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PHYLOGENETICS AND SYSTEMAUCS OF THE WORLD STEPHANIDAE (HYMENOPTERA)
Volume I
DISSERTATION
Presented in Partial Fulfillment o f the Requirements fbr
the Degree of Doctor of Philosophy in the Graduate
School of The Ohio State University
By
Alexandre Pires Aguiar, M.S.
*****
The Ohio State University 2000
Dissertation Committee: Approved by Norman F. Johnson, Adviser
Wbodbridge A. Foster
Hans Klompen Entomolo uate Program
JohnW. Wenzel UMi Number 9971503
Copyright 2000 by Aguiar, Alexandre Pires
All rights reserved.
UMT
UMI Microform9971503 Copyright 2000 by Bell & Howell Information and Leaming Company. All rights reserved. This microform edition is protected against unauthorized copying under Title 17, United States Code.
Bell & Howell Information and Leaming Company 300 North Zeeb Road P.O. Box 1346 Ann Arbor MI 48106-1346 Copyright by Alexandre Pires Aguiar 2000 ABSTRACT
The generic relationships within the 6mily Stephanidae are examined on the basis
of a cladistic analysis, the fossil species and genera are revised and its phylogeny
discussed, and the Australian fauna, of importance fbr the classification of the entire
fiunEy, is thoroughly investigated and revised. The relationships o f the family with other
Hymenoptera is also investigated, based on a réévaluation of literature data.
For the phylogenetic analysis, morphological observations are presented and discussed, and additional information and illustrations are provided for the majority of characters and their states. The analysis o f 57 species and 176 informative characters resulted in relationships and groupings which differ significantly fi'om traditional hypotheses. Results support the monophyly of the subfamily Foenatopinae Enderlein, but its original sense is expanded to mclude an aberrant new taxon, Xanthostephanus n. gen.
The subâmily Stephaninae Enderlein including Schlettereriinae Orfila) is paraphyletic according to the analysis, but was maintained as a valid taxon for practical reasons. The monophyly o fthe genus Schlettererius Ashmead could not be assessed, but it is mamtained as a valid taxon based on its “distantly basal” position in relation to other stephanids. Newly discovered characters strongly support each of the following genera as
u monophyletîc; Stephanus Jurine,Hemistephanus Enderlein^ Parastephanelhis Enderleiiv and Foenatopus Smith. Results suggest t h a t Benoit, Neostepfumus^ The morphological terminology for the family is revised and illustrated, and a key and descriptions to all valid genera are provided. Analysis of combined literature data did not support the ofren-coghated sister group relationship of Stephanidae and Orussidae, suggesting instead a basal position of stephanids, together with non-ceraphronoid evaniomorphs, within the Microhymenoptera clade of a biphyletic Apocrita. The fossil o f Protostephanus ashmeadi Cockerell is confirmed to be a stephanid, and illustrated. The taxonomic status oîProtostephanus Cockerell and Electrostephanus Brues, their relationships with mctant genera of Stephanidae, and problems in defining or synonymizing fossil genera of the family are discussed. Electrostephanus is provîsorûy red^ned, two fossil species, E sulcatus sp. n. and E neovenatus sp. n. are described and illustrated from male specimens preserved in Eocene Baltic amber, and a key to species provided. The Australian stephanids are revised and k^ed. Three genera and 21 species (15 new) are recognized. The taxonomic importance of over 300 morphological characters, includmg 45 morphometric ratios, is assessed. The history of introduction and establishment o f Schlettererius cinctfpes in Australia is summarized.Stephanus in crassicauda M orl^ is transferred to Meffschusr, the female is redescribed, the male reported fer the first time and compared with the female. Occurrence of the Neotropical genus Hemistephanus in Australia is ruled out,H. giganteus Girault is synonymised with M crassicauda ^ o r l ^ ) , m A H granulatus Elliott is transferred to Parastephanellus. The groundplan o f AasXx?Siasi P arastephanellus is described in detail, and a new character set is proposed fbr taxonomic studies and descriptions of its species; P, albicepsW&oXt is proposed as a junior synonym for P. damellicus (Westwood), and P. picticeps Roman is placed as species inquirendum. New taxa areM ceneonatrix, P, albigenuis, P, adiazetos, P. brevis, P. callidictyon, P. denotatus, P, isogfyptos, P. laevigatus, P. maculatus, P. platyceratus, P. squamatus, P. strigosus, P. tanytarsus, P. trifasciatus, and P. vertipictus. Other valid species for Australia are P. caudatus Elliott, and P. ntfoom a tus (Cameron). IV ACKNOWLEDGMENTS I am deeply grateful to Norman F. Johnson, my advisor, for providing first-rate intellectual and material support to my research, and fi)r allowing me full, unconditional liberty to pursue not only my own ideas, but my own ideals, to think and materialize whatever I could, to write as much or as unconventionally as I wanted, to test, to experiment, to fail, and to succeed. For encouraging and supporting the flapping of my ambitious wings, my most smcere thank you. John W. Wenzel put the “PhJD.” in my degree. With his admirable knowledge, powerful teaching skills, and genius, John shuns away all mediocrity, allowing not only himself but also all the lucky ones around him, to aim for bright achievements under an unclouded sity. I thank him fbr so accessibily sharing his diamond talents with me. Luciana Musetti provided indispensable advice and professional help, both o f the highest quality, in nearly all parts, and fbr nearly all phases of my work: past, present, and future. Also, more sharply and intelligently tunned to the world and its people than any mommg newspaper, Luciana is an oasis of lucidity, one fiom which L fbr some luclty reason, always had the privilege o f enjoying and benefitting fi’om indefinitely. Ave, Lucianal Wbodbridge A. Foster and Hans Klompen raised the level of my committee to peak quality» and it is with pride that I acknowledge their input to my work and formation. I warmly thank Andrey Sharkov for his superb rendermg of Figure 1 (chapter 1), and for sharing with me his artistry, knowledge, witty sense of humor, and passionate eloquence. Without Jens-WOhebn Janzen the revision of the fossil stephanids would never materialize. After finding two extremely rare fossfls m this fomily, Jens kindly contacted m^ and for two years we enthusiastically cooperated in a joint paper, published while I was editing this dissertation. The holotypes of the new taxa described were made available for this study, and later donated to the %mburg museum, by Jens von Holt (Amberworld, Hamburg) (£. sulcatus sp. n.), and Carsten GrOhn (Oinde, near Hamburg) (& neovenatus sp. n.), and their kindness is also greatly acknowledged. Brian J. Armhage and Steven Passoa, although not directly involved in my work, were always receptive, always mterested, and showed such a constant and distinct aura of confidence m me, that th ^ too significantly contributed to the appearance of the final woric, and to my confidence m it. A large number of curators and other university^ scholars, listed on pages 7-13, promptly invested their time m my enterprise, and I am deeply grateful to all of them for this, hfichael J. Sharks kmdly helped with all my questions about wing venation nomenclature. Ian Naumann and Keith L. Taylor helped with several reprints of papers on Sdiiettererius cmctipeSy and Elame DeConick kindly copied and mailed me several VI important reprints of P. L. G. Benoit's papers. Theron Ellinger skillfîilly developed the films and customized all photographs that appear in this worl^ without ever frowning at any of my multiple and demanding requests. I also benefitted slgnfficantly from interacting with many other colleagues, each of which with their own admirable talent, and all shanng an mspirin& unstoppable willpower: Todd Blackledge, Mark Branham, Eric Dotseth, Istvan Karsai, IStesh Khanna, Cheol-Min Khn, Chi-Feng Le^ T. Keith Philips, and Kurt Picket, to d te only those with whom I talked more frequently. This project was supported m its majority^ by a generous scholarship from CAPES (“FundaçSo CoordenaçSo de Aperfeiçoamento de Pessoal de Nivel Superior”, Brazil), and by a grant from NSF to my advisor. Without then financial support, my wish and will to generate knowledge would never have been executed b^ond the title page. vu VTTA March20,1967 ...... Bom >Guaçui,Esptrito Santo, Brazil I985-I990 ______B_A. Biology, UniversidadeFederal do Espirito Santo, Brazil 1991-1994 ...... M.S. Entomology, UniversidadeFederal do Paranà, Brazil PUBLICATIONS L Aguiar, A P., and Janzen, J.-W. 2000. An overview of fossil Stephanidae hymenoptera), with description of two new taxa from Baltic amber, and key to species o f ElectrostephanusBmes. Ejrtomologica Sccondmœnca 30: 443-452. 2. Aguiar, A. P. 1998. Revisio do Gênero Hemistephanus’EvAeà&xi, 1906 (Hymenoptera, Stephanidae), com consideraçdes metodolôgicas. R evista hrasilefradeEntomolo^a 41:343-429. 