A WORLD REVIEW OF RONDANI (DIPTERA;

SPHAEROCERIDAE) WITH A REVISION OF THE NEW WORLD SPECIES

A Thesis

Presented to

The Faculty of Graduate Studies

of

The University of Guelph

by

MATTHEW DAVID BERGERON

In partial fulfilment of requirements

For the degree of

Master of Science

December, 2009

© Matthew Bergeron, 2009 Library and Archives Bibliotheque et 1*1 Canada Archives Canada Published Heritage Direction du Branch Patrimoine de I'edition 395 Wellington Street 395, rue Wellington OttawaONK1A0N4 Ottawa ON K1A0N4 Canada Canada

Your file Votre reference ISBN: 978-0-494-58389-0 Our file Notre reference ISBN: 978-0-494-58389-0

NOTICE: AVIS:

The author has granted a non­ L'auteur a accorde une licence non exclusive exclusive license allowing Library and permettant a la Bibliotheque et Archives Archives Canada to reproduce, Canada de reproduire, publier, archiver, publish, archive, preserve, conserve, sauvegarder, conserver, transmettre au public communicate to the public by par telecommunication ou par Nnternet, preter, telecommunication or on the Internet, distribuer et vendre des theses partout dans le loan, distribute and sell theses monde, a des fins commerciales ou autres, sur worldwide, for commercial or non­ support microforme, papier, electronique et/ou commercial purposes, in microform, autres formats. paper, electronic and/or any other formats.

The author retains copyright L'auteur conserve la propriete du droit d'auteur ownership and moral rights in this et des droits moraux qui protege cette these. Ni thesis. Neither the thesis nor la these ni des extraits substantiels de celle-ci substantial extracts from it may be ne doivent etre imprimes ou autrement printed or otherwise reproduced reproduits sans son autorisation. without the author's permission.

In compliance with the Canadian Conformement a la loi canadienne sur la Privacy Act some supporting forms protection de la vie privee, quelques may have been removed from this formulaires secondaires ont ete enleves de thesis. cette these.

While these forms may be included Bien que ces formulaires aient inclus dans in the document page count, their la pagination, il n'y aura aucun contenu removal does not represent any loss manquant. of content from the thesis.

•+• Canada ABSTRACT

A WORLD REVIEW OF COPROICA RONDANI (DIPTERA; ) WITH A REVISION OF THE NEW WORLD SPECIES

Matthew David Bergeron Advisor: University of Guelph, 2009 Professor S.A. Marshall

Coproica Rondani, 1861 (Diptera: Sphaeroceridae) is here redefined and the New

World species revised on the basis of over 16,000 examined specimens. The genus is divides into three major clades. Eight new species (C bifurcata, C. bispatha, C. brachystyla, C. diabolica, C. emarginata, C. galapagos, C. novacula, and C. testudined) are described from the New World, and redescriptions are provided for 16 additional species. Included are two keys (one for the New World species and another to cover all species), reviews of all species in the genus, expanded New World distribution records, and illustrations of male and female genitalic structures. Table of Contents

Introduction 1 Introduction 1 Natural History of Coproica (Biology and Behaviour) 3

Materials and Methods 7 Sources of Material 7 Specimens Examined 9 Adult Morphological Characters and Terminology 11 Measurements and Ratios 12 Phylogenetic Approach 13

Phylogeny 15 Phylogeny of Coproica 15 List of Characters 20 Discussion 24

Taxonomy 28 Discussion of Generic Nomenclature 28 Generic Diagnosis 30 Generic Description 32 Key to New World Species 37 Key to World Species 41

Species Accounts 48 (Zetterstedt, 1847) 48 Coproica albiseta Papp, 2008 56 Coproica aliena Papp, 2008 56 Coproica bifurcata sp. nov 57 Coproica bispatha sp. nov 60 Coproica bispinosa Papp, 2008 64 Coproica brachystyla sp.nov 65 Coproica brevivenosa Papp, 2008 69 Coproica cacti (Richards, 1960) 70 Coproica coreanaFapp, 1979 72 Coproica demeteri Papp, 2008 75 Coproica dentataPapp, 1979 76 Coproica diabolica sp. nov 78 Coproica digitata (Duda, 1918) 83 Coproica emarginata sp. nov 86 (Stenhammer, 1855) 91 Coproicaflavifacies Papp, 2008 98 Coproica galapagos sp. nov 99 Coproica ghanensis Papp, 1979 101 Coproica hirticula Collin, 1956 102

i Coproica hirtula (Rondani, 1880) 109 Coproica hirtuloidea (Duda, 1925) 114 Coproica insulaepasqualis Enderlein, 1938 120 Coproica lacteipennis Hayashi, 2005 120 Coproica lugubris (Haliday, 1835) 121 Coproica microps Papp, 2008 124 Coproica mitchelli (Malloch, 1913) 125 Coproica novacula sp. nov 131 Coproicapappi Carles-Tolra, 1990 134 Coproicaperlugubris Papp, 2008 136 Coproicapseudolacteipennis Papp, 2008 136 Coproicapusio (Zetterstedt, 1847) 137 Coproica rohaceki Carles-Tolra, 1990 140 Coproica rufifrons Hayashi, 1991 143 Coproica ruwenzoriensis (Vanschuytbroeck, 1950) 147 Coproica saprophaga Papp, 2008 147 Coproica serra (Richards, 1938) 148 Coproica setulosa (Duda, 1929) 150 Coproica testudinea sp. nov 154 Coproica thaii Papp, 2008 157 Coproica unispinosa Papp, 2008 157 Coproica urbana (Richards, 1960) 158 Coproica vagans (Haliday, 1833) 166

Works Cited 171

Matrix (Tables) 176

Figures 178

n List of Tables

Table 1: Character state matrix (characters 1 -25) 176 Table 2: Character state matrix (characters 26 - 50) 177

111 List of Figures

Figurel: Phylogeny 178 Figure 2: Jacknife supports 179 Figure 3: Bootsrap supports 179 Figure 4: Bremer supports 179 Figure 5: Coproica acutangula left lateral 180 Figure 6: Coproica acutangula scutellum 180 Figure 7: Coproica acutangula head, dorsal 181 Figure 8: Coproica diabolica katepisternum 181 Figure 9: Coproica ferruginata katepisternum 181 Figure 10: Coproica novacula katepisternum 182 Figure 11: Coproica setulosa katepisternum 182 Figure 12: Coproica rufifrons katepisternum 182 Figure 13: Coproica vagans katepisternum 182 Figure 14: Coproica acutangula fore basitarsus 183 Figure 15: Coproica acutangula hind basitarsus 183 Figure 16: Coproica acutangula mid tibia, dorsal 183 Figure 17: Coproica bifurcata mid tibia, dorsal 183 Figure 18: Coproica ferruginata mid tibia, dorsal 183 Figure 19: Coproica lugubris mid tibia, dorsal 183 Figure 20: Coproica rufifrons mid tibia, dorsal 183 Figure 21: Coproica bifurcata mid basitarsus, ventral 184 Figure 22: Coproica brachystyla mid basitarsus, ventral 184 Figure 23: Coproica hirticula mid basitarsus, ventral 184 Figure 24: Coproica lugubris mid basitarsus, ventral 184 Figure 25: Coproica mitchelli mid basitarsus, ventral 184 Figure 26: Coproica novacula mid basitarsus, ventral 184 Figure 27: Coproica rufifrons mid basitarsus, ventral 184 Figure 28: Coproica setulosa mid basitarsus, ventral 184 Figure 29: Coproica acutangula male left wing 185 Figure 30: Coproica acutangula female left wing 185 Figure 31: Coproica acutangula epandrium, posterior 186 Figure 32: Coproica acutangula epandrium, left lateral 186 Figure 33: Coproica acutangula aedeagus and associated structures 187 Figure 34: Coproica acutangula sternite 5 and synsternite 6+7 ...187 Figure 35: Coproica acutangula left postgonite 187 Figure 36: Coproica acutangula female terminalia, dorsal ; 188 Figure 37: Coproica acutangula female terminalia, left lateral 188 Figure 38: Coproica acutangula female terminalia, ventral 188 Figure 39: Coproica acutangula female sternite 8 188 Figure 40: Coproica acutangula spermathecae 188 Figure 41: Coproica bifurcata male left wing 189 Figure 42: Coproica bifurcata epandrium, left lateral 189 Figure 43: Coproica bifurcata epandrium, posterior 189 Figure 44: Coproica bifurcata left postgonite 190

IV Figure 45: Coproica bifurcata right postgonite 190 Figure 46: Coproica bifurcata aedeagus and associated structures 190 Figure 47: Coproica bifurcata sternite 5 and synsteraite 6+7 190 Figure 48: Coproica bifurcata female terminalia, dorsal 191 Figure 49: Coproica bifurcata female terminalia, ventral 191 Figure 50: Coproica bifurcata female terminalia, left lateral 191 Figure 51: Coproica bifurcata female sternite 8 191 Figure 52: Coproica bifurcata spermathecae 191 Figure 53: Coproica bispatha male left wing 192 Figure 54: Coproica bispatha epandrium, left lateral 192 Figure 55: Coproica bispatha epandrium, posterior 192 Figure 56: Coproica bispatha left postgonite 193 Figure 57: Coproica bispatha right postgonite 193 Figure 58: Coproica bispatha aedeagus and associated structures 193 Figure 59: Coproica bispatha sternite 5 and synsternite 6+7 193 Figure 60: Coproica bispatha female terminalia, left lateral 194 Figure 61: Coproica bispatha female terminalia, dorsal 194 Figure 62: Coproica bispatha female terminalia, ventral 194 Figure 63: Coproica bispatha spermathecae 194 Figure 64: Coproica bispatha female sternite 8 194 Figure 65: Coproica brachystyla male left wing, dorsal 195 Figure 66: Coproica brachystyla epandrium, left lateral 195 Figure 67: Coproica brachystyla epandrium, posterior 195 Figure 68: Coproica brachystyla left postgonite 196 Figure 69: Coproica brachystyla right postgonite 196 Figure 70: Coproica brachystyla aedeagus and associated structures 196 Figure 71: Coproica brachystyla sternite 5 and synsternite 6+7 196 Figure 72: Coproica brachystyla female terminalia, dorsal 197 Figure 73: Coproica brachystyla female terminalia, ventral 197 Figure 74: Coproica brachystyla female terminalia, left lateral 197 Figure 75: Coproica brachystyla female sternite 8 197 Figure 76: Coproica brachystyla spermathecae 197 Figure 77: Coproica cacti male left wing 198 Figure 78: Coproica coreana male left wing 198 Figure 79: Coproica coreana epandrium, left lateral 199 Figure 80: Coproica coreana epandrium, posterior 199 Figure 81: Coproica coreana left postgonite 200 Figure 82: Coproica coreana aedeagus and associated structures 200 Figure 83: Coproica coreana sternite 5 and synsternite 6+7 200 Figure 84: Coproica coreana female terminalia, dorsal 201 Figure 85: Coproica coreana female terminalia, ventral 201 Figure 86: Coproica coreana female terminalia, left lateral 201 Figure 87: Coproica coreana female sternite 8 201 Figure 88: Coproica coreana spermathecae 201 Figure 89: Coproica diabolica male left wing 202 Figure 90: Coproica diabolica epandrium, left lateral 202

v Figure 91: Coproica diabolica epandrium, posterior 202 Figure 92: Coproica diabolica left postgonite 203 Figure 93: Coproica diabolica right postgonite 203 Figure 94: Coproica diabolica aedeagus and associated structures 203 Figure 95: Coproica diabolica sternite 5 and synsternite 6+7 203 Figure 96: Coproica diabolica female terminalia, dorsal 204 Figure 97: Coproica diabolica female terminalia, ventral 204 Figure 98: Coproica diabolica female terminalia, left lateral 204 Figure 99: Coproica diabolica female sternite 8 204 Figure 100: Coproica diabolica spermathecae 204 Figure 101: Coproica digitata male left wing 205 Figure 102: Coproica digitata sternite 5 and synsternite 6+7 205 Figure 103: Coproica digitata epandrium left lateral 206 Figure 104: Coproica digitata epandrium, posterior 206 Figure 105: Coproica digitata left postgonite 207 Figure 106: Coproica digitata right postgonite 207 Figure 107: Coproica digitata aedeagus and associated structures 207 Figure 108: Coproica digitata spermathecae 207 Figure 109: Coproica digitata female sternite 8 207 Figure 110: Coproica digitata female, dorsal 208 Figure 111: Coproica digitata female, left lateral 208 Figure 112: Coproica emarginata left male wing 209 Figure 113: Coproica emarginata epandrium, posterior 209 Figure 114: Coproica emarginata epandrium, left lateral 209 Figure 115: Coproica emarginata left postgonite 210 Figure 116: Coproica emarginata right postgonite 210 Figure 117: Coproica emarginata aedeagus and associated structures 210 Figure 118: Coproica emarginata right surstylus 210 Figure 119: Coproica emarginata sternite 5 and synsternite 6+7 210 Figure 120: Coproica emarginata female terminalia, dorsal 211 Figure 121: Coproica emarginata female terminalia, ventral 211 Figure 122: Coproica emarginata female sternite 8 211 Figure 123: Coproica emarginata spermathecae 211 Figure 124: Coproica ferruginata male left wing 212 Figure 125: Coproica ferruginata epandrium, posterior 212 Figure 126: Coproica ferruginata epandrium, left lateral 212 Figure 127: Coproica ferruginata left postgonite 213 Figure 128: Coproica ferruginata aedeagus and associated structures 213 Figure 129: Coproica ferruginata sternite 5 and synsternite 6+7 213 Figure 130: Coproica ferruginata female terminalia, dorsal 214 Figure 131: Coproica ferruginata female terminalia, ventral 214 Figure 132: Coproica ferruginata female terminalia, left lateral 214 Figure 133: Coproica ferruginata spermathecae 214 Figure 134: Coproica galapagos male left wing 215 Figure 135: Coproica galapagos sternite 5 and synsternite 6+7 215 Figure 136: Coproica galapagos aedeagus and associated structures 215

VI Figure 137: Coproica galapagos left postgonite 215 Figure 138: Coproica galapagos epandrium, posterior 216 Figure 139: Coproica galapagos epandrium, left lateral 216 Figure 140: Coproica galapagos female terminalia, dorsal 217 Figure 141: Coproica galapagos female terminalia, ventral 217 Figure 142: Coproica galapagos female terminalia, left lateral 217 Figure 143: Coproica galapagos spermathecae 217 Figure 144: Coproica hirticula male left wing 218 Figure 145: Coproica hirticula epandrium, left lateral 218 Figure 146: Coproica hirticula epandrium, posterior 218 Figure 147: Coproica hirticula spermathecae 219 Figure 148: Coproica hirticula aedeagus and associated structures 219 Figure 149: Coproica hirticula sternite 5 and synsternite 6+7 219 Figure 150: Coproica hirticula left postgonite 219 Figure 151: Coproica hirticula female terminalia, dorsal 220 Figure 152: Coproica hirticula female terminalia, ventral 220 Figure 153: Coproica hirticula female terminalia, left lateral 220 Figure 154: Coproica hirticula female sternite 8 220 Figure 155: Coproica hirtula male left wing 221 Figure 156: Coproica hirtula epandrium, left lateral 221 Figure 157: Coproica hirtula epandrium, posterior 221 Figure 158: Coproica hirtula left postgonite 222 Figure 159: Coproica hirtula aedeagus and associated structures 222 Figure 160: Coproica hirtula sternite 5 and synsternite 6+7 222 Figure 161: Coproica hirtula female terminalia, dorsal 223 Figure 162: Coproica hirtula female terminalia, ventral 223 Figure 163: Coproica hirtula female terminalia, left lateral 223 Figure 164: Coproica hirtula spermathecae 223 Figure 165: Coproica hirtuloidea male left wing 224 Figure 166: Coproica hirtuloidea epandrium, left lateral 224 Figure 167: Coproica hirtuloidea epandrium, posterior 224 Figure 168: Coproica hirtuloidea left postgonite 225 Figure 169: Coproica hirtuloidea right postgonite 225 Figure 170: Coproica hirtuloidea aedeagus and associated structures 225 Figure 171: Coproica hirtuloidea sternite 5 and synsternite 6+7 225 Figure 172: Coproica hirtuloidea female sternite 8 225 Figure 173: Coproica hirtuloidea female terminalia, dorsal 226 Figure 174: Coproica hirtuloidea female terminalia, ventral 226 Figure 175: Coproica hirtuloidea female terminalia, left lateral 226 Figure 176: Coproica hirtuloidea spermathecae 226 Figure 177: Coproica lugubris male left wing 227 Figure 178: Coproica lugubris epandrium, left lateral 227 Figure 179: Coproica lugubris epandrium, posterior 227 Figure 180: Coproica lugubris aedeagus and associated structures 228 Figure 181: Coproica lugubris female sternite 8 228 Figure 182: Coproica lugubris spermathecae 228

vn Figure 183: Coproica lugubris left postgonite 228 Figure 184: Coproica lugubris female terminalia, left lateral 229 Figure 185: Coproica lugubris sternite 5 and synsternite 6+7 229 Figure 186: Coproica lugubris female terminalia, dorsal 230 Figure 187: Coproica lugubris female terminalia, ventral 230 Figure 188: Coproica mitchelli male left wing 231 Figure 189: Coproica mitchelli epandrium, left lateral 231 Figure 190: Coproica mitchelli epandrium, posterior 231 Figure 191: Coproica mitchelli left surstylus 232 Figure 192: Coproica mitchelli aedeagus and associated structures 232 Figure 193: Coproica mitchelli sternite 5 and synsternite 6+7 232 Figure 194: Coproica mitchelli left postgonite 232 Figure 195: Coproica mitchelli female terminalia, dorsal 233 Figure 196: Coproica mitchelli female terminalia, ventral 233 Figure 197: Coproica mitchelli female terminalia, left lateral 233 Figure 198: Coproica mitchelli female sternite 8 233 Figure 199: Coproica mitchelli spermathecae 233 Figure 200: Coproica novacula male left wing 234 Figure 201: Coproica novacula epandrium, left lateral 234 Figure 202: Coproica novacula epandrium, posterior 234 Figure 203: Coproica novacula female sternite 8 235 Figure 204: Coproica novacula aedeagus and associated structures 235 Figure 205: Coproica novacula sternite 5 and synsternite 6+7 235 Figure 206: Coproica novacula left postgonite 235 Figure 207: Coproica novacula female terminalia, dorsal 236 Figure 208: Coproica novacula female terminalia, ventral 236 Figure 209: Coproica novacula female terminalia, left lateral 236 Figure 210: Coproica novacula spermathecae 236 Figure 211: Coproicapappi male left wing 237 Figure 212: Coproica pappi female sternite 8 237 Figure 213: Coproica pappi aedeagus and associated structures 237 Figure 214: Coproica pappi sternite 5 and synsternite 6+7 237 Figure 215: Coproica pappi epandrium, left lateral 238 Figure 216: Coproica pappi epandrium, posterior 238 Figure 217: Coproica pappi left postgonite 238 Figure 218: Coproica pappi female terminalia, dorsal 239 Figure 219: Coproica pappi female terminalia, ventral 239 Figure 220: Coproica pappi female terminalia, left lateral 239 Figure 221: Coproicapappi spermathecae 239 Figure 222: Coproicapusio male left wing 240 Figure 223: Coproica pusio epandrium, left lateral 240 Figure 224: Coproica pusio epandrium, posterior 240 Figure 225: Coproica pusio left postgonite 241 Figure 226: Coproica pusio aedeagus and associated structures 241 Figure 227: Coproica pusio sternite 5 and synsternite 6+7 241 Figure 228: Coproica pusio female sternite 8 241

vm Figure 229: Coproicapusio female terminalia, dorsal 242 Figure 230: Coproica pusio female terminalia, ventral 242 Figure 231: Coproica pusio female terminalia, left lateral 242 Figure 232: Coproica pusio spermathecae 242 Figure 233: Coproica rohaceki male left wing 243 Figure 234: Coproica rohaceki epandrium, left lateral 243 Figure 235: Coproica rohaceki epandrium, posterior 243 Figure 236: Coproica rohaceki left postgonite 244 Figure 237: Coproica rohaceki aedeagus and associated structures 244 Figure 238: Coproica rohaceki sternite 5 and synsternite 6+7 244 Figure 239: Coproica rohaceki female terminalia, dorsal 245 Figure 240: Coproica rohaceki female terminalia, ventral 245 Figure 241: Coproica rohaceki female terminalia, left lateral 245 Figure 242: Coproica rohaceki spermathecae 245 Figure 243: Coproica rufifrons male left wing 246 Figure 244: Coproica rufifrons epandrium, left lateral 246 Figure 245: Coproica rufifrons epandrium, posterior 246 Figure 246: Coproica rufifrons left postgonite 247 Figure 247: Coproica rufifrons aedeagus and associated structures 247 Figure 248: Coproica rufifrons sternite 5 and synsternite 6+7 247 Figure 249: Coproica rufifrons female sternite 8 247 Figure 250: Coproica rufifrons female terminalia, dorsal 248 Figure 251: Coproica rufifrons female terminalia, ventral 248 Figure 252: Coproica rufifrons female terminalia, left lateral 248 Figure 253: Coproica rufifrons spermathecae 248 Figure 254: Coproica serra male left wing 249 Figure 255: Coproica setulosa male left wing 249 Figure 256: Coproica setulosa epandrium, left lateral 250 Figure 257: Coproica setulosa epandrium, posterior 250 Figure 258: Coproica setulosa left postgonite 250 Figure 259: Coproica setulosa spermathecae 251 Figure 260: Coproica setulosa aedeagus and associated structures 251 Figure 261: Coproica setulosa sternite 5 and synsternite 6+7 251 Figure 262: Coproica setulosa female sternite8 251 Figure 263: Coproica setulosa female terminalia, dorsal 252 Figure 264: Coproica setulosa female terminalia, ventral 252 Figure 265: Coproica setulosa female terminalis, left lateral 252 Figure 266: Coproica testudinea male left wing 253 Figure 267: Coproica testudinea epandrium, left lateral 253 Figure 268: Coproica testudinea epandrium, posterior 253 Figure 269: Coproica testudinea left postgonite 254 Figure 270: Coproica testudinea female sternite 8 , 254 Figure 271: Coproica testudinea aedeagus and associated structures 254 Figure 272: Coproica testudinea sternite 5 and synsternite 6+7 254 Figure 273: Coproica testudinea female terminalia, dorsal ..255 Figure 274: Coproica testudinea female terminalia, left lateral 255

IX Figure 275: Coproica testudinea spermathecae 255 Figure 276: Coproica urbana male left wing 256 Figure 277: Coproica urbana sternite 5 and synsternite 6+7 256 Figure 278: Coproica urbana aedeagus and associated structures 256 Figure 279: Coproica urbana female sternite 8 257 Figure 280: Coproica urbana epandrium, left lateral 257 Figure 281: Coproica urbana epandrium, posterior 257 Figure 282: Coproica urbana left postgonite 257 Figure 283: Coproica urbana right postgonite 257 Figure 284: Coproica urbana female terminalia, dorsal 258 Figure 285: Coproica urbana female terminalia, ventral 258 Figure 286: Coproica urbana female left lateral 258 Figure 287: Coproica urbana spermathecae 258 Figure 288: Coproica vagans male left wing 259 Figure 289: Coproica vagans epandrium, left lateral 259 Figure 290: Coproica vagans epandrium, posterior 259 Figure 291: Coproica vagans spermathecae 260 Figure 292: Coproica vagans distiphallus and basiphallus 260 Figure 293: Coproica vagans sternite 5 and synsternite 6+7 260 Figure 294: Coproica vagans female terminalia, dorsal 261 Figure 295: Coproica vagans female terminalia, left lateral 261

x Introduction

The cosmopolitan genus Coproica Rondani includes 43 species of small (0.9 to

2.4mm), generally dull brown distinguishable from most other sphaerocerids by the presence of setae on the scutellar disc (Fig: 6). Most species of Coproica are

associated with ungulate dung and are often spectacularly abundant on the dung of horses

and cows. Several species associated with the dung of domesticated have become secondarily widespread, and 6 of the 43 known species of Coproica are now

cosmopolitan. Other species are now recognized as occurring in widely separated areas;

for example Coproica rohaceki was described from the Mediterranean but is here newly recorded from Brazil, Argentina, and Chile.

The Coproica of the Old World are relatively well known, with 24 recognized

species, in contrast, only five New World species were formally described prior to this

study. There have been many regional studies specifically focused on Coproica in the

Old World (Carles-Tolra, 1990; Hayashi, 1991, 2005; Papp, 1973, 1979, 2008), but

relatively little attention has been given to the New World species. Malloch (1913)

described C. mitchelli from Texas, Duda (1925, 1929) described C. hirtuloidea and C. setulosa, respectively, from South America, and Richards (1960) reviewed Coproica and

described C. cacti and C urbana from the Nearctic region. An unpublished MSc thesis by Swann (1993) dealt with the New World species of Coproica, but because it was

incomplete and had several serious deficiencies it was left unpublished while the rest of

the Nearctic Sphaerocerid genera have been revised (as summarized in Rohacek et al.,

2001). The genus Coproica is now the only significant group in a Nearctic sphaerocerid

1 fauna of 42 genera and 312 species for which there is no modern revision or functional identification tools.

Richards (1960) included the only identification key to consider the Coproica species from all zoogeographic regions, and the only previously published key to New

World species. That paper omitted C. mitchelli (Malloch, 1913) since the paratypes

Richards examined were in poor condition and he was not certain whether the species belonged in Coproica or Rachispoda. Including those described here, 28 species of

Coproica have been described since Richards' revision (Papp, 1973, 1979, 2008; Carles-

Tolra, 1990; Hayashi, 1991, 2005). Papp (2008) revised the Old World species and included a key to the species of Coproica from regions other than the Nearctic and

Neotropical.

We here provide a key to the New World species as well as a key to all described species in the genus (C insulaepasqualis excepted), across all zoogeographic regions.

2 Natural History of Coproica (Biology and Behaviour)

Coproica adults are most abundant on feces from ungulates but some species have been recorded from dog dung, bear scats, poultry manure, pig feces, grass piles, decaying

fungus and various types of vertebrate and invertebrate carrion. Coproica species are

typically the first to arrive at fresh horse dung, showing up in as little as 1 or 2

minutes after defecation (Hafez, 1949). The adult are typically found either on the

surface of the dung or the grass and surfaces surrounding it (Lachmann, 1990).

Lachmann (1990) observed that males of C lugubris establish their territories on blades

of grass surrounding feces, and this was where most copulating pairs were found. Hafez

(1949) noted that Coproica would take refuge in tunnels in the dung made by scarab

beetles, probably seeking higher temperatures and humidity.

A few Coproica species can be found in very specific microhabitats. C.

testudinea sp. nov. has been reared from the contents of gopher tortoise burrows in

Florida, yet it has never been collected from pan traps outside the nest entrances.

Individuals of this species have reduced eyes, as is typical for many burrow or cave-

dwelling sphaerocerid species (Marshall and Peck, 1984, Papp and Plachter, 1976), but

are fully macropterous. The only records of C cacti (Richards, 1960) are reared

specimens from decaying cactus in the southwestern United States and Mexico (Richards,

1960).

Many Coproica species are synanthropic. This is especially evident for C. ferruginata, C. vagans and C. acutangula, that have been recorded as very abundant on

ranches and farms where large quantities of manure are stored (Hafez, 1949; Papp 1975,

3 1979, 1992). Many of the most common species on ungulate dung (C. ferruginata, C. acutangula, C. hirticula, C. vagans, and C. hirtula) are cosmopolitan in distribution.

Despite the high level of synanthropy amongst some very common Coproica species, they are rarely of any direct economic importance. Considering their habitat, it is possible that Coproica species may contaminate surfaces or cause human illness.

Greenberg et al. (1963) recorded two types of Salmonellae on Coproica species collected from a chicken processing plant in Mexico, and Hayashi (1986) placed eight species of

Coproica on his list of synanthropic sphaerocerids of hygienic importance in Japan.

Since Coproica species rarely far from the dung in which they breed they are unlikely to transmit disease organisms to human food.

There is little known about the feeding habits of adult Coproica, but they probably visit dung for adult feeding as well as mate location and oviposition. Many of the sphaerocerids found at feces breed elsewhere and visit dung only as adults, but

Coproica larvae develop in feces (Goddard, 1938; Okely, 1974). Like all Sphaeroceridae larvae, they are microbial grazers feeding on a mixture of decomposing material and microorganisms.

Lachmann (1990, 1996, 1997) documented the mating behaviour of several species in the genus. Courtship behaviours consist of a male approaching a female head to head. If the female moved, the male would match her movement, and in C. acutangula, C lugubris, and C. hirticula, the male would occasionally tap the female with his fore tarsi. The male would then mount the female and drum her abdomen with his mid and hind legs before and after achieving genital contact. Lachmann (1990, 1996) found that unreceptive females of C acutangula and C. lugubris either crossed their hind

4 legs over their abdomen or by "jumping around like a bucking horse until he dismounts"

(Lachman, 1990). The copulatory displays in Coproica are relatively short compared to some other Limosinines such as Chaetopodella scutellaris, in which males have a complex precopulatory visual display (Lachmann, 1990).

Lachmann's (1990, 1996, 1997) work showed that Coproica males utilize two differnt methods of insemination. The time spent in copula varies from 8.0 - 10.4 minutes in C. ferruginata to 29.0 - 38.6 minutes in C. lugubris, and that sperm transfer only occurs near the end of this period in C. hirticula, C. lugubris, C. ferruginata and C. vagans, yet occurs within the first 2 minutes in C. acutangula. Sperm is deposited either within the vagina of the female (C. ferruginata and C. vagans), or is directly pumped into the spermathecae (C. acutangula, C. hirticula, and C. lugubris). Species that deposit a sperm mass within the vagina of the female also deposit a mating plug post insemination.

The mating plug in these species reduces sperm competition by preventing the female from remating. Mating plugs were not deposited by males in the other three species studied by Lachmann. In all species the spermathecal ducts are surrounded by muscle and have a distal valve, so the female could feasibly control access of sperm to the spermathecae. This is important, because due to the lack of an elaborate precopulatory ritual much of the female's evaluation of the male appears to be based on internal male structures assessed during copulation (Lachmann, 1996, 1997).

The use of a sperm plug in C ferruginata and C. vagans and its absence in both the C. acutangula species group and C. hirticula suggest that it is a derived adaptation related to sperm transfer in Coproica. The presence of two different modes of insemination in Coproica suggests that the split between these modes has played a

5 significant role in the evolution of Coproica species. The use of a sperm plug appears to have been derived within the C. vagans species group (Fig: 1), although more complete information on the copulatory behaviours of the species in Coproica are required to support this hypothesis.

Coproica females will lay eggs on both fresh and old dung with a preference for fresh dung (Hafez 1939, 1949). Most eggs are forced into the dung by the female. The eggs of C. ferruginata have a pair of respiratory appendages projecting towards the surface (Okely, 1974). The eggs hatch in approximately 8 hours at 28°C and remain in the larval stage for 3 to 4 days. The larvae tend to aggregate on the ground beneath the dung for pupation, which lasts 2 days at 28°C (Hafez, 1949).

Hafez (1949) reported Staphylinidae {Philonthus) and Histeridae {Hister bimaculatus) as important predators of Coproica larvae. Philonthus is generally more abundant than Hister so Hafez (1949) speculated that Philonthus has a greater impact on larval populations, although this was not measured empirically. The beetle larva makes an incision near the centre of a fly larva with its mandible and then sucks out the contents. Philonthus species are predators of the larvae of a variety of coprophagous

Diptera including species from the Muscidae (Harris and Oliver, 1979), Sepsidae (Hafez,

1948), and Sphaeroceridae (Hafez, 1949). Studies suggest that species of Philonthus may have a significant effect on the populations of these flies (Harris and Oliver, 1979).

6 Materials and Methods Sources of Material

More than 16 000 adult Coproica specimens were examined for this study. Most of the specimens examined were from the University of Guelph Collection.

Material that was borrowed from or deposited in other institutions is listed in the body of the text associated with the following acronyms.

ABS - Archbold Biological Station, Lake Placid, Florida.

AMNH - Department of Entomology Collection, American Museum of Natural History, New York, New York.

BCPM - Royal British Columbia Museum, Victoria, British Columbia.

BMNH - Department of Entomology, The Natural History Museum, London, United Kingdom.

BPBM - J. Linsley Gressitt Center for Research in Entomology, Department of Entomology, Bernice P. Bishop Museum, Honolulu, Hawaii.

CASC - Department of Entomology, California Academy of Sciences, Golden Gate Park, San Francisco, California.

CBFC - Coleccion Boliviana de Fauna, La Paz, Bolivia

CMNH - Section of Insects and Spiders, Carnegie Museum of Natural History, Pittsburgh, Pennsylvania.

CNCI - Canadian National Collection of Insects, Centre for Land and Biological Resources Research, Biological Research Division Agriculture Canada, Ottawa, Ontario.

DEBU - Department of Environmental Biology, University of Guelph, Guelph, Ontario.

IAVH - Instituto Alexander von Humboldt, Santafe de Bogota DC, Colombia.

INBC - Instituto Nacional de Biodiversidad, Santo Domingo de Heredia, Costa Rica.

IZAV - Museo del Instituto de Zoologia Agricola Universidad Central de Venezuela, Maracay, Venezuela.

7 MZLU - Museum of Zoology, Lund University, Lund, Sweden.

MZSP - Museu de Zoologia da Universidade de Sao Paulo (formerly Departamento de Zoologia, Secretaria de Agricultura; see depositories in Prado, 1969), Sao Paulo, Brazil

PMAE - Provincial Museum of Alberta, Edmonton, Alberta.

QCAZ - Universidad Catolica del Ecuador, Quito, Ecuador

ROME - Department of Entomology, Royal Ontario Museum, Toronto, Ontario.

SEMC - Snow Entomological Museum, University of Kansas, Lawrence, Kansas.

UASC - Museo de Historia Natural "Noel Kempff Mercado," Santa Cruz de la Sierra, Bolivia.

USNM - United States National Entomological Collection, Department of Entomology, U.S. National Museum of Natural History, Washington DC.

UWIC - Insect Collection of the Department of Biological Sciences, University of the West Indies, St. Augustine, Trinidad.

8 Specimens examined

Most specimens examined were either air-dried or critical point dried and affixed with glue to small, white cardboard points.

Wings were mounted for photography and close examination of characters.

Wings were removed with fine forceps, dipped in warm 10% KOH for 1 second and rinsed in cold water, neutralized in 10% acetic acid and then given another water bath.

Wings were placed on a flat slide with a drop of glycerin, and then placed under a cover slip for observation.

Genitalia dissections were often required for accurate identification, and were necessary for the formulation and testing of species hypotheses. Whole abdomens were cleared by immersion in hot 10% KOH for 3 to 5 minutes before being rinsed in cold water, placed in 10% acetic acid for 5 or more minutes, then given a final rinse of cold water. Cleared genitalia were placed in glycerin on depression slides to be observed and illustrated. Genitalia and wings were stored in glycerin in microvials pinned below the specimens.

Sequence data from mitochondrial DNA (Cytochrome Oxidase 1, the "barcode region") were obtained for C. urbana and related species in order to aid in the assessment of intraspecific variation. The sequence data ("barcodes") and associated specimen data are stored in the Barcode of Life Data Systems (BOLD) database

(http://www.barcodinglife.org/views/login.php). Specimens submitted for barcoding are flagged within this manuscript with 'BOLD,' and with yellow barcode of life labels in the

University of Guelph Insect Collection.

9 Label data for all specimens examined are formatted and errors in spelling have been corrected. Specimens borrowed from or deposited in institutions other than the

University of Guelph Insect Collection (DEBU) are followed by the acronyms for their respective institutions. Specimens that have been dissected are noted with an asterisk (*).

Data entries for specimens used for illustrations or wing photographs are noted with "- illustr" or "-wing" respectively.

10 Adult Morphological Characters and Terminology

The terminology used here follows McAlpine (1981) and Marshall and Richards

(1987) with the exception of a few abdominal characters, where Marshall and Langstaff

(1998) is used. The term subepandrial sclerite (Marshall and Langstaff, 1998) is used for the structure referred to as sternite 10 of the male abdomen by McAlpine (1981). The term postgonite is used in the place of paramere, and tergite 10 and sternite 10 of the female abdomen are used in the place of the terms epiproct and hypoproct.

The term pollinose is used to describe the minute setae covering the abdominal sclerites of Coproica species; it is used primarily as a contrast to the glabrous female sclerites present in some species of Coproica. The term setulose has been used to describe surfaces with small socketed macrosetae, primarily for the surface of the scutellum.

11 Measurements and Ratios

All measurements were averaged over five specimens of each sex if available.

Where there is substantial variation, the measurement is presented as a range; otherwise all measurements are presented as an average for the species. In many cases the drying process alters body length, and for this reason, body lengths are considered approximate.

1.) Body Length: Distance between tip of first flagellomere and tip of abdomen.

2.) Eye to gena ratio: Distance between dorsal most point of eye and ventral most point divided by the continuation of that line to most ventral point of gena.

3.) Head ratio: Distance between most dorsal point of eye and ventral margin of gena divided by distance between vibrissa and posterior margin of gena.

4.) Scutellar Ratio: Width of scutellum at base divided by length of scutellum.

5.) Costal sector ratio: Length of second costal sector divided by length of third costal sector.

12 Phylogenetic Approach

Character polarity was determined using Elachisoma Rondani, 1880 and

Gonioneura Rondani, 1880 as a generalized outgroup. These genera are likely candidates as Coproica's closest relatives based on many similarities in external and internal morphology. Both Elachisoma and Gonioneura share a very similar chaetotaxy and distiphallus structure to Coproica although Gonioneura is likely a closer relative, due to the presence of many strong synapomorphies (see Generic Diagnosis). To polarize characters unique within Coproica the "functional ingroup, functional outgroup" method was used (Watrous and Wheeler, 1981; Maddison et al., 1984).

An a posteriori system of character weighting was used following the method outlined by Marshall (1987). The system of weighting was used to indicate the level of confidence that the various clades on the tree are accepted, and not to affect the phylogenetic construction. Additional tree statistics (bootstrap, jackknife, and Bremer) were calculated to provide further information pertaining to the clades presented on the tree (Fig: 2).

All multi state characters are left unordered.

The software TNT (Goloboff et. al, 2008; freely available from the Willi Hennig

Society website (http://www.cladistics.org/index.html)) was used for all phylogenetic analyses. The most parsimonious trees were obtained using the 'New Technologies

Search' with the 'Ratchet', 'Drift' and 'Tree fusing' options selected (Driven Search:

200,000 add sequences, random seed =1). Jackknife and bootstrap (standard) supports were calculated using 100,000 replicates each. Bremer supports were calculated by comparing the most parsimonious tree (tree length = 321) to 7508 suboptimal trees (50

13 steps longer, 0.50 relative fit distance) using a heuristic search for 10,000 trees (saving 10 trees, random seed =1). The macro BREMER.run (Pefia & Wahlberg, 2008) was used to calculate the final values.

14 Phylogeny

Phylogeny of Coproica

Complete descriptions of the characters referenced herein are available on pages

20-23.

Fifty characters were used in the phylogenetic analysis of Coproica and one most parsimonious tree was obtained (length 321, consistency index 0.399, retention index

0.651). The only synapomorphies identified for Coproica are the setae evenly distributed on the anterior two thirds of the scutellum or arranged in two or more anterior rows and always continuous medially (character 6), a pair of sub-apical and mid-apical setae on the scutellum (characters 8 and 9), and the distinct arrangement of dorsal and ventroapical bristles on the mid tibia (character 10).

Gonioneura is likely a closer relative to Coproica than Elachisoma due to the presence of several shared characters, such as thickened surstylar bristles (character 29), patches of microsetulae on the wing (character 19), and a strikingly similar mid tibial chaetotaxy.

Coproica, excluding C. serra (the plesiomorph sister group to the rest of the genus), and C. coreana (an unplaced species between the C. acutangula and C. urbanalC. vagans species groups) can be divided into 3 major clades: the Coproica acutangula species group, the C. vagans species group, and the C. urbana species group

(Fig: 1). The C. acutangula species group includes mostly Old World species, the C. urbana species group is composed entirely of New World species, and the C. vagans species group is an assemblage of species with various New World and Old World origins.

15 C. coreana is difficult to place in a species group because it seems to share characters from all groups. C. coreana most closely resembles species in the C. acutangula species group, though it lacks the major synapomorphies of the group.

The Coproica acutangula species group is recognized by an angulate anal vein

(character 18), a posterior row of distinctly enlarged bristles on male sternite 5 (character

23), a sinuate bend in male sternite 6 (character 24), and the presence of thickened spur­ like or nail-like bristles on the surstylus (character 29). Two species not included in the

C. acutangula species group, C. coreana and C. serra, possess thickened surstylar bristles. Thickened bristles on the surstylus occur throughout the Limosininae, including one of the two genera used in the outgroup analysis (Gonioneura), and is therefore set as the ancestral state in Coproica.

The Coproica acutangula species group consists of six species: C. acutangula, C. digitata, C. lugubris, C. mitchelli, C. pappi, and C. pusio. Two distinct groupings of species occur within the species group: the species pair C. acutangula and C. digitata and the clade composed of C lugubris, C. mitchelli, and C. pusio. The sister species C. acutangula and C. digitata make up the best supported species pair on the tree (jackknife

97, bootstrap 98, Bremer 11). A ventral apical process on the male hind basitarsus

(character 15), the second costal sector longer than the third (character 16), and a sharp posterior process on the postgonite (character 33) are all synapomorphies for the species pair C. acutangula and C. digitata. Coproica lugubris, C. mitchelli, and C. pusio form a clade defined by the absence of posterodorsal bristles on the mid tibia (a reversal of character 11), an elongate, subapical bristle on the postgonite (character 32), and female sternite 8 at least three times longer than wide (character 42). These species all have a

16 heavy chaetotaxy and distinctly shaped, elongate surstylus. C. mitchelli seems out of place in the C. acutangula species group as the only exclusively New World species in an otherwise strictly Old World clade. We are confident, though, in C. mitchellfs current placement in the tree, due to the presence of several unequivocal synapomorphies that link C. mitchelli to the C. acutangula species group.

The Coproica vagans species group is composed of ten species and is recognized on the basis of three synapomorphies: posterior katepisternal bristle long (character 5), posterior row of scutellar setae elongate (character 7), and anal vein with an angulate bend (character 18). The species group can also be characterized by the sub-basal ventral bristles on the mid basitarsus, which are either in a pair [character 13(2)] or a cluster of three [character 13(3)]. A well-supported monophyletic group within the C. vagans species group is made up of C. ferruginata, C. rohaceki, and C. setulosa, all of which have female tergite 10 and cerci flat, glabrous, and orange (character 38) and female cerci with apical bristles modified into stout spines (character 37). Coproica hirtula, C. lacteipennis, and C. rufifrons represent another well supported lineage within the C. vagans species group. These species share a broad female sternite 8, which is at least 3/4 the width of sternite 7 (character 40). The group is zoogeographically heterogeneous, and includes New World, Old World, and cosmopolitan species.

The Coproica urbana species group is an entirely New World lineage of nine species. The synapomorphies defining the C. urbana species group are a pointed anterior margin on female sternite 8 (character 41) and a tapered or conical spermathecal base

(character 48). Excluding C testudinea sp. nov., this lineage is strongly supported by several more synapomorphies, including asymmetrical postgonites (character 31), and the

17 reduced dorsal surface of female tergite 8. The reduction of female tergite 8 is likely correlated with the absence of a modified male sternite 5 in this group. When a male

Coproica mounts a female he uses sternite 5 to clamp down on the membranous area between female tergites 8 and 10 (Lachmann, 1996). The expanded membranous region on the dorsum of the female may act to allow males with a plain sternite 5 to obtain a better grasp on conspecific females.

Coproica bifurcata sp. nov., C. emarginata, sp. nov., and C. hirtuloidea constitute a small clade within the C. urbana species group characterized by an anteroventral row of small setae along the entire length of the mid basitarsus (character 14), and by similarly shaped surstyli. This group is primarily Neotropical but C emarginata occurs in the southern Nearctic. Coproica cacti, C. bispatha sp. nov., C brachystyla sp. nov., and C. urbana belong to a strongly supported group characterized by the fusion of female tergite

10 to the cerci. In this clade, the right postgonite is hooked (Figs: 69, 283), while the left is spoon-shaped (Figs: 68, 282), except in C. bispatha, which has spoon-shaped left and right postgonites (Figs: 56, 57). The reversal of the asymmetrical postgonites in C. bispatha is a highly improbable change in character state. The postgonites in C. bispatha, however, are very similar in shape to those in C. brachystyla, C. cacti, and C. urbana.

The right postgonite of C. bispatha has a small apical notch (Fig: 57), absent on the left postgonite (Fig: 56), which is likely reminiscent of the hooked right postgonite of the other species (Figs: 69, 283).

Two species in the Coproica urbana species group have highly specific habitats.

Coproica testudinea is only found within gopher tortoise burrows in Florida, and C. cacti is closely associated with decaying cactus in the southwest United States.

18 DNA barcodes were obtained for C. urbana and C. diabolica sp. nov. Due to the age and collection method of most of the specimens submitted, only 5 specimens of C. urbana and the 1 C. diabolica sp. nov. returned with complete sequence data. Coproica diabolica sp. nov. has a 10% divergence in sequence data from C. urbana, and there was less than 1% divergence between all specimens of C. urbana. Although the data made available by this process was not used in the phylogenetic analysis it was useful while assessing the variation found in the widespread C. urbana.

19 List of Characters

The plesiomorphic state is indicated by (0), apomorphic states are indicated by (1) or greater. The lowest weight characters are labeled with (+), the highest weight characters are accompanied by (+++).

1 Gena - colour, (O)uniform in colour, same colour as head and thorax; (1) yellow on posterior half, contrasting with dark head and thorax (+)

2 Interfrontal bristles, (0) 4 or more pairs; (1)3 pairs (++)

3 Eye size, (0) small, eye to gena ratio 1.5 or less; (1) large, eye to gena ratio 2.0 or greater (+)

4 Anterior katepisternal bristle length, (0) extending less than half the distance between katepisternum and notopleuron; (1) extending half the distance between katepisternum and notopleuron, or more; (2) absent or minute (++)

5 Posterior katepisternal bristle length, (0) short, extending less than half the distance between katepisternum and notopleuron; (1) long, extending half the distance between katepisternum and notopleuron, or more (++)

6 Scutellum - discal setae, (0) absent; (1) present but discontinuous medially, in lateral patches; (2) present in 2 anterior rows; (3) present in 3 or more anterior rows (+++)

7 Scutellum - discal setae, (0) all setae on disc same length; (1) posterior row of scutellar setae at least 1.5 times length of rest of the scutellar setae (+)

8 Scutellum - pair of enlarged posterior discal setae (subapical), (0) absent; (1) present (+++)

9 Scutellum - midapical setae, (0) absent; (1) single pair present between enlarged posterior marginal bristles (+++)

10 Dorsal mid tibia - chaetotaxy, (0) all posterodorsal bristles absent; (1) posterodorsal bristle present at 3/4 (+++)

11 Dorsal mid tibia - chaetotaxy, (0) proximal posterodorsal bristles absent; (1) 1 proximal posterodorsal bristle present; (2) 2 proximal posterodorsal bristles present (++)

12 Mid basitarsus - anterior row of setae, (0) 2 strong anterior setae, 1 apical and 1 at 1/2; (1) 4 or more evenly spaced setae on distal half (+)

20 13 Mid basitarsus - ventral basal setae, (0) absent; (1) single anteroventral bristle; (2) paired ventrobasal bristles, 1 anteroventral, 1 posteroventral; (3) 2 anteroventral bristles and 1 posteroventral bristle, proximal anteroventral seta shorter than other 2 (++)

14 Mid basitarsus - anteroventral row of setae, (0) setae typically elongate, never forming a continuous row along length of midbasitarsus; (1) row of small setae along entire length of midbasitarsus (++)

15 Male hind basitarsus - ventral apical process, (0) absent; (1) present (+++)

16 Costal sector ratio, (0) C2 less than or equal to C3, C2 0.7 to 1.0 times as long as C3; (1) C2 greater than C3, C2 1.1 to 1.6 times longer than C3 (+)

17 Costal bypass, (0) costa extends at least 7 vein widths past its intersection with R4+5; (1) Costa terminates within 5 vein widths of its intersection with R4+5 (+)

18 Anal vein - sinuosity, (0) shallowly sinuate, without a cornered bend; (1) angularly sinuate, distinct 90° to 135° angle present (++)

19 Patches of thickened microsetae, (0) absent or not in following positions; (1) enlarged setae present between R2+3 and costa and on posterior margin of wing across from the terminus of the anal vein (+++)

Male Genitalia

20 Male sternite 5 - posteromedial spines, (0) absent; (1) patch of dense setae posteromedially (++)

21 Male sternite 5 - posteromedial tab, (0) absent; (1) present in a pair; (2) present in a pair, right tab larger than the left; (3) distinct, single medial tab (+)

22 Male sternite 5 - surface bristles, (0) bristles uniform in size; (1) enlarged laterally (+)

23 Male sternite 5 - surface bristles, (0) bristles relatively uniform in size over surface; (1) posterior row of distinctly enlarged bristles (+++)

24 Male sternite 6, (0) narrow and tapered, straight; (1) narrow with sinuate bend below centre of sternite 5 (+++)

25 Epandrium - enlarged dorsal setae, (0) absent; (1) enlarged pair of dorsolateral setae present (++)

26 Epandrium - fusion ventral to anal opening, (0) discontinuous, with large rounded lobes almost touching each other; (1) narrowly fused, broadly fused in C. testudinea (+++)

21 27 Male cerci, (0) simple, with a medial protuberance; (1) flat, unmodified; (2) small, pointed sub-apical process; (3) sub-apical medial process just dorsal to an additional distinct, elongate process; (4) pointed medial process, and additional setose process; (5) pointed medial process, and additional setose process with distinct constriction at base (+)

28 Hypandrium, (0) basal lateral arms of hypandrium loosely articulating with epandrium; (1) basal lateral arms of hypandrium fused to epandrium (+++)

29 Surstylus - bristles, (0) thickened into broad, heavy spurs or nail-like processes; (1) thin and unmodified (++)

30 Postgonite - shape, (0) curved anteriorly; (1) simple, straight; (2) curved posteriorly; (3) s-shaped, with sharp posterior angle and apex pointing anteriorly (++)

31 Postgonites, (0) symmetrical; (1) asymmetrical; (2) asymmetrical with left postgonite broad (+++)

32 Postgonite - elongate, subapical anterior bristle, (0) absent; (1) present (++)

33 Postgonite - posterior process, (0) absent; (1) process present halfway along postgonite (+++)

34 Basiphallus - posteroventral surface, (0) undeveloped; (1) posteroventral crest or keel present (+++)

Female Genitalia

35 Female tergite 8 - dorsal surface, (0) sclerotized; (1) only present as a thin anterior strip, broad laterally; (2) only present as a thin membrane, with lateral sclerotized plates (++)

36 Female tergite 10 - fusion, (0) tergite 10 free from cerci; (1) tergite 10 fused to cerci (+++)

37 Female cerci - bristles, (0) thin and sinuate; (1) short, modified into thick spines (++)

38 Female tergite 10 and cerci - surface, (0) pruinose; (1) glabrous and shiny with a few outstanding microsetae; (2) tergite 10 and anterior half of cerci glabrous, rest pollinose (++)

39 Female sternite 7 - posterior margin, (0) straight or rounded; (1) with paired mediolateral processes; (2) with medial point; triangular; (3) with central notch (+)

40 Female sternite 8, (0) small, at most 1/2 width of sternite 7; (1) large, at least 3/4 width of sternite 7 (+++)

22 41 Female steraite 8 - anterior margin, (0) anterior margin straight; (1) anterior margin with pointed medial process (+++)

42 Female steraite 8, (0) broad or equally as long as broad; (1) at least three times as long as broad (+++)

43 Female sternite 8 - row of 4 posterior setae, (0) absent; (1) present (++)

44 Female sternite 8 - pair of elongate bristles, (0) absent; (1) present (+)

45 Female steraite 10 orientation, (0) relatively flat, resting dorsally beneath epiproct; (1) curved, wrapping around the tip of the abdomen, oriented posteriorly (++)

46 Female sternite 10 - enlarged pair of medial setae, (0) absent; (1) present (++)

47 Spermathecae - length, (0) wider than long; (1) approximately as long as wide; (2) approximately 1.5 times as long as wide (++)

48 Spermathecae - shape, (0) evenly rounded; (1) base tapering, conical; (2) cylindrical (++) 49 Spermathecae - duct length, (0) duct short, 1 to 2 times the length of spermathecal bulb; (1) duct very long, 3 to 7 times length of spermathecal bulb (++)

50 Spermathecae - duct constrictions, (0) spermathecal duct completely membranous, no constrictions present; (1) a short portion of spermathecal duct sclerotized below spermathecal bulb, with a constriction at its base. (++)

23 Discussion

Coproica is a genus of widespread and common flies that make up a significant

component of the fauna of many ecosystems, including urban and agricultural settings.

