeographic Summary of the Mammals of Southwestern Colombia 106 Key to the Fleas of Southwestern Colombia 113 Taxonomic Treatment of the Fleas of Southwestern Colombia . 117 Analysis of Host-Parasite Relationships 165 Acknowledgements 177 References 178 INTRODUCTION The Departamento del Valle, one of several political divisions of the Republic of Colombia, comprises an area of about 20,430 km2, located in the southwest sector of that country. It contains contrasting ecological situations, including very wet rainforest of the Pacific lowlands, western and eastern slopes of the Cordillera Occidental, basin of the Rio Cauca in the rainsha- dow of Cordillera Occidental at 1000 meters, and the ascending western slope of the high Andean Cordillera Central. This territory is bordered on the north by the departamentos del Choco and Risaralda, on the south by the Departamento del Cauca, on the east by the departamentos del Tolima and Quindio, and on the west by the Pacific Ocean. The Departamento del Valle contains a great variety of plant and animal life, representing more than half of the total number of species of mammals known to occur in Colombia. There is little information about the fauna and the importance that many of these animals have as positive or negative elements in the health of humans and domesticated animals. This need for basic research in biology is indicative of the whole of Colombia as well as other areas of South America. During the past two decades, the Universidad del Valle, in conjunction with the Rockefeller Foundation and the Tulane University International Center for Medical Research, have contri- buted significantly to knowledge of arthropods and mammals associated with zoonoses in Col- ombia (Cali Virus Faboratory, 1965, 1968, 1970). Through the study of ectoparasites collect- ed by these research units, an interest developed for investigation of fleas and their mammalian hosts in the Departamento del Valle and other regions of Colombia. The importance of fleas in the epidemiology of plague, murine typhus and other diseases is well known, thus primary consideration of these medically important arthropods is warran- ted. In some instances, knowledge of the ectoparasitic fauna of an area may also provide un- derstanding of relationships among animal host groups (Clay, 1951 ; Hopkins, 1942; Patterson, 1957). Although plague, which remains a major concern, has not yet been found in Colombia, it is present in the adjacent countries of Ecuador, Venezuela, Peru and Brazil. There is, how- ever, a strong possibility that the disease could become established in Colombia, since ideal climatic factors occur in many areas of the country. In addition, both wild rodent species and those commensal with man are present; these have been implicated elsewhere either as actual or potential reservoirs of plague. Several species of fleas occurring in Colombia, such as Xeno- psylla cheopis, the classical vector of plague, as well as Pulex irritans and certain species of Polygenis, have been implicated in transmission of plague in other countries (Chabaud, 1947; Quaest. Ent., 1977 13 (2) 94 Mendez Macchiavello, 1948, 1954, 1958; Moll and O’Leary, 1945; Panamerican Health Organization, 1956; Pollitzer, 1954). Presently, the Siphonaptera fauna of Colombia is poorly known. In a survey prior to be- ginning this study, I found that Colombian material was not well represented in collections of several institutions, including the British Museum (Natural History) and the United States National Museum of Natural History. The literature contains few reports concerning Siphonaptera from Colombia. Dunn (1929) and Patiho-Camargo (1940) briefly mention some common species. Fuller (1942) and Macch- iavello (1948) gave several Colombian records of fleas. Costa Lima and Hathaway (1946) pre- sent information on several Colombian species. Gast Galvis (1950) in his list of fleas from Col- ombia considered 19 species and subspecies. Johnson (1954 and 1957) has most significantly contributed to our knowledge of the fleas of Colombia. In 1954, she described a new species of Pleochaetis from that country. Her outstanding monograph “Fleas of South America”, published in 1957, contains several new distributional records of species for Colombia and, in addition, the description of six new species. Mendez (1968) described a new genus and spe- cies of Colombian flea, and more recently, Mendez and Hanssen (1975) reported a new Col- ombian taxon discovered in the Departamento del Meta. Tamsitt and I. Fox (1970) recorded Rhynchopsyllus pulex Haller. This work is largely based on information obtained from collections made in the Departa- mento del Valle and neighboring political divisions of southwestern Colombia, such as Nar- ino, Cauca, Putumayo and Huila. In view of the ecological affinities of these territories and the similarity of their mammalian fauna, it is likely that these areas have virtually the same ectoparasitic fauna. The present account contains descriptions of four new species of Polygenis Jordan. The fe- male of Sphinctopsylla diomedes Johnson, known previously from the male, and the male of Ctenidiosomus traubi Johnson, are also described. A key to species of fleas known or presump- tively existing in the southwestern portion of Colombia, and new records for the country, are also presented. Rhynchopsyllus megastigmata Traub and Gammons is considered here to be a synonym of R. pulex Haller. Conventional figures for the majority of the species concerned are included. The nucleus of the material was from collections made by Harold Trapido while engaged in virus research sponsored by the Rockefeller Foundation. Additional specimens were collec- ted personally or obtained from the Universidad del Valle and other sources. PHYSICAL ENVIRONMENT OF THE DEPARTAMENTO DEL VALLE Topography Inasmuch as most of the siphonapteran material used in this study is from the Departamen- to del Valle, the following geographical discussion is limited to this territory. No other poli- tical division of Colombia displays such a diversity of geographic features and, consequently, ecological conditions. The topographic information given below is derived mainly from Es- pinal (1968), Sanchez (1965) and Sauer (1950). Generally, the physiographic and faunistic elements of this area are characteristic of the southwestern zone of Colombia, which consists of the Cauca Valley territory, surrounding mountains, and Pacific coastline within the limits of the Departamentos del Valle, Cauca, Huila, Narino, and Putumayo. The landscape is dominated by the Western and Central ranges of the Andean mountain system. These mountains contain some of the higher peaks of Colombia. The Western Cor- dillera extends north-northeastward parallel to Central Cordillera and is almost parallel to the Mammalian-Siphonapteran Associations 95 REPUBLIC OF COLOMBIA Fig. 1. Map of Colombia. Quaest. Ent., 1977 13 (2) 96 Mendez Pacific coast. It is the lowest of the three ranges forming the Andean system. The Central Cordillera in the eastern sector of El Valle contains high peaks such as the mountains of Huila and Barragan, which exceed 3000 meters. Several rivers and numerous streams form a Pacific watershed which originates in the moun- tains and empties into the Pacific Ocean. The more important rivers are the Anchicaya, Dagua, J Naya and Cauca. This last river is the largest and courses through the Departamento from south : to north. Mangrove swamps flooded by high tides are along the shoreline of the Pacific coasts. Warm tropical rain forests follow the swamps and occupy extensive inland zones of dense vegetation, which is impenetrable in many areas. The rich plant life, water, and cover, offer optimal con- ditions for an abundance of animal forms. The pocket-like arid valley of Dagua, however, with its grassy and brushy cover, introduces a contrasting zone in this lowland territory. A subtropical zone of humid temperate climate follows the lowland cloud forests. This tem- perate zone is characteristic of the middle slopes of the mountains, where vegetation is rich but not very luxuriant. Many of the faunal elements of these subtropical forests are similar to those found in lowlands, since many species evolved from ancestors which moved to the upper zones. Apparently climate and other geographical conditions have been limiting factors in es- tablishment of some species that have more restricted tolerances. The Cauca Valley, an agriculture area of the temperate zone, is a narrow isolated region con- sisting of approximately 400,000 hectares between the Western and Central Cordillera. It is nearly 160 kilometers long and only 12 kilometers wide, and is drained by the Cauca River. Although much of its territory is occupied by pasture, mainly Para ( Panicum barbinode ) and Guinea {P. maximum ) grasses, the Cauca Valley represents the most fertile and productive agricultural land of Colombia. Lush montane cloud forests occupy extensive zones of moderate and high elevations (from below 1000 to about 3000 meters) throughout most of the Central and Western Cordilleras. The uppermost reaches of the mountains, from over 3000 meters, are largely unforested, con- sisting primarily of extensive grass plains characteristic of paramos (such as Las Hermosas, Chinche, Miraflores and Barragan). These areas support more selected types of plants and ani- mals; indeed the paucity of animal life in the paramos is directly correlated to the poor diver- sity of cover. As in other areas of Colombia, the Departamento del Valle is suffering from much defores- tation, due primarily to agricultural development, hydroelectric projects and raising of cattle. Geology By virtue of its rock composition, Colombia comprises two geological regions: the Oriental plains; and the Andean geosynclinal regions (Biirgl, 1961). Only the latter region is in south- western Colombia, including the Departamento del Valle. The vast Oriental plains are in east and southeastern Colombia. The Andean geological region was apparently submerged during long periods from the be- ginning of the Cretaceous, and accumulated large deposits of marine, continental and volcan- ic sediment. The complicated tectonic movements experienced by these lands were responsi- ble for formation of the present Andean Mountains, which constitute the “backbone” of Colombia. The Andean geosynclinal region was consolidated before the Cambrian period, and includes the following mountain systems: 1. Central Cordillera, 2. Western Cordillera and Coastal Cor- dillera (Serrania de Baudo), and 3. Eastern Cordillera. The Lower and Middle Magdalena Val- ley basins separate the Central and Eastern Cordillera. The Lower Valley contains non-marine Mammalian-Siphonapteran Associations 97 Tertiary rocks, while, the Tertiary reservoirs of the Middle Valley are non-marine. Western and Central Cordilleras are separated by the valleys of the Upper Cauca and Upper Patia rivers. Other mountains of Colombia, such as the Santa Marta Mountains, Perija Mountains and the Pacific Coast Range, have affinities with the Andean system. The coastal zone of the Andean Region is represented by the Bolivar Geosyncline, a lowland area of Tertiary Marine formation, west of the Andean mountains and extending from South- ern Ecuador to the Gulf of Uraba in northern Colombia. This strip of land has been interpre- ted as a seaway which apparently permitted movement of terrestrial animals during periods ranging from upper Cretaceous times to Recent (Nygren, 1930; Hershkovitz, 1966). No marine formations from the Upper Cretaceous have been found in this zone. The Eastern Cordillera displays abundant deposits of Cretaceous and Tertiary rocks, with little or no recent volcanic elements. In addition, Jurassic, Triassic and Paleozoic rocks are found in these mountains. Western and Central Cordilleras, as well as the Pacific Coast Range, are primarily formed of igneous and metamorphic rocks, having only subordinate sedimentary beds. Each Cordillera has several high volcanic peaks. The Precambrian sedimentary history of Colombia is not known (Jacobs, Biirgl and Conley, 1963). According to these authors, during a great part of Cambrian and Ordovician time, the actual territory of the Eastern and Central Cordilleras, and at least the western aspect of the Llanos and Putumayo-Caqueta lowlands, were occupied by seas. Later, the area came under the influence of volcanic and diastrophic activity, which destroyed the Cambrian-Ordovician marine deposits. Rock shields, characteristic of the Andean system during the Cambrian, ap- parently were greatly disturbed by erosion and their detritus filled marine and terrestrial de- pressions of the region. The most important fossils from the lower Paleozoic in Colombia are brachiopods, trilobites and graptolites. The territory west of Central Cordillera shows no in- dication of Paleozoic marine sediment. Some Middle Ordovician fossils have been found in Colombia. However, Upper Ordovician and Silurian apparently are not represented by fossiliferous layers in this country. Fossils from the Devonian of Colombia are represented by brachiopods, bryozoans and tri- lobites, among other invertebrates. Many plant and animal fossils, primarily marine forms, are known from the Carboniferous. Evidence obtained from fossils indicates that the lower Carboniferous sea invaded the eas- tern part of the Andean Region, which was covered by sediment thereafter. During the upper Carboniferous, the sea gradually retreated and large semi-swampy forests moved to sections previously occupied by water. Abundant Permian fossils have been found in only a few areas of Colombia, primarily at Serrania de Perija. They represent sponges, crinoids, brachiopods, gasteropods, cephalopods, and other marine animals. The eastern portion of the Central Cor- dillera contains Marine Upper Triassic rock (the Payande Formation), and represents a com- bination of sandstone, limestone and shale. The Eastern and Central Cordilleras contain low- er Jurassic sedimentary rocks, principally of continental origin. Evidently much igneous activity occurred in the Andean Region during the Mesozoic. Also, it is apparent that the major part of western Colombia was occupied primarily by seas during most of Late Jurassic and Cretaceous time. Fossils from these periods are scant and consist mainly of ammonites and other molluscs. The Cretaceous Colombian and Peruvian faunas dis- played a strong relationship with those of Southern Europe (Olsson, 1956). Considerable tec- tonic movements that occurred during Late Cretaceous and early Tertiary, produced constant changes in the landscape of this region and in the formation of marine and non marine depo- sits. The Tertiary Continental deposits contain an excellent vertebrate fauna, including num- erous mammals. There is also evidence of considerable faulting and folding during the late Paleocene. Quaest. Ent., 1977 13 (2) 98 Mendez Throughout the Eocene, land forms of this area were somewhat different from the present. However, the principal elements of the Andes and the Pacific Coast basin originated during this period, and Weeks (1947) points out that the Bolivar Geosyncline introduced important chan- ges in Western Colombia and Ecuador. Much tectonic and volcanic activity took place during