3. Aguiar, A P., and Musetti, L. 1998. The Stephanidae Homepage. URL: http:/Jmsbiosci,ohiostate.edu/projects/stephanids.html 4. Aguiar, A P., and Sharkov, A 1997. Blue pan traps as a potential method for collecting Stephanidae. Journal o fHymenoptera Research 6:422-3. 5. Aguiar, A P. 1997. Mating behavior of Pelechmspofyturator hymenoptera: Pelecinidae). Entomologiccd News 108:117-121. MATOR AREA OF STUDY Entom olo^ viii TABLE OF CONTENTS Abstract...... ii Acknowledgments ...... v Vita...... vin LîstofTables...... xii ListofHgures...... xiv Litroduction...... 1 Chapters: 1. Phylogenetics and classffîcation ofthe Stephanidae ...... 6 Material and methods ...... 6 Specimens and museums...... 6 Identification...... 13 Selection oftaxafbrtheanalysis ...... 15 Male specimens ...... 16 Selection of outgroups...... 16 Sdectionofdiaracters ...... 18 Character coding and polarization ...... 19 Observation and Hlustratioa of characters ...... 21 Drawing conventions ...... 22 Termmology ...... 23 Mbrphologcal characters and observations ...... 33 Phylogenetic analysis ...... 123 Monophyly and classMcationofstephanidtaxa ...... 132 Monophyly oftheStephanidae ...... 132 Classîfîcationinsubâinilies...... 135 Stephaninae Enderlein ...... 135 Schlettererius Ashsaeai...... 137 StephanusJvcàat ...... 139 A/egûcAusBrullé...... 141 IX Hemistephanm-^Parastephanellm+^o&caio^mz& ------146 HemistephcmusBTLà&Atàa...... 148 form&pAoMeZ&y+Foenatopinae ...... 153 ParastephaneUusBoàsAàxi ...... 156 FoenatopmaeEndedem ...... 159 Xanthostephamsnav.gea ...... 162 FoemüopusSmüi...... 165 General considerations ...... 171 Key to subfamilies and genera ...... 173 Descriptions ...... 176 Schlettererius ^Yaaezd...... 176 StephanusJ\xtvR& ...... 180 MeffschusBrvSLè...... 184 Hemistephamts'Endtclàa...... 191 ParastephaneUus'EnicA&xi ...... 197 Xcanhostephanusnov. gen ...... 202 Foenatopus Smith...... 207 Relationships of Stephanidae with otherHymenoptera ...... 214 Historic...... 214 Combined analysis ofliterature data ...... 217 Results and discussion...... 220 Conclusion...... 227 PlatesofiUustrations...... 228 Cladograms ...... 262 2. The fossil Stephanidae, with description ofnew taxa ...... 288 Litroduction...... 288 Material and methods ...... 289 Results and discussion...... 290 GenmProstostephanusCodises^...... 290 Gem&ElectrostephamsBru^ss...... 291 Key to species ...... 292 Descriptions ...... 293 ElectrostephanussulcatusTL sp ...... 293 ElectrostephanusneavenatusxL sp ...... 296 Taxonomic status and relationships offbssil genera ...... 299 Conclusions...... 304 Plates ofillustrations...... 305 3. Revisionofthe Australian Stephanidae ...... 311 Introduction...... 311 Material and methods ...... 313 General ...... 313 Morphometricanalysis ...... 315 Sphericity indec ...... 316 Identification ...... 317 Sexrecognition ...... 317 Morphometnc ratios ...... 318 Body size and interpretation of character states ...... 319 K ^to Australian genera of Stephanidae ...... 320 Gtaixs&:hlettererius...... 321 GecmMegischus...... 323 Key to Australian species ofMegischus...... 323 M crassicauda ^ o rle y ) ...... 324 M. ceneonatrixn. sp ...... 328 GemsParastephanelhis ...... 331 Groundplan ...... 333 Key to species ...... 338 Descriptions ...... 349 P. maculatustL sp ...... 349 P^brevtsxLSç ...... 354 P. laevigatusxL sp ...... 358 P. isoglyptosn. sp ...... 363 P. dbOTe///cay(Westwood) ...... 368 P. albigemrisTL sp ...... 3 76 P, adiazetos n. sp ...... 380 P. callidictyon n. sp ...... 384 P . tanytarsusn. sp ...... 388 P. gyamlatus (EVaaVl) ...... 393 P. platyceratusTL sp ...... 397 P. strigosusn, sp ...... 401 P. trifasciatusvL sp ...... 406 P. squcmatusn, sp ...... 411 P. rM^omflrar(Cameron)...... 417 P. vertipictusn. sp ...... 423 P. denotatusn. sp ...... 427 P. cautknusEMloiX...... 431 Species inquirendum ...... 436 Stephanid taxa mistakenly reported in Australia ...... 43 8 Plates oOllustrations ...... 440 ListofReferences...... 496 APPENDIX Rangesofmorphometric ratios fbrspecies ofAustralianPams/ep^ane/ftfs ...... 506 XI USX OF TABLES Table Page LI Species examined for the cladistic analysis, with database number and depository museum for correspondent voucher or type specimens 14 1.2 Distinctly unspecialized features ofSchlettererius, compared with remaining extant genera of Stephanidae ...... 18 1.3 Data matrix fbr 55 species and 176 morphological characters used in the analysis...... 117 1.4 Character state changes on the branches of the shortest cladograms obtained when the constant of concavity, K, in PIWE, is set to I 131 3.1 Selected morphometric ratios evaluated for taxonomic usefulness with species of Australian Parastephanellus ...... 318 3j2 Morphometric ratios to be avoided in taxonomic decisions with species of Austn!&an. Parastephanellus ...... 318 3.3 Number of specimens studied (n) fbr each species o f Australian Parastephanellus, correspondent percentage (%) in relation to total number o f specimens, and indication of species for which males are unknown...... 332 3.4 Main differences between morphotypes of P, squamatus n. sp ...... 416 3.5 Intermediate characters in the holotype ofP. p ictice p s Roman...... 43 7 3.6 Mmimum and maximum values of taxonomically important morphometric ratios fbr Australian species o îParastephanellus, Females...... 506 3.7 Minimum and maximum values of taxonomically important morphometric ratios for Australian species o î Parastephanellus, Females...... 507 XII 3.8 Minimum and maximum values of taxonomically important morphometric ratios for Australian species o f Parastephanellus, Females...... 508 3.9 Minimum and maximum values of taxonomically important morphometric ratios for Australian species o f Parastephanellus, Females...... 509 3.10 Minimum and maximum values of taxonomically important morphometric ratios for Australian species o fParastephanellus, Females...... 510 xui LIST OF FIGURES Figure Page 1 HabitusofParasiepfumellusrvfoomatusQZameTon) ...... 228 General morphoIo©r, Parastephanellus damellicus (Westwood): 2 Head,frontai ...... 231 3 Head, dorsal ...... 231 4 Pronotum, dorsal...... 231 5 Head, postenor\new ...... 231 6 Prothorax, lateral ...... 231 7 Propodeumand metapleuron ...... 231 8 Stepfwmtsserrator,arXjaaa...... 233 9 Hemistephanus cylindricus, antenna...... 23 3 10 H. cylindricuSy flagellomeres 1-2 ...... 233 11 S(dilettererhtscmctipeSydMBxm&...... 233 12 Mepschustibiatory^a%f&omstes\~l ...... 233 13 FoerxnopusaurantiicepSy pedicell and flagellomeres 1-3 ...... 233 14 FœnatopusÇ^-Diastephanus) sp. A, scape, pedicell, flagellomeres 1-3 ...... 233 15 Foenatopus sp.y flagellomeres 1-3 ...... 233 16 Megischusn. sp., scap^ pedicell, flagellomeres 1-2 ...... 233 17 Paros/gp^oneZrus/v/oom a^, scape;, pedicel], flagellomeres 1-5 ...... 233 18 H. cg/mÿTow, pedicell and flagellomeres 1-3 ...... 233 19 P. nigricaucbiSy pedicell and flagellomeres 1-3 ...... 233 20 M. texanuSy scape, pedicell, flagellomeres 1-5 ...... 23 3 21 P. granulatuSy flagellomeres 1-3 ...... 233 22 Parastephanellussp.y pedicell and flagellomeres 1-4 ...... 233 23 M, califomiaiSy scape, pedicell, flagellomeres 1-4 ...... 233 24 MegischustL sp., pedicell and flagellomeres 1-6 ...... 233 25 H limpidipenniSy scape, pedicell, flagellomeres 1-2 ...... 233 26 HtyUndricuSyZsAema...... 233 XIV Head: 27 FoenatopiisflavitkTUeOuSyXaXBsai ...... 235 28 F. «ovflguffwesri; lateral...... 235 29 F (=i?/as/epAanas)sp.B, lateral ...... 235 30 HemistephcmusorfilanuSy^oAedorYai!^...... 235 31 H.cyliruMcuSyîmtiaX...... 235 32 Megischmoccipittalis^ frontal...... 23 5 33 M. maculipemis^ dorsal ...... 235 34 Schlettererius cmctipes^ dorsal ...... 23 5 35 F.camulipeSyàatsalL...... 235 36 F. aurantiiceps, dorsal ...... 235 37 F,amtulipeSy\ateai...... 235 38 ventro-Iateral, . right ...... 235 39 M texarmSy ventro-Iateral, right ...... 235 40 fbgnoAgwrsp. A, ventro-Iateral, right ...... 237 41 Hemistephanus elimatuSy ventral...... 237 MaxOlar palpus: 42 Meg^chustihiatory articles 3-5...... 237 43 M. tibiator, article3...... 237 44 Megischusn. sp., articles 2-3 ...... 237 45 Schlettererius cmctipes, articles 2-3...... 237 46 M maculipermiSy articles 2-3...... 237 47 M texanuSy articles 2-3...... 237 48 H validuSy mandibleandclypeus ...... 237 49 M egischus n. sp., head, latero-ventral ...... 237 50 sp., maxHlar palpus, articles 2-6 ...... 237 51 Stephanusserratory mandible and clypeus ...... 237 52 Mmacv/rpenn», mandible and clypeus ...... 237 53 H macnmtSy mandibleandclypeus ...... 237 54 ScMettereriuscinctipeSypcQïiQVijaxL ...... 239 55 Schlettererius cinctipeSy pronotum...... 239 56 MegischustibitUory colo...... 23 9 57 StephanusserratorypTonotum ...... 239 58 StephanusxrreUoryÇtonotum ...... 239 59 HemistephanuserugatuSypronotmi ...... 239 60 M . texanuSy pronotum...... 239 61 M texanus, pronotum...... 239 62 K limpidipemiSy pronotum...... 239 63 M.macuIipenniSyptonotam ...... 239 64 Mmocii/ipenmsr,pronotum ...... 239 65 JdrnthostephanusxanthocephaluSypronotum ...... 239 XV 66 Parastephanellusisogfyptosy pronotum...... 239 67 P. isogfyptos^ pronotum...... 239 68 XcmthostephanusxanthocephaluSy pronotum...... 23 9 69 FoemtopusamuIipes^ptonoXxxca...... 241 70 F.annu/pcy, pronotum...... 241 71 Foenaiopussg.y^xonotxim...... 241 72 F, abOaceus, pronotum...... 241 73 F , alufacettSy pronotum...... 241 74 F. novagufneensis, pronotum...... 