Coproica is also the last major genus of Sphaeroceridae in the Nearctic region to be revised. With the keys, descriptions of new species, and re-descriptions of previously

recognized species in this thesis, the tools are now available for researchers to identify all

currently recognized sphaerocerid flies collected in the Nearctic Region except for those

in the small genus Rudolfina. Taxonomic revisions such as this thesis facilitate biological

surveys by providing the means to identify or recognize species. For example, three

species of Coproica (C. mitchelli, C. hirticula, and C. diabolica sp. nov.) were collected

during a recent survey of the Bruce Peninsula (Canada: Ontario) (Marshall et al., 2001), yet were unidentifiable previous to this revision.

The known ranges of many species have here been significantly expanded.

Coproica rohaceki is newly recorded from the New World, and C. hirtuloidea, previously

only recognized from Bolivia and Peru, is here recorded as common throughout much of

South America. In other cases, gaps in the distributions of many other common species

such as C. acutangula, C. ferruginata, and C. hirticula have been filled.

The status of Coproica urbana (e.g., single species or a complex comprised of

multiple species) is an issue that could benefit from additional collection efforts. The

morphological variation apparent within C. urbana is here considered to be intraspecific

as I was unable to discern geographic patterns for any morphological characters (see

species comments for C. urbana). Moreover, attempts to analyze genetic differences

were relatively unsuccessful. I examined mostly old, dried specimens, many of which

24 had been collected using pan traps. The age of the specimens and collection methods had resulted in degradation of the DNA in all but five of 16 individuals analyzed. There was

<1% variation in the DNA barcode sequences obtained from these specimens (one from

Texas, four from Ontario). Although my assessment of C. urbana indicates that it is a

single widespread species, it would be useful to perform morphological and genetic

analyses on large series of specimens collected from across the range of the species to allow this question to be settled.

Two species, C. testudinea sp. nov. and C. cacti, provide examples of extreme habitat specialization in Coproica. Because these species within Coproica have become highly specialized and restricted to very specific microhabitats, other specialized species may be awaiting discovery.

Once a revision such as this has been conducted, biogeographical patterns become more evident. Coproica is split into three main species groups: the C. acutangula, C.

urbana, and C. vagans species groups (Fig: 1). Two of these have relatively simple biogeographic patterns. The C. acutangula species group is primarily found in the Old

World, with one New World species (C. mitchelli) and one cosmopolitan species (C. acutangula). The C. urbana species group consists of only New World species. More

interesting zoogeographic patterns emerge when examining the distribution of species in the zoogeographically heterogeneous C. vagans species group.

The C. vagans species group contains three New World species (C. novacula, C. setulosa, and C. galapagos), one Old World species (C. lacteipennis), and six widespread

or cosmopolitan species (C. hirticula, C. rohaceki, C. ferruginata, C. vagans, C. hirtula,

and C. rufifrons). The presence of so many cosmopolitan species muddles the

25 zoogeographic origins of the flies in this species group. Coproica hirticula, one of the cosmopolitan species, is likely New World in origin based on its exclusively New World sister species C. novacula. The location of the C. hirticula and C. novacula species pair as the sister clade to the rest of the C. vagans species group suggests that the C. vagans species group is New World in origin. The simplest solution, therefore, is that the cosmopolitan species within the C. vagans species group are New World in origin and secondarily distributed by human activities. This solution is corroborated by the results of my thesis that place the New World-exclusive C. urbana species group, as the sister clade to the C. vagans species group.

The conclusion that the C. vagans species group is a New World clade puts the distribution of C. lacteipennis, an Old World species, out of place. C. lacteipennis belongs to a clade including C. rufifrons, a widespread species in both the Oriental and

Neotropical regions, and C. hirtula, a cosmopolitan species. The sister species for this clade is the New World C. galapagos, which demonstrates a cross-Pacific disjunction between these sister taxa. Similar patterns for sister taxa occur elsewhere in the

Sphaeroceridae. For example, Rudolfina digitata (Marshall) from the Nearctic region has

a close sister species, Rudolfina rozynski (Rohacek), from central Europe and the Kola peninsula (Marshall, 1991), and species from and Phthitia have been shown

to have disjoint populations in the Nearctic and Palearctic (Marshall, 1997). The eastern

Palearctic distribution of C. lacteipennis (Japan: Ryukyu Islands) makes it likely that the ancestor of C. lacteipennis dispersed westward across the Bering Strait. This pattern may

also explain the New World distribution of C. mitchelli, a species in the C. acutangula

26 species group, though the dispersal in this case would have occurred in the opposite

direction.

The development of a phylogeny is instrumental in putting the behaviours and

adaptations of organisms into an evolutionary context. For example, the use of a sperm

plug is absent in Coproica acutangula, C. lugubris, and C. hirticula, and present in C. ferruginata and C. vagans (Lachmann, 1997). When these states are mapped onto the

phylogeny, it becomes apparent that the use of a sperm plug is a derived state within the

genus. With the ancestral and derived states of this character inferred, more complex

hypotheses can be made about the function of the sperm plug in Coproica and what

selection pressures may have led to its presence in C. ferruginata and C. vagans.

Moreover, within the framework provided by the phylogeny, it is possible to predict the

presence/absence of sperm plugs in other species within the genus; these hypothesized

character states can be confirmed through actual observation.

With this revision, the New World Coproica are now accessible for researchers to

document and expand our knowledge about these ubiquitous flies. The revision allows

for accurate identification of species, an essential aspect of all biological sciences, and

allows observations and hypotheses about Coproica biology, behaviour, and

zoogeography to be put into an evolutionary perspective.

27

Discussion of Generic Nomenclature

Rondani (1861) named Coproica to replace the name Heteroptera Macquart,

1835, preoccupied by Heteroptera Rafinesque, 1814. Macquart (1835) had erected the genus Heteroptera for specimens labeled as Limosina pusilla (Fallen, 1820). However, these specimens had been misidentified by Meigen in 1830, and were actually an undescribed species of Sphaeroceridae, later to be described as Limosina acutangula

Zetterstedt, 1847. Limosina pusilla belongs in the genus Ischiolepta, not Coproica, and its correct placement would have left Coproica without a type species. To amend this, a case was made to the International Commission of Zoological Nomenclature (ICZN) to conserve the names Coproica Rondani, 1861 and Ischiolepta Lioy, 1864 {fox Ischiolepta pusilla (Fallen)) (Wheeler and Swann, 1994). Limosina acutangula Zetterstedt, 1847 was designated as the type species of Coproica and Heteroptera Macquart, 1835 was placed on The Official Index of Rejected and Invalid Generic Names in Zoology (Wheeler and

Swann, 1996).

Coproica was originally a monotypic genus as defined by Macquart (1835) (as

Heteroptera) and later Rondani (1861). The presence of enlarged setae on the dorsal margin of the male wing in C. acutangula was used as the primary defining character for the genus. Species belonging to Coproica were later placed within the massive genus

Limosina. Duda's (1918) revision of the European Limosina split it into subgenera, one of which was Limosina (Coprophila) (type species: Borborus vagans Haliday, 1833), a junior subjective synonym of Coproica. The subgenus was erected for species having a setulose scutellum, apical and preapical mid tibial bristles absent, and enlarged ventral

28 setae on the mid basitarsus. Duda (1925) later expanded this definition to include an angularly sinuate anal vein. This expanded definition facilitated the exclusion of the few species of Rachispoda that have sparse setae on the scutellum. Later, Vanschuytbroeck

(1950) incorrectly incorporated the subgenus Elachisoma Rondani, which lacks the setulose scutellum, into Leptocera (Coprophila) on the basis of the presence of the angularly sinuate anal vein. Richards (1960) described two species of Nearctic Coproica,

C. cacti and C. urbana, which both lack the angulate anal vein. Richards noted that these species had two patches of lengthened and differently oriented microchaetae on the wings, 1 on the posterior margin of the wing and another between wing veins C and R2+3.

Coproica was treated as a subgenus of Limosina by Duda (1918, 1925, 1938), then as a subgenus of Leptocera Olivier, 1913 by Spuler (1924, 1925) and Richards

(1960). Frey (1945) and Collin (1956) treated Coproica as a genus in their respective reviews, yet Coproica was not consistently given generic status until Hackman (1969).

29 Generic Diagnosis

Small (0.9mm to 2.4mm) Limosininae, recognizable by the following features.

Synapomorphies of Coproica are in boldface.

1.) Scutellar disc with setae evenly distributed on anterior 2/3 of scutellum or arranged in 2 or more anterior rows, setae always continuous medially.

This is the traditional defining character for Coproica. It remains the best defining and diagnostic character for the genus. In Coproica, as redefined here, these setae form a patch that is always continuous medially, and never forms lateral patches as in some species of Philocoprella, Pullimosina (Dahlimosina), and Rachispoda.

2.) Subapical scutellar setae enlarged:

This is another characteristic of the scutellum that separates Coproica from all other species of Sphaeroceridae with a setulose scutellum.

3.) Midapical scutellar setae present:

Another characteristic of the scutellum that separates Coproica from all other species of

Sphaeroceridae with a setulose scutellum.

4.) Wing with patches of enlarged microsetulae between R2+3 and costa, and where the anal vein would intersect the posterior margin:

This was introduced as a defining character for Coproica by Richards (1960), and the patches of microsetulae in the above positions are reliably found in all members of the

30 genus. This character is difficult to see under a regular dissecting microscope, and frequently requires high magnification and careful angling of the wing for viewing. The microsetulae are clearly visible on slide-mounted wings. Similar patches of setulae are found in Philocoprella and Gonioneura.

5.) Mid tibia with anterodorsal bristles at 1/4, 1/2, and 3/4, and a posterodorsal bristle at

3/4, bristles at 3/4 stronger than others:

All Coproica have the basic mid tibial bristle arrangement outlined above, with or without additional posterodorsal bristles. A very similar bristle pattern is found in

Gonioneura, although, speciess of Gonioneura have the second and third anterodorsal bristles closer together than in Coproica.

6.) Mid tibia with enlarged bristles at anterior and posterior ventrolateral apices:

Gonioneura has a ventral-apical patch of bold setae on the mid tibia. Though different in appearance, these bristles are likely homologous, due to the presence of the correlated characters outlined above.

31 Generic Description

Genus Coproica Rondani, 1861

Coproica Rondani, 1861: 10 (feminine) [nom.n. for Heteroptera Macquart, 1835]. Type species: Limosina acutangula Zetterstedt, 1847, subsequent designation by ICZN, 1996: 136. - Richards, 1960c: 199-208 [as subgenus of Leptocera Olivier, 1813; diagnosis, review of species, key]; Richards, 1965c: 725 [as subgenus of Leptocera; Nearctic catalog]; Richards, 1967b: 16 [as subgenus of Leptocera; Neotropical catalog]. Hackman, 1969a: 202, 207 [as genus; phylogenetic notes, biogeography]; Papp, 1973c: 24-30 [as genus; diagnosis, key to Hungarian species, illustr.]; Hackman, 1977: 400 [as genus; Oriental catalog]; Richards, 1980: 618 [as subgenus of Leptocera; Afrotropical catalog]; Papp, 1984: 80-82 [as genus; Palaearctic catalog]; Papp & Rohacek, 1987: 105-107 [as genus; key to Mediterranean species]; Marshall & Richards, 1987: 999 [as genus, diagnosis in key, illustr.]; Marshall, 1989b: 603 [as genus; Australasian/ Oceanian catalog]; Hayashi, 1991c: 235-236 [as genus; key to species from Pakistan]; Hayashi, 1995a: 229-232 [as genus; key to Japanese species, illustr.]; Rohacek, 1998a: 477 [diagnosis in key, illustr.]. Hayashi, 2005a [illustr.]; Papp, 2008 [key to Palearctic species, illustr.]. Heteroptera Macquart, 1835: 570 (feminine) [a junior homonym of Heteroptera Rafinesque, 1814 and Heteroptera Latreille, 1817, Hemiptera, cf. Richards, 1960c: 199]. Type species: Copromyzapusilla auct. nee Fallen, 1820 [= Coproica acutangula (Zetterstedt, 1847)], monotypy. - Haliday, 1836: 334 [diagnosis]; Rondani, 1856: 124 [recognition of type species]; Rondani, 1861: 10 [homonymy]; Richards, 1930: 306-308 [as subgenus of Leptocera; diagnosis, review of British species]. Coprophila Duda, 1918: 45 (feminine) [as subgenus of Limosina Macquart, 1835]. Type species: Borborus vagans Haliday, 1833, subsequent designation by Spuler, 1925a: 123. - Duda, 1918: 45-47 [as subgenus of Limosina; diagnosis, key]; Spuler, 1924d: 377 [as subgenus of Leptocera Olivier, 1813]; Spuler, 1925a: 122- 124 [as subgenus of Leptocera; diagnosis, key]; Richards, 1930: 266 [synonymy]; Duda, 1938: 161-168 [as subgenus of Limosina; review of Palaearctic species, key].

Always macropterous. Body length 0.9mm to 2.4mm. Colour brown or orange. Head typically wider than long, always taller than long, ratio of height to length typically 1.3:1

- 1.5:1, but longitudinally flattened in C. coreana and C. testudinea (height:length approximately 1.7:1). Frons convex, micropubescent. Eye with row of small setulae running from anterior vertex to a point medial of orbital bristles, and with 2 subequal

32 lateroclinate orbital bristles. Inner vertical bristles approximately 1.3 times length of orbital bristles; Outer vertical bristle approximately same length as orbital bristles; interfrontal bristles in 3 to 6 sub-equal pairs; ocellar bristles proclinate, length subequal to inner vertical bristles; inner occipital bristles slightly convergent; 0-2 pairs of small, convergent postocellar setulae. Eye with anterodorsal notch. Eye to gena ratio variable between species. Row of setae present along ventral margin of gena; genal setae arranged into 1-2 rows. Face tuberculate dorsally, concave ventrally, with anteriorly projecting ventral margin and faint carina extending ventrally from tubercle. Antennae divergent; arista uniformly pubescent, situated dorsolaterally in middle of first flagellomere. Scutum approximately as wide as long, with a pair of of postsutural

(prescutellar) dorsocentral bristles, and sometimes an additional postsutural pair of dorsocentrals (C. setulosa). Postpronotum with 1 anterior and 2 basal setae, lateral basal postpronotal seta twice the length of other postpronotal setae. 2 enlarged intra-alar and 2 enlarged supra-alar bristles. Scutellum typically evenly rounded, width at base of scutellum 1.1 to 1.5 times length. Scutellum with four large marginal bristles, a smaller midapical pair of setae, 1 to 4 additional marginal setae between larger lateral and apical marginal bristles. Discal setae evenly distributed on anterior 2/3 of scutellum or arranged in 2 or more anterior rows, always continuous medially, never forming 2 lateral patches;

1 pair of convergent discal bristles on posterior 1/3. Thoracic pleurites except katepisternum bare; katepisternum with 2-3 strong dorsal bristles, a ventral patch of ventrally facing setae, and a single central dorsally oriented bristle. Presternum narrow, with slightly expanded anterior sclerotized region. Coxae yellow to brown; mid coxa often darker. Legs yellow to brown, fore leg occasionally darker than other legs. Fore

33 femur with enlarged posterodorsal seta at 1/2, or with posterodorsal row of enlarged setae. Mid femur with 2 sub-apical anterior bristles and 1 sub-apical posterior bristle.

Mid tibia with anterodorsal bristles at 1/4, 1/2, and 3/4; posterodorsal bristles variable, one relatively strong posterodorsal at 3/4; ventral bristle at 2/3. Mid basitarsus chaetotaxy highly variable, anteroventral and posteroventral apical bristles always present. Hind basitarsus sometimes with apical ventral process in males (C. acutangula and C. digitata). First costal sector with 2 rows of bristles approximately 2 times width of costa and an enlarged pair of bristles near base, second and third costal sectors with rows of setulae 1-1.5 times width of costa. Length of second costal sector typically equal to or slightly less than third costal sector, some species with second costal sector longer.

Anal vein never reaching posterior wing margin, fading approximately half-way, with a cornered angle or shallow bend. Abdominal tergites well-sclerotized, sparsely and uniformly setulose, each tergite with an enlarged bristle at posterolateral corners. Sternite

1 absent. Sternite 2 present or reduced to a lightly sclerotized patch or strip. Sternites 3 and 4 broadly rectangular.

Male terminalia:

Sternite 5 broad and rectangular or crescent-shaped, posterior margin plain or with 1 or 2 medial tabs, with or without posteromedial patch of dense microsetulae, bristles typically denser laterally, bare medially in some species. Synsternite 6+7 simple, straight, or with distinct medial bend; small pair of setulae followed by enlarged pair of bristles on left side; synsternite 6+7 broadly fused to sternite 8. Sternite 8 narrowly articulating with epandrium dorsally, with small pair of setulae followed by enlarged pair of bristles similar to those on synsternite 6+7. Epandrium with enlarged pair of anterodorsal bristles

34 in all species except C. galapagos sp. nov., C. mitchelli, and C. pusio; subanal plate present or absent, narrow or thick if present. Cercus fused to epandrium, unmodified

(straight ventral margin), with a simple medial protuberance, or elaborately modified.

Hypandrial arms well sclerotized, solidly fused to hypandrial apodeme and to anteroventral corners of epandrium; subepandrial sclerite articulating with hypandrium

(at its point of fusion with epandrium) anteriorly and with cerci posteriorly. Surstylus highly variable, shape and chaetotaxy species-specific, typically with rounded anterior lobe (pointed in some species); articulating anterodorsally with epandrial-hypandrial junction and posterodorsally with subepandrial sclerite. Left and right postgonites typically symmetrical, but asymmetrical in some species; articulating posteriorly at joint between aedeagal apodeme and basiphallus, and anteriorly with posteromedial process on hypandrium. Aedeagal apodeme (phallapodeme) simple, elongate. Ejaculatory apodeme small, narrow, slightly expanded at apex. Basiphallus elongate and tubular, straight or elbowed; with posterodorsal opening behind articulation with aedeagal apodeme; posteroventral keel-like process present in many species. Distiphallus with proximal tubular sclerite articulating with basiphallus; tubular sclerite with proximal opening and split distally, forming dorsal and ventral sclerites. Ventral sclerite produced into pair of large lateral sclerites, each with 0-3 dorsal processes of species-specific shapes. Dorsal sclerite produced distally between lateral sclerites, typically bifurcate, often fusing apically with, or in centre of, lateral sclerites. Paired membranous sheets extending distally from ventral sclerite, usually with lightly sclerotized knobs or ridges on outer surface, sometimes surrounding tip of distiphallus.

35 Female terminalia:

Tergite 7 unmodified. Tergite 8 broad and rectangular, narrowed dorsomedially, or broken into 2 separate lateral sclerites connected by a dorsomedial membrane. Tergite 9 absent. Tergite 10 pollinose or glabrous; with a pair of anterolateral processes and pair of central bristles; narrowly fused/articulating with cerci laterally, or broadly fused to cerci in some species. Cercus pollinose or glabrous; rounded, rectangular, or triangular.

Sternite 7 broad and rectangular, some species with posteromedial invagination.

Sternite 8 highly variable, shape species-specific, typically with a posterior row of

4 bristles or spines, most species with an enlarged pair of central bristles. Sternite 10 pollinose, shape variable, lying flat ventrally or 'wrapped' posteriorly around tip of abdomen. Spermathecae spherical, cylindrical, or pear-shaped, apical invaginaton typically cylindrical, surface with fine, fingerprint-like etchings; spermathecal duct length between spermathecal bulb and valve variable between species; spermathecal valve sclerotized distally and membranous proximally.

36 Key to New World species

1 Mid tibia without a proximal posterodorsal bristle (Fig: 19) 2 - Mid tibia with 1 - 3 proximal posterodorsal bristles (Figs: 16-18, 20) 3

2 Posterior katepisternal bristle approximately 3 times length of anterior katepisternal bristle (Fig: 10). Mid basitarsus with pair of strong sub-basal ventral bristles (Fig: 26). Surstylus simple, curved posteriorly with sharp apical process (Fig: 145). Female sternite 8 small, oval (Figs: 152, 154). Widespread in North and South America Coproica hirticula Collin, 1956 - Katepisternal bristles subequal (Fig: 8). Mid basitarsus with 1 strong sub-basal anteroventral bristle (Fig: 25). Surstylus with elongate posterior process with thickened, black bristle at apex (Figs: 189, 191). Female sternite 8 elongate, 3 times longer than wide (Figs: 196, 198). Widespread in North and Central America Coproica mitchelli (Malloch, 1913)

3 Mid basitarsus without a strong sub-basal ventral bristle, with row of uniform, small anteroventral bristles (Fig: 21) 4 - Mid basitarsus with 1 or more strong sub-basal ventral bristles (Figs: 22-28) and 1-4 bold bristles on anteroventral margin 9

4 Male S 5 - Female $ 7

5 Surstyli asymmetrical; left surstylus with small notch at apex; right surstylus concave posteriorly, with blunt processes bordering concavity (Figs: 114, 119). Central America, Florida, Caribbean Coproica emarginata S sp. nov. - Surstyli symmetrical, neither notched nor concave (Figs: 42, 166) 6

6 Surstylus with strong angles on posterior lobe and strongly concave ventrally (Fig: 166). Right postgonite without expanded apex (Fig: 169). Central and South America Coproica hirtuloidea $ (Duda, 1925) - Surstylus with rounded posterior lobe, without strong angles, and shallowly concave ventrally (Fig: 42). Right postgonite with expanded apex (Fig: 45). Argentina, Bolivia, southern Brazil Coproica bifurcata S sp. nov.

7 Female sternite 7 with paired posterolateral lobes (Fig: 49). Anterior process on sternite 8 broad, with small lateral projections (Figs: 49, 51). Argentina, Bolivia, southern Brazi Coproica bifurcata $ sp. nov. - Female sternite 7 with straight posterior margin (Figs: 122, 176). Anterior process on sternite 8 narrow and tapering, or evenly rounded (Figs: 121, 122, 172, 174). South America or southern United States 8

8 Anterior point on sternite 8 rounded, pollinose (Figs: 172, 174). Central and South America Coproica hirtuloidea 9 (Duda, 1925)

37 - Anterior point on sternite 8 sharp, glabrous, (Figs: 121, 122). Central America, Florida, Caribbean Coproica emarginata $ sp. nov.

9 Posterior katepisternal bristle long, extending at least 1/2 distance from base of bristle to wing base, anterior and posterior katepisternal bristles subequal, posterior bristle distinctly longer and thicker than anterior bristle, or with 3 katepisternal bristles (Figs: 9- 13) 10 - Posterior katepisternal bristle short, no longer than 1/3 the distance from base of bristle to wing base, anterior and posterior katepisternal bristles always subequal (Fig 8) 17

10 Katepisternum with 3 dorsal bristles (Fig: 9). Scutum orange, with medial brown line. Widespread Coproica ferruginata (Stenhammer, 1855) - Katepisternum with 2 dorsal bristles (Fig: 10-13). Scutum colour black or brown ..11

11 Two to four pairs of postsutural dorsocentral bristles present. Anterior and posterior katepisternal bristles extending almost to or past level of wing base (Fig: 11). Female cercus and tergite 10 glabrous, cercus with stout apical spine (Fig: 263). Southern United States and South America Coproica setulosa (Duda, 1929) - One pair of postsutural, prescutellar dorsocentrals present. Posterior katepisternal bristle at least 2 times longer than anterior bristle or both bristles never reaching level of notopleuron (Figs: 10, 12, 13). Female cercus and tergite 10 glabrous or pollinose, never with stout apical spine (Figs: 207, 250, 294) 12

12 Second costal sector 1.1 to 1.6 times longer than third costal sector (Fig: 288). Frons always uniform brown to dark brown, never with anterior orange band. Mid basitarsus with cluster of 3 enlarged sub basal ventral bristles (Fig: 27). Epandrium always with pair of enlarged dorsal bristles (Figs: 234, 235, 289, 290). Female tergite 10 and cercus pollinose or glabrous and orange (C. rohaceki, Fig: 239). Female sternite 8 always less than 1/2 width of sternite 7 (Fig: 240) 13 - Second costal sector 0.7 to 1.0 times as long as third (Fig: 243). Frons uniform brown to dark brown or with bright anterior orange band (C. rufifrons). Mid basitarsus with 2 or 3 sub basal ventral bristles (Figs: 26, 27). Epandrium with or without (C. galapagos, Figs: 138, 139) pair of enlarged dorsal bristles. Female tergite 10 and cercus always pollinose. Female sternite 10 less than 1/2 width of sternite 7 or greater than 1/2 width of sternite 7 (C. hirtula, Fig: 162; C. rufifrons, Fig: 251) 14

13 Posterior katepisternal bristle 1.5-2.0 times longer than anterior katepisternal bristle (Fig: 12). Anterodorsal and posterodorsal mid tibial bristles equal in length. Male surstylus quadrate, lacking a distinct posterior process (Fig: 256). Female cercus and tergite 10 glabrous and orange (Fig: 263). South America Coproica rohaceki Carles-Tolra, 1990 - Anterior and posterior katepisternal bristles subequal (Fig: 13). Anterodorsal mid tibial bristles longer than posterodorsal bristles. Male surstylus elongate posteriorly, claw- shaped at apex (Fig: 289). Female cercus and tergite 10 pollinose (Fig: 294). Widespread Coproica vagans (Haliday, 1833)

38 14 Mid tibia with 1 proximal posterodorsal bristle (Fig: 17). Mid basitarsus with 1 pair of sub basal ventral bristles (Fig: 26). Southern United States and South America Coproica novacula sp. nov. - Mid tibia with 2 or more proximal posterodorsal bristles (Fig: 20). Mid basitarsus with cluster of 3 sub-basal ventral bristles, 1 small and 1 large anteroventral bristle and 1 large posteroventral bristle (Fig: 27) 15

15 Epandrium with only uniformly small bristles (Figs: 138, 139). Width of female sternite 8 less than half width of sternite 7 (Fig: 141). Ecuador (Galapagos) Coproica galapagos sp. nov. - Epandrium with pair of enlarged dorsolateral bristles (Figs: 156, 157, 244, 245). Width of female sternite 8 greater than half width of sternite 7 (Figs: 162, 251) 16

16 Anterior margin of frons with orange band. Surstylus with 2 posteroventral processes, proximal process deflected medially (Figs: 244, 245). Female sternite 8 with small posteromedial invagination (Figs: 249, 251). Southern United States and South America. Coproica rufifrons Hayashi, 1991 - Frons uniformly dark brown. Surstylus without ventral processes (Fig: 156). Posterior margin of female sternite 8 straight (Fig: 162). Widespread Coproica hirtula (Rondani, 1880)

17 Scutellum with 3 or more anterior rows of small setae. Interfrontal bristles in 4 or more pairs (Fig: 7). Second costal sector longer than third costal sector (Figs: 29, 30); male wing with crossvein dm-cu forming sharp angle with CuAi, hind margin with patch of elongate setulae (Fig: 29). Male hind basitarsus with apicoventral process (Fig: 15). Surstylus with 1 or more distinctly thickened apical bristles (Fig: 32). Widespread Coproica acutangula (Zetterstedt, 1847) - Scutellum with 2 anterior rows of small setae. Three pairs of interfrontal bristles, occasionally 4 on one side. Second costal sector shorter than third; crossvein dm-cu and CuAi perpendicular, hind margin of male wing unmodified (Fig: 276). Male hind basitarsus unmodified. Surstylus without a thick apical bristle or spur (Fig: 280). New World only 18

18 Eye small, eye to gena ratio typically 1.5:1 or less. Associated with gopher tortoise burrows or cacti. Nearctic 19 - Eye large, eye to gena ratio 2:1 or greater. No known association with above habitats. Nearctic or Neotropical 20

19 Interfrontal bristles in 3-4 short, subequal pairs. Mid basitarsus with paired sub-basal ventral bristles. Associated with gopher tortoise burrows. Florida Coproica testudinea sp. nov. - Interfrontal bristles in 3 thick, erect pairs, with middle pair stronger than other 2 pairs. Mid basitarsus with 1 sub-basal ventral bristle (Fig: 22). Associated with cacti. Southwest United States Coproica cacti (Richards, 1960)

39 20 Male cercus with small but distinct hooked processes (Figs: 90, 91); posterior lobe on surstylus broad and directed ventrally (Fig: 90). Female cerci free from tergite 10 (Fig: 96). Eastern North America and Caribbean Coproica diabolica sp. nov. - Male cercus without hooked processes (Figs: 280, 281); posterior process on surstylus narrow or small, often dorsally curved (Figs: 54, 66, 280). Female cerci fused to tergite 10 (Figs: 61, 72, 284) 21

21 Female cerci and tergite 10 glabrous on anterior half; apical and sub-apical bristles on cerci modified into short, thick spines (Fig: 72). Posterior process on surstylus small and pointed (Fig: 66). Central and South America Coproica brachystyla sp. nov. - Female cerci and tergite 10 completely pollinose; apical bristles sinuate, not modified into spines (Figs: 61, 284). Posterior process on surstylus elongate, slightly up curved (Figs: 54, 280) 22

22 Left and right postgonites symmetrical, both rounded at tip (Fig: 56, 57). Posterior process on surstylus short (Fig: 54). Sternite 10 complete, without floating sclerite between lateral arms (Fig: 62). Central and South America Coproica bispatha sp. nov. - Postgonites asymmetrical, left postgonite rounded apically, right postgonite hooked apically (Figs: 282, 283). Surstylus with long posterior process with rounded tip (Fig: 280). Sternite 10 with small floating sclerite between lateral arms (Fig: 285). Widespread in North and Central America Coproica urbana (Richards, 1960)

40 Key to World species Species described by Papp (2008) were not examined, and are included in this key based only on their descriptions.

1 Mid tibia without a proximal posterodorsal bristle (Fig: 19) 2 - Mid tibia with 1 - 3 proximal posterodorsal bristles (Figs: 16, 17, 18, 20) 14

2 Mid basitarsus without strong ventral setae. Discal cell short; dm-cu equal in length or shorter than segment of M between rm and dm-cu. Surstylus narrow, with thin, short posterior process. Oriental or Afrotropical 3 - Mid basitarsus with at least 1 strong ventral seta (Figs: 22-28). Crossvein dm-cu equal in length or longer than segment of M between rm and dm-cu. Shape of surstylus variable (Figs: 145, 178, 191,223). Distribution variable 4

3 Faded apical portion of M with a sinuate, S-shaped curve. Thailand Coproica aliena Papp, 2008 - Faded apical portion of M curved, but not sinuate. Afrotropical Coproica brevivenosa Papp, 2008

4 Posterior katepisternal bristle approximately 3 times the length of anterior katepisternal bristle. Mid basitarsus with pair of strong ventral basal bristles. Cosmopolitan Coproica hirticula Collin, 1956 - Anterior and posterior katepisternal bristles subequal. One strong anteroventral basal bristle on mid basitarsus or strong ventral basal bristles absent 5

5 Mid tibia with a distinct, long apical ventral bristle 6 - Mid tibia without enlarged apical ventral bristle 8

6 Head uniformly dark brown. Mid basitarsus lacking strong ventral bristles but with anteroventral and posteroventral rows of small setae. Surstylus with thickened posterior bristle, all bristles on surstylus short. Nepal, Thailand Coproica thaii Papp, 2008 - Anterior 1/3 of frons, face and gena yellow. Mid basitarsus with 2 or more strong ventral bristles. Surstylus either without a thickened posterior bristle or with bristles greatly elongate. Afrotropical 7

7 Femora dark brown. Bristles on surstylus short Coproica ruwenzoriensis (Vanschuytbroeck, 1950) - Femora yellow. Surstylus with 2 elongate setae, 2-3 times as long as width of surstylus Coproica flavifacies Papp, 2008

8 Ventral half of first flagellomere orange in females. Surstylus with venteromedial comb of thick, black bristles, without elongate, narrow, posterior process. Afrotropical Coproica serra (Richards, 1938)

41 - First flagellomere completely black or brown. Surstylus without ventral row of thick black bristles, C. pusio has a venteromedial row of long wavy setae (Figs: 178, 191, 223) 9

9 Surstylus short, rounded, without large thickened bristles at tip of posterior process. Postgonite with only small setae apically. Oriental 10 - Surstylus greatly elongate, narrow posterior process with a single thickened toenail-like bristle at its apex (Figs: 178, 191, 223). Postgonite with elongate subapical anterior bristle (Figs: 183, 194, 225). Distribution variable 11

10 Mid basitarsus with 2 strong anteroventral bristles, 1 sub-basal bristle and another atl/2. Surstylus with only small, uniformly spaced bristles posteroventrally. Nepal, Thailand Coproica bispinosa Papp, 2008 - Mid basitarsus with 1 large anteroventral bristle at 1/3. Surstylus with cluster of 3 enlarged posteroventral bristles. Oriental, Beijing Coproica unispinosa Papp, 2008

11 Mid basitarsus with 1 anteroventral bristle 2 times length of other ventral bristles (Fig: 24). Surstylus with thick anterior process approximately equal in length to posterior process (Fig: 178). Male steraite 5 with enlarged, central, spinose process (Fig: 185). Female sternite 8 broad anteriorly (Fig: 181). Widespread in Old World Coproica lugubris (Haliday, 1835) - Bristles on ventral surface of mid basitarsus uniform in size (Fig: 25). Surstylus with or without anterior process, but never greater than half as long as of anterior process (Figs: 194, 225). Male sternite 5 without posteromedial process (Figs: 193, 227). Female sternite 8 narrow along entire length (Figs: 198, 228). New or Old World 12

12 Ventral margin of surstylus straight, without anterior process, or bumps; row of elongate ventral setae along entire length. Female cercus with short, spine-like apical and sub apical bristles. Female sternite 8 with small bulge at posterior 1/3. Afrotropical.... Coproica perlugubris Papp, 2008 - Surstylus with anteroventral process or ventral wavy bump, never flat (Figs: 191, 223). Bristles on female cerci elongate, sinuate (Figs: 195, 229). Female sternite 8 straight (Figs: 198, 228). Distribution Variable 13

13 Surstylus with anterior process absent or reduced to a wavy bump, and with sparse setulae (Fig: 191). Female sternite 10 with straight anterior margin (Fig: 196). North and Central America Coproica mitchelli (Malloch, 1913) - Surstylus with anterior process half the length of posterior process and with heavy rows of long setulae. Anterior margin of female sternite 10 with 2 invaginations flanking a medial lobe (Fig: 230). Widespread in Old World Coproica pusio (Zetterstedt, 1847)

14 Mid basitarsus with row of uniform, small anteroventral bristles, without a strong sub- basal ventral bristle (Fig: 21) 15

42 - Mid basitarsus with 1-4 bold bristles on anteroventral margin and 1 or more strong sub- basal ventral bristles (Figs: 22-28) 20

15 Male $ 16 -Female? 18

16 Surstyli asymmetrical; left surstylus with small notch at apex; right surstylus concave posteriorly, with blunt processes bordering concavity (Figs: 114, 119). Central America, Florida, Carribean Coproica emarginata $ sp. nov. - Surstyli symmetrical, neither notched nor concave (Figs: 42, 166) 17

17 Surstylus with strong angles on posterior lobe and strongly concave ventrally (Fig: 166). Right postgonite not bifurcate (Fig: 169). Central and South America Coproica hirtuloidea S (Duda, 1925) - Surstylus with rounded posterior lobe, without strong angles, and shallowly concave ventrally (Fig: 42). Right postgonite bifurcate (Fig: 45). Argentina, Bolivia, southern Brazil Coproica bifurcata S sp. nov.

18 Female sternite 7 with paired posterolateral lobes (Fig: 49). Anterior process on sternite 8 broad, with small lateral projections (Figs: 49, 51). Argentina, Bolivia, southern Brazil Coproica bifurcata $ sp. nov. - Female sternite 7 with straight posterior margin (Figs: 122, 176). Anterior process on sternite 8 narrow and tapering, or evenly rounded (Figs: 121, 122, 172, 174). South America or southern United States 19

19 Anterior point on sternite 8 rounded, pollinose (Figs: 172, 174). Central and South America Coproica hirtuloidea $ (Duda, 1925) - Anterior point on sternite 8 sharp, glabrous, (Figs: 121, 122). Central America, Florida, Caribbean Coproica emarginata $ sp. nov.

20 Posterior katepisteraal bristle long, extending at least 1/2 distance from base of bristle to wing base (Figs: 9-13) 21 - Posterior katepisternal bristle short, no longer than 1/3 as long as distance from base of bristle to wing base (Fig: 8) 35

21 Katepisternum with 3 dorsal bristles (Fig: 9). Scutum orange with medial brown line. Cosmopolitan Coproica ferruginata (Stenhammer, 1855) - Katepisternum with only 2 dorsal bristles (Figs: 10-13). Scutum colour variable 22

22 2-4 pairs of postsutural dorsocentral bristles present. Anterior and posterior katepisternal bristles extending almost to or past level of wing base (Fig: 11). Female cerci and tergite 10 glabrous, with stout apical spine (Fig: 263). Southern United States and South America Coproica setulosa (Duda, 1929)

43 - 1 pair of postsutural, prescutellar dorsocentrals present. Posterior katepisternal bristle at least 2 times as long as anterior bristle or both bristles never reaching level of wing base (Figs: 10, 12, 13). Female cerci and tergite 10 glabrous or pollinose, never with stout apical spine (Figs: 140, 161, 207, 239, 250, 294) 23

23 Second costal sector 1.1 - 1.6 times as long as than third (Figs: 288) 24 - Second costal sector 0.7 - 1.0 times as long as third (Figs: 134, 155, 200, 243) 27

24 Postocellar, occipital and postocular setae white. Surstylus with elongate, narrow posterior process. South Africa Coproica albiseta Papp, 2008 - All head bristles brown or black. Surstylus without narrow process 25

25 Hind tibia with long preapical dorsal bristle. Large, reddish species Coproica demeteri Papp, 2008 - Hind tibia lacking a long preapical dorsal bristle. Body colour brown or black, never orange or red 26

26 Posterior katepisternal bristle 1.5 - 2.0 times longer than anterior katepisternal bristle. Posterodorsal mid tibial bristles equal in length to anterodorsal midtibial bristles. Heavily bristled flies. Female cerci and tergite 10 glabrous and orange (Fig: 239). Male surstylus quadrate, lacking a distinct posterior process (Fig: 233). Mediterranean, Neotropical Coproica rohaceki Carles-Tolra, 1990 - Anterior and posterior katepisternal bristles subequal in length (Fig: 13). Posterodorsal mid tibial bristles shorter than anterodorsal midtibial bristles. Without a distinctly heavy chaetotaxy. Female cerci and tergite 10 pollinose (Fig: 293). Male surstylus elongate posteriorly, claw-shaped at apex (Fig: 289). Cosmopolitan Coproica vagans (Haliday, 1833)

27 Mid basitarsus without a distinct basal ventral bristle, anteroventral bristle present at 1/3, posteroventral bristle present at approximately 1/2. Afrotropical 28 - Mid basitarsus with 2 or more basal or sub-basal ventral bristles, additional midbasitarsal bristles not in above combination (Figs: 26, 27) 29

28 Eye to gena ratio 2:1 or greater. Mid basitarsus with anteroventral bristle at 1/2, paired with posteroventral bristle. Male sternite with an asymmetrical posteromedial process Coproica ghanensis Papp, 1979 - Eye to gena ratio 1.5:1 or less. Mid basitarsus without additional anteroventral bristles. Male unknown Coproica microps Papp, 2008

29 Mid tibia with 1 proximal posterodorsal bristle (Fig: 17). Mid basitarsus with 1 pair of sub-basal ventral bristles (Fig: 26). Southern United States and South America Coproica novacula sp. nov. - Mid tibia with 2 or more proximal posterodorsal bristles (Fig: 20). Mid basitarsus with sub-basal ventral cluster of 3 bristles present at base of mid basitarsus, 1 small and 1 large anteroventral bristle and 1 large posteroventral bristle (Fig: 27) 30

44 30 Epandrium with only uniformly small bristles (Figs: 138, 139). Width of female steraite 8 less than half width of steraite 7 (Fig: 141). Ecuador (Galapagos) Coproica galapagos sp. nov. - Epandrium with enlarged pair of dorsolateral bristles (Figs: 156, 157, 244, 245). Width of female sternite 8 greater than half width of sternite 7 (Figs: 162, 251) 31

31 Frons uniformly dark brown or with orange anterior band. Surstylus with 2 posteroventral processes, proximal process short and blunt (Fig: 244) 32 - Frons uniformly dark brown. Surstylus without ventral processes (Fig: 156) 33

32 Frons with anterior orange band. Cosmopolitan...Coproica rufifrons Hayashi, 1991 - Frons uniformly dark brown. Oriental Coproica saprophaga Papp, 2008 33 Wing clear. Male surstylus elongate (Fig: 156). Cosmopolitan Coproica hirtula (Rondani, 1880) - Wing whitish, milky. Male surstylus short and quadrate. Oriental or Afrotropical 34

34 Postgonite geniculate, with distinct posterior angle. Surstylus quadrate, plain along posterior margin. Oriental Coproica lacteipennis Hayashi, 2005 - Postgonite straight, narrow. Surstylus rounded, with posterior hook-like process. Afrotropical Coproica pseudolacteipennis Papp, 2008

35 Interfrontal bristles in 4 or more pairs (Fig: 7). Surstylus with 1 or more distinctly thickened apical bristles (Figs: 32, 79, 103, 215) 36 - Interfrontal bristles in 3 pairs, occasionally four bristles on one side. Surstylus without a bold apical bristle (Figs: 54, 66, 90, 267, 280) 40

36 Male hind basitarsus with a distinct apicoventral process (Fig: 15); cercus never with large, conspicuous process. Female sternite 8 with all bristles on approximately equal in length (Figs 38, 111) 37 - Male hind basitarsus lacking a distinct apical process, if process is present, then male cerci with a conspicuous, large ventral process (in C. dentata but refer to figures 215 and 216). Female steraite 8 with 2 elongate bristles, distinctly longer than other bristles on sternite (Figs: 85, 219) 38

37 Gena entirely bright yellow. Posterior margin of male wing unmodified (Fig: 101). Surstylus divided into long, thin, finger-like processes (Fig: 103). Palearctic Coproica digitata (Duda, 1918) - Gena dark brown along posterior margin. Posterior margin of male wing with long, anteriorly directed bristles (Fig: 29). Surstylus with 2 blunt processes, without deep invaginations dividing them (Fig: 32). Cosmopolitan Coproica acutangula (Zetterstedt, 1847)

38 Surstylus with thickened posterior bristle curved medially or dorsally (Fig: 79); male cercus unmodified, without large ventral processes (Figs: 79, 80). Oriental, Eastern Palearctic Coproica coreana Papp, 1979

45 - Surstylus with thickened ventral bristle short and straight (Fig: 216); male cercus with a pair of conspicuous, large, curved processes (Figs: 215, 216) 39

39 Surstylus with ventral row of 4-7 bristles lateral to 2 distinctly thickened bristles; process on male cercus with 2 small setae along ventral margin, 1 small seta on small dorsal tooth. Mongolia Coproica dentataVapp, 1979 - Ventral margin of surstylus with patch of 20 or more small bristles (Fig: 215); process on male cercus without small setae on ventral margin or dorsal tooth (Figs: 215, 216). Palearctic Coproica pappi Carles-Tolra, 1990

40 Eyes small, eye to gena ratio typically 1.5:1 or less. Associated with gopher tortoise burrows or cacti. Nearctic 41 - Eyes large, eye to gena ratio 2:1 or greater. No known association with the above habitats. Nearctic or Neotropical 42

41 Interfrontal bristles in 3-4 subequal pairs. Mid basitarsus with paired sub basal ventral bristles (Fig: 23, 26). Associated with gopher tortoise burrows. Florida Coproica testudinea sp. nov. - 3 pairs of thick, erect interfrontal bristles, middle pair stronger than other 2 pairs. Mid basitarsus with 1 sub basal ventral bristle (Fig: 22). Associated with cacti. Southwest United States Coproica cacti (Richards, 1960)

42 Male cercus with small but distinct hooked processes (Figs: 90-91). Posterior lobe of surstylus broad and directed ventrally (Figs: 90). Female cerci free from tergite 10 (Fig: 96). Eastern North America, Caribbean Coproica diabolica sp. nov. - Male cerci without hooked processes. Posterior process of surstylus narrow or small, often dorsally curved (Figs: 54, 66, 280). Female cerci fused to tergite 10 (Figs: 61, 72, 284) 43

43 Cerci and tergite 10 glabrous on anterior half; apical bristles on cerci modified into short, thick spines (Fig: 72). Posterior process on surstylus very small and pointed (Fig: 72). Central and South America Coproica brachystyla sp. nov. - Cerci and tergite 10 completely pollinose; apical bristles sinuate, not modified into spines (Figs: 61, 284). Posterior process on surstylus elongate, slightly up curved (Figs: 54,280) 44

44 Posterior process on male surstylus very short (Fig: 54); postgonites symmetrical, rounded at tip (Figs: 56, 57). Sternite 10 complete, without floating sclerite between lateral arms (Fig: 62). Central and South America Coproica bispatha sp. nov.

46 - Surstylus with long distal process with rounded tip (Fig: 280); postgonites asymmetrical, left postgonite rounded apically (Fig: 282), right postgonite hooked apically (Fig: 283). Sternite 10 with a small floating sclerite between lateral arms (Fig: 284). Widespread in North and Central America Coproica urbana (Richards, 1960)

47 Species Accounts Coproica acutangula (Zetterstedt, 1847) (Figures: 7, 14-16,29-40)

Distr.: Afrotropical: Zaire; Australasian/ Oceanian: Hawaii (USA); Nearctic: Canada (AB, AT, BC, MB, ON, QC), USA (AK, AR, AZ, CA, CO, FL, GA, ID, IN, KY, LA, MD, MN, NC, NH, NJ, NM, OH, OK, OR, PA, TN, TX, UT, VA, WA, WI, WY); Neotropical: Bermuda, Colombia, Costa Rica, Dominican Republic, Ecuador, Guatemala, Honduras, Jamaica, Mexico (BCS, CAM, CHU, COA, DUR, JAL, MEX, MOR, NAY, VRC), Puerto Rico, Venezuela; Oriental: Pakistan, Taiwan; Palaearctic: Afghanistan, Algeria, Andorra, Austria, Belgium, Bulgaria, Croatia, Czech Republic, Denmark, Estonia, Finland, France, Germany, Georgia, Great Britain, Hungary, Italy, Japan, Kazakhstan, Kirghistan, Latvia, Macedonia, Mongolia, Netherlands, Norway, Poland, Roumania, Russia (CET, FE, NET, SET, WS), Slovakia, Spain, Sweden, Switzerland, Tadjikistan, Uzbekistan, Yugoslavia. Limosina acutangula Zetterstedt, 1847: 2499 [male]. Type locality: Sweden, Paradislyckan nr. Lund. ST male (MZLU, cf. Kim, 1972a: 208). Coproica acutangula. - Rondani, 1880: 42 [generic combination, male, diagnosis]; Papp, 1973c: 27 [key, illustr.]; Papp, 1984: 81 [Palaearctic catalog]; Marshall, 1989b: 603 [Australasian/Oceanian catalog]; Lachmann, 1996: 759-768 [morphology, male & female genitalia, copulation, illustr.]; Rohacek, 2007b: 114-116. Limosina (Coprophila) acutangula. - Duda, 1938: 162-163 [subgeneric combination, redescription, illustr.]. Leptocera (Coproica) acutangula. - Richards. 1960c: 205 [generic combination, diagnosis, illustr.]; Richards, 1965c: 725 [Nearctic catalog]; Tenorio, 1968: 196, 199-200 [diagnosis, male genitalia, illustr.]; Richards, 1980: 618 [Afrotropical catalog]. Coprophila acutangula. - Schumann, 1961: 298-299 [generic combination, egg, illustr.]; Schumann, 1962: 422, 440 [larva, illustr.]; Hinton, 1981: 747 [egg, illustr.]. Limosina (Coprophila) acutangulata. - Vanschuytbroeck, 1942: 11 [misspelling, list]. Borborus pusillus. - Meigen, 1830: 296 [misidentification, nee Copromyzapusilla Fallen, 1820]; Morge, 1976: PI. 163, Figs 12a,b [Meigen's original illustr.]. Heteroptera pusilla. - Macquart, 1835: 570 [generic combination]. Limosina (Coprophila) pusilla. - Duda, 1918: 206-208 [generic combination, redescription, illustr.]; 1938: 162-163 [redescription]. Leptocera (Coprophila)pusilla. - Duda, 1925: 205 [generic combination]. Leptocera (Heteroptera)pusilla. - Richards, 1930: 306 [subgeneric combination]. Coprophila pusilla. - Hammer, 1941: 250-251 [generic combination, egg, illustr.].

48 Body length 1.5 - 2.5 mm. Body colour dark brown. Face brown, frons dark brown.

Interfrontal bristles in 4-5 (usually 5) pairs; postocellar bristles slightly convergent; 2 pairs of small postocellar setulae. Gena frequently yellow on anterior 1/3 to 2/3, occasionally brown to dark brown; eye to gena ratio approximately 2.3:1; 8-10 genal setae with anterior seta twice as long as posterior. Scutellum with 25-30 discal setae distributed evenly between basal 1/6 and basal 1/4; 3-4 lateral bristles between larger anterior and posterior marginal scutellar bristles. Katepisternum with 2 small subequal dorsal bristles, each approximately 1/5 as long as the distance between dorsal margin of katepisternum and wing base. Legs brown, fore coxa yellow. Distal 1/2 of male fore basitarsus conspicuously quadrate. Mid tibia with additional small anterodorsal bristle at

1/8; proximal posterodorsal bristles present at approximately 1/4 and 1/2. Mid basitarsus with small, stout anterior bristles at 1/2 and 3/4, 1 sub apical bristle, 1 stout, sub-basal anteroventral bristle, and a strong anteroventral bristle at 1/2 with a smaller posteroventral bristle just distal to it. Male hind basitarsus with conspicuous ventral process. Second costal sector 1.1-1.9 times length of third costal sector. Costal bypass short, terminating within 5 vein widths of R4+5. Anal vein with distinct angulate bend.

Crossvein dm-cu on male wing with kink at anterior 1/3, intersection with cu acute and conspicuous. Posterior margin of male wing with distinct concavity at 3/4, row of enlarged ventral hairs along concavity, directed anteriorly.

Male terminalia:

Sternite 5 crescent-shaped, anterior margin concave, posterior margin slightly produced medially with posteromedial patch of dense microsetae and a posterior row of enlarged bristles; lateral bristles enlarged. Synsternite 6+7 with distinct medial bend. Epandrium

49 with enlarged pair of dorsolateral bristles; subanal plate absent, cerci and epandrium medially separate below anal opening; cercus fused to epandrium, ventral margin relatively flat and straight, 2 or 3 enlarged bristles present in a cluster posteromedially on.

Surstylus with posterior and posteroventral processes; posterior process with elongate thickened black bristle at base, 2 subapical thickened, toenail-like bristles, and 4-6 apical bristles; posteroventral process short, rounded, heavily bristled. Postgonites symmetrical,

straight, with distinct pointed posterior process half way along its length, and row of 3

small anterior setae distal to posterior process. Basiphallus gently curved, tubular, open posteriorly; posteroventral process elongate, blade-like. Lateral sclerites rounded, with

elongate, anteriorly directed process; central sclerite fusing with lateral sclerites apically;

additional sclerite ventral to central sclerite, fused ventrally with central sclerite apically and in centre; ventral surface of distiphallus bulbous. Apical membranes with sparse,

lightly sclerotized spines.

Female terminalia:

Tergite 8 long laterally, short dorsally. Tergite 10 pollinose, narrowly fused to cerci posterolaterally; cerci rounded-triangular, pollinose, with long and sinuate bristles.

Sternite 7 broad, rounded-rectangular. Sternite 8 simple, oval shaped, with a posterior

row of 4 small setae. Sternite 10 not wrapped posteriorly around tip of abdomen, lateral

arms curved dorsally, medial and lateral pairs of enlarged bristles. Spermathecae

spherical; duct length approximately equal to length of spermathecal bulb.