the Eocene and Oligocene (Hammen, 1961; Vuilleumier, 1971). Very few fossil vertebrates of Eocene age are known from Colombia (Stirton, 1953). Layers corresponding to the Eocene and Oligocene are richer in Foraminifera, molluscs and other marine fossils. The Cauca Valley is particularly rich in Oligocene marine rocks, mainly consisting of algal, and foraminiferal lime- stone. This area also has coal-bearing Tertiary jocks, that probably originated during the Mio- cene or before. A large number of fossil vertebrates has been discovered in Miocene deposits. The fossil mam- mal fauna of this period is very interesting and contains some relics of families believed to have disappeared in earlier times. Numerous findings of mammalian Pleistocene fossils have taken place in Colombia (Stirton, 1953; Patterson and Pascual, 1968). Many of these fossils represent species that became extinct at the end of the Pleistocene. Jacobs, Biirgl and Conley (1963) con- sider that the most recent stage of tectonic movement and volcanic activity was initiated in the early Miocene and continued into Recent time, particularly affecting the Central Cordil- lera and the southernmost part of the Western Cordillera. Some of the volcanoes existing to- day are still active. Soils The following considerations of soil distribution in the Departamento del Valle are based primarily on two major sources: the FAO/UNESCO Soil Map of South America (1961), and the account by Beek and Bramao (1968). A great deal of work has been done in South America to determine distribution of major soils as important patterns in development of agricultural zones and exploitation of minerals and other natural resources significant to the general economic progress of the continent. How- ever, despite the knowledge that has been accumulated during many years, the study of struc- ture, composition, and distribution of South American soils is far from complete. The nature of the soils of the Departamento del Valle is linked to patterns of ecological factors that govern the life zones, such as climate, geomorphology, topography, and vegeta- tion. Departamento del Valle, and other areas of southwestern Colombia, from which mater- ial used in the present study has been collected, consist of Lowlands and Andes soils, two of the major structural elements of soil distribution that have been established for South Ameri- ca. Uplands soils, the other element considered in this segregation, are found in the eastern part of South America. Criteria assumed for establishment of these general categories are based on a complex association of factors such as geography, climate, vegetation, physiography, etc. The Pacific Coastal Lowlands form a high portion of the western side of the Departamento and are primarily characterized by contiguous areas of tropical evergreen and deciduous for- ests. The alluvial soils predominating in this territory are stratified with little organic matter and little profile development. The Pacific littoral is particularly dominated by dense mangro- ve forests, interspersed with swamp forests. This zone contains Quaternary marine and fluvio- marine deposits with alluvial plains and terraces, estuarine and delta alluvial deposits, and lo- cal areas of coastal sand dunes. The Northern Andes is the other soil region of the Departamento del Valle. The soil com- position is more diverse and the profile of the area interrupted by western and central ranges of the Andean mountains. These are separated by the Cauca Valley, a strip of land which also includes areas of the Departamento de Cauca, Narino, Caldas and Antioquia. RAIN SHADOW IN TRANSECT OF DEPARTAMENTO DEL VALLE Mammalian-Siphonapteran Associations 99 Quaest. Ent, 1977 13 (2) Fig. 2. Rainshadow in transect of Departamento del Valle. 100 Mendez In the western sector of the Northern Andes region of the Departamento del Valle, volcan- ic rocks are the most important elements of the soil structure. This condition reflects the vol- canic origin of these mountains. Most soils in areas of moderate elevation in the northern An- des are Laterosols, derived from volcanic material. Laterosols are dark colored surface soil with lighter subsoil, and, in addition to being slightly acid, contain a high amount of organic matter. Because of their low fertility level, they have limited use for farming. Extensive areas of the northern Andes are represented by Andosols. Such soils consist primarily of volcanic ash with dark surfaces, combined with organic matter and minerals such as nitrogen, phosphorus, cal- cium and potassium. Those that are not too acid are relatively fertile and excellent for grow- ing different crops. Sectors of the Departamento del Valle, dominated by Andosols, have ex- cellent pasture lands and areas devoted to agriculture. Dark paramo soils, which may be derived from heavy clays, perhaps of glacial origin are found adjacent to Andosols. They consist of some volcanic ash and are characterized by a high degree of acidity and paucity of nutrients. The paramo lands have high humidity, low temper- ature, the vegetation is poorly diversified and consists of pastures and forests of secondary growth. The northern part of the Departamento del Valle is characterized by Reddish Brown Lat- eritics. These soils contain dark, reddish brown, granular clay surface soil, with yellowish-brown, clay subsoil. Aluminum silicate, its principal mineral compound, is mixed with iron and other ingredients of inorganic and organic origin. They occur below 2000 meters on which are pri- mary and secondary forests in addition to pastures, coffee plantations and other cultivated areas. Climate The complexity of its geography contributes to the variety of climates existing in the De- partamento del Valle, since these elements are intimately associated. It is interesting to note that all four of the climatic regions outlined by Eidt (1969) for South America, are represen- ted in the Departamento del Valle, namely, tropical rain, temperate, arid and tundra. Tropical rain climates are confined to the Pacific territory, from sea level to the mountain bases reaching an elevation of close to 1 ,000 meters. These areas are hot and humid, with tem- perature ranges from 24°C to 30°C. The annual rainfall is heavy, exceeding 760 cm a year. It rains almost daily, thus this area does not have a true dry season. Humid warm winds of the Pacific Ocean and Andean chain contribute to the heavy precipitation; Figure 3, adapted from Espinal (1968), illustrates the rainshadow influence. According to Espinal (1968), with- in the Western and Central Cordillera there are two rainy periods during the year: the first from April to June, and the second from September to November. Between these wet sea- sons there is notably decreased precipitation. Temperate climates are characteristic of the subtropical areas extending from 1,000 m to 2,000 m, which have a temperature range of 18°C to 24°C. These areas are the first table lands above the lowlands, and are influenced by higher masses of the Andes and by action of two kinds of winds, interpreted as mountain-valley breezes and land-sea breezes. These winds af- fect climate and precipitation of the Cauca Valley system, and rivers, and other subtropical areas of this territory. The eastern part of the Departamento maintains a cooler climate typical of areas above 2,000 meters, (in which various paramos are found). The northern sector of the Central Cor- dillera contains an arid zone, subjected to strong winds. This zone contrasts with the cloudy and rainy forests dominating the Andean landscape. Some of the higher peaks of Central Cor- dillera are cool and damp paramos where the temperature is below 12°C. Mammalian-Siphonapteran Associations 101 SOIL MAP OF THE DEPARTAMENTO DEL VALLE Fig. 3. Soil map of the Departamento del Valle. Quaest. Ent., 1977 13 (2) 102 Mendez VEGETATIONAL FORMATIONS OF THE DEPARTAMENTO DEL VALLE Most of the information presented below, including Fig. 4, is based on Espinal (1968), and Espinal and Montenegro (1963). These studies were made according to the classification esta- blished by Holdridge (1947), which considers aspects of temperature and humidity in analysis of life zones. Ecological parameters of rodents and other mammals, as well as of the ectoparasites affect- ing them are highly dependent on biotic conditions, soils, climates and other factors. Vegeta- tional formations are therefore of primary importance in understanding habitat requirements and distribution patterns of these animals. However, such formations in time and space usu- ally grade into one another and therefore are not permanent. In addition, many animals and plants are able to tolerate a moderate range of climatic conditions. A brief description of all of the vegetational zones outlined for the Departamento del Valle follows. Tropical Very Dry Forest This formation is in two areas. One is Loboguerrero, at the upper part of Rio Dagua in the center of the Departamento. The other region represents a flat belt of forest platform in the Central Cordillera, extended from Cali to near San Francisco at the left shore of the Cauca River, reaching an elevation of 1200 to 1400 m. The prevalent climate of this zone is dry and the mean temperature is over 24°C. Mean rainfall during the year ranges from 500 to 1000 mm. Characteristic plants of this zone are figs {Ficus spp.), Vachelia farnesiana, spurge {Euphorbia caracasana ), Pithcellobium dulce, Desmanthus virgatus, Achatocarpus nigricans, mesquite {Pro- sopis juliflora ), Jatropha go ssypii folia, Fugatera pterota, Ocimum micranthum, Heliotropium sp., Lantana canescens, Citharexylum sp., Portulaca pilosa and Talinum paniculatum. Tropical Dry Forest It is distributed along the Central Valley crossed by the Cauca River and along a strip of land surrounding the Tropical Very Dry Forest of Loboguerrero. Mean temperature of the area is over 24°C and rainfall fluctuates between 1000 and 2000 mm. The characteristic vege- tation of this zone grows in lands not over 1000 m high and consists of cabuyas {Furcraea sp.), Croton sp., Turnera almifolia, Cephalocereus colombianus and other plants. Tropical Moist Forest This zone occurs in three areas below 1 000 m in the upper half of the Departamento. They are the canyon of Rio Garrapatas near El Cairo and Versalles, a strip near Rio Dagua and pro- bably an area in the mid portion of Rio Calima. The mean temperature is higher than 24°C and the mean rainfall is from 2000 to 4000 mm. Some plants found in this type of forest are the balsa {Ochroma lagopus ), roble {Tabebuia pentaphylla ), and hogplum {Spondias mombiri). Tropical Very Wet Forest This formation extends from the basin of Rio Anchicaya as a belt that runs from south to north along the Western Cordillera. The area is dominated by a variety of plants such as ceiba {Ceiba pentandra), balsa {Ochroma lagopus), black rubber {Castilla elastica), guarumos {Cec- ropia spp.), calabash {Crescentia cujete), cedars {Cedrela spp.), annato {Bixa orellana ), sand- box {Hura crepitans), and others. These lands are not above 1000 m. The temperature exceeds 24°C and there is a mean rainfall of 4000 to 8000 mm per year. Mammalian-Siphonapteran Associations 103 VEGETATIONAL FORMATIONS TROPICAL VERY DRY FOREST tropical dry forest III TROPICAL MOIST FOREST TROPICAL VERY WET FOREST [ | TROPICAL RAIN FOREST | SUBTROPICAL DRY FOREST jT' : F 1 SUBTROPICAL MOIST FOREST j;- SUBTROPICAL VERY WET FOREST j ' "-"j SUBTROPICAL RAIN FOREST LOWER MONTANE DRY FOREST jjfeSI LOWER MONTANE MOIST FOREST h I LOWER MONTANE VERY WET FOREST !££££§ LOWER MONTANE RAIN FOREST MONTANE VERY WET FOREST MONTANE RAIN FOREST PACI F OCE A DEPARTAMENTO DEL VALLE Fig. 4. Vegetational formations of Departamento del Valle. Quaest. Ent., 1977 13 (2) 104 Mendez Tropical Rain Forest This is the largest of the vegetational formations of the Departamento, extending from the very wet lowland coast line to the margins of the Western Cordillera. Dense mangrove forests consist primarily of red mangrove ( Rhizophora brevistyla) in association with black mangrove (. Avicennia marina ), white mangrove ( Laguncularia racemosa), and buttonwood ( Conocarpus ). Inland forests, beyond the Pacific littoral, contain a diversity of plants such as guarumo ( Cec - ropia sp.),wild figs (Ficus), mammagua (Inophleum armatum), membrillo ( Gustavia superba), Acacia melanoceras, sensitive-plant (Mimosa), cashew (Anacardium excelsum), balsa (O chroma spp.), and many others. The tropical rain forests are below 1000 m, in a climate with mean temperature above 24°C and mean annual rainfall exceeding 8000 mm. Subtropical Dry Forest This includes isolated areas on the eastern half of the Departamento. The major area is along the oriental baseline of the Western Cordillera from Cali to San Francisco. One patch is in El Dovio’s ravine and two other patches occur in the Central Cordillera. Typical plants in this zone are figs (Ficus spp.), fiques (Fourcraea sp.), mosquero (Croton sp.), big belly tree (Wigandia caracasana), indigo (Indigobera sp.), cuipo (Cavanillesia platanifolia), among others. It registers a mean temperature of 17°C to 24°C and the mean annual rainfall is between 500 to 1000 mm. Subtropical Very Humid Forest This formation encompasses very humid areas of the Western and Central Cordillera having an elevation between 1 100 to 1900 m. The mean temperature is from 17°C to 24°C and the annual rainfall fluctuates between 2000 to 4000 m. The vegetation of this zone is represented by guamo (Inga spp.), guayacan (Tabebuia chrysantha), balsa (Ochroma lagopus), guarumo (Cec- ropia), yaragua (Melinis minulti flora), pigs fern (Pteridium aquilinum), fox tail (Andropogon sp.), and others. Subtropical Moist Forest It is distributed as an extensive area surrounding the dry Rio Cauca valley and extending to the edges in the Western and Central cordilleras. Other patches of this zone are located in areas surrounding El Dovio and in the upper portion of the rivers Dagua, El Carmen and El Treinta. The zone reaches an altitude between 1 100 to 2000 m. The mean temperature ranges from 17°C to 24°C, while the annual rainfall is between 1000 to 2000 mm. Typical plants of this zone are the macedero (Trichantera gigantea), surrumbo (Trema micrantha), cordoncillo (Piper adunCun), balsa (Ochroma lagopus), iraca (Carludovica palmata), trompet (Bocconia frutescens), coralito (Hamelia patens), rubber (Ficus sp.), chayote (Sechium edule) and many others. Subtropical Rain Forest It is particularly represented by a strip of watered land of the Western Cordillera with ele- vations between 900 to 1900 m. The humidity in this vegetation formation is very high and the temperature reaches from 17°C to 24°C. The annual rainfall exceeds 4000 mm. This zone combines some virgin forests as well as agriculture lands. Some of its characteristic plants are platanillos (Heliconia), ferns (Gleicheniaceas), guarumo ( Cecropia ), rubber (Castilla elastica), paco (Cespedesia macrophylla), pejibaye (Guilielma gasipals), and avocado (Persea americana). Mammalian-Siphonapteran Associations 105 Lower Montane Dry Forest This formation is limited to an area of high elevations (from 2000 to 3000 m) of Barragan in the Central Cordillera. The region possesses a dry climate with a mean temperature of 1 2°C to 