241 75 Foenatopussp^'ptonotam...... 241 76 F. novaguineensis, pronotum...... 241 77 Kpew/Aerr, pronotum ...... 241 78 F , (^Neostephanus) sp., pronotum ...... 241 79 F, pentheri, pronotum...... 241 80 F. (^Diastephanus) sp. D, pronotum ...... 241 81 fbenafcyKsn. sp., pronotum ...... 241 82 F, seyrigi, pronotum...... 241 83 Schlettererius cinctipes, apexofprostemum ...... 243 84 apex .ofprostemum...... 243 85 Meffschusmaculipermis, connection or pro- and mesothorax ...... 243 86 Schlettererius cinctipes, tibia! spur and vellum...... 243 87 Stephanusserrator, tibial spur and vellum...... 243 88 fbena/cpKs sp. D, tibial spur and vellum ...... 243 89 M maculipermiSy tibial spur and vellum...... 243 90 F./orojosarras; tibial spur and vellum ...... 243 91 F . annulipeSy posterior margim ofpronotum ...... 243 92 Schlettererius cmctipes, mesoscutum, parapsides, and axillae ...... 243 93 Stephanusserrator, mesoscutum and parapsides ...... 243 94 M texanus, mesoscutum and parapsides ...... 243 95 M . maculipermis, mesoscutum and parapsides ...... 243 96 ParastephanellusTL sp., mesoscutum, parapsides, and axillae ...... 243 97 K albomaculatus, mesoscutum, parapsides, andaxHlae ...... 243 98 Hemistephanus cylindricus, posteriormargimofpronotumand mesoscutum 243 99 F . aurantiiceps, mesoscutum and parapsides ...... 243 100 M fUrcatus, mesepistemum...... 243 101 F.rmran/f/ceps, mesepistemum ...... 243 102 Foenatopus (j=Diastephanus) sp.B, thorax ...... 245 103 fbenorqpKssp. A, mesepistemum ...... 245 104 F.Ç^Neostepharrus) sp.Aythorax ...... 245 105 Megischusfitrcatus, propodeum+metapleuron ...... 245 106 thorax ...... 245 107 Parastephanellus n. sp., posterior half of mesepistemum, mid coxa, propodeum + metapleuron...... 245 XVI I0& M coronator, articulation ofpetioleand hind legs with thorax, posterior view 245 109 Foenatopussç, propodeum+metapleuron, and mid coxa ...... 245 110 M bases ofmid coxae, ventral view ...... 245 111 F. Q=Nèostephamis) sp. A, articulation o f petiole and hind legs with thorax, posterior view ...... 245 112 Parastephanellus n. sp., mid tibia, mesal side ...... 245 113 Detail of spines on mid tibia oîParastephanellus n. sp ...... 245 114 articulation ofhind coxa with thorax ...... 247 115 Foenatopus (j=Neostephanus) sp. A, articulation ofhind coxa with thorax ...... 247 116 Parastephanellusgranulatus, Mndcoxa ...... 247 117 Meglschustexanus,imdcoxa...... 247 118 H.elinuitus,)mdcoxa...... 247 119 F.aurantffceps,lmdcaxa ...... 247 120 F, aurantiiceps, bind coxa ...... 247 121 F, aurantiiceps, base ofhind femur, mesal ...... 247 122 Foenatopussç. A, baseofhind femur, mesal ...... 247 123 Schlettererius cinctipes, hind coxa ...... 247 124 Foenatopussp. F, base ofhind femur, mesal ...... 247 125 F. albom aculatus, base ofhind femur, mesal ...... 247 126 F. annulipes, hindfemur ...... 247 127 F. aw w /y es, mid-transversal section ofhind femur ...... 247 128 Stephanusserrator, hind femur ...... 247 129 Schlettereriusdeterminatoris, hindfemur ...... 247 130 Hemistephanusmacrurus, hind femur ...... 249 131 Foenatopus aurantiiceps, hind femur ...... 249 132 Foenatopussp. I, hind femur ...... 249 133 Schlettererius cinctipes, hind tibia, lateral ...... 249 134 Stephanusserrator, hind tibia, lateral ...... 249 135 A/ggwcAifcpmflCK/ipenwtf, hind tibia, lateral ...... 249 136 Parastephaneïïusn.sp.,\m&\Mdi,\sXetü ...... 249 137 M/fdmr/or, hind tibia, lateral ...... 249 138 F.anmr/rjpes, hind tibia, lateral ...... 249 139 M^raimews, hind tibia, lateral ...... 249 140 M tibiato r, hind tibia, lateral ...... 249 141 H. cylindricus, hind tibia, ventro-Iateral ...... 249 142 Mju rc a tu s, hind tibia, ventro-Iateral ...... 249 143 Schlettererius cinctipes, apex ofthorax and basal abdominal segments ...... 251 144 Foenatopussp. A, apex ofpetiole and abdominal segments3-5, latero-ventral 251 145 Foenatopussp. A, apex ofthorax and petiole ...... 251 146 Stephanusserrator,Stonttarsus ...... 251 147 K(=i>/asre/>Aawas)sp.A,ap«cofabdomen ...... 251 148 F.annulipes,apexofàbdoiaen ...... 251 149 Afegischuscrassicauda, hind wing hamuli and apex ofveinSc^R ...... 251 xvn 150 M Jurcatus, front wing, posteriormargin apical to retinaculum ...... 251 151 A/.^rcortto, detail ofshorthairs,Mjî»rca/to...... 251 152 M ^ rc a fts , detail oflong hairs ...... 251 153 M tibiator, front wm& posterior margin apical to retinaculum, with detail o f short hairs.: ...... 251 154 Mrfh/fl/or, detail oflong hairs ...... 251 Front wing, 4-vein nodus and setae distribution at apex of M+Cu: 155 M bekilyanus ...... 251 156 Hernistepfumuselimatus...... 251 157 ParastephanelhisTLsp ...... 251 158 P.rtigricaudus ...... 251 159 F.Jlcmdmtatus ...... 251 160 Stephanus serrator, hind win& ventro-apical portion, showing veins M+Cu and 2M, and apex of ventral wing fold ...... 251 161 MegischuscrassicaudaMorley, front win& vein and ceil nomenclature...... 253 162 6kAWgrgrmscmcfÿes(Cameron), hind wing, veinand cell nomenclature ...... 253 Front wing: 163 Schletterius cinctipes...... 255 164 Stephcoms serrator...... 255 165 Megischus anomalipes ...... 255 166 M . texanus ...... 255 167 M tibiator ...... 255 168 Hemistephanusmacrurus ...... 257 169 H. cylindricus ...... 257 170 Xanthostephanusxanthocephalus ...... 257 171 ParastepkmellusTvfoomatus...... 257 172 Foenatopuspentheri...... 257 173 Foenatopus aurantiiceps ...... 259 174 F. (j=Neostephanus) sp ...... 259 175 F.novaguineensis...... 259 176 Foenatopussp...... 259 177 Foenatopussp...... 259 178 F. alutaceus ...... 259 Hind wing: 179 Schlettereriuscinctipes...... 261 180 Stephanusserrator ...... 261 181 Meffschusanomalipes ...... 261 182 M . texanus ...... 261 183 M~ crassicauda ...... 261 xviu 184 M tib ia to r ...... 261 185 HemistephcmaUmpidipermis ...... 261 186 Xanthostephanusxanthocephalus ...... 261 187 Parastephanelhisrufoomatus ...... 261 188 Foenatopussp...... 261 189 ParastephaneHusn. sp ...... 261 190 F. aurantiiceps ...... 261 191 F. (^Neostephanus) sp ...... 261 192 F.rujvaguineensis...... 261 193 Foenatopussp...... 261 194 F. alutaceus ...... 261 195 Strict consensus tree cf36 MFCs found by NONA...... 263 196 Strict consensus tree of36 MFCs found by NONA with aid ofRatchet ...... 265 197 Single MFC cladogram found by NONA following the application of successive approximations character weighting ...... 267 198 Strict consensus of four MFCs obtained with implied weighting implemented using FIWE with /T set to 1 ...... 269 199 Cladogram obtained using wing venation characters only ...... 271 200 Restricted analysis of Foenatopinae taxa. Strict consensus of 3 MFCs obtained with implied weighting implemented using FIWE with A" set to 1 ...... 273 201 Réévaluation ofliterature data for Hymenoptera. All taxa. Strict consensus tree of 411 MFCs found by NONA with aid ofRatchet ...... 275 202 Réévaluation ofliterature data for Hymenoptera. All taxa. Single MFC cladogram found by NONA following the application of successive approximations character weighting ...... 277 203 Réévaluation ofliterature data for Hymenoptera. All taxa. Strict consensus of 4 MFCs obtained with implied weighting implemented using FIWE under A=1.. 279 204 Réévaluation ofliterature data for Hymenoptera. All taxa. Strict consensus of 3 MFCs obtained with implied weighting implemented using FIWE under itr=2 (non- ceraphronoid evaniomorpha only) ...... 279 205 Réévaluation ofliterature data for Hymenoptera. All taxa. Strict consensus of 5 MFCs obtained with implied weighting implemented using FIWE under ^=3 (non- ceraphronoid evaniomorpha only) ...... 279 206 Réévaluation ofliterature data for Hymenoptera. All taxa. Strict consensus of 4 MFCs obtained with implied weighting implemented using FIWE under ^=5 (non- ceraphronoid evaniomorpha only) ...... 279 207 Réévaluation ofliterature data for Hymenoptera. All taxa. Single MFC obtained with implied weighting implemented using FIWE under A=6 (non-ceraphronoid evaniomorpha only) ...... 279 208 Réévaluation ofliterature data for Hymenoptera. Extant taxa only. Strict consensus tree of 107 MFCs found by NONA with aid ofRatchet ...... 281 XIX 209 Réévaluation ofliterature data for Hymenoptera. Extant taxa only. Single MPC cladogram found by NONA following the application of successive approximations character weighting ...... 283 210 Réévaluation ofliterature data for Hymenoptera. Extant taxa only. Strict consensus of all MFCs obtained with implied weighting implemented using PIWE under all possible values fo r K ...... 285 211 Réévaluation ofliterature data for Hymenoptera. Shared taxa only. Strict consensus tree of 6 MPCs found by NONA with aid ofRatchet ...... 287 212 Réévaluation ofliterature data fbr Hymenoptera. Shared taxa only. Single MPC cladogram found by NONA following the application o f successive approximations character weighting ...... 287 213 Réévaluation ofliterature data for Hymenoptera. Shared taxa only. Strict consensus of two MPCs obtained whb implied weighting implemented using PIWE under all possible values for ^ ...... 287 Protostephanus asmeadi Cockerell: 214 Head ...... 306 215 Body ventrally, from prostemum to near apex of abdomen ...... 306 216 Frontleft wing, ventral side ...... 306 217 Detail ofstructure of cell IM and related venation...... 