Specimens Examined:

CANADA: AB: Stettler, 30.viii.1980, S.A. Marshall (8& 6$); Wagner Bog, 2- 9.viii.l985, pan tp. 2, leg. A.T. Finnamore; (1<$, PMAE); same data except 23- 27.viii.1985, pan tp. 8 (1$, PMAE); lOmi. N of Nordegg, 20.vii.87, around cow parsnip roots, S.A. Marshall (1$); Jumping Pound Ck., 7.viii.l980, dung vac, S.A. Marshall

50 (1$); lOmi. E of Patricia, 16-17.viii.1980, prairie, R.S. Anderson, S.A. Marshall (1$); Cypress Hills, 14.vi.1980, cow dung S.A. Marshall (1$). BC: Cranbrook, l.viii.1980, horse dung, S.A. Marshall (11(5, 21$); near Cranbrook, 16.vii.1980, sweeps, G. Gibson (1$, CNCI); Wyndell, viii.1980, bear dung vac, S.A. Marshall (1$); Ainsworth, Woodbury Creek, 5.vii.l980, S.A. Marshall (1$). ON: Horse C. acutangula {AS); C. acutangula, 6.vi, v IN (1$); Algonquin Prov. Pk., Swan Lk. stn., 23-30.vi.1994, Scott Lk. Survey, A5 shore pans (1$); Algonquin Prov. Pk., Swan Lk. stn., 45°29'15"N, 76°43'20"W, 2-12.vii.1993, Scott Lk. Survey, pan A2, lakeshore, Larson, Marshall, Barr (lcJ); Algonquin Prov. Pk., 0.5 km S of Billy Lk., 21-28.vii.1984, bog, moose dung, Kpendreigh (1 $); Arkell, 20.vi.1956, D.H. Pengelly (1(5); Arkell, 18.ix.1993, on horse dung, CD. Jones (1$); Belfountain, 10.vi.1981, mushroom, S.A. Marshall (1$); Blair, RARE, The Dells, 43°22'55"N, 80°20'32"W, 20.vi-17.vii.2006, malaise, Bergeron & Cheung (1$: debu00268440); Chaffeys Locks, Queens Univ. Bio. Stn., 12.ix.80, human dung, field, S.A. Marshall (1$); Damascus Wylde Lk., 25-3l.viii. 1992, pan tps., S.A. Marshall (1(5); Guelph, Hanlon Pkwy. & Laird Rd., 18.iv.2005, on deer carcass, J. Klymko (1$: debu00242722); Guelph, 3.vii.l978, L. Cartwright (2(5); Guelph, Arboretum, 9.viii.l984, field, mushroom tps., B.V. Brown (\<$, 19); Guelph, Arboretum, 3-13.X.1984, deciduous, mushroom tps., B.V. Brown (1$); Guelph, 30.ix.1979, on window, S.A. Marshall (1$); Guelph, 5-8.vii.1982, pan tp., K.W. Barber (1$); Guelph, University, 23.vi.1984, pan tp., K. Gilbert (1$); Hamilton, Royal Botanical Gardens, 15.vi.98, RET - carrion, V. Carney & K. Scott (1$); Healy Falls, 18.vii.1979, W.A. Attwater (1(5, 2$); Healy Falls, 7.vii.l979, horse dung, W.A. Attwater (3

51 (IS); Washago, 44°44.76'N, 85°18.73'W, PAN 6B: carrion tp., 10-19.V.98, Cristopherson (1$); Windsor, Ojibway Prairie, 18-19.vi.2002, on dung, O. Lonsdale (1 $: debuOl 118682); Wylde Lk. Bog, 8km E of Arthur, 16-22.vi.1987, W4E pt4, floating mat, D. Blades (1<5*). QC: Hull, 21.ix.1983, wet flooded forest, L. Dumouchei (1?*). COLOMBIA: Antioquia, Venecia, 1335m, ii.1984, manure pile, O. Piedrahita (\S, CNCI). COSTA RICA: Guanacaste, Estac. Cacao, 1000-1400m, Lado suroeste del Volcan Cacao, II curso Parataxon, vi.1990, LN823300,375700 (\S, INBC); Guanacaste, Estac. Mengo, SW side Volcan Cacao, 85 28'10"W, 10 55'43"N, 1100m, ii.1989, GNP Biodiversity Survey (1$, INBC); Guanacaste Ref. Nac. Fauna Silvestre Rafael Lucas Rodriguez, Palo Verde, 10m, Nov. 1990, D. Acevedo, LN259000,388400 (AS, 7?, INBC); Limon, Circuito Reviente Pecho Espavel, R.B. Hitoy Cerere, 150-650m, 12.X.1993, LS184200_643300#2377, G. Carballo, M.A. Zumbado (14$, 6$, INBC); San Jose, San Gerardo de Dota, 9°33'N, 83°48'W, 9.viii.l995, roadside, sweep, S.A. Marshall (1$: debu00206925); San Jose, San Carlos, Riosparaiso Res., Pecari Stn., 16km NNE of Quepos, 9°33'53"N, 84°7'32"W, 400m, 22-27.ii.2006, S.A. Marshall (4$: debu00259741, 00259643, 00259671, 00259601); San Jose, San Gerardo de Dota, near lodge, 2000m, 9°33'N, 83°48'W, 9.viii.l995. stream in pasture, sweeping, S.A. Marshall (IS: debu00176578; 2$: 00176580, 00176586); San Jose, San Gerardo de Dota, 2000m, 9.viii.95, stream, in pasture near lodge, S.A. Marshall (2S)', San Jose, Zurqui de Moravia, 1600m, v.1991, P. Hanson (\S). DOMINICAN REPUBLIC: Barahona, 4km E of Barahona, 28-29.ix.1985, dung, R.F. Woodruff (IS); La Cienega 2nd refugia, 12.U989, dung, J.E. Swaan (5$, 5$); Monte Cristi, 3km N of Villa Elisa, 1-3.X.85, dung, R.E. Woodruff (AS); Pedernales, La Abeja, 38 km NNW of Cabo Rojo, 18-09N, 71-38W, 1250m, 15.vii.1987, J.E. Rawlins & R.E. Davidson (2c?, 1$, CMNH); Puerto Plata, 22.U989, sweep on wet trail, S.A. Marshall (3cT, 3$); San Juan, El Capa, 1km off rd. to Vallejuela, 21.V.1985, 81 tp., Nunez, Woodruff & Stanger (5S). ECUADOR: Napo, SierrAzul Res., 14km W of Cosanga, 2200m, 0°40'55S, 77°56'9W, 9.V.2002, O. Lonsdale (2$: debu00195604, 00186777); same as previous except 9-11.V.2002 (2$: debu00180380, 00180320); 9-10.V.2002, malaise, Buck & Paiero (2$: debu00202516, 00202664); Napo, SierrAzul Res., 14km W of Cosanga, 5.xi.l999, sweeping disturbed trail, S.A. Marshall (1$: debu00115156*-illustr; 3$: 00115155, 00115162-wing, 00115116); Napo/Pichincha, Papallacta Pass, 4200 m, 0 19'15S, 78 11'51W, 29.iv- ll.v.2002, paramo, yellow pans, Marshall & Buck, (\S, debu00202177); Pichincha, Maquipucuna Biol. Res., 1200m 0°7'34N, 78°37'57W, 27-28.iv.2002, sweep, M. Buck, (\S: debu00195256-wing; 1$: 00202864). GUATEMALA: Baha Verapaz 6.6km W of Chilasco, 1700m, 30.V.1991, dung, H. Howden (1$). HONDURAS: L. de Yajoa, 2600', l-2.vi.1994, dung tp., Howden? (\S). JAMAICA: Montego Bay, 18.U985, ex. horse manure, J.E. Corrigan (10$, 6$); Olar. Portland Ridge, ll-15.viii.1974, forest, S. Peck (IcT); St. Ann, Discovery Bay, 3-6.ix.1974, dung, S. Peck (1$); St. Catherine, 28.vii- l.viii.1974, dung under trees in horse pasture, Peck (IS)- MEXICO: Campeche, 87 mi E of Escarcega, 800', 8-14.viii.1971, trop. semi-evergreen, human dung, A. Newton (1$); Coahuila, Boquillas, 24-26.ix.1979, pig dung tp., R.E. Woodruff, G. Fincher (18c?, 29$); Jalisco, 14 mi W of Atenquique, hardwood forest, 7900', dung, A. Newton (AS); Jalisco, 18 mi W Atenquique fir forest, 9300', dung, A. Newton (2S); Mexico State, 1 mi E of Ixtapan de la Sal, 6200', km 78, 31.viii-6.ix.1971, trop/Dec/Jun, human dung, Newton (IcT); Mexico State, 1 mi NE of Tenancingo, 7100', 31.viii-6.ix.1971, oak pine medr.,

52 human dung, A. Newton (3$); Morelos, 4 mi W of Tres Cumbres, 9000', km 6 3/4, 29.viii.-4.ix. 1971, oak, human dung, A. Newton (lc?); Morelos, 4 mi S of Tres Cumbres, km 57 3/4, 8500', 29.viii-4.ix. 1971. human dung, A. Newton (1$); Morelos, 7 mi W of Tres Cumbres, 9600', km 12, 29.viii-4.ix. 1971, oak fine fir dung, A. Newton (1$). PUERTO RICO: Toro Nepro Nat. For., Rd.143, km 2.5, 3-4.vii.1977, human dung tp.., J. Micheli & R.E. Woodruff (lc?). UNITED STATES OF AMERICA: AZ: Cochise Co., 5100', Coronado Nat. For., Chiricahau Mt. Stewart cpgd., 16-18.viii.1984, road dung, L.B. Carlson (13$, 19$); Cochise Co., Coronado Natl. Mon., ll-15.viii.1984, dung tp., B.V. Brown (30(5*, 61$); Cochise Co., Coronado Nat. For., Coronado Nat. Mem., ll-15.viii.1984, dung tp., L.B. Carlson (lc?, 5$); Cochise Co., Chiricahua Mts., 6mi ESE of Portal, 4150', l.ix.1970, desert grass, human dung, A. Newton (4c?, 8$); Cochise Co., Chiracahua Mts., 7000', 15-2l.vii. 1978, dung, O. Kukal (1$); Cochise Co., Chiracahua Mts., 15-21.viii. 1978, dung, O. Kukal (3c?); Cochise Co., Huachuca Mts,, 6000', Miller Canyon, ix.1970, oak woodland, dung, A. Newton (1$, 1$); Cochise Co., Huachuca Mts., Carr Canyon., 7100', 1.x. 1970, oak & evergreen, human dung, A. Newton (24c?, 18$); Cochise Co., Huachuca Mts., Ramsey Canyon, 5600', ix.1970, human dung, A. Newton (1$); Cochise Co., near Portal, Stewart cpgd, 16-19.viii.1984, dung tps., B.V. Brown (5c?); Cochise Co., Miller Canyon, 3.vi.l991, sweep along trail, J.E. Swann (lc?); Cochise Co., SWRS, 18-23.viii.1984, dung tp., B.V. Brown (2c?, 3$); Cochise Co., Portal, SWRS, 18-23.viii.1984, mushroom tps., B.V. Brown (1$); Cochise Co., Portal, SWRS, Chiricahua Mts., 18-23.viii. 1984, FIT, B.V. Brown (lc?, 5$); same as previous except yellow pan (2$); Coronado Co., Bonito Pk., near Flagstaff, 5- 9.viii.l984, dung tp., B.V. Brown (5c?); Graham Co., 2.4mi W on Hwy 336 from Hwy 191, 3800', 19-26.viii.1993, J.B. O'Hara (1$); Pima Co., Santa Catalina Mts., Mt. Lemmon, 9000', dung, A. Newton (1$); Pima Co., Santa Catalina Mts., Mt. Lemmon, 9000', viii.72, fir forest, dung tps., A. Newton (lc?); Santa Cruz Co., 7mi. WNW of Nogales, Calabase Canyon, 3850', ix.1972, desert grass dung, A. Newton (3$); same as previous except desert grass dung (2c?); Santa Cruz Co., 8 mi NNW of Nogales, Walker Canyon, 3900', ix.1970, oak woodland, dung, A. Newton (7c?, 7$); Santa Cruz Co., Patagonia Lk. St. Pk., 9-11.viii. 1984, dung tp., B.V. Brown (lie?, 17$); Santa Cruz Co., Patagonia Lk. Rec. Area, 4500', 9-11.viii. 1984, dung tp., L. Carlson (2c?, 3$); Santa Cruz Co., Santa Rita Mts., Madera Canyon, 5500', ix.1970 oak, human dung, A. Newton (15c?*, 4$). AR: Garland Co., 1.2mi N, 6-8.iii.1977, pig dung tp., Woodruff & Wiley (3c?, 5$); Montgomery Co., 20mi W Glenwood, T45 R27W, 15.vi.-28.vii.1994, H.W. Robinson (1$); Montgomery Co., 5.4mi E IdaHwy 270, 6-8.iii.1977, pig dung tp., Woodruff & Wiley (4c?, 20$); Polk Co., 13mi NW of Mena Rich Mt., 2800', 1- 3.vi.l979, mesic oak hickory, S.&J. Peck (lc?); Scott Co., 7mi E Y City, Jet., Hwy270 & Rt71 on 270, 6-8.iii.1977, pig dung tp., Woodruff & Wiley (4c?, 4$); Wash Co., Devils Den St. Pk., Devils Den Cave, 28-31.V.1979, S.&J. Peck (2c?, 1$*); same as previous except dung (1$); Wash Co., 3mi S of Devils Den St. Pk., 28-31.V.1979, oak-hickory, S.&J. Peck (lie?*, 6$). CO: Roatte Co., 5mi NE of Clark Hinman cpgd., 7600', 23- 25.vi.1972, dung tp. (1$). FL: Alachua CCo., 6.iii.l980, on cow dung, J. Givinski (1$); Alachua Co., Gainesville, 2-5.vii.1982, pig dung, horse pasture, K.L. Rench (lc?, 7$); Alachua Co., Gainesville, Hogtown Ck., 12.X.1976, P.M. Choate & R.E. Woodruff (3c?, 3$); Alachua Co., Rd. 136., 8.V.1983, pig dung tp., K.W. Vick (3c?, 1$); Archbold Bio. Sta., 12-16.xii.1985, dung pan, scrub-sand, S.A. Marshall (lc?, 2$); Archbold Bio. Sta.,

53 17-21.xii.1985, scrub dung, S.A. Marshall; (3c?, 2$); Archbold Bio. Sta., 12-16.xii.1985, scrub, dung pitfall tp., S.A. Marshall (lc?, 1$); Highlands Co., 3mi N of Avon Pk., 14- 18.vi.1982, pig dung tps. in sabal scrub, R.E. Woodruff & K.L. Rench (lc?, 4$); same as previous except pig dung tps. in gopher burrow (2c?, 3$); Highlands Co., 7mi NE of Cornwell, 14-18.vi.1982, pig dung tps. in pasture R.E. Woodruff & K.L. Rench (1$); Highlands Co., Highlands Hammock St. Pk., orange grove, 14-18.vi.1982, pig dung, Woodruff & Rench (3$); Highlands Co., Lk. Placid, 2-3.xi. 1983, pig dung tp., R.E. Woodruff (2$); Polk Co., Rt.27, 7mi N of 14, 2-3.xi.1983, pig dung tp., R.E. Woodruff (4c?, 5$); Manatee Co., Longboat Key, 3-4.vii.1983, pig dung tps., R.E. Woodruff (2c?, 2$); Marion Co., Ocala Nat. For., Oklawaha Swamp, ll-12.vi.1984, dung tp., S.A. Marshall (1$); Liberty Co., lOmi SW of Juniper, Rt. 12, 26.iii.1983, pig dung, turkey- oak, Woodruff & Thomas (lc?, 1 $); Okaloosa Co., 4.5mi N of Holt, Blackwater R. State For., A&M Res. Sta., 15-17.vi.1978, pig dung, L. Stange; (2$*); Orange Co., E Orlando, 2-3.vii.1982, pig dung, sand-pine scrub, Woodruff (1$); Suwannes Co., 8.7mi W of O'Brien, S349, 25.iv. 1977, pig dung tp. #6, gopher tortoise burrow, Woodruff & Woley, (Id1). GA: Clinch Co., 8mi S of Fargo, US 441, 5-25.vi.1984, dung tp., S.A. Marshall (1$); Rabun Co., Black Rock Mtn. St. Pk., 26-27.vi.1984, human dung, S.A. Marshall (lc?). HI: Maui Is., Haleakala, Crater Paliku pasture, 1980m, 16.iv.1998 (1$); Waipio Valley, 26.xii.1996, sweep along road, S.A. Marshall (1$, 1$). IN: Lawrence Co., Hoosier Nat. For., Donaldson Trail, 19.V.1991, J.E. Swann, (lc?, 38$). KY: Whitley Co., Cumberland Falls St. Pk., 29-30.vi.1984, dung tp., S.A. Marshall (2$). LA: East Baton Rouge Par., 16.iv.1999, black bear carrion, E.J. Watson (lc?, debu00153121). NH: Strafford CCo., lmi SW of Durham, 24.vii-6.viii. 1987, D.S. Chandler (1$). NJ: Burlington Co., Delanco, Hawk Is., 2-16.vi.1969, scrub hardwoods, dung, A. Newton (lc?). NM: Bernalillo Co., 7mi S of Tijerus, 2400', 23-26.viii.1975, car. tp., pinyon- juniper, S. Peck (46*, 4$); Bernalillo Co., Isleta, 4900', 16-20.vi.1979, floodplain forest, carrion, S&J Peck (3c?, 5?). NC: Bladen Co., Singletary Lk. St. Pk., 34°35'00"N, 78°27'30"W, 19-22.V.2003, pan tps., Marshall & Paiero (lc?: debu00252170); same as previous except malaise (1$: debu00227364); Jackson Co., Blue Ridge Pkwy., Grassy Ridge Mine Overlook, 5250', 5-27.vi.1984, mushroom tp., S.A. Marshall (1$). OH: Champaign Co., 6mi NW of Urbana Hume's Woods, l-5.vi.1977, pig dung, R.E. Woodruff (2$); Champaign Co., 6mi NW of Urbana Hume's Woods, l-5.vi.1977, R.E. Woodruff (2c?, 1$). OK: Lattimer Co., 5mi W of Red Oak, 15-21.V.1977, dung tp., K. Stephen (3c?, 12$); same as previous except 5-ll.vi.1977 (2$*-illustr.); 8-14.V.1977, dung (2$, 7$); Murray Co., Sulphur, Chickasaw Rec. area, 3-5.vi.1979, woody ck. bottom, S.&J. Peck (lc?); Wichita Mts., 5-7.vi.1979, dung tps. (1$). PA: Trout Run, 28.ix.1979, dog shit, S.A. Marshall (lc?). SC: Columbia, Riverbanks Zoo & Botanical Gardens, N34°00.582', W81°04.560', 22.vi.2007, M.P. Nelder (lc?, 1$); Greenville, Greenville Zoo, N34°50.581\ W82°23.237\ 14.ix.04, M.P. Nelder (lc?, 4$); same as previous except dated, 29.V.2007 (lc?). TX: Angelina Co., Angelina Natl. For., Boykin Cemetery, 13-17.vi.1993, spring fen, dung, S.A. Marshall (2c?: debu00128175, 00128205; 3c?); Angelina St. For., Bouton Lk. cpgd., 12-14.vi.1993, dung/vac. tps., S.A. Marshall; (1$); Big Bend Nat. Pk., Bouquilias Crossing, 25-26.ix.1975, pig dung tp., R. Woodruff & G.Fincher (3c?); Bouton Lk., 13-17.vi.1993, dung/vac. tps., S.A. Marshall, (lc?, 2$); Brewster Co., 3.5mi E of Marathon, 26-27.ix.1979, dung tp., R. Woodruff & G. Fincher (1$, 1$); Terrell Co., 30mi E of Sanderson, 23-27.ix.1979, on pig dung, B.

54 Woodruff & B. Fincher (1$); Warren, Kirby St. For., 12-17.vi.1993, dung tps., S.A. Marshall (1$). UT: Duchesne Co., Mirror Lk., 10300', 30.vii-12.viii.1979, S.&J. Peck, spruce fir meadow, carrion malaise (12$, 24$); Daggett Co., Dutch John, 6300', 2- 8.viii.l979, pinyon juniper, woodland, carrion, S.&J. Peck (1$); Summit Co., Henrys Fork cpgd., l-10.viii.1979, pine-aspen at stream, willows carrion-malaise, S.&J. Peck, (1$); Summit Co., Wasatch For., Bear Riv. cpgd., 8400', 30.vii-l 1 .viii.1979, malaise, streamside, S.&J. Peck (1$); Summit Co., NW side of Gilbert Peak, 11500-12500', 1- 10.viii.1979, tundra, carrion, S.&J. Peck (3$*, 5$). WY: Big Horn Co., 10 mi E of Lovell, 3800', 17-26.viii.1979, sagegrass-barren carrion, S.&J. Peck (1$); Big Horn Co., 30 mi E of Lovell, 9000', 18-26.viii.1979, spruce fir carrion, S.&J. Peck, (1$); Crook Co., Black Hills Nat. For., Cook Lk. Cons. Area Rd., 2.5km E of cpgd, 4-2l.viii. 1980, pan tps., cow dung, rocky hillside, pines, J.E. Swann (1$); Uinta Co., 6mi E of Mt. View, 6800', 1-1 l.viii. 1979, sagebrush, S.&J. Peck (3$); Uinta Co., 8mi E of Mt. View, 6800', 1-1 l.viii. 1979, sagebrush, S.&J. Peck (2$); Uinta Co., 8mi SE of Evanston, 7100', 30.vii-11.viii.1979, carrion, sage-grass, riparian, S.&J. Peck (8$*, 3$); Sublette Co., Daniel, 7200', 16-22.viii.1979, carrion sagebrush steppe, S.&J. Peck (1$). RUSSIA: SIBERIA: Altai Reg., Gorno Altai, 9.vii.l991, roadside sweeps near riv., S.A. Marshall (\

Comments:

Coproica acutangula was established as the type species for Coproica in 1996

(Wheeler and Swann, 1996). It is cosmopolitan in distribution and belongs to Clade 1, an

Old World group. Male Coproica acutangula can be distinguished from all other species of Coproica by a cluster of elongate setae on the posterior wing margin, dm-cu forming acute angle with CuA], and a ventral process on the hind basitarsus, (the latter character is shared with its sister species, C. digitata). The process on the hind basitarsus may also occur in C. dentata, yet this is unconfirmed (see comments under C. dentata Papp, 1979).

55 Coproica albiseta Papp, 2008

Distr.: Afrotropical: South Africa. Coproica albiseta Papp, 2008: 6 [both sexes, phylogenetic notes, illustr.]. Type locality: Republic of South Africa, Eastern Cape Pr. HT male (HNHM).

Coproica aliena Papp, 2008

Distr.: Oriental: Thailand. Coproica aliena Papp, 2008: 41 [both sexes, phylogenetic notes, illustr.]. Type locality: Thailand, Mae Taeng Elephant Camp. HT male (HNHM).

56 Coproica bifurcata sp. nov. (Figures: 17,21,41-52)

Distr.: Neotropical: Argentina, Bolivia, Brazil.

Body length 1.2 - 1.7. Body colour brown. Face and frons brown. Interfrontal bristles in 3-4 pairs; postocellar bristles convergent; postocellar setulae absent. Eye to gena ratio approximately 2:1; 4 genal setae in longitudinal row posterior to vibrissa. Scutellum with approximately 10 bristles loosely arranged in 2 rows between basal 1/4 and basal 1/2, posterior row approximately 1.3 times length of anterior row; 2 lateral bristles between larger anterior and posterior marginal scutellar bristles. Katepisternum with 2 small dorsal bristles, each approximately 1/5 as long as the distance between dorsal margin of katepisternum and wing base. Legs and fore coxa brown. Mid tibia with 1 proximal posterodorsal bristle at 2/5. Mid basitarsus with row of small, stout anterior bristles beginning at 1/5 and extending to apex, continuous row of small, stout anteroventral bristles, 2-3 small posteroventral bristles, variable in position. Second costal sector 0.8-

1.0 times length of third costal sector. Costal bypass short, terminating within 5 vein widths of R4+5. Anal vein sinuate, lacking an angulate bend.

Male terminalia:

Sternite 5 rounded rectangular, posteromedial patch of slightly denser microsetae; all surface bristles uniform in size. Synsternite 6+7 straight medially. Epandrium with enlarged pair of dorsolateral bristles; subanal plate absent, cerci and epandrium medially separate below anal opening; cercus fused to epandrium, ventral margin with strong posteromedial bristled processes, process on left cercus 2 times longer than process on right cercus. Surstylus with rounded posteroventral lobe lined with short bristles.

57 Postgonites asymmetrical, curved anteriorly; left postgonite thick, with pointed apex; right postgonite narrow distally, bifurcate at apex. Basiphallus gently curved, tubular, closed posteriorly, ventral margin unmodified. Lateral sclerites with sharp posterodorsal angle, row of 3 dorsal processes, proximal process sharp with narrow, pointed medial process, central process broad, blunt, distal process, narrow, sharp, and sinuate; distal ventral angle rounded, protruding. Central sclerite fused with lateral sclerites ventral to central process.

Female terminalia:

Tergite 8 reduced to 2 long lateral sclerites with medial membrane connecting lateral sclerites. Tergite 10 pollinose, narrowly fused to cerci posterolaterally; cerci rounded triangular, pollinose, cereal bristles long and sinuate. Sternite 7 broad, rounded rectangular, posterior projection, broad, deeply concave, appearing like 2 processes.

Sternite 8 with broad anterior process with 2 small lateral points, large pair of central bristles and a row of four small setae anterior and posterior to it. Sternite 10 not wrapped posteriorly around tip of abdomen, posterior margin bluntly pointed, medial and lateral pairs of enlarged bristles present. Spermathecae elongate, pear-shaped; duct length approximately 3 times length of spermathecal bulb.

Type Material:

HOLOTYPE: ARGENTINA: Salta, Rosario de Lerma, 24.ii.1992, swept over bulldozed road, S.A. Marshall (1(3*).

PARATYPES: ARGENTINA: Same as holotype (10^**-illustr-wing, 13$**-illustr.); same as holotype except swept over foul ditch (1$); Jujuy, Cochinoca, 3700m, 26.ii.1992, sweep small stream, llama dung, debris, S.A. Marshall (2(5**, 2$*); Jujuy, Tres Cruces lakeside, 3700m, 26.ii.1992, sweep, S.A. Marshall (1$, 1$); Jujuy, Abra Pampa, 3500m, 26.ii.1992, llama dung, S.A. Marshall (1$); Salta, Rosario de Lerma, INESALT, 17-28.ii.1992, yard pan tps., S.A. Marshall (1$); Salta, El Ray Nat. Pk., 900m, Aguas Negras Tr., ll-15.xii.1987, forest malaise tp., FIT, S.&J. Peck {AS*, 10

58 $***); Salta, El Ray Nat. PL, 900m, Rio La Salsa, 5-10.xii.1987, malaise tp., FIT, open stream side in forest, S.&J. Peck (9^***, 4?*); Salta, El Ray Nat. Pk., 1000m, Pozo Verde tr. km 7, 5-15.xii.1987, malaise FIT, yungas forest, S.&J. Peck (1$, 1 ?).

Other Specimens Examined:

ARGENTINA: Salta, Canyada la Gotera, 19.ii.1992, sweep/wet litter, S.A. Marshall (4c?, 1?); Salta, 22km N of La Caldera, 1550m, El Ucumar, 2-30.xii.1987, malaise, subtropical humid forest, S.&J. Peck (1$); Salta, 1500m, La Caldera Camino, la Cornisa, 27.ii.1992, roadside forest sweeps, S.A. Marshall (2<5*); Salta, Rosario de Lerma, INESALT, 16-28.ii.1992, yard malaise, S.A. Marshall (2

Etymology:

The specific name means "two-forked" and references the forked tip of the right paramere.

Comments:

Coproica bifurcata can be distinguished from its closest relative, C. hirtuloidea, by its bifurcate and apically broad right postgonite (narrow and sinuate apically in C.

hirtuloidea). These two species are difficult to separate on the basis of external

characters, and dissections of male specimens may be required for accurate identification

of specimens from Chile, Argentina, and Bolivia where the ranges of C. hirtuloidea and

C. bifurcata overlap.

59 Coproica bispatha sp. nov. (Figures: 53-64)

Distr.: Nearctic: USA (AZ, CO, TX); Neotropical: Barbados, Brazil, Costa Rica, Dominican Republic, Galapagos Is. (Ecuador), Honduras, Jamaica, Mexico (CHP, ROO, TAB), St. Kitts, Venezuela.

Body length 1.1 - 1.5 mm. Body colour light brown to brown. Face light brown, frons brown. Interfrontal bristles in 3 pairs; postocellar bristles convergent, postocellar serulae absent. Eye to gena ratio approximately 2:1; 3 genal setae in longitudinal row posterior to vibrissa with anterior seta twice as long as posterior setae. Scutellum with 10-12 discal scutellar setae arranged in 2 rows between basal 1/4 and basal 1/2, setae in posterior row approximately 1.5 times length of anterior row; 2 lateral bristles between larger anterior and posterior marginal scutellar bristles. Katepisternum with 2 small, subequal dorsal bristles, each approximately 1/4 the distance between katepisternum and wing base in length. Legs and fore coxa brown. Mid tibia with 1 proximal posterodorsal bristle at 2/5.

Mid basitarsus with row of small, stout bristles extending from basal 2/5 to apex, stout anteroventral basal bristle, stout anteroventral bristles at 2/5 and 1/2 with small paired posteroventral bristles, specific arrangement of bristles on ventral surface of mid basitarsus may vary between individuals. Second costal sector 2 0.9-1.2 times length of third costal sector. Costal bypass long, extending 7 or more vein widths beyond R4+5.

Anal vein sinuate, lacking an angulate bend.

Male terminalia:

Sternite 5 rounded rectangular, bristles on surface uniform in size. Synsternite 6+7 straight medially. Epandrium with enlarged pair of dorsolateral bristles; subanal plate absent, cerci and epandrium discontinuous below anal opening; cercus fused to

60 epandrium, ventral margin unmodified, rounded, with 1 enlarged bristle. Surstylus with blunt, short posterior process with several short bristles at apex, ventral margin lined with small setae. Postgonites slightly asymmetrical, postgonites straight, narrow, with rounded spoon-like apex, right postgonite with small apical notch. Basiphallus gently curved, tubular, posterior margin open, very short posteroventral keel. Lateral sclerites rounded, 2 short dorsal processes near apex; central sclerite fused with lateral sclerites apically. Apical membranes broad.

Female terminalia:

Tergite 8 reduced to 2 lateral sclerites with medial connecting membrane. Tergite 10 pollinose, broadly fused to cerci; cerci pollinose, apical setae short and sinuate. Sternite 7 broad with deeply concave posterior projection, appearing like 2 lateral processes.

Sternite 8 with narrow anterior process, 2 lateral patches of 3 small setae, row of 4 posterior setae separated medially into 2 pairs. Sternite 10 wrapped around tip of abdomen with one enlarged pair of medial bristles. Spermathecae conical; duct length 3-

4 times length of spermathecal bulb, short sclerotized portion of duct adjacent to spermathecal bulb, with small constriction at its base.

Type material:

HOLOTYPE: ECUADOR: Galapagos, Floreana, Agriculture zone, 360m, 22.ii.1989, sweeping, B.J. Sinclair (ltf, CNCI).

PARATYPES: BARBADOS: Welchman Hall, Gully ii.1979, dung tp., leg, S. Peck (2$). BRAZIL: Minas Geras, Congonhas Serra, 22.ii.1990, sweep along creek, horse dung, S.A. Marshall (1$, MZSP). COSTA RICA: San Jose, Zurqul de Moravia, 1600m, v.1991, P. Hanson (1$, INBC). DOMINICAN REPUBLIC: Alcoa Rd., 600m, 17- 20.1.1989, FIT, carrion, S.A. Marshall & J. Swann (1^); Barahona, 7km NW of Paraiso, 200m, rainforest remnant, 27.xi.1991, fungus on logs, 91-338, S&J Peck (2& 29); Pedernales, 28km N of Cabo Rojo, 760m, evergreen dry forest, 29.xi-3.xii. 1981, FIT, 91- 349, Masner & Peck (39, 4$); same as previous except carrion tp., 91-348 (3$, 79); carrion tp., 91-343, S.&J. Peck (1$); Pedernales, La Abeja, 38 km NNW of Cabo Rojo,

61 (18-09N, 71-38W), 1250m, 15.vii.1987 J.E. Rawlins & R.E. Davidson (lc?, CMNH); near Cabo Rojo, Alcoa Rd., km 26, 17-20.i.l989, FIT, S. Marshall & J. Swann, (1$). ECUADOR: Galapagos: Same as holotype (13c?*, 18$, CNCI); Isabela, 7km WNW ofSt. Tomas, near Campanento Pumas, 640m, 8.vii.l985, sifted horse dung, 83-212, S.&J. Peck (lc?, QCAZ); San Cristobal, Casetta, pasture, 560m, 19.ii.1989, horse dung, B.J. Sinclair (lc?, CNCI); San Cristobal, Casetta, 560 m, pampa, 16-19.ii. 1989, dung tp., B.J. Sinclair (30c?, 20$, CNCI); St. Cruz CDRS, Arid zone, 3l.i-9.ii. 1989, tortoise dung tp., B.J. Sinclair (6c?, 3$, CNCI); St. Cruz, 2km N of Bellavista, 360m, 2.vii.l985, guava thicket, agricultural area, FIT, S.&J. Peck (8c?-BOLDx2, 13$, QCAZ); same as previous except dated 14.v-13.vii.1986 (5c?, 4$, QCAZ); St. Cruz, 25km N of Bellavista, 360m, 13.vii.1985, guava thicket agricultural area, malaise, FIT, S.&J. Peck (lc?*-illustr., 2$*- illustr., QCAZ); St. Cruz, 4km N of Bellavista, Media Luna, 620m, 14.v.-13.vii.l985, 85- 158, S&J Peck (3$, QCAZ); same as previous except Miconia zone, FIT (lc?, 2$, QCAZ); miconia zone, FIT (2.vii) (3c?, 7$**-BOLDxl, QCAZ); St. Cruz, 4km N of Bellavista, Media Luna, 620m, 21-31.V.1985, Miconia zone, dung tps., S.&J. Peck (lie?, 6$, QCAZ); St. Cruz, agricultural zone, Bellavista, 160m, l-9.iv.1989, field, FIT, 89- 205, Peck & Sinclair (lc?, 5$, CNCI); St. Cruz, agricultural zone, l-9.iv.1989, malaise, 89-204, Peck & Sinclair (1$, CNCI); St. Cruz, Los Gemelos, 600m, 31.i-4.ii.1989, forest dung tp., B.J. Sinclair (lc?, CNCI); same as previous except add Scalesia, BS43 (2$, CNCI); St. Cruz, Los Gemelos, 31km N of Sta. Ros., 570m, 15.vii.1985, Scalesia forest, FIT & malaise, S.&J. Peck (1$, QCAZ); same as previous except 23.vi.1985, 85-188b (2$, QCAZ); St. Cruz, Media Luna, 500m, Miconia, l-29.ii.1989, FIT-trough, 89-39, Peck & Sinclair (lc?, 1$, CNCI). HONDURAS: Dpto. F. Morazan, El Zamorano, 22- 29.vii.1970, blacklight tp., G.F. Freytag (1$, debu00020364); Zamorano, 30.V.1994, H.&A. Howden (1

62 Etymology:

The name bispatha means "two spoons" and refers to the postgonites, which have a rounded spoon-like terminus and are symmetrical, unlike C. urbana and C. brachystyla.

Comments:

Coproica bispatha is nearly identical to C. urbana with the exception of the symmetrical, spoon-like postgonites, and the absence of a faint, floating sclerite anterior to female sternite 10. The postgonites in these species are typically large and exposed, so dissections are rarely necessary to distinguish males.

63 Coproica bispinosa Papp, 2008

Distr.: Oriental: Nepal, Thailand. Coproica bispinosa Papp, 2008: 32 [both sexes, phylogenetic notes, illustr.]. Type locality: Nepal, Royal Chitwan NP, Bandarjhola Is.. HT male (HNHM).

64 Coproica brachystyla sp. nov. (Figures: 22, 65-76)

Distr.: Nearctic: Canada (ON; data suspect); USA (FL, SC, TX, UT); Neotropical: Argentina, Barbados, Belize, Bolivia, Brazil, Costa Rica, Dominican Republic, Ecuador, Guatemala, Honduras, Mexico (CAM, CHP, COA, ROO, TAB), Panama, Paraguay, Puerto Rico, St. Kitts, St. Martin, Venezuela.

Body length 1.1 - 1.5mm. Body colour brown. Face light brown, frons brown.

Interfrontal bristles in 3 pairs, postocellar bristles convergent, postocellar setulae absent.

Eye to gena ratio approximately 2:1, 3 genal setae in longitudinal row posterior to vibrissa with anterior seta approximately 1.5 times the length of posterior setae.

Scutellum with 10-12 setae loosely arranged in 2 rows between basal 1/4 and basal 1/2; 2 lateral bristles between larger anterior and posterior marginal scutellar bristles.

Katepisternum with 2 small, subequal dorsal bristles, each approximately 1/5 as long as the distance between dorsal margin of katepisternum and wing base. Fore coxae brown, legs light brown. Mid tibia with a small proximal posterodorsal bristle at 2/5. Mid basitarsus with row of 5-7small, stout anterior bristles extending from 1/3 to apex, strong sub-basal anteroventral bristle, stout anteroventral bristle at 1/2, and small posteroventral bristles at 1/3 and 2/3, additional setae often present on ventral surface, variable. Second costal sector 0.8-1.3 times length of third costal sector. Costal bypass long, extending 7 or more vein widths beyond R4-1-5. Anal vein sinuate, lacking an angulate bend.

Male terminalia:

Sternite 5 unmodified, all surface bristles uniform in size. Synsternite 6+7 straight medially. Epandrium with enlarged pair of dorsolateral bristles; subanal plate absent, cerci and epandrium medially separate below anal opening; cercus fused to epandrium, ventral margin with short medial process, 2 large bristles on cercus. Surstylus with short,

65 blunt posterior process. Postgonites asymmetrical, straight, left postgonite with rounded, spoon-like apex, right postgonite shorter, with shallow apical hook. Basiphallus re­ shaped, open posteriorly, posteroventral margin unmodified. Lateral sclerites rounded, 2 short dorsal processes near apex, elongate, narrow process on distal ventral angle.

Central sclerite fused with lateral sclerites just before apex, distal series of 3 small blunt processes.

Female terminalia:

Tergite 8 reduced to 2 lateral sclerites with medial connecting membrane. Tergite 10 glabrous, broadly fused to cerci; cerci glabrous on anterior half, pruinose on posterior half, apical and subapical setae modified into spines. Sternite 7 rounded rectangular, posterior margin with strong medial invagination. Sternite 8 oval with strong anterior process, and anterior medially divided row of 4 small setae. Sternite 10 wrapped posteriorly around tip of abdomen. Spermathecae spherical, slightly tapered towards duct; duct length 2-3 times length of spermathecal bulb.

Type material:

HOLOTYPE: COSTA RICA: San Jose, San Carlos, Riosparaiso Res., Pecari Stn., 16km NNE Quepos, 9°33'53"N, 84°7'32"W, 400m, 22-27.ii.2006, S.A. Marshall (1$: debu00259735, INBC).

PARATYPES: ARGENTINA: Salta, 30km E of Salta Campo Quijano, 18-20.ii.1992, dung tp. in forest remnant, S.A. Marshall (2c?); Salta Prov., El Ray Nat. Pk., 900m, Rio La Salsa, 5-10.xii.1987, FIT, malaise tp., open stream side in forest, S.&J. Peck (1?); Salta, El Ray Nat. Pk., 1000m, Pozo Verde Tr. km 7, 5-15.xii.1987, malaise-FIT, yungas forest, S.&J. Peck (2& 1$); Salta, Rosario de Lerma, INESALT yard, 16-28.ii.1992, malaise tp., S.A. Marshall (2 <$). BARBADOS: Weichman Hall Gully, 250m, 20- 25.ii.1979, carrion, moist woods, S.&J. Peck (IcT). BELIZE: Mt. Pine Ridge, Hidden Valley inst., 2500', 10-15.U991, dung tps., grass/pine, S.A. Marshall (IcT, 1$); BOLIVIA: La Paz, Chulumani, Apa Apa Reserve, 2000m, 16°21'15 S, 67°30'21 W, l.iv.2001, sweep, S.A. Marshall (1<$: debu00208859, CBFC); Santa Cruz, Campo Guairuy, 24km S of Camiri, 870m, 18.viii-2.ix.2000, malaise tp., Irwin & Hauser (3cT: debu00144466, 00144586, 00144735; 1$: 00144123, UASC); Santa Cruz, Poterillo de

66 Guerda, 17°40.49'S, 63°27.36W, 4-7.iv.1998, H.&A. Howden (1$, UASC); Santa Cruz, Quebrada, isiri, 20km S of Carniri, 865 m, 18.viii-2.ix.2000, malaise, Irwin & Hauser (2$: debu00147621, 00147836, UASC). BRAZIL: Bahia, 15km NE of Porto Seguro (Ecological Reserva "Pau-Brasil"), 1' Atlantic Forest, 19-27.ii.1986, Shannon-tp., Daltoas & Vera Cristina S. Amaorim (2c?, MZSP); Minas Gerais, Institute Estadual De Florstes Parque Florestes Do Ibitipoca, 1700m, 12.ii.1990, swept near fresh dung, S.A. Marshall (2c?, MZSP); Parana, near Londrina, Mata Dos Godoy, 28-3l.i. 1990, malaise (dry head), S.A. Marshall (lc?, MZSP); Mato Grosso, Pocone, Centro (14:25 hs), 16.V.1990, BPF Group col. (1$, 1?, MZSP); Rio de Janeiro, Terespolis Pedra do Sino, ~2000m, 14.iii.1990, human dung, S.A. Marshall (1$, MZSP). BRITISH HONDURAS: Belmopan, 26-30.vii.1972, palm for. carrion, S. Peck (1$). CANADA: ON: Bruce Co., Bruce Penninsula Nat. Pk., Emmett Lk. Rd., 9-12.vii.1996, car net, (1$: debu00105022, data out of normal range). COSTA RICA: Same as holotype (25c?: debu00259713, 00259715, 00259732, 00259750, 00259725, 00259743, 00259711, 00259704, 00259734, 00259640, 00259739, 00259747, 00259722, 00259731, 00259748, 00259745, 00259749, 00259716, 00259708, 00259714, 00259701, 00259755, 00259756, 00259738, 00259702; 33$: 00259683, 00259672, 00259740, 00259724, 00259707, 00259699, 00259698, 00259679, 00259635, 00259754, 00259726, 00259710, 00259727, 00259728, 00259730, 00259746, 00259758, 00259712, 00259678, 00259729, 00259717, 00259733, 00259718, 00259703, 00259720, 00259736, 00259751, 00259721, 00259705, 00259753, 00259757, 00259706, 00259737, INBC); same as holoype except 13-15.iv.2006, pans in slash near springs (3c?: debu00259938, 00259918, 00258890; 2$: 00258534, 00258851, INBC); Florencia For., 28.ii.1980, dung tp.., H. Howden (2$, INBC); Heredia, La Selva Biol. Stn., 3km S of Puerto Viejo, 80m, 10°26'N, 84°1'W, 26.vi-27.vii.2001, FIT, S. Chatzimanolis (1$: debu00228869-wing, INBC); Heredia, La Selva, 9.V.1989, black light tp., B.V. Brown (2$, INBC); Heredia, Puerto Viejo, La Selva Biological Station, 7.V.1989, black light, B.V. Brown (lc?, INBC); Heredia, Rara Avis, El Plastico 1st Forest Mar. house, 580m, 19-20.V.1987, human dung, Norrbom (1$, INBC); Limon, Siquerres, Pacuarito, Las Brisas, Estacion Rio Dantas, 22-25.viii.1996, malaise tp., Gustafsson et al., (1$: debuOOl 17245, INBC); Puntarenas, 5km W of Rincon Punta EL, 15m, 6-iii- 1968, iguana carrion, 0-6-1-124, Iguana, Barney & Cornaby (lc?, INBC); Puntarenas, R.F., Golfo Dulce, 24km W Piedras Blancas, 200m, xii.1990, P. Honson (IcT, 2?, INBC); Puntarenas, Osa Penninsula, Rincon, 2.5 km S ~50m, 8°42'1N, 83°30'50W, ll.viii.2001, 2° forest, dung pans, M. Buck (1$: debu00188278, INBC); San Jose, Zurqui de Moravia, 1600m, v.1981, P. Hanson (lc?: debu00100587, INBC). DOMINICAN REPUBLIC: Monte Cristi, 3km N of Villa Elisa, 1-3.X.1985, dung, R.E. Woodruff (1$); Pedernales, 30km N of Cabo Rojo, 18-07N, 71-39W, 1070m, 23- 24.vii.1990, C. Young, J.E. Rawlins, & S. Thompson (1$); ECUADOR: Napo, Jatun Sacha Res., 6km E of Misahualli, 450m, 1°4'S, 77°37'W, 2-7.V.2002, varzea, dung pans, M. Buck (lc?: debu00196107, QCAZ); Napo, Yasuni Nat. PL, 3-20.xi.1998, forest malaise tp., Pape & Viklund (lc?: debuOOl 16953, QCAZ). GUATEMALA: Peten, Tikal, 28-30.vii.1978, dung tp., Heleva & Kukal (1$). HONDURAS: Lago de Yajoa, 2600', 1- 2.vi.l994, dung tp., Howden? (3c?**-illustr.xl, 4$*-illustr.). MEXICO: Campeche, 53 mi E of Escarcega, 500', 8-14.viii.1971, tropical semi-evergreen., dung, A. Newton (1$); same as previous except 87 mi E of Escarcega, 800', human dung (1$); Chiapas, 5 mi S of Bochil, 5000', 18-24.viii.1971, oak, human dung, A. Newton (lc?); Chiapas, 11 mi E

67 of Trinitaria, 5200', km 18, 5.viii. 1971, tropical eciduous for., human dung, A. Newton (2<$); Chiapas, Lagunas de Montebello Parque Nacional, Aqua Tinta, 4900', 21- 24.viii.1971, oak-pine, human dung, A. Newton (2$); Chiapas, Lagunas de Montebello Parque Nacional, La Eucantada, 4900', 21-24.viii.1971, oak/pine/liquid amber, human dung, A. Newton (39); Chiapas, Ocosingo Rd. 76km S of Palenque, Rt. 195, 760m, 5- 29.vii.1983, rainfor. window tp., S.&J. Peck & R. Anderson (1$); Coahuila Boquillas, 24-26.ix.1979, pig dung tp., R.E. Woodruff & G. Fincher (l<$) Quintana Roo, 160m, Kohunlich, 68km W of Chetumai, 14-17.vii 1982, tropical seasonal for., FIT, S.&J. Peck (2

Etymology:

The name brachystyla means "short stylus" referring to the very short pointed posterior process on the surstylus.

68 Coproica brevivenosa Papp, 2008

Distr.: Afrotropical: Kenya, Nigeria, Tanzania, Zanzibar. Coproica brevivenosa Papp, 2008: 42 [both sexes, phylogenetic notes, illustr.]. Type locality: Nigeria, Yankari Reserve. HT male (HNHM).

69 Coproica cacti (Richards, 1960) (Figure: 77)

Distr.: Nearctic: USA (AZ, CA); Neotropical: Mexico (BCN). Leptocera (Coproica) cacti Richards, 1960c: 203 [both sexes, illustr.]. Type locality: USA, Arizona, Maricopa Co., 2.2 mi NE Morristown. HT male (USNM). - Richards, 1965c: 725 [Nearctic catalog]; Richards, 1967b: 16 [Neotropical catalog]. Coproica cacti. - Papp, 1979a: 104 [generic combination].

Body length 1.1 mm. Body colour brown. Face and frons brown. Interfrontals in 3 strong, erect pairs, 2nd pair stronger than others; postocellar bristles slightly convergent, small postocellar setulae absent. Eye small, eye to gena ratio approximately 1.4:1; gena with longitudinal row of 3 small setae posterior to vibrissa. Scutellum with approximately 12 setae arranged in 2 loose rows between basal 1/4 and basal 1/2; 1 lateral scutellar bristle between larger anterior and posterior marginal scutellar bristles.

Katepisternum with 2 small, subequal dorsal bristles, each approximately 1/8 as long as the distance between dorsal margin of katepisternum and wing base. Legs and fore coxae light brown. Mid tibia with proximal posterodorsal bristle at 2/5. Mid basitarsus with row of 6 anterior bristles extending from 3/4 to apex, 1 posteroventral basal bristle, 2 anterior bristles at 3/5, and 1 posteroventral bristle between 2 anteroventrals. Second costal sector 1.0-1.1 times length of third costal sector. Costal bypass long, extending 7 or more vein widths beyond R4+5. Anal vein sinuate, lacking an angulate bend.

Type Material:

PARATYPES: UNITED STATES OF AMERICA: AZ: Pinal Co., 33mi E of Gila Bend, ll.ii.1958, Reared: ADM 15 ll.iii.1958, R.E. Ryckman & E.T. Ryckman, Paratype No. 65054 USNM (1 (sex unknown)*-wing, USNM)

70 Specimens Examined:

UNITED STATES OF AMERICA: AZ: Tucson, 10.vi.1959, barrel cactus rot, M. Adachi;(l(5**, 1$).

Comments:

Coproica cacti is only known from specimens reared from broken or decaying cacti. Although very similar to C. urbana, the species is easily distinguished from all other species of Coproica by a reduced eye to gena ratio and a thickened central interfrontal bristle.

Male and female genitalia have not been illustrated or included in the redescription of Coproica cacti due to the low availability of specimens. Among these specimens individuals were either already missing abdomens (previously cleared and disintegrated), or considered too valuable to risk dissection.

71 Coproica coreana Papp, 1979 (Figures: 78-88)

Distr.: Oriental: China (HKG), Pakistan, Taiwan, Thailand; Palaearctic: Japan, North Korea, South Korea. Coproica coreana Papp, 1979a: 98 [both sexes, phylogenetic notes, illustr.]. Type locality: North Korea, Prov. South Pyongan, Changlyong san, 50 km N of Pyongyang and 15 km E from from Sa-gam. HT male (HNHM). - Papp, 1984: 81 [Palaearctic catalog]; Hayashi, 1986a: 117, 119 [key, illustr.].

Body length 1.1 - 1.5mm. Body dark brown. Face brown, frons dark brown; head appearing longitudinally flattened, height 1.6-1.8 times length. Interfrontal bristles in 4-5

(usually 5) subequal pairs; postocellar bristles slightly convergent, 1 pair of small postocellar setulae present. Gena brown; eye to gena ratio 2.0:1-2.5:1. 4 genal setae in longitudinal row posterior to vibrissa, anterior seta approximately 1.3 times length of posterior setae, and a single seta present just ventral to row. Scutellum with 10-15 discal setae arranged into 2 loose rows between basal 1/4 and basal 1/2; 2 lateral setae between larger anterior and posterior marginal scutellar bristles. Katepisternum with 2 small subequal dorsal bristles, each approximately 1/7 as long as the distance between dorsal margin of katepisternum and wing base. Fore coxa yellow, legs brown. Mid tibia with additional small anterodorsal bristle at 1/6, proximal posterodorsal bristle at 2/5. Mid basitarsus with small, stout sub-apical bristle, an anterior bristle at 1/2, a strong sub-basal anteroventral bristle , a stout anteroventral bristle at 3/5, and 2-3small, posteroventral bristlesin variable positions. Second costal sector 0.8-0.9 times length of third costal sector. Costal bypass short, terminating within 5 vein widths of R4+5. Anal vein sinuate, lacking an angulate bend.

72 Male terminalia:

Sternite 5 rounded rectangular, posteromedial patch of microsetae, surface bristles enlarged laterally. Synsternite 6+7 straight medially. Epandrium with enlarged pair of anterodorsal bristles and ventrolateral bristles; subanal plate absent, cerci and epandrium discontinuous below anal opening; cercusfused to epandrium, ventral margin relatively flat and straight, with small, bristled medial process. Surstylus with elongate, thickened, toenail-like, dorsomedially curved bristle at posterior apex, cluster of 3-4 elongate posteroventral bristles. Postgonites symmetrical, straight, and narrow. Basiphallus gently curved, tubular, posteroventral margin with elongate rounded crest open posteriorly dorsal to crest. Lateral sclerites rounded, 2 small dorsal processes near apex, apex medially deflected. Central sclerite apically fusing with lateral sclerites.

Female terminalia:

Tergite 8 long laterally, short dorsally. Tergite 10 pollinose, narrowly fused to cerci posterolaterally; cerci rounded, pollinose, apical bristles long and sinuate. Sternite 7 rounded rectangular. Sternite 8 trapezoidal in shape, broadest posteriorly, rounded, 2 enlarged bristles within a row of 4 smaller setae, and posterior row of 4 setae. Sternite 10 not wrapped posteriorly around abdomen, with posteromedial blunt point and pair of enlarged central bristles. Spermathecae spherical; duct length approximately equal to bulb length.

Specimens Examined:

THAILAND: Kanchanaburi, 19.viii.1989, T. Hayashi (6cj?**-illustr.-wing, 6?**- illustr.).

73 Comments:

Coproica coreana is an unplaced species between the C. acutangula species group and the C. vagans and C. urbana species groups. It shares several weak putative synapomorphies with each species group, and these conflicting apomorphies contribute to its placement as the sister species to the C. vagans and C. urbana species groups.

