, f ! j ) j . C' " , • :.CLASSIFICATION, ZOOGEOGRAPHY, AND PHYLOGENY OF THE GENUS SINOPHORUS' FOERSTER (HYMENOPTERA: ICHNEUMONIDAE) • By' © PAUL MICHA!l- SANBORNE, B.~c.·, M.Sc. , .A .Thes I s " Submitted to the School of Graduate Studies "\ . i"n Partial Fulfi lment of the Requirements , . for .the Deg ree Doctor •of Philosophy r \ McMaster U~;versity '. Hanllton, Ontario, Canada .. January, 1983 ',. o ) (I) f / n , '. .. j' THE GENUS SI tlOPHORUS FOERSTER, (HYMENOPTERA) i , . " o • I ( , -' • DOCTOR OF- PHILOSOPHY (1983) HcMA~TER UNIVERSITY (Bio)ogy) Hamilton, O~t~rio TITLE: Classific~tion, Zoogeography, and Phylogeny'of the Genus Sinophorus Foer.ster (Hymenoptera : l~hrielll1onidae) : • "(, \ \ \ -, • (i I) .I o TABLE OF CONTENTS ,~ \ L~T ~ OF TABLES. ••.••.••.• .•••.••.•", ••.•••••....•.•..•....•••••••. vi .L1ST Of' FIGURES ••.••••.•••••••.••••••..••...•..••..•.. .... ; .•....••.••• vi i ~llSTRACT : ; .•.••..•••.•...•••••.•. ~V i j i . .' . AC KNOWLE DG EHEHTS ••••••..•'.•••.••••••••••••••.•.•••.••..••....••• ix I NTRODUCT ION. '.' .'••.• ~ , •••••.••..•••••..•••.••••• > ••••• • . • . '.:. 1, d MATERIALS AND METHODS •• : ; V . 2 . i or Collections Studied••..•.•••••••••••••••••.•••••••••.•••••• 2 Term i nology' •••••••.•••••••,' .•• ' '•.•.••• 3 Taxonomic Characters and. thei r Measurement •••• : .• : •.•..•••• '. 3 11'Iustrati9ns '•••••. , ! :.:. 10 ZOOGEOGRAPHY •••••••.•••••.•.' ••••.••.••.....•• : ••••••••.•••••••.•.••. 13 PHYlOGEtlY. : '.••.....•••••••••.'••.••••.•••.•••••.••.••• 28 BIOLOGy •••••••••••••.••••••••••.•.••••••••••••••.•••.••.•.••.•••• 40 T,I\XONOHY ••••.J•••••••..••••••••••••• , •••••••••••••••••••••••••••••• 46 . \ oeSCrllJtlon.. 0 f s·'Inoph'orus Foer.ster . 46 Keys t~ the North and Hiddle American' Species' of Sinophorus ••.•• 49 . .' . Fern a Jes ;' :. .1.. : •••• ' ••• : • , •••••••••••••• 50 Males •........... : •..........'..........• 0''';" •••••••••••• : ••• ". 64 Keys to the Palearctic Species·of Sinophoru~•••.••••. , •••••••••• 79 . Females , ; .. 79 , Hales •• , •••••••• : ••••·••••••••••• : . 88 C~SSIFEMUR GROUP •••••••••••••••• ; 94 II. arm;'l latus. .. 129 1. crass i femur •••••.••.•••••• 95 12. nigricoxa•• " 131 2. brunnefemur.............. 97 )3. nigridontis.. 135 3. megadon tis••••••••••••••• 100 MONTANUS GROUP ••••••••. 1)8 , . COSTALI S GROUP ..................... 104 14. tur i onus ••••• OS 4. renOOlinatus •• o •••••••••••. 105'/ 15. lat I fossus ••• 142 5. ·sapporoensis••••••••••••• 108 ! 16. m~:mtanus••••• ' 144b 6. pectinatus •............•• l1T i 17. albipalpus••• 150 7. hydr i ae•••••••••••••••••• 115 18. curvatus ••••• 153 8. 9 raec i ens is.. " .... ; ••••••• 120 1 ~. dil·atus ........ .155. 9. Jun iperinus•••••••••••••• 122 20. pal1'idus •••••. 157 v 10. costa f is ................. 126 21. boreus •••• : •• 160 (Hi) ( / \ r .) 22. indistinctus 163 51. longifaci!,s 261 23. geni~ulatus7.•..: ...•• 166 • 52. eruficinctus •. : ...• 264 ..f. .. 24. J apon lOCUS ••••••••••••• 16~' 53,' ·ventosus.~ .....•. ;. 266 25. alpinus . 171 54. furus 21'9 ,/ 26. orient·is 174 55. ·splendidus•........• 272 26a. o. orient is 176 56 •. neof.urus •.·.. ~ ... : .. 275 26b. o. euronotis•.•• T77 57. heliot:,a~•........• 277 26c. o. occidental is. 178 .DESERTUS GROUP •............•• 280 27. i dahoens is ••. e._ .. ;;3. sabulum . 281 . --..;" 179 . ' . 28. popof.en sis••.••..••.•• 183 •. ·d,sertus.·.•...•.••. 283 • 29'• va~ianae••.•...•.•.••• 185. OBTUSUS .G UP ..... :. 286 30. val idus 187 60. 286 31. rhyaCioniae•..•••....• 191 1, 60a. b. ·brochus•.. 288 32. canadens is 195 I GOb. ·b. brev i'p i 1us 289 .' 33 • tunidus - 1971 ·61. obtusus 290' 34. vadabi 1is 201.1 62. 'Infimus 292 . : 35. mason I •••••• '•••••••• " 204 I I 63. sarissophorus•.•.•• 295 36. robustus 207 ., SUj.CATELLUS GROUP ••.••...••.•.299 37. minutus ·..•• 210 64. sulcatel1us••.•••.• 299 33. cotidianus •... : 213 oJ 65. inflatus 3,02 39. l<lnceolatus•••.••••••• 216 66. tahoens is 305 40. androsae...... ,' ....... 221 ALB I DUS GROUP 308. .PRISMATIS GROUP •....'.•••.••.•. " 226 67. fuscicarpus •. ; .•••• 309 ... .. 41. brevi stylus•..•••..• ".226 68. :alfidus 312 42. betulel1ae 230 69. ~nigritel1us 315 43. pri smat i s 234 70•. villosus 319 44. townesorum 237 71. b i 1imbus, .••••••••• 323 RUGOSUS GROUP 24T PLEURAL I5 GROUP •••••••••.•..• 326 45. ru~sus : 242 72. pleura1Is 326 46. striatus•.•••.. ~ .••••• 245 73. punctifrons 330 RUF' COXA GROUP 248 74. colunbiensis• 333 .47. ruficoxa ; ••.•• ,' 2Jr8. 75. \~oodor.um•..••.•..•• 336 " 48. triangularis•••.•••••• 251 76. pciropleurqn •••..••• 338 49. fa1 catus 254 VERICULUS GROUP >43 FURUS GROUP •..••• : •••••••...•••• 257 I 77. sticticaJae···· .. ··.343 I . 50. rufitergum 258· 78. vericu1us ••·•. · .•• ·.346 <iv) r .. \ . 7J. constrictus •••••.•••••• 350 89. caradrinae•..•• '..•• 383 -:. 1 30. punctatus., •.•...•.. :;. 3531 ,90. re Ia t i vus •.• ;' •.•••• 386 81. an i sodon tis........ '.' .•• 356 1 91. planus••.•..•..•.;. 339 XANTHOSTOHUS GflOUP,' ••••.••.••.••• 360 92. erectus•.•.. ; .••..• 393 82. kasp~ryani •••. : ••.•.••• 360 1 93. 1eptofemur. •••..•• .'. 396 33. nitidus••••••••••••••.• 364;.1 94. xanthocoxa • .' •..••.• 398 84. exartemae.. .............. 367! 95. .hall i .............. 401 85. pet io Ius ...............'. 370 96. proceorus ..........' .• 403 x~~tnost~us ~73 I' • 86. •••• 97. neoprocerus ••.••••• 406 87. ' ~ibial is.............',. 376 98. platyfacies ••••. ; .• '409 RELATIVUS GROUP •. ; •••••••••••.•••• 380 C8. PO~()sus. •••'••••••••••••• 380 . AW£NDA 99. inclivous (Costal is Group) ...,•• ,.,...... : •••.:.••••••• , •••••••• 414 100. rot~ndus tFurus Group) •••••••.•••.•••.••••: ••••••• , •••••••: 417 101. peteri (Furus Group); : ~ 419 102, apiculus (Pleural is Group) •••.••••••••••••~ •..••••••••••••• 422 lll3:. m0!'.9ccoensis ePleural is Group) 424 -. ." e ' REFERENCES ••• , .••••••\ .••..•••••• '.' •••••••••••.•••..••.•••..•••••.••• 428 flAps •.....................•.......................• ' .... : ...............•..• 433- FIGURES •••••••••••••••••••••••••••••••.·••.•••·•••• , ••••••.••• , •••••••• 5 45 • INDEX TO HOSTS : , ••.• : 631 ... ( (v) .. r LI ST 9F TABLES Frequency distribution of maximum I)rie~r range, in km., o( speClies of Sinophorus in Ilorth and l1iddle lVnerica., P. 18. \ Total number of "species and subspecies, average landmass "1 Interval values (ALlV) and total interval villues (TIV) 'l/ derived from map 3. P. 19. 3. Distribution of Ilorttl and Hlddle lVnerlcan species and 'I subspe,:'ies of S inophorus In relat ion to centers 'tlf ~oncentra- ---I tlon (Map 4). P. 20. , . 4. Dlsslmllarl'ty values among centers of concentration of SiJ+ophorus s~P. in Ncrth and :Iiddlc Jlt.1erica. P. 23. 5. (ndex of disslmi larlty among centers of concentratl,on of Sinophorus ,spp. in North and I1lddle lVnerlca, determined from Table 4. P. 23. ) .. 6-. Postulated ancdstra] and derived character states of selected I structures of species of Sinophorus. P. 29. 7. Sinophorus spp. and their hosts: A comparison of the relative lengths of ovipositors and the feeding 'behaviour of hosts, P. 41. (vI) , , LI ST OF FIGURES FIGURES - 9 Frontal view of heads of Sinophor~s spp. , 10 - 17 Dorsal view of heads of ?inophorus spp.. • 18 - 26. Mandibles of Sinophorus spp. 27 - 35 Mesopleural foveae of Sinophorus spp. ) 36 - 51.. Hind 1egs of §..0E'0rus ·spp. ' • , 52 - 56 Lateral vie.. of 1st abdominal segments 'of Sinophorus spp. 57 E !19. Dorsal viel<.of 1st abdominal segments of Sinophorus spp. # 60 - 62 Dorsa I vIew of terg ites 2 • 3 of abdomens of ina les of ~nophorus spp. • 63 73 • Lateral view of ovipositors of females of Sinophorus ... spp. ~ , 74 - 79 • LateraI view of ovi pes Itor tips of females of Sinophorus spp. ;- • 80 - 86 • Fore and hind wings of the genus ~JP-0plex and the genus Sinophorus. ~ . G7 -'169., Propodea of iinoph~ spp. 170 - 171. Phylogenetic relationships among species groups of Sinopi,or:Js. (v Ii) \ \, I ABSTRACT The world species of the genus Sinqphorus Foerster (Porizontinae= , Campopleginae) are'revised. One hundred and three species and S~ subspecies are recognized, Eighty species and 5 subspecies are described as new: albipalpus, alpinus, androsae, anisodontis, apiculus, armilliltus, betulellae, bilimbus, boreus, br. Ius, brochus (.!:.. brochus,--, .!:. brevipilus), brunl)efemur, columbiensis, co ictus, cotidianus,' curvatus, dilatus', erectus, falcatus, graeciensis, halli, heliothae, hyd,riae, idahoensis, incl ivous, indistinctus, infimus, inflatus, japonicus, kasparyani, lanceolatus, latifossus, leptofemur, longifacies, masoni, megadontis, minutus, moroccoensis, neofurus, ~~rocerus, nigricQxa, nigridontis, obtusus, orientis ,(0. euronotis, £. occidental is, • - j ~. orientis), pall idus, pectinatus, peteri, petiolus, planus"platyfacies, 'poropleuron, porosus, prismatis, procerus, punctatus, punctifrons, rhyaci,onJae, Tobustus, ~ndus, ruficoxa: rUfit~rgum, rugosus, sabulum, ,sapporoensis, sa.rrssOphorus, splendidus" sticticalae, striatus, .. tahoensis, tibial is', townesorum, ,triangularis, tumidus, variabil is, varianae, ventosus, vericulus, vi llosus, "'codorum, xanthocoxa. TI.enty- ~ three species are re-described: albldus (Gmel in), canadensis (Viereck), caradrinae (Viereck), costalis (Thomson), desertus Kasparyan, eruficinctus (\Jalkl~y), exartemae (Uchida), furus (Cresson), fuscicarpus (Thomson), geniculatus (Go-........,,), juniperinus (Holmgren), !!!£!!.~ (Cresson), nlgritellus (Thomson), nitidus (Brischke), pleural is (Thomson), popojensis (Ashmead), relativus (Viereck), renomlnatus (Morley), sulfatel1'us (Viereck), turionus