17°C, and an annual rainfall of 500 to 1000 mm. The native vegetation is poor and consists primarily of capers ( Cassia sp.) and mosqueros ( Croton sp.); cultivated species include maize ( Zea mays), onion {Allium cepa), wheat {Triticum), and potatoes {Solanum tuberosum). Lower Montane Moist Forest This region includes two areas in the Central Cordillera, where the altitude is from 1 800 to 3000 m. It is a cold formation, humidity is high and mean temperature varies from 12°C to 17°C. Annual rainfall is over 4000 mm. Some of the plants found in this territory are the following: carbonero {Befaria sp.), chilco {Baccharis sp.), capers {Cassia sp.), and mortino {Miconia albi- cans). Lower Montane Very Wet Forest This formation occupies extensive subtropical lands of the Western and Central Cordillera at elevations between 1800 and 3000 m. The cool climate registers a mean temperature of 12°C to 17°C. Annual precipitation is calculated to be between 2000 to 4000 mm per year. These forests contain many different plants such as guarumos {Cecropia spp.), lichens {Cora pavonia, Cetraria sp.), mosses {Polytricum sp.), horse tail {Equisetum sp.), encimo {Weinmannia balbi- siana), cascarillo {Ladenbergia), etc. Lower Montane Rain Forest This is a wet formation distributed as occasional areas in the Western and Central Cordillera. The elevation ranges from 1800 to 2900 m and the mean temperature varies from 12°C to 1 7°C. The annual rainfall registers over 4000 mm. Typical plants of this zone are the white guarumo {Cecropia), capers {Cassia sp.), quassia {Quassia sp.), cherry {Freziera sericea), ced- rillo {Brunellia sp.), lulo {Solanum quitoense), horse tail {Equisetum bogotense), berries {Ru- bus sp.), strawberries {Fragaria sp.), and others. Montane Very Wet Forest This formation occupies a broad portion of the typical paramo distributed from Santa Lucia to Barragan in the Central Cordillera. It has an elevation above 3000 m and a mean temperature from 6°C to 12°C. The annual rainfall fluctuates between 1000 to 2000 mm. The plant-life is represented by frailejones {Espeletia), capers {Cassia sp.), espadero {Rapanea sp.), charcoal mak- er {Befaria sp.), morteno {Hesperomeles sp.), grasses such as Calamagrotis and Festuca, etc. Montane Rain Forest These areas are very humid, over 3000 m in altitude, localized in the Western and Central Cordillera. Mean temperature is between 6°C and 12°C and rainfall is over 2000 mm. This type of vegetation formation contains cultivated territories as well as undisturbed forests. Among the plants found in these lands are frailejones {Espeletia sp.), blackberries {Rubus sp.), encen- illo {Weinmannia sp.), Senecio sp., Vaccinium sp., cherry-tree {Prunus sp.) and Bacharis sp. Quaest. Ent., 1977 13 (2) 106 Mendez HISTORIC AND ZOOGEOGRAPHIC SUMMARY OF THE MAMMALS OF SOUTHWESTERN COLOMBIA The ecological parameters of mammals and their ectoparasites in Colombia and other South American countries is of interest to investigators concerned with a variety of zoonoses linked to those animals. However, little has been reported on their biology and distribution. Even sys- tematic studies of these vertebrates are not complete and names and taxonomic status of a number of forms are not settled. To understand the origin and distribution of mammals presently occurring in southwest Colombia one must read an array of discussions by authorities on South American mammals. To fit the limited scope contemplated here, I have tried to be selective in interpretation of possible events that, in my opinion, have a logical foundation. References concerning evolu- tion and zoogeography of South American mammals are Hershkovitz (1962, 1966, 1969, 1972); Keast (1968); Loomis (1914); Patterson and Pascual (1963, 1968); Savage (1974); Scott (1937); Simpson (1940, 1943, 1945, 1950, 1969); Stirton (1950). The theory of plate tectonics has introduced a new argument for exploring the origin and distribution of Latin American fauna and flora (Raven and Axelrod, 1975; Valentine and Moores, 1974). According to this theory, movement of land masses or plates constituting the earth’s crust produces cataclysms and volcanic action with subsequent complicated modifica- tions of the elements involved. As applied to the southern continents, and particularly to South America, some intriguing perspectives occur with regard to the origin and affinities of the biota. In view of the geological facts, during the Middle Cretaceous South America was connected with Africa, and via Antarctica, with Madagascar, India and Australia. It was sep- arated from those eastern lands over an extended period of time. Apparently the connection with Australia lasted longer. It has been speculated on this basis that an exchange of fauna, at least in part by island-hopping, took place between Australia and South America until early Oligocene. On the other hand, a more effective connection between South America and Afri- ca existed until the end of the Cretaceous. Probably during late Eocene, interchange of tropical and warm-temperate animals between South America and Africa was greater than between Africa and North America. Some groups of animals, such as the river turtles, family Pelomedusidae, which occur in South America, Madagascar and Africa, also seem to afford a logical basis for this assumption. It is also belie- ved that routes for the movement of cool-temperate organisms between southern South Am- erica and Australia existed until nearly 4 million years ago. The argument appears to be sub- stantiated by certain elements of the flora of angiosperms common in Africa and South Am- erica. Events of a different nature, primarily geological and climatic, occurring in the millions of years since separation of the southern continents during the Cretaceous, have interrupted fau- nal development and geographical ranges of taxa in South America. This has introduced many gaps that obscure the entire historical sequence. Geological changes during the Cenozoic, are very important in attempting to understand some of the general history of the mammals of the Departamento del Valle, and other parts of western Colombia. The mammalian faunal structure of South American began to evolve with primitive forms recorded from the Paleocene, such as marsupials, members of the orders Condylarthra, Not- ungulata, Litopterna and other ungulates (Hershkovitz, 1968; Patterson and Pascual, 1968; Simpson, 1950). Apparently during the beginning of the Tertiary, the Middle American bridge permitted mammals to move to and from North and South America. The many fossil discoveries from Mammalian-Siphonapteran Associations 107 southern South America have revealed that, early in the Paleocene, the most important nuclei of South American mammals were concentrated in this portion of the continent. The earliest mammal records from northern South America and Middle America are represented only by marsupials and condylarths. During the Eocene and Oligocene epochs, and presumably during most of Cenozoic, North America was isolated from South America by a sea barrier since the connection represented at present by the Isthmian link did not exist. Fossils of monkeys and caviomorph rodents have been found in Oligocene beds of South America. However, the evolution and dispersal of these mammals prior to this time remain a mystery. Hershkovitz (1968) and Simpson (1943, 1950) believe that these mammals invaded South America probably as island-hoppers in the early Oligocene. According to Raven and Axelrod (1975), it is more probable that primates and cav- iomorph rodents reached South America and Africa during the Oligocene. It is also probable that the ancestors of those mammals arrived in this Southern continent during the Cretaceous. However, fossils from this epoch that would support this theory have not been found in South America. From the fossil history, it is evident that armadillos, ant-eaters and ground sloths inhabited South America since the Tertiary and were probably distributed in areas of mild climate, at least close to our present tropical conditions. It is possible that they originated from a Creta- ceous stock. According to Simpson and other authors, they gradually moved into South Am- erica when the Middle American land bridge was re-established in the late Pliocene. There is no indication that these mammals migrated between South and North America before this time, with the probable exception of the ancient armadillos. Fossil armadillos have been dis- covered in early Tertiary beds of North America (Kiirten, 1969). Hershkovitz (1972) discus- sed the probability that migration of terrestrial animals started during the period of isolation of the continents. Other authors (Scott, 1937; Simpson, 1940, among them) maintain that during the period of isolation of the continents, the South American rodent fauna was appar- ently represented only by caviomorphs. They also believe that the North American fauna then contained myomorph and sciuromorph rodents, but no caviomorphs. According to Hershkov- itz (1972) the tribe Sigmodontini, of the murid subfamily Cricetinae, distinguished by the possession of a complicated glans penis with a three digitated baculum, originated in this con- tinent from ancestors that rafted there from Africa. From South America they subsequently spread northward into North America and Eurasia. On the other hand, the Peromyscini, char- acterized by a simple glans penis, with an unbranched and normally elongated baculum, is essentially North American and apparently replaced the Sigmodontini, when they probably disappeared from boreal America during Oligocene. The complex-penis type Cricetinae seem to have more recently invaded Central and North America. This hypothesis appears to be sup- ported by some ectoparasites, such as fleas, occurring on these mammals. The flea genus Poly- genis, which has moved into Middle America with its complex-penis type Cricetinae and Cavi- omorph hosts from South America, is a good indicator of such an event (Wenzel and Tipton, 1966). In South America, as well as in other areas of the world, the early history of bats is obscure because of the paucity of fossils. However, it is estimated that they probably appeared in this continent after the Paleocene (Patterson and Pascual, 1972). Hershkovitz (1969) believes that bats were probably well established in north-western South America and the Isthmian region during most of the Tertiary. Storms may have played an important role in interchange of bats between the Americas when they were separated. During Pleistocene times, South America received from North America a diversity of mam- mals such as horses, mastodonts, tapirs, peccaries, camels, deer, some cricetine rodents, squir- rels, rabbits, canids, bears and other carnivores. Faunal interchange between North and South Quaes t. Ent., 1977 13 (2) 108 Mendez America during the late Tertiary, as well as radiation of many mammal groups has been the subject of much speculation and even today no causal explanation is generally accepted. How- ever, it appears that the majority of the families and genera of South American mammals or- iginated on this continent when it was completely or partially isolated. At the present time the northern sector of South America contains the major portion of the mammalian fauna, while the southern sector is exceedingly poor in number of species (Fittkau, j 1969; Osgood, 1942). Information based on Baker (1967), Cabrera and Yepes (1940) and other j authors, as well as on personal data, reveals that the major portion of southwestern Colombia, j corresponding to Pacific Coastal Lowlands, maintains a number of ubiquitious mammal spe- cies in addition to those forms possessing a more restricted distribution. The remaining mam- malian fauna of the southwest region of Colombia is fundamentally represented by a limited number of Andean elements. The recent biogeographical approach for the Neotropical region presented by Muller (1973) is followed for the brief zoogeographic description of mammals of the Departamento del Valle and the rest of southwestern Colombia. Muller based his interpretation of zoogeographical pat- terns on dispersal centers devised from comparative studies of plant and animal distribution. Colombia and several other territories of tropical America belong to the Brazilian Subregion of the Neutropical Region (Hershkovitz, 1958). The complexity of ecological situations in this nation has provided for the existence of a diverse mammalian fauna. Many areas, particu- larly those humid zones rich in vegetation and not yet disturbed by man, offer optimal con- dition for some populations of mammals, especially rodents and bats. They possess good re- productive potential and have been able to occupy a number of ecological niches through the development of specialized features. Those lands characterized by drier conditions and a low productivity of plants, particularly of fruit trees, as well as those somewhat disturbed by man, show, as a rule, less diversity and smaller populations of mammals. In a broad sense, sylvan species are not too selective in their food and habitat requirements, while pastoral species seem to display a higher level of adaptation by their food habits and locomotor organs. Among the Neotropical dispersal centers outlined by Muller, the following directly concern the Departamento del Valle and other major portions of southwestern Colombia: 1. the Cauca Center, 2. The Colombian Montane Forest Center, 3. The Colombian Pacific Center, and 4. The North Andean Center. For this discussion it is convenient to follow these zoogeographic divisions. Figure 5 has been prepared using as a source the map given by Muller (1973). The Cauca Center. — As defined by Muller, this zone is represented by the Cauca and Patia valleys, between the west and central Cordilleras of Northern Colombia, and are separated from each other by the Popayan plateau at an elevation of 1750 meters. Like many other areas of Colombia, this territory is being progressively deforested; however, it still harbors many mam- malian species, some of which are widely distributed in other areas of the Continent. Some of the faunal elements that previously existed in this territory probably disappeared many years ago. Faunal indicators of this zone are the wooly opossum, Caluromys derbianus derbianus, and red squirrel, Sciurus granatensis valdiviae, which are found in the Cauca and Patia valleys. The ancestors of these mammals evidently came from the north. According to Muller, the slight differentiation between the populations of these subspecies in the two valleys suggests that the Popayan plateau is not a strong barrier to species adapted to open habitats. In addi- tion, the forest biome of the southern part of the Cauca Valley did not prevent a spread of fauna during post-glacial times. Available information indicates that South America experienced drastic climatic changes during the Quaternary (Hammen, 1961 ; Patterson and Pascual, 1963; Vanzolini, 1973). Haffer (1967) considers that climatic changes during the Pleistocene and post-Pleistocene had more influence than orogenic events on the fauna west of the Andes. He also states that isolation Mammalian-Siphonapteran Associations 109 Fig. 5. Dispersal Centers related to the southwest Colombian