74 Coproica demeteri Papp, 2008

Distr.: Afrotropical: Nigeria, Tanzania. Coproica demeteri Papp, 2008: 8 [both sexes, phylogenetic notes, illustr.]. Type locality: Nigeria, Yangukari Reserve. HT male (HNHM).

75 Coprica dentata Papp, 1973

Distr.: Palaearctic: Mongolia. Coproica dentata Papp, 1973a: 381 [both sexes, phylogenetic notes, illustr.]. Type locality: Mongolia, Archangaj aimak, "Ih-Tamir" [= Ichtamir]. HT male (HNHM). - Papp, 1984: 81 [Palaearctic catalog].

Comments:

Coproica dentata is described as possessing a tooth-like process on the male hind basitarsus, suggesting a close relationship with the clade including C. acutangula and C. digitata. Coproica dentata also seems to share several complex male genitalic characters with C. pappi, including the almost identical shape of the surstylus and an elongate process on the male cercus, and the two species may be synonyms (see below).

Examination of the holotype of C. dentata will be necessary to clarify the possibility of homology between the tooth-like bristle and that in C. digitata and relatives, and to check for the presence of this bristle in both C. dentata and the identical (or almost identical) C. pappi.

The identity of Coproica dentata remains less than clear, even though 418 paratypes were designated in the original description. All the Hungarian specimens were later considered misidentified and were actually a species that had been described by

Carles-Tolra, C. pappi Carles-Tolra, 1990 (Papp, 1999). Illustrations of the male genitalic characters were still later provided for both C. dentata and C. pappi in Papp,

2008 but when we examined specimens identified as C. dentata from Hungary they were found to match the illustrations provided for C. dentata in Papp, 2008, yet lacked the process on the hind basitarsus, which should be diagnostic of C. dentata. Careful comparison of the holotypes of these two Old World species is needed to resolve the issue, and to assess whether or not these two species are actually synonyms.

76 Due to the striking similarity between Coproica dentata and C. pappi and the lack of a process on the hind basitarsus of all male specimens observed, specimens observed have been identified and treated as C. pappi. Therefore, no specimens of C. dentata were

examined.

The structure referred to as the "terminal process of the male hypopygium" in

Papp, 1979 and as the "postgonite" in Papp, 2008 is in fact a process on the male cercus.

77 Coproica diabolica sp. nov. (Figures: 8, 88-100)

Distr.: Nearctic: Canada (ON), USA (AR, FL, GA, IL, MS, NC, OH, OK, TN, TX); Neotropical: Brazil, Costa Rica, Dominica, Dominican Republic, Ecuador, Guyana, Grenada, Jamaica, Mexico (CHP), St. Kitts, St. Lucia, St. Vincent, Trinidad, Venezuela.

Body length 1.1 - 1.3 mm. Body brown. Face and frons dark brown. Interfrontal bristles in 3 subequal pairs, postocellar bristles slightly convergent, postocellar setulae absent. Eye to gena ratio approximately 2.0. Longitudinal row of three genal setae posterior to vibrissa, anterior seta twice the length of posterior 2. Scutal setae stout, thickened. Scutellum with approximately 14 discal setae arranged in 2 rows between basal 1/3 and basal 1/2; 1 lateral seta between larger anterior and posterior marginal scutellar bristles. Katepisternum with 2 small, subequal dorsal bristles, each approximately 1/7 as long as the distance between dorsal margin of katepisternum to wing base. Fore coxa and legs brown to light brown. Mid tibia with proximal posterodorsal bristle at 2/5. Mid basitarsus with a row of 4-5 anterior bristles along extending from 2/3 to apex, 1 ventral basal bristle, 1 posteroventral bristle at 1/2, and anteroventral bristles at 2/5 and 3/5. Second costal sector 0.6-0.1 times length of third costal sector. Costal bypass long, extending 7 or more vein widths beyond R4-1-5. Anal vein sinuate, lacking an angulate bend.

Male terminalia:

Sternite 5 rounded, with short paired processes along posterior margin with lightly sclerotized surfaces laterally, a small posteromedial patch of microsetae, and all surface bristles uniform in size. Synsternite 6+7 with slight medial bend. Epandrium with enlarged pair of dorsolateral bristles; subanal plate absent, cerci and epandrium

78 discontinuous below anal opening; cercus fused to epandrium, ventral margin with short, rounded medial processes, and a small sub-apical pointed process. Surstylus with rounded posteroventral lobe and several bristles along posterior margin. Postgonites asymmetrical, straight, left postgonite broad, with pointed apex, right postgonite narrow on distal half, with hook at apex. Basiphallus straight, with sharp posterodorsal curve, posteroventral margin unmodified, open posteriorly. Lateral sclerites with expanded membranous posterodorsal corner, small, pointed dorsal processes near apex, and apical margin deflected medially with subtle ribbing. Central sclerite fused apically with lateral sclerites. Apical membranes short, broad.

Female terminalia:

Tergite 8 long laterally, short dorsally. Tergite 10 pollinose, narrowly fused to cerci posterolaterally; cerci elongate, rounded, and pollinose, apical bristles long and sinuate.

Sternite 7 broad, rounded, and concave along posterior margin. Sternite 8 with 3 pronged anterior process, notched posteriorly, and a row of 4 small setae anterior and posterior to an additional small pair of setae. Sternite 10 wrapped posteriorly around tip of abdomen, posterior margin with blunt point, pair of enlarged medial setae. Spermathecae slightly conical; duct length 2-3 times length of spermathecal bulb, short sclerotized portion of duct adjacent to spermathecal bulb, with small constriction at its base.

Type Material:

HOLOTYPE: CANADA: ON: Bruce Co., Dorcas Bay Dunes, 2-25.viii.1999, malaise, S.A. Marshall, (ltf*: debu00081266*).

PARATYPES: BRAZIL: Bahia, 15km NE of Porto Seguro (Ecological Reserve "Pau- Brasil"), 1' Atlantic Forest, 19-27.ii.1986, Shannon-tp., D.&V. Cristina & S.Amaorim (13$, 11$, MZSP); Parana, Curitiba, ponds near Zoo, 7.ii.l990, sweep, S.A. Marshall (16\ MZSP); Rio Jacara, Pagua, 22.U990, wet litter by stream, S.A. Marshall (1

79 open wet area, S.A. Marshall (1

80 M. Buck (1$: debuOO 195249, QCAZ); Pichincha, 47km S of Santo Domingo, Rio Palenque Res. Stn., l.v.1987, light, B. Brown (1$, QCAZ); GUYANA: Rupununi Dist., 200\ Kurupukari, E side Essequibo, 7-11.X.1990, riv. savannah scrub/1* rainfor. edge, malaise, ROM 905049, B.Hubley, L.Coote (1$, ROME); GRENADA: Grand Etang, 6.U995, sweep wet trail, S.A. Marshall (1(J). JAMAICA: 4000', Hardwar Gap, 10.vii.1966, Howden & Becker (1$). MEXICO: Chiapas, Palenque, 28.vii.1983, rainforest, S. & J. Peck (1$). ST. KITTS & NEVIS: Nevis Dasents Estate, 220m, 28.viii-21.ix. 1985, malaise, citrus orchard, L. Coote (IcT). ST. LUCIA: Anse La Raye, Anse Galet, 1 km SSW Anse La Raye, 13-56N, 61-03W, 50m, 21-30.vi.1991, J.E. Rawlins & S.A. Thompson (IcT*, CMNH). ST. VINCENT: Kingstown, St. Andrews Botanical Garden, 1-3.X.1991, M.T.&R.E. Woodruff (1$*). TRINIDAD: St. George Co., Arima Ward, NY Zool. Soc. Sta., ll.vi.1977, pig dung tp., R. Woodruff; (1$, UWIC). UNITED STATES OF AMERICA: AR: Garland Co., 1.2mi N of Crystal Springs, Hwy 270, 6-8.iii.1977, pig dung tp., Woodruff & Wiley (1 ?); Montgomery Co., 20mi W of Glenwood, T45 R27W, 15.vi.-28.vii.1994, H.W. Robinson (1$); Scott Co., 7mi E Y City, Jet. of Hwy270 & Rt71 on 270, 6-8.iii.1977, pig dung tp., Woodruff & Wiley (1 $). FL: Archbold Bio. Sta., 12-16.xii.1985, dung pan, scrub-sand, S.A. Marshall (1?); Archbold Bio. Stn., 13-18.xii.1985. rotten fungi, S. Marshall; 2 $s, 1 $ Archbold Bio. Sta., 14.xii.1985, vac. under carrion, S.A. Marshall (1$, lc?); Archbold Res. Stn., 15-16.iv.1989, dung vac, K.N. Barber (1$*); Archbold Bio. Stn., 16.xii.1985, carrion residue, S.A. Marshall (2$, 1$); Archbold Bio. Stn., 17.iv.1989, dung vac, S.A. Marshall (1$); Archbold Bio. Sta., 17-2l.xii. 1985, scrub dung, S.A. Marshall (1$); Archbold Bio. Sta., 18.xii.1985, on old carrion site, S.A. Marshall (2<51*-illustr, 2$*- illustr); Archbold Bio. Sta., along Lk. Placid, 19.xii.1985, dec. veg., S.A. Marshall, (2$, lc?); Archbold Bio. Stn., Lk. Placid, xi.1979, D. Hardwick, (1$); Archbold Bio. Sta. Lk. Annie, 13-18.xii.1985, yellow pan near shoreline, S.A. Marshall (lef); Archbold Bio. Sta., Lk. Placid, 6.xi.l985, M. Deyrup (1$); Collier Co., Collier-Seminole St. Pk., iv.1971, pine, palm, human dung, A. Newton (1

81 holotype except malaise (2^*: debu00227332, 00227397*; 4$: 00227380, 00227348, 00227333, 00227395). OH: Morgan Co., Wildcat Hollow, Wayne Nat. For., 14.vii.1989, wet grass clippings, S.A. Marshall (1$). OK: Latimer Co., 5mi W of Red Oak, 5- ll.vi.1977, dung tp., K. Stephen (1$*, 1?). TN: Sevier Co., Great Smokey Mts. Nat. Pk., W trib. Of Porter's Creek 0.8km SW of access gate, 18.vii.1999, J. Cooper (1$: debu00167109). TX: Angelina Co., Angelina Nat. For., Boykin Cemetery, 13- 17.vi.1993, spring fen, dung, S.A. Marshall (4c?*: debu00128250, 00128268, 00128306, 00128351*, wing; 1$: 00128267). Angelina St. For., Boykin Cemetery, 13-17.vi.1993, spring fen on dung, S.A. Marshall; San Jacinto Co., Coldspring, Double Lake cpgd, 22.v-16.viii.1983, FIT, 83-14, S.&J. Peck (ltf). VENEZUELA: Amazonas, Rio Mavaca Camp, 65°06'W, 2°2'N, 150m, 16-27.iii.1989, Phipps-FUDECI Expedition by Amer. Mus. Nat. Hist. D.A. Grimaldi, coll. (2cT, AMNH). Maracay, 20km N of Rancho Grande Bio Stn., La Trilla, 100m, 26.ii.1996, riv. sweep, S.A. Marshall (1 $, IZAV).

Etymology:

The name diabolica means "devil" and refers to the shape of the female sternite 8, which looks like it has horns anteriorly.

Comments:

Coproica diabolica is widely distributed, though uncommon, throughout eastern

North America, Central America and northern South America.

82 Coproica digitata (Duda, 1918) (Figures: 101-111)

Distr.: Oriental: Pakistan; Palaearctic: Afghanistan, Bulgaria, Canary Is. (Spain), Czech Republic, Egypt, France, Hungary, Israel, Kirghizstan, Macedonia, Pakistan, Romania, Slovakia, Slovenia, Spain (incl. Balearic Is.), Tunisia, Turkestan, Uzbekistan, Yugoslavia. Limosina (Coprophila) digitata Duda, 1918: 209 [male, illustr.]. Type locality: Yugoslavia, Novi. LT male, designated by Papp, 1979a: 100 (HNHM). - Duda, 1938: 163-164 [male, redescription, illustr.]. Leptocera (Coprophila) digitata. - Duda, 1925: 205 [generic combination]. Leptocera digitata. - Hafez, 1939: 327-331 [preimaginal stages, life history, illustr.]. Leptocera (Coproica) digitata. - Richards, 1960c: 206 [subgeneric combination, diagnosis, both sexes, key]. Coproica digitata. - Papp, 1973c: 27 [generic combination, key, illustr.]; Papp, 1984: 81 [Palaearctic catalog]; Rohacek, Baez & Buck, 2003: 56 [new record from Canary Is.].

Body length 1.5 - 2.0 mm. Body dark brown. Face yellow, frons dark brown.

Interfrontal bristles in 5 subequal pairsrfrontal bristles, postocellar bristles erect, 1 pair of

small postocellar setulae. Gena bright yellow, eye to gena ratio approximately 2.5:1;,

Gena with row of five small setae. Scutellum with approximately 40 discal setae between

base and basal 4/5; 4 lateral setae between larger anterior and posterior marginal scutellar

bristles. Katepisternum with 2 small, subequal bristles, each approximately 1/4 as long as

the distance between dorsal margin of katepisternum and wing base. Fore coxa yellow,

legs dark brown. Mid tibia with additional small anterodorsal bristle at 1/9, proximal

posterodorsal bristles at 1/5 and 1/2. Mid basitarsus with stout anterior bristles at 1/2 and

sub-apical, a small anteroventral sub-basal bristle, a stout anteroventral bristle at 1/2, and

3-4 short posteroventral bristles. Second costal sector 1.2-1.5 times length of third costal

sector. Costal bypass short, terminating within 5 vein widths of R4+5. Anal vein with

distinct angulate bend.

83 Male terminalia:

Steraite 5 crescent-shaped, with small paired posteromedial tabs, posteromedial patch of microsetae, posterior row of enlarged bristles, and bristles enlarged laterally. Synsternite

6+7 with strong medial bend. Epandrium with enlarged pair of dorsolateral bristles, ventrolateral row of 4 or more conspicuously large bristles; subanal plate absent, cerci and epandrium discontinuous below anal opening though tightly associated; cercus fused to epandrium, with 1 enlarged bristle, ventral margin with short, rounded medial process.

Surstylus with constriction at base of anterior lobe, with broad ventral process and bifurcate posterior process; ventral process with anterolateral row of elongate bristles, dense row of posteroventral setae, and pair of thickened, elongate, toenail-like bristles at apex, apical bristles curved medially; posterior process bifurcate, each fork with a thickened toenail-like bristle at apex, dorsal fork with a central, blunt bristle, deflected medially. Postgonites asymmetrical, each with a small pointed posterior process at 1/2 and a row of 3 small setae distal to posterior process, left postgonite rounded and spoon­ like at apex, right postgonite slightly bifurcate at apex. Basiphallus L-shaped, tubular, with strong posteroventral crest, ridged posteriorly, blade-like ventrally. Lateral sclerites rounded, with 2 dorsal processes, anterior process sharp and deflected medially, distal process blunt; central sclerite apically fused with lateral sclerites; additional sclerite ventral to central sclerite, fused ventrally with central sclerite apically and at base; ventral surface of distiphallus bulbous. Apical membranes heavily covered with small sclerotized spines.

84 Female terminalia:

Tergite 8 long laterally, short dorsally. Tergite 10 pollinose, narrowly fused to cerci posterolateral^; cerci rounded, pollinose, apical bristles long and sinuate. Sternite 7 broad, rounded. Sternite 8 rounded, trapezoidal, broadest anteriorly, notch along anterior margin, 2 small lateral setae, and anterior row of 4 small setae. Sternite 10 not wrapped posteriorly around tip of abdomen, lateral arms curved dorsally, and medial pair of enlarged bristles. Spermathecae spherical; duct length approximately equal to length of spermathecal bulb.

Specimens Examined:

HUNGARY: Tata erdo Oreg-to, faeces-csapda, 22.viii.1959, leg., Mihalyi (l<$*, 1$). PAKISTAN: Nathiagali, NWFP, 27-30.vii.1988, T. Hayashi; (6c?***-illustnc2-wing, 6$***-illustr).

85 Coproica emarginata sp. nov. (Figures: 112-123)

Distr.: Nearctic: USA (FL, MI, MA, TX); Neotropical: Belize, Brazil, Costa Rica, Dominican Republic, Grenada, Mexico (ROO), Puerto Rico, St. Kitts, St. Vincent, Trinidad, Venzuela.

Body length 1.0 - 1.8 mm. Dark brown to brown. Face and frons dark brown.

Interfrontals in 4 subequal pairs, postocellar bristles slightly convergent, 0-1 pairs of small postocellar setulae. Gena narrow, eye to gena ratio approximately 2.5:1; 2-4 genal setae in longitudinal row posterior to vibrissa, anterior seta twice the length of posterior setae. Scutellum with 10-12 discal setae arranged into 2 loose rows between basal 1/3 and basal 1/2; 2 lateral setae between larger anterior and posterior marginal scutellar bristles. Katepisternum with 2 small, subequal dorsal bristles, each approximately 1/8 as long as the distance between dorsal margin of katepisternum and wing base. Fore coxa brown to light brown, legs light brown, apices of tibiae and tarsi yellow. Mid tibia with additional small proximal posterodorsal bristle between 1/2 and 3/5. Mid basitarsus with row of small anterior bristles setae extending from 1/3 to apex, row of small anteroventral setae extending from base to apex, and row of posteroventral setae extending from 1/3 to apex. Second costal sector 0.6-1.0 times length of third costal sector. Costal bypass short, terminating within 5 vein widths of R4+5. Anal vein sinuate, lacking an angulate bend.

Male terminalia:

Sternite 5 rounded rectangular, posteromedial patch of microsetae bristles slightly enlarged laterally. Synsternite 6+7 straight medially. Epandrium with enlarged pair of dorsolateral bristles; subanal plate absent, light membrane in its place, cerci and

86 epandrium discontinuous below anal opening; cercus fused to epandrium, ventral margin with rounded medial process. Surstyli asymmetrical, with dense short setae along posterior margin, left surstylus with small posteroventral cleft, right surstylus with large concavity. Postgonites asymmetrical, broad, and straight, left postgonite slightly sinuate and pointed at apex, right postgonite narrow along distal 1/5 and pointed at apex.

Basiphallus straight, with abrupt posterior angle, tubular, posteroventral margin unmodified, open posteriorly. Lateral sclerites with sharp, elongate process at posterodorsal angle, row of 3 additional dorsal processes, proximal process sharp and

deflected medially, central process broad and blunt, distal process narrow, sharp and

deflected medially, distal ventral angle rounded and protruding; central sclerite fused with lateral sclerites ventral to central process.

Female terminalia:

Tergite 8 long laterally and short dorsally. Tergite 10 pollinose, narrowly fused to cerci posterolaterally; cerci rounded triangular, pollinose, cereal bristles long and sinuate.

Sternite 7 broad, rectangular, with 2 faint membranous lobes along posterior margin.

Sternite 8 rounded, anterior margin with a pointed process, rows of 4 small bristles

anterior and posterior to pair of enlarged medial bristles. Sternite 10 not wrapped posteriorly around tip of abdomen, with a pair of enlarged medial bristles. Spermathecae

elongate; duct approximately 2 times the length of spermathecal bulb.

Type Material:

HOLOTYPE: COSTA RICA: Alajuela, Volcan Tenorio, Bijagua Biol. Sta., 700m, 20.vi.2000, cut wet field, pan tps., M. Buck (1$: debuOO 132680, INBC).

PARATYPES: BELIZE: San Ignacio, Maya Mtn. Lodge, 7.1.1991, malaise, S.A. Marshall (3$); tr. to Xunanrunich, 18.U991, S.A. Marshall (1

87 Mato Grosso, Varea Grande, Mapim, 15.iii.1990, (5:30 PM) BPF Group col. (1& 1$, MZSP). COSTA RICA: Same as holotype (256": debuOO 132821, 00132820, 00132853, 00132838, 00132861, 00132873, 00132734, 00132686, 00132659, 00132649, 00132609, 00132674, 00125883, 00132652, 00125884, 00132665, 00132864, 00132697, 00132715, 00132822, 00132735, 00125871, 00132608, 00125893, 00132826; 26$: debu00132716, 00132670, 00132668, 00125869, 00125894, 00132720, 00132626, 00132695, 00132723, 00132865, 00132862, 00125887, 00132690, 00132634, 00132660, 00132823, 00132808, 00132859, 00132848, 00132842, 00132863, 00132824, 00132692, 00132664, 00132630, 00132623, INBC); Alajuela, La Virgen del Socorro, near Carriblanca ca., 700m, 16.ii.1989, Grimaldi & DeVries (1<3\ AMNH) Alajuela, Volcan Tenorio, Bijagua Biol Sta., 700m, 16.vi.2000, M. Buck (84c?*: debu00130727, 00130565, 00130564, 00130562, 00130559, 00130555, 00130552, 00130548, 00130546, 00130544, 00130543, 00130609, 00130652, 00130728, 00130728, 00130761, 00130765, 00130776, 00130778, 00130664, 00130680, 00130902, 00130796, 00130683, 00130803, 00130684, 00130721, 00130720, 00130901, 00130896, 00130980, 00130970, 00130967, 00130802, 00130797, 00130982, 00130964, 00130936, 00125951, 00126040, 00126082, 00126091, 00126093, 00125952, 00125958, 00125974, 00125983, 00126105, 00126110, 00126114, 00126117, 00126121, 00126122, 00126123, 00126127, 00125926, 00125929, 00125950, 00125932, 00125934, 00130931, 00130974, 00126080, 00126079, 00130333, 00130327, 00130352, 00130414, 00130416, 00130420, 00130428, 00130462, 00130479, 00130480, 00130482, 00130503, 00130506, 00130515, 00130516, 00130524, 00130525, 00130533, 00130538, 00130540, 00130541, 00130920*-illustr; 33?*: 00125925, 00130829, 00125928, 00130801, 00130962, 00126135, 00126116, 00126056, 00126054, 00126011, 00125990, 00125975, 00125933, 00130726, 00130751, 00130912, 00130795, 00130910, 00130698, 00130677, 00130636, 00130605, 00130598, 00130553, 00130361, 00130311, 00130413, 00130437, 00130447, 00130448, 00130501, 00130308, 00130534, 00130450*-illustr, INBC); Alajuela, Volcan Tenorio, N slope near Bijagua Biol. Sta., 700m, 7.vi.2000, rainfor., on tree fungus, M. Buck (lc?: debu00126281, INBC); Estacion Santa Rosa, Guanacaste, 300m, 16.ii.1996, S.A. Marshall, Borde del Rio. L_N_313000_359800 #6920 (1$, INBC); Estrella Valley, Pandora, 19.ii.1984, malaise tp., A.&A. Howden (4$, INBC); Guanacaste, Guanacaste Cons. Area, Pitilla Field. Stn., 29.U996, malaise, J. Noyes (1$, INBC); Guanacaste, Guanacaste Nat. Pk., Ptilla Biol. Stn., 800m, ll.ii.1995, by stream, yellow pans, L. Masner (1$: debu00206088, INBC); Heredia, 16km SSE of La Virgen, 1050-1150m, 10°16'N, 84°5'W, 21.ii-9.v.2001, transect (1$: debu00230616, INBC); Heredia, Santo Domingo, INBio Pk., 5-6.X.1999, leaf litter, M. Buck, (8c?: debu00103732, 00103752, 00103584, 00103578, 00103582, 00103594, 00103580, 00103747; 8$: debu00103579, 00103593, 00103572, 00103583, 00103597, 00103577, 00103755, 00103745, INBC); Heredia , Santo Domingo, INBio Pk., 9°58'23N, 84°5'30W, 21.viii.2001, grass clippings, M. Buck (11$: debu00199778, 00199774, 00199781, 00199729, 00199766, 00199738, 00199767, 00199753, 00199713, 00199734; 23$: 00199770, 00199765, 00199757, 00199772, 00199777, 00199731, 00199763, 00199746, 00199739, 00199762, 00199718, 00199769, 00199758, 00199760, 00199755, 00199779, 00199740, 00199747, 00199754, 00199744, 00199749, 00199728, 00199748, INBC); Heredia, Sto. Domingo, INBio Pk., 9°58'23N, 84°5'30W, 18- 19.viii.2001, on compost, Buck & Barber (2c?: debu00187508, 00187495; 5$: 00187504, 00187499, 00187443, 00187460, 00187473, INBC); Pt. Viejo La Selva Verde, 3.iii.l991,

88 FIT & dung tps., S.Peck (1$, INBC); Puntarenas, Penas Blancas, E of Monteverde Res., 10°18'N, 84°49'W, 900m, 25.V.1987, on rotten banana pits, A. Norrbom (1$, INBC); San Jose, San Carlos, Riosparaiso Res., Pecari Stn., 16 km NNE of Quepos, 13- 15.iv.2006, S.A. Marshall (8

89 Borneo, ROM 870031 (5<$, 2$, ROME); same as previous except 18-21.x. 1987, ROM 870030 (ROME). VENEZUELA: Aragua, El Limon Poso del Diablo, 570m, 13.iv.1994, L.Masner, 94-13 (4

Etymology:

The specific name is based on the latin word emarginatus which means notched at the apex, and refers to the notched apex of the male surstylus.

Comments:

Males have asymmetrical surstyli, an unusual character in the genus. Females of

C. emarginata are difficult or impossible to distinguish morphologically from C. hirtuloidea in Central America, where their ranges overlap.

90 Coproica ferruginata (Stenhammer, 1855) (Figures: 9, 18, 124-133)

Distr.: Afrotropical: Congo, Ethiopia, Ghana, Morocco, Nigeria, Seychelles, South Africa, Tanzania, Togo, Zaire; Australasian/ Oceanian: Australia (ACT, NSW, NT, QLD), Fiji, Guam (USA), Hawaii (USA), Japan, New Zealand, Niue, Samoa, Tonga; Nearctic: Canada (AB, BC, MB, NB, NS, ON, QC, SK), USA (AR, CA, DC, FL, GA, IA, ID, IL, IN, KS, KY, MA, MD, MI, MN, MT, NC, ND, NE, NH, NM, NY, OH, PA, SC, SD, TN, WA, WI, WV); Neotropical: Argentina, Bahamas, Bermuda, Bolivia, Brazil, Chile, Colombia, Costa Rica, Jamaica, Honduras, Mexico (CHI, NAY), St. Kitts, Surinam; Oriental: China (HKG, XZ), India (Uttaranchal), Indonesia (Flores, Sumbawa), Nepal, Pakistan, Sri Lanka, Taiwan; Palaearctic: Afghanistan, Algeria, Andorra, Austria, Azores (Portugal), Belgium, Bulgaria, Canary Is. (Spain), China (BEI, TIB), Cyprus, Czech Republic, Denmark, Egypt, Finland, France (incl. Corsica), Germany, Great Britain, Greece (incl. Crete), Hungary, Italy, Israel, Japan, Latvia, Madeira (Portugal), Malta, Mongolia, Netherlands, North Korea, Norway, Poland, Roumania, Russia (CET, NET), Slovakia, Spain (incl. Balearic Is.), Sweden, Switzerland, Tadjikistan, Tunisia, UAE, Yugoslavia (Serbia). Limosina ferruginata Stenhammar, 1855: 397 [both sexes]. Type localities: Sweden, "in Ostrogothia ad Haradshammar, Grebo et urbem Lincopiam haud parce", "in Scania ad urgem Lund et prope Kullen". LT female, designated by Gapasin & Kim, 1972: 1248 (UZIU) [no specimen of type series is labelled and, therefore, no definite type locality can be selected on the basis of lectotype]. Limosina (Coprophila) ferruginata. - Duda, 1918: 215-219 [subgeneric combination, redescription, illustr.]; Duda, 1938: 164 [redescription, illustr.]. Leptocera (Coprophila) ferruginata. - Duda, 1925: 205 [generic combination]. Leptocera (Heteroptera) ferruginata. - Richards, 1930: 307 [subgeneric combination]. Coprophila ferruginata. - Vanschuytbroeck, 1951b: 19 [generic combination]. Limosina (Coproica) ferruginata. - Frey, 1936: 109 [subgeneric combination]. Leptocera (Coproica) ferruginata. - Hardy, 1952: 462 [subgeneric combination]; Richards, 1960c: 295 [diagnosis, key]; Richards, 1965c: 725 [Nearctic catalog]; Richards, 1967b: 16 [Neotropical catalog]; Tenorio, 1968: 196, 200 [diagnosis, male genitalia, illustr.]; Richards, 1980: 618 [Afrotropical catalog]. Coproica ferruginata. - Frey, 1945: 80 [generic combination]; Papp, 1973c: 26 [key, illustr.]; Hackman, 1977: 400 [Oriental catalog]; Papp, 1984: 81 [Palaearctic catalog]; Marshall, 1989b: 603 [Australasian/Oceanian catalog]; Lachmann, 1996: 759-768 [morphology, male & female genitalia, copulation, illustr.]; Gatt, 2008: 698 [new record from UAE]. Leptocera ferruginata. - Gapasin & Kim, 1972: 1248-1250 [redescription, illustr.]. Borborus illotus Williston, 1896a: 434 [female, illustr.]. Type locality: West Indies, St. Vincent I. ST(s) female(s) (BMNH ?). - Collin, 1912: 101 [synonymy]. Limosina illota. - Coquillett, 1900b: 269 [generic combination]. Leptocera (Leptocera) illota. - Spuler, 1924b: 115 [generic combination]. Leptocera (Coprophila) illota. - Duda, 1925: 208 [subgeneric combination].

91 Body length 1.5 - 2.5 mm. Body colour orange and brown. Face and gena orange, frons dark brown. Interfrontals in 4-6 subequal pairs, postocellar bristles convergent, 1 pair of

small postoceallar setulae. Eye to gena ratio approximately 2:1; 5 genal setae arranged

into a loose longitudinal row, anterior most seta twice length of posterior setae. Scutum

and scutellum orange, brown streak down centre of scutum; Scutum with additional small pair of dorsocentral bristles just anterior to prescutellar pair. Scutellum with 40-50 discal

scutellar setae, posterior row of discal setae approximately 2 times the length of other

discal setae; 4-5 lateral setae between larger anterior and posterior marginal scutellar bristles. Anterior half of anepimeron and posterior half of katepisternum light orange, rest of thoracic pleurites dark brown; Katepisternum with 3 long dorsal katepisternal bristles, each approximately 2/3 to 3/4 as long as distance between dorsal margin of

katepisternum and wing base, posterior katepisternal bristle slightly longer than anterior

2. Fore coxae light brown or yellow, fore tibia and fore tarsi dark brown, rest of legs

brownish orange. Mid Tibia with additional pair of small bristles at 1/10, an additional

small anterodorsal bristle at 2/5, and large proximal posterodorsal bristles present at 1/5

and 1/2, each equal in length to anterodorsal bristles. Mid basitarsus with row of 5-6

stout anterior bristles extending from 3/5 to apex, a small, stout sub-basal anteroventral

bristle, an anteroventral bristle just distal to sub-basal bristle and approximately 3 times

its length, a posteroventral sub-basal bristle, and 3 anteroventral and 2 posteroventral

small bristles on distal half. Second costal sector 1.6-2.2 times length of third costal

sector. Costal bypass short, terminating within 5 vein widths of 1*4+5. Anal vein with

distinct angulate bend.

92 Male terminalia:

Sternite 5 broad, rounded rectangular, with posteromedial patch of microsetae.

Synsternite 6+7 straight or slightly curved. Epandrium with dorsolateral and ventrolateral pairs of large bristles; subanal plate very narrow, cercus or epandrium barely connected below anal opening; cercus fused to epandrium, ventral margin relatively flat and straight, with posteromedial cluster of 3-4 bristles. Surstylus simple, elongate, trapezoidal, and narrow distally, with posterior cluster of 10-15 bristles and small bristles scattered over lateral and medial surfaces. Postgonites symmetrical, straight, and with subapical lip. Basiphallus with L-shaped, tubular, posteroventral margin with acute crest, open posteriorly dorsal to crest. Lateral sclerites with 2 dorsal processes, proximal process sharp and deflected anteromedially, distal process rounded and deflected posteriorly, apex acute, rounded, and slightly spinose; central sclerite with pair of small apical processes, fused to lateral sclerites between dorsal processes. Apical membranes short.

Female terminalia:

Tergite 8 short dorsally. Tergite 10 glabrous, orange, and narrowly fused to cerci posterolaterally; cerci glabrous, orange, short setae scattered on posterior half, and tuft of short bristles at apex. Sternite 7 broad, rounded rectangular, slightly concave anteriorly,

4 elongate posterior bristles. Sternite 8 trapezoidal, width approximately 2/3 width of sternite 7, posterior row of 4 elongate bristles. Sternite 10 with central pair of enlarged bristles. Spermathecae elongate, rounded; duct length approximately 2/3 length of bulb.

Specimens examined:

ARGENTINA: Santa Fe, 26 km S of Reconquista, Rt. 11, 2km NE of Berna, 20- 21.ii.1980, pig dung tps., R.E. Woodruff (l<$). BRAZIL: Rio de Janeiro, Teresopolis,

93 Pedra do Sino, ~2000m, 14.iii.1990, human dung, S.A. Marshall (1$). CANADA: AB: Jumping Pound Creek, lO.viii. 1980, S.A. Marshall, (ltf); Wagner Bog, 10-16.vii.1985, pan tp. 13, Leg, A.T. Finnamore (lc?*, PMAE); same as previous except pan tp. 2 (1$, PMAE); 28.viii.-10.ix.1985, pan tp. 1 (1<3\ PMAE); 28.viii.-10.ix.1985, pan tp. 2 (1$, PMAE); 28.viii.-10.ix.1985, pan tp. 3 (1$, PMAE). MB: 17 km N of Woodbridge on Hwy 210, Sandilands Prov. For., 5.vii.l987, R. Roughley (2c?). NB: St. Andrews, 11- 13.vi.1978, old bedroom, SA Marshall (1$); St. Andrews, 15.vii.1978, S.A. Marshall (I<$); St. Andrews, Gibson Lake, 1 l.viii.1978, pitfall tps., S.A. Marshall (2c?). NS: Cape Breton Highland Nat. Pk., North Mt., 400m, 10.viii.1983, PG766864 fen pan tp., J. Martin (1$). ON: 3km S of Canfield, 7-13.ix.1986, J. Troubridge (ltf); 12.4km NNE of Searchmont, mi 10 Whitman Dam Rd., 7.vii.l986, dead Marmota, K.N. Barber (1$, 2$); same as previous except human dung {\S)\ Alfred, lO.v.1981, bog, moose dung, Peck & Anderson (1 $); Algoma Distr., Hilton Beach, 10.vii-20.viii.1989, cedar swamp, pig dung pans, J.E. Swann (1$: debuOO 184634); same as previous except red oak forest, pig dung pans (2cT: debu00184353, 00184803; 1$: debuOO 184297); Algonquin Prov. Pk., Swan Lk. Stn., Scott Lk. Survey, 15-3l.viii. 1994, A5 Sphagnum (IcT); same as previous except C5 Shore Pan tp. (ltf); 23-30.vi.1994, A5 shore pans (3c?-wing, 3$); 23-30.vi.1994, A3 Sphagnum malaise tp., (1$); 23-30.vi.1994, C3 Sphagnum (1$); 28.vii.1995, A3 Transition, S.A. Marshall (1$); 28.vii.1995, D2 Hemlock pan tps., S.A. Marshall & D.C. Caloren (2$); Algonquin Prov. PL, Swan Lk. Stn., Scott Lk. Survey, 45°29'15"N, 78°43'20"W, 2-12.vii.1993, malaise C9, hardwood, Larson, Marshall, & Bar (1$); Algonquin Prov. PL, Arowhon Rd., Simm's Pit, 45°34'19"N, 78°42'41"W, 15- 16.viii.2002, yellow pans, M. Buck (ltf: debu00225846); Arkell, 20.vi.1956, D.H. Pengelly (1

94 Res., 28.V.1992, R.W. Burgess (2c?); Guelph, Arboretum, 12-14.vii.1984, field, mushroom tps., B.V. Brown (13)', Guelph, Arboretum, 26.iv.-2.v.l984, mushroom tps., deciduous, B.V. Brown (1$); Guelph, Hanlon Conservation Area, 25.vi.1984, mushroom bait tp., K. Gilbert (13); Guelph, UofG Arboretum (south), 01-10.viii.1993, malaise tp. at edge of field/mixed for., D.C. Caloren (1$); Guelph. UofG Arboretum, 21-27.viii.1984, dead insect baited tps., B.V. Brown (13)', Haldimand-Norfolk Reg., Delhi-Simcoe Railway, site 2, ll-14.vii.2001, yellow pans, S. Paiero (ltf: debuOl 105587); Hald- Norfolk Reg., Manester Tract, 6 km NNW of St. Williams, 8.vi.2001, for., dung pans, M. Buck (6c?: debu00158239, 00158116, 00158205, 00158210, 00158217, 00158170; 6$: 00158131, 00158198, 00158165, 00158153*, 00158178*-illustr, 00158138*); Hald- Norfolk Reg., Manester Tract, 6 km NNW of St. Williams, 15.vi.2001, forest, decayed shrimps pans, Buck, Paiero & McKendry (1$*: debuOOl58568*); Halton Reg., Milton, Deny Rd. & 4th Line, 12-16.vii.2001, grass field, yellow pans, S. Paiero (1$: debu00172154); same as previous except 23-24.vi.2001 (1$: debuOO 174970); Hamilton- Wentworth Reg., Flamborough, 5th Cone, Lawson Farm, 43 18'58"N, 80 2'26"W, 25- 26.ix.2002, M. Buck (1$*: debu00224756*-illustr); Hamilton Royal Botanical Gardens, 15.vi.1998, RET- carrion, V. Carney & K. Scott (23, 29); same as previous except dated Ol.v.1998 (1$); Hamilton, l.vii.1980; malaise tp., M. Sanborne (1$); Heckston, 20 km SE of Kemptville, 24.vi-21.vii.1984, FIT, M. Kaulbars (1$); Hilton Beach, 6.v.- 10.vi.1989, sugar maple forest pan tps.., pig dung, J.E. Swann (13)', Hilton Beach yB, 6.v.-10.vi.l989, sugar maple for. pan tps., pig dung, J.E. Swann (1$); Hilton Township, Tenby Bay, 27.vi.1992, malaise at edge of lakeshore & poplar for., Leg, J.E. Swann (\3)', Iron Bridge, 5.viii. 1981, bear dung, S. Marshall (3$); Kent Co., Rondeau Prov. Pk., South Point Tr. near east parking lot, 42°15'42"N, 81°50'49"W, 17.vi-3.vii.2003, savanna, malaise, S.A. Marshall (1$: debuOl 137071); Kent Co., Rondeau Prov. Pk., Visitor Centre, 42°46'50N, 81°50'38W, 16-17.vi.2003, MV light, M. Buck (1$: debuOl 125190); Lambton Co., Port Franks, Watson Property near Ik., 31.vii.- 06.viii. 1996, pans, J. Skevington (\3, 1 $); same as previous except dated 25-3l.vii. 1996 (1$*); Lambton Co., Port Franks, Karner Blue Sanctuary, 19-26.viii.1996, pans, J. Skevington (13)', same as previous except dated 10-15.vii.1996 (1<3**); 25-30.vii.1996 (13)', Marmora, 27.vi.1957, K.D. Southern (1$); same as previous except dated 18.vi.1957 (1$); Mattawa, Champlain Prov. Pk., 2-30.ix.1979, mixed for. carrion, S.&J. Peck (1?*); Meaford, 20.V.1981, compost box, S.A. Marshall (1$); Mer Bleue, 21.v- 3.vi.l982, L. Dumouchel (1$); Mer Bleue, 23-28.vi.1982, intercept tp., L. Dumouchel & G. Bisdee (13)', same data as previous except dated 14-23.vi.1982 (33)', Muskoka Distr., Muldrew Lk., near Gravenhurst, 29.v-4.vi.2006, O. Lonsdale (23: debu00263569, 00263601); Oliver Bog, 3km S of Gait 25.vi-16.vii.1988 pt4 - forest V4C D. Blades; (23)', same as previous except 26.ix-25.xi.1988, ptl, sedge pond, V1G (1$); Ottawa, vii.1990, bus shelter, J.R. Vockeroth; (13*, CNCI); Ottawa, x.1990, J.R. Vockeroth (13); Ottawa, 6.ix.l984, swept from liquid manure pit, J.R. Vockeroth (43, 4$); Owell, 14.vi.1978 D. Morris; (1$); Point Pelee, 30.vi.1978, W.A. Attwater (2$); Point Pelee, 22.vii.1979, K.N. Barber (1$); Point Pelee Nat. Pk., forested area by west beach, 10- 21.viii.1999, malaise/pan tps., O. Lonsdale (1$: debuOOO 11244); Point Pelee Nat. Pk., Visitor's Centre, 22-29.V.2000, malaise & pan tps., O. Lonsdale (1$: debuOl001986); Puslinch, l.vii.1983, malaise, Coote & Marshall (1$, \3); Puslinch, 20.viii.1982, G.M. Grant (1$); Puslinch, 30.vi-2.vii.1984, maple swamp malaise tp., Coote & Marshall

95 (lcJ); Sarnia, 8.vii.l976, J. Heraty (1$); Shirleys Bay, 15km W of Ottawa, 15- 30.vi.1984, FIT, M. Kaulbars (lc?*); same as previous except dated l-15.vii.1984 (1$); Simcoe, 9.vi.l987, snap beans pan tp., B. Rathwell (2cT*); Simcoe Co., Washago, 44°44.76'N, 85°18.73'W, 10-19.ix.1998, PAN 5B: carrion tp., Christopherson (1

96 (2(5*); same as previous except 7.ix-5.x.l969, deciduous for. (lc?)- MN: Clearwater Co., Itasca St. Pk., 28-29.viii.1979, carrion deciduous forest., S.&J. Peck (1$). NH: Coos Co., 3mi NE East Inlet Dam, Norton Pool, 25.vi.-9.vii. 1986, FIT, D.S.Chandler (1$); Coos Co., lmi NE East Inlet Dam, 8-2l.viii. 1986, FIT, D.S.Chandler (1$); Durham Co., Source Hole Bog, 22-25.vi.1986, pan tp., S.A. Marshall (1& 1 $); Rock Co., 1 mi SW of Durham, 7-20.vii.1987, FIT, D.S. Chandler (1$); Strafford Co., Spruce Hole, 3mi SW of Durham, 15-17.X.1987, FIT, D.S. Chandler (1$). NM: Catron Co., 8mi SE of Luna, 7500', 9-14.vii.1979, pond, pine at stream, S.&J. Peck (1$). NY: Hamilton Co., Speculator, 5.viii.l983, dead worms, S.A. Marshall (1$). NC: Raleigh, 13.ix.1963 from thrips culture (1$); Swain Co., NE of Mt. Collins, 5900', 17-22.V, carrion, Newton (1$). ND: Billings Co., Medora, Roosevelt Nat. Pk., 27.viii.1979, sage grassland, carrion, S.&J. Peck (1$). OH: Champaign Co., 6mi NW of Urbana, Hume Wood, l-5.vi.1977, pig dung, R.E. Woodruff (lc?). SC: Cotteton St. Pk., 27.ix.-5.x.l983, pig dung tps., R.E. Woodruff (1$); Greenville, Greenville Zoo, N34°50.581\ W82°23.237', 12.ix.2005, ex. elephant dung, M.P. Nelder (1$). TN: Sevier Co., Greenbriar Cove, Ramsey Cascade Tr., 2700' 18-23.V.1972, dung, cove for., A. Newton (ltf*); Sevier Co., Greenbriar Cove, Ramsey Cascade Tr., 2900', cove for. (1<3).

97 Coproica flavifacies Papp, 2008

Distr.: Afrotropical: Tanzania. Coproica flavifacies Papp, 2008: 13 [male, phylogenetic notes, illustr.]. Type locality: Tanzania, Mem . HT male (HNHM).

98 Coproica galapagos sp. nov. (Figures: 134-143)

Distr.: Neotropical: Galapagos Is. (Ecuador).

Body length 1.6-2.1 mm. Body brown or light brown. Face light brown, frons brown.

Interfrontals in 4 pairs, postocellar bristles slightly convergent, 1 pair of small postocellar setulae. Eye to gena ratio approximately 2:1; 3 small genal setae arranged in a longitudinal row posterior to vibrissa with anterior seta approximately 1.5 times as long as posterior 2. Scutellum with approximately 15 setae arranged in 2-3 loose rows between basal 1/4 and basal 1/2; 2 lateral setae between larger anterior and posterior marginal scutellar bristles. Katepisternum with 2 dorsal bristles, anterior and posterior bristles each approximately 1/5 and 1/2 as long as the distance between dorsal margin of katepisternum and wing base respectively. Legs and fore coxa light brown. Mid tibia with proximal posterodorsal bristles at 1/4 and 1/2. Mid basitarsus with stout sub-apical anterior bristle, a stout sub-basal posteroventral bristle paired with a smaller anteroventral sub-basal bristle, and stout anteroventral bristle at 1/2 paired with a smaller posteroventral bristle. Second costal sector 0.8-1.0 times length of third costal sector.

Costal bypass short, terminating within 5 vein widths of R4+5. Anal vein with distinct angulate bend.

Male terminalia:

Sternite 5 rounded rectangular with posteromedial patch of microsetae, surface bristles uniform in size. Synsternite 6+7 without medial bend, but apex gently curved posteriorly. Epandrium lacking enlarged pair of dorsal bristles; subanal plate apparently absent, epandrium merging with membranous portion of anal opening ventromedially; cerci fused to epandrium, separate medially, ventral margin with long rounded medial 99 processes. Surstylus with claw-like posteroventral process, posterior and ventral margins lined with short bristles, claw-like process with pointed posteroventral process, and rounded anterior process. Postgonites symmetrical, curved posteriorly, curled at apex.

Basiphallus gently curved, posteroventral margin with elongate blade-like crest, open posteriorly. Lateral sclerite with posterodorsal angle reduced; 1 sharp medially deflected dorsal process, lateral surface covered in small membranous warts distally; central sclerite not fused to lateral sclerites, and with paired, dorsally deflected apical processes.

Membranous sheets narrow, with few sclerotized ridges sub-apically.

Female terminalia:

Tergite 8 broad laterally, narrower dorsally. Tergite 10 pollinose, narrowly fused to cerci posterolaterally; cerci rounded triangular, and pollinose, apical bristles long and sinuate.

Sternite 7 broad and rounded. Sternite 8 rounded, trapezoidal, and wider anteriorly, with

3-4 anterior bristles. Tergite 10 wrapped around posterior tip of abdomen, with medial pair of large bristles. Spermathecae cylindrical; duct length 5-7 times the length of spermathecal bulb.

Type Material:

HOLOTYPE: ECUADOR: Galapagos, Espanola Bahia Manzanillo, 7-10.vi.1985, sand beach, carrion tp., S.&J. Peck (16\ QCAZ).

PARATYPES: ECUADOR: Same as holotype (6c?*-illustr-wing, 8$*-illustr, QCAZ).

Etymology:

The specific name refers to the Galapagos Islands, the only collection locality for this species.

100 Coproica ghanensis Papp, 1979

Distr.: Afrotropical: Ghana, Nigeria, Zanzibar. Coproica ghanensis Papp, 1979a: 101 [female, taxonomic notes, illustr.]. Type locality: Ghana, Nakpanduri. HT female (HNHM). Papp, 2008: 29 [male, taxonomic notes, illustr.].

101 Coproica hirticula Collin, 1956 (Figures: 23, 144-154)

Distr.: Australasian/Oceanian: Australia (ACT, NSW, QLD), New Zealand; Nearctic: Canada (AB, BC, NB, ON, SK), USA (AK, AR, AZ, CA, FL, GA, IL, IN, KY, MA, MI, MN, MO, NM, NC, OK, SC, TN, TX, UT, WY); Neotropical: Argentina, Barbados, Bolivia, Brazil, Chile, Colombia, Costa Rica, Ecuador, Guatemala, Jamaica, Mexico (CHP, COA, HID, ROO, VER), Peru, Trinidad; Oriental: China (HKG), Taiwan; Palaearctic: Belgium, Bulgaria, Canary Is. (Spain), Czech Republic, Denmark, Germany, Great Britain, Greece (Crete), Finland, Hungary, Italy, Israel, Japan, Madeira (Portugal), Malta, Norway, Poland, Roumania, Slovakia, Spain (incl. Balearic Is.), Sweden, Switzerland, UAE. Coproica hirticula Collin, 1956: 178 [both sexes]. Type locality: England, Cambs., Kirtling. STs (OXUM, see Pont 1995: 80). - Papp, 1973c: 30 [diagnosis, key, illustr.]; Papp, 1984: 82 [Palaearctic catalog]; Marshall, 1989b: 603 [Australasian/Oceanian catalog]; Lachmann, 1996: 759-768 [morphology, male & female genitalia, copulation, illustr.]; Gatt, 2008: 698 [new record from UAE]; Rohacek, 2007b: 104 [new record from Madeira (Portugal)]. Leptocera (Coproica) hirticula. - Richards, 1960c: 202, 207 [generic combination, key].

Body length 1.2-2.1 mm. Body brown to light brown. Face light brown, frons brown.

Interfrontal bristles in 4 pairs, postocellar bristles slightly convergent, 1 pair of small postocellar setulae. Eye to gena ratio approximately 2:1; 3 genal setae in longitudinal row posterior to vibrissa. Scutellum with approximately 15 discal setae in 2 rows between basal 1/4 and basl 1/2; 3 lateral setae between larger anterior and posterior marginal scutellar bristles. Katepisternum with 2 dorsal bristles, anterior and posterior bristles each approximately 1/5 and 2/3 as long as the distance between dorsal margin of katepisternum and wing base respectively. Legs light brown, fore coxa light brown or yellow. Mid tibia without proximal posterodorsal bristles. Mid basitarsus with row of stout, small anterior setae extending from 3/5 to apex, pair of ventral basal bristles, and pair of small ventral bristles at 3/5. Second costal sector 0.7-1.0 times length of third

102 costal sector. Costa extending 7 or more vein widths beyond R4+5. Anal vein with distinct angulate bend.

Male tertninalia:

Sternite 5 rounded rectangular, with paired posteromedial tabs, right tab larger, with posteromedial patch of dense microsetae surface bristles enlarged laterally. Synsternite

6+7 straight medially. Epandrium with enlarged pair of dorsolateral bristles; subanal plate present, narrow, with medial suture; cercus fused to epandrium, ventral margin with large rounded lateral process. Surstylus curved along anteroventral margin, with row of small anteroventral bristles, and pointed posteroventral process. Postgonites symmetrical, straight, narrow, and unmodified. Basiphallus straight,tubular, posteroventral angle with small, elongate process, posterior open dorsal to crest. Lateral sclerite with bifurcate proximal dorsal process extending laterally and medially, ventral apical portion cup shaped, and with additional sharp, sinuate ventral process; central sclerite short, fused to lateral sclerites beneath proximal dorsal process. Membranous sheets elongate and broad.

Female terminalia:

Tergite 8 long laterally and short dorsally. Tergite 10 pollinose, narrowly fused to cerci posterolaterally; cerci pollinose, apicall bristles long and sinuate. Sternite 7 broad, with large triangular point on posterior margin. Sternite 8 small, with 4 minute setae along posterior margin. Sternite 10 not wrapped around posterior tip of abdomen, with lateral arms curved dorsally, and 1 pair of enlarged medial bristles. Spermathecae spherical, tapered at base; duct length 7-8 times length of spermathecal bulb.