306 Electrostephanus sulcatus SÇ. n.: 218 Habitus...... 308 219 Frontwing ...... 308 220 Hind trochantellus, trochanter, and femur ...... 308 221 Metaplairon and propodeum ...... 308 222 Propodeum ...... 308 Electrostephanus neovenatus sp. n.r 223 Head, fronto-lateral ...... 310 224 Lefr hind leg, external side ...... 310 225 Lefr front wing ...... 310 226 Apical 0.75 of right hind wing ...... 310 227 Metapleuron andbasal section o f‘petiole” ...... 310 Setae on apec o f vein MKZu: 228 Electrostephanussulcatussg.n...... 310 229 Electrostephanusneovenatus sp. n ...... 310 Megischus crassicaucài M ort^: 230 Vertex...... 441 231 Pronotum...... 441 232 Propodeum ...... 441 XX 233 Metapleuron...... 441 234 Pronotum, mesoscutum, and axillae...... 441 235 Hind leg view oftrochanters trough tibia ...... 441 236 Axillae, scutellum, and crenulate sulcus...... 441 Megischus ceneonatrix n. sp.: 237 Vertex...... 443 238 Pronotum...... 443 239 Propodeum ...... 443 240 Metapleuron and propodeum ...... 443 241 Pronotum, mesoscutum, axillae, andpartofscutellum...... 443 242 Hind leg, outer side view offemur and tibia ...... 443 243 Detail ofliind femur, outerside...... 443 244 Locality records fbrM crassicauda M orl^ andM ceneonatrix n. sp ...... 445 245 Locality recordsfbr&^Algffgrerwgcmcfipes (Cameron) ...... 445 246 Seazonal distribution oSMegischuscrassicaudaMoiXe^ ...... 445 247 Seazonal dÀsXribv&onofSchlettererius cinctipes (tZdsasxod) ...... 445 Measurements used to calculate morphometric ratios, Parastephanellus vertipictus n. sp.: 248 Body in lateral view ...... 447 249 Head ...... 447 250 Pronotum...... 447 251 Tergites 3-4 ...... 447 Values of morphometric ratios fbr all examined female specimens of all species: 252 Ratio Ov/Ptl...... 449 253 Ratio Ov/Tt...... 449 254 Ratio Ov/Hdl ...... 449 255 Ratio Ov/Saw...... 449 256 Ratio Cxl/h ...... 449 257 Ratio Ct/Fml...... 449 258 Ratio Fm SI ...... 449 259 Ratio twAEE ...... 449 260 Ratio btl/w...... 451 261 Ratio Pnl/Saw...... 451 262 Ratio twrio...... 451 263 Ratio Pnl/Ew...... 451 264 Ratio Gsl/Ptl...... 451 265 Ratio F tl/ni ...... 4SI 266 Ratio Hdl/Ew ...... 451 267 Ratio Ew/tw...... 451 268 RarioiEE/eEE ...... 453 XXI 269 Ratio îEE/io ...... 453 270 Ratio Ppl/Ew ...... 453 271 Ratio io/oo...... 453 Parastephanellus maculatus n. sp.: 272 Head sculpture...... 455 273 Pronotum and mesoscutum...... 455 274 Axilla^ scutellum, basai section of propodeum...... 455 275 Pronotum...... 455 276 Ax3Ia^scuteIIuin,propodeum...... 455 277 Posteriorhalfofmesepistemum, metapleuron, propodeum ...... 455 278 HLid &murand tibia, outer side ...... 455 279 Tergites 6-9 and pygidium ...... 455 Parastephanellus brevis n. sp.: 280 Head sculpture ...... 457 281 Pronotum and mesoscutum...... 457 282 Tergites 7-9 and pygidium ...... 457 283 Mesoscutum sculpture...... 457 284 Pronotum...... 457 285 Apex ofscutellum, propodeum ...... 457 286 Propodeum and metapleuron ...... 457 287 Right &ont wmg, venationcentrally ...... 457 288 Hind fémur and tibia, detail ofsculpture, outer side ...... 457 Parastephanellus laevigatus n. sp.: 289 Head sculpture and colour, m orpho^e 1 ...... 459 290 Same, m orp ho^e H ...... 459 291 Pronotum...... 459 292 Axillaeand scutellum sculpture...... 459 293 Same, different specimen ...... 459 294 Pronotum...... 459 295 Propodeum ...... 459 296 Thorax, except pronotum ...... 459 297 Propodeum, detail of sculpture ...... 459 298 Mesepistemum ...... 459 Parastephanellus isoglyptos n. sp.: 299 Head sculpture ...... 461 300 Pronotum...... 461 301 Scutellum and propodeum...... 461 302 Metapleuronand propodeum ...... 461 303 Pronotum andmesoscutum...... 461 xxu Parcatephaneiïus damellicus (Westwood): 304 Head sculpture...... 463 305 Pronotum...... 463 306 Pronotum, detail ofsculpture...... 463 307 Axillae and scutellum...... 463 308 Pronotum...... 463 309 Axillaeand scutellum, second specimen ...... 463 310 Scutellum, detad ofsculpture, third specimen ...... 463 311 Apex of scutellum and propodeum ...... 463 312 Mesepistemum...... 463 313 Propodeum andmetapleuron ...... 463 314 Propodeum, detail ofsculpture...... 463 Parastephanellusalbigenuis n. sp.: 315 Head sculpture...... 465 316 Pronotum...... 465 317 Mesepistemum posterior halfand metapleuron ...... 465 318 Head posteriorly and pronotum ...... 465 319 Thorax, except anterior portion ofpronotum ...... 465 320 Axillae, scutellum, basal halfofpropodeum ...... 465 321 Caster...... 465 Parastephanellus adiazetos n, sp.: 322 Head sculpture...... 467 323 Pronotum and mesoscutum...... 467 324 Scutellum and basai halfofpropodeum ...... 467 325 Pronotum...... 467 326 Thorax ...... 467 327 Mesepistemum posteriorhalÇ propodeum and metapleuron ...... 467 Parastephanelluscallidîctyon n. sp.: 328 Head sculpture ...... 469 329 Vertex sculpture, detail ...... 469 330 Frons...... 469 331 Scutellum...... 469 332 Pronotum and mesoscutum...... 469 333 Prostemum...... 469 334 Pronotum, detail ofsculpture...... 469 335 Scutellumapecand propodeum ...... 469 336 Mesepistemum posteriorhaK propodeumand metapleuron ...... 469 337 Pronotum...... 469 xxm Parastephanellus tanytarsus n. sp.: 338 Head sculpture ...... 471 339 Head sculpture, anteriorview ...... 471 340 Pronotum and mesoscutum...... 471 341 Rightfront wing, venation centrally ...... 471 342 Pronotum...... 471 343 Scutellum apex and propodeum ...... 471 344 Axillaeand scutellum...... 471 345 Prostemum, ventral...... 471 346 Mesopseudosteraum, discrimen...... 471 347 Hind fëmurand tibia, outer side...... 471 Parastephanellus granulatus (EUiott) : 348 Head sculpture ...... 473 349 Pronotum and mesoscutum...... 473 350 Right front wing, venation centrally ...... 473 351 Pronotum and anteriorhalfofmesepistemum ...... 473 352 Prostemum, ventral...... 473 353 Mesepistemum...... 473 354 Scutellum and propodeum...... 473 Parastephanellus platyceratus n. sp.: 355 Head sculpture...... 475 356 Head posteriorly and pronotum ...... 475 357 Pterothorax and propodeum ...... 475 358 Thorax, except pronotum ...... 475 Parastephanellus strigosus n. sp.: 359 Head sculpture ...... 477 360 Pronotum, détail...... 477 361 Propodeumand metapleuron ...... 477 362 Head sculpture, detail...... 477 363 Pronotum...... 477 364 Axillae and scutellum...... 477 365 Right front wing ...... 477 Parastephanellus trifasciatus n. sp.: 366 Head sculpture...... 479 367 Pronotum...... 479 368 Head, anteriorview ...... 479 369 Pronotum...... 479 370 Mesoscutum, axilla^ scutellum...... 479 371 Same, different specimen ...... 479 XXIV 372 Mesepistemum...... 479 373 Mesepistemum posteriorhalÇ propodeum and metapleuron ...... 479 Parastephanellus squœnatusn. sp.: 374 Head-sculpture...... 481 375 Pronotum and mesoscutum, m orpho^el...... 481 376 Head sculpture, posterior view ...... 481 377 Pronotum, morphotypeü ...... 481 378 Pronotumposteriorly, mesoscutum, axillae, scutellum...... 481 379 Pronotum...... 481 380 Thorax, except pronotum ...... 481 381 Head and thorax ...... 481 Parastephanellus rufoomatus (Cameron): 382 Head sculpture ...... 483 383 Pronotum and mesoscutum...... 483 384 Thorax ...... 483 385 Pronotum...... 483 386 Mesepistemum ...... 483 387 Mesothorax ...... 483 388 Mesothorax ...... 483 389 Mesepistemum posterior halÇ propodeum and metapleuron ...... 483 390 Scutellum and propodeum ...... 483 Parastephanellus vertipictus n. sp.: 391 Head sculpture ...... 485 392 Temple^ pronotum, mesoscutum...... 485 393 Scutellum...... 485 394 Pronotum and mesothorax ...... 485 395 Prostemum, ventral...... 485 396 Thorax, except pronotum ...... 485 397 Mesepistemum posterior hal( propodeumand metapleuron ...... 485 Parastephanellus eknotatus n. sp. : 398 Head sculpture ...... 487 399 Pronotum and mesoscutum...... 487 400 Thorax ...... 487 401 Scutellum and propodeum...... 487 402 Hlndfemurand tibia, outerside...... 487 403 Tergtes7-9 andpygidîum, detad ...... 487 XXV Parasieplumellus caudatus n. sp.: 404 Head sculpture...... 489 405 Pronotum and mesoscutum...... 489 406 Axillae and scutellum...... 489 407 Pronotum...... 489 408 Propodeum ...... 489 409 Propodeum andmetapleuron ...... 489 410 Tergites 6-9 and pygidium ...... 489 Locali^ records for species of Paarastephamllus occurring in Australia: 411 AUspp ...... 491 412 P. isogfyptosiL sp ...... 491 413 P, maculatusn. sp ...... 491 414 P. brem n. sp ...... 491 415 P, laevigatusn. sp ...... 491 416 P, albigemisn^ sp...... 491 417 P. adiazetosn. sp ...... 491 418 P.callicüctyonrLSÇ ...... 491 419 P. flSwne///cay(WestwGod) ...... 491 420 P. tanytarsusxL sp ...... 491 421 P. granuijto^lliott)...... 491 422 P. platyceratus n. sp ...... 491 423 P.strigosusn.sç...... 491 Locality records for species of Parastephanellus occurring in Australia: 424 P. U^asciatusTi, sp ...... 493 425 P. vertipictusn^ sp ...... 493 426 P, squamatusn, sp ...... 493 427 P,denotatusxi.sp ...... 493 428 P. rH^o/7Krttts(Cameron)...... 493 429 P. caudatusWiati ...... 493 430 Seasonal distribution of AustralianParas/epAane/7z£s...... 495 Seasonal distribution for species o f Australian Paras/e/7Aane//u.s known from seven or more specimens: 431 P.rufoomatus...... 495 432 P. d a m e llic u s...... 495 433 PMoglyptosn^sp...... 495 434 P. trifaxiatusTL sp ...... 