fauna. Quaes t. Ent., 1977 13 (2) 110 Mendez and differentiation of most Pacific species occurred after the early Pleistocene uplift of the Northern Andes, probably under orographic conditions similar to those existing today. Ecological relationships of mammal species inhabiting this area are indicated in Table 1. Table 1. Ecological Relationships of Some Mammals of the Cauca Center HABITAT LIVING SPACE Surface or Aquatic or Roosting Arboreal or subsurface semiaquatic areas semiarboreal Meadow Zygodontomys brevicauda brunneus Sigmodon hispidus bogotensis Sylvilagus brasiliensis fulvescens Urocyon cinereoargenteus Odocoileus virginianus Forest: Oryzomys alfaroi palmirae Chironectes minimus Artibeus jamaicensis Didelphis m. marsupialis open or Oryzomys munchiquensis Lutra annectens Artibeus lituratus Philander opossum close canopy Oryzomys albigularis Peropteryx kappleri griscescens kappleri Thomasomys aureus Noctilio labialis Marmosa impavida caucae Thomasomys fuscatus Glossophaga soricina Rhipidomys latimanus Dasyprocta punctata Tadarida brasiliensis Reithrodontomys mexicanus Agouti paca guanta Dasypus novemcinctus Nasua nasua candace Conepatus semistriatus Mustela frenata Felis yagouaroundi Felis tigrina pardinoides Mazama americana zetti Lasiurus ega fuscatus Potos flavus megalotus Domestic Rattus rattus Molossus major or semi- Rattus norvegicus Eumops bonariensis domestic Mus musculus The Colombian Montane Forest Center. — This dispersal center seems to be confined to the forest biomes of Colombia (with the exception of the Sierra Nevada de Santa Marta), Ecuador and Venezuela. It includes two subcenters, namely, the West Andean and the East Andean. Inasmuch as affinities have been found between the animal populations of the Central Andes and those of the West Andean Subcenter, the Central Andes is included with the West Andean Subcenter (Muller, 1973). The fauna of the Colombian Montane Forest Center may be considered subtropical and dis- plays similarities with those of the Colombian Pacific Center, which is essentially tropical. This condition results from most of the species of the Colombian Montane Forest Center having evolved originally from ancestors that came from the lower lands. Characteristic mammals of the Colombian Montane Forest Center and their ecological re- lationships are indicated in Table 2. Mammalian-Siphonapteran Associations 111 Table 2. Ecological Relationships of Some Mammals of the Colombian Montane Forest Center HABITAT LIVING SPACE Surface or subsurface Aquatic or Roosting semiaquatic areas Arboreal or semiarboreal Forest: Cryptotis thomasi open or medallinius close canopy Oryzomys caliginosus monticola Nectomys alfari esmeraldarum Thomasomys aureus popayanus Thomasomys cinereiventer Chylomys instans Dasyprocta fuliginosa candelensis Sylvilagus brasiliensis fulvescens Nasua olivacea Mazama americana Domestic Rattus rattus or semi- Rattus norvegicus domestic Mus musculus Chironectes minimus Eptesicus brasiliensis Didelphis azarae andinus Ichthyomys hydrobates Metachirus nudicaudatus nicefori Myotis chiloensis colombianus Philander opossum griscescens Sciurus pucherani caucensis Reithrodontomys mexicanus milled Echinoprocta rufescens Aotus trivirgatus lemurinus The Colombian Pacific Center. - The extensive territory of lowlands in the western part of Colombia involves the major portion of the center. This territory continues to the north along the base of the mountains and ends on lands watered by the Magdalena River. To the south, this center encroaches upon a portion of the northern part of Ecuador. This center includes two subcenters: the Nechi, which encompasses the area between the Rio Sinu and the lower reaches of the Rio Cauca; and the Choco, which includes the area west of the Andes. Possibly during the Tertiary, the gap separating Central and South America divided the ter- ritory now known as the Colombian Pacific Center. This thesis stems from the existence at that time, of a seaway south of Panama and the Gulf of Uraba, which connected the Caribbean Sea and the Pacific Choco basin of Western Colombia (Haffer, 1967). This seaway was closed during the late Pliocene and seems to correspond to the Bolivar Geosyncline discussed by Ny- gren (1950) and Hershkovitz (1968). Glacial and interglacial periods of the Pleistocene evidently influenced the Pacific lowlands of Colombia. Glaciation of the mountains produced considerable temperature reduction and high humidity. At this time, sea level lowered about 1 00 m and extensive movements of fauna occurred between the Amazonian region and the trans-Andean area of western Colombia and Central America. The interglacial periods were particularly drier in the northern part of Col- ombia, when the humid forest moved southward due to influence of winds, and sea level rose about 30 to 50 m. As a result of this condition the Maracaibo basin and other parts of the Northern Colombian plains were flooded (Haffer, 1967). Studies of birds, lizards and amphibians presently inhabiting this center, have indicated the origin and distribution patterns of some of these vertebrates. A strong relationship of the Pac- ific lowland fauna of Colombia to that of the Amazon region suggests the interchange of ani- mals during remote times of the Pleistocene. Today, the Pacific mammalian fauna of the Quaes t. Ent., 1977 13 (2) 112 Mendez Departamentos del Valle, Choco, Cauca and Narino, inhabiting dense forests of the coastal lands, are, with some exceptions, similar to those of the Pacific lands of Eastern Panama and Northern Ecuador. Some elements of the mammal fauna of the Colombian Pacific Center and their ecological relationships are indicated in Table 3. Table 3. Ecological Relationships of Some Mammals of the Colombian Pacific Center HABITAT LIVING SPACE ,1 Surface or Aquatic or Roosting Arboreal or subsurface semiaquatic areas semiarboreal Marsh Procyon cancrivorus Hydrochaeris hydrochaeris panamensis isthmius Nectomys alfari esmeraldarum Meadow Zygodontomys brevicauda Sigmodon hispidus Odocoileus virginianus v tropicalis Forest: Oryzomys capito Chironectes minimus Carollia subrufa Didelphis marsupialis open or Oryzomys caliginosus Hydrochaeris hydrochaeris Carollia castanea Caluromys d. derbianus close canopy isthmius 1 Neacomys tenuipes tenuipes Chiroderma villosum Philander opossum Lutra annectes melanurus Heteromys australis Phyllostomus hastatus Proechimys guyannensis Phyllostomus discolor Marmosa robinsoni colombianus isthmica Proechimys semispinosus Sturnira lilium Cebus capucinus Hoplomys gymnurus Uroderma b. bilobatum Dasyprocta punctata Glossophaga soricina A teles fusciceps chocoensis Artibeus cincereus Tamandua tetradactyla Agouti paca Vampyrops heller i Cyclopes didactylus Nasua nasua Vampyrops dorsalis Bradypus griseus Mustela frenata Diaemus youngi Choloepus hoffmanni Eira barbara Desmodus rotundus Oryzomys concolor Galictis vittata canaster Tylomys mirae Felis yagouaroundi Dyplomys caniceps Felis pardalis Microsciurus flaviventer isthmius Felis onca Bassaricyon gabbi medius Tayassu pecari Potos flavus Tayassu tajacu Mazama americana Tapirus bairdii Domestic Rattus rattus Molossus major or semi- Rattus norvegicus Eumops auripendulus domestic Mus musculus The North Andean Center. — This area is the Central and Eastern Cordillera in Colombia and the Andean mountains in Ecuador and Peru. The most characteristic biome of this center is the paramos with its selected plant and animal life. To this center belong such mammals as whose names and ecological relationships are indicated in Table 4. According to Muller, it is apparent that 28 species of the North Andean faunal elements (more than 75%) belong to families of North and Central American origin. Chapman (1917), referring to birds from the paramo zone, indicated that the majority were Mammalian-Siphonapteran Associations 113 derived from the sea level equivalent of this zone in southern South America. It seems logical to assume that the same circumstance occurred with the mammals also. Table 4. Ecological Relationships of Some Mammals of the North Andean Center HABITAT LIVING SPACE Surface or subsurface Aquatic or semiaquatic Roosting areas Arboreal or semiarboreal Meadow Sylvilagus brasiliensis andinus Cerdocyon thorn Odocoileus virginianus goudotii Forest: open or close canopy Caenolestes obscurus Thomasomys cinereiventer cinereiventer Tremarctos ornatus Felis concolor Mazama americana Pudu mephistophiles Tapirus pinchaque My otis sp. Lasiurus sp. Marmosa dryas FLEAS OF SOUTHWESTERN COLOMBIA In this presentation descriptions are limited to those forms that are either new to science or have been previously described from one sex. Only citation of the original description for every taxon is given. For complete synonymy, type data and other distributional records the reader is referred to Johnson (1957), Tipton and Mendez (1966) and Tipton and Machado- Allison (1972). The principal sources of information of Colombian mammal hosts have been Allen (1912, 1913, 1915, 1916), Borrero (1967), Cabrera (1958 and 1961), Cabrera and Yepes (1940), Hershkovitz (1941, 1947, 1948, 1949, 1960, 1962), Gyldenstolpe (1932), Tate (1932a, b, 1935), and Osgood (1912). The following report pertains to taxa from southwestern Colombia available to us for study and forms that are not represented in our material but have been reported before from that territory or are likely to occur there. The order followed is that of Johnson (1957). To conserve space, I have abbreviated locality data for material of previously described species by omitting collection number, name(s) of collector(s), and designating only month of collection by Roman numeral. These details are available on request. For new species des- cribed here, complete data are given for each specimen. The acronym “HTC” preceding a num- ber in parentheses represents Harold Trapido Collection. Other numbers in parentheses refer to material in the collection of the Gorgas Memorial Laboratory. 3 KEY TO SPECIES OF FLEAS OF SOUTHWESTERN COLOMBIA 1 Thorax very reduced or compressed; antesensilial bristles absent; female burrowed into skin 2 1' Thorax not reduced or compressed; antesensilial bristles present; females not burrowed into skin 3 3. The following papers were helpful in the preparation of this key: Johnson (1954 and 1957), Ewing (1929), and Tipton and Mdndez (1966). Rhopalopsyllus lugubris, R. cacicus saevus, Polygenis dunni, and P. roberti beebei have not been re- ported from southwestern Colombia; however, they are likely to occur there and are included in this key. Quaest. Ent., 1977 13 (2) Mendez Head frons angular (Fig. 44); anterior lower margin of hind coxa with tooth-like projection; host not bat Tunga penetrans (Linnaeus), p. Head frons rounded; hind coxa without tooth-like projection; host-bat . . Rhynchopsyllus pulex Haller, p. Sword-like ridge of mesocoxa absent; mesonotum always lacking pseudo- setae; metanotum rectangular, not markedly broader dorsally than ven- trally; not more than one row of bristles on abdominal terga II- VI; meta- coxa with patch of spinelets on inside Sword-like ridge of mesocoxa present in most specimens; mesonotum with or without pseudosetae; metanotum much broader dorsally than ventrally; metacoxa without patch of spinelets on inside , Genal and pronotal comb present (Fig. 41) Ctenocephalides fells (Bouche), p. Genal and pronotal comb absent Mesothoracic pleural rod present; tergum VIII of female complete dor- sally; bulga of spermatheca (Fig. 40E) not globular, heavily pigmented . . Xenopsylla cheopis (Rothschild), p. Mesothoracic pleural rod absent; tergum VIII of female divided dorsally; bulga of spermatheca globular, lightly pigmented Dorsal aedeagal sclerite broad throughout; crochets small and elongate, rodlike; sternum VII of most females with 7-9 bristles Pulex simulans Baker, p. Dorsal aedeagal sclerite relatively long and slender; crochets expanded apically, not rodlike; sternum VII of most females with 4-5 bristles .... Pulex irritans Linnaeus, p. Head with true helmet (Fig. 1 1A, 1 19A); ctenidia on helmet, gena and pronotum Head without helmet (Fig. 16A); ctenidia absent or present, limited to gena and/or pronotum Helmet separated from head dorsally; vertical comb posteromarginal .... Helmet not separated from rest of head (Fig. 1 1 A); vertical comb mesad . Cleopsylla monticola Smit , p. Two large genal bristles anterior to cibarial pump; tergum VIII of male with apodeme; female spermatheca barrel-shaped, without internal tub- ercle, and hilla not enlarged basally One or two large genal bristles posterior to or vertically in line with cib- arial pump; tergum VIII of male without apodeme; female spermatheca not barrel-shaped, with internal tubercle or with hilla enlarged basally . . Genal comb teeth reduced, slightly longer than broad (Fig. 1 19A) Plocopsylla phyllisae Smit , p. Genal comb teeth normal, more than twice as long as broad (Fig. 14) ... . Plocopsylla thor Johnson, p. Pronotal comb spines about 18, not pointed (Fig. 12A); male with vertical row of four close-set spiniforms on sternum VII; female with subcaudal row of heavy spiniform setae on sternum IX (Fig. 12A) Sphinctopsylla diomedes Johnson, p. Pronotal comb spines about 30, pointed; male only with normal bristles on sternum VII; female without subcaudal row of heavy spiniform setae on sternum IX Sphinctopsylla tolmera (Jordan), p. 165 164 . 4 . 7 164 . 5 158 . 