103 Specimens Examined:

ARGENTINA: Jujuy, Abra Pampa, 3500m, 26.ii.1992, llama dung, S.A. Marshall (16c?, 19$); Jujuy, Cochinoca, 3700m, 26.ii. 1992, sweep, small stream, llama dung, debris, S.A. Marshall (111c?, 120$); Santa Fe, 12km S of Rosario, Rt 9, 19-2l.iii. 1980, dung tps., R.F. Woodruff & H. Cordo (lc?, 1$); Santa Fe, 26km S of Reconquista, Rt.ll, 2km NE of Berna, 20-2l.ii. 1980, pig dung tps., R.E. Woodruff (2c?**, 1$); Santa Fe, Cabal 65km N of Santa Fe, Rt 11, 20-2l.iii. 1980, dung, Cordo & Woodruff (1$); Salta Rosario de Lerma, INESALT yard, 16-28.ii.1992, malaise, S.A. Marshall (7c?, 12$); same as previous except 17-28.ii.1992, pan tps. (1$) ; Salta Rosario de Lerma, 24.ii.1992, swept over foul ditch, S.A. Marshall (4c?, 3$); same as previous except 29.ii.1992, cow dung (lc?, 1$). BARBADOS: Turner Hall Woods, 21.ii.1979, dung tp., S. Peck (lc?); Welchman Hall, Gully, ii.1979, Leg, dung tp., S. Peck (lc?, 1 $). BOLIVIA: Santa Cruz, Campo Guairuy, 24km S Camiri, 870m, 18.viii-2.ix.2000, malaise tp., Irwin & Hauser (lc?: debuOO 146577). BRAZIL: Parana, Curitiba, behind Nat. Hist. Museum, 5- 9.ii. 1990, FIT in woods, S.A. Marshall (lc?*); Rio e Janeiro, Nova Friburgo Mury, 1000m, 1 l.ii.1990, swept off goat dung, open wet area, S.A. Marshall (3c?*, 2$); Rio de Janeiro, Teresopolis, Pedra do Sino, 2000m, 14.iii. 1990, human dung, S.A. Marshall (26c?*****, 40$); same as previous except residue sweep (lc?*); CANADA: AB: 10 mi E of Patricia, 16-17.viii.1980, prairie, S.A. Marshall & R.S. Anderson (lc?*); Calgary, 13.vii.1980, compost heap, S.A. Marshall (5c?***, 1$); Stettler, 30.viii.1980, S.A. Marshall (3$); Wagner Bog, 25.vii.l.viii.l985, pan tp. 1, Leg, A.T. Finnamore (2c?, PMAE); same as previous except l-7.viii.1985 (lc?, PMAE); 28.viii.-10.ix.1985, pan tp. 7 (lc?, PMAE); BC: Ainsworth, 31.vii.1980, sweep, S.A. Marshall (lc?); N Shore of Shuswap lake, 50 59'N, 119 06'W, 400m, 23-3l.vii. 1987, malaise tp., O'Hara (lc?*). NB: St.Andrews, 28.viii.1978, dead seagull, S.A. Marshall (lc?). NS: Kingsport, 5.ix.l980, Dondale & Redner (1$). ON: 12.4km NNE of Searchmont, mi 10 Whiteman Dam Rd., 2.vii.l986, human dung, K.N. Barber (2c?*); Algoma Distr., Hilton Beach, 10.vi-8.vii.1989, cedar swamp, pig dung pans, J.E. Swann (1$: debu00148388); same as previous except 10.vii-18.viii. 1989, beech forest, pig dung pans (lc?: debu00176142; 1$: 00184751); Algonquin Prov. PL, Lake Sasejewan, Davies Bog, 15-17.viii.1988, dung, S.A. Marshall (lc?, 1$*); Algonquin Prov. Pk., Swan Lk. Stn., Scott Lk. Survey, 1- 30.ix.1994, A5 Sphagnum, pans (lc?); same as previous except 01-10.vii.1994, A3 Sphagnum malaise tp. (1$); Arkell, 2.vi.l956, D.H. Pengelly (lc?*); Arkell, x.1984, horse (2c?, 1$); Bruce Co., Bears Rump Is., 45 18'N, 81 32W, 26.vii-31.vii.1995, malaise, S.A. Marshall (1$); Bruce Co., Inverhuron Prov. Pk., front dunes, 44°17'33"N, 81°35'28"W, 10.ix.2003, white pans, Buck & Paiero (lc?: debu00236647); Chaffeys Locks, Queens Univ. Bio. Stn., 12.ix.1980, human dung, field, S.A. Marshall (4c?*, 7$); same as previous except field day (1$); field carrion (5c?***, 7$); Courtland, 23.vi.1980, car net, S.A. Marshall (lc?*); Crieff Fen 3km W of Puslinch, 22.vii- ll.viii.1988, pt2x, forest edge, F2xC, D. Blades (2c?); Essex Co., East Sister Is. Nat. Res., 41°49'N, 82°51'W, 30.vi.2003, shore, yellow pans, S.A. Marshall (lc?: debu00238457); Fergus, 03-06.viii.1992, black composter, S.A. Marshall (3$); Guelph, 1984, composter, B. Higgins (13c?, 9$); Guelph, 12.X.1979, carrion, S.A. Marshall (2c?); same as previous except 21.ix.1979 (1$); 18.i.l979, dead squirrel (1$); x.1981, grass clippings (lc?*); 18-i-1979, dead squirrel (lc?); Guelph, 14.vii.-16.viii.1981, pan tps.,

104 K.N. Barber (1$); Guelph, 24.vi-14.vii. 1981, K.N. Barber (1$); Guelph, Arboretum, 13- 27.X.1984, mushroom tps., deciduous, B.V. Brown (1$); same as previous except mushroom tps., field (1$*, 1?); 3-13.X.1984, field, mushroom tps. (1$); Guelph, Hanlon Conservation Area, 29.vii.-7.vii. 1984, bait tp., carrion, K. Gilbert (2c?); Hald-Norfolk Reg., Manester Tract, 6km NNW of St. Williams, 15.vi.2001, forest, decayed shrimps pans, Buck, Paiero & McKendry (1$: debuOOl58587); Hamilton, Royal Botanical Gardens, Ol.v.1998, RET - carrion, Carney & Scott (1$); Hamilton, Royal Botanical Gardens, 03.vi.1998, pan tp., carrion, V. Carney & K. Scott (4c?, 2$); Hilton Beach, 10.x-24.xii.1987, pan tps., hardwood forest, J.E. Swann (1$); Hilton Beach, 26.ix- 13.X.1987, hardwood forest, fish entrails pan tps., J.E. Swann (2$); Icewater Creek, WS, 46°53.7'N 84°03.4'W, 4-22.ix.1998, pitfalls, mixed forest, K.N. Barber (ltf); Kenora, Acid Bog Project, 25.vi-2.vii.1984, Tp. 14-17E3, D. Rosenburg (lc?*); Lambton Co., Pinery Prov. Pk., behind Meeting Centre, 05.ix. 1994, ex. deer bones, J. Skevington (l£); Meaford, ll.x.1981, compost heap, S. Marshall (1$*); Mer Bleue, 22-29.ix.1982, L. Dumouchel, Oliver Bog, 3km S of Gait, 24-30.vii.1987, pt2 expanse hollow, 02bK, D. Blades (l<$); same as previous except 8-25.ix.1988, ptl sedge pond, V1F (1$); Ottawa, 6.ix.l984, swept from liquid manure pit, J.R. Vockeroth (236*, 2$); Ottawa, vii.1990, Ex. Bus shelter, J,R, Vockeroth (1$, CNCI); Ottawa, viii.1979, grass piles, A. Telka (4c?, 1$); Pt. Pelee, 7-10.vii.1980, pan tp. on dunes, K. Barber (2c?*, 1$); Point Pelee Nat. Pk., wooded area by west beach, 21 .viii-10.ix. 1999, malaise/pan tps., O. Lonsdale (IcT, debu0001184); same as previous except dated 10-21.viii. 1999 (1

105 00112020*-illustr, 00112028); Pichincha, Valley near Hosteria San Jorge, 10km NW of Quito, 3000m, 23.X.1999, pans in grasspile, S.A. Marshall (15$*: debu00108694, 00108784, 00108782, 00108775, 00108768, 00108745, 00108675, 00108663, 00108730, 00108629, 00108628, 00108603, 00108624, 00108610, 0010860 l*-illustr; 24$: debu00108618, 00108625, 00108666, 00108652, 00108649, 00108645, 00108638, 00108635, 00108677, 00108671, 00108737, 00108729, 00108720, 00108713, 00108631, 00108633, 00108741, 00108750, 00108749, 00108743, 00108772, 00108783, 00108790, 00108687). JAMAICA: Blue Mt., Whitfield Hall, 4200', 10.ix.1988, goat dung, T.A. Wheeler (1$*, 1?); Kingston, nr. Mt. Royal Hotel, 16.iv.1959, sweeping, TB Sexton Jr. Cal. Acad. Sci. Coll. (1$, CASC). MEXICO: Chiapas, 21 mi N of Bochil, 5500', 18- 24.viii.1971, pine, oak liquid amber, human dung, A. Newton (1$); Coahuila, Boquillas, 24-26.ix.1979, pig dung tp., R.E. Woodruff & G. Fincher (1$); Hidalgo, 7 mi NE of Pachuca, 9300', vii.1971, oak, pine, human dung, A. Newton (1$); Hidalgo, 4 mi W of Tulancingo, 7600', l-6.vii.1971, oak, human dung, A. Newton (3$); Quintana Roo, Xcaret, 13.L1992, J.R. Vockeroth (3$, 1$); Veracruz, 1.7 mi N of Teocelo, 3700', 22- 24.vii.1973, human dung, A. Newton (1$). NEW ZEALAND: South Is., Banks Pen., Port Levy, 22-26.ii.1999, near sea streambed, pan tps., S.A. Marshall (2$: debu00102090, 00102034); South Is., Christ church, ca. 15km S, 'Macrocarpa', 43°42'31"S, 172°33'58"E, 1998, fungus tp. #6, exotic tree plantation, S. Hodge (1$: debu00104576; 2$: 00104588, 00104587); South Is., Otago, Lake Hawea, Hunter Valley Sta., ll-15.ii.1999, stream, pan tps., S.A. Marshall (3$: debu00102261, 00102282, 00102275); South Is., Takaka Pupu Springs, 60m, 6.ii.l999, dung tps., S.A. Marshall (23$: debu00101926, 00101914, 00101915, 00101931, 00101922, 00101903, 00101900, 00101898, 00101956, 00101904, 00101890, 00101902, 00101953, 00101969, 00101972, 00101968, 00101963, 00101887, 00101964, 00101959, 00101957, 00101960, 00101961; 23$: debu00101896, 00101952, 00101923, 00101919, 00101967, 00101962, 00101975, 00101973, 00101916, 00101954, 00101955, 00101905, 00101888, 00101965, 00101899, 00101895, 00101897, 00101889, 00101927, 00101913, 00101912, 00101894, 00101929); South Is., Wier Rd., SSW of Culverden, 42°49'53"S, 172°48'7"E, 1998, fungus tp. #13, exotic trees plantation, S. Hodge (2$: debu00104537, 00104555). PERU: Pirua, Peru Fr., 19-20-41 251, cage-cotton buds, P.A. Berry (1$*, STRI). SPAIN: Canary Is., Isla Gomera, El Cedro, 1100m, l-3.ix.1992, laurel forest dung tps., S.&J. Peck, 92-338 (1$). TRINIDAD: St. George Co., Arma Ward, NY Zool. Soc. Sta., ll.vi.1977, pig dung tp., R. Woodruff (1$); St. George Co., Curepe, 10 38'0N, 61 24'0W, 17.xii.1978, malaise, F.D. Bennett, (1$: debu00151627). UNITED STATES OF AMERICA: AR: Wash Co., 3mi S of Devils Den St. Pk., 28-31.v. 1979, oak- hickory, S.&J. Peck (1$). AZ: Cochise Co., Huachuca Mts., Carr Canyon, 7100', oak & evergreen, ix.1970, human dung, A. Newton (1$); Coconino Co., Flagstaff, 7100', 18- 25.vii.1979, pond pine meadow malaise, S.&J. Peck (1$); Santa Cruz Co., Patagonia Lk. St. PL, 9-1 l.viii. 1984, dung tp.., B.V. Brown (2$, 2$). CA: Los Angeles Co., National Forest Lightning Ridge trail (near Big Pines) 738, 12.viii.1985, horse dung, G.J. Umphrey (1$*)- FL: Alachua Co., Gainseville, 2-5.vii.1982, pig dung, horse pasture, K.L. Rench (12$, 8$); Broward Co., Hwy. 84, mi. 59, 12.xii 1985, old carrion, S. Marshall (1$, 1$); Cary St. Park, 17.iv.1989, dung, T.A.Wheeler (3$); Clay Co., Gold Head Branch State Pk., 14.iv.1971, dung, A. Newton (1$); Dade Co., Hialeah, 9.vii.l971, at human feces, C.E. Stegmaler, Jr. (1$); Liberty Co., Torreva SP., 6.vi.l982,

106 pig dung tp., R.E. Woodruff (1$); Liberty Co., lOmi SW of Juniper, Rt.12, 26.iii.1983, pig dung, turkey-oak, Woodruff & Thomas (IcT, 4$); Manatee Co., Longboat Key, 3- 4.vii.l983, pig dung tps., R.E. Woodruff (2$); Marion Co., 175, 4mi N exit67, 12- 18.vi.1983, pig dung tps., R.E. Woodruff (lc?); Marion Co., Ocala NE Rd.65, 1.5mi W of St. Rd. 19, 15-16.iii.1984, dung, R. Woodruff (1$); same as previous except Rd 90, 8mi N St. Rd. 19 (1$); Marion Co., Rt. 316 near Eureka, 15-16.iii.1984, pig dung tp., Woodruff (1(J); Okaloosa Co., 4.5mi N of Holt, Blackwater R. State For., Fla. A&M Res. Sta., 15-17.vi.1978, pig dung, L. Stange (5c?*, 3$); Polk Co., 2mi N of Lake Wales, 25- 29.iv.1983, pig dung tp., R.E. Woodruff (1$); Suwannee Co., 8.7 mi W of O'Brien, S- 349, 25.iv.1977, pig dung tp. #6 gopher tortoise burrow, Woodruff & Woley (1$, 1$). GA: 14mi N of Folkston, 10-18.iv.1989, pan tp., human dung, J.E. Swann (1$); Rabun Co., Terrora Pk., Tallulah Falls, 4-ll.vi.1983, pig dung tps., KW Vick (1$). IL: Cook Co., Palos Pk., 13.X.1965, leg, H.S. Dybas (1$). IN: Lawrence Co., Hoosier Nat. For., Donaldson Tr., 19.V.1991, J.E. Swann (1$); Putnam Co., near Cunot. Lieber St. Pk., 31.vii.1984, vac. dog dung, L.B. Carlson (1& 3$). KY: Rowan Co., 24km SW of Morehead, Cave Run Lk., 14.v-20.vii.1983, Fagus forest, FIT, S.&J. Peck (1$). MA: Middlesex Co., Medford, 7.ix-5.x.l969, deciduous for., A. Newton (1$); same as previous except ll.viii-5.x.l969, pine for., carrion (lc?); Middlesex Co., Bedford, vii.1969, mixed for., carrion, A. Newton (lc?)- MI: Gogebic Co., Crooked L. Bog, 15.viii.1977, Leg: J. Wagner (lc?); same as previous except 23.vi.1978, Ber(?) otter dung & root mat subst. (1$). NM: Santa Fe Co., Santa Fe Baldy, 10800-11800', 19.vi.- 4.vii.l979, spruce-fir-tundra, S.&J. Peck (1$, 1$); Socorro Co., 20mi W of Socorro, Water Canyon, 7000', 28.vi-7.vii. 1979, mixed "mesic" forest, S.&J. Peck (3c?**); Valencia Co., 2mi W of Los Lunas, 4900', 16-30.vi.1979, desert grassland carrion tps., S.&J. Peck (1$). NC: Bladen Co., Singletary Lk. St. Pk., 34°35'N, 78°27'3W, 19- 22.V.2003, pans, Marshall & Paiero (5c?-wing: debu00227146, 00252099, 00252090, 00227354-wing, 00227347; 3$: 00252121, 00227378, 00227374); Swain Co., NE slope of Mt. Collins, 5900', Pices-Abies, 17-22.V.1972, human dung, A. Newton (1$). OK: Lattimer Co., 5mi W, 5-1 l.vi.1977, red oak, dung tp., K. Stephen (lc?). SC: Barnwell St. Pk., 10-18.iv.1989, pan tps., mushrooms in areas near Lake, S.A. Marshall (lc?); Richland Co., Columbia, Riverbanks Zoo & Botanical Gardens, N34°00.582', W81°04.560', 22.vi.2007, M.P. Nelder (1$, lc?). TN: Sevier Co., Greenbriar Cove, 1700', 19-23.V.1972, A. Newton (1$); Sevier Co., Greenbriar Cove, Ramsey Cascade Tr., 2900', cove forest (16*)- TX: Bastrop Co., Bastrop St. Pk., muddy lake shorepan tps., 8.iv.2000, M. Buck (1$: debuOO 114052). UT: Daggett Co., Dutch John, 6300', 2- 8.viii.l979, pinyon juniper, woodland, carrion, S.&J. Peck (3c?, 4$); Duchesne Co., Mirror Lk., 10300', 30.vii-12.viii.1979, spruce fir meadow, carrion-malaise, S.&J. Peck (lc?); Duchesne Co., Uinta Mts., Rocky Sea Pass, 11200', 30.vii-12.viii.1979, carrion, rocky tundra, S.&J. Peck (1$). WY: Big Horn Co., lOmi E of Lovell, 3800', 17- 26.viii.1979, sagegrass-barren carrion, S.&J. Peck (3c?, 1$); Big Horn Co., lOmi E of Lovell, 3800', 17-26.viii.1979, sage-grass-barren carrion, S.&J. Peck (2$); Red Gulch Road, Hwy. 14, 5-19.viii.1990, near shell pans in desert area, cow dung; J.E. Swann; same as previous except in barren area near cottonwoods (1$); in grass hummock in desert area, cow dung (1$); Sublette Co., 7 mi N of Pinedale, 9000', 19-24.viii.1979, pine meadow, carrion malaise, S.&J. Peck (1$); Teton Co., 6800' Moran Jet., 17-

107 24.viii.1979, sage steppe, carrion, S.&J. Peck (1$); Uinta Co., 8mi SE Evanston, 7100', 30.vii-11 .viii. 1979 carrion, sage-grass, riparian, S.&J. Peck (1$*, 1 $)•

Comments:

Coproica hirticula has a cosmopolitan distribution and is one of the most common species of Coproica in the New World. Since C. hirticula's sister species, C. novacula, is an exclusively New World species, it is likely that C. hirticula is native to the New World.

108 Coproica hirtula (Rondani, 1880) (Figures: 155-164)

Distr.: Afrotropical: Congo, Ethiopia, Ghana, Kenya, Madagascar, Nigeria, Seychelles, Yemen, Zaire; Australasian/ Oceanian: Bonin Is. (Japan), Guam (USA), Hawaii (USA), Kiribati (Onotoa Atoll), Marshall Is., Micronesia (Kapingamarangi Atoll, Satawal I., Yap Is.), New Zealand, Northern Mariana Is. (Saipan), Papua New Guinea, Palau, Pitcairn Is.; Nearctic: Canada (AB, BC, NB, ON, QC), USA (AR, AZ, CA, FL, GA, IA, ID, IL, KS, MA, MI, MN, NC, NE, NJ, NM, NY, OR, PA, UT, WA, WY); Neotropical: Argentina, Chile , Costa Rica, Galapagos Is., Jamaica, Mexico (NLE), (other records: Bermuda and Bolivia, are misidentifications of C. rufifrons); Oriental: China (HKG), India, Malaysia, Nepal, Pakistan, Sri Lanka, Taiwan, Vietnam; Palaearctic: Afghanistan, Andorra, Austria, Azores (Portugal), Belgium, Bulgaria, Canary Is. (Spain), Czech Republic, Egypt, Finland, Germany, Great Britain, Greece (Thira), Hungary, Italy, Israel, Japan, Latvia, Madeira (Portugal), Malta, Netherlands, North Korea, Norway, Poland, Romania, Russia (CET), Slovakia, Spain (incl. Balearic Is.), Sweden, Switzerland, Tunisia, Uzbekistan, Yugoslavia (Serbia); South Atlantic: Saint Helena. Limosina hirtula Rondani, 1880: 40 [sex not stated]. Type locality: Italy, nr. Parma. STs, sex unknown (MZUF). Limosina (Coprophila) hirtula. - Duda, 1918: 222-225 [subgeneric combination, redescription, both sexes, illustr.]; Duda, 1938: 164-165 [redescription, illustr.]. Leptocera (Coprophila) hirtula. - Duda, 1925: 206 [generic combination]. Leptocera (Heteroptera) hirtula. - Richards, 1930: 308 [subgeneric combination]. Coprophila hirtula. - Vanschuytbroeck, 1951b: 20 [generic combination]. Limosina (Coproica) hirtula. - Richards, 1952a: 431 [subgeneric combination]. Leptocera (Coproica) hirtula. - Harrison, 1959: 261 [subgeneric combination, diagnosis]; Richards, 1960c: 202, 206-297 [key, taxonomic notes]; Richards, 1965c: 725 [Nearctic catalog]; Richards, 1967b: 16 [Neotropical catalog]; Tenorio, 1968: 196, 200-201 [diagnosis, male genitalia, illustr.]; Okely, 1974: 55-56 [puparium, illustr.]; Richards, 1980: 618 [Afrotropical catalog]. Coproica hirtula. - Frey, 1945: 80 [generic combination]; Papp, 1973c: 28 [key, illustr.]; Hackman, 1977: 400 [Oriental catalog]; Papp, 1984: 82 [Palaearctic catalog]; Marshall, 1989b: 603 [Australasian/Oceanian catalog]; Kobayashi et al., 1990: 111 [bacterial flora on adults]. Limosina exigua Adams, 1904: 454 [a primary junior homonym of Limosina exigua Rondani, 1880 = Minilimosina fungicola (Haliday, 1836); both sexes]. Type locality: USA, New Mexico, Las Cruces. STs, both sexes (? SEMC). - Spuler, 1925a: 123 [homonymy]; Richards, 1960c: 206 [synonymy]. Limosina (Coprophila) hirtula var. crinita Duda, 1918: 224 [both sexes]. Type locality: Germany, Usedom. STs (?ZMHB). - Papp, 1984: 82 [synonymy]. Coprophila hirtula var. crinita. - Vanschuytbroeck, 1951b: 20 [generic combination]. Leptocera (Coproica) hirtula var. crinita. - Richards, 1960c: 207 [taxonomic notes]. Leptocera (Coprophila) exiguella Spuler, 1925: 123 [nom.n. for Limosina exigua Adams,

109 1904]. Type locality: USA, New Mexico, Las Cruces. - Richards, 1960c: 206 [synonymy].

Body length 1.4 - 2.0 mm. Body colour brown. Face brown, frons dark brown.

Interfrontal bristles in subequal 4 pairs, postocellar bristles convergent or cruciate, 1 pair of small postocellar setulae. Eyes small, eye to gena ratio approximately 2:1; 3-4 small genal setae in longitudinal row, anterior seta approximately 2 times length of posterior setae. Scutellum with approximately 15 setae in 2 loose rows between basal 1/4 and basal 1/2; 2-3 lateral setae between larger anterior and posterior marginal scutellar bristles. Katepisternum with 2 dorsal bristles, anterior and posterior bristles each approximately 1/3 and 2/3 as long as the distance between dorsal margin of katepisternum and wing base respectively. Legs and fore coxa brown to light brown.

Mid tibia with additional small anterodorsal bristle at 1/7 and small proximal posteroventral bristles at 1/5 and 2/5. Mid basitarsus with 2-3 (usually 2) stout anterior bristles at 3/5 and sub-basally, 2 anteroventral sub-basal bristles with distal anteroventral

2 times length of proximal bristle, a posteroventral sub-basal bristle equal in length to distal anteroventral bristle, and 2 small ventral pairs of setae on distal half. Second costal sector 0.8-1.1 times length of third costal sector. Costal bypass long, extending 7 or more vein widths beyond R4-1-5. Anal vein sinuate, lacking an angulate bend.

Male terminalia:

Sternite 5 rounded rectangular, surface bristles uniform in size. Synsternite 6+7 straight medially. Epandrium with enlarged pair of dorsolateral bristles; subanal plate apparently absent, epandrium merging with membranous portion of anal opening ventromedially, cerci discontinuous medially; cercus fused to epandrium, ventral margin with 2 rounded

110 medial processes. Surstylus with quadrate posterior process, ventral margin straight, with

12 or more elongate posterior and ventral marginal bristles. Postgonites symmetrical, curved anteriorly, strong posterior angle, narrow on apical 1/3, apically blunt.

Basiphallus gently curved, tubular, posteroventral margin with shallow rounded crest, open posteriorly dorsal to crest. Lateral sclerites elongate, rounded, and extended apically, with 2 dorsal processes, second process positioned ventral and distal to first; central sclerite fused to lateral sclerites near second process. Membranes long with small sclerotized spines on surface.

Female terminalia:

Tergite 8 long laterally and only short dorsally. Tergite 10 pollinose, narrowly fused with cerci posterolaterally; cerci rounded rectangular, pollinose, apical bristles long and sinuate. Sternite 7 broad, rounded rectangular. Sternite 8 rounded rectangular, approximately as wide as sternite 7. Tergite 10 wrapped around posterior tip of abdomen with pairs of medial and lateral enlarged bristles. Spermathecae elongate, ovoid; duct length 2-3 times length of spermathecal bulb.

Speciemens Examined:

ARGENTINA: Jujuy, Abra Pampa, 3500m, 26.ii.1992, llama dung, S.A. Marshall (1$). CANADA: AB: Onefour, 930m, 49°2'14N, 110°28'15W, 29.vi.2001, creek bed, sweep, S.A. Marshall (ltf: debu00170163); Wagner Bog, l-7.viii.1985, pan tp. 1, Leg, A.T. Finnamore (1$*, PMAE); same as previous except 12-25.vi.1985, pan tp. 2 (1$, PMAE); 26.vi-2.vii. 1985, pan tp. 2 (ltf*, PMAE); 25.vii-l.viii. 1985, pan tp. 9 (1& PMAE); 12-25.vi.1985, pan tp. 10 (1^*, PMAE); Writing On Stone Prov. Pk., Willow N, 30.viii-10.ix.1990, M. Klassen (1$). BC: lOmi W of Hope Jones Ck., 8-28.vii.1980, wet for. carrion, S.A. Marshall (l<$). ON: 12.4km NNE Searchmont, mi 10 Whitman Dam Rd., 7.vii.l986, dead Marmota, K.N. Barber (2$); Algonquin Prov. Pk., Swan Lk. Stn., Scott Lk. Survey, 23-30.vi.1994, A5 shore pans (1§); Bruce Co., Covels, 45°18'N, 81°46'W, 26.vii-31.vii. 1996, malaise, S.A. Marshall (1$); Crieff Bog, 3k N of Puslinch, 4-17.V.1988, pt3b sedge is., D. Blades, C36W (1$); Essex Co., Middle Is., 41°41'N, 82°41'W, 4.vii.2003, yellow pans, Marshall & Cheung (1$: debu00222061; 1$*: debu00222065*-illustr); Guelph, 1984, composter, R. Higgins (lie?*, 3?*, 8?); Guelph,

111 4.iii. 1989, chicken manure, S. Gray (12$*, 6$*); same as previous except 11 .ii. 1989, M. Ross (lowing); 3.iii.l989, C. Grinton (2$**, 2?*); Guelph, 17.vi.1983, grass pile, S.A. Marshall (1$*); Guelph, 21.X.1959, D.H. Pengelly (1$*, 3$); Guelph, 21-25.vii.1993, black composter, D.C. Caloren (1$); Guelph, 28.V.1980, carrion, S.A. Marshall (1$); Guelph, Arboretum, 9-1 l.vi. 1984, coniferous for. mushroom tps., B.V. Brown (1$); Guelph, Hanlon Conservation Area, 25.vi.1984, mushroom bait tp., K. Gilbert (2$); Guelph, Hanlon Conservation Area, 29.vi-7.vii. 1984, bait tp. carrion, K. Gilbert (3$, 1$); Hamilton, Royal Botanical Gardens, 3.vi.l998, pan tp., carrion, V. Carney & K. Scott (3$); same as previous except 8.vii.l998 (1$); 3.vii.l998 (1$); 15.vi.1998, pan tp., RBT control (1$); Hilton Beach, 14.X.1989, ex lichen and moss, J. Swann (1$, 4$); Hilton Beach, 16.ix.-14.x. 1989, sugar maple forest pan tps., J.E. Swann (1$, 1 $); Hilton Township, Tanby Bay, 30.vi.1992, malaise at edge of lakeshore & poplar forest, leg, J.E. Swann (1$*); Kars 24.xii.1978, M. Kaulbars (1$, 2$); Kent Co., Rondeau Prov. Pk., South Point Tr., east parking lot, 42°15'42N, 81°50'49W, 10.vi.2003, oak savanna, S.A. Marshall (1$: debuOl 125367); Marmora, 9.vii.l957, K.D. Southern (1$); Ottawa, viii.1979, grass piles (1$); Pt. Pelee, 7-10.vii.1980, pan tp. on dunes, K. Barber (1$); Pt. Pelee, 31.vii.1978, W.A. Attwater (1$); Pt. Pelee, Nat. Pk., Visitor Centre, 22- 30.vii.2000, malaise & pans, O. Lonsdale (1$: debuO 1004922); Sault Ste Marie, Bristol Place, 21.viii.1987, grass clippings, K. Barber (1$); Simcoe, 19.viii.1985, light tp., S. Enrich (1$); Simcoe Co., Washago, 44°44.76'N, 85°18.73'W, 28.v-4.vi. 1998, PAN5B, carrion, Christopherson (1$); Wainfleet Bog, 8km S of Welland, 28.vi-3.vii.1988, pt 1, north ditch, A. Stirling, NIB (2$*-illustr); same as previous except 4-1 l.vii. 1988, pt 2, 1962 zone (1$); Waterloo Reg., Blair, RARE, Blair Tr., 43°22'38" N, 80°20'34"W, 9- 12.vi.2006, yellow pans in grass clipping, Marshall & Cheung (1$: debu00271173); Wylde Lk., Bog 8k E of Arthur, 16-22.vi.1987, W4E pt4 floating mat, D. Blades (2$). NB: St. Andrews, 15.viii.1979, dead seagull, S.A. Marshall (1$); St. Andrews, MP, 30.vi.1978, S.A. Marshall (1$). QC: Hull, 21.ix.1983, wet flooded forest, L. Dumouchel (1$). COSTA RICA: Guanacaste, Est. Pitilla, 700m, 9km S Sta. Cecilia, II curso Parataxon., v.1990, L-N-330200,380200 (1$). ECUADOR: Galapagos, Espanola, Bahia Manzanillo, 7-10.vi.1985, sand beach carrion tp., S.&J. Peck (2$, 16$); Galapagos, Espanola, Bahia Manzanillo, 5-10.vi.1985, Prosopis grove behind beach, carrion tps., S.&J. Peck (3$); Galapagos, Santa Fe Is., 3-6.iv.1988, sea lion dung tp., littoral, B.J. Sinclair, 89-180 (8$, 8$). JAMAICA: St. Catherine, 28.vii-l.viii.1974, dung under trees in horse pasture, Peck (2$). MEXICO: Nuevo Leon, Surface at Cueva del Camino, 8500', 23.viii.1973, R. Jameson (1$). UNITED STATES OF AMERICA: AR: Wash Co., 3mi S of Devils Den St. Pk., 28-31.V.1979, oak-hickory, S.&J. Peck (3$, 2$); Wash Co., Devils Den St. Pk., Devils Den Cave, 28-31.v. 1979, S.&J. Peck (3c?*). AZ: Coconino Co., Tuba City, 4700', 22-25.vii.1979, desert grass shrub, carrion tps., S.&J. Peck (2$); Logan Co., Ozark Nat. For., Magazine Mt., 23.v-8.vi.1991, pans, mushroom baited, J.E. Swann (1$); Navajo Co., 15mi SE Holbrook, 14-16.vii.1979, 5300', grassland carrion tps., S.&J. Peck (1$). CA: Santa Clara Co., Del Peurto Rd., (Rdl30), 28.V.1992, J. Skevington (1$). FL: Gray Co., Gold Head Branch St. PL, 4-14.iv.1971, carrion, A. Newton (1$). GA: Rabun Co., Terrora Pk., Tallulah Falls, 4-ll.vi.1983, pig dung tps., K.W. Vick (1$). MA: Culture ex. chicken shit, Hubbard stn. (1$). NM: Bernalillo Co., 7mi S of Tijerus, 2400', 23-26.viii.1975, carrion tp. pinyon-juniper, S. Peck (1$, 1$); Bernalillo Co., Isleta, 4900', 16-20.vi.1979, floodplain forest, carrion,

112 S.&J. Peck (3$); Eddy Co., 30mi WSW of Carlsbad, Sitting Bull Falls, 4600', 23- 27.vii.1975, dung 15, S. Peck (1$, 2$); Guadalupe Co., Santa Rosa Lk. St. Pk., 3.viii.l984, desert, blacklite, B.V. Brown (1$); Guadalupe Co., Santa Rosa St. Pk., 3- 4.viii.l984, dung tp., B.V. Brown (1$); Lincoln Co., 1 mi N of Sierra Blanca Ski, 11 000', 10-26.vi.1979, alpine grass meadow, carrion, S.&J. Peck (14$); Santa Fe Co., 14mi NE of Santa Fe, 9600', 18.vi-3.vii.1979, aspen w/stream, S.&J. Peck (1$); Santa Fe Co., Santa Fe Baldy, 10800-11800', 19.vi-4.vii.1979, spruce-fir-tundra, S.&J. Peck (1$, 1$); Socorro Co., 20mi W of Socorro, Water Canyon, 7000', 28.vi.-7.vii. 1979, mixed "mesic" forest, S.&J. Peck (1$); Socorro Co., 3mi N of Socorro, 4700', 28.vi-6.vii. 1979, Larrea desert, S.&J. Peck {\<$, 1$); Torrance Co., 6mi SW of Manzano, Red Can. Camp, 75- 9000', 17-30.vi.1979, spruce-fir litter, S.&J. Peck (ltf); Valencia Co., 2mi W of Los Lunas, 4900', 16-30.vi.1979, desert grassland, carrion tps., S.&J. Peck (4$). NY: Syracuse, 12.iii.1976, ex. nest of screech owl, J.R. Philips, CNC det. lot 77-654 (\<$, CNCI); Syracuse, 24.vi.1976, ex. nest of American Kestrel; J.R. Philips, CNC det. lot 77- 654 (2

Comments:

Papp (2008) speculated that the species currently accepted as Coproica hirtula in fact pertains to C. hirtula var. crinita (Duda, 1918), and that C. hirtula (Rondani) was redescribed as C. rufifrons by Hayashi (1991). C. hirtula is treated here according to the widely accepted concept of the species, following Hayashi's concept of C. rufifrons.

This is in the best interest of taxonomic stability, but it is possible that nomenclatorial changes will be required if the types of C. hirtula and C. hirtula var. crinita are critically examined.

113 Coproica hirtuloidea (Duda, 1925) (Figures: 165-176)

Distr.: Nearctic: USA (GA); Neotropical: Argentina, Bolivia, Brazil, Colombia, Costa Rica, Peru, Venezuela. Leptocera (Coprophila) hirtuloidea Duda, 1925: 206 [both sexes]. Type locality: Peru, Cuzco [3,200-4,000 m]; Bolivia, Mapiri, Lorenzopata. STs (SMTD). Leptocera (Coproica) hirtuloidea. - Richards, 1960c: 206 [subgeneric combination]; Richards, 1967b: 16 [Neotropical catalog]. Coproica hirtuloidea. - Papp, 1979a: 102 [generic combination].

Body length 1.2 - 2.0mm. Body brown. Face and frons brown. Interfrontals in 3 subequal pairs, postocellar bristles convergent, postocellar setae absent. Eye to gena ratio

2.1:1; 3 genal setae in longitudinal row posterior to vibrissa, anterior seta approximately

1.5 times length of posterior setae. Scutellum with approximately 10 setae in 2 loose rows between basal 1/4 and basal 1/2; 2 lateral setae between larger anterior and posterior marginal scutellar bristles. Katepisternum with 2 small, subequal dorsal bristles, each approximately 1/6 as long as the distance between dorsal margin of katepisternum and wing base. Legs and fore coxa brown. Mid tibia with small, proximal posterodorsal bristle at 1/3. Mid basitarsus with row of 3-4 small anterior bristles between 2/3 and apex, anteroventral row of 5-6 small bristles along entire length, often 2-3 small posteroventral bristles in variable positions. Second costal sector 0.7-1.0 times length of third costal sector. Costal bypass short, terminating within 5 vein widths of R4+5. Anal vein with distinct angulate bend.

Male terminalia:

Sternite 5 rounded rectangular with posteromedial patch of denser setae posteromedially, surface bristles uniform in size. Synsternite 6+7 straight medially. Epandrium with enlarged pair of dorsolateral bristles; subanal plate apparently absent, epandrium merging

114 with membranous portion of anal opening ventral medially, cerci discontinuous medially; cercus fused to epandrium, ventral margin with pointed medial process, left process twice the length of right process. Surstylus with quadrate posteroventral lobe with patch of small ventral setae, ventral margin straight. Postgonites asymmetrical, curved anteriorly, left postgonite broad, blunt at apex, right postgonite narrow and sinuate in distal 1/2, blunt at apex. Basiphallus L-shaped, tubular, posteroventral margin unmodified, closed posteriorly. Lateral sclerites with membranous process at proximal dorsal angle, 2 small dorsal processes subapically, an additional small medial projection on proximal process, and a small pointed process on ventral apical angle; central sclerite fused to lateral sclerites subapically. Membranous sheets short, broad.

Female terminalia:

Tergite 8 long laterally reduced to medial connecting membrane dorsally. Tergite 10 pollinose, narrowly fused to cerci posterolaterally; cerci rounded and triangular, apical bristles long and sinuate. Sternite 7 broad, rounded rectangular. Sternite 8 rounded with a blunt anteromedial process, and a row of 4 small setae anterior and posterior to a pair of enlarged medial setae. Sternite 10 not wrapped around posterior tip of abdomen, with a blunt anteromedial process, and pair of enlarged medial bristles. Spermathecae pear- shaped; duct approximately 3-4 times length of spermathecal bulb.

Specimens Examined:

ARGENTINA: Jujuy, Cochinoca, 3700m, 26.ii.1992, sweep small stream llama dung, debris, S.A. Marshall (6<3\ 7$*); same as previous except dated 29.ii.1992 (l<$); Salta, 4.5km W of Salta, 1950m, El Alisal, l-29.xii.1987, moist ravine thicket; malaise FIT, S.&J. Peck (1$); Salta, 17 km N SBP, S7-134 La Caldera, Altodela Sierra, 1550m, 2- 30.xii.1987, malaise-FIT, sub tropical humid for., S.&J. Peck (1$); Salta, 30km E of Salta Campo Quijano, 20.ii.1992, sweep, dung, S.A. Marshall (14c?**, 17$**); same as previous except stream debris, leaf litter (IcT); sweep, forest vestige (IcT, 1 $); sweep wet mud (1$); Salta, 100 km W of Salta, Santa Rosa de Tastil, 3000m, 21.ii.1992, very dry

115 along creek, S.A. Marshall (2c?, 1$); Salta, 1500m, La Caldera Camino la Cornisa, 27.ii.1992, roadside forest sweeps, S.A. Marshall (2$*); Salta Campo Quijano, road to San Antonio do los Cobres, 18.ii.1992, above forest level, site 2&3, S.A. Marshall (1$); Salta, Canyada la Gotera, 15km W of Chicoana, 16-28.ii.1992, for. remnant, S.A. Marshall (3$); Salta, El Maray, 50km W of Chicoana, 28.ii.1992, dry area sweep, pampas grass along creek, S.A. Marshall (1$); Salta, El Maray 50km W of Chicoana, 2500m, 18-28.ii.1992, dry, pan tps., S.A. Marshall (5c?, 1$); Salta, El Ray Nat. Pk., 900m, Rio La Salsa, 5-10.xii.1987, FIT malaise tp., open stream side in for., S.&J. Peck (3c?, 3$); same as previous except ll-15.xii.1987, Aguas Negres Trail, for., malaise tp., FIT, (2c?*, 1$); Salta Sumalao, 19.ii.1992, sweep, wet muddy area along field, S.A. Marshall (lc?); Salta, Rosario de Lerma, 24.ii. 1992, swept over foul ditch, S.A. Marshall (2c?, 1$); Salta, Rosario de Lerma, INESALT yard, 16-28.ii.1992, malaise, S.A. Marshall (18c?, 6$*); same as previous except malaise overflow (lc?); 17-28.ii.1992, pan tps. (1$*). BOLIVIA: La Paz, Chulumani, Apa Apa Reserve, 2000 m, 16°21, 15S, 67°30'21W, l.iv.2001, sweep, S.A. Marshall (1$: debu00208895); La Paz, Zongo id., -3800 m, 16°10'4S, 68°7'49W, 18.iv.2001, alpine shrub, on cow dung, S.A. Marshall (16c?: debu00188793, 00188773, 00188746, 00190078, 00190133, 00188796, 0018702, 00188852, 00188855, 00188821, 00188807, 00188868, 00188867, 00188723, 00188714, 00188713; 12$: 00190095, 00190137, 00190090, 00190052, 00188733, 00190156, 00188812, 00188860, 00190109, 00190159, 00188695, 00188701); Santa Cruz, Rio Piray near Aguaclara, 3.X.1996, P. Bettella (5c?: debu00101403, 00101419, 00101410, 00101087, 00101335). BRAZIL: Parana, Londrina, Mata dos Godoy, 30.i.-2.ii.l990, S.A. Marshall (1$); Rio Jacara, Pagua 22.U990, wet litter by stream, S.A. Marshall (lc?); Rio de Janiero, Nova Friburgo Mury, 1000m, 11 .ii. 1990, swept off goat dung, open wet area, S.A. Marshall (1?*); Sao Paulo., Sao Jose do Barreiro, 1650m, i.1969, malaise tp., M. Alvarenga (2c?**). COLOMBIA: Antioquia, La Ceja, 2100mt., 3.ii.l984, manure pile (lc*, 4$). COSTA RICA: Heredia, Santo Domingo, INBio Pk., 3.x. 1999, leaf litter, Marshall & Buck (1?: debuOO 107267); San Jose, Zurqui de Moravia, 1600m, iii.1981, P. Hanson (1$: debu00100688). ECUADOR: Cachi, Guandera For. Res., 15km E of San Gabriel, 3300 m, l.xi.1999, trail to station, sweeping, S.A. Marshall (2c?: debu00108138, 00108133); Cotopaxi, 45km NNE of Latacunga, 3700m, 19-25.vii.1985, shrub grass, paramo, carrion, S.&J. Peck (14c?*: 6$*); Cotopaxi, 40km N of Latacunga, 3350m, 17- 25.vii.1985, parano, carrion, S.&J. Peck (37c?*x7, 29?**); Napo, above Papallacta, 4000m, ii.1983, paramo & elfin forest, M. Sharkey & L. Masner (2c?, 1 $); Napo, Baeza, 1900m, 12-15.ii.1983, FIT, L. Masner (1$); Napo, Baeza, 1500m, 16-19.V.1987, wet montane rain forest, malaise tp., Coote & Brown (1$: debuOO 140734); Napo, Quito- Baeza Rd., above thermal spgs., Papallacta, 3200m, 22-24.ii.1983, pan tps., L. Masner (1$); Napo, SierrAzul Res., 14km W of Cosanga, 2200m, 0°40'55S, 77°56'9W, 9.V.2002, sweep tr., M. Buck (7c?: debuOOl86860, 00186536, 00185909, 00185908, 00185907, 00186859, 00185844; 3?: 00186534, 00185905, 00186535); same as previous except 9-10.V.2002, treefall, yellow pans, (6c?**: debu00202488, 00202385*, 00202465, 00202340, 00202513, 00202356*-illustr; 4$**: debu00202401, 00202370, 00202358*- illustr, 00202403*-illustr); same as previous except lO.v.2002, treefall (29c?*: debu00195406, 00195372, 00195357, 00195412, 00195413, 00195416, 00195415, 00195394, 00195363, 00195383, 00195361, 00195424, 00195408, 00195405, 00195396, 00195392, 00195378, 00195385, 00195393, 00195370, 00195354, 00195402, 00195403,

116 00195404, 00195390, 00195381, 00195399, 00195377, 00195380*; 27$*: 00195358, 00195411, 00195422, 001954221, 00195409, 00195423, 00195425, 00195355, 00195374, 00195387, 00195400, 00195391, 00195386, 00195368, 00195401, 00195362, 00195371, 00195384, 00195373, 00195375, 00195360, 00195352, 00195359, 00195353, 00195365, 00195379, 00195364*); Napo, SierrAzul Lodge, 14km W of Cosanga, 2200 m, 5.xi.l999, for. sweep, S.A. Marshall (lc?: debuOO 140343; 1$: debu00140294); same as previous except sweeping disturbed tr. (10c?*: debu00115097, 00115115, 00115148, 00115151, 00115153, 00115074, 00115064, 00115172, 00115168, 00115133*; 25$*: 00115079, 00115083, 00115092, 00115098, 00115099, 00115101, 00115108, 00115110, 00115111, 00115120, 00115126, 00115159, 00115165, 00115138, 00115141, 00115143, 00115144, 00115145, 00115075, 00115158, 00115166, 00115171, 00115167, 00115132*); Napo, SierrAzul Lodge, 14km W of Cosanga, 2200m, 0°40'55S, 77°56'9W, 9-10.V.2002, malaise, Buck & Paiero (44c?***: 00202656, 00202658, 00202663, 00202514, 00202538, 00202651, 00202652, 00202542, 00202672, 00202673, 00202675, 00202662, 00202561, 00202565, 00202638, 00202630, 00202699, 00202709, 00202627, 00202682, 00202691, 00202635, 00202603, 00202611, 00202620, 00202619, 00202681, 00202578, 00202616, 00202683, 00202557, 00202554, 00202567, 00202660, 00202655, 00202654, 00202648, 00202646, 00202618, 00202686, 00202604*, 00202598*, 00202625*; 46$**: debu00202669, 00202644, 00202659, 00202668, 00202667, 00202665, 00202553, 00202649, 00202536, 00202530, 00202526, 00202711, 00202564, 00202558, 00202522, 00202521, 00202671, 00202700, 00202705, 00202697, 00202696, 00202703, 00202692, 00202592, 00202704, 00202690, 00202684, 00202622, 00202623, 00202621, 00202624, 00202626, 00202610, 00202609, 00202614, 00202615, 00202602, 00202678, 00202573, 00202579, 00202587, 00202575, 00202591, 00202677, 00202529*, 00202617*); Napo, SierrAzul Res., 14km W of Cosanga, 2200 m, 0°40'55S, 77°56'9W, lO-ll.v.2002, treefall, yellow pans, Marshall & Paiero (1$: debuOO 194799); Pichincha, Bellavista Res., trail 'B', 30.x. 1999, sweeping, S.A. Marshall (lc?: debuOOl 16316; 3$: debuOOl 16336, 00116286, 00116240); Pichincha, Cotopaxi Nat. PL, Quebrada Mishahuaicu, 3600 m, 26.x-8.xi. 1999, pans along stream, S.A. Marshall (1$: debuOO 142674); Pinchincha, Nanegalito, 7km SE of trout farm 'San Jose', 1500 m, 27- 30.X.1999, riverine for., sweeping tree falls, S.A. Marshall (2$: debu00140007, 00140098); Pichincha, Nono, 8 km N road to Mindo, 25.ix.1999, roadside, sweeping, S.A. Marshall (2c?*: debuOOl 12077, 00112083*; 2$*: 00112067, 00112084*); same as previous except 15 km NW road to Mindo, 24.X.1999, roadside sweep, cow dung (8c?*: debuOOl15301, 00115286, 00115282, 00115276, 00115275, 00115290, 00115298, 00115273*; 4$*: 00115299, 00115281, 00115266, 00115274*); Pichincha, Quito, 10km NW, Valley near Hosteria San Jorge, 23.x. 1999, pans in grasspile, S.A. Marshall (3c?- wing: debuOOl 15594-wing, 00115595, 00115577; 2$: 00115590, 00115597); same as previous except 3050m, 22-25.X.1999, creek bed, pan tps. (2c?: debuOOl 12034, 00112023; 2$: 00112026, 00112019); Pichincha, 34.5 km E Tandapi, 2835 m, 24- 29.vi.1975, bamboo shrub, dung tp., S. Peck (lc?: debu00261171; 4$: 00261132, 00261153, 00261154, 00261180); Pichincha, 3.5 km SE of Tandayapa, 28.X.1999, in green leaf litter (tree fall), S.A. Marshall (4c?*: debuOOl 16070, 00116071, 00116077, 00116657*; 2$*: 00116634, 00116104*); Pichincha, valley near Hosteria San Jorge, 10km NW of Quito, 3000 m, 23.X.1999, pans in grasspile, S.A. Marshall (79c?**: debu00108727, 00108725, 00108721, 00108719, 00108718, 00108717, 00108715,

117 00108714, 00108711, 00108608, 00108641, 00108640, 00108636, 00108732, 00108673, 00108672, 00108670, 00108669, 00108668, 00108665, 00108662, 00108661, 00115665, 00108630, 00108627, 00108616, 00108615, 00108613, 00115585, 00115602, 00115673, 00115666, 00108598, 00115667, 00115572, 00108777, 00108778, 00108779, 00108781, 00108696, 00108788, 00108691, 00108693, 00108710, 00108751, 00108758, 00108699, 00108760, 00108763, 00108766, 00108771, 00115658, 00115660, 00115661, 00115662, 00108698, 00108703, 00108701, 00108704, 00115663, 00115634, 00115635, 00115637, 00115639, 00115641, 00115642, 00115651, 00115654, 00115655, 00115657, 00115604, 00115605, 00115608, 00115619, 00115623, 00115616, 00115630, 00115633, 00108650*, 00115624*; 32?**: 00115636, 00108748, 00108705, 00108767, 00108762, 00108757, 00108756, 00108702, 00108785, 00108780, 00115569, 00108597, 00108593, 00108592, 00108589, 00108587, 00108683, 00115589, 00108611, 00108626, 00108674, 00108679, 00108682, 00108731, 00108642, 00108646, 00115584, 00108602, 00108660, 00108722, 00108648, 00108653*, 0115612*); Pichincha Pk. 4km W of Aloag, 3000m, 19-25.vii.1985, carrion tp., pasture at ravine edge, S.&J. Peck (10c?***, 11$*); Zamora- Chinchipe, 8km NW of Zamora, mouth Rio Sabanilla, 1420m, l.xi.1987 (1$). UNITED STATES OF AMERICA: GA: Wilkinson Co., Big Sandy Ck., 8mi S of Irwinton, US441, 5-25.vi.1984, FIT near dung, S. Marshall (lc?*; outside of normal range). VENEZUELA: Arugua, Henri Pittier NP, tr. above Rancho Grande, 1225m, 22- 25.i.l996, malaise tp., J.&A. Skevington (lc?); Maracay, Rancho Grande Bio. Stn., 1250m, 5. iii.1995, sweep along trail, S.A. Marshall (1$); Merida, Call Santa Rosa, 25- 30.iv.1988, malaise, pasture/cloud for. edge, S.A. Marshall (3$); Merida, Jaji to La Azulita, 3.V.1988, sweeps along road, S.A. Marshall (lc?*, 2$); Merida, Los Chorros 23.iv.88 sweeps S.A. Marshall (1$); Merida, Los Chorros, 1200m, 5.V.1988, sweep at trail head, S.A. Marshall (1$); Merida, St. Mucuchies Tr., 5km S of Paramo near Ik., 4000m, 28.iv.1988, under Espeletia debris, S.A. Marshall (1$); Rancho Grande, 12- 30.xii.1987, malaise intercept, M. Sanborne (1$); Trujillo, 10km E of Bocono, Laguna de Lucerdo, 1760m, 3.iii.1995, slash/compost, S. Marshall (13c?, 9$); Trujilla, Bocono, rd. to Guaramacal, 2200m, 2.iii.l995, under dung, S.A. Marshall (3c?, 2$); Trujillo, Guaramacal Nat. Pk., 2000~3000m, 26.viii-l.ix. 1992, car net, L. Masner (2c?: debu00152902, 00152985; 2$: 00152939, 00152342).

Comments:

See comments under Coproica bifurcata sp. nov. and Coproica emarginata sp. nov.

The known range of C. hirtuloidea within South America has been greatly expanded. C. hirtuloidea was previously recorded from Bolivia and Peru, but is now also recognized from Argentina, Brazil, Colombia, Costa Rica, and Venezuela. The single

118 male specimen collected in Georgia (USA) is outside the normal range for this species - it is unlikely that C. hirtuloidea is established in Georgia.

119 Coproica insulaepasqualis Enderlein, 1938.

Distr.: Australasian/Oceanian: Easter I. (= Isla de Pascua). Coprophila ferruginata var. insulae-pasqualis Enderlein, 1938: 678 [sex not given, illustr.]. Type locality: "Oster-Insel" (= Easter Island). STs, sex unknown (?NHRS). Coprophila insulaepaschalis. - Enderlein, 1938: 678 [incorrect original spelling]. Coproica insulaepasqualis. - Marshall, 1989b: 603 [generic combination, Australasian/ Oceanian catalog].

Comments:

Coproica insulaepasqualis was described as a subspecies of C. ferruginata, and was subsequently raised to the species level. This species is likely a junior subjective synonym of C. ferruginata.

Coproica lacteipennis (Hayashi, 2005)

Distr.: Japan. Coproica lacteipennis Hayashi, 2005a: 208 [both sexes, illustr.]. Type locality: Japan, Okinawa Pref., Iriomote, I., Uehara. HT male (NIID).

120 Coproica lugubris (Haliday, 1835) (Figures: 19,24,177-187)

Distr.: Australasian/Oceanian: Papua New Guinea; Oriental: China (HKG), India (Uttar Pradesh), Pakistan, Taiwan; Palaearctic: Afghanistan, Andorra, Austria, Belgium, Bulgaria, Czech Republic, Denmark, Finland, France (incl. Corsica), Germany, Great Britain, Greece (Crete), Hungary, Israel, Italy (incl. Pantelleria I., Sardinia), Japan, Kazakhstan, Kirghizstan, Latvia, Lithuania, Macedonia, Madeira (Portugal), Netherlands, North Korea, Norway, Poland, Romania, Russia (CET, FE, NET, SET), Slovakia, Spain (incl. Balearic Is.), Sweden, Switzerland, Tadjikistan, Tunisia, Yugoslavia (Serbia). Limosina lugubris Haliday, 1836: 332 [both sexes]. Type locality: not given (probably Ireland). STs (NMID). - Laurence 1955a: 189 [biology, preimaginal stages, illustr.]. Limosina (Coprophila) lugubris. - Duda, 1918: 219-222 [subgeneric combination, redescription, taxonomic notes, illustr.]; 1938: 165-166 [redescription].l Leptocera (Coprophila) lugubris. - Duda, 1925: 205 [generic combination]. Leptocera (Heteroptera) lugubris. - Richards, 1930: 307 [subgeneric combination]. Leptocera (Coproica) lugubris. - Richards, 1960c: 202, 207 [subgeneric combination, key]. Coprophila lugubris. - Hammer, 1941: 250-251 [generic combination, egg, illustr.]; Schumann, 1961: 298-299 [egg, illustr.]; Schumann, 1962: 421-422, 438-440 [larva, illustr.]; Hinton, 1981: 747 [egg, illustr.]. Coproica lugubris. - Papp, 1971c: 434 [generic combination]; Papp, 1973c: 28 [key, illustr.]; Papp, 1984: 82 [Palaearctic catalog]; Lachmann, 1990: 163-168 [behaviour, illustr.]; Lachmann, 1991: 198-206 [biology, preimaginal stages, illustr.]; Lachmann, 1996: 759-768 [morphology, male & female genitalia, copulation, illustr.]; Rohacek, 2007b: 104 [new record from Madeira (Portugal)]. Limosina Stenhammari Zetterstedt, 1860: 6400 [both sexes]. Type locality: Sweden, nr. Illstorp. LT male, designated by Rohacek, 1977c: 177 (MZLU). - Rohacek, 1977c: 177 [synonymy]. Limosina Thalhammeri Strobl, 1898: 276 [both sexes]. Type localities: Austria, Admont; Hungary, Kalocsa. STs [probably NMBA]. - Duda, 1918: 219 [synonymy]. Limosina monfalconensis Strobl, 1909: 300 {male]. Type locality: Italy, Monfalcone. ST male [probably NMBA]. - Duda, 1918: 219 [synonymy]. Limosina (Coprophila) lugubris var. cilicrus Duda, 1938: 166 [? male]. Type locality: Russia (SET), Kuban, "Anlegestelle Tiberdii", "Asgen-Bach". HT, ? male (ZISP).

Body length 1.3 - 2.0mm. Body brown. Face and frons brown. Interfrontal bristles in 4 subequal pairs , postocellar bristles convergent, 1 pair of small postocellar setulae. Eye to gena ratio approximately 2:1; 4 genal setae in longitudinal row posterior to vibrissa, anterior seta approximately 1.5 times length of posterior setae. Scutellum with 40-45

121 discal setae evenly distributed over anterior 4/5; 3 lateral setae between larger anterior and posterior marginal scutellar bristles. Katepisteraum with 2 long dorsal bristles, each approximately 1/3 to 1/2 as long as the distance between dorsal margin of katepisternum and wing base, posterior katepisternal bristle slightly longer than anterior bristle. Legs brown, fore coxa brown or light brown. Mid tibia without proximal posterodorsal bristles. Mid basitarsus with small, stout anterior setae at 1/3, 2/3 and 5/6, anteroventral setae at 1/5, 1/2 and 4/5, length of seta at 1/2 approximately 2 times width of mid basitarsus, and 2-3 posteroventral setae between 1/3 and 2/3. Second costal sector 0.7-1.1 times length of third costal sector. Costal bypass short, terminating within 5 vein widths of R4+5. Anal vein with distinct angulate bend.

Male Terminalia:

Sternite 5 rounded rectangular, with strongposteromedial tab with lateral spines and small knobby setae, posterior row of enlarged bristles, and surface bristles larger laterally.

Synsterntie 6+7 straight medially. Epandrium with enlarged pair of dorsolateral bristles; subanal plate present, very narrow; cercus fused to epandrium, ventral margin with pointed medial process, and rounded, heavily bristled lateral process. Surstylus with elongate, quadrate anteroventral process with a patch of short bristles on anterior margin, an elongate, narrow posteroventral process with a toenail-like bristle at its apex, and 2 strong ventral bristles between anterior and posterior processes. Postgonites symmetrical, anteriorly curved, pointed at apex, with subapical dorsal bristle present. Basiphallus straight, tubular, posteroventral margin with elongate, sickle-shaped crest approximately equal in length to rest of basiphallus and open posteriorly dorsal to crest. Tubular sclerite tall, with distinct dorsal medial ridge, and dorsal lateral wing-like processes. Lateral

122 sclerites rounded and elongate, with dorsal margin medially deflected, and a sharp dorsal process at 1/2; Central sclerite fused with lateral sclerites just beyond dorsal process;

additional, faint sclerite ventral to central sclerite and fused at apex and centre of central

sclerite. Membranous sheets broad, covered in soft membranous spines.

Female Terminalia:

Tergite 8 long laterally, reduced to a thin connecting membrane medially. Tergite 10 pollinose, narrowly fused to cerci posterolateral^; cerci pollinose, apical bristles short

and sinuate. Sternite 7 broad, rounded rectangular. Sternite 8 triangular and broad

anteriorly, with cluster of four small sub-apical setae. Sternite 10 not wrapped posteriorly

around tip of abdomen, with blunt process on posterior margin and medial pair of

enlarged bristles. Spermathecae spherical; duct length approximately twice the length of

spermathecal bulb.

Specimens Examined:

???: koiiogs fahl Sihijaea 16 8 20 AL Melander Collection 1961 (1& USNM); Jifeld S- Harz Duda 28 8 9 (1(5"); 6 4 17 St. Wendal Rheinl. Duda (IcT*). AUSTRIA: Austria, Tirol, Kleines Walsertal, Widderstein, ~1800m, 24.ix.1994, old cow pats, M. Buck (4c?*- wing: debu00154167, 00154168, 00154169*, 00154170-wing, 00154171; 1$: 00154172). FRANCE: Besse, Puy de Doma, l.viii.1926, O.W. Richards (IcT, 1 ?); Haras de Jardy, Fr. Amer., 14.x. 1961, McG x Cow Dung, S&O Vaucresson. HUNGARY: South Hungary, Kiskunsagi Nat. Pk. Kunfeherto (lake), 19..viii.l981, leg, on donkey excrement, J. Rohacek (1?*). PAKISTAN: Northwest Frontier Prov., Nathiagali, 27- 30.vi.1988, T. Hayashi; (6c?**-illustr, 6$*-illustr). POLAND: 4 10 K Wustune b. Habelschwerdt I. Duda AL Melander Collection 1961 (1$, USNM). SLOVAKIA: Slovakia cr., Slovensky kras Brzotin nr. Rozneva, 4.ix.l980, leg, on cow excrement, J. Rohacek (2c?**); Slovakia cr., Slovensky kras Turna n. B., l.ix.1980, leg. on cow excrement, J.Rohacek. SWITZERLAND: Canton Luzern, Entlebuch Graben Matteli, 800m,3-4.viii.l974(4$).

123 Coproica microps Papp, 2008

Distr.: Afrotropical: Guinea. Coproica microps Papp, 2008: 22 [female, taxonomic notes, illustr.]. Type Locality: Guinea, Coyah . HT female (HNHM).

124 Coproica mitchelli (Malloch, 1913) (Figures: 25, 188-199)

Distr.: Nearctic: Canada (AB, BC, ON, SK), USA (AR, AZ, CA, FL, ID, KY, MD, MN, MT, NC, ND, NM, OK, OR, PA, SD, TX, UT, VA, WA, WY); Neotropical: Costa Rica, Mexico (COA, JAL). Leptocera mitchelli Malloch, 1913c: 135 [both sexes]. Type locality: USA, Texas, Victoria. HT male (USNM). Leptocera (Coprophila) mitchelli. - Spuler, 1925a: 124 [subgeneric combination]. Leptocera (Coproica) mitchelli. - Richards, 1965c: 725 [subgeneric combination, Nearctic catalog]; Richards, 1967b: 16 [Neotropical catalog].

Body length 1.3 - 2.2mm. Body brown. Face light brown, frons brown. Interfrontal britles in 4 subequal pairs, postocellar bristles erect, with 1 pair of postocellar setulae.

Eye to gena ratio approximately 2.2:1; 4 genal setae in longitudinal row posterior to vibrissa with anterior seta approximately 2 times as long as posterior 3. Scutellum with

20-25 discal setae arranged into 3-4 loose rows between basal 1/5 and basal 3/5; 3 lateral bristles between larger anterior and posterior marginal scutellar bristles. Katepisternum with 2 small subequal bristles, each approximately 1/3 as long as the distance between dorsal margin of katepisternum and wing base. Legs brown, fore coxa brown to light brown. Mid tibia lacking proximal posterodorsal bristles. Mid basitarsus with row of four stout anterior bristles extending from 2/3 to apex, 1 anteroventral sub-basal bristle, and pairs of stout ventral bristles at 2/5 and 2/3. Second costal sector 0.7-1.1 times length of third costal sector. Costal bypass short, terminating within 5 vein widths of R4-1.5. Anal vein with distinct angulate bend.

Male Terminalia:

Sternite 5 rounded rectangular, with pair of subtle posterior lobes, posteromedial patch of dense microsetae, and a posterior row of enlarged bristles; lateral bristles enlarged.

Synsternite 6+7 with slight medial bend. Epandrium with enlarged pair of dorsolateral

125 bristles (though often difficult to distinguish from other epandrial bristles due to heavy chaetotaxy), bristles denser ventrally; subanal plate present, narrow; cerci fused toepandrium, ventral margin with pointed medial process and lateral process, lateral process heavily bristled and with distinct constriction at base. Surstylus deflected medially, with pointed anterior process, posteroventral process with thickened toenail- like bristle at apex, 2 ventral lobes, 4-5 bristles along on lobes, and 2 additional ventral bristles between posterior lobe and posteroventral process. Postgonites symmetrical, straight, broad, flattened at apex, and with subapical anterior bristle. Basiphallus straight, tubular, with short, blade-like posteroventral process, and open posteriorly. Tubular sclerite with small lateral processes. Lateral sclerites rounded, with membranous process at proximal dorsal angle, 2 broad, rounded dorsal processes, and medially deflected dorsal margin. Central sclerite not fused with lateral sclerites; additional sclerite, ventral to central sclerite, fused with central sclerite apically and in centre. Ventral most sclerite extended. Apical membranes reduced.

Female Terminalia:

Tergite 8 broad, almost connecting ventrally, long laterally, short dorsally. Tergite 10 pollinose, narrowly fused to cerci posterolaterally; cerci pollinose, apical bristles short and sinuate. Sternite 7 broad, rectangular. Sternite 8 elongate with posterior notch, and 4 small posterior setae. Sternite 10 not wrapped posteriorly around tip of abdomen, with blunt posterior point, and pair of enlarged medial bristles. Spermathecae spherical; duct length 2-3 times length of spermathecal bulb.

Specimens Examined:

CANADA: AB: Cypress Hills, 14.vi.1980, cow dung, S. Marshall (23& 28$); Lower Kananaska Hwy., 5.viii.l980, cow dung, S.A. Marshall (1$, 1$); Wagner Bog, 26.vi.-

126 2.vii.l985, pan tp. 2, Leg, A.T. Finnamore (1 $, PMAE); same as previous except pan tp. 5, leg (2$, PMAE); pan tp. 7, leg (1& PMAE); pan tp. 10, leg (1& PMAE); pan tp. 3, leg (1 $, PMAE); Writing-On-Stone Prov. Pk., 30.viii-10.ix.1990, malaise tp., M. Klassan (1 $); same as previous except birch N. (1(5*). BC: Ainsworth, Woodbury Ck., 5.vii.l980, dung, S.A. Marshall (1$*); Cranbrook, l.viii.1980, horse dung, S. Marshall (1$); Osoyoos, Mt. Kobau, 560m, 24-28.viii.1991, D. Blades & C. Maier, SOCAP - LMS (7c?, 13$, BCPM). ON: Aberfoyle, 25.vi.1956, D.H. Pengelly (IcT); Alfred, lO.v.1981, bog, moose dung, Peck & Anderson (2$); Alfred, Alfred Bog, 25.ix.1983, sifted moss under fungi & moose dung, Berl, S. Peck (IcT); Bruce Co., Inverhuron Prov. Pk., front dunes, 44 17'33"N, 81 35'28"W, 22.viii.2003, yellow pans, M. Buck (1$: debu00236605); Chaffeys Locks, Queens Univ. Bio. Stn., 12.ix.1980, human dung, field, S.A. Marshall (IcT, 4$); Fergus, 23-30.vii.1992, black composter, S.A. Marshall (IcT); Glen Huron, see page spring, 19.vi.1985, B. Sinclair (IcT); Grenville Co., 2km SW of Bishops Mills, 2.vi.l989, B. Gill & Cook (IcT); 7mi E of Griffith, 19.vii.1990, B.E. Cooper (2$); Guelph, 12-15.viii.1993, black composter, D.C. Caloren (1$); Guelph, 24.vi-14.vii.1981, malaise tp., K.N. Barber (1$); Guelph, 27.iv-21.v.l980, pan tp., K.N. Barber (IcT); Guelph, 28.V.1980, carrion, S.A. Marshall (1$); Guelph, Arboretum, 9.viii.l984, field, mushroom tps., B.V. Brown (IcT); Hilton Beach, 6.v-10.vi.l989, red oak forest pan tps., pig dung, J.E. Swann (IcT); same as previous except forest pan tps., pig dung (IcT); cedar swamp pan tps., pig dung (IcT); Kent Co., Rondeau Prov. Pk., South Point Trail, near, east parking lot, 42 15'42"N, 81 50'49"W, 9-10.viii.2003, forest clearing, yellow pans, M. Buck (lcT*-illustr: debu01139538*-illustr); Mer Bleue, 21-26.vii.1987, intercept tp., L. Dumouchel & J. Denis (IcT); Norval, 16.V.1980, carrion, S.A. Marshall (ltf*, 2$); Ottawa, 2.ix.l984, marshy shore of Ottawa River, J.R. Vockeroth (1$, CNCI); Ottawa, viii.1979, grass piles, A. Telka (2c?*); Ottawa, 17.vii.1979, S.A. Marshall (IcT); same as previous except dry cow pat (1$); Prescott, 30.xi.1977, K. Barber (IcT); Simcoe Co., Washago, 44 44.76"N, 85 18.73"W, 12-19.iv.1998, pan4b, carrion tp., C. Christopherson (1 $); Wainfleet Bog, 8km S of Welland, 3-8.viii.1988, pt 2 -1962 zone, N2G, A. Stirling (IcT). SK: Grasslands Nat. Pk., 20.vii.1997, P2, S.A. Marshall (2c?, 1$); Grasslands Nat. Pk., Val Marie, E329782 N5451113, 10.vii.1996, pitfall tp. (2$); Grasslands Nat. Pk., Val Marie; 22.vii.1997, prairie dog RET, Marshall & Finnamore, P4RBT1 (IcT). MEXICO: Coahuila, Boquillas, 24-26.ix.1979, pig dung tp., R.E. Woodruff & G. Fincher (7c?*, 10$); Jalisco, 18mi W of Atenquique, 9300', fir for., dung, A. Newton (1 $). UNITED STATES OF AMERICA: AR: Garland Co., 1.2mi N of Crystal Springs Hwy. 270, 6-8.iii.1977, pig dung tp., Woodruff & Wiley (2c?, 10$); Logan Co., Ozark Nat. For., Magazine Mt., 23.v-8.vi.1991, pans, mushroom baited. J.E. Swann (1$); Scott Co., 7mi E Y City, Jet. Hwy. 270 & Rt. 71 on 270, 6-8.iii.1977, pig dung tp., Woodruff & Wiley (16<^, 22$); Wash Co., 3mi S of Devils Den St. Pk., 28-31.V.1979, oak-hickory, S.&J. Peck (IcT); same as previous except 10 mi S of Devils Den St. Pk. (1 $). AZ: lOmi NW of Flagstaff, San Francisco Mts., 9500', 18-24.vii.1979, spruce-fir-aspen meadow malaise, S.&J. Peck (4c?, 1$); Apache Co., Alpine, Luna Lake, 7900\ 9-14.vii.1979, pine meadows, S.&J. Peck (IcT: debu00104152; 8c?, 3$); Cochise Co., Chiracuhua Mts., 7000', 15-2l.vii. 1978, dung, O. Kukal (3c?, 2$); same as previous except Turkey Ck. (IcT); Martyr cpgd, 15-20.vii.1978 (IcT); Cochise Co., Chiricahua Mts., 6mi ESE of Portal, 4150', l.ix.1970, desert grass, human dung, A. Newton (lc?); Cochise Co., Chiracahua Mts. 15-2l.viii. 1978, dung, O. Kukal (lc?); same as previous except E of

127 Turkey Ck., 6500', dung tps. (3c?, 2$); Cochise Co., 5100', Coronado NF., Chiricahau Mt., Stewart cpgd., 16-18.viii.1984 road dung, L.B. Carlson (5c?, 2$); Cochise Co., Coronado Nat. For., Coronado Nat. Mem., ll-15.viii.1984, carrion, L.B. Carlson (43)', Cochise Co., Coronado Nat. Mem., ll-15.viii.1984, FIT, B.V. Brown (1$); same as previous except dung tp. (203, 24$); Cochise Co., Huachuca Mts., Carr Canyon, 7100' ix.1970, oak & evergreen, human dung, A. Newton (23, 2$); Cochise Co., Huachuca Mts., Miller Canyon, 6000', ix.1970, oak woodland, dung, A. Newton (13)', Cochise Co., Huachuca Mts., Ramsey Canyon, 5600', ix.1970, human dung, A. Newton (13)', Cochise Co., Portal, 5.2mi W, Southwest Res. Stn., 1650m 31°52'58N, 109°12'20W, malaise, 5 Jul 1995, S.D. Gaimari, debu00200276; 1 3 Cochise Co., Portal, SWRS., Chiricahua Mts., 18-23.viii.1984, FIT, B.V. Brown (43, 3$); same as previous except dung tp. (13, 2$); mushroom tp.. (3$); intercept tp.. (1$); yellow pan (2$); Coconino Co., Coconino Nat. For., Bonito PL, llmi N of Flagstaff, 7000', 7-8.viii 1984, mt. meadow, carrion, L.B. Carlson (\3, 1$); Coconino Co., Coconino Nat. For., Bonito PL, 8-25.viii.1984, FIT, B.V. Brown (3c?, 5$); Coconino Co., Flagstaff, 7100', 18-25.vii.1979, pond pine meadow malaise, S.&J. Peck (\3, 19); Coronado Co., Bonito PL, near Flagstaff, 5- 9.viii.l984, dung tp.., B.V. Brown (2c?); Flagstaff Oak, "ok. Can.,"(?) 5900', 17- 25.vii.1979, S.&J. Peck (43, 1$); same as previous except near Sterling Canyon, 3900\ riparian woods (23, 1$); Graham Co., 2.4mi W on Hwy 336 from Hwy 191, 3800', 19- 26.viii.1993, J.B. O'Hara (\3, 1?); Navajo Co., 15mi SE of Holbrook, 5300', 14- 16.vii.1979, grassland carrion tps., S.&J. Peck (lc?); Santa Cruz Co., 7mi WNW Nogales, Calabasas Canyon, 3850', ix.1972, desert grass, dung, A. Newton (33*, 1 $); Santa Cruz Co., 8mi NNW Nogales, Walker Canyon, 3900', ix.1970, dung, oak woodland, A.Newton (43); Santa Cruz Co., Madera Canyon, Madera picnic ground, 12.vi.1992, malaise river edge, J. Skevington & A. Goering (lc?); Santa Cruz Co., Patagonia Lk. St. PL, 9-1 l.viii. 1984, dung tp., B.V. Brown (1$, 3$); Santa Cruz Co., Santa Rita Mts., Madera Canyon, 5500', ix.1970, oak, human dung, A. Newton (3c?, 6$). CA: Kent Co., Mt. Pinos, 8000', 3.vi.l992, sweep, J. Skevington & A. Goering (1$); Santa Barabara Co., Santa Cruz Is., Christy's Beach & Canyon, 19-22.iii.1982, pan tps., J.T. Huber (3c?- wing, 4$, CNCI); Santa Clara Co., Del Peurto Rd. (Rd 130), 28.V.1992, J. Skevington (1$); same as previous except pan tps., J. Skevington & A. Goering (1$). FL: Alachua Co., Gainesville, 2-5.vii.1982, pig dung, horse pasture, K.L. Rench (2$); Gainesville, UF Beef Farm, 13.vii.1991, G.Y. Hu, 1A (lc?, 2$); Highlands Hammock St. PL, 15- 21.xii.1985, cypress swamp dung pan, S.A. Marshall (1$); Marion Co., Rd65, Ocala Nat. For., 18-23.vi.1984, sand-pine, human dung tp., S.A. Marshall (1$); Marion Co., Sand trail to Oklawaha Swamp, 10-15.vi.1984, mushroom tp., S.A. Marshall (\3); Marion Co., Zay Prairie, Ocala Nat. For., 14-18.vi.1984, human dung tp., S.A. Marshall (23, 2$); same as previous except 14-19.vi.1984, pan tp. (lc?*); 10-14.vi.1984, mushroom tps. by shore, (2$). ID: Custer Co., 25mi W of Stanley, 9-17.viii. 1990, pans in pine forest human dung, J.E. Swann (2c?); Kootenai Co., Wolf Lodge Bay, KOA, 12.5km E of Coeur d'Alene, 29.viii.1981, P.H. Arnaud Jr. (1$, CASC); Nez Perce Co., Top of Lewiston Grade, 30.vi.1975, Edward Rogers (1$, CASC). KY: Edmonson Co., Mammoth Cave Nat. PL, 15.v-20.viii.1983, mesic for., FIT, S.&J. Peck (1$); Rowan Co., 24km SW of Morehead, Cave Run Lk., 14.v-20.vii.1983, Fagus forest, FIT, S.&J. Peck (43); MD: Ocean City, 9.V.1991, seashore, S.A. Marshall (1$). MN: Clearwater Co., Itasca St. PL, 28-29.viii.1979, carrion, deciduous forest, S.&J. Peck (1$). MT:

128 Carbon PL, Beartooth Mts., Rt.212, 10000', 18-25.viii.1979, carrion, S. Peck (1$); Carbon-Pk. Cos, Beartooth Mts., Rt. 212, 10900', 18-25.viii.1979, carrion, tundra, S.&J. Peck (1$); Ravalli Co., Bitterroot Nat. For., Sam Billings Mem. Cpgd., 8-17.viii.1990, pans along stream human dung, J.E. Swann (1$). NM: Bernalillo Co., 7mi S of Tijerus, 2400', 23-26.viii.1975, carrion tp., Pinyon-juniper, S. Peck (2c?, 4$); Catron Co., 5mi W Luna, San Francisco Riv. pond, 7400', 9-14.vii.1979, pine-meadows, S.&J. Peck (4$, 5$); Catron Co., 6mi S Lupa, 8000', 9-14.vii.1979, pond, pine forest, S.&J. Peck (1$); Dona Ana Co., Jornada Expm. Range, 12.x.1984 reared from cow manure, 7-Manure, 8- 10, J. Kinzer (lc?); Eddy Co., 30mi WSW of Carlsbad, Sitting Bull Falls, 4600', 23- 27.vii.1975, dung, S. Peck, 15 {2S, 2$); Guadalupe Co., Santa Rosa Lk. St. Pk., 3- 4.viii.l984, dung tp., B.V. Brown (3c?, 1$); Lincoln Co., lmi N of Sierra Blanca Ski, 11000', 10-26.vi.1979, alpine grass meadow, carrion, S.&J. Peck (20c?*, 16$); Mora Co., W of Ocale Lefebros, Lyn. Ca., 7500', 36°12'N, 105°07'W, 10-1 l.ix. 1994, J.B. O'Hara (lc?); Santa Fe Co., Santa Fe Baldy, 10600-11800', 19.vi-4.vii.1979, spruce-fir- tundra, S.&J. Peck (2d1, 2$*); Socorro Co., 20mi W of Socorro, Water Canyon, 7000', 28.vi-7.vii. 1979 mixed "mesic" forest, S.&J. Peck (1$); Torrance Co., 6mi SW of Manzano, Red Can. cpgd, 75-9000', 17-30.vi.1979, spruce-fir litter, S.&J. Peck (7$*, 4$); Torrence Co., Manzano Mts., 12mi W of Manzano, Red Canyon cpgd., 8000', 27- 30.viii.1993, malaise, J.B. O'Hara (lc?*-illustr, 2$*-illustr). NC: Bladen Co., Singletary Lk. St. Pk., 34°35'N, 78°27'3W, 19-22.V.2003, malaise, Marshall & Paiero (1?: debu00227308). ND: Billings Co., Medora, Roosevelt Nat. Pk., 27.viii.1979, sage grassland, carrion, S.&J. Peck (lc?, 7$); McHenry Co., W side Buffalo Lodge Lk., 28 mi N of Granville 14-28.ix.1971, carrion tp., shoreline, P.D. Tomlin (1$); Roosevelt Nat. Pk., 28.viii.1979, carrion (1$). OK: Wichita Mts., 5-7.vi.1979, dung tps., (lc?, 1$); Wichita Mts., Refuge, 5-7.vi.1979, shortgrass prairie, S.&J. Peck (2c?, 1$); Lattimer Co., 5mi W, 15-21.V.1977, red oak, dung tp., K. Stephen (1$). SD: Custer Co., Wind Cave Nat. Pk., 27.vi.1957, bison dung, H.S. Dybas (lc?, 2$, 1?). TX: Angelina Co., Angelina Nat. For., Boykin Cemetery, 13-17.vi.1993, spring fen, dung, S.A. Marshall (1$: debuOO 128160); Brewster Co., 3.5mi E of Marathon, 26-27.ix.1979, dung tp., R. Woodruff & G. Fincher (3c?); Kimble Co., Llano Riv. at Co. rd. 385, l-2.iv.2000, floodplain for., pig dung pitfalls, E.G. Riley (1$: debuOO 110637); Medina Co., 8mi E of Sabinal, Rt.90, 22-28.ix.1979, dung tp., R. Woodruff & G. Fincher (lc?); Randall Co., Paio Duro Canyon St. Pk., 26.v-7.vi.1991, pan tps., J.E. Swann (lc?); Terrell Co., 30mi E of Sanderson, 23-27.ix.1979, on pig dung, B. Woodruff & G. Fincher (3c?); Val Verde Co., Baker's Crossing, 20.5mi N of Comstock, 2.iv.2000, on/near dung, M. Buck (lc?*: debuOOl 10221*; 1$: debuOO 110220). UT: Daggett Co., Dutch John, 6300', 2- 8.viii.l979, pinyon juniper, woodland, carrion, S.&J. Peck (3$); Duchesne Co., Mirror Lk., 10300', 30.vii-12.viii. 1979, spruce fir meadow, carrion-malaise, S.&J. Peck (lc?); Summit Co., NW side of Gilbert Peak, 11500-12500', l-10.viii.1979, tundra carrion, S.&J. Peck (lc?, 1$). VA: Giles Co., vie Mt Lk. Biol. Stn., 14.V.1998, Bear Cliff Tr. sweep, S.A. Marshall (1?). WY: Big Horn Co., 30mi E of Lovell, 9000', 18- 26.viii.1979, spruce fir carrion, S.&J. Peck (lc?, 3$); same as previous except 34mi E of Lovell, 9400', meadow edge, malaise (2c?*); lOmi E of Lovell, 3800', sagegrass-barren, carrion (lc?, 1?); Crook Co., Black Hills Nat. For. Cook Lk. Rec. Area Rd., 2.5km E of cpgd, 4-2l.viii. 1990, pan tp., cow dung, logged pine area, J.E. Swann (2$); same as previous except pans, cow dung, oak/hawthorn scrub (1 $); pan tps., cow dung, scrub oak

129 (lc?, 59); Crook Co., Black Hills Nat. For. Cook Lk. Rec. Area, 4-2l.viii. 1990, FIT on poplar ridge, J.E. Swann (2c?, 1$); Fremont-Teton Co., Togwotee Pass, 9600', 16- 24.viii.1979, carrion meadow, malaise, S.&J. Peck (1$); Red Gulch Rd., off Hwy. 14, 5- 19.viii.1990, near shell pans in sagebrush, in desert area, cow dung, J.E. Swann (59c?*, 699); same as previous except near shell pans in grass hummock, in desert area, cow dung (10c?, 17$); near shell pan tps. in barren, in area near cotton woods, cow dung, (3c?*, 7$); near shell pan tps. in barren area, near cotton woods, (9c?, 139); near sneU pans in barren area near cottonwoods (21c?, 23c?); near shell pans in desert area, cow dung (12c?, 159); Sheridan Co., 20.3km W of Burgess Jet., Antelope Butte Rec. Area, 5- 20.viii.1990, pans along stream cow manure, J.E. Swann (4c?, 29); Sheridan Co., Black Mt., 5-20.viii.1990, pine for. pans, cow dung, J.E. Swann (5c?, 139); Sheridan Co., Black Mt, off Hwy 14, 5-20.viii.1990, pine forest pans cow dung, J.E. Swann (4c?*, 199); Sublette Co., 7mi N of Pinedale, 9000', 16-24.viii.1979, pine meadow, carrion malaise, S.&J. Peck (19); Uinta Co., 6mi E of Mt. View, 6800', 1-1 l.viii. 1979, sagebrush, S.&J. Peck (2c?, 39); Unita Co., 8mi SE of Evanston, 7100', 30.vii.-ll.viii. 1979, carrion, sage-grass, riparian, S.&J. Peck (lc?, 19); Wind Riv. Mts., 20mi NE of Pinedale near Nelson Lake, 10400-11000', 15-23.viii.1979, carrion tundra, S. Peck (19).

130 Coproica novacula sp. nov. (Figures: 10,26,200-210)

Distr.: Nearctic: USA (AZ, TX); Neotropical: Argentina, Bolivia, Brazil, Costa Rica, Ecuador, Guatemala, Honduras, Mexico (MEX, MOR, OAX).

Body length 1.1 - 1.8mm. Body colour brown. Face and frons brown. Interfrontal bristles in 4 subequal pairs, postocellar bristles erect, one pair of small postocellar setulae. Eye to gena ratio approximately 2:1; 4 genal setae in longitudinal row posterior to vibrissa, anterior bristle 2-3 times longer than posterior 3. Scutellum with approximately 18 discal setae arranged in 2 loose rows between basal 1/4 and basal 1/2; 2 lateral bristles between larger anterior and posterior marginal scutellar bristles.

Katepisternum with 2 dorsal bristles, anterior and posterior bristles each approximately

1/7 and 2/3 as long as the distance between dorsal margin of katepisternum and wing base respectively. Legs brown, fore coxa brown to light brown. Mid tibia with 1 proximal posterodorsal bristle between 2/5 ando 1/2. Mid basitarsus with row of stout anterior bristles extending from 2/3 to apex, a pair of bold sub-basal bristles, and three small pairs of bristles between 2/5 and 4/5. Second costal sector 0.7-0.9 times length of third costal sector. Costal bypass long, extending 7 or more vein widths beyond R4+5. Anal vein with distinct angulate bend.

Male Terminalia:

Sternite 5 rounded rectangular, with paired posteromedial tabs, a posteromedial patch of dense microsetae; surface bristles uniform in size. Synsternite 6+7 with slight bend posteromedially. Epandrium with enlarged pair of dorsolateral bristles; subanal plate present, narrow; cerci fused to epandrium, ventral margin relatively flat and straight, with

1 large bristle in centre. Surstylus elongate, triangular, pointed posteroventrally, and

131 sharp point at apex, with row of small anteroventral setae and elongate posterodorsal bristle. Postgonites symmetrical, straight, and narrow. Basiphallus straight, tubular, with rounded posteroventral crest, open posteriorly dorsal to crest. Lateral sclerite deflected medially, with plateau-like process at proximal dorsal angle, 2 small sharp dorsal processes, apex cup-shaped and shallow, and subtly ribbed along apical dorsal margin.

Central sclerite fused to lateral sclerites ventral to proximal process and with sharp apical process.

Female Tertninalia:

Tergite 8 long laterally and short dorsally. Tergite 10 pollinose, narrowly fused with cerci posterolaterally; cerci rounded and triangular, apical setae long and sinuate.

Sternite 7 broad and rectangular, with posterior margin concave. Sternite 8 broad, with rounded posterior margin, and posterior row of 4 small setae. Tergite 10 not wrapped posteriorly around tip of abdomen, with pointed posterior margin and medial pair of elongate bristles. Spermathecae spherical; duct length approximately 4 times length of spermathecal bulb.

Type material:

HOLOTYPE: COSTA RICA: Cartago, Rio Grande de Orosi, near Tapanti Nat. Pk., 1100-1150 m, 9.X.1999, floodplain and forest, Marshall and Buck (1$: debu00103948, INBC).

PARATYPES: ARGENTINA: Salta, El Ray Nat. Pk., 900m, Rio La Salsa, 5- 10.xii.1987, FIT, malaise tp., open stream side in forest, S.&J. Peck (l<$, 1$); Salta, El Rey Nat. Pk., 900m, Aguas Negras Tr., ll-15.xii.1987, forest, malaise tp., FIT, S.&J. Peck (1$); Salta, El Rey Nat. Pk., 1000m, POZO, Verde Tr., km 7, 5-15.xii.1987, malaise, FIT, yungas forest, S.&J. Peck (1$); Salta Rosario de Lerma, INESALT yard, 16-28.ii.1992, malaise tp., S.A. Marshall (1$); Santa Fe, 26km S of Reconquista, Rt.ll, 2km NE Berna, 20-21.ii.1980, pig dung tps., R.E. Woodruff (2$). BOLIVIA: Santa Cruz, Acite Cuevo, 12km W of Boyuibe, 890m, 18.viii-2.ix.2000, malaise tp., Irwin & Hauser (l<$: debu00147387, UASC); Santa Cruz, Campo Guairuy, 24km S of Camiri, 870m, 18.viii-2.ix.2000, malaise tp., Irwin & Hauser (1$*: debu00143116*; 2$:

132 00144564, 00144119, UASC). BRAZIL: Rio Grande do Sul, Pelotas, vii.1998, C.L. Bicho (lcJ*: debu00105615*, MZSP). COSTA RICA: Cartago, Purisil, -1200 m, lO.x.1999, sweeping coffee waste, S.A. Marshall (43*: debu00127435, 00127436, 00127437, 00127438*, INBC); Cartago, Rio Macho For. Res., 2200m, 10.x. 1999, sweeping, S.A. Marshall (4c?: debu00103883, 00103884, 00103879, 00103876; 3$: 00103844, 00103875, 00103897, INBC); Heredia, Sto. Domingo, INBio Pk., 9°58'23N, 84°5'30W, 18-19.viii.2001, on compost, Buck & Barber (1$: debu00187432, INBC). ECUADOR: 12-23.iii.1992, S. Peck (1& QCAZ). GUATEMALA: Fraijanes, Finca San Antonio, 1800m, 14.ii.1987, J. Manger (ltf: debu00263008); Quetzalten, Ango Prov., 8km SE of Zumil, 2450m, 19-21.vi.1993 #069, ex FIT, tp.#l, J. Ashe & R. Brooks (1

Etymology:

The specific name is derived from the latin word novacula which means "sharp knife," and refers to the blade-like appearance of the surstyli.

Comments:

Coproica novacula is known from relatively few specimens, all from South and

Central America, and southwest United States. A unique mid basitarsal chaetotaxy and surstylar shape each make this species easily recognizable.

133 Coproica pappi Carles-Tolra, 1990 (Figures: 211-221)

Distr.: Palaearctic: Hungary, Spain. Coproica pappi Carles-Tolra, 1990a: 35 [both sexes, phylogenetic notes, illustr.]. Type locality: Spain, Province Gerona, Mieres. HT male (CTB). Coproica dentata. - Papp, 1973a: 382 [misidentiflcation of Hungarian specimens, see Papp, 1999: 536].

Body colour dark brown. Face and frons brown. Interfrontal bristles in 4 subequal pairs, postocellar bristles erect, 1 pair of small postocellar serulae. Eye to gena ratio 2.0:1 -

2.7:1; 4 genal setae in longitudinal row posterior to vibrissa. Scutellum with approximately 18 setae arranged in 3 or more rows between basal 1/4 and basal 2/3; 2 lateral bristles between larger anterior and posterior marginal scutellar bristles.

Katepisternum with 2 small, subequal dorsal bristles, each approximately 1/6 as long as the distance between dorsal margin of katepisternum and wing base. Legs and fore coxa brown. Mid tibia with proximal posterodorsal bristle at 1/2. Mid basitarsus with stout anterior bristles sub-apically and at 1/2, 1 sub-basal anteroventral bristle, 4-6 small scattered ventral bristles on distal 1/2 with variable arrangement. Second costal sector

0.8-0.9 times length of third costal sector. Costal bypass short, terminating within 5 vein widths of R4+5. Anal vein with angulate bend.

Male Terminalia:

Sternite 5 crescent-shaped, with rounded anterior cornersposteromedial patch of dense microsetae, and posterior row of enlarged bristles; surface bristles enlarged laterally.

Synsternite 6+7 with a distinct posterior bulge medially. Epandrium with enlarged pair of dorsolateral bristles; Subanal plate present and narrow, with small dorsal process; cerci fused to epandrium, ventral margin strongly modified, with small sub-apical medial

134 processes with 2 small posterior setae, enlarged medial process ventral to sub-apical process, enlarged medial process with distinct sub-apical angle, and 2 posterior setae.

Surstylus with anterior point, 2-3 thickened ventral bristles, dorsally curved posterior process with basal constriction and medially deflected toenail-like bristle at apex.

Postgonites symmetrical, curved posteriorly, and flattened and shovel-like at apex.

Basiphallus straight, elongate, tubular, and closed posteriorly with rounded posteroventral crest. Lateral sclerites rounded, with 2 dorsal medially deflected processes; central sclerite fused with lateral sclerites ventral to dorsal processes, with a pair of rounded apical lobes; ventral surface of distiphallus slightly bulbous. Membranous sheets narrow.

Female Terminalia:

Tergite 8 long laterally, and short dorsally. Tergite 10 pollinose, narrowly fused to cerci posterolaterally. Sternite 7 broad and rounded rectangular. Sternite 8 rounded posteriorly and square anteriorly, with 2 posterior rows of small setae, and pair of enlarged posterior bristles. Sternite 10 not wrapped around posterior tip of abdomen, with wavy posterior margin, and pair of enlarged medial bristles. Spermathecae spherical; duct length approximately equal in length with spermathecal bulb.

Specimens Examined:

HUNGARY: Kiskunsag N.P. Bugac, 18.ix.1980, on horse excrement, leg, L. Papp (3(5l***-illustr-wing, l$-illustr).

Comments:

The structure referred to as the "postgonite" in Papp, 2008 is in fact the male cercus. See comments under Coproica dentata Papp, 1979.

135 Coproica perlugubris Papp, 2008

Distr.: Afrotropical: Ethiopia, South Africa. Coproica perlugubris Papp, 2008: 37 [both sexes, taxonomic notes, illustr.]. Type Locality: Republic of South Africa. Eastern Cape Pr., Shamwari Game Reserve. HT male (HNHM).

Coproica pseudolacteipennis Papp, 2008

Distr.: Afrotropical: South Africa. Coproica pseudolacteipennis Papp, 2008: 24 [male, taxonomic notes, illustr.]. Type locality: Republic of South Africa, Eastern Cape Pr., Shamwari Game Reserve. HT male (HNHM).

136 Coproica pusio (Zetterstedt, 1847) (Figures: 222-232)

Distr.: Afrotropical: all records are doubtful; Oriental: Pakistan; Palaearctic: Afghanistan, Andorra, Austria, Belgium, Bulgaria, Czech Republic, Denmark, Finland, Germany, Great Britain, Hungary, Japan, Italy, Latvia, Norway, Mongolia, Russia (NET, ES, SET), Slovakia, Spain, Sweden, Switzerland, Yugoslavia (Serbia). Limosinapusio Zetterstedt, 1847: 2496 [female]. Type locality: Sweden, Esperod. LT male, designated by Rohacek, 1983e: 159 (MZLU). Coproica pusio. - Rohacek, 1983e: 159 [generic combination]; Papp, 1984: 82 [Palaearctic catalog]. Limosina (Coprophila)pseudolugubris Duda, 1924b: 179 [both sexes]. Type locality: Hungary, Gyon. LT female, designated by Papp, 1979a: 103 (HNHM). - Rohacek, 1983e: 159 [synonymy]. Leptocera (Coprophila)pseudolugubris. - Duda, 1925: 206 [generic combination]; 1938: 166-167 [redescription]. Leptocera (Heteroptera) pseudolugubris. - Richards, 1930: 308 [subgeneric combination]. Leptocera (Coproica) pseudolugubris. - Richards, 1960c: 202, 207 [subgeneric combination]; Okely, 1974: 55-56 [puparium, illustr.]; Richards, 1980: 618 [Afrotropical catalog]. Coproica pseudolugubris. - Hackman, 1965a: 502 [generic combination].

Body length 1.3 - 1.9mm. Body colour brown or light brown. Face and frons light brown to brown. Interfrontal bristles in 4-5 subequal pairs, postocellar bristles convergent, postocellar setulae absent. Gena either entirely yellow or only on anterior

1/2; eye to gena ratio 2:1 to 2.2:1; 4 genal setae in longitudinal row posterior to vibrissa, anterior seta longer than posterior 3. Scutellum with approximately 20 discal setae between basal 1/5 and basal 3/5; 3 lateral bristles between larger anterior and posterior marginal scutellar bristles. Katepisternum with 2 small, subequal dorsal bristles, each approximately 1/4 as long as the distance between dorsal margin of katepisternum and wing base. Legs and fore coxa light brown to brown. Mid tibia without proximal posterodorsal bristles. Mid basitarsus with row of 3 small, stouut anterior bristles along distal 1/2, 1 sub-basal anteroventral bristle , and pairs of ventral bristles at 1/3 and 2/3.

137 Second costal sector 0.8-1.0 times length of third costal sector. Costal bypass short, terminating within 5 vein widths of R4-1-5. Anal vein with distinct angulate bend.

Male Terminalia:

Steraite 5 rounded, with posteromedial patch of dense microsetae medially, and posterior row of enlarged bristles; surface bristles enlarged laterally. Synsternite 6+7 with slight medial bend. Epandrium with enlarged dorsolateral bristles absent or indistinguishable from other epandrial bristles, bristles denser ventrally; subanal plate present, narrow; cerci fused to epandrium, ventral margin with pointed medial process and lateral process, lateral process with distinct constriction at base, deflected medially, and heavily bristled.

Surstylus with pointed anterior process, a medially deflected posteroventral process with thickened apical toenail-like bristle, 2 ventral wavy lobes, posterior lobe approximately 2 times size of anterior lobe, 8-10 bristles on ventral lobes, and 2 bristles set within the invagination between posterior lobe and posteroventral process. Postgonites symmetrical, anteriorly curved, broad at apex, and with subapical dorsal bristle.

Basiphallus elongate, straight, tubular, with small posteroventral crest, and open posteriorly dorsal to crest. Lateral sclerites rounded and elongate, dorsal margin deflected medially, with single sharp dorsal process; central sclerite with apical process; an additional, faint sclerite ventral to central sclerite fused with central sclerite apically and centrally. Apical membranes reduced.

Female Terminalia:

Tergite 8 long laterally, shorter dorsally. Tergite 10 pollinose, narrowly fused with cerci posterolaterally; cerci rounded and pollinose, apical bristles long and sinuate. Sternite 7

138 broad, rounded rectangular. Sternite 8 elongate and narrow, with posterior row of 4 small setae. Sternite 10 not wrapped around posterior tip of abdomen, anterior and posterior margin with broad rounded lobes, with pair of enlarged medial bristles. Spermathecae pear-shaped; duct 1-2 times length of spermathecal bulb.

Specimens examined:

???: 16 8 20 Kimigati (ltf); 4 10 K; AL Melander Collection, 1961 (1$, USNM). GERMANY: Bavaria, Distr. Neu-Ulm, Senden Hittistetten, 1.5km E, 487m, 48 20'0N, 10 7'0E, 12-19.viii.1993, arable field carrion tp., M. Buck (2

139 Coproica rohaceki Carles-Tolra, 1990 (Figures: 233-242)

Distr.: Neotropical: Argentina, Brazil, Chile; Palaearctic: Canary Is. (Spain), Italy, Japan, Malta, Norway, Spain (incl. Balearic Is.). Coproica rohaceki Carles-Tolra, 1990a: 37 [both sexes, phylogenetic notes, illustr.]. Type locality: Spain, Province Barcelona, Cabrils. HT male (CTB). - Hayashi, 1995a: 230-232 [diagnosis, key, illustr.]. Coproica rohaceki Carles-Tolra, 1986: 32 [nomen nudum].

Body length 1.6 - 2.3mm. Body dark brown, ventral portion of thorax orange-brown.

Face and frons dark brown. Interfrontal bristles in 4 subequal pairs, postocellar bristles slightly convergent, with 2 pairs of small postocellar setulae. Gena brown with orange tinge; eye to gena ratio 1.8:1 - 2.0:1; patch of 6 small genal setae, anterior most seta 2-3 times length of posterior setae. Scutellum with approximately 25 discal setae between base and basal 3/4, setae denser laterally, with additional strong pair of medial setae along posterior row, approximately 1.5 times length of other discal setae; 5 lateral bristles between larger anterior and posterior marginal scutellar setae. Katepisternum with 2 dorsal bristles, anterior and posterior bristles each approximately 1/2 and 1/3 as long as the distance between dorsal margin of katepisternum and wing base respectively. Legs brown to light brown, fore coxa light brown. Mid tibia with strong proximal posterdorsal bristles at 1/4 and 1/2. Mid basitarsus with stout anterior bristles sub-basally and at 1/2 and 3/4, 2 sub-basal anteroventral bristles with proximal bristle shorter, a long posteroventral sub-basal bristle, and 2-4 small anteroventral and posteroventral bristles on distal half in variable arrangement. Second costal sector 1.0-1.5 times length of third costal sector. Costal bypass short, terminating within 5 vein widths of Pv4+5. Anal vein with distinct angulate bend.

140 Male Terminalia:

Sternite 5 rounded rectangular, posterior margin with a small medial process and posteromedial patch of microsetae; all surface bristles uniform in size. Synsternite 6+7 straight medially. Epandrium with enlarged pair of anterodorsal bristles; subanal plate absent, epandrium and cerci discontinuous below anal opening; cerci fused to epandrium, with 1 enlarged bristle, ventral margin flat and straight. Surstylus quadrate, with dense ventral row of short bristles. Postgonites symmetrical, anteriorly curved, with distinct posterior angle, broad except for narrow, pointed apical process. Basiphallus gently curved, tubular, with posteroventral crest, open posteriorly dorsal to crest. Lateral sclerites narrow, rounded dorsally; central sclerite pointed at apex. Membranes elongate, narrow, and bulbous ventrally, with rows of lightly sclerotized ridges and warts.

Female Terminalia:

Tergite 8 fully sclerotized dorsally. Tergite 10 glabrous, orange, narrowly fused to cerci posterolaterally; cerci orange, glabrous on anterior half, pollinose on posterior half, apical bristles short. Sternite 7 broad, rounded, broader along anterior margin. Stenite 8 broad, wider than 1/2 the width of sternite 7, row of 4 small setae closely associated with posterior row of 6 larger setae. Stenite 10 not wrapped posteriorly around tip of abdomen, with lateral arms curved dorsally, pointed posterior margin, and a pair of enlarged lateral bristles. Spermathecae spherical; duct length 1-2 times length of spermathecal bulb.

Type material:

PARATYPES: SPAIN: Barcelona, Cardedeu, 5.i.l983 on cow dung, leg, M. Carles- Tolra (1$*); Barcelona, Cabrils, 8.iv.l982, on sheep dung, leg, M. Carles Tolra (1$).

141 Specimens Examined:

ARGENTINA: Salta, 30km E of Salta Campo Quijano, 18-20.ii.1992, dung tp. in forest remnant, S.A. Marshall (1$). BRAZIL: Rio de Janeiro, Teresopolis, Pedra do Sino, 2000m, 14.iii.1990, human dung, S.A. Marshall (1$*, 1$); Rio Grande do Sul Pelotas, vii.1998, C.L. Bicho (10^***-illustr-wing: debu00105584, 00105619, 00105590, 00105604, 00105608, 00105609, 00105610, 00105586, 00105617, 00105585; 16$*- illustr: 00105602, 00105594, 00105595, 00105605, 00105597, 00105599, 00105598, 00105611, 00105616, 00105587, 00105601, 00105593, 00105618, 00105600, 00105612, 00105603); same as previous except vii.1999 (8$: debuOO 105634, 00105646, 00105645, 00105644, 00105641, 00105632, 00105624, 00105637; 16$: 00105630, 00105638, 00105633, 00105629, 00105623, 00105639, 00105640, 00105631, 00105626, 00105625, 00105642, 00105643, 00105636, 00105627, 00105628, 00105635). CHILE: San Sebastian, 28.xii.1992, dry river, S.A. Marshall (1$). SPAIN: Canary Is., Isla Gomera, El Cedro, 1100m, 26.viii-2.ix. 1992, laurel forest, FIT, S.&J. Peck, 92-330 (1$); correct data probably: Canary Is., Gomera, 1992, carrion tps., S.&J. Peck (1$*).

Comments:

Coproica rohaceki is here newly recorded from the New World, with records from Argentina, Brazil and Chile.

142 Coproica rufifrons Hayashi, 1991 (Figures: 12, 20, 27, 243-253)

Distr.: Australasian/Oceanian: American Samoa, Australia (QLD), Cook Is., Fiji, France Polynesia (Tahiti), Kiribati, Marshall Is., Micronesia (Kapingamarangi, Satawa Is., Yap Is.), Palau, Papua New Guinea, Solomon Is., Tonga, Vanuatu, Western Samoa; Nearctic: Canada (NS, data suspect), USA (FL, NY [data suspect], TX); Neotropical: Argentina, Bermuda, Brazil, ?Bolivia, Dominican Republic, Ecuador, Galapagos Is. (Ecuador), Grenada, Guatemala, Jamaica, ?Mexico, St. Kitts, Trinidad, Virgin Is.; Oriental: China (HKG), Pakistan, Taiwan; Palaearctic: Canary Is. (Spain), Japan, Madeira (Portugal), Saudia Arabia, UAE. Coproica rufifrons Hayashi, 1991c: 237 [both sexes, phylogenetic notes, illustr.]. Type locality: Pakistan, Chilas. HT male (ITU). - Hayashi, 1995a: 232 [diagnosis, taxonomic remarks, illustr.]; Hayashi, 2001: 253 [new record from American Samoa, Australia (QLD, ), Cook Is., FIJI, France Polynesia (Tahiti), Kiribati, Marshall Is., Micronesia (Kapingamarangi, Satawa; I., Yap Is.), Palau, Tonga, Vanuatu, Western Samoa]; Rohacek, 2007b: 104 [new record from Madeira (Portugal)]; Gatt, 2008: 700 [photograph, new record from UAE].

Body length 1.1 - 1.9 mm. Body colour brown to dark brown. Face brown, frons dark brown, anterior 1/5 of frons orange. Interfrontal bristles in 4-5 subequal pairs, postocellar bristles convergent or cruciate, 1 pair of small postocellar setulae. Eye to gena ratio approximately 2:1, 3-4 genal setae in longitudinal row, anterior seta approximately 2 times length of posterior 3 setae. Scutellum with approximately 15 setae arranged in 2 loose rows between basal 1/4 and basal 1/2; 3 lateral bristles between larger anterior and posterior marginal scutellar bristles. Katepisternum with 2 dorsal bristles, posterior bristle distinctly longer than anterior bristle, anterior and posterior bristles each approximately 1/3 and 2/3 as long as the distance between dorsal margin of katepisternum and wing base respectively. Legs and fore coxa brown to light brown.

Mid tibia with additional small anterodorsal bristle at 1/7, and small proximal posteroventral bristles at 2/5 and 1/5. Mid basitarsus with 2-3 (usually 2) stout anterior bristles, 1 at 3/5 the other sub-basal, 2 sub-basal anteroventral bristles, distal

143 anteroventral bristle twice length of proximal bristle, 1 posteroventral bristle equal in length to distal anteroventral bristle, and 2 small ventral pairs of bristles setae on distal

1/2 half. Second costal sector 0.7-1.1 times length of third costal sector. Costal bypass long, extending 7 or more vein widths beyond R4+5. Anal vein sinuate, lacking an angulate bend.