495 435 P.strigosusti,sp...... 495 436 P,platyceratusn,sp ...... 495 437 P , laevigatusn. sp ...... 495 438 P.sqaamatusn.sp...... 495 INTRODUCTION Stephanids are a group wasps with about 300 described species worldwide. Adults can be easily recognized by the presence of five distinct tubercles on the top of the head (hence the name stephanus, Greek for crown), hind leg segments highly modified (as in Rg. 1), and a long and narrow body. Most stephanids develop as ectoparasitoids on wood boring insect larvae, especially those o f Cerambycidae ^ e n o it 1951a, 1984a, BiQthgen 1953, Meyer e t aL 1978, Pagliano 1986, VôDger 1994, Visipamch 1994) and Buprestidae (Townes 1949, Chao 1964, Mateu 1972, Pagliano 1986, Braza 1989, Carison 1972), but several other 6mHîes of Coleoptera, as well as larvae of siridd wasps (Taylor 1967a, Kirk 1975) and (^patently o f solitary bees (Beriand 1951, MalySev 1966, Kdnigsmann 1978) can also be attacked. Indirect evidence suggest that host spedfid^ is low (e.g., Roman 1917, Benoit 1949,1984a), but unambiguous data on polypha^ is available only for Schlettererius cinctipes QSjuAi 1975) mà Stephanus senator OBlOthgen 1953, Pagliano 1986). Stephanids are found mostly m tropical fi)rests, but many live in desert or semi- desert areas (see for example Benoit 1984b), reach high latitudes (e.g., Megischus bicolor m Canada, S. serrator m Germany, Hemistephanus artiosulcatus in Argentina), or are 1 fnHpmic m many oceanic islands throughout the world. Most stephanids are regarded as rar^ but this may be related to the lack of eflScient collecting techniques for the group (Aguiar & Sharicov 1997). Adults seem to be predominantly diurnal, but some species are attracted to light tr^ s ^aum ann 1992, and personal observation), suggesting that flight actmty may start in, or extend to crepuscular hours. Flight and other movements are usually slow and weak, but cryptic coloration and behavior contribute to make many species extremely mconspicuous in their habitaL Although this may help to avoid predators, concentration of mdtviduals in exposed conditions for mating or oviposMon occurs at least for a few species in the genera Stephanus (VôOger 1994), Hemistephanus (personal observation), Parastephanellus (Rodd 1951), and Foenatopus (Cheesman 1932, Gauld 1995). Further non-taxonomic information about stephanids is mostly limited to fiagmented behavioral and biological observations on S. serrator (Blüthgen 1953, Jansen e t a l. 1988, VôOger 1994), and seasonal data for Megischus spp. in California (Eblstead 1986). The recognition and definition o f stephanids as a separate fomily was a persistent problem for nearly a century and a hal^ mamly because of similarities with members of Ichneumonoidea. Classification doubts started with the first stephanid in the scientHc literature, described by Zschach (1788) in the Ichneumonidae, but later transferred to Braconidae by Fabricius (1804), and then placed among the Evanioidea by Jurme (1807). Leach (1815) was the first to propose classifying stephanids (then known firom a single species) in. a separate fiunily, the ‘*Stephanida” (first used as ‘*Stephanidae” by Hialiday 1839), but he also mcluded the ichnoimonid genus X orides m his new taxon, and again classified the group as part of Ichneumonoidea. Westwood (1841) was the first author to formally investigate the problem, dissecting and comparing specimens, but he concluded that stephanids should be transferred to Braconidae. Westwood’s proposition was not fiiHy adopted, but seems to have been mfluential in that ichneumonids were not associated with stephanids thereafter. Some doryctine braconids were later treated as stephanids (e g., Kiefter 1908, Enderlein 1901, Kiefter 191 la. Strand 1911), but Elliott (1922) transferred these taxa to a difterent fiunily, the “Stenophasmidae,” and provided the first accurate diagnosis for the family Stephanidae. The notion that stephanids were somehow related to the Ichneumonoidea, however, persisted until recently (e.g., Townes 1949, Carlson 1979, De Santis 1980), being convincmgly dismissed only under phylogenetic «amination (Rasnitsyn 1980, Shark^ & Wahl 1992). Currently, the mostly widely adopted hypothesis suggests that stephanids are best classified in then own superfiunily, the Stephanoidea Rasnitsyn 1980 & 1988, Gibson 1985, Naumann 1992). The family has also been treated as part o ( or closely related to the Evanioidea (forme 1807; Smith 1861a, b; Cameron 1903), Megafyridae (Ashmead 1900b, Riek 1973), and Orussidae (Ashmead 1900b, Hanson & Gauld 1995). Some authors suggest that the fomily may be the sister group of all other ^o crita, or at least one of its most basal taxa (Gibson 1985, WMtfidd 1992), but this idea apparently is not unambiguously supported when all available evidence is considered (Ronquist 1999). This problem is addressed in greater detaü m the present work. Stephanids have been important in sthnulatmg discussions about the classification of the Hymenoptera, and, more recently, about the phylogeny of the order, hi this latter area, th ^ are mteresting both as a taxon for which origin and relationships remain to be convincingly theorized, and as a group composed of what seems to be a peculiar combination o f highly derived and highly ancestral features. Such characteristics contribute to generate ambiguous results in pf^logenetic investigations, which test and encourage more research. For example, stephanids appear as part of Symphyta in Dowton & Austin (1994), as an intermediate taxon between Symphyta and Apocrita in VOhelmsen (1997), and as a group apical to the Ichneumonoidea in Ronquist e t a i (1999). No other fiunily in the order is interpreted in such variable way under cladistic analysis, and the wide range of possibilities indioite that whatever the &ial conclusion is, important implications may derive fbr the understanding of the evolution of other Hymenoptera. In more pragmatic terms, however, the problem of most immediate concern, and which also remains mostly unresolved, is the establishment of a sound classification for stephanid taxa, especially a well supported mterpretation fbr its genera. Opinions about the delimitation and ^onym y of the 11 genera proposed m the literature vary enormously, ranging firom the adoption (Schletterer 1889a) or defense (Ceballos 1926) of a single genus in the fiunily, to intermediate (Townes 1949) or consenting (Orfila 1956) mterpretations, or proposition of genera based on octreme remterpretations o f the relevance of taxonomic characters m stephanids @enolt I9Sla), Expectedely, such conflicting literature Is often confusing and published data on misldentlfied stephanid taxa remams a common & ct This is particular true, for example, with the genera Hemistephanus and ParastephanelluSy which have been ftequentfy^ confounded with each other (see chapter 3). The problems discussed above, and the now evident limitations of traditional taxonomy in generatmg a meanmgfiil and well-supported classification for the group, show that intuitive interpretations o f characters are particularly inefiScient when applied to stephanids. This situation cleaT invites a réévaluation under the more rigorous principles and methods o f cladistics. CHAPTER I PHYLOŒNETICS AND CLASSMCAnON OF THE FAMILY STEPHANIDAE (HYMENOPTERA) The main objective ofthis chapter is to mvestigite and discuss the intemal phylogeny of the 6mily Stephanidae accordmg to cladistic principles and methods, and to review the genus level classffication of the family based in these results. MATERIALS AND METHODS Specimens and museums This work is based on the examination of over 3000 specimens of Stephanidae borrowed 6om 112 muswims and private collections worldwide listed below with codens from Arnett e t aL (1993). Codens between brackets refer to collections not mentioned m Arnett's book. ABSC Archbold Biological Station Collection, USA D^rop) AEIC American Entomological Institut^ USA Wahl) AMNH American Museum of Natural History, USA ^ Smethurst) AMNZ Auckland Institute and Miseum, New ZIealandQ. Eariy) AMSA The Australian Museum, Sydney South N.S.W. Moulds & B. Day) ANIC Australian National Insect Collection, CSIRO (I. Cardale) ANSP Academy of Natural Sciences of Philadelphia, USA (D. Azuma) BHMH Universidade Federal de Minas Gerais, Brazil (P. Silveira) BMNH British Museum of Natural History, England (L. Tarel, S. Lewis, C. Taylor, D. Notton) BPBM Bishop Museum, Hawaii, USA (G. Samuelson) CASC Califismia Academy of Sciences, USA OD. Ubick & W. Pulawsld) CDAE California State Collection of Arthropods, USA (I. Sorensen and T. Tyler) CHAH Henry A Hespenheide private collection, Los Angeles, California, USA CIDA Albertson College of Idaho, Caldwell, USA (W. Clark) [CIRD] Centre de Coopération Internationale en Recherche Agronomique, France (G. Delvare) CMNH The Carnegie Museum ofNatural Hstory, USA (I. Rawlins) CNCI Canadian National Collection of Insects Q. Ihiber) CSUC Colorado State University, USA (B. Kondratieff & S. Rtzgerald) CDASC] Clement E. Dasch private collection, New Concord, Ohio, USA DEIC Deutsches Entomologisches bstitut, Germany (S. Blank) DZUP Universidade Federal do Paranà, Brazil (V. Graf & G. Rosado-Neto) EAPC Escuela Agrfcola