6 164 164 8 12 9 124 10 11 127 127 124 127 Mammalian-Siphonapteran Associations 115 12 ( 7') 12' 13 02') 13' 14 03) 14' 15 (14') 15' 16 (15') 16' 17 (16) 17' 18 (17') 18' 19 (18') 19' 20 (19) 20' 21 (20') 21' 22 (19') 22' 23 (22) Anterior margin of head conical, with two short spiniform bristles (Fig. 16A); tibia with dorsal comb of short, stout bristles Leptopsylla segnis (Schonherr), p. 131 Anterior margin of head not conical, without short spiniform bristles; tibia without dorsal comb of short, stout bristles 13 Several abdominal terga with well developed combs 14 Abdominal terga without combs 15 Crochets with convex margin dorsad; female sternum VII with caudal margin truncate Ctenidiosomus rex Johnson, p. 121 Crochets with convex margin ventrad; female sternum VII with caudal margin not truncate Ctenidiosomus traubi Johnson, p. 121 Anteroventral margin head with two large genal spines (Fig. 15A); on bats Sternopsylla distincta speciosa Johnson, p. 127 Anteroventral margin of head with more than two or without genal spines; not on bats 16 With combination of: anterior tentorial arm present, inserted anterior to eyes; mesopleural rod not dorsally bifurcated; ventral margin of pronotum not bilobed; tarsal segment V with four pairs of plantar bristles 17 Combination not as above 31 Antennal club symmetrical; without apical spinelets on metanotum .... Tetrapsyllus comis Jordan, p. 132 Antennal club asymmetrical; apical spinelets on metanotum 18 Frontoclypeal margin of head subconical; pronotal comb present (Fig. 36A) Scolopsyllus colombianus Mendez, p. 142 Frontoclypeal margin of head rounded; pronotal comb absent 19 Prosternosome projected downward between coxae, mesocoxa rectangular, margins parallel Rhopalopsyllus 20 Prosternosome not projected downward between coxae; mesocoxa asym- metrical, obviously broadest basally Polygenis 4 22 Spiracle of metepimere oblong, prolonged dorsally; bulga of spermatheca globular (Fig. 39D) . . Rhopalopsyllus lugubris Jordan and Rothschild, p. 158 Spiracle of metepimere rounded or ovoid; bulga not globular 21 Labial palpus extended to apex of coxa I or beyond (Fig. 38A); movable process of clasper longer than distal arm of sternum IX (Fig. 38B); sperm- atheca somewhat boomerang-shaped (Fig. 38C) Rhopalopsyllus cacicus saevus Jordan and Rothschild, p. 158 Labial palpus not extended to apex of coxa I (Fig. 37A); movable process of clasper about as long as distal arm of sternum IX (Fig. 37B). Sperma- theca strongly S-shaped (Fig. 37C) Rhopalopsyllus australis tupinus Jordan and Rothschild, p. 142 Distal arm of sternum IX much shorter than proximal arm; heel of sternum IX strongly angular; inner tube of aedeagus reflexed dorsally, not coiled apically 23 Distal arm of sternum IX about equal to length of, or longer than, proxi- mal arm; heel of sternum IX weakly angular; inner tube of aedeagus re- flexed ventrally, coiled apically 24 Clypeal tubercle above eye level (Fig. 32A); posterior margin of fixed process convex; spermatheca sinuate, without separation between bulga and hilla (Fig. 32E) Polygenis thurmani Johnson, p. 141 4. Females of Polygenis are difficult to identify and the characters used in this key pertain almost entirely to males. The females of P. trapidoi, n. sp. and/*, hopkinsi, n. sp., are unknown. Quaest. Ent., 1977 13 (2) 116 Mendez 23' 24 24' 25 25' 26 26' 27 27' 28 28' 29 29' 30 30' 31 31' 32 32' 33 Clypeal tubercle at eye level (Fig. 29 A); posterior margin of fixed pro- cess sinuate; spermatheca humped, with distinct separation between bulga and hilla (Fig. 29C) Polygenis klagesi (Rothschild), p. 140 (22') Distolateral lobes of aedeagus with angular projection; sternum VIII of male divided in half ventrally, with short dorsocaudal extension Polygenis trapidoi, new species, p. 141 Distolateral lobes of aedeagus without angular projection; sternum VIII of male not divided in half ventrally, without dorsocaudal extension 25 (24') Apex of distal arm of male sternum IX with distinct group of stout bristles Polygenis bohlsi bohlsi (Wagner), p. 132 Apex of distal arm of male sternum IX without distinct group of stout bristles 26 (25') Aedeagal fender present; dorsal margin "of fixed process of clasper sin- uate; spermatheca with cribose bulga, its ventral margin interrupted at bulga-hilla junction (with probable exception of P. hopkinsi , n. sp.) . . 27 Aedeagal fender absent (Fig. 30C); dorsal margin of fixed process of clasper usually slightly convex; spermatheca not cribose, its ventral margin continuous at bulga-hilla junction (Fig. 30E) Polygenis pradoi (Wagner), p. 140 (26 ) Aedeagal side piece above basal part of inner tube; aedeagal ribs not numerous (Fig. 25B) Polygenis delpontei, new species, p. 138 Aedeagal side piece absent or below basal part of inner tube; aedeagal ribs very numerous 28 (27') Aedeagal lateral lobes very reticulate; subapical ridge of median dorsal lobes of aedeagus present (Fig. 2 1C) Polygenis caucensis, new species, p. 137 Aedeagal lateral lobes not reticulate or faintly reticulate; subapical ridge of median dorsal lobes of aedeagus absent 29 (28') Labial palpus extended to trochanter I (Fig. 26 A); several distal arm bristles of sternum IX very long, approximately four times maximum width of distal arm (Fig. 26B) . . . Polygenis dunni (Jordan and Rothschild), p. 139 Labial palpus not extended to trochanter I; distal arm bristles of sternum IX short or as moderate length, not very long 30 (29') Aedeagal fender well developed, half-moon shaped; distal arm of sternum IX distinctly broad medially (Fig. 3 1 B) Polygenis roberti beebei (I. Fox), p. 140 Aedeagal fender reduced to slender, arched, inconspicuous structure; distal arm of sternum IX not distinctly broad medially (Fig. 27C) Polygenis hopkinsi, new species, p. 139 (16') Genal comb present 32 Genal comb absent 33 (31 ) First spine of genal comb almost overlapped by second (Fig. 8A); trabecula centralis present; labial palpus 5-segmented; both sexes with three antesensilial bristles. . . . Neotyphloceras rosenbergi (Rothschild), p. 117 First spine of genal comb not overlapped by second (Fig. 9A); trabecula centralis absent; labial palpus 4-segmented; both sexes with two antesen- silial bristles Adoratopsylla intermedia copha Jordan, p. 117 (31') Pronotal comb with more than 24 spines; bristles of antennal segment 2 long in both sexes (Fig. 17A) Dasypsyllus gallinulae perpinnatus ( Baker), p. 131 Mammalian-Siphonapteran Associations 117 33' Pronotal comb of most specimens with less than 24 spines; bristles of antennal segment 2 short in male, not extended to apex of club in female 34 34 (33') Protibia with seven dorsal notches with paired bristles; meso- and meta- tibia with six dorsal notches with paired bristles proximal to only single dorsal bristle Pleochaetis smiti Johnson, p. 131 34' Protibia with five or six dorsal notches with paired bristles; meso- and metatibia with five dorsal notches with paired bristles proximal to only single dorsal bristle Pleochaetis equato ris equatoris (Jordan), p. 131 SUPERFAMILY CERATOPHYLLOIDEA FAMILY HY STRICHOPS YLLIDAE SUBFAMILY CTENOPHTALMINAE TRIBE NEOTYPHLOCERATINI Neotyphloceras rosenbergi (Rothschild) (Figure 8) Typhloceras rosenbergi Rothschild, 1904, Novit. Zool., 11: 639, PI. 13, Fig. 68-69; PI. 14, Fig. 71, 74. Material examined. — Ex Didelphis marsupialis. Depto. del Valle, Municipio de Cali - 26, Pichindd, 1600m, VIII. Ex Didelphis azarae. Depto. del Valle - 6 , Finca Holanda (nr. Paramo de Chinche), 2700m, X. Ex Oryzomys caliginosus. Depto. del Valle, Municipio de Cali - 3 6, 4? Quebrada Honda, nr. Pichindd, 1800m, I, III, VIII, IX. Ex Oryzomys alfaroi. Depto. del Valle, Municipio de Cali - 2? Quebrada Honda, nr. Pichinde', 1800 m, III, IX; 9 , Florida, 8 km. S.E. “La Diana”, 1700m., X. Ex Oryzomys albigularis. Depto. del Valle, Municipio de Cali - 4(3, 4? , Valle del Rib Pichinde', 1700 - 1900 m., I, X, XII; 2(3, Cerro Munchique, 60 km. by road W. Popayan, Pena del Perro, 2160m., V; 9, Finca La Flora, Quebrada Norte, Pichindd, 1900m., VIII; 5(3, Saladito, Km. 12, 2000m., Ill; 4<3, 59, Finca Holanda (nr. Pdramo de Chinche), 2700 m., X. Depto. del Cauca - 9, Cerro Munchique, Finca El Retiro, 2200m., V. Ex Oryzomys (Oligoryzomys) species. Depto. del Valle, Municipio del Cali - 9, Quebrada Honda, nr. Pichindd, 1800m., X. Ex Rhipidomys latimanus. Depto. del Valle, Municipio de Cali - 9, Quebrada Honda, nr. Pichindd, 1800m., X; <3, 29, Valle del Ri'o Pichindd, 1700 - 1900 m., VII, XII. Ex Rhipidomys similis. Depto. del Cauca — 9, Cerro Munchique (60 km. by road W. Popayan, sitio No. 1), 2000m., V. Ex Thomasomys aureus. Depto. del Cauca — 9 > Pilimbala, 3100m., V. Ex Thomasomys cineriventer. Depto. del Cauca, Cerro Munchique -9,60 km. by road W. Popayan, Sitio. No. 1, 2500m., V; (3, sitto No. 3, V; 6, 60 km. by road W. Popayrfn, Pena del Perro, 2160 m., V. Depto. del Narino - 6(3, 9 , Laguna de La Cocha, 2700 m., V; <3, Km 38, between Pasto & Sibundoy, Comisario de Putumayo, 3100 m., V; <3, Km. 33, between Pasto & Sibundoy, 2900 m., V. Ex Thomasomys fuscatus. Depto. del Valle, Municipio de Cali - 3 <3, Valle del Rib Pichindd, 1700 - 1900 m., I, III, XI; 2(3, 29, Pichindd, 1900 m., V; 2(3, 9, Pichindd, Rincon del Yarumal, V; <3, Pichindd, Finca La Flora, VII; 6(3, 8 9, Saladito, Km. 12, 2000 m., II. Depto. del Narino - 9, Laguna de La Cocha, 2700 m. Remarks. - In addition to Colombia, this species is distributed in Venezuela, Ecuador and Peru. Specimens are regularly found on a variety of rodents and less commonly on marsupials. Our material contains specimens from 1 2 species of mammals, indicated above. Other hosts in the literature for this species are the following: Philander opossum, Marmosa, Akodon, Chil- omys, Rheomys, Stictomys, and Sigmodon. The variety of hosts on which fleas of this species are found indicates a low degree of host specificity. TRIBE ADORATOPSYLLINI Adoratopsylla (Tritopsylla) intermedia copha Jordan (Figure 9) Stenopsylla intermedia copha Jordan, 1926, Novit. Zool., 33: 391, Fig. 15. Quaest. Ent., 1977 13 (2) 118 Mendez Fig. 6. General appearance of female of Ctenocephalides felis (Bouche). AN. - Antenna, A.S. - Antesensilial seta, A.S.T. - Anal stylet of female, B.C. - Bursa copulatrix, C. - Coxa, CL. - Claw of tar- sus, D.A.L. - Dorsal anal lobe, E. - Eye, EP. - Epipharynx, F. - Femur, FR. - Frons, G.CT. - Genal ctenidium, L. - Lacinia, L.P. - Labial palp, L.P.B. - Lateral plantar bristles, M.L. - Maxillary lobe, M.P. - Maxillary palp, MPM. - Mesepimere, MPS. - Mesepisternum, MSN. - Mesonotum, MTM. - Metepimere, MTN. - Metanotum, MTS. - Metasternum, O. - Occiput, O.B. - Ocular bristle, P. - Prono- tum, P.CT. - Pronotal ctenidium, PRM. - Proepimere, S. - Sensilium, SP. - Spermatheca, ST. - Sternum, T. - Tergum, TA. - Tarsus, TI. - Tibia, V.A.L. - Ventral anal lobe. Mammalian-Siphonapteran Associations 119 DLL Fig. 7. Structures of apex of aedeagus of Scolopsyllus colombianus Mdndez. A.M.S. - Apico-median sclerite, AP.S. - Apodemal strut, CR. - Crochet, Cr.P. - Crochet processes, C.S. - Crescent sclerite, DL.L. - Distolateral lobes, F.M. - Fluted membrane, H.L. - Heel at base of aedeagal pouch, I.T.-A - Apical portion of sclerotized inner tube, I.T.-B - Basal portion of inner tube, L.L. - Lateral lobes, M.D.L. - Median dorsal lobes, P.R. - Penis rods, PS.T. - Pseudotube. Quaes t. Ent., 1977 13 (2) 120 Mendez Fig. 8. Neotyphloceras rosenbergi (Rothschild). Female. A. Head, prothorax and procoxa; B. Modified abdominal segments; C. Spermatheca. Mammalian-Siphonapteran Associations 121 Material examined. — Ex Didelphis azarae. Depto. del Valle - 3(5, 3?, vie. Cali, X. Ex Didelphis marsupialis. Depto. del Valle, Municipio de la Cumbre - (5, 9 , La Maria, 1400 m, XI; Municipio de Cali - 7(5, 79, Pichindd, 1600 m., VIII; 10(5, 159, Lago Calima, 1450 m., II. Ex Philander opossum. Depto. del Valle - Pichindd, 1600 m., VIII; 39, Alto Anchicaya', 650 m., II. Ex Oryzomys caliginosus. Depto. del Valle - (5, Pichindd, 1600 m., I. Ex Thomasomys fuscatus. Depto. del Valle - 2(5, 39, Saladito, (Km. 12), 2000 m., II. Remarks. — This subspecies has been reported from Colombia, Panama, Ecuador and Peru, from sea level to over 3000 meters. It is a common parasite of marsupials and displays a high degree of infestation on some single host animals (Tipton and Mendez, 1966). In southwestern Colombia specimens have been obtained from the hosts indicated above. Elsewhere specimens have been recorded from Oryzomys caliginosus and Proechimys semispinosus. FAMILY PYGIOPSYLLIDAE SUBFAMILY PYGIOPSILLINAE Ctenidiosomus rex Johnson Ctenidiosomus rex Johnson, 1957, Mem. Ent. Soc. Wash. 5:50, PI. 20. Remarks. — Type material (2 males and 2 females) from San Agustin, Departamento de Huila, Colombia, represents the only specimens known of this taxon. These specimens were collected from Thomasomys, Oryzomys and Rhipidomys. It seems probable that a species of Thomasomys (probably T. laniger ), is the natural host of this flea species. Ctenidiosomus traubi Johnson (Figure 10) Ctenidiosomus traubi Johnson, 1957, Mem. Ent. Soc. Wash. 5:49-50, PI. 17; PI. 18, Fig. 5; PI. 19, Fig. 1, 2. The original description is based on the holotype female, ex Caenolestes obscurus, Colom- bia, Depto. de Antioquia, Sanson, 7 km. E. of Paramo, 3,160 m., 18 Oct. 1950, P. Hershkovitz collector. Males are described below. Material examined. — Ex Caenolestes obscurus. Depto. del Cauca - 9 , Puracd Park, 3520 m., V. Ex Thomasomys aureus. Depto. del Cauca - (5, Pilimbala, 3100 m., V. Ex Thomasomys cinereiv enter. Depto. del Narino - (5, Comisam Putumayo, Km. 77, between Sibundoy & Mocoa, 2200 m., V. Description of male. — Head (Fig. 10A). Strongly fracticipit, with frons evenly rounded, without clypeal tub- ercle. Preantennal region with 3 distinct discs and abundant micropores; principal bristles of preantennal region arranged in 2 rows, secondary bristles very short, scattered mainly on preocular area. Eye deeply excised ventrally, weakly pigmented. Postocular area with 2 pits just below eye. Genal area bilobed, its anterior lobe or genal process broadly rounded, not accum- inate. Posterior lobe of gena evenly rounded, moderately broad. Postantennal region with anterior micropores, several pits profusely distributed, 2 rows of bristles, in addition to short bristles on dorsal margin and in front of antenna, mainly on antennal fossa. Pedicellus of antenna covered with short bristles. Thorax. Very setose. Pronotum with 2 rows of bristles preceding comb of about 26 spines. Mesonotum at least with 3 well defined rows of bristles, remaining bristles short, concentrated on anterior pronotal region. Mesepisternum with few non-prominent bristles on anterodorsal area near pleural ridge of mesothorax. Mesepimere with bristles of different sizes. Metanotum with about 3 or 4 defined rows of bristles, posteriormost row of long bristles and short intercalaries. Other bristles not arranged in rows in anterior metanotal area. Lateral metanotal area apparently with no more than 1 bristle. Met- episternum with oval outline interrupted by anterior projection of metasternum, provided with few bristles. Metepimere with 3 rows of uneven bristles. Abdomen. Combs on terga II- V, and with various number of spines (in the two specimens examined), respectively, 14-15; 14-15; 12-13; 14-15. Upper antesensilial birstle about 1/2 as long as lower bristle. Modified abdominal segments. Tergum VIII reduced to subtriangular plate provided with broad spiracle, with group of short bristles near antesensilial bristles. Sternum VIII large, ensheating principal structures of genitalia, provided with num- erous marginal and inner bristles, caudal margin entire, without sinus. Clasper large, somewhat pyriform, projected anterior- ly into short manubrium curved upward. Process with subrounded apical expansion, largely squamose, with group of bristles on outer and inner surfaces. Posterocaudal margin of process with four long bristles. Movable process of clasper broadest near basal area, narrowed apically, with bristles of varied size. Sternum IX (Fig. 10B) like those of other males of genus. Distal arm club-shaped, with broad and rounded apex with 4 stout bristles on caudal margin; remaining bristles smaller, Quaest. Ent., 1977 13 (2) 122 Mendez Fig. 9. Adoratopsylla intermedia copha Jordan. Male. A. Head, prothorax and procoxa; B. Genitalia; C. Apex of aedeagus, A.I.T. - Armature of inner tube, CR. - Crochet, C.S. - Crescent sclerite, L.L. - lateral lobes, L.S.I. - Lateral sclerotization of inner tube, S.I.T. - Sclerotized inner tube. Female. D. Spermatheca and 7th abdominal segment. From “The Fleas (Siphonap- tera) of Panama” by Tipton and Mdndez, in “Ectoparasites of Panama”, Field Museum of Natural History, Chicago (1966). Mammalian-Siphonapteran Associations 123 Fig. 10. Ctenidiosomus traubi Johnson. Male. A. Head, prothorax and procoxa; B. Ninth sternum; C. Clasper; D. Apex of aedeagus. Quaest. Ent., 1977 13 (2) 124 Mendez scattered over most of arm. Proximal arm of sternum IX narrow at base, broad at apex. Aedeagus like that of Ctenidiosomus perplexus Tipton and Machado-Allison. Median dorsal lobe broad, with rounded dorsal margin, produced as caudal subacum- inate blade. Lateral lobes