Male Terminalia:

Sternite 5 rounded rectangular; all surface bristles uniform in size. Synsternite 6+7 straight medially. Epandrium with enlarged pair of dorsolateral bristles; subanal plate absent; epandrium and cerci discontinuous below anal opening, though epandrium merging ventrally with membranous portion of anal opening; cerci fused to epandrium, ventral margin with rounded medial processes, and 1 enlarged bristle. Surstylus with 2 posteroventral processes, each medially deflected, posterior most process pointed and 2 times length of other process, with ventral row of 5 or more bristles. Postgonites symmetrical, anteriorly curved, distinct posterior angle, apical 1/3 narrow and blunt at apex. Basiphallus gently curved, tubular, with rounded posteroventral crest, open posteriorly dorsal to crest. Lateral sclerites rounded, with medial process and small lateral warts; central sclerite narrowly fused with lateral sclerites sub-apically, with elongate apical process. Membranes elongate, narrow, with lightly sclerotized ridges.

Female Terminalia:

Tergite 8 long laterally, very short dorsally. Tergite 10 pollinose, narrowly articulating with cerci posterolaterally; cerci rounded, pollinose, apical bristles long and sinuate.

Sternite 7 broad, rounded rectangular. Sternite 8 rounded, approximately as wide as sternite 7, with posteromedial notch. Tergite 10 wrapped posteriorly around tip of

144 abdomen, with pairs of medial and lateral enlarged bristles. Spermathecae spherical; duct length 7-8 times length of spermathecal bulb.

Type material:

PARATYPES: PAKISTAN: Northwest Frontier Prov., Ayubia, 22.vii.1988, T. Hayashi (1$); Northwest Frontier Prov., Balakot, 2.viii.l988, T. Hayashi (4<$*); Northwest Frontier Prov., Chilas, 2.ix.l988, T. Hayashi (4<$, 4$); Northwest Frontier Prov., Kalam, Swat, 6-8.viii.1988, T. Hayashi (1$); Northwest Frontier Prov., Miandam, Swat, 3.viii.l988, T. Hayashi (3c?); Northwest Frontier Prov., Shangla Pass, 9.viii.l988, T. Hayashi (4$).

Specimens examined:

ARGENTINA: Salta, Rosario de Lerma, INESALT yard, 16-28.ii.1992, malaise, S.A. Marshall. BRAZIL: 15km NE of Porto Seguro, (Ecological Reserve "Pau-Brasil"), 19- 27.ii.1986, D.&V. Cristina & S. Amaorim (1$*); Mato Grosso, Cuiaba, Bairro Barbado, 17.xi.1990, Paganelli (1$); Mato Grosso, Cuiaba, Sao Geronimo, 4.iv.l990, BPF Group (2(J, 2$); Mato Grosso, Coxipo do Ouro, 17.xi.1990, C.H. Paganelli (2$); Mato Grosso, Pocone, Centro (14:25 hs), 16.V.1990, BDF Group (1$); Rio de Janeiro, Nova Friburgo Mury, 1000m, 11.ii.1990, swept off goat dung, open wet area, S.A. Marshall (1$); Mato Grosso, Varzea Grande, Mapim, 15.iii.1990, (5:30 PM), BPF Group (1$, 1$). CANADA: NS: Lockpont, 4.ix.l958, J.R. Vockeroth (1<$, data out of normal range). DOMINICAN REPUBLIC: Barahona, 4km E Barahona, 28-29.ix.1985, dung, R.E. Woodruff (1$); Monte Cristi, 3km N of Villa Elisa, 1-3.X.1985, dung, R.E. Woodruff (2$); Pedernales, 14.5km N of Cabo Rojo, 18-03N, 71-39W, 165m, 13-19.vii.1990, arid thorn-scrub, intercept tp., L. Masner, J. Rawlins, & C. Young (1<$, CMNH). ECUADOR: GALAPAGOS: Espanola, Bahia Manzanilla, 7-10.vi.1985, sand beach carrion tp., S.&J. Peck (\<$, 3$); Isabela, Sierra Negra, pampa, 1000m, 4.iii. 1989, horse dung - swept, B.J. Sinclair (6$, 6$); Santiago, 3km NE of Aquacate Camp, 740m, 5- 9.vi.l991, moss elfin forest, FIT, S. Peck, 91-201 (1$); St. Cruz, 4km N of Bellavista, Media Luna, 620m, 14.v-13.vii.1985, FIT Miconia zone (2.vii), S.&J. Peck (1$*, 1?); same as previous except 21-31.V.1985, Miconia zone, dung tps. (1$); 17-21.V.1985, Miconia zone, carrion tps. (1$); St. Cruz, Academy Bay, ECCD, 10.v-14.vii.1985, 30m arid zone, thorascrub malaise-FIT, S.&J. Peck (1$, 1 $); St. Cruz, Academy Bay, ECCD "Cuerva De Iguana", 8km N of Grieta, with water, lO.v.1985, bait tp., S.&J. Peck (3$- illustr); St. Cruz, agri.zone, l-9.iv.1989, malaise, Peck & Sinclair, 89-204 (1$); St. Cruz, CDRS, arid zone, 5-9.ii.1989, dung tp., B.J. Sinclair (\S, CNCI); same as previous except 31.i-9.ii.1989, tortoise dung tp. (2$-wing, 2$); St. Cruz, CDRS, littoral zone, 4- 9.H.1989, dung tp., B.J. Sinclair (16

145 Peck (2$); St. Catherine, 28.vii-l.viii.1974, dung under trees in horse pasture, Peck (1$). SAUDI ARABIA: Quwayzah, E of Jeddah, 24.X.2000, around camel urine, N. Gadallah (13

Comments:

See comments under Coproica hirtula (Rondani, 1880).

Coproica rufifrons is widespread and abundant in northern South America and southern United States, and is the most common species on the Galapagos Islands. Two specimens examined had label data indicating occurrences well north of the main range of this species: one from Washington D.C. and the other from Nova Scotia.

146 Coproica ruwenzoriensis (Vanschuytbroeck, 1950)

Distr.: Afrotropical: Kenya, Nigeria, Tanzania, Zaire. Limosina (Coprophila) ruwenzoriensis Vanschuytbroeck, 1950a: 41 [both sexes]. Type locality: Zaire, W. Ruwenzori, Kalonge. HT male (MRAC). Leptocera (Coproica) ruwenzoriensis. - Richards, 1960c: 201, 206 [generic combination, key]; Richards, 1980: 618 [Afrotropical catalog]. Coproica ruwenzoriensis. - Papp, 1979a: 104 [generic combination]. Papp, 2008: 13 [redescription, taxonomic notes, illustr., distr.].

Coproica saprophaga Papp, 2008

Distr.: Oriental: Thailand, Vietnam. Coproica saprophaga Papp, 2008: 25 [both sexes, taxonomic notes, illustr.]. Type locality: Thailand, Thung Khai Botanical Gardens. HT male (HNHM).

147 Coproica serra (Richards, 1938) (Figure: 254)

Distr.: Afrotropical: Ethiopia, Kenya, Uganda. Leptocera (Heteroptera) serra Richards, 1938a: 391 [male, illustr.]. Type locality: Kenya, E side Mount Elgon, Elgon Saw Mill, Camp II [2,470 m]. HT male (MNHN). Leptocera (Coproica) serra. - Richards, 1960c: 201, 206 [subgeneric combination, key]. Coproica serra. - Papp, 1979a: 103 [generic combination, female, diagnosis]. Papp, 2008: 16 [taxonomic notes, illustr.].

Body length 1.4 to 1.7. Body dark brown. Face and frons dark brown. Interfrontal bristles in 4 subequal pairs, postocellar bristles short, thick, and erect, 1 pair of small postocellar setulae. Eye to gena ratio 2:1 - 2.3:1; 4-5 genal setae in longitudinal row posterior to vibrissa, anterior seta approximately 1.5 times longer than posterior setae.

Scutellum with 20-25 setae evenly distributed between basal 1/6 and basal 2/3; 2 lateral bristles between larger anterior and posterior marginal scutellar bristles. Katepisternum with 2 small subequal dorsal bristles, each approximately 1/10 as long as the distance between dorsal margin of katepisternum and wing base. Legs and fore coxa brown. Mid tibia lacking proximal posterodorsal bristles. Mid basitarsus with anterior row of small, stout setae on extending from 1/2 to apex, a strong sub-basal ventral bristle, a strong anteroventral bristle at 1/2, and 2-3 small bristles along both posteroventral and anteroventral margins, with variable positions. Second costal sector 0.7-0.9 times length of third costal sector. Costal bypass short, terminating within 5 vein widths of R4-1-5. Anal vein sinuate, lacking distinct angulate bend.

Specimens examined:

KENYA: Masai Mara Nat. Pk., 10-ll.vii.1988, ex. buffalo dung, yellow pan tp., B.J. Sinclair (2c?*-wing, 4$*).

148 Comments:

Specimens of Coproica serra were identified on the basis of genitalic figures in

Papp, 2008. The ventral row of thickened bristles on the surstylus makes this species easily recognizable. The species also appears to a have ventral orange patch on the first flagellomere, a character that was overlooked by both Richards (1938) and Papp (2008).

This orange patch was very faint in some specimens examined, yet was distinct in others.

The anterior katepisternal bristle was noted as absent by both Richards (1938) and Papp

(2008), but is here determined to be present, though minute.

Male and female genitalia have not been illustrated or included in the redescription due to the low quantity and quality of observed specimens.

149 Coproica setulosa (Duda, 1929) (Figures: 11,28,255-265)

Distr.: Nearctic: USA (AZ, FL, MI, NM, TX); Neotropical: Antigua, Bolivia, Brazil, Colombia, Costa Rica, Dominican Republic, Ecuador, Honduras, Jamaica, Mexico (CHP, COA, GUE, HID, JAL, MOR, NAY, NUL, QRE, ROO, SIN, SON), Panama, St. Kitts, St. Lucia, Trinidad, Venezuela, Virgin Is.. Leptocera (Coprophila) setulosa Duda, 1929: 38 [both sexes]. Type locality: Bolivia, 60 km N San Jose de Chiquitos. STs, both sexes (SMNS). Leptocera (Coproica) setulosa. - Richards, 1960c: 202, 206 [subgeneric combination, key]; Richards, 1967b: 16 [Neotropical catalog]. Coproica setulosa. - Papp, 1979a: 104 [generic combination].

Body length 1.3 - 2.1mm. Body light brown to brown. Face light brown, frons brown.

Interfrontals in 4 subequal pairs, postocellar bristles slightly convergent, 1 pair of small postocellar setulae. Eye to gena ratio approximately 2.5:1; 10-15 small genal setae, anterior seta approximately 1.5 times length of posterior setae. Scutum with 3 additional pairs of dorsocentral bristles anterior to prescutellar pair, each approximately 1/3 the length of prescutellar. Scutellum with 25-35 discal setae between basal 1/5 and basal 1/2, row of 4 or more bristles at 1/2 approximately 5 times length of other discal setae; 4 lateral bristles between larger anterior and posterior marginal scutellar bristles, second lateral bristle long, approximately 2/3 length of anterior marginal bristle. Katepisternum with 2 long subequal dorsal bristles, anterior bristle extending to just beneath wing base, posterior bristle extending just beyond wing base. Legs and fore coxa brown to light brown. Mid tibia with additional small anterodorsal bristle at 1/5 with paired posterodorsal bristle, and strong proximal posterodorsal bristles at 1/3 and 1/2. Mid tibia with small, stout anterior setae at 1/2, 3/4, and sub-apical, a strong, sub-basal posteroventral bristle approximately 3 times width of mid basitarsus in length, an equally long anteroventral bristle at 1/2, and a small posteroventral bristle at 2/3. Second costal

150 sector 0.7-1.0 times length of third costal sector. Costal bypass long, extending 7 or more vein widths beyond R4+5. Anal vein with distinct angulate bend.

Male Terminalia:

Sternite 5 rounded rectangular, with posteromedial patch of microsetae; surface bristles enlarged laterally. Synsternite 6+7 straight medially. Epandrium with enlarged pair of dorsolateral and ventrolateral bristles; subanal plate present, narrow; cerci fused to epandrium, ventral margin rounded, with 1 enlarged central bristle. Surstylus with rounded posterior and ventral lobes, and row of short ventral bristles. Postgonites symmetrical, straight, broad, blunt at apex. Basiphallus elongate, gently curved, tubular, with blunt, elongate posteroventral crest, open posteriorly dorsal to crest. Lateral sclerites with 2 dorsal processes, dorsal margin deflected medially, apex rounded, with several small spines; central sclerite fused with lateral sclerites ventral to second process.

Membranes elongate, narrow, with apical spines.

Female Terminalia:

Tergite 8 reduced to long lateral sclerites, connected by a dorsomedial membrane.

Tergite 10 and cerci dorsally flattened; Tergite 10 glabrous, orange, central bristles minute, fused to cerci posterolaterally and tightly associated medially; cerci orange, glabrous anteriorly, pollinose posteriorly, cereal bristles short and straight, apical bristle modified into a stout spine. Sternite 7 broad, short, and concave posteriorly. Sternite 8 with pointed anterior margin, concave posterior margin, with 2 loose rows of 4 small bristles bordering a pair of larger bristles. Tergite 10 wrapped posteriorly around tip of abdomen. Spermathecae spherical; duct length approximately 2 times length of spermathecal bulb.

151 Specimens Examined:

BRAZIL: 15km NE Porto Seguro, (Ecological Reserve "Pau-Brasil"), 19-27.ii.1986, primary Atlantic forest, Shannon-tp., D.&V. Cristina & S. Amaorim (1$); Minas Gerais, Instituto Estadual De Florstes, Parque Florestal Do Ibitipoca, 1700m, 12.ii.1990, swept near fresh dung, S.A. Marshall (lc?, 1$). COLOMBIA: Antioquia, La Ceja, 2100 mt., 3.ii.l984, manure pile (2c?s, 6$). COSTA RICA: Heredia, Rara Avis, El Plastico 1st Forest Mar. house, 580m, 19-20.V.1987, human dung, Norrbom (1$); Puntarenas, Osa Peninsula, 2.5 km S of Rincon, ~50m, 8°42'1N, 83°30'50W, 10-ll.viii.2001, treefall, yellow pans, M. Buck (lc?: debuOO 199840); Guanacaste, Ref. Nac. Fauna Silvestre, Rafael Lucas Rodriguez, Palo Verde, 10m, xi.1990, D. Acevedo, L-N-259000, 888400 (3c?, 1$). DOMINICAN REPUBLIC: Barahona, 4km E of Barahona, 28-29.ix.1985, dung, R.F. Woodruff (2c?, 4$); Monte Cristi, 3km N of Villa Elisa, 1-3.X.1985, dung, R.E. Woodruff (13c?, 9$); Puerto Plata, 22.U989, sweep on wet tr., S.A. Marshall (lc?); San Juan, El Capa, 1km off rd. to Vallejuela, 21.v. 1985, Bl tp., Nunez, Woodruff, & Stanger (1$). ECUADOR: Napo Baeza, 2000m, l-10.iii.1979, S.A. Marshall (4c?, 2$); Cotopaxi, 40km N of Latacunga, 3350m, 17-25.vii.1985, paramo, carrion, S.&J. Peck (5$); Cotopaxi, 45km NNE of Latacunga, 3700m, 19-25.vii.1985, carrion tp., shrub grass, paramo, S.&J. Peck (lc?); Pichincha Nono, 15km NW of rd. to Mindo, 24.x. 1999, roadside sweep, cow dung, S.A. Marshall (lc?: debuOOl 15283; 1$: debuOO 115285); Pichincha Pk., 4km W of Aloag, 3000m, 19-25.vii.1985, carrion tp., pasture at ravine edge, S.&J. Peck (2c?*, 3$); Rio Palenque, 25-26.ii.1976, dung tp., S. Peck (lc?). HONDURAS: Dpto. F. Morazan, El Zamorano, 22-29.vii.1970, blacklight tp., G.F. Freytag (1$: debu00020362); Uyuca, 5900', 3.vi.l994, malaise, H. Howden (1$); F.M., 6km SE of Zamorano, 7.vi.l994, dung tp., H. Howden (lc?); Zamorano, 30.V.1994, H.&A. Howden (4c?, 3$). JAMAICA: St. Ann, Discovery Bay, 3-6.ix.1974, dung, S. Peck, (7c?, 16$); Montego Bay, ex. horse manure, 18.U985, J.E. Corrigan (6c?, 4$); St. Catherine, 28.vii-l.viii. 1974, dung under trees in horse pasture, Peck (4c?, 3$). MEXICO: 6 mi S junction, Mex. 60, 68, 6500', N 6 Mex, 27-30.V.1971, human dung, A. Newton (lc?); Chiapas, 2 mi S of Trinitaria, 5100', 21-24.viii.1971, oak-tropical deciduous human dung, A. Newton (3c?, 5$*); Coahuila, Boquillas, 24-26.ix.1979, pig dung tp., R.E. Woodruff, G. Fincher (23c?, 22$); Hidalgo, 12 mi SW of Jacala, 6200', 23-30.vi.1971, walnut, oak pine, dung, A. Newton (lc?); Quintana Roo, 45 km NE of Ixtapa, 10-12.vii.1987, B. Gill (2c?). ST. KITTS & NEVIS: Bayford Dairy, 300m, 1- 15.xii.1985, malaise old field/for., L.D. Coote (lc?); Nevis, Dasents Estate, 220m, 28.viii- 21.ix.1985, malaise, citrus orchard, L. Coote (14c?, 10$). TRINIDAD: St. George Co., Arima Ward, NY Zool. Soc. Sta., 1 l.vi.1977, pig dung tp., R. Woodruff (lc?); UNITED STATES OF AMERICA: AZ: Cochise Co., Chiricahua Mts., 6mi ESE of Portal, 4150', ix.1970, desert grass human dung, A. Newton (lc?); St.Cruz. Co., 7mi WNW of Nogales, Calabasas Canyon, 3850', ix.1972, desert grass, dung, A. Newton (6c?*, 4$*); St. Cruz Co., 8mi NNW of Nogales, Walker Canyon, 3900', ix.1970, oak woodland, dung, A. Newton (28c?*, 14$); St. Cruz Co., Santa Rita Mts., Madera Canyon, 5500' ix.1970, oak, human dung, A. Newton (2c?, 3$). FL: Alachua Co., Gainesville (Hogtown Creek), 12.x. 1976, P.M. Choate & R.E. Woodruff (lc?*, 4$); Dade Co., Hialeah, 9.vii.l971, at human feces, C.E. Stegmaier Jr. (lc?); Gainesville, UF Beef Farm, 13.vii.1991, 1A, G.Y. Hu (3c?, 3$); Marion Co., Rd 65, Ocala Nat. For., 18-23.vi. 1984, sand-pine, human dung

152 tp., S. Marshall. MI: Gogebic Co., Crooked Lk. Bog, 23.vi.1978, leg, Ber, otter dung & root mat. subst., J. Wagner (lc?). NM: Torrance Co., 6mi SW of Manzano, Red Can. cpgd, 75-9000', 17-30.vi.1979, spruce-fir litter, S.&J. Peck (1$). TX: Big Bend Nat. Pk., Bouquillas Crossing, 25-26.ix.1979, pig dung tp., R. Woodruff & G. Fincher (3<$, 3$); Brewster Co., 3.5mi E of Marathon, 26-27.ix.1979, dung tp., R. Woodruff & G. Fincher (8<3\ 11$); Cameron Co., Laguna Atascosa Nat. Wildlife Ref, vii.1972, thorn forest, dung, A. Newton (2c?*-illustr, 8$*-illustr); Hidalgo Co., Bentsen-Rio Grande Valley St. PL, 21-25.V.1971, willow-thorn for. near riv., dung, Newton {!<$*, 3$); Medina Co., 8mi E of Sabinal, Rt.90, 22-28.ix.1979, dung tp., R. Woodruff & G. Fincher (1$); San Patricio Co., Welder Wildlife Ref, site V, 21-23.V.1993, H.&A. Howden (ltf-wing: debuOO 117844-wing); Terrell Co., 30mi E of Sanderson, 23-27.ixl979, on pig dung, B. Woodruff & G. Fincher (4c?, 6$); Uvalde Co., lOmi W of Uvalde, Nueces riv., Rt.90, 22-27.ix.1979, pig dung tp., Woodruff & Fincher (3$); Val Verde Co., 17mi W Langtry, 23-27.ix.1979, pig dung tp., H. Woodruff & G. Fincher (21c?, 13$). VENEZUELA: Bergentina, 18.iv-7.v.l988, dry forest near ck., FIT, S.A. Marshall (1$); Trujillo, Guaramacal Nat. Pk., 2000-3000m, 26.viii-l.ix. 1992, car net, L. Masner (1$: debuOO 152906). VIRGIN ISLANDS: St. Thomas, Est. Bordeaux, lO.viii. 1980, molasses & dung pit fall tp., M. Ivie (ltf).

153 Coproica testudinea sp. no v. (Figures: 266-275)

Distr.: Nearctic: USA (FL).

Body length 0.9 - 1.3mm. Brown along dorsum, yellow to light brown and ventrally.

Face and frons brown; head appearing longitudinally flattened, head height 1.5 to 1.9 times head length. Interfrontal bristles in 4 subequal pairs, postocellar bristles straight or slightly convergent, 2 pairs of small postocellar setulae. Eye small, gena striate and free of microsetae on dorsal half, light reflective polinosity present on ventral half, eye to gena ratio approximately 1.4:1; 2 enlarged genal setae, 2-3 more scattered genal setae.

Thorax light brown dorsally, yellow ventrally. Scutellum with 10-13 discal setae arranged in 2 rows between basal 1/3 and basal 2/3; 2 lateral setae between anterior and posterior marginal bristles. Katepisternum with 2 small, subequal dorsal bristles, each approximately 1/5 as long as distance between katepisternum and wing base. Legs and fore coxa yellow. Mid tibia with 1 proximal posterodorsal bristle at 1/2. Mid basitarsus with anterior row of small, stout bristles, a sub-basal ventral pair of bristles, and 2 ventral rows of small bristles setae along entire length of mid basitarsus. Second costal sector

0.7-0.9 times length of third costal sector. Costal bypass short, terminating within 5 vein widths of R4+5. Anal vein indistinct.

Male Terminalia:

Sternite 5 rounded rectangular, patch of small setae posteromedially; surface bristles uniform in size. Synsternite 6+7 straight medially. Epandrium with enlarged pair of dorsolateral bristles; subanal plate present, thick; cercus fused to base of epandrium, ventral margin rounded but unmodified. Surstylus with quadrate posterior lobe, lobe covered in dense setae and separated from rest of surstylus by ridge, anterior portion with 154 ventral 90° angle. Postgonites symmetrical, straight, and narrow on distal 2/5.

Basiphallus gently curved, tubular, posteroventral margin bulbous, closed posteriorly.

Lateral sclerites narrow, with elongate, membranous proximal dorsal process, a bifurcate dorsal process, and a subapical process emerging from medial surface; central process fused with lateral sclerites ventral to bifurcate process. Apical membranes short.

Female Terminalia:

Tergite 8 unmodified, sclerotized dorsally. Tergite 10 pollinose, narrowly fused to cerci posterolaterally; cerci pollinose, apical bristles short and sinuate. Sternite 7 broad, with blunt posterior point. Sternite 8 rounded, triangular, with lateral patches of 4 small bristles. Tergite 10 not wrapped posteriorly around tip of abdomen, bulbous and rounded, with pair of enlarged medial bristles. Spermathecae pear-shaped; duct length 3-

4 times length of spermathecal bulb.

Type specimens:

HOLOTYPE: UNITED STATES OF AMERICA: FL: Putnam Co., Hollister, 17- 23.vii.1984, Gopher tortoise Burrow, E.G. Milstrey (IS).

PARATYPES: UNITED STATES OF AMERICA: FL: Same as holotype (ltf*- illustr); Alachua Co., Lochioosa GMA, 3.x. 1984, gopher tortoise burrow, Eric G. Milstrey (lc^-wing); same as previous except dated 21.X.1984 (2$**-illustr); Putnam Co., Hollister/Robert's ranch, 5.xii.l984, gopher tortoise burrow, emerged from material, Eric G. Milstrey (\$, 3$*); Putnam Co., Hollister/Robert's ranch, 5.xii.l984, gopher tortoise burrow, Eric G. Milstrey (1$); same as previous except dated 15.xi.1984 (1$); 20.vi.l984(lc?).

Etymology:

The specific name is derived from the Latin word testudo, which means "tortoise," and is given based on the only known habitat of this species, within gopher tortoise burrows.

155 Comments:

Coproica testudinea is a rare species only known from gopher tortoise burrows in

Florida.

156 Coproica thaii Papp, 2008 Distr.: Oriental: Nepal, Thailand Coproica thaii Papp, 2008: 17 [both sexes, taxonomic notes, illustr.]. Type locality: Thailand, Mae Fang NP.. HT male (HNHM).

Coproica unispinosa Papp, 2008

Distr.: Oriental: India, Thailand, Vietnam; Palearctic: China (Beijing) Coproica unispinosa Papp, 2008: 34 [both sexes, taxonomic notes, illustr.]. Type locality: Thailand: Mae Taeng Elephant Camp. HT male (HNHM).

157 Coproica urbana (Richards, 1960) (Figures: 276-287)

Distr.: Nearctic: Canada (AB, BC, MB, NB, NS, ON, SK); USA (AL, AR, AZ, CO, FL, GA, ID, IL, IN, KY, MA, MI, MN, MS, MT, NC, ND, NH, NM, NY, OH, OK, PA, SC, TN, TX, UT, VA, WV, WY). Neotropical: Costa Rica, Cuba, Mexico (CHP, OAX). Leptocera (Coproica) urbana Richards, 1960c: 204 [both sexes]. Type locality: USA, Illinois, Urbana. HT male (USNM, lost). - Richards, 1965c: 725 [Nearctic catalog]. Coproica urbana. - Papp, 1979a: 104 [generic combination].

Body length 1.1 - 1.8mm. Uniformly light to dark brown. Face and frons brown.

Interfrontal bristles with 3 subequal pairs, postocellar bristles cruciate, small postocellar setae absent. Eye to gena ratio approximately 2:1; 3 small dorsally curved genal bristles in longitudinal row, additional small setae between longitudinal row and ventral margin of gena. Scutellum with 10-14 discal setae arranged in 2 rows between basal 1/4 and 1/2;

2 lateral setae between larger anterior and posterior marginal scutellar bristles.

Katepisternum with 2 small, subequal dorsal, each approximately 1/6 as long as the distance between dorsal margin of katepisternum and wing base. Legs and fore coxa light brown to yellow. Mid tibia with 1 proximal posterodorsal bristle at 1/2. Mid basitarsus with anterior row of small bristles extending from 1/3 to apex, a stout sub-basal anteroventral bristle (occasionally with additional anteroventral bristle distal to sub-basal bristle), and a cluster of 2-4bristles at 1/2 (typically 2 anteroventral and 1 posteroventral bristle, or 1 anteroventral and 1 posteroventral bristle). Second costal sector 0.8-1.0 times length of third costal sector. Costal bypass long, extending 7 or more vein widths beyond

R4+5. Anal vein sinuate, lacking an angulate bend.

158 Male Terminalia:

Sternite 5 rounded rectangular; surface bristles uniform in size. Synsternite 6+7 straight medially. Epandrium with enlarged pair of dorsolateral bristles; subanal plate absent, epandrium and cerci discontinuous below anal opening; cercus fused to epandrium, ventral margin relatively flat and straight, deflected medially, with 1 enlarged central bristle. Surstylus with rounded posteroventral process with slight constriction at base and apical patch of setae. Postgonites asymmetrical, straight, left postgonite narrow, and rounded and spoon-like at apex, right postgonite narrow along distal 1/3, with shallow hook at apex. Basiphallus elongate, straight, tubular, with small posteroventral process, open posteriorly. Lateral sclerites rounded, narrow, with sharp dorsomedial process, and short, narrow apicoventral process; central sclerite fused apically with lateral sclerites.

Female Terminalia:

Tergite 8 reduced to 2 lateral sclerites connected by dorsomedial membrane. Tergite 10 pollinose, broadly fused to cerci; cercus pollinose, apical setae short and sinuate. Sternite

7 broad, rounded, with slight posteromedial notch. Sternite 8 rounded, with anteromedial point, a pair of anterior setae, and central and posterior rows of 4 small setae. Sternite 10 wrapped posteriorly around tip of abdomen, and with 1 enlarged pair of medial bristles.

Spermathecae spherical, slightly tapering towards duct; short portion of duct sclerotized, with sharp constriction at base of bulb, duct length 4-5 times length of spermathecal bulb.

Specimens examined:

CANADA: AB: Wagner Bog, 2-9.viii.1985, pan tp. 1, Leg, A.T. Finnamore (1$, PMAE); same as previous except pan tp. 2 (2<$, 1 $, PMAE); pan tp. 7 (3<3\ PMAE); pan tp. 9 (1<3\ PMAE); Wagner Bog, 12-25.vi.1985, pan tp. 7, leg, A.T. Finnamore (1& PMAE); same as previous except pan tp. 8 (1$, PMAE); Wagner Bog, 26.vi.-2.vii. 1985, pan tp. 2, leg, A.T. Finnamore (3$*, PMAE); same as previous except pan tp. 5 (\<$, PMAE); Wagner Bog, l-7.viii.1985, pan tp. 1, leg, A.T. Finnamore (1$, PMAE); same

159 as previous except 3-ll.viii.1985, pan tp. 10 (2<$, PMAE); 6-13.viii.1985, pan tp. 3 (16\ PMAE); 10-16.viii.1985, pan tp. 6 (1& PMAE); 16-22.viii.1985, pan tp. 7 (lc?, PMAE); 17-24.viii.1985, pan tp. 8 (1& PMAE); 25.vii-l.viii. 1985, pan tp. 1 (lc?, PMAE). BC: Boundary Ck. Prov. Pk., 30.vii.1980, sweeping, G. Gibson (3c?*, CNCI); Osoyoos, Mt. Kobau, 560m, 4.vi-8.vii.l991, D. Blades & C. Maier, SOCAP - LA2 (17*$, 11*?, BCPM); same as previous except 8-13.vii.1991, SOCAP - LA3 (1$, BCPM); 14.vii- 23.viii.1991, SOCAP - LA4 (1$, 2$, BCPM); 24-28.viii.1991, SOCAP - LM5 (1$, BCPM); Osoyoos, Mt. Kobau, 1080m, l.vi.1991, D. Blades & C. Maier, SOCAP - OS1D1 (IcT, BCPM); Osoyoos, Mt. Kobau, 1100m, 8-12.vii.1991, D. Blades & C. Maier, SOCAP - OP32 (1$, BCPM). MB: Devils Lk., "100km S Grand Rapids"(?), 7.vi- 16.ix.1984, pine/aspen, S.&J. Peck (1$); Erickson, l-5.viii.1983, mushroom pitfalls, DH Pengely & KN Barber (2$). NB: St. Andrews, Gibson Lk., viii.1978, pitfall tps., S. Marshall (3c?***, 1 $*). NS: Antigonish Co., 1km N of Antigonish, 2.vi-5.viii.l984, FIT, L. MacMillan (1$); Cape Breton Highlands Nat. Pk., Lone Shielding, PG729861, 19- 21.vii.1983, FIT, D.&J. Bright (1$). ON: Algonquin Prov. Pk., Swan Lk. stn., Scout Lk. Survey C7, 28.vii-05.viii. 1994, maple pans, S.A. Marshall (IcT*); Bruce Co., Cameron Lk. Rd., S of spring fen, 20-29.vii.1999, pan tp., S.A. Marshall (2$*: debu00081687, 00081688*; 1$*: 00081680*); Bruce Co., Dunks Bay, 5-12.vii.1999, dune malaise, S.A. Marshall (1

160 Juarez, 7600', 10-18.viii.1973, oak woodland, dung, A. Newton (let). UNITED STATES OF AMERICA: AL: Bon Secour, NWR, 6.V.1993, vac. off rotting mushroom, S.A. Marshall (3c?***, 2$***); Monroe Co., 2mi E of Perdue Hill, Claiborne Mem. Br., iii.1977, dung tp., Wiley & Woodruff (let*, 1?*). AZ: 10 mi NW of Flagstaff, San Francisco Mts., 9500', 18-24.vii.1979, spruce-fir-aspen meadow malaise, S.&J. Peck (IcT*); Apache Co., Alpine, Luna Lk., 9-14.vii.1979, pine meadows, S.&J. Peck (lc?*, 1$*); Cochise Co. Chiracahua Mts., E Turkey Ck., 6500\ 15-2l.vii. 1978, dung tps., O. Kukal (let*, 1$); Cochise Co., 5100\ Coronado Nat. For., Chiricahua, Mt. Stewart cpgd., 16-18.viii.1984, rd. dung, L.B. Carlson (let*); Cochise Co., Huachuca Mts., Miller Canyon, 6000', ix.1970, dung, oak woodland, R. Newton; Cochise Co., Huachuca Mts., Carr Canyon, 7100', ix.1970, oak & evergreen, human dung, A. Newton (let*); Coronado Co., Bonito Pk., near Flagstaff, 5-9.viii.1984, dung tp., B.V. Brown (let*); Santa Cruz Co., 7mi WNW of Nogales, Calabasas Canyon, 3850\ ix.1972, desert grass, dung, A. Newton (4cT***-BOLD, 2$*-BOLD). AR: Garland Co., 1.2mi N of Crystal Springs, Hwy 270, 6-8.iii.1977, pig dung tp., Woodruff & Wiley (let*, 1?*); Madison Co., 3mi S Brashers, 1600', 19.vi-12.vii.1972, mixed hdwd. for., dung, A. Newton (let*, 1$*); Montgomery Co., 5.4mi E Ida, Hwy 270, 6-8.iii.1977, pig dung tp., Woodruff & Wiley (let*, 2?**); Newton Co., 20.viii.1976, Edward Rogers (1$, CASC); Wash Co., 3mi S Devils Den St. Pk., 28-31.V.1979, oak hickory, S.&J. Peck (2c?**, 1?*); Wash. Co., Devils Den St. Pk., Devils Den Cave, 28-31.V.1979, S.&J. Peck (let*, 1?*). CO: Roatte Co., 5mi NE of Clark Hinman cpgd., 7600', 23-25.vi.1972, dung tp., (15et). FL: Alachua Co., Gainesville, Hogtown Creek, 12.x. 1976, P.M. Choate & R.E. Woodruff (let*, 1?*); Alachua Co., Gainesville, 2-5.vii.1982, pig dung, horse pasture, K.L. Rench (IcT*, 1?*); Alachua Co., Rd.176, ll.v.1983, carrion tp., K.W. Vick (let*, 1?); Broward Co., Hwy84 mi59, 12.xii.1985, old carrion, S. Marshall (2ct-BOLDx2); Broward Co., 12.xii.1985, S.A. Marshall (IcT-BOLD); same as previous except under cow dung (let*); Clay Co., Gold Head Branch St. Pk., iv.1971, mixed hard wd., human dung, A. Newton (2c?**, 1?*); Clay Co., Gold Head Branch St Pk., 4-14.iv.1971, carrion, A. Newton (let*, 1?*); same as previous except dung (let*, 1$*); Highlands Co., Archbold Res. Stn., 15-16.iv.1989, dung vac, K.N. Barber (let*); Highlands Co., Archbold Bio. Stn., Lk. Placid, xi.1979, D. Hardwick (let*, 1$*); Highlands Co., Archbold Bio. Stn., Lk. Placid, xi.1978, D.&V. Hardwick (1$*, 1?*); Highlands Co., Archbold Biol. Sta., Lk. Placid, Highlands Co., 10.V.1985, M. Deyrup (let, 1$, ABS); Highlands Co., Archbold Biol. Sta., Lk. Placid, Highlands Co., 4.V.1983, malaise tp., tr. 1,550, M. Deyrup (let, ABS); same as previous except on unidentified scats (let, ABS); Highlands Co., Archbold Res. Stn., 17.iv.1989, dung vac, S.A. Marshall (2et**); same as previous except 14.xii.1985, vac. under carrion (2ct*-BOLD); 12-16.xii.1985, scrub, dung pitfall (1?*); 18.xii.1985, on old carrion site (let*, 1?*); 12-16.xii.1985, dung pan, scrub-sand (3et**, 1 ?); 13.xii.1985, at light (let*); 17-21.xii.1985, scrub dung (let*, 1$*); Hendry Co., La Belle Capt, Hendry Rd., iv.1971, pine, hardwood, near riv., human dung, A. Newton (let*, 1?*); Highlands Co., Highlands Hammock St. Pk., Old Hammock Tr., 15-2l.xii. 1985, dung tp., S.A. Marshall & B.V. Brown (let*, 1?*); Highlands Co., Archbold Bio. Stn., 13.xii.1985, rot fungi, S. Marshall (let*, 1?*); same as previous except 13-18.xii.1985 (let*, 1$*); 16.xii.1985, carrion residue (let*, 1?*); Highlands Co., Archbold Biol. Stn., 6.X.1964, P.H. Arnaud Jr. (2$, CASC); Highlands Co., Highlands Hammock St. Pk., (orange grove), 14-18.vi.1982, pig dung, Woodruff &

161 Rench (1$*, 1?*); Highlands co, 3mi N of Avon PL, 14-18.vi.1982, pig dung tps. in gopher burrow, R.E. Woodruff & K.L. Rench (1$*, 1?*); Lee Co., Sanibel Is., 27.iv- 3.V.1983, dung, Carlson & Brown (1$*, 1?*); Marion Co., Ocala Nat. For., Oklawaha, 1 l-12.vi.1984, swamp, dung tp., S. Marshall (16**); Marion Co., Ocala Nat. For., Rd. 65, 1.5mi W of St. Rd. 19, 15-16.iii.1984, dung, R. Woodruff (1$*); same as previous except Rd.90, 8mi N of St. Rdl9, pig dung tp., (ltf*, 1?*); Marion Co., Ocala Nat. For.., Silver Springs 1 l.vi. 1984, woods, human dung sand pine, S. Marshall (16**, 1?*); same as previous except 7-ll.vi.1984, FIT (16**, 1$*); 22.vi.1984, vac. wild mushroom (1$*, 1$*); Marion Co., Silver Springs Woods, Ocala Nat. For., 8-1 l.vi.1984, human dung, S. Marshall (16**, 1?*); Marion Co., Rd.65, Ocala Nat. For., 18-20.vi.1984, sand-pine, human dung tp., S.A. Marshall (1<$*, 1 $*); same as previous except 18-23.vi.1984 (1$*, 1$*); Marion Co., Salt Springs, Ocala Nat. For., 14-18.vi.1984, pine, human dung, S. Marshall (16**, 1$*); Marion Co., Zay Prairie, Ocala Nat. For., 10-14.vi.1984, mushroom tps. by shore, S. Marshall (16**); Marion Co., Zay Prairie, Ocala Nat. For., 8- 14.vi.1984, carrion tp., S. Marshall (1$*, 1?*); Okaloosa Co., 4.5mi N of Holt, Blackwater R. State For., A&M Res. Stn., pig dung, L. Stange (1?*); Okaloosa Co., 4.5mi N of Holt, Blackwater R. State For., A&M Res. Stn., 15-17.vi.1978, pig dung, L. Stange (16**); Polk Co., 2 mi N of Lake Wales, 25-29.iv.1983, pig dung tp., R.E. Woodruff (IcH; Polk Co., Rt.27, 7mi N of 14, 25-29.iv.1983, pig dung tp., R.E. Woodruff (1(5**, 1?*); Putnam Co., Ocala Nat. For., Johnson Field cpgd, iv.1971, mixed hardwood, human dung, A. Newton (2$**, 2$**); Sarasota Co., Myakka River St. Pk., 22-23.V.1982, pig dung tp., R.E. Woodruff (2$**, 2$**); Suwannee Co., 8.7mi W of O'Brien, S-349, 25.iv.1977, pig dung tp. #6, gopher tortoise burrow, Woodruff & Woley (16*, 1$); Volusia Co., Tomoka St. Pk., 20.vi.1984, dung tp., S.A. Marshall (1$*, 1$*). GA: Camden Co., 14mi N of Folkston, 10-18.iv.1989, pan tp., human dung, J.E. Swann (16**); Rabun Co., Terrora Pk., Tallulah Falls, 4-ll.vi.1983, pig dung tps., K.W. Vick (26***, 2$**); Wilkinson Co., Big Sandy Ck., 4mi S of Irwinton, US441, 5-25.vi.1984, dung tp., S.A. Marshall (1$*, 1?*). ID: Clark Co., 2mi E of Memphis, 6-10.vi.1983, pig dung, K.W. Vick (16**, 1?*); Lawrence Co., Hoosier Nat. For., Donaldson Tr., 19.V.1991, J.E. Swann (16**, 1?*); Putnam Co., nr. Cunot, Lieber St. Pk., 31.vii.1984, vac. dog dung, L.B. Carlson (16**)- KY: Edmonson Co., Mammoth Cave Nat. Pk., 15.v- 20.viii.1983, mesic for. FIT, S.&J. Peck (16**, 2?*); Rowan Co., 24km SW of Morehead Cave, 14.v-20.viii.1983, Fagus for. FIT, S.&J. Peck (16**, 1 $*); same as previous except Run Lake (16**, 1?*). LA: Grant Parish, 18 km N of Alexandria, Stuart Lk. cpgd., 19.v- 17.viii.1983, for. FIT, S.&J. Peck (lc?*, 1?*). MI: Gogebic Co., Crooked Lk. Bog, 15.viii.1977, leg, J. Wagner (266*, 14$). MS: Forrest Co., 6mi W of Wiggins, Sweet Bay Bog, 5-8.V.1994, Sphagnum, dung tps., S.A. Marshall (26***, 1?*); Pontotoc Co., 32km SW of Tupelo "Tackshish"(?), mi 250, 17.v-19.viii.1983, oak for., FIT, Peck (16**). MT: Ravalli Co., Bitterroot Nat. For., Sam Billings Mem. cpgd., 8-17.viii.1990, pans along stream human dung, J.E. Swann (46*). NC: Bladen Co., Singletary Lk. St. Pk., 34°35'N, 78°27'3W, pans, Marshall & Paiero (16*: debu00227144; 1$: 00166568); Jackson Co., 2300', Cullowhee, Cane Ck., 5-25.vi.1984, riparian, dung tp., S. Marshall (16**, 1?*); Jackson Co., Blue Ridge Pkwy, Bear Paw Gap, mi 29, 4500', 5-27.vi.1984, mushroom tp., S.A. Marshall (16**). ND: Billings Co., Medora, Roosevelt Nat. Pk., 27.viii.1979, sage grassland, carrion, S.&J. Peck (76*, 1$); Billings Co., Roosevelt Nat. Pk., 28.viii.1979, carrion (1& 1$). NH: Coos Co., 1 mi NE, vii-7.viii.1986, FIT, D.S.

162 Chandler (1$). NM: Bernalillo Co., Isleta, 4900', 16-20.vi.1979, floodplain for., carrion, S.&J. Peck (1(?*, 1$); Catron Co., 5mi W Luna, 7400', San Francisco River Pond, 9- 14.vii.1979, pine-meadow, S.&J. Peck (1$*); Eddy Co., 30mi WSW of Carlsbad, Sitting Bull Falls, 4600', 23-27.vii.1975, dung 15, S. Peck (ltf*); Lincoln Co., South Fork cpgd. 7, 7mi W of Angus, 7600', 6-8.vii.1972, dung (2c?**); Santa Fe Nat. For., near Windsor cpgd., 27.V.1991, spring near tr., J.E. Swann (lc?*); Socorro Co., 20mi W Socorro, Water Canyon, 7000', 28.vi.-4.vii. 1979, mixed "mesic" forest, S.&J. Peck (1$*, 1?*). NY: Lk. Placid, 30.viii.1979, dung tp., S.A. Marshall (1$*, 1?*). OK: Comanche Co., Wichita Mts. Refuge, 5-7.vi.1979, shortgrass prairie, S.&J. Peck (1$*, 19*); Latimer Co., 5mi W Red Oak, 5-1 l.vi. 1977, dung tp., K. Stephen (7c?*x7). PA: Blue knob St. Pk., 3000', 5.V.1991, dear dung, S.A. Marshall (1$*); Trout Run, 28.ix.1979, dog shit, S.A. Marshall (\<3*, 1$*). SC: 13mi W Orangeburg, 27.iii.1980, S.A. Marshall & S.C. Weeses (lc?*). TN: Servier Co., 7.9mi S of Gatlingburg, 3000', 21-24.V.1977, carrion, A. Newton (19*). TX: Angelina Co., Angelina Nat. For., Boykin Cemetery, 13-17.vii.1993, spring fen, dung, S.A. Marshall (2c?**: debu00128224*, 00128192*; 4$*x4: 00128230*, 00128347*, 00128191*, 00128195*); Bastrop Co., Bastrop St. Pk., 24-27.V.1983, oak woods, carrion tps., S.&J. Peck (lc?*, 19*); Bastrop Co., Bastrop St. Pk., 8.iv.2000, mud lake shorepan tps., M. Buck (lc?*: debuOOl 14100*; 19*: 00114050*); Bouton Lk., 13- 17.vi.1993, dung & vac. tps., S.A. Marshall (2c?**, 29**); Brazos Co., College Station, Lick Ck. Pk., 5-9.iv.2000, post oak savannah by ck. malaise tp., M. Buck (2c?-BOLDx2: debuOO 117643-BOLD, 00117645-BOLD); same as previous except 28-30.iii.2000, bottomland for. near ck., malaise tp. (lc?-BOLD: debu00109688-BOLD); Cameron Co., Laguna Atacosa Nat. Wildlife Refuge, vii.1972, thorn forest, dung, A. Newton (29**); Hidalgo Co., Bentsen-Rio Grande Valley St. Pk., 21-25.V.1971, willow thorn for. near riv., dung, Newton (2c?**, 19*); Kimble Co., Llano River at Co. rd. 385, l-2.iv.2000, floodplain for., pig dung pitfalls, E.G. Riley (lc?*: debuOO 110642*; 19*: 00110635*); Montgomery Co., 4.5 mi N of Montgomery, 2.v-17.vi.l987, FIT, pine/black walnut for., R.S. Anderson (lc?*); San Jacincto Co., Coldspring, Double Lk. cpgd, 22.v-16.viii. 1983, FIT, S.&J. Peck, 83-14 (lc?*, 19*); San Jacincto Co., 5km S Coldspring, Double Lk. cpgd., 22.v-16.viii.1983, for. FIT, S.&J. Peck (lc?*,l9*); same as previous except 22- 24.V.1983, for. carrion tp., Peck (lc?*, 19*); San Patricio Co., Welder Wildlife Ref., site V, 21-23.V.1999, H.&A. Howden (lc?*: debuOOl 17822*; 19*: 00117830*); Uvalde Co., lOmi W of Uvalde, Nueces Riv., Rt.90, 22-27.ix.1979, pig dung tp., Woodruff & Fincher (lc?*,l9*); Warren, Kirby St. For., 12-17.vi. 1993, dung tps., S.A. Marshall (3c?*, 39*). UT: Dagget Co., Dutch John, 6300', 2-8.viii.1979, pinyon juniper, woodland, carrion, S.&J. Peck (3c?***). VA: Bath Co., 9.6km N of Clifton Forge, 13.v-21.viii.1983, Tsuga, Fagus for., FIT, S.&J. Peck (lc?*); Prince William Co., 0.5km NE of Thorofare Gap, vie. Haymarket, 25.vi.1966, P.H. Arnaud Jr. (19*, CASC); Shenandoah Co., 16km S of Strasburg, 12.v-22.viii.1983, for. FIT, S.&J. Peck (lc?*, 19*). WV: Pendleton Co., Spruce Knob, 4600', 27.vii-9.viii, conifer for., C.S. & A. Newton. WY: Crook Co., Black Hills Nat. For., Cook Lk. Conservation Area Rd., 4.2km E of cpgd, 4-2 l.viii. 1990, pans; cow dung, stream in pine for. with oak scrub, J.E. Swann (lc?*); Sheridan Co., Black Mt, off Hwy 14, 5-20.viii.1990, pine for., pans cow dung, J.E. Swann (2c?**); Sublette Co., Daniel, 7200', 16-22.viii.1979, carrion, sagebrush steppe, S.&J. Peck (lc?*); Uinta Co., 6mi E Mt. View, 6800', 1-1 l.viii. 1979, sagebrush, S.&J. Peck (2c?**, 19*); Uinta

163 Co., 8mi SE of Evanston, 7100', 30.vii-ll.viii.1979, carrion, sage-grass, riparian, S.&J. Peck(lc?*).

Comments:

Almost half of the specimens examined were Coproica urbana. The slight variations in surstylus shape and high variation in ventral mid basitarsus bristle patterns, a character considered important within Coproica (Collin, 1956; Richards, 1960), made the assessment of the variation in this species exceptionally difficult. C. urbana was divided into four morphospecies in an unpublished MSc thesis (Swann, 1993) but we could find no correlated, discontinuous characters that reliably separate those putative morphospecies. The character states proposed by Swann were either not measurable, or were represented in every long series of C. urbana examined.

The morphological evidence that C. urbana represents a single biological species was supported by limited mitochondrial DNA evidence, as 5 specimens "barcoded"

(sequenced for Cytochrome Oxidase 1) showed very little variation, falling within 1% of each other. Four specimens were from Ontario and 1 from Texas, indicating that the

Texas specimen is the same species as the Ontario specimens. Additional sequence data for C. urbana specimens, especially from other regions, would be of interest, but the specimens on which this revision was based were collected several years ago, mostly in pan traps or dung traps, rendering the extraction of molecular data difficult to impossible.

Further testing of the limits of C. urbana using molecular data will require the collection of fresh C. urbana.

Label data for all specimens of C. urbana examined over the course of this study are not included here, but data from specimens that were dissected, illustrated, used for

164 gathering morphometric data, borrowed from other institutions, or for new state, province and country records, are included.

165 Coproica vagans (Haliday, 1833) (Figures: 13,288-295)

Distr.: Afrotropical: Ethiopia, South Africa, Tanzania, Zaire; Australasian/Oceanian: Australia (QLD), Guam (USA), Hawaii (USA); Nearctic: Canada (AB, BC, NS, ON, SK), USA (AR, AZ, CA, CO, CT, DC, FL, ID, MI, MO, NC, NJ, NM, OK, OR, PA, SD, TX, UT, WA, WY); Neotropical: Argentina, Bermuda, Bolivia, Chile, Costa Rica, Jamaica, Mexico (BCS, MEX); Oriental: Taiwan; Palaearctic: Afghanistan, Algeria, Andorra, Austria, Azerbaidjan, Belgium, Bulgaria, Canary Is. (Spain), Cyprus, Czech Republic, Denmark, Egypt, Finland, France (incl. Corsica), Georgia, Germany, Great Britain, Greece (incl. Crete, Thira), Hungary, Iceland, Italy (incl. Sardinia), Ireland, Israel, Japan, Latvia, Macedonia, Malta, Mongolia, Netherlands, Norway, Poland, Romania, Russia (CET, NET, SET), Saudia Arabia, Slovakia, Spain (incl. Balearic Is.), Sweden, Switzerland, Tadjikistan, Tunisia, Turkey, Yugoslavia (Serbia); South Atlantic: Saint Helena, UAE. Borborus vagans Haliday, 1833: 178 [both sexes ?]. Type locality: Ireland, Downshire, Holywood. STs (NMID). Limosina vagans. - Haliday, 1836: 332 [generic combination, redescription]. Limosina (Coprophila) vagans. - Duda, 1918: 210-214 [subgeneric combination, redescription, illustr.]; Duda, 1938: 167 [redescription, illustr.]. Leptocera (Coprophila) vagans. - Duda, 1925: 205 [generic combination]. Leptocera (Heteroptera) vagans. - Richards, 1930: 307 [subgeneric combination]. Leptocera vagans. - Goddard, 1938: 250-251 [puparium, illustr.]. Limosina (Coproica) vagans. - Frey, 1936: 110 [subgeneric combination]. Leptocera (Coproica) vagans. - Richards, 1960c: 201, 205 [subgeneric combination, taxonomic notes, key]; Richards, 1967b: 16 [Neotropical catalog]; Tenorio, 1968: 196, 201 [diagnosis, male genitalia, illustr.]; Richards, 1980: 618 [Afrotropical catalog]. Coprophila vagans. - Vanschuytbroeck, 1951b: 20 [generic combination]. Coproica vagans. - Frey, 1949: 36 [generic combination]; Papp, 1973c: 26 [diagnosis, key, illustr.]; Hackman, 1977: 401 [Oriental catalog]; Papp, 1984: 82 [Palaearctic catalog]; Marshall, 1989b: 603 [Australasian/Oceanian catalog]; Lachmann, 1996: 759-768 [morphology, internal genitalia, copulation, illustr.]; Rohacek, 2007b: 116 [delete the distribution of Madeira (Portugal)]; Gatt, 2008: 700 [new record from UAE, photograph]. Limosina opacula Stenhammar, 1855: 389 [both sexes]. Type localities: Sweden, "Ostrogothia" and "Gottlandia". LT male, designated by Rohacek, 1983e: 159 (UZIU) [no specimen of type series is labelled and, therefore, no definite type locality can be selected on the basis of lectotype]. Limosina albipennis Rondani, 1880: 41 [both sexes ?]. Type locality: Italy, nr. Parma. STs (MZUF). - Duda, 1918: 210 [synonymy]. Limosina (Coprophila) vagans var. flava Duda, 1918: 210 [male, illustr.]. Type locality: Tunisia, Gafsa. STs males (ZMHB). - Papp, 1979a: 104 [synonymy]. Coprophila vagans var. flava. - Vanschuytbroeck, 1951b: 20 [generic combination]. Leptocera (Coproica) vagans var. flava. - Richards, 1960c: 201, 205 [generic

166 combination, key]. Limosina vagans var. rufa. - Collin, 1949: 201 [lapsus]. - Richards, 1960c: 205 [synonymy].

Body length 1.3 - 2.3mm. Body brown. Face light brown, frons brown. Interfrontal bristles in 4-5 subequal pairs, postocellar bristles convergent, 1 pair of small postocellar setulae. Eye small, eye to gena ratio 1.3:1 - 1.5:1; 6 genal setae, anterior seta 2 times length of posterior bristles. Scutellum with 20-25 discal setae arranged in 3 loose rows between basal 1/5 and basal 3/5, posterior row of setae approximately 1.5 times length of other discal setae; 3 lateral setae between larger anterior and posterior marginal scutellar bristles. Katepisternum with 2 long, subequal dorsalbristles, anterior and posterior bristles each approximately 3/5 and 4/5 as long as distance between dorsal margin of katepisternum and wing base respectively. Legs and fore coxa brown to light brown.

Mid tibia with small additional anterodorsal bristle at 1/10, and proximal posterodorsal bristles present at 1/5 and 2/5. Mid bastitarsus with bold anteriory setae sub-basally and at 1/2, a sub-basal ventral pair of bristles, a posteroventral sub-basal bristle approximately

1.3 times length of anteroventral bristle, an anteroventral bristle at 2/5 and approximately as long as 2 times width of mid basitarsus, an anteroventral bristle at 4/5, and small posteroventral bristles at 2/5 and 3/5. Second costal sector 1.1-1.7 times length of third costal sector. Costal bypass short, terminating within 5 vein widths of JLH-S. Anal vein with distinct angulate bend.

Male Terminalia:

Sternite 5 rounded rectangular; surface bristles enlarged laterally. Synsternite 6+7 straight medially. Epandrium with enlarged pair of dorsolateral bristles; subanal plate present, narrow; cercus fused to epandrium, ventral margin with broad rounded medial

167 processes, and with 1 outstanding bristle. Surstylus with 2 small dorsally curved processes separated by a concave invagination, 3 or more enlarged anteroventral bristles, and several small bristles along ventral margin. Lateral sclerites rounded, with blunt subapical dorsal process, an apicoventral posteromedially deflected process; central sclerite fused with lateral sclerites on apicoventral process. Membranous sheets broad.

Female Terminalia:

Tergite 8 long laterally and short dorsally. Tergite 10 pollinose, narrowly fused to cerci posterolaterally; cerci rounded and pollinose, apical bristles long and sinuate. Steraite 7 broad, rounded. Sternite 10 not wrapped posteriorly around tip of abdomen, bulbous, with medial and lateral pairs of enlarged bristles. Spermathecae spherical; duct length approximately 2 times length of spermathecal bulb.

Specimens examined:

CANADA: AB: Writing-On-Stone Prov. Pk., Sedges, 16-24.viii.1990, M. Klassen (1$); Writing-On-Stone Prov. Pk., 30.viii-10.ix.1990, malaise, T.S. & M. Klassen (1$); Hailstone Butte, 601m, W of Nanton, 21-23.vii.1987, dung pans, alpine creek, sedge, willow moss, wet, S. Marshall (1$); Jumping Pound Ck., 7.viii.l980, dung vac, S.A. Marshall (1?). BC: Haney, UBC research forest, 3.vii.l988, sweep near stream, S.A. Marshall (1#*). NS: Cape Breton Highlands Nat. Pk., Lone Shielding, PG861731, 19.viii.1983, pan tps., M.J. Sharkey (1 $). ON: Algonquin Prov. Pk., Swan Lk. Stn., Scott Lk. Survey, 10-17.vii.1994, A5 Sphagnum pans (1 $); Essex Co., East Sister Is. Nat. Res., 41° 49'N, 82° 51'W, 30.vii.2003, S.A. Marshall (1$: debu00238524); Essex Co., Point Pelee Nat. Pk. De Laurier Tr., 8.vi.2000, black light, E. Reichert & O. Lonsdale (1$: debuO 1001031); Kent Co., Rondeau Prov. Pk., Spicebush Tr., 42 18'9"N, 81 51'6"W, 15.viii-7.ix.2003, Carolinian for., malaise, Marshall et al. (1$: debuOl 139007); Kent Co., Rondeau Prov. Pk.., Visitor Centre, 42 46'50N, 81 50'38W, 16-17.vi.2003, MV. light, M. Buck (1?*: debuO 1125162*); Ottawa, 6.ix.l984, swept from liquid manure pit, {\S, 1$). SK: Grant N, 08.viii.1998, dead cow, C. Christopherson (l^-wing, 1$); Grant E, Grasslands Nat. Pk., 26.viii.1998, dead cow H, leg, C. Christopherson (1$); same as previous except dated 08.viii.1998 (2& 1$). COSTA RICA: Puntarenas, 2000m, 12.vii.1995, S.A. Marshall (1$). JAMAICA: St. Catherine, 28.vii-l.viii. 1974, dung under trees in horse pasture, Peck (6$). MEXICO: Baja California Sur, 5 mi E of Santa Marta, 10100', km 8.5, 29.viii-4.ix. 1971, fir for., human dung, A. Newton (1$, 1$). SAUDIA ARABIA: Quwayzah, E of Jeddah, 24.X.2000, around camel urine, N. Gadallah (9$**: debuOO 185776*, 00185777*, 00185778, 00185779, 00185780,

168 00185781, 00185782, 00185783, 00185784; 4$: debu00185785, 00185786, 00185787, 00185788). UNITED STATES OF AMERICA: AZ: Apache Co., Alpine Luna Lk., 7900', 9-14.vii.1979, pine-meadows, S.&J. Peck (1$); Buckeye, Buckeye Rec. Area, 21.iv.1979, K.N. Barber (1$); Cochise Co., Chiricahua Mt., 6mi ESE of Portal, 4150', ix.1970, desert grassland, human dung, A. Newton (lc?); Cochise Co. Chiracahua Mts., 7000', 15.vii.1978, dung, O. Kukal, (1$); Cochise Co., Huachuca Mts., Carr Canyon, 7100', ix.1970, oak & evergreen, human dung, A. Newton (3c?); Coronado Co., Bonito Pk., near Flagstaff, 5-9.viii.1984, dung tp., B.V. Brown (lc?); Flagstaff, Oak "ok. osn."(?), 5900', 17-25.vii.1979, S.&J. Peck (1$); Madera Canyon, 14.V.1977, dung, J. Helaud (1$); same as previous except dung, dung tp. (1$); Navajo Co., 15mi SE of Holbrook, 5300', 14-16.vii.1979, grassland carrion tps., S.&J. Peck (5$); Santa Cruz Co., 7mi WNW of Nogales, Calabasas Canyon, 3850', ix.1972, desert grass, dung, A. Newton (1$). AR: Wash Co., 3mi S of Devils Den St. Pk., 28-31.V.1979, oak-hickory, S.&J. Peck (1$). CA: Marin Co., San Anselmo, 17.iv.1980, S.A. Marshall (1$); Santa Clara Co., Del Peurto Rd. (Rdl30), 28.V.1992, J. Skevington (1$). FL: Okaloosa Co., 4.5mi N of Holt Blackwater R. St. For., Fla. A&M Res. Stn., 15-17.vi.1978, pig dung, L. Stange (Id1*). NM: Bernalillo Co., Isleta, 4900', 16-20.vi.1979, floodplain for., carrion, S.&J. Peck (6c?***, 5$); Catron Co., 6mi S of Luna, 8000', 9-14.vii.1979, pond, pine for., S.&J. Peck (1$); Catron Co., 5mi W of Luna, San Francisco River Pond, 7400', 9- 14.vii.1979, pine-meadows, S.&J. Peck (lc?); Eddy Co., 30mi WSW of Carlsbad, Sitting Bull Falls, 4600', 23-27.vii.1975, dung 15, S. Peck (lc?); Lincoln Co., lmi N of Sierra Blanca Ski, 11000', 10-26.vi.1979, alpine grass meadow, carrion, S.&J. Peck (16c?, 19$); Sandoval Co., Bernalillo, 5100', 20-24.vi.1979, grassland carrion, S.&J. Peck (6c?, 10$); Santa Fe Co., 14mi NE of Santa Fe, 9600', 18.vi-3.vii.1979, aspen w/stream, S.&J. Peck (6c?, 4$); Santa Fe Co., Santa Fe, Baldy, 10800-11800', 19.vi-4.vii.1979, spruce- fir-tundra, S.&J. Peck (20c?**, 28$); Socorro Co., 20mi W of Socorro, Water Canyon, 7000', 28.vi-7.vii. 1979, mixed "mesic" for., S.&J. Peck (10c?, 6?*); Socorro Co., S. Baldy Pk., 10400', 20mi W Socorro, 28.vi-7.vii.1979, alpine meadow, S.&J. Peck (lc?); Torrance Co., 6mi SW of Manzano, Red Can. cpgd, 75-9000', 17-30.vi.1979, spruce-fir litter, S.&J. Peck (20c?*x4, 24$, 1?); same as previous except 7500, malaise pine fir for. (lc?); Valencia Co., 2mi W of Los Lunas, 4900', 16-30.vi.1979, desert grassland carrion tps., S.&J. Peck (33c?**-illustr, 52$**-illustr). NC: Swain Co., Great Smokey Mts. Nat. Pk., Ravens Fork/ Oconoluftee River confluence, 4.V.2001, W. Reeves (1$: debuOO 166807). OK: Comanche Co., Wichita Mts. Refuge, 5-7.vi.1979, shortgrass prairie, S.&J. Peck (2?); Lattimer Co., 5mi W of Red Oak, 5-1 l.vi. 1977, dung tp., K. Stephen (2$). TX: Cameron Co., Laguna Atascosa Nat. Wildlife Refuge, vii.1972, thorn forest, dung, A. Newton (1$); Randall Co., 3420', Palo Duro Can. St. Pk., 7-9.vi.1979, mesquite-prairie, S.&J. Peck (2c?, 7$); Randall Co., Palo Duro Canyon St. Pk., 26.v.- 7.vi.l991, pans baited with Opuntia, J.E. Swann (9c?, 1$). UT: Dagget Co., Dutch John, 6300', 2-8.viii.1979, pinyon juniper, woodland, carrion, S.&J. Peck (4c?**, 5?*, 2?); Duchesne Co., Mirror Lk., 10300', 30.vii-12.viii. 1979, spruce fir meadow, carrion- malaise, S.&J. Peck (6$); Summit Co., Henry's Fork Park, 9000', l-10.viii.1979, meadow w/willow malaise, S.&J. Peck (1$); Summit Co., NW side of Gilbert Peak, 11500-12500', l-10.viii.1979, tundra carrion, S.&J. Peck (2c?, 6$). WY: Big Horn Co., lOmi E of Lovell, 3800', 17-26.viii.1979, sagegrass-barren carrion, S.&J. Peck (1$); Red Gulch Rd. off Hwy.14, 5-19.viii.1990, near shell pans in grass hummock in desert area,

169 cow dung, J.E. Swann (3$); same as previous except near shell pans in desert area, cow dung (3<5\ 2$); near shell pans in barren area near cottonwoods (2$, 4$); near shell pan tps. in barren area near cottonwoods, cow dung (l<3); Uinta Co., 6mi E of Mt. View, 6800', 1-1 l.viii. 1979, sagebrush, S.&J. Peck (2$**, 7?); Uinta Co., 8mi SE of Evanston, 7100', 30.vii-ll.viii. 1979, carrion, sage-grass, riparian, S.&J. Peck (2$*, 1$). RUSSIA: Siberia Novosibirsk Region, Chany Lk. Bio. Stn., 5.vii.l991, swept over dirty mud/ cow dung/ algae, S.A. Marshall (lc?).

170 References

Carles-Tolra, M. 1990. New species and records of Sphaeroceridae (Dipt.) from Spain. Entomologist's Monthly Magazine. 126: 33-46.

Collin, J.E. 1956. Some New British Borboridae (Diptera). J. Soc. Brit. Ent. 5: 172-178.

Duda, O. 1918. Revision der europaischen Arten der Gattung Limosina Maquart (Dipteren). Abh. zool.-bot. Ges. Wein. 10: 1-240.

Duda, O. 1925. Die aussereuropaischen Arten der Gattung Leptocera Olivier = Limosina Macquart (Dipteren) mit Berucksichtigung der europaischen Arten. Arch. f. Naturag. 90A (11)1924: 5-215.

Duda, O. 1929. Die Ausbeute der deutschen Chako-Expedition 1925/1926 (Diptera). VI. Sepsidae, VII. Piophilidae, VIII. Cypselidae, IX. Drosophilidae und X. Chloropidae. Konowia. 8(1): 33-50.

Duda, O. 1938. 57. Sphaeroceridae (Cypselidae). In Lindner, E. (ed.): DieFliegen der palaearktischen Region. E. Schweizerbart.sche Verlagsbuchhandlung, Stuttgart. Vol.6, 182 pp.

Enderlein, G. 1938. 60. Die Dipterenfauna der Juan-Fernandez-Inseln und der Oster- Insel. Almqvist & Wiksell Printing Co. 643-680.

Frey, R. 1945. Tiergeographische Studien uber die Dipteren-fauna der Azoren. I. Verzeichnis der bisher von den Azoren bekannten Dipteren. Soc. Scient. Fenn., Comm. Biol. VIII. 10: 1-114.

Goddard, W.H. 1938. The description of the puparia of fourteen British species of Sphaeroceridae (Borboridae, Diptera). Trans. Soc. Br. Ent. 5: 235-258.

Gordh, G. & Hall, J.C. 1979. A critical-point drier used as a method of mounting insects from alcohol. Entomol. News. 90: 57-59.

Goloboff, P.A., Carpenter, J.M., Arias, J.S., & Miranda Esquivel, D.R. 2008a. Weighting against homoplasy improves phylogenetic analysis of morphological data sets. Cladistics. 24: 758-773.

Goloboff, P.A., Farris, J., & Nixon, K. 2008b. TNT: a free program for phylogenetic analysis. Cladistics. 24: 774-786.

Greenberg, B., Valela, G., Bornstein, A., & Hernandez, H. 1963. Salmonellae from flies in a Mexican slaughterhouse. Am. J. Hyg. 77: 177-183.

171 Hackman, W. 1969. A review of the zoogeography and classification of the Sphaeroceridae (Borboridae, Diptera). Notulae Entomologicae 49: 193-210.

Hafez, M. 1939. The life history of Leptocera digitata Duda. Bulletin de la Societe Fouadler d.Entomologie, Le Caire 23: 326-332.

Hafez, M. 1948. Ecological and biological observations on some coprophagus Sepsidae (Diptera). Ibid. 31:307-312.

Hafez, M. 1949. Observations on the biology of some coprophagous Borboridae. Proc. Roy. Entomol. Soc. Lon. (A), 24: 1-5.

Harris, R.L. & Oliver, L.M. 1979. Predation of Philonthus flavolimbatus on the horn fly. Env. Ent. 8:259-260.

Hayashi, T. 1986. Studies on the sphaerocerid flies of synanthropy and hygenic importance in Japan (Diptera) I. The key to species of the flies. Jpn. J. Sanit. Zool. Vol. 37, No. 2: 113-120.

Hayashi, T. 1986. Studies on the sphaerocerid flies of synanthropy and hygenic importance in Japan (Diptera) II. Records of the forty species. Jpn. J. Sanit. Zool. Vol. 37, No. 3: 193-204.

Hayashi, T. 1991. The genus Coproica Rondani from Pakistan (Diptera, Sphaeroceridae). Jpn. J. Sanit. Zool. Vol.42, No. 3: 235-240.

Hayashi, T. 1992. New records of the genus Coproica from the Australian region (Diptera, Sphaeroceridae). Jpn. J. Entomol. 60(2): 332.

Hayashi, T. 1995. New records of the genus Coproica Rondani (Diptera: Sphaeroceridae) from Japan. Jpn. J. Sanit. Zool. 46: 229-233.

Hayashi, T. 2001. Distributional notes on Coproica rufifrons Hayashi (Diptera: Sphaeroceridae) in the Australasian and Oceanian regions. Med. Entomol. Zool. 52:253-254.

Hayashi, T. 2005. The genus Coproica Rondani (Diptera: Sphaeroceridae) from the Ryukyu Islands, Japan. Med. Entomol. Zool. 56: 207-210.

Lachmann, A.D. 1990. Courtship behaviour and mating in two cow dung inhabiting Sphaerocerid flies (Diptera: Sphaeroceridae). Ethology. 86: 161-169.

Lachmann, A.D. 1991. Vergleichende Unterschung zum Lebenszyklus der kuhdungbewohnenden Sphaeroceridenarten Chaetopodella scutellaris (Haliday, 1836) und Coproica lugubris (Haliday, 1836). Dtsch. Ent. Z. 38: 197-210.

172 Lachmann, A.D. 1996. Copulation and engagement of male and female genitalia in five Coproica Rondani species (Diptera: Sphaeroceridae). Ann. Entomol. Soc. Am. 89: 759-769.

Lachmann, A.D. 1997. Sperm transfer during copulation in five Coproica species (Diptera: Sphaeroceridae). Eur. J. Entomol. 94: 271-286.

Lachmann, A.D. 1998. Sexual receptivity and post-emergence ovarian development in females of Coproica vagans (Diptera: Sphaeroceridae). Phys. Ent. 23: 360-368.

Macquart, J. 1835. Histoire naturelle des Insectes. Diptty an Collection des suites Insectes. Vol. 2, 710 pp., N.E. Roret, Paris.

Maddison, W.P., Donoghue, M.J. & Maddison, D.R. 1984. Outgroup analysis and parsimony. Syst. Zool. 33: 83-103.

Malloch, J.R. 1913. Two new species of Borboridae from Texas. Proc. Ent. Soc. Wash. 15: 135-136.

Marshall, S.A. 1982. A revision of Halidayina Duda (Diptera: Sphaeroceridae). Can. Ent. 114:841-847.

Marshall, S.A. 1987. Systemastics otBitheca, a new genus of New World Sphaeroceridae (Diptera). Syst. Ent. 12: 355-380.

Marshall, S.A. 1991. Rudolfina digitata sp. nov., a new Nearctic sphaerocerid with a disjunct alpine-arctic distribution. Can. Ent. 123: 621-626.

Marshall, S.A. 1997. Sphaerocerid Flies (Diptera: Sphaeroceridae) of the Yukon. In: Danks, H.V. and Downes, J. A. (eds.). Insects of the Yukon. Biological Survey of Canada (Terrestrial ), Ottawa, (p. 663-685).

Marshall, S.A., Buddie, C, Sinclair, B., & Buckle, D. 2001. Spiders, flies and some other arthropods of the Fathom Five Islands and the upper Bruce Peninsula, pp 191-229 in: M. Munawar (Ed.). Biology of the Fathom Five Islands.

Marshall, S.A. & Langstaff, R. 1998. Revision of the New World species oiOpacifrons Duda (Diptera, Sphaeroceridae, Limosininae). Contrib. Sci. Nat. Hist. Los Angeles Co. All: 1-27.

Marshall, S.A. & Peck, S. B. 1984. Distribution of cave-dwelling Sphaeroceridae (Diptera) in eastern North America. Proc. Entomol. Soc. On. 115: 37-41.

173 Marshall, S.A. & Richards, O.W. 1987. 93. Sphaeroceridae. In McAlpine J. F., Peterson, B. V., Shewell, G. E., Teskey, H. J., Vockeroth, J. R., Wood, D. M. (eds): Manual ofNearctic Diptera. Agriculture Canada Monograph No. 28, Vol. 2, vi+ 675 1332 pp. (p. 993-1006). Minister of Supply and Services Canada, Ottawa.

Okely, E.F. 1974. Description of the puparia of twenty-three British species of Sphaeroceridae (Diptera, Acalyptratae). Trans Roy. Entomol. Soc. Lon. 126(1): 41-56.

Papp, L. 1973. Sphaeroceridae (Diptera) from Mongolia. Acta Zool. Hung. 19: 369-425.

Papp, L. 1975. Ecological data on flies breeding in dung heaps. Acta Zool. Hung. 21: 425-433.

Papp, L. 1979. A contribution to the knowledge on the species of the genus Coproica Rondani, 1861 (Diptera: Sphaeroceridae). Opusc. Zool. Budapest. XVI, 1-2: 97- 105.

Papp, L. 1992. Fly communities in pasture dung: some results and problems (Diptera). Acta Zool. Hungarica. 38 (1-2): 75-88.

Papp, L. 1999. Species of 22 Diptera families of the Aggtelek National Park. In Mahunka, S. (ed.): The Fauna of the Aggteleg National Park 2, pp. 525-546, Hungarian Natural History Museum, Budapest.

Papp, L. 2008. A review of the Old World Coproica Rondani, 1861 (Diptera: Sphaeroceridae), with descriptions of twelve new species. Acta Zool. Hung. 54 (Suppl. 2):l-45.

Papp, L. 2008. 2008. New genera of the Old World Limosininae (Diptera: Sphaeroceridae). Acta Zool. Hung. 54 (Suppl. 2): 47-209.

Papp, L. & Plachter, H. 1976. On cave-dwelling Sphaeroceridae from Hungary and Germany (Diptera). Ann. Hist. Nat. Mus. Nat. Hung. 68: 195 207.

Pefia, C. & Wahlberg, N. 2008. Prehistorical climate change increased diversification of a group of butterflies. Biol. Let. 4(3): 274-278.

Richards, O.W. 1930. The British species of Sphaeroceridae (Borboridae, Diptera). Proc. Zool. Soc. Lon. 1930: 261-345.

Richards, O.W. 1938. Diptera Sphaeroceridae (Borboridae, Cypselidae). Mission scientifique de l'Omo. Mem. Mus. Nat. Hist. Paris. 4(40): 381-405.

174 Richards, O.W. 1960. On two N. American species of Leptocera Oliv., subgenus Coproica Rdi., with a review of the subgenus (Dipt., Sphaeroceridae). Ann. and Mag. Nat. Hist. Ser. 13, 2/16: 199-208.

Rohacek, J. 1982. A monograph and re-classification of the previous genus Limosina Maquart (Diptera: Sphaeroceridae) of Europe: Part I. Beitr. Ent. Berlin. 32: 195- 282.

Rohacek, J., Marshall, S.A., Norrbom, A.L., Buck, M., Quiros, D.I., & Smith, I. 2001. World catalog of Sphaeroceridae (Diptera). Slezske zemske museum, Opava, 414 pp.

Rondani, C. 1861. Dipterologiae italicae prodromus. Vol. 4. Species italicae ordinis dipterorum in genera characteribus definita, ordinatim collectae, methodo analitica distinctae, et novis vel minus cognitis descriptis. 174 pp., Parmae.

Sabrosky, C.W. 1948. A dipteron new to North America. Proc. Entomol. Soc. Wash. 50: 85.

Smith, LP. & Marshall, S.A. 2004. A Review of the New World genus Pterogramma Spuler and a revision of the Pterogramma sublugubrinum group (Diptera: Sphaeroceridae: Limosininae). Contrib. Sci. Nat. Hist. Los Angeles Co. 499: 1- 163.

Spuler, A. 1925. Studies in North American Borboridae (Diptera). Can. Ent. 57: 99-104, 116-124.

Swann, J.E. 1993. Systematics of the new world Coproica Rondani (Diptera: Sphaeroceridae). University ofGuelph, M.Sc. Thesis. 222p.

Vanschuytbroeck, P. 1950. Dipteres Sphaeroceridae du Musee du Congo beige. Annales du Musee du Congo Beige, Tervuren, Serie in 8o (Sciences Zoologiques), 5: 5-46.

Watrous, J.E. & Wheeler, Q.D. 1981. The out-group comparison method of character analysis. Syst. Zool. 30: 1-11.

Wheeler, T.A. & Swann, J.E. 1994. Case 2917. Coproica Rondani, 1861 and Ischiolepta Lioy, 1864 (Insecta, Diptera): proposed conservation of usage by the designation of Limosina acutangula Zetterstedt, 1847 as the type species of Coproica. Bull. Zool. Norn. 51:316-319.

Wheeler, T.A. & Swann, J.E. 1996. Opinion 1839: Coproica Rondani, 1861 and Ischiolepta Lioy, 1864 (Insecta: Diptera): conserved by the designation of Limosina acutangula Zetterstedt, 1847 as the type species of Coproica. Bull. Zool. Norn. 53(2): 136-137.

175 .O-.i-i O. 0:'Pi©^©:'i-<:'-r© ;•<£>.• ©!<$!'©!;&>- C";-©1»--H*-4J*-t ?-»-*•;©©'; c-i©:© ~&\ ©

©/; ^H ::Q •

o i©!o |p^ OSHO irr*i wl*^.;HH\oi:*ii;H5*!* ©; o ^ ^ :^ ©|0;0 ;j»rtj© [Hi-< l^i f-t^?'-':t?(:oi^lH ;©]• O'-H I:I:

O jijSjl O :£j8 O j«%0 [Hi oM|i O ia&! 0|<3rJ0 jfS o so:; -H :o; o so: o J©;; -H

olo|o:lo^og:oi|oii(o;o;o||!oi9o:0:0^oii;o

ojp]oliio^rt|i|io|y:oi|*o?9ijO!^o:s!o^|oj«^o.'|elo

§H Mi!

©if© i*i; T-H i<^:rH ;^S>H »"**:©: •"* ii**; »"H ii*"* © iHi fHiiHjTH ii: © >ii© ©hi »"* ^O^ *"* i***; ©

, S»| ^ ijcsu c-* i?©i c-v^r* ** *-H iOiiM isj:i©>*-^i-t :t*;(S

iof o o © «"H *-H 1-H i-H © O ii © © HH OH © W ^H

&° M'o, o o o o o t-H © rH © O <-* O © t-l H © © ©On o © o o o o O —I O © o © o o © © o © o © r< -H © o — OHO (jjj1 ^j ^=G5'- m o o: © © © © © — Ijol; :i o!li?o ibioi©; i:i ill « Hii ill SB Hi si* 11 U lit 2:1 &l£i;£l IIP uidliolid

176 e QOOrHQ^-0<-iOOOOOOOOOOOOOOQe"OOi-lO

: O C) *H r-H i-l ^v© © © © © ^H :*H:i ©<•-* ^H :© ©•-<: O © O »H C" :©'.^H::'*H O

OOrtrt—IC^. OOO—lOMtSMrH.-HOOOOrtOOC'. o^HrtO

ID OOO-H^t^. OOOOO-HO-HOC^. OOOOOOrtC^. mO»HO

3 0©»-i©i—ic^»-40©t-i©©©i-Hi-(e"©©©i-i©T-i©C" •-*©©©

OOOOOe-'OOOOOOOOOe-

0O»H^H*-tc^. ©^©^HO©©O»-IC^ ©©©©©©© c^-,-i.-i»-4© e C-OOOOC-OOOOOOOIHOC-OOI-IOO'H'NC-OOOO

« 00*-*^HOC^. OMOOOO€SOOC^. OOOOOOOC^. OOC^O M oe eoeor4c-oooo<-ioooooooaoO'Hoe">Hooo

© 0:0 ^^:i-H c^« © © © © »-H © © © iO O r* ©:©© ^ ~H:© c^:*-4i© .;*<;© a 000**i-iC~OOOOOOOOOOOOOOOOOc>-<-lO<-<0 in ootstSM^-o^HOi-ii-ioo-HM^-riorlrsooo^-rsoMO s ^T-H©©OOl-HO»-IOO^H*-i^H©C^^**-H»-li-l^*i-lT-ii-li-tf-l»-lO a OrfOOOOOO^OOOOOOOOOOOOOOOOOOO

M OONOHriONxNOOOONOOOeOOOOOOOriO

O

S0*^»-t»-ti-H©i-t©»-Hi-li-^^-<»^»-*»^©©»-H0©f—( *"* © »H i-H »-( i-H

OS

IS OHOHHHOMffOHOOOOOT^HniHOOrtHHO

^-OOOOC-'OOOOOOOOOOi-ii-ii-i^-if-ioOc^i-ii-iOi-i ihliiii(ltailtayirii« ouuuuuuuyuuduuouuduuuuuuuuuu

177 - Outgroup •C. serra -D-D- C. digitata dHhHHtt^ 3 Cacutangula c 3 i-WH+H 2 P rO-HH-O- 11 J7B) 3QU) 3SU) « Cpappi 3 on| i D-0- C. mitchelli Q .2 HH-H- Cpusio § a

•—HHHh r C, lugubris •^HHh -C. coreana 1 D-tHHHHl- • C. novjail J il 24 36 27 3S(2) 40 riHHH}- • C himcula ) i>i au»4oi39a) 4 ! rHHHHHHJ-f rahrfMki L+HH HHM-0- 4HHHHHHHK. wtulosa 5 16 17 26 3SUI 16 41 4S, 1 i-HHHH HHHHH] C. fcrruginata II 21(31 34 3S 47{2! -a-f-Q-B-Ch • C.vagans HHh i-HHJ-0-0 ; C. rjdldp-iqo^ 7 12 1; 4soi as "—D-D— 1—HHH-Q- 1C. hirhita LHHHl- -0- • C. rufifrons ! Lltfl-H MH>- * C. IdcteipcnniH -B-0-D-D-D- G testudinea 1—D-0- C. hirtuloidea 3 l s HmHH at [ «> i+HHH —M- C. bifurcata .2 4HH u L-HH 22 47(2) 49 C. emarginata a. 1 270) 31(2) 390) * C. diabolica § HHH-H C. cacti -p LHHHH I—D— Curbarta wm-\ 390)

•HH1 Cbrachystyia

Figure 1. Most parsimonious tree generated from the analysis {Length = 321, CI = 0.399, Rl = 0.651). I = Synapomorphy/autapomorphy, 0 =homplasy, 1 = reversal.

178 - Outgroup

- C.coreana

• C. serra 98 - C.digitata \-2- r "C - C. acutangula L-4H- • C. pappi • C. mite hell i L47-[ - C. pusio 34-C H«H - C. lugubris - C.vagans

- C. novacula

- C.hirticula

• Crohaceki h-t - C. setulosa -c • C ferruginata •-H - Cgalapagos - C.hirtula bt - C mfifrons I-2S-C - C. lacteipennis

- C testudinea

• C. cacti

• C.urbana

- C. braehystyla

. C. bispatha 4^ > C.diabolica - C. emarginata H: C. hirtuloidea 3.-C C. bifurcata

- Outgroup

- C.serra

-C.i dlgitata

n-C acuta ngula

pappi A . c. mitchelli ^ .pUSiO < -c . lugubris "-3H .coreana 2 . novae u la r -C . hirticula -c vagans >-2H I-2H . ferruginata .setulosa •-2H . rohaceki . galapagos «-3- . hirtula . rufifrons

. lacteipennis

. testudinea

. emarginata

. hirtuloidea -c. bifurcata H -C.i diabolica {?u -C. cacti bispatha £ .urbana -Ebraehystyl a

Figures 2-4: 2. Jacknife supports. 3. Bootstrap supports. 4. Bremer supports.

179 Mid tibia

Hind basitarsus

Setulose scutellum Scutellum

Enlarged lateral bristles

Figures 5-6: Coproica acutangulo (Zetterstedt). 5. left lateral. 6. Setulosity on scutellum. Abbreviations: SAB = subapical scutellar bristle, MAB = midapical scutellar bristle.

180 nterfrontal bristles orbital bristles ocellar bristle inner vertical bristle ostocellar bristle inner occipital bristle outer vertical bristle—^j\ T outer occipital bristle postocellar setuiae

dorsal katepisternal bristles anepimeron

9

katepistenum

mid coxa

Figures 7-9: 7. Coproica acutangula (Zetterstedt) head dorsal. 8. Coproica diabolica sp. nov. left katepisternum. 9. Coproica ferruginata (Stenhammer) left katepisternum.

181 Figures 10 -13 left katepisternum. 10. Coproica novacula sp. nov. 11. Coproica setulosa (Duda). 12. Coproica rufifrons (Hayashi). 13. Coproica vagans (Haliday).

182 Figures 14 - 20:14. Coproica acutangula (Zetterstedt) male left fore basitarsus poster­ ior. 15. Coproica acutangula male left hind basitarsus anterior. 16. Coproica acutangula mid tibia dorsal. 17. Coproica bifurcata sp. nov. mid tibia dorsal. Ml.Coproica ferruginata (Stenhammer) mid tibia dorsal. 19. Coproica lugubris {Haliday) mid tibia dorsal. 20. Cop­ roica rufifrons (Hayashi) mid tibia dorsal.

183 21 22 23 24

Figures 21 - 28 Mid basitarsus ventral. 21. Coproica bifurcata sp. nov. 22. Coproica bra- chystyla sp. nov. 23. Coproica hirticula Collin. 24. Coproica lugubris (Haliday). 25. Copro­ ica mitchelli (Malloch). 26. Coproica novacula sp. nov. 27. Coproica rufifrons (Hayashi). 28. Coproica setulosa (Duda).

184 costal sesloti costal sector 2 :EMil l 29 ^^«^SsW

/

y? aluL

/ "**--_ bulla angulate _.. analveirv&j / / ES / / t wing fold ? EMS

cell r 2+3 R2+3

30 ^'^Wa^v*

CB-O"' crossvein r-m

M-- ms crossvein dm-cu"

aumpveMQiA, ^(§$$1*

Figures 29 - 30 Coproica acutangula (Zetterstedt). 29. Male left wing, dorsal. 30. Fe­ male left wing, dorsal. Abbreviations: CB = costal bypass, EMS = enlarged microset- ulae, ES = enlarged setulae along posterior margin of male hind wing.

185 Figures 31-32 Coproica acutangula (Zetterstedt). 31. Male terminalia, posterior. 32. Male terminalia, left lateral. Abbreviations: AA = aedeagal apodeme, C = cer- cus, DLB = dorsolateral bristle, Ep = Epandrium, Hyp = hypandrium, Pg = postgo- nite, Ss = surstylus.

186 Figures 33 - 35 Coproica acutangula {Zetterstedt). 33. Aedeagus and associated structures with postgonites removed. 34. Male sternite 5 and synsternite6+7. 35. Left postgonite. Abbreviations: AA = aedeagal apodeme, 8a = basiphallus, CS = central sclerite, DP = dorsal process, EA = ejaculatory apodeme, LS = lateral sclerite, MS = membranous sheets, PP = posterior process, PVC = posteroventral crest, S5 = sternite 5, S6+7 = synsternite 6+7, TS = tubular sclerite, VS = ventral scl­ erite 187 Figures 36 - 40 Coproica acutangula (Zetterstedt). 36. Female terminalia, ventral. 37. Female terminalia, left lateral. 38. Female terminalia, ventral. 39. Female stemite 8. 40. Spermathecae. Abbreviations: C = cercus, S7 = sternite 7, S8= stemite 8, Si0 = stemite 10, T8 = tergite 8, T10 = tergite 10.

188 41 f ^'

Figures 41 - 43 Coproica bifurcata sp. nov. 41. Male left wing, dorsal. 42. Male terminalia, left lateral. 43. Male terminalia, posterior. Abbreviations: C = cercus, Pg = postgonite, Ss = surstylus.

189 Figures 44 - 47 Coproica bifurcata sp. nov. 44. Left postgonite. 45. Right postgonite. 46. Aedeagus and associated structures with postgonites removed. 47. Male sternite 5 and synsternite 6+7. Abbreviations: CS = central sclerite, LS = lateral sclerite, MS = membranous sheets, S6+7 = synsternite 6+7, TS = tubular sclerite.

190 Figures 48 - 52 Coproica bifurcata sp. nov. 48. Female terminalia, dorsal. 49. Female te- rminalia, ventral. 50. Female terminalia, left lateral. 51.Femalesternite8. 52.5perma- thecae. Abbreviation: T8 = tergite 8.

191 53

Figures 53 - 55 Coproica bispatha sp. nov. 53. Male left wing, dorsal. 54. Male terminalia, left lateral. 55. Male terminalia, posterior.

192 Figures 56 - 59 Coproica bispatha sp. nov. 56. Left postgonite. 57. Right postgonite. 58. Aedeagus and associated structures with postgonites removed. 59. Male sternite 5 and synstemite 6+7.

193 Figures 60 - 64 Coproica bispatha sp. nov. 60. Female terminalia, left lateral. 61. Fem­ ale terminalia, dorsal. 62. Female terminalia, ventral. 63. Spermathecae (depressions are an artifact of the drying process). 64. Female sternite 8. Abbreviations: AP = ant­ erior process on sternite 8, C = cercus,T8 = tergite 8,T10 = tergite 10.

194 65

66

Figures 65 - 67 Coproica brachystyla sp. nov. 65. Male left wing, dorsal. 66. Male termin­ al, left lateral. 67. Male terminalia, posterior. Abbreviation: C = cercus.

195 Figures 68 - 71 Coproica brachystyla sp. nov. 68. Left postgonite. 69. Right postgonite. 70. Aedeagus and associated structures with postgonites removed. 71. Male stemite 5 and synsternite 6+7. Abbreviation: AP = apical process.

196 Figures 72 - 76 Coproica brachystyla sp. nov. 72. Female terminalia, dorsal. 73. Female term- inalia, ventral. 74. Female terminalia, lateral 75. Female stemite 8. 76. Spermathecae. Abb­ reviation: CS = cereal spines.

197 77 * \

^8 ^^,.v-^s^tf^^ ^.yV^v

\

X.

\ !

Figure 77. Coproica cacti (Richards), male left wing, dorsal. 78. Coproica coreana Papp, male left wing, dorsal.

198 Figures 79 - 80 Coproica coreana Papp. 79. Male terminaia, left lateral. 80. Male ter- minalia, posterior.

199 Figures 81 - 83 Coproica coreana Papp. 81. Left postgonite. 82. Aedeagus and associ­ ated structures with postgonites removed. 83. Male sternite 5 and synsternite 6+7. Abbreviations: CS = central sclerite, LS = lateral sclerite, MS = membranous sheet, PVC = posteroventral crest, TS = tubular sclerite.

200 Figures 84 - 88 Coproica coreana Papp. 84. Female terminalia, dorsal. 85. Female ter- minalia, ventral. 86. Female terminalia, left lateral. 87. Female sternite 8. 88. Sperma- thecae.

201 89 \.

Figures 89 - 91 Coproka diabetica sp. nov. 89. Male left wing, dorsal. 90. Male termina ^f,eraL91-Male,erminalia'P°^

202 Figures 92 - 95 Coproica diabolica sp. nov. 92. Left postgonite. 93. Right postgonite. 94. Aedeagus and associated structures with postgonites removed. 95. Sternite 5 synsternite 6+7. 203 Figures 96-100 Coproica diabolica sp. nov. 96. Female terminalia, dorsal. 97. Female ter- minalia, ventral. 98. Female terminalia, left lateral. 99. Female sternite 8. 100. Spermath- ecae.

204 101 .-dr-i^N****- I 'JliiiifV"'

Figures 101-102 Coproica digitata

205 Figures 103-104 Coproica digitata (Duda). 103. Male terminalia, left lateral. 104. Male terminalia, posterior. Abbreviations: C = cercus, EVB = enlarged ventral bristles, Pg = po- stgonite, Ss = surstylus.

206 Figures 105-109 Coproica digitata (Duda). 105. Left postgonite. 106. Right postgonite. 107. Aedeagus and associated structures with postgonites removed. 108. Spermathecae. 109. Female sternite 8. Abbreviations: CS = central process, LS = lateral sclerite, MS = membranous sheet, PP = posterior process

207 Figures 110-111 Coproica digitata (Duda). 110. Female terminalia, dorsal. 111. Female terminalia, left lateral.

208 112

-..-*. > *,•. \, \ .

Figures 112-114 Coproica emarginata sp. nov. 112. Male left wing, dorsal. 113. Male term- inalia, posterior. 114. Male terminalia, left lateral.

209 Figures 115-119 Coproica emarginata sp. nov. 115. Left postgonite. 116. Right pos- tgonite. 117. Distiphallus and basiphallus. 118.Sternite5andsynsternite6+7. 119. Right surstylus. 210 Figures 120 -123 Coproica emarginata sp. nov. 120. Female terminalia, dorsal. 121. Fe­ male terminalia, ventral. 122. Female sternite 8. 123. Spermathecae (depressions are an artifact of the drying process). Abbreviations: AP = anterior process on sternite 8, ML = membranous lobe.

211 124 x •<

125 126

Figures 124 -126 Coproica ferruginata (Stenhammer). 124. Male left wing, dorsal. 125. Male terminalia, posterior. 126. Male terminalia, left lateral.

212 Figures 127 -129 Coproka ferruginata (Stenhammer). 127. Left postgonite. 128. Aed- eagus and associated structures with postgonites removed. 129. Male sternite 5 and synsternite 6+7.

213 Figures 130 -133 Coproica ferruginato {Stenhammer). 130. Female terminalia, dorsal. 131. Female terminalia, ventral. 132. Female terminalia, left lateral. 133. Spermathecae. Abbr­ eviation: S8 = sternite 8.

214 134 ,v..,.- ^^^i'i^S^^ X-- 'J' * ' *> J t-

135

Figures 134 -137 Coproica galapagos sp. nov. 134. Male left wing, dorsal. 135. Male ster- nite 5 and synsternite 6+7. 136. Distiphallus and basiphallus. 137. Left postgonite.

215 Figures 138 -139 Coproica galapagos sp. nov. 138. Male terminalia, posterior. 139. Male terminalia, left lateral.

216 Figures 140-143 Coproica galapagos sp. nov. 140. Female terminalia, dorsal. 141.Fema leterminalia, ventral. 142. Female terminalia, left lateral. 143. Spermathecae (the depres­ sion is an artifact of the drying process). Abbreviations: C = cercus, S8 = stemite 8, S10 = sternite 10, T8 = tergite 8, T10 = tergite 10.

217 144 -;-•-: -vi^^i^^w^^^AW

145 146

Figures 144 -146 Coproica hirticula Collin. 144. Male left wing, dorsal. 145. Male terminalia, left lateral. 146. Male terminalia, posterior.

218 Figures 147 -150 Coproica hirticula Collin. 147. Spermathecae. 148. Aedeagus and asso­ ciated structures with postgonites removed. 149. Male sternite 5 and synsternite 6+7. 150. Left postgonite. Abbreviations: AT = asymmetrical tab, BP = bifurcate process, LS = lateral sclerite, MS = membranous sheet, VP = ventral process.

219 Figures 151 -154 Coproica hirticula Collin. 151. Female terminalia, dorsal. 152. Female terminalia, ventral. 153. Female terminalia, left lateral. 154. Female sternite 8.

220 155

156 157

Figures 155 -157 Coproica hirtula (Rondani). 155. Male left wing, dorsal. 156. Male ter- minalia, left lateral. 157. Male terminalia, posterior.

221 Figures 158 -160 Coproica hirtula (Rondani). 158. Left postgonite. 159. Aedeagus and associated structures with postgonites removed. 160. Male sternite 5 and synsternite 6+7.

222 163 164

Figures 161 -164 Coproica hirtula (Rondani). 161. Female terminalia, dorsal. 162. Fema­ le terminalia, ventral. 163. Female terminalia, left lateral. 164. Spermathecae. Abbrevia­ tion: 58 = sternite 8.

223 165 x

Figures 165-167 Coproica hirtuloidea (Duda). 165. Male left wing, dorsal. 166. Male ter- minalia, left lateral. 167. Male terminalia, posterior.

224 Figures 168 -172 Coproica hirtuloidea (Duda). 168. Left postgonite. 169. Right postgo- nite. 170. Distiphallusandbasiphallus. 171.Sternite5 synstemite 6+7. 172. Female st- ernite 8. Abbreviations: CS = central sclerite, LS = lateral sclerite, MP = membranous process, MS = membranous sheet.

225 175

Figures 173 -176 Coproica hirtuloidea (Duda). 173. Female terminalia, dorsal. 174. Fe­ male terminalia, ventral. 175. Female terminalia, left lateral. 176. Spermathecae.

226 177 ^^*^^"S**J- ^ --srf"- 2 'ie*kp

/>'•.

Figures 177 -178 Coproica lugubris

227 Figures 180 -183 Coproica lugubris {Haliday). 180. Aedeagus and associated structures with postgonites removed. 181. Female sternite 8. 182. Spermathecae {depressions are an artifact of the drying process). 183. Left postgonite. Abbreviations: CS = central sclerite, LS = lateral sclerite, MS = membranous sheet, PVC = posteroventral crest, SAB = subapical anterior bristle, VS = ventral sclerite, WLP = wing-like process.

228 Figures 184-185 Coproica lugubris

229 Figures 186 -187 Coproica lugubris (Haliday). 186. Female terminalia, dorsal. 187. Fe­ male terminalia, ventral. Abbreviations: S8 = sternite 8, T8 = tergite 8.

230 188

: : 4kp~- '. •' • •*—t-^ijgjftv^ rf ._ r-.

189 190

Figures 188 -190 Coproica mitchelli (Malloch). 188. Male left wing, dorsal. 189. Male te rminalia, left lateral. 190. Male terminalia, posterior.

231 Figures 191-194 Coproica mitchelli (MaHoch). 191. Left surstylus. 192. Aedeagus and as­ sociated structures with postgonites removed. 193. Male sternite 5 and synsternite 6+7. 194. Left postgonite. Abbreviations: LP = lateral process, MP = membranous process, MS = membranous sheet, SAB = subapical anterior bristle, VS = ventral sclerite.

232 Figures 195-199 Coproica mitchelli (Malloch). 195. Female terminalia, dorsal. 196. Fem­ ale terminalia, ventral. 197. Female terminalia, left lateral. 198. Female stemite 8. 199. Spermathecae.

233 200

201 202

T Pg

Figures 200 - 202 Coproica novacula sp. nov. 200. Male left wing, dorsal. 201. Male term- inalia, left lateral. 202. Male terminalia, posterior. Abbreviations: Pg = postgonite, Ss = surstylus.

234 Figures 203 - 206 Coproica novacula sp. nov. 203. Female sternite 8. 204. Aedeagus and associated structures with postgonites removed. 205. Male sternite 5 and synst- ernite 6+7. 206. Left postgonite. Abbreviation: PLP = plateau-like process.

235 209 210

Figures 207 - 210 Coproica novacula sp. nov. 207. Female terminalia, dorsal. 208. Female terminalia, ventral. 209. Female terminalia, left lateral. 210. Spermathecae.

236 Figures 211-212 Coproica pappi Carles-Tolra. 211. Male left wing, dorsal. 212. Female st- ernite 8. 213. Aedeagus and associated structures with postgonites removed. 214. MaJe sternite 5 and synsternite 6+7.

237 Figures 215 - 216 Coproica pappi Carles-Tolra. 215. Male terminalia, left lateral. 216. Male terminalia, posterior. 217. Left postgonite. Abbreviations: C =cercus, Pg = postgonite, SAP = subapical process on cercus, Ss = surstyJus.

238 Figures 218 - 221 Coproica papp/Carles-Tolra. 218. Female terminalia, dorsal. 219. Fem­ ale terminalia, ventral. 220. Female terminalia, left lateral. 221.Spermathecae.

239 222 ^^'ssssas^^isasaa^a^s^<^sa^9^ m

.73;„,7', "J-

Figures 222 - 224 Coproica pusio (Zetterstedt). 222. Male left wing, dorsal. 223. Male te- rminalia, left lateral. 224. Male terminalia, posterior.

240 Figures 225 - 228 Coproica pusio (Zetterstedt). 225. Left postgonite. 226. Aedeagus and associated structures with postgonites removed. 227. Male sternite 5 and synsternite 6+7. 228. Female sternite 8. Abbreviation: SAB = subapical anterior bristle.

241 Figures 229 - 232 Coproica push (Zetterstedt). 229. Female terminalia, dorsal. 230. Female terminalia, ventral. 231. Female terminalia, left lateral. 232. Spermathecae.

242 233 I.

234

Figures 233 - 235 Coproica rohaceki Carles-Tolra. 233. Male left wing, dorsal. 234. Male terminalia, left lateral. 235. Male terminalia, posterior.

243 Figures 236 - 238 Coproica rohaceki Carles-Tolra. 236. Left postgonite. 237. Aedeagus and associated structures with postgonites removed. 238. Male sternite 5 and synsternite 6+7. Abbreviations: CS = central sderite, LS = lateral sclerite, MS = membranous sheet.

244 Figures 239 - 242 Coproica rohaceki Carles-Tolra. 239. Female terminalia, dorsal. 240. Fe­ male terminalia, ventral. 241. Female terminalia, left lateral. 242. Spermathecae.

245 243

Figures 243 - 245 Coproica rufifrons Hayashi. 243. Male left wing, dorsal. 244. Male term- inalia, left lateral. 245. Male terminalia, posterior.

246 Figures 246 - 249 Coproica rufifrons Hayashi. 246. Left postgonite. 247. Aedeagus and associated structures with postgonites removed. 248. Male sternite 5 and synsternite 6+7. 249. Female sternite 8.

247 Figures 250 - 253 Coproica rufifrons Hayashi. 250. Female terminalia, dorsal. 251. Female terminalia, ventral. 252. Female terminalia, left lateral. 253. Spermathecae. Abbreviat­ ion: S8 = sternite 8.

248 255

•VA*- ,*•*.-

Figure 254. Coproica serra (Richards), male left wing, dorsal. 255. Coproica setulosa (Duda), male left wing, dorsal.

249 Figures 256 - 257 Coproica setulosa (Duda). 256. Male terminalia, left lateral. 257. Male terminalia, posterior. 258. Left postgonite.

250 Figures 259 - 262 Coproica setulosa (Duda). 259. Spermathecae (the depression is an artifact of the drying process). 260. Aedeagus and associated structures with postgo- nites removed. 261. Male sternite 5 and synsternite 6+7. 262. Female sternite 8.

251 Figures 263 - 265 Coproica setulosa {Duda). 263. Female terminalia, dorsal. 264. Female terminalia, ventral. 265. Female terminalia, left lateral. Abbreviation: CS = cereal spines.

252 266

Figures 266 - 268 Coproica testudinea sp. nov. 266. Male left wing, dorsal. 267. Male ter- minalia, left lateral. 268. Male terminalia, posterior.

253 Figures 269 - 272 Coproica testudinea sp. nov. 269. Left postgonite. 270. Female ster- nite 8. 271. Aedeagus and associated structures with postgonites removed. 272. Male sternite 5 and synsternite 6+7. Abbreviations: LS = lateral sclerite, MP = membranous process, MS = membranous sheet, SPM = subapical process on medial surface of lateral sclerite. 254 Figures 273 - 275 Coproica testudinea sp. nov. 273. Female terminalia, dorsal. 274. Fem­ ale terminalia, left lateral. 275. Spermathecae.

255 276

Figures 276 - 278 Coproica urbana (Richards). 276. Male left wing, dorsal. 277. Male ster- nite 5 and synsternite 6+7 278. Aedeagus and associated structures with postgonites re­ moved. Abbreviations: AP = apical process, DPM = dorsal process on medial surface of lateral sclerite.

256 Figures 279 - 283 Coproica urbana (Richards). 279. Female sternite 8. 280. Male terminalia, left lateral. 281. Male terminalia, posterior. 282. Left postgonite. 283. Right postgonite.

257 Figures 284 - 287 Coproica urbana^Richards). 284. Female terminalia, dorsal. 285. Fem­ ale terminalia, ventral. 286. Female terminalia, left lateral. 287. Spermathecae. Abbrev­ iations: AP = anterior process, BC = basal constriction, f S = floating sclerite.

258 288

Figures 288 - 290 Coproica vagans (Haliday). 288. Male left wing, dorsal. 289. Male term- inalia, left lateral. 290. Male terminalia, posterior.

259 Figures 291 - 293 Coproica vagans (Haliday). 291. Spermathecae. 292. Distiphallus. 293. Male sternite 5 and synsternite 6+7.

260 Figures 294 - 295 Coproica vagans (Haliday). 294. Female terminalia, dorsal. 295. Female terminalia, left lateral. 261