Panamericana, Honduras (% Cave) EBCC Estaciôn de Biologia “Chamella,” Mexico (A Rodrigues) EDNC North Carolina Dept of Agriculture, USA (K. Ahlstrom) EMEC Essig Museum of Entomology - California, USA (R. Zuparko) ^fU S Utah State University, USA (W. Hanson) FCDA Fresno County Dept, of Agriculture, USA (N. Smith) FCNI Forestry Comission NSW, Australia (C. Urquhart) FCTH Forestry Tasmania, Australia (D. Bashford) HOC Fundaçâo Instituto Oswaldo Cruz, Brazil (S. de Oliveira) FMNH Held Museum ofNatural Hstory, Chicago, USA (P. ParrEo) FSCA Florida State Collection of Arthropods, USA (J. Wley) GBFM Unwersidad de Panama (R. Cambra & D. Quintero) HNHM Magyar Termeszettudomanyi Muzeum, Hungary (Z. Lajos) IMLA Fundacion Mguel LEo, Argentina ^ De Correa & C. Berta) INBC Instituto Nadonal de Biodiversidad, Costa Rica (I. Carvajal) INKS Ulihois Natural History Survty, USA Methven) INPA Distituto Nadonal de Pesquisas da Amazonia, Brazil (J. Ra6eQ 8 IRCW University of Wisconsin, USA (S. Krauth) ISUI Iowa State University, USA (R. Lewis) IZAV Distituto de Zoologie Agricola, Venezuela (J. Garcia) LACM Los Angeles County Museum ofNatural History, USA (EL Snelling) LSUC Louisiana State University, USA (V. Moseley) MAFC ManAedo Fritz private collection, Argentina (incorporated to IMLA) MCPM Milwaukee City Public Museum, USA (P. Quistorf) MEMU Mississippi Entomological NDiseum, USA (T. Schiefer) MLPA Universidad Nadonal de La Plata, Argentina (J. Schnack & M. Loiacono) MLUH Martin-Luther-Universitat, Germany (K. Schneider) MNHN Museum National dlffistoire Naturelle, France (C. VQemant) MNMS Museo Nadonal de Ciendas Naturales, Madrid, Spain (I. Izquierdo) MNNC Museo Nadonal de Historia Natural, Chile (Fresia Rojas A) MTEC Montana State University, USA (M. Ivie) MUCR Universidad de Costa Rica, Cd Universitaria “Rodrigo Fado” (EL Lezama) MVMA Museum of Victoria, Australia (K. Walker) MZUP Museu de Zoologie da Universidade de Sio Paulo, Brazil (C. Brandio) NCSU North Carolina State University, USA (R. Blinn) NHMW Naturhistorisches Museum Wien, Austria (S. Schôdl) NHRS Naturiiistoriska RDcsmuseet, Sweden (L.-A. Janzon) NMNH National Museum ofNatural Hstory (Smithsonian), USA (D. Smith & D. Furth) NSWA Biological & Chemical Research Institute, Australia Fletcher) ODAC Oregon Department of Agriculture USA (R. Westcott) OSEC Oklahoma State University, USA Arnold) OSUC The Ohio State University Insect Collection, USA (A Sharkov & N. Johnson) OXÜM Oxford University Museum, England (C. O’Toole) PMNH Peabody Museum ofNatural Hstory, USA (L. Munstermann) PPCD Plant Industries Division, WV Dept, of Agriculture USA (S. Clark) PSUC Pensylvania State University, USA ^ Kim &D. Skipper) QBUM Miseu Nadonal, Rio de Janeiro, Brazil (M. Monn^ QCAZ Quito Catholic Zoology Museum, Ecuador (G. Onore) QDPC Queensland Department of Prhnary Industries, Australia Q. Donaldson) QMBA Queensland Miseum, Australia (C. Burwell) 10 RMNH Nationaal Natuurhistorische Museum^ The Netheriands (K, van Achterberg) ROME Royal Ontario Museum, Canada(B. H ubl^) RÜIC Rutgers - The State Universfty of New Jersq r, USA (M. May) RWSC Robert W. Surdick personal collection. Bethel Park, Pennsylvania, USA SAMA South Australiam XAiseum, Adelaide, Australia (E. Matthews) SAMC South Afiican Nbiseum, Cape Town, South Afiica (M. Cochrane) SANC National Collection of Insects, South Afiica (C. Eardley & R. Urban) SCSC St Cloud State University, USA (R Gundersen) SEAN Servicio Entomologico Autônomo, Nicaragua (J.-M. Maes) SEMC Snow Entomological Musoim, Kansas, USA (R. Brooks) SFVS Cal%)mia State Unwersity, Northridge, USA (P. Bellinger) SMFD Forschungsinstitut Nat Senckenberg, Germany ^-P . Kopelke) SMNS Staatliches Museum fiir Naturkunde, Germany (T. Osten) SMTD Staatliches Museum fOr Tierkund^ Germany OR Eck) SOIC Sultanate of Oman Insect Collection (M. Gallagher, S. Al-Farsi, K. Salim) STRI Smithsonian Tropical Research Distitute (A. Aiello) TAMU Texas A&M University, College Station, USA CB. Ril^) UADE University of Aransas, Fayetteville, USA (C. Carlton) 11 UAIC University ofArizona, Tucson, USA (C Olson) UCDC University of California - Davis, Bohart Museum, USA (S. Hqrdon) ÜCFC Unwershy of Central Florida, USA (S. Fullerton) UCMC University of Colorado Museum, USA (V. Scott) UCMS University of Connecticut, USA (J. O’Donnel) UCRC University of California - Riverside, USA (S. Frommer) UGCA University of Georgia, USA (C. Smith) ULKY University of Louisville, Kentucky, USA (C. Covell) UMMZ University of Michigan, Museum of Zoology, USA (Name not provided) UMRM University of hfissouri, Columbia, USA ^ Sites) UNCB Universidad Nadonal de Colombia (Eduardo Florez D.) UQIC University of Queensland, Australia (M. Schneider & G. Daniels) UWIC Trinidad and Tobago, University of West Indies (C. Starr) WAMP Western Australian Museum (T. Houston & D. Jones) WARI Duncan Swan Insect Collection, Australia (P. Dangerfield) WFBM University of Idaho, WJF. Barr Entomological Museum, USA (F. MerickeO WSUC Washington State University, USA Zack) ZFMK Zoologisches Forschungsinstitut und Museum Alexandre Koenig Germany ^ -H . Lampe) 12 ZMAN Instituut voor Taxonomische Zoologie, The Netherlands (W. Hogenes & A. StoeQ 2^MHB Zoologisches Museum an der Humboldt Universitât, Germany (Name not provided) ZMUC Zoologisk Museum, University of Copenhagen, Denmark (L. VHhelmsen) ZMUH UnivershSt Hamburg Germany (EL Abraham) ZSMC Zoologische Staatssammlun^ Germany (J, Schuberth & H. Fechter) All specimens were barcoded, and databased using theM opsus relational database system developed by Johnson & Musetti (2000). Liformation and literature associated with the spechnens or taxa can be assessed via http://iris.biosci.ohio> state.edu/projects/stepfumids. Identification Taxa examined for this work are listed in Table 1.1. Identification of species was Ihnited by the feet that about one tfiird of the type spechnens of stephanids are now missing or were inaccessible. Since indhddual morphological variation in stephanids is higft, and since only a few spechnens could be exammed fer most taxa, I judged appropriate to designate voucher specimens for all species included in the cladistic analysis (Table 1.1). Interpretation o f character states were based nuiinly (or, in some cases, exclusively) on the study o f these specimens. 13 Sdccms Vouchers Depositories “Diastephanus’^alutaemu OSUC2I648,21649,216S0 Continued on next page. Table 1.1. Spedeaeauninedfbrthe cladistic analyst^ database number, and depository museum for correspondent voucher or type specimens. Spechnens are females, except where indicated whh the mate ^mbol. HT, ho lo ^e; NT, neotype; LT, lectotype. Gmeric names used as in Elliott (1922), to mdicate representatweness o f exammed taxa. Genera or generic placements considered invalid m this woric are cited between quotation maks. 14 Table I.l (Continued): Species Vouchers Depositories ’TUtosteplumus"'alluamii OSÜC22733 NT, 22734-22740 MNHN “NeosU^iantts''sp. A OSÜC21392 CASC PanutephmeUus denotatus OSÜC21042,21043.21044.2l045rf. 21046 QMBA.AMSA Parastephanellus granulatus OSUC20941.20942 ANIC. WAMP Parastephanettus nigrieauthts OSUC2I364.21365 BPBM, CASC Parastephanellus pygmaeus OSUC2I575«f, 21576 UQIC, SAMA Parastephanelba rujbonuitus OSÜC20912, 209l9«f, 20930 ANIC PantsUphaneUus sp. A OSUC 21362. 21363 CASC Panutephaneilus n. sp. OSUC2137S BPBM SeUettererts emetipes OSUC20695,20827cr, 20834 CNCI, EMEC SM ettm ris determmatoris OSUC2I616 H r HNHM “Sttphamu" eUiotti OSUC21665 NHMW “Stephamts" gigfu 0SUC21657HT NHMW “Stephanus " pachylomerus 08UC21622.21O3 CMNH.C1RAD Stephanus serrator OSUC20587,20588.20589cf. 20593cf, 21617 ZMAN. NHMW. SMNS “Stephanus" seyrigi 0SUC21663HT.21662f MNHN “Stephanus" tibiator OSUC20951,20952 CASC, NHMW “Stephanus " xanthocephalbis OSUC20944.20945,20946.20947 Selection o f taxa for the amfysis Cladistic anafysis was performed at the species level. Coding and evaluating characters at genus level was not a practical option, since there is not a reasonable consensus on the delimitation most stephanid genera in the literature (e g., a single genus recognized in Schletterer 1899a versus eight genera in Elliott 1922). G enom e species of all genera described m the literature were mcluded in the analysis, but the selection of species was expanded to mclude a widely representative sample o f the morphological types within the ânûly, basal species, and groups from distinct geographical locations. No special attention was devoted to achieve a thorough geographical representation, but selected taxa are from all major biogeographical regions. 