narrower than distal lobe, slightly arched. Crochets crescent-shaped. Aedeagal apodemal rod arched apically. Penis rods strongly coiled, extensively fimbriate on apical portion. Remarks. — Presently, Ctenidiosomus traubi is known only from Colombia, where it has been collected in localities with elevation ranging from 2200 m to 3500 m. The scant material available has been collected from one marsupial and two cricetine rodent species. As Lewis (1974) pointed out, it is likely that the preferred hosts of C. traubi are rodents. FAMILY STEPHANOCIRCIDAE SUBFAMILY CRANEOPSYLLINAE TRIBE CRANEOPSYLLINI Cleopsylla monticola Smit (Figure 1 1) Cleopsylla monticola Smit, 1953, Bull. Brit. Mus. (Nat. Hist.) Entomol., 3(5): 193, Fig. 13, 15, 17, 19, 20. Material examined. — Ex Oryzomys albigularis. Depto. del Cauca - <5, 9 , Cerro Munchique, 60 km. by road W. Popaydn, sitio No. 1, 2500 m., V. Depto. del Valle - 13 <5, 69, Finca Holanda (nr. Paramo de Chinche), 2700 m., X. Ex Rhipidomys similis. Depto. del Valle - 2(5, 9 , Finca Holanda, 2700 m., V. Ex Thomasomys cinereiv enter. Depto. del Narino — 2c5, 9 , Laguna de La Cocha, 2700 m., V. Ex Thomasomys fuscatus. Depto. del Valle - (5, Pichindd, Finca La Flora, 1900 m., V. Remarks. — Reports of this helmet flea are from Ecuador, Colombia and Venezuela. In Colombia, specimens have not been collected above 2700 meters elevation. The range of ver- tical distribution for this species in Venezuela is from 120 meters to 1443 meters (Tipton and Machado-Allison, 1971). Five species of cricitines harbor C. monticola in southwestern Colom- bia. (see above for details) Other hosts recorded in the literature are Caenolestes fuliginosus, Didelphis marsupialis, Marmosa fuscata, M. dryas, Oryzomys minutus, Rhipidomys venustus, Rhipidomys sp., Thomasomys hylophilus, T. laniger, T. vestibus, Chilomys ins tans, and birds. Sphinctopsylla diomedes Johnson (Figure 12) Sphinctopsylla diomedes Johnson, 1957, Mem. Ent. Soc. Wash. 5:68, PI. 32. This species was originally described from two male specimens ex Caenolestes obscurus, Colombia: Depto. of Huila, San Agustin, San Antonio, left bank of Rio Magdalena (Cordillera Central), 2200 m, 24 Aug. 1950. P. Hershkovitz collector. Material examined. — Ex Caenolestes obscurus. Depto. del Cauca - 8(5, 79, PuracdPark, 3500 m., IV- VI. Depto. de Cundinamarca, Municipio de Soacha - 9, Soche, 2700 m., IX. Description of female. — Head (Fig. 12A). Frons margin moderately rounded. Helmet comb of 13 spines. Genal comb of 5 spines. Thorax. Pronotum (Fig. 12 A) with 2 rows of bristles and conspicuous comb of 9 spines per side. Remaining thoracic structures and legs as in male. Abdomen. Terga I-IV with apical spineletes. All terga with two rows of bristles but anteriormost row more reduced. Ter- gum VII with 2 subequal antesensilial bristles. Modified abdominal segments (Fig. 12B). Posterior margin of tergum VIII sinuous, with 2 groups of large, stout spini- form bristles, upper group of 4-6 bristles, lower group of 1-3. Both groups preceded by scattered bristles of various size and location, in addition to short and moderate size marginal bristles. Spiracle of tergum VIII with very broad basal portion. Sterna II- VI with 1 row of 6 bristles. Sternum VII with group of 3-4 stout spiniform bristles on each side, in addition to sev- eral inconspicuous bristles. Posterior margin of this sternum almost straight, not indented. Sensillum with about 11 sensory pits per side. Dorsal anal lobe with several bristles. Ventral anal lobe with only 1 or 2 bristles. Anal stylet short and stout, dorsally and ventrally convex, its apical bristle about twice the length of stylet body, with minute ventral bristle and mesal bristle of moderate size. Spermatheca (Fig. 12B, 12C) with divided bulga, anterior section globular, fairly reticulate, and pos- terior section not reticulate, followed by short, upturned unpigmented hilla. Main body of bursa copulatrix short, sinuous, weakly sclerotized. Mammalian-Siphonapteran Associations 125 B Fig. 11. Cleopsylla monticola Smit. Male. A. Head, prothorax and procoxa; B. Mesothorax, metathorax and tergum I. Quaest. Ent., 1977 13 (2) 126 Mendez B Fig. 12. Sphinctopsylla diomedes Johnson. Female. A. Head, prothorax and procoxa; B. Modified abdominal segments; C Spermatheca. Mammalian-Siphonapteran Associations 127 Remarks. — This species is endemic to southwestern Colombia where specimens have been found in areas between 2200 and 3500 meters. All specimens presently existing in collections are from Caenolestes obscurus individuals, which seem to be the natural host. Sphinctopsylla tolmera (Jordan) Craneopsylla tolmera Jordan, 1931, Novit. Zool., 36:314, Fig. 5. Material examined. — Ex Thomasomys cinereiventer. Depto. del Narino -(5,9, Laguna de La Cocha, 2700 m., V. Depto. del Valle - <5, Finca Holanda (nr. Paramo de Chinche), 2700 m., X. Remarks. - The geographical range of S. tolmera involves Colombia, Ecuador and Venezuela, in some areas exceeding 2000 meters elevation. Rodents of the genus Thomasomys probably are the preferred hosts in Colombia and Ecuador. In Venezuela this flea species seems to be more associated with Oryzomys minutus. Of 36 males and 76 females recorded by Tipton and Machado- Allison (1972), 32 males and 63 females were recovered from 46 specimens of this rodent species. Plocopsylla phyllisae Smit (Figure 13) Plocopsylla phyllisae Smit, 1953, Bull. Brit. Mus. (Nat. Hist.) Entomol., 3:197, Fig. 25, 26, 28, 30. Material examined. — Ex Caenolestes obscurus. Depto. del Cauca — 7(5, ll9, PuracdPark, 3500 m., IV, V. Depto. de Cundinamarca, Municipio de Soache — 29, El Soche, 2700 m., IX. Remarks. — Some populations of this species live in territories over 3000 meters elevation in Ecuador and Colombia. The holotype male was secured from Oryzomys sp; nevertheless, the material (11 <5c5 and 2199) available to us for study came from six specimens of Caenoles- tes obscurus, which is probably the preferred host. Individuals of this terrestrial marsupial live in dark damp forests of paramos and are crepuscular or nocturnal. They seem to be pri- marily insectivorous (Osgood, 1921 ; Tate, 1931). Plocopsylla thor Johnson (Figure 14) Plocopsylla thor Johnson, 1957, Mem. Ent. Soc. Wash., 5:73-74, PI. 38 (Fig. 1 , 2, 3, 6, 7), PI. 39 (Fig. 1, 2, 3). Material examined. — Ex Thomasomys cinereiventer. Depto. del Cauca - 9, Cerro Munchique, 60 km. by road W. Popaya'n, sitio No. 3, V. Depto. de Narino - Laguna de La Cocha, 2700 m., V. Remarks. - Plocopsylla thor seems to be restricted to areas of high elevation (particularly between 2000 and 3000 meters) in Colombia. Specimens have been found associated with cricetine rodents of the species Oryzomys albigularis and Thomasomys spp. The typical host for this flea is probably Thomasomys cinereiventer. FAMILY ISCHNOPSYLLIDAE SUBFAMILY ISCHNOPSYLLINAE Sternopsylla distincta speciosa Johnson (Figure 15) Sternopsylla distincta speciosa Johnson, 1957, Mem. Ent. Soc. Wash. 5: 100; PI. 48, Fig. 3, 4; PI. 50, Fig. 3, 8. Remarks. - The description of this subspecies is based on a holotype male, an allotype fe- male, and three female paratypes ex Tadarida brasiliensis, Peru: Dept, of Cuzco, Quince Mil, 19 June 1950, C. Kalinowski collector. One male and three female paratypes ex Tadarida sp., Colombia: Dept, of Huila, Pitalico, 1350 m, 28 Nov. 1951, P. Hershkovitz collector. No other Quaest. Ent., 1977 13 (2) 128 Mendez Fig. 13. Plocopsylla phyllisae Smit. Male. A. Head, prothorax and procoxa; B. Mesothorax, metathorax and tergum I; C Modified abdominal segments; D. Spermatheca. Mammalian-Siphonapteran Associations 129 Fig. 14. Plocopsylla thor Johnson. Male. Head, prothorax and procoxa. Quaest. Ent., 1977 13 (2) 130 Mendez Fig. 15. Sternopsylla distincta speciosa Johnson. Male. A. Head, prothorax and procoxa; B. Eight sternum; C. Process and movable finger of clasper; D. Distal arm of ninth sternum. Female. E. Modified abdominal segments; F. Anal stylet and ven- tral anal lobe; G. Spermatheca. From “The Fleas (Siphonaptera) of Panama” by Tipton and Mdndez, in “Ectoparasites of Panama”, Field Museum of Natural History, Chicago (1966). Mammalian-Siphonapteran Associations 131 Colombian records are mentioned in the literature. FAMILY CERATOPHYLLIDAE SUBFAMILY LEPTOPSYLLINAE Leptopsylla segnis (Schonherr) (Figure 16) Pulex segnis Schonherr, 1811, K. svenska Ventenskakad. Handl. (2) 32:98, PI. 5, Fig. A. B. Material examined. — Ex Rhipidomys latimanus. Depto. del Valle, Municipio de Cali - 9, Quebrada Honda, nr. Pichindd, 1800 m., IX. Ex Rattus rattus. (3, same locality as above. Remarks. — Leptopsylla segnis is a cosmopolitan species which has been introduced with commensal rodent hosts to many parts of the world. In South America it seems to be confined to areas of high elevation in Colombia, Ecuador, Peru, Venezuela, Brazil, Chile and Argentina. Typical hosts are various species of Muridae; however, the true host is the house mouse, Mus mus cuius. SUBFAMILY CERATOPHYLLINAE Dasypsyllus gallinulae perpinnatus (Baker) (Figure 17) Ceratophyllus perpinnatus Baker, 1904, Proc. U.S. Nat. Mus., 27: 386, 391, 445, Fig. 1-6. Material examined. — Ex Thomasomys cineriventer. Departo. del Cauca, Cerro Munchique - (3, 60 km. by road W. Popay^n, sitio No. 1, 2500 m., V; <3, sitio No. 3, 2500 m., V; 9, Pena del Perro, 2160 m., V. Remarks. - Dasypsyllus gallinulae perpinnatus is a widespread bird flea species, recorded from several countries in the New World. Our specimens represent the first report of this tax- on for Colombia. It is also known from Canada, United States of America, Panama, Venezuela and Argentina. Our specimens were obtained from different sites and dates on three Thomasomys cinerei- venter specimens. These rodents were trapped on the ground; however, the fact that they were parasitized by this bird flea suggests that T. cinereiventer is perhaps partly arboreal. The fleas were probably obtained from bird nests located on trees visited by the rodents. At the present time little is known about the habits of T. cinereiventer. Pleochaetis equatoris equatoris (Jordan) Ceratophyllus equatoris Jordan, 1933, Novit. Zool., 38: 344, Fig. 63, (partim ). Material examined. — Ex Thomasomys cinereiventer. Depto. del Narino 9, Laguna de La Cocha, 2700 m., V; 9, Km. 77, between Sibundoy & Mocoa, Comisaria de Putumayo, 2200 m., V. Remarks. — Pleochaetis equatoris equatoris has been reported from Peru, Ecuador and Col- ombia. According to Johnson (1957), it is likely that specimens from Peru assigned by Macch- iavello (1948) to P. equatoris equatoris are P. dolens quitanus. Pleochaetis smiti Johnson (Figure 18) Pleochaetis smiti Johnson, 1954, Jour. Wash. Acad. Sci., 44(9): 291, 295, Fig. 1, 3, 6-8, 10, 12, 13, 16, 21, 25, 26, 31. Material examined. — Ex Caenolestes obscurus. Depto. del Cauca — 9, Puracd Park, 3500 m., IV. Ex Oryzomys albigularis. Depto. del Cauca - <3, Cerro Munchique, 60 km. by road W. Popayan, sitio No. 3, 2500 m., V. Depto del Valle - 2(3, 29, Finca Holanda (nr. Paramo de Chinche), 2700 m., X. Quaest. Ent., 1977 13 (2) 132 Mendez Ex Thomasomys cinereiventer. Depto. de Narino - <3, Laguna de La Cocha, 2700 m., V; Comisaria de Putumayo - 9, Km. 38, between Pasto & Sibundoy, 3100 m, V; <3, Km. 77, between Sibundoy and Mocoa, 2200 m., V. Remarks. - P. smiti is to date known from Colombia, Ecuador and Venezuela. Judging from our collection and the data available in the literature, the vertical distribution of this species ex- tends from 1980 to 3810 meters. Tipton and Machado- Allison (1972) report abundant mat- erial (203 males and 208 females) from Venezuela. Evidence is presented by these authors that the characteristic host of P. smiti in Venezuela is Oryzomys minutus. They recovered 364 spe- cimens of P. smiti from 158 specimens of this rodent. I suspect that in Colombia, P. smiti is probably more specific on Oryzomys albigularis\ however, flea specimens collected in this country are few and do not allow final interpretation as to preferred host species. SUPERFAMILY RHOPALOPSYLLOIDEA FAMILY RHOPALOPSYLLIDAE SUBFAMILY RHOPALOPSYLLINAE TRIBE PARAPSYLLINI Tetrapsyllus comis Jordan (Figure 19) Tetrapsyllus comis Jordan, 1931, Novit. Zool., 37:135, Fig. 1. Material examined. — Ex Caenolestes obscurus. Depto. del Cauca - 2?Purace Park, 3500 m., V. Remarks. — Tetrapsyllus comis was hitherto known only from Ecuador. Our record is the first for the Republic of Colombia. The scant information about this flea does not allow for determination of host preference. We have two females from Caenolestes obscurus while the Ecuadorian holotype female was taken on Sigmodon sp. Apparently, T. comis is a typical member of the Andean fauna of the northwest portion of South America, perhaps limited in its distribution to Colombia and Ecuador. The male of this species remains unknown. TRIBE RHOPALOPSYLLINI Polygenis bohlsi bohlsi (Wagner) (Figure 20) Pulex bohlsi Wagner, 1901, Hor. Soc. Ent. Ross., 35:21, PL 1, Fig. 6. Material examined. — Ex Oryzomys albigularis. Depto. del Valle - <3, Valle del Rio Pichindd, 1700 - 1900 m., Ill; Saladito (Km. 12), 2000 m., II. Ex Oryzomys alfaroi. Depto. del Valle, Municipio de Buga - 5(3, 79 , Sonso, 1000 m., V. Ex Oryzomys caliginosus. Depto. del Valle, Municipio de Buga - 14(3, 149, Sonso, 1000 m., Ill, VI. Depto. del Valle, Municipio de Cali - 4(3, 79, Valle del Rio Pichindd, 1700 - 1900 m., Ill, IV, VI, VII, 6, 59 , Pichindd, 1800 - 1880 m., I, III, IV, VII. Depto. del Valle - 39 , La Buitrera, 1000 m., II; 35(3, 479 , Lago Calima, 1450 m., II, III; (3, Florida, 8 km. S.E. “La Diana”, 1700 m., XII. Ex Thomasomys fuscatus. Depto. del Valle, Municipio de Cali - 9 , Pichindef, IV. Remarks. — Our records of P. bohlsi bohlsi extend the range of this taxon which is now known from Colombia, Ecuador, Venezuela, Trinidad, Brazil, Argentina and Paraguay. The majority of our southwestern Colombia specimens are from oryzomine rodents obtained from 1000 to 2000 meters elevation. Tipton and Machado-Allison (1972) suggest that the optimum habitat of this flea species is at elevations between 1000 and 1500 meters and the preferred hosts are cricetine rodents and perhaps more specifically of akodont stock. Mammalian-Siphonapteran Associations 133 Fig. 16. Leptopsylla segnis (Schonherr). Male. A. Head, prothorax and procoxa; B. Mesothorax, metathorax and tergum I; C. Process and movable finger of clasper; D. Hind femur and hind tibia. Female. E. Modified abdominal segments; F. Sperma- theca. Quaest. Ent, 1977 13 (2) 134 Mendez Fig. 17. Dasypsyllus gallinulae perpinnatus (Baker). Male. A. Head, prothorax and procoxa; B. Genitalia. Female. C. Sperma- theca and 7th abdominal segment. From “The Fleas (Siphonaptera) of Panama” by Tipton and Mdndez, in “Ectoparasites of Panama”, Field Museum of Natural History, Chicago (1966). Mammalian-Siphonapteran Associations 135 B D Fig. 18. Pleochaetis smiti Johnson. Male. A. Head, prothorax and procoxa; B. Process and movable finger of clasper; C. Distal arm of 9th sternum. Female. D. Modified abdominal segments; E. Spermatheca. Quaest. Ent., 1977 13 (2) 136 Mendez Fig. 19. Tetrapsyllus comis Jordan. Female. A. Head, prothorax and procoxa; B. Modified abdominal segments; C. Spermathe- Mammalian-Siphonapteran Associations 137 Polygenis caucensis, new species (Fig. 21,22) Type Material. — Holotype 6 ex Oryzomys caliginosus, (00036), Alto Anchicaya', 650 m, Depto. del Valle, Colom- bia, 11.11.1974, E. Mdndez & L. Vela'squez; allotype 9, (00032), same locality, date and collector; 9paratype, (00042), same locality, date and collector; <3 paratype (Acc. No. B-571), ex Didelphis marsupialis, Curiche River, Depto. del Choco, Colom- bia, 19. VI. 1967, D.G. Young Holotype and allotype are in the National Museum of Natural History, Washington, D.C.; I paratype is in the British Museum (Natural History) and 1 paratype is in the Gorgas Memor- ial Laboratory’s collection. Diagnosis. - This species appears to be closest to Polygenis roberti beebei I. Fox. Males are differentiated by the more reticulate aedeagal lateral lobes. In addition, P. caucensis males have an aedeagal median dorsal lobe with a subapical ridge, a structure apparently absent from males of P. r. beebei. Description. — Male. Head (Fig. 21A). Frons fairly rounded, interrupted by short, angular tubercle protruded up- ward. Preantennal region with moderate number of micropores and 2 pits in front of antennal scape. Preocular row of 2 bri- stles inserted near eye. Post-ocular row with few minute bristles on lower and upper portions of preocular region. Arch of tentorium not conspicuous. Oral angle well defined. Genal lobe semiangular. Eyes subovate, not incised, large and well pig- mented. Maxilla with acuminate tip extended to last segment of maxillary palp. Post-antennal region with 3 rows of bristles behind antennal scape. Antennae densely covered with minute and prominent bristles irregularly distributed. Thorax. Pronotum and mesonotum with 2 rows of bristles. Mesepisternum with 2 large bristles per side. Mesepimeron apparently with 2 or 3 bristles per side. Metanotum with 3 rows of bristles. Metanotal flange with about 6 spinelets. Metep- isternum with single bristle. Metepimeron with 9 bristles in 2 rows. Legs. Posteromarginal notches of metatibia with strong bristles with following distribution: 2-2-2-3-2-3. Abdomen. Terga I-V with spinelets and 2 rows of bristles. Unmodified sterna with single row of bristles, those of sterna II and III preceded by very few marginal and submarginal bristles. Modified abdominal segments (Fig. 21C). Sternum VIII somewhat reduced, with several unequal bristles. Fixed process of clasper (Fig. 21B) broad, with its total length in excess of maximum width, and with barely projected apical lobe, dorsal margin shallowly sinuate, posterior and ventral margins strongly sinuate, last one moderately indented. Chaetotaxy as illus- trated. Movable finger of clasper (Fig. 21B) not extended to apical lobe, with several marginal and inconspicuous inner bris- tles. Proximal arm of sternum IX shorter than distal arm, of irregular shape, narrowed at basal portion, then considerably ex- panded and sinuate, terminated in subangular projection. Distal arm of sternum IX (Fig. 22A) curved cephalad, more dilated medially but gradually tapered toward its subrounded tip. Chaetotaxy of this arm limited to some apical bristles and postero- marginal bristles of various sizes. Heel of sternum IX prominent, with slender, straight terminal tendon. Sternum VIII with wide subrounded caudal lobe and single row of unequal bristles. Apodeme of aedeagus (Fig. 22B) broad, shorter than termi- nal portion of aedeagus, with subrounded apex. Terminal portion of aedeagus (Fig. 22B) with semirounded distolateral lobes. Median dorsal lobe large, almost reaching posterior portion of distolateral lobes, with upper margin shallowly convex. Crochet not apparent. Lateral lobe gently reticulated, broadly curved, except for short basal angular prominence. Side piece prominent, almost triangular, its longest side fairly convex. Fender well sclerotized, arched and conspicuous. Apical portion of inner tube larger than basal portion, strongly convoluted, with final portion slender. Basal portion of inner tube of moderate proportions, with narrow foramen on anterior half. Crescent sclerite short, indistinct. Side piece broad, of irregular shape, with anterior margin convex, posterior margin angulate. Fulcral latero-ventral lobe represented by short, slightly curved knob-like structure. Pseudo tube long, sinuate, well sclerotized. Lateral thickening of end chamber very long, sinuous, reaching heel at base of ae- deagal pouch. Heel conspicuous, slightly curved upward, with apex subrounded. Fluted membrane prominent. Female. Head, thorax, legs and unmodified abdominal somites essentially as in male. Modified abdominal segments. Tergum VII ventrally expanded beyond longitudinal axis of abdomen, with 2 rows of bri- stles. Tergum VIII large, with sinuous posterior margin and bristles of different length. Sternum VII with posterior margin subtruncate, with several uneven bristles apparently in single row. Sternum VII broad but not conspicuous, with subangular caudal margin. Dorsal anal lobe and ventral anal lobe both with subtruncate apex and several bristles. Anal stylet about two times as long as maximum width, with 2 minute ventral bristles preceding long apical bristle. Spermatheca (Fig. 22C) with humped bulga well delimited from short, upturned hilla. Dorsal margin of bulga and apex of hilla respectively with short pro- jection. Length. Holotype, 2.96 mm; allotype, 3.31 mm. Remarks. — The trivial epithet of this taxon has been adopted from the Depto. del Cauca, where part of the type material was obtained. Quaest. Ent., 1977 13 (2) 138 Mendez Polygenis delpontei, new species (Fig. 23, 24, 25) Type material. — Holotype 6 and allotype 9(HTC-315) ex Oryzomys caliginosus, Colombia, Depto. del Valle, Municipio de Cali, Quebrada Honda near Pichinde', elevation 1800 m, 7. X. 1967, H. Trapido. Paratypes: 3 9 with same data as holotype; 1 (5(HTC-146) with same host, locality and collector as holotype but 13. VIII. 1965; 1<3, 1 9(HTC-212) with same data as holotype but 8.IX.1965; 1 9(HTC-213) ex Reithrodontomys mexicanus, other data as HTC-212; 1 <3(HTC- 243) with same data as HTC-212 but 16.IX.1965; 1 9(HTC-302 ) ex Oryzomys (Oligoryzomys) sp. (?) with same data as HTC-212 but 4.X.1965. Following paratypes with same data as holotype except date: 2 5 (HTC-317) 11. X. 1963; l9(HTC- 330) 19.X.1965; l9 (HTC-393) 17. XI. 1965; 2 6 (HTC-397) 18.XI.1965; l5(HTC-429) with same data as holotype but 29.XI.1965; 1 9(HTC-454) with same data as holotype but 14.XII.1965; 1 5(HTC-1307) with same data as holotype but 13.XII.1966, 2 (5(00007) same host, Rincon del Yarumal, Pichinde', Depto. del Valle, 21-25. I. 1974, M. Thomas & L. Vel- a'squez; 15(00010), La Buitrera, Depto. del Valle, 7. II. 1974, E. Mendez & L. Velasquez; ex Thomasomys fuscatus, 1(5, (HTC-2955), Rincdn del Yarumal, Pichinde', Depto. del Valle, 8.V.1969, H. Trapido, id, with same host and locality but 28.V.1969; l9 (00064), Saladito (Km 12), 2000 m, Depto. del Valle, 17.11.1974, E. Mdndez & L. Vela'squez; l5(HTC-805) with same data as holotype but elevation 1700 - 1900 m, 5. II. 1966; l5(HTC-1317) with same data as HTC-605 but 13. XII. 1966. Holotype and allotype are in the U.S. National Museum of Natural History. Paratypes are in the collections of the following institutions and specialists; British Museum (Natural History), Universidad del Valle, Colombia, Gorgas Memorial Laboratory, Robert Traub and Phyllis T. Johnson. Diagnosis. — Polygenis delpontei is closely related to P. brachinus Jordan. It differs from this species primarily in that it has the terminal portion of the aedeagal sclerotized inner tube rod-like and upturned. In P. brachinus the terminal portion of the sclerotized inner tube is flat and bent downward. Description. — Male. Head (Fig. 23A). Frons evenly rounded. Frontal tubercle barely projected out of margin, sub- rounded by wide sclerotized area. Preantennal region profusely covered with micropores. Preantennal row of 7 unequal bri- stles evenly spaced. Preocular row of 3 medium size bristles and 2 or 3 secondary short bristles. Premarginal bristles of anten- nal fossa weak, inconspicuous. Oral angle acute but not prominent. Genal lobe subangular. Eye oval, with small ventral in- dentation and moderate pigmentation. Post antennal region essentially with 3 rows but posteriormost bristle somewhat dis- placed, thus suggesting 4th row. Thorax. Pronotum and mesonotum each with 2 rows of bristles, last row with intercalaries. Mesepisternum somewhat re- ticulated, with 2 bristles. Mesepimeron with 3 bristles. Metanotum with 4 rows of bristles; anterior row reduced to about 3 or 4 bristles. Flange of metanotum with 9 to 11 spinelets. Legs. Typical of genus. Metatibia with posteromarginal notches with strong bristles arranged from upper to lower as fol- lows: 2-2-2- 3-2-3. Modified abdominal segments (Fig. 23C). Tergum VIII small, with single row of bristles. Sternum VIII relatively large, caudally in form of subangular lobe, with 1 row of 5 to 7 unequal bristles. Caudal flaps of sternum VIII opened at 0.33 of ventral margin. Dorsal and anal lobes of proctiger as in other Polygenis. Fixed process of clasper (Fig. 23B) with subangular apical lobe. Dorsal margin barely sinuate, ventral and posterior margins strongly sinuate but without true indentations. Dor- sal area of fixed process very setose; rest of bristles of this structure scantly distributed over ventral, posterior and inner ar- eas. Movable process of clasper (Fig. 23B) slightly shorter than distal arm, with broad apex, usually of 3 lobes. Distal arm of sternum IX (Fig. 23A) curved cephalad, with broad base, but with about equal width throughout most of its length, numer- ous bristles oriented posteriorly. Heel of sternum IX small, with short slender terminal tendon. Apodeme of aedeagus (Fig. 24B) with very broad portion before subrounded apex. Terminal portion of aedeagus (Fig. 24B) with prominent rounded dis- tolateral lobes. Median dorsal lobe with posterior semiangular projection. Crochet indistinct. Lateral lobe broadly convex. Dorsal area of terminal portion of aedeagus with distinct striated section facing apical portion of sclerotized inner tube. Lat- eral thickening of end chamber sinuous, ended almost at level with basal segment of sclerotized inner tube. Fender long, sin- uous and slender. Basal portion of sclerotized inner tube much shorter than apical portion of this tube, with large foramen on posterior half. Ribs scant, with irregular distribution. Crescent sclerite short, sinuous. F ulcral latero-ventral lobe curved caudad, with rounded apex. Heel at base of aedeagal pouch subacute, connected with apodemal rod. Vesicle ventrally ex- panded into thick ridge of irregular shape. Final section of this ridge apparently receiving anterior portion of fluted mem- brane. Penis rod coiled, ending on thick nob-like portion. Fluted membrane moderately developed. Female. Head (Fig. 25A). Frons more evenly rounded than in male. Thoracic structures and legs similar to those of male. Tergum I with 2 or 3 rows of bristles and posteromarginal row of spinelets. Other unmodified abdominal somites as in male. Modified abdominal segments (Fig. 25B). Sternum VIII moderately developed, extended beyond longitudinal axis of ab- domen, with 2 rows of bristles and single antepygidial bristles. Tergum VIII long, with posterior margin somewhat angular, with several posteromarginal and submarginal bristles preceded by long irregular row of bristles. Sternum VII with postero- caudal margin abruptly truncate, not incised, with combination of long and short bristles. Sternum VIII reduced, caudally Mammalian-Siphonapteran Associations 139 truncate. Sternum IX short, with posterior margin straight or barely arched. Dorsal anal lobe of proctiger (Fig. 25C) with truncate apex and prominent apical and subapical bristles. Anal stylet with 2 minute ventral bristles before long apical bris- tle. Spermatheca (Fig. 25 B, D) with cribose bulga clearly separated from short hilla with terminal portion upturned, dilated, semi-globular. Length. Holotype, 2.04 mm; allotype, 2.01 mm. Remarks. — This species is dedicated to the memory of the late Dr. Eduardo Del Ponte, whose excellent work on Siphonaptera and other blood-sucking insects contributed to the foundation of medical entomology in South America. Polygenis dunni (Jordan & Rothschild) (Figure 26) Rhopalopsyllus dunni Jordan & Rothschild, 1922, Ectop., 1:269, Fig. 261, 262. Remarks. - Although this species has not been yet recorded from Colombia, there is a strong possibility that it occurs in this country. It is presently known from Panama, Venezuela and Trinidad, and parasitizes an assemblage of hosts (see Tipton and Mendez, 1966 and Tip- ton and Machado- Allison, 1972). Polygenis hopkinsi, new species (Fig. 27, 28) Type material. — Holotype male ex Oryzomys albigularis (HTC-1838), Cerro Munchique (60 kms by road west of Popayin); Pena del Cerro, elevation 2160 m, Departamento del Cauca, Colombia, 11.V.1967, H. Trapido. The holotype is in the U.S. National Museum of Natural History. Diagnosis. — Males of Polygenis hopkinsi are similar to those of P. litargus Jordan & Roths- child, from which they are readily differentiated by having the terminal portion of the scler- otized inner tube bent upward. In P. litargus males, this structure ends barely sinuate and or- iented cephalad. Description. — Male. Head (Fig. 27A). Frons rounded, its contour not interrupted by unpronounced clypeal tubercle. Preantennal area above clypeal tubercle with micropores. Preantennal row of bristles of 5 bristles, one near antenna exceeds others in size. Preocular row of 3 principal long bristles and short secondary bristles separated from displaced minute bristle inserted near ventral margin. Eye subovate, moderately pigmented, with deep ventral indentation. Tentorium with arched anterior arm preceding principal stem. Oral angle short, acuminate. Tip of maxilla extended to anterior portion of last seg- ment of maxillary palpus. Genal angle subacuminate. Postantennal region with 3 rows of gradually increased number of bri- stles. Micropores limited to space between antennal base and first row of bristles. Group of short and thin bristles located near scape of antenna. Thorax. Mesonotum with 3 rows of bristles, first row reduced, of 2 or 3 bristles. Mesepisternum with 2 bristles. Mesepi- meron with 3 bristles. Lateral metanotal area with 2 large bristles preceded by 2 short bristles. Metepisternum with single long bristle and group of short bristles on anteromarginal projection. Legs. As in other Polygenis. Metatibia with strong bristles inserted in notches as follows: 2-2-2-3-2-3. Abdomen. Tergum I with 3 rows of bristles, first row reduced to about 4 bristles. Remaining unmodified terga with 2 rows of bristles. Sternum I with scattered ventral, subventral and inner bristles. Other unmodified sterna with single row of bristles. Modified abdominal segments (Fig. 27C). Tergum VIII short but extended to abdominal axis. Sternum VIII large, with truncate caudomarginal expansion, its ventral division clearly posterior to row of bristles. Fixed process of clasper (Fig. 27B) with dorsal margin moderately sinuate, gradually elevated toward apical portion. Posterior margin of fixed process very un- dulate, with short semiangular protrusion proximad to fovea of fixed process. Ventral margin deeply indented. Bristles of fixed process as illustrated. Movable finger (Fig. 27B) slightly bent upwards, not extended to apex of fixed process of clas- per, clothed with several marginal, submarginal and inner bristles. Distal arm of sternum IX (Fig. 28A) semi-falcate, almost as long as proximal arm, with anterior margin devoid of bristles and posterior margin with bristles of moderate size distri- buted over half its length. Heel of sternum IX heavily sclerotized, with relatively short tendon. Distolateral lobes of aedeagus (Fig. 28B) with anterior portion subrounded and posterior portion obtuse, prolonged into large ventromarginal extension. Median dorsal lobe with subangular apical portion. Apicomedian sclerite of aedeagus striated. Crochet imperceptible. Later- al lobe undulate, with distinct superior subangular projection. Dorsal area of terminal portion of aedeagus with small stria- ted section facing junction of apical and basal portions of sclerotized inner tube. Lateral thickening of end chamber sinuous, ended at level of vesicle. Lateroventral sclerite sickle-shaped but with rounded tip. Heel at base of aedeagal pouch semi-acum- inate, attached to apodemal rod. Fender semi-arched, slender and elongate. Apical portion of sclerotized inner tube with 2 Quaest. Ent., 1977 13 (2) 140 Mendez loops, its terminal section wide, upturned, oriented cephalad, with semiglobular apex. Basal portion of sclerotized inner tube with mesal foramen of moderate size. Ventral nodular section of basal portion large and prominent. Ribs distributed from area near vesicle to fender. Crescent sclerite arched, not prominent. Side pieces semi-triangular, with opposite ends acuminate. Length. Holotype, 2.31 mm. Remarks. — This species is named for the late G.H.E. Hopkins in recognition of the outstand- ing contributions he made to the systematics of Siphonaptera and Anoplura. Polygenis klagesi (Rothschild) (Figure 29) Pulex klagesi Rothschild, 1904, Novit. Zool., 11:620, PI. 9, Fig. 28; PI. 10, Fig. 35, 39. Material examined. — Ex Proechimys semispinosus. Departo. del Valle - <5, Rio Raposo, XI. Ex Hoplomys gymnurus. Depto. del Valle - 11(5, 11? , Alto Anchicaya', 650 m., II. In addition to material from the southwest part of Colombia, I have examined 1 1 males and 17 females from Carimagua, Depto. del Meta, 8 males and 1 1 females from the Depto. de Antio- quia and large series of males and females from Depto. del Choco. Remarks. - Our specimens of Polygenis klagesi taken on Hoplomys gymnurus from Alto Anchicaya, Depto. del Valle, and Curiche, Depto. del Choco, do not agree in certain features with specimens taken on Proechimys semispinosus which we are tentatively interpreting as typ- ical Polygenis klagesi samuelis. In view of the considerable amount of variation displayed by P. klagesi , it is advisable not to try to segregate Colombian populations into subspecies until an adequate study of material obtained throughout the geographical range of these fleas is done. Polygenis klagesi is presently known from Brazil, Panama, Costa Rica, Colombia, Venezuela, Trinidad and Ecuador, from sea level to altitudes below 900 meters. The P. klagesi complex occurs on a spectrum of hosts; however, these fleas are more natur- ally associated with the spiny rat family Echimyidae, particularly with Proechimys. Polygenis pradoi (Wagner) (Figure 30) Rhopalopsyllus pradoi Wagner, 1937, Zeits. Parasit., 9:420, Fig. 4. Material examined. — Ex Oryzomys albigularis. Depto. del Valle, Municipio de Cali — 2(5, 29, Pichindd(La Esperanza), 1900 m., I, VIII. Ex Oryzomys caliginosus. Depto. del Valle, Municipio de Cali - 39, Quebrada Honda, nr. Pichinde', 1800 m., IX, X; 9(5, 189, Valle del Rio Pichindd, 1700- 1900 m„ I, III, VI-XI;4(5, 1 1 9 , Pichindd, 1780-1900 m„ X. Ex Rhipidomys latimanus. Depto. del Valle, Municipio de Cali — (5, Pichindd, 1900 m., XI. Ex Thomasomys fuscatus. Depto. del Valle, Municipio de Cali - 9, Valle del Rio Pichindd, 1700 - 1900 m., XI. Remarks. — Polygenis pradoi has been reported from Brazil. Our specimens from southwes- tern Colombia represent the first records of this flea species for the country. Oryzomys (Mel- anomys) caliginosus , probably the commonest rodent in southwestern Colombia, stands out as the more favored host in this territory. Of 48 specimens of P. pradoi obtained, 42 were from that host while four were taken on Oryzomys albigularis , one on Rhipidomys latimanus and one on Thomasomys fuscatus. The reports of this species from Brazil concern the following hosts: Nasua socialis, Didelphis cancrivora, Oryzomys physodes, Akodon sp. (possibly cursor ), a wild rat and wild mouse (Johnson, 1957). Polygenis roberti beebei (I. Fox) (Figure 31) Rhopalopsyllus beebei I. Fox, 1947, Zool: N.Y. Zool. Soc., 32:117 , Fig. 2. Mammalian-Siphonapteran Associations 141 Remarks. — Polygenis roberti beebei has not been yet collected in the southwest corner of Colombia; nonetheless, we consider its presence there as probable. This taxon has not been re- ported from Colombia in the literature; however, we have examined 36 specimens (15 males and 21 females) from the Departamento de Antioquia, kindly loaned by Dr. V.J. Tipton. The zoogeography of P. roberti beebei presently involves Venezuela, Peru, Colombia, Trini- dad and Panama. It has been collected at low and moderate altitudes. The normal hosts of this flea are marsupials and rodents, notably some members of the cricetine genus Oryzomys. Polygenis thurmani Johnson (Figure 32) Polygenis thurmani Johnson, 1957, Mem. Ent. Soc. Wash. 5:169-170, Pis. 84, 85. Material examined. — Ex Didelphis marsupialis. Depto. del Valle - 6, 29, Lago Calima, 1450 m., II. Ex Oryzomys albigularis. Depto. del Valle, Municipio de Cali - (5, Quebrada Norte, Finca La Flora, Pichindd, 1900 m., VIII; 6, La Esperanza, Pichindd, VIII; <3, 29, Quebrada Honda, nr. Pichindd, 1800 m., X; 3d, 9, Pichindd, 1600 m., I. Ex Oryzomys species (probably O. caliginosus). Depto. del Valle, Municipio de Cali - 9 , Quebrada Honda, nr. Pichindd, 1800 m., XII. Ex Rhipidomys latimanus. Depto. del Valle, Municipio de Cali - d, Valle del Rio Pichindd, 1700 - 1900 m., VII; 9, La Cumbre, Finca La Maria, 1500 m., XI. Remarks. — Polygenis thurmani is now known from Peru and Colombia. The vertical distri- bution indicated by our material from southwestern Colombia fluctuates from 1450 meters to 1900 meters elevation. Our present information shows that in every one of these countries the hosts favored by P. thurmani are different. In Peru this flea has been taken from Akodon pulcherrimus inambari, Phyllotis phaeus, Oryzomys stolzmanni or Oxymycteris p. nigrifrons. Our Colombian specimens are from Didelphis marsupialis, Oryzomys albigularis, Oryzomys sp. (probably O. caliginosus ), and Rhipidomys latimanus. The data on this species is still in- adequate for drawing any conclusions on host preference. However, the premise that Oryzo- mys albigularis is the typical host might be reasonably founded on the basis of our collection: ten of 16 specimens of P. thurmani were found on this rodent. Polygenis trapidoi, new species (Fig. 33, 34) Type material. — Holotype d(HTC-1236) from Oryzomys caliginosus, Colombia, Depto. del Valle, Municipio de Cali, Valle del Rio Pichindd, elevation 1700 - 1900 m, 31. X. 1966, H. Trapido. Paratypes as follows: ldwith same data as holotype: ld(HTC-1629) with other data as holotype but 31. III. 1967, ld(HTC-1603), represented by mounted genitalia only, with same data as holotype but 17. III. 1967; ld(00005), same host and locality but 21-25.1.1974, M. Thomas & L. Vel- asquez; ld(00007), same data as 00005, 1(5(00008), same data as (00005). The holotype male is in the U.S. National Museum of Natural History. One paratype male is in the British Museum (Natural History) and the other paratype males remain in the Gorgas Memorial Laboratory’s collection. Diagnosis. — Near P. dunni Jordan & Rothschild from which it is separable by the subangu- lar caudomarginal expansion of sternum VIII. Other diagnostic structures are contained in the genitalia illustrated. Description. — Male. Head (Fig. 33A). Frontal tubercle not exserted. Micropores dispersed from area before falx to proximity of frontal tubercle. Preantennal row of 5 bristles, one near antenna longest. Preocular row with 3 prominent bris- tles and about equal number of minute bristles. Oral angle acuminate. Eye moderately pigmented, with profound ventral sin- us. Tentorium well defined, with anterior stem strongly arched. Maxillary lobe acuminate, extended to 4th segment of max- illary palpus. Genal lobe definitely angular. Post antennal area with 3 rows of bristles and group of minute bristles close to antennal scape. Pronotum wide, its posterior margin extended beyond level of proepimeron. Thorax. Pronotum and mesonotum each with 2 rows of bristles. Metanotum having 3 rows of bristles. Lateral metanotal area with 3 short anterior bristles followed by 2 large posterior bristles. Metepisternum with single bristle near pleural arch. Metepimeron with 6 bristles distributed in 2 rows. Quaest. Ent., 1977 13 (2) 142 Mendez Legs. Metatibia with 6 posteromarginal notches with strong bristles as follows: 2-2-2-3-2-3. Modified abdominal segments (Fig. 33C). Tergum VIII small but. extended to abdominal axis. Sternum VIII with poster- ior margin undulant but entire, with appreciably expanded subangular posterocaudal portion, having 10 bristles of diverse sizes oriented on mesal area. Fixed process of clasper (Fig. 33B) with subangular posterolateral elevation. Posterior margin of fixed process with angular prominence at about 0.25 of distance from top. Ventral margin notably undulate; distribution of bristles as illustrated. Movable process of clasper (Fig. 33B) elongate but not extended to apical portion of fixed process of clasper; barely wider on medial region with lateral margins slightly sinuate, with bristles scattered along margins and inner areas. Proximal arm of sternum IX (Fig. 33C) shorter than distal arm, constricted before bilobed apex. Distal arm of sternum IX (Fig. 34A) elongate, slightly curved upward, with numerous marginal and submarginal bristles of various sizes. Basal spur of sternum IX projected, with thick and highly sclerotized basal portion, provided with slender tendon. Distolateral lobes of terminal portion of aedeagus (Fig. 34B) somewhat falcate. Median dorsal lobe angular. Apico-median sclerite of aedeagus re- duced and striated. Crochet indistinct. Lateral lobe wide, irregularly sinuated. Fender reduced and arched. Apical portion of inner tube slightly larger than basal portion of sclerotized inner tube, with coiled section of 2 or 3 loops, ended in free un- dulated portion directed upward. Basal portion of sclerotized inner tube compact, with most of superomarginal area straight, with very narrow foramen almost limited to anterior half but extended beyond mid point into fraction of posterior half. Ven- tral nodular section of this basal portion small. Ribs very numerous, on most of anterior section of terminal portion of aedea- gus. Crescent sclerite sinuated, inconspicuous. Side pieces elongate, sinuous, with 2 opposite acuminate extremes. Latero- ventral sclerite represented by curved knob-like structure with broad obtuse apex. Heel at base of aedeagal pouch semi-fal- cate, very prominent. Fluted membrane well developed. Length. Holotype, 2.04 mm. Remarks. — I take great pleasure in dedicating this species to Dr. Harold Trapido in appre- ciation for his kind collaboration and his contributions to the knowledge of ectoparasites. Scolopsyllus colombianus Mendez (Fig. 35, 36) Scolopsyllus colombianus Mdndez, 1968, J. Med. Ent. 5:405-410, Fig. 1-14. Material examined. — Ex Oryzomys caliginosus. Depto. del Valle, Municipio de Cali - dholotype, 9 allotype, Quebrada Honda, nr. Pichindd, 1800 m, IX; 9 paratype, same locality, XII; 29 paratypes, Valle del Rio Pichindd, 1700 - 1900 m., HI, V. Ex Oryzomys alfaroi. Depto. del Valle, Municipio de Cali — (5 paratype, Valle del Rio Pichindd, XI. Ex Didelphis marsupialis. Depto. del Valle, Municipio de Cali - Pichindd, 1600 m., VIII. Remarks. — Scolopsyllus colombianus is probably endemic to subtropical mountains of moderate elevations surrounding the Cauca Valley. The preferred host appears to be Oryzom- ys caliginosus , which is the most abundant and widespread rodent in the Departamento del Valle. Rhopalopsyllus australis tupinus Jordan & Rothschild (Figure 37) Rhopalopsyllus australis tupinus Jordan & Rothschild, 1923, Ectop., 1:328, Fig. 339. Material examined. — Ex Eira barbara. Depto. del Valle, Municipio de Cali - (3, Pichinde', VI. Remarks. — Data gathered in Panama (Tipton and Mendez, 1966) indicated that R. austra- lis tupinus is normally found in association with the caviomorph rodent species Dasyprocta punctata and Agouti paca, which are the more selected hosts. The Collared Peccary, Tayassu tajacu, and the White-lipped Peccary, Tayassu pecari , follow those rodents in host preference. Other animals such as Didelphis marsupialis, Chironectes minimus, Proechimys semispinosus and other rodents, are also favored to a lower degree. Occasional hosts are carnivores like Eira barbara, Galictis allamandi, Canis familiaris, and others, that are infested by fleas obtain- ed from their prey. Rhopallopsyllus australis tupinus has not been previously reported from Colombia. We have also examined one male and five females from Carimangua, Departamento del Meta. This spe- cies is known from Panama, Bolivia, Brazil and Peru. At least in Panama, specimens of this taxon have been collected from sea level to elevations close to 1600 meters. Mammalian-Siphonapteran Associations 143 Fig. 20. Polygenis bohlsi bohlsi (Wagner). Male. A. Head, prothorax and procoxa; B. Process and movable finger of clasper; C. Apex of aedeagus. Female. D. Distal arm of 9th sternum; E. Spermatheca. Quaes t. Ent., 1977 13 (2) 144 Mendez Fig. 21. Polygenis caucensis, n. sp. Male. A. Head and prothorax; B. Process and movable finger of clasper; C. Modified abdo- minal segments. Mammalian-Siphonapteran Associations 145 Fig. 22. Polygenis caucensis, n. sp. Male. A. Distal arm of 9th sternum; B. Apex of aedeagus. Female. C. Spermatheca. Quaest. Ent., 1977 13 (2) 146 Mendez Fig. 23. Polygems delpontei, n. sp. Male. A. Head and prothorax; B. Process and movable finger of clasper; C. Modified abdo- minal segments. Mammalian-Siphonapteran Associations 147 Fig. 24. Polygenis delpontei, n. sp. Male. A. Distal arm of 9th sternum; B. Apex of aedeagus. Quaest. Ent., 1977 13 (2) 148 Mendez B D Fig. 25. Polygenis delpontei, n. sp. Female. A. Head and prothorax; B. Modified abdominal segments; C. Dorsal and anal lobes of proctiger; D. Spermatheca. Mammalian-Siphonapteran Associations 149 Fig. 26. Polygenis dunni (Jordan & Rothschild). Male. A. Head, prothorax and procoxa; B. Genitalia. Female. C. Spermatheca and 7th abdominal segment. From “The Fleas (Siphonaptera) of Panama” by Tipton and Mdndez, in “Ectoparasites of Pana- ma”, Field Museum of Natural History, Chicago (1966). Quaest. Ent., 1977 13 (2) 150 Mendez Fig. 27. Polygenis hopkinsi, n. sp. Male. A. Head and prothorax; B. Process and movable finger of clasper; C. Modified abdo- minal segments. Mammalian-Siphonapteran Associations 151 Fig. 28. Polygenis hopkinsi, n. sp. Male. A. Distal arm of 9th sternum; B. Apex of aedeagus. Quaest. Ent., 1977 13 (2) /