15 Selection ofParastepfumelhts taxa was aided by a detailed taxonomic mvestigation o f the Australian species o f this genus (chapter 3), Many species of ParastephamUus other than those from Australia also had to be examined for the mentioned analysis, and this genus could therefore be comprehensively evaluated, providmg the basis for an accurate choice of representative taxa. Male specimens Males o f the selected spedes, when availably were used only in an od hoc basis, generally to help confirm ambiguous character states in female specimens. Characters from males, however, were not dhectly considered because data associated with this sex m stephanids are excessively mcomplete or inaccurate due to the following problems partially discussed in Aguiar 1998); (I) much greater and more irregular variability in suifrtce sculpture, body proportions, and color patterns than in females; (2) average size generally much smaller than females, resultmg usually in more ambiguous character state mterpretation than may be achieved with females; and (3) males of most species are unknown. Selection o foutgroups Exact relationships of stephanids with other Hymenoptera are unknown (see Roquist e t aL 1999, and discussion in this woric). Consequently, there are no obvious sister groups known for this fiunily. One commonly adopted solution for this situation is the consideration o f a hypothetical outgroup (e.^ Shaw 1990). Within Stephanidae, however, there is some evidence mdicating a basal position for Schlettererius P^hmeaây as well as a considerably 16 large evolutionary gap between the species in this genus and the remaining extant stephanids. The basal position of Schlettererius emetipes was initially suggested by Townes (1949), and is further supported by the Act that a series of distinctly unspecialized features o fHymenoptera occur in both known species ofSchlettererius. These same features are distinctly specialized or extremely rare in other stephanids (Table 1.2). At the same tune, a prelhnmary analysis indicated that numerous morphological features characterize Schlettererius as a taxon which is intensely distinct from other actant stephanids, an observation which was confirmed in this woric (see discussion fbr this genus). Although only the correct rooting of the tree can define whether such features are synapomorphies fer Schlettererius or fer the remaining stephanids, the feet remains that in either case a long series of synapomorphies separate one group from the other. Combined, these considerations suggest that a “distantly basal” position of Schlettererius within Stephanidae can be reasonably hypothesized a prio ri, making this genus a credible reference pomt to root the tree (fr was also used as a reference to the generation of preliminary polarfeation hypotheses, explained m the hem “Character coding and polarization,” below). Note, however, that the final cladogram was rooted between Schlettererius emetipes and S. detemrinatoris. This is an artffîce to allow the length of the branch (ecpiivalent to number of ^napomorphies) between Schlettererius and other stephanids to be visualized. 17 Sdikttererius______Other extant stephanids ______1.T2 and S2shoit and broad, isolated L Fused into a long cylindrical petiole 2. Pronotum short, blunt 2. Pioiiotum distinctly elongate, with several socialized regions and structures (colo, preannutar, semiannular, pronotalfold, etc.) 3. venation conqilete^g. 163), vehis tubular 3. Mary ndmdous and speüral veins 4. ffiid tibia normal, onffirmify shaped 4. From strongly consiessedbasalty and dilated Sically> to extremely specialized in many shapes 5. Both sexes with five tarsomeres on legs, all with S. Females with three tarsomeres on hind leg, each similar shise and length one with a distinct shape and length (except in £ ______serrator)______ Table 1% Distinctly unspedalized features of Schlettererius, compared with remaming extant genera o f Stephanidae. Selection o fcharacters Only external morphology was considered, but all genus-level and most species- level characters ever used in the taxonomy of stephanids were investigated. The search was also aimed at finding and investigatmg new characters, and for this reason an inclusive approach was adopted: characters were abandoned only in case of great difificulty or unpossibOity^ in defining reasonably discrete states. Li spite of some opposition about the use of biometric characters in phylogenetic analyses (eg., Pimentel & Riggins 1987), a few o f them have been considered here (chs. 3,5,47,72, 84,96,110, 115,150,151,168). It seems obvious that structures with notorious variation in shape or relative proportions between taxa may be phylogenetically significant. The major problem is that character states cannot be objectively defined unless a distmctly non-overlapping, or at least bhnodal or polymodal distribution of the 1 8 proportions is observed. However» ^two extreme studies occur (e.g., all characters mentioned above except chs. 5 and 150), then it seems evident that at least two character states do exist, and can be ecplored. In such context, measurements and ratios are probably more useful not in ‘‘reducing” the subjectiveness o f decisions on how to delimit each stat^ but rather in allowing these decisions to be used consistently. The consideration o f biometric characters also seems important in that evidence is not being ignored. Albeit imprécisions can be generated by transitional forms, usefol information is also generated by the foct that distinctly different states will be separated by deferent coding and therefore contribute to the final result. Note that transitional forms are also common for qualitative characters, contributing to make initial hypotheses about them shnilariy subjective. For both types of characters, however, there are no reasons preventing the data of being repeatedly reinterpreted according to the results generated in each step o f the analysis, thus generating better supported results each time. This process corresponds to the “checking correcting and re-checking approach of Hennig (1965). Character coding avipolarization Characters were initially interpreted separately, with each one coded for all taxa before the next character was considered. Character states occurring sporadically, or in relatively fow specimens o f a taxon, or which represented anomalous developments (e.g., spurious veins) were not recorded as polymorphisms. Proportions or shapes could be interpreted dh-ectly in most cases (chs. 5,47,72, 115, ISO, 168), but some presented ectreme and contmuous variation between taxa, and were interpreted with objective mrasurements, made with an ocular micrometer (chs. 3, 19 84,96,110,151). During character search and codm& and before any cladogram was obtained, the ordering and polarization of states for each character were hypothesized. Polarization hypotheses could also be proposed for all biometric characters because in all cases only one state was present m the outgroup taxa (see also item “Selection of outgroup taxa” above). The completed matrix was then submitted to heuristic searches in Hennig86 v. 1.5 (Farris 1988), first whh all characters unpolarized, then whh characters polarized according to the mhial hypotheses. Character distribution on the resulting cladograms were then examined in Clados version 1.6.1. (Mxon 1998) and evaluated. New possibilities for ordering and polarization hypotheses were then considered, implemented in revised matrices, and resubmitted to Henm*g86. The results were compared again, and the process repeated several times, until polarization hypotheses could be decidedly sustamed (or not) for all characters. Tree searching in Hennig86 was performed whh the commands “mh*; bb*;” which perform the most exhaustive heuristic search available in Hennig86 (Lipscomb 1994). The resultmg most parsimonious tre% were submitted to successive approximations character weighting with the commands “xs w; mh*; bb*;” until results did not change m three consecutwe herations. Hermig86 was used only as a prelhninary way of assessing the coding scheme devised for the characters, its mam adavantage bemg that results for heuristic searches could be obtained quickly. Polymorphic entries were changed to Q.e., “missing data”) b ^ r e the matrnc was submitted to Hennig86, since h does not process this Qrpe of data. 20 As polymorphic entries were extremely rare m the matron their overall influence was negligible for the mtended purpose at this stage. All programs used were executed from withm Dada version 1.2.7. (b&con 1995). Observation and illustration o fcharacters The intricate sculpture patterns and the complex structure of the pronotum provide taxonomically hnportant characters for stephanids, but are also considerably dffîcult to mterpret and illustrate accurately. For this reason, special attention must be given to the use of adequate illumination while observing, drawing or photographing stephanid specimens through the stereomicroscope. Of many dËforent ^ e s of lights I tested, none proved more efiScient than conventional fluorescent illumination, which can improve resolution dramatically when compared, for example, with the light used in fiber optics light ^rstems. Similar results are discussed by Vincent & HofSnan (1974). Cuticular sculpture and püosity must be evaluated with great care on greasy specimens, which is often the case with dried stephanids. Grease in these insects is usually not immediately evident, but mteiferes optically with the microsculpture, and easily collapses hairs, especially the small ones (e.g, pruinosity), making microsculpture and pilostty patterns often totally invisible. For this reason, a 6-12 hours immersion in pure detergent might be necessary for adequate cleaning o f some specimens. After the detergent treatment, specimens must be rinsed at least twice in water, and then dehydrated in pure alcohol or acetone before it can be safely dried again. Cuticular colors do not seem 21 to be dnectly affected, but prolonged (2-3 days) or repeated treatments can afect or destroy structures or tissues attached mtemalfy, and therefore m od^ the way colors are perceived (e.g., genal stripe). Wing illustrations were prepared from slide mounted wings. Slide preparation and illustration of stephanid wings demand particular attention in two aspects. (1) The wing membrane on the costal area of the front wing is always rolled or folded back and upwards, and is also considerably sclerotized, making it difBcult to flatten out during dide mounting, frnmersing the wmg in a 20% solution o f KOH for one or two minutes wül soften the membrane, making it much easier to unfold. Unfolding is important because it allows the observation of the pfloshy on the costal cell, as well as its shape. (2) In many stephanid species nearly half of the front wing veins, and usually all hind wing veins except Sc+R, are entirely spectral, and will disappear completely m conventional slide preparations. To solve this problem, the following technique was developed: A drop of any water-soluble glue is placed on a slide, a clean and dried wing placed over the glue (without anything covering it), and the preparation left for 24 hr in a humid chamber. The humidity allows the glue to slowty and un^rm ly spread under the the wing. The different refraction indexes from the air on the dorsal surfoce of the wmg and the glue on its ventral surfoce produces an optical ^fect that fecOitates the observation of spectral veins. Drawing conventions Spectral veins (Mason 1986), by definition, do not have pigmentation and therefore can only be illustrated in a schematic way. T h ^ are represented here by light stippling (Figs. I7S-I78) mdicatingthe trajectory and approximate thickness o fthe vein. 2 2 Nebulous veins are indicated by & distinctly denser stippling (e.g.. Fig, 173-174). Folding lines on wings are represented by single longitudinal dotted lines (e g., CuP in Figs 173- 178, dorsal and ventral wing folds in Figs 184, 185). Illustrations o fwings or details of venation and other structures correspond to the right win& dorsal side; other illustrations correspond to the left side or structure, except otherwise mdicated. TERMINOLOGY The followmg terminology was revised and expanded from Aguiar (1998), based mostly on Richards (1956) and Gibson (1985). Wing venation terminology follows Sharks & Wharton (1997), with the different ^ e s of venation described with the terminology proposed by Mason (1986). Vein CuP is interpreted according to Mason (1990). Terms used to describe cuticular sculpture follow Harris (1979). Antennal sensillae (Figs 8-26). Two types of multiporous plate sensOlae are observed on stephanids, and th ^ are referred here as follows. Type Isem illaei always large and conspicuous, usually circular or elliptical (Kg. 18), but somethnes shaped as a transverse stripe ^ig. 14). Type UsensiUae: very small, often similar to “punctures” in between Type n sensillae (e.g.. Figs 13,23), each one bearing a single mmute hair ^ g 15). Frontal carma (Fig. 2). A weak centro-longitudinal carina on ffons, usually formed by convergence o f the general sculpture. Paraocular strication. (Kg. 2; see also chapter 3). A series of short, almost regular, transverse strigae on ffons, along ^ e margm. 23 Coronal area (Rgs 2,31-36). The raised area atop the frons, delimited by the five coronal tubercles, and containing the central ocellus. Tubercles (Figs 2 ,3 ,3I-3Q. Conspicuous cuticular projections on top of head, which delunit the coronal area; one anterior and two lateral tubercles always stout and acuminate; posterior pair usually rounded, sometimes connected to each other mesally. Anterior ocellus (“coronal ocellus” of some authors) ^ g s 2,3,31,32). The ocellus within the coronal area. Literocellar carinae (Figs 3,33, 35, 26). The transverse carinae in between or immediately behind the posterior ocelli; at least the most anterior one strong, blade-like, as wide as the distance fi"om one posterior ocellus to the other. Vertac (Rgs 3,33-36). The very large area between qres, posterior ocelli, and occipital carma, nearly always covered with conspicuous, extremely variable sculpture. The interocellar carinae are described independently, so that “anterior” on vertex refers to the area immediately behind them. The vertex of Stephanidae is commonly referred to as “occiput” in earlier works (e.g., Elliott 1922, Orfila 1949). Centro-longitudinal impression (Fig. 3). A narrow depression that crosses the vertex centro-Iongitudmally; usually seen only with tangent fllummation. Post-vertex (Aguiar 1998:380-1, Figs 55,59). The posterior region o f vertex; usually mdistinct, but sometimes fiat, negatwely inclmed in relation to the main area of the vertex, with weak sculpture or polished smooth (ixttaSxiHemistephams spp.) Genal angle (Rgs 2,3). The angle produced by a bulging or acummate gena. 24 Genal stripe illustrations in chapter 3). A usuaify yellowish or ivory stripe that crosses the gena from base of mandible to templ^ extending alon^ but not connected to, the e y t margin (most Parastephanellus spp.) Occipital carina (Figs S, 27,28,33-39, etc). A conspicuous carina between vertec+gena and occiput; always present in stephanids, but width and apical (= ventral) d^ferentiation highly variable, hi the literature often described as “bordered” when very narrow, looking as a linear structure, and “not bordered” or “collar-like” vdien projected as a sharp, wide or very wide edge. Occiput (Rgs 5,40,41). A relatively small and deeply concave area between occipital carma and fbrame magnum, contiguous with postgena and postgenal bridge. Postgena (Fig. S). The area ventral to the occiput and posterior to the gena, between bypostomal and occipital carinae. Postgenal bridge (Richards 1956) (Figs 5,38,39,40). Area formed by the connection of the right and Idt postgenae. The postgena in stephanids is exceptionally elongate when compared with that of other Hymenoptera. Postgenal line (Rgs 5,38,39,40). The centro-longitudmal divisory line on postgenal bridge fiirmed along the connection (not fiiston) of the right and left postgenae; distinct ftom the base of the l^ostom al carma to the foramen magnum. Hvpostomal carina ^ g s 5,49 ). A blade^like carma bordermg the oral fossa, getting progressively wider anteriorly. Pronotum ^ g s 4,6,54-82). hi stephanids, normally a very elongate and highly specialized structure with several distinct regons and structures, deRied below. 25 Apical carina (Kg. 4). The upturned apical margin of the pronotum; varies from a simple dorsally refiexed border (as Figs 60,61,66,67) to a convoluted, broad and microsculptured structure, sometimes much wider laterally than centrally (as Figs 59, 62) Pronotal fold (or “central fi)ld” of some authors) ^ g s 4,6). A variously differentiated transverse cuticular elevation, which is shallowly to deeply invaginated anteriorly, located subapically or submedially on the pronotum. Sometimes collapsed centrally, an in Parastephanellus spp. Colo ^ g s 4,6). The area of the pronotum between apical carma and pronotal fold; usually moderately concave dorso-longitudinally, contiguously with the invagination of pronotal fold. A variable number o f transverse carinae are normally present on both sides of the colo. These are described always from apex to pronotal fold, so that the first pair is the one hnmediately behind the ^ical carina; when continuous from side to sid^ thus appearmg as a single structure, a carma is said to be complete; fr^interrupted dorsally, or not reaching the opposite side, incomplete. Semiannular (Elliott 1922, Orfila 1949) (Rgs 4 ,6).The large, mfiated, round (i^proximately a 1/4 sphere) posterior part of the pronotum. Preannular (Rgs 4,6). The short, usually transverse area between (and contiguous with) pronotal fold and semiannular. Preannular ruga (Aguiar 1998:380, Rg. 56). A strong transverse rugosity between preannular and semiannular areas. Preannular impression (Aguiar 1998:380, Rg. 56). A transverse depression behind the preannular ruga (some Hemistephanus spp. and M egischus spp.) 2 6 Pronotai lobe ^ g s 4,6). The dilated upper posterior corner of semiannular, preceded by a depression which is usually coarsely transverse^ strigate. Femoral rnipression (Fig, 6), A longitudinal, and slightly oblique depression which nins lateralfy, from immediately behind the pronotal fold to the ventro-posterior margin of the pronotum, Ading in its final third. Ventral area ^ g . 6). The narrow, longitudinal, fiat area behind pronotal fr>ld and ventrad o f femoral hnpression. Prostemum (Fig. Q. The two ventral elongate hemistemites of the prothorax, which in stephanids are apically globularly or triangularly prominent. Prosternai depression (see Aguiar 1998:381, Fig. 61). A central depression on prostemum, immediately behind its prominent apex, usually as long as 0.2-0.4 of the posteraum length. Mesoscutum (Figs 92,95, M e). A semicfrcular or narrowly transverse dorsal area between the semiannular and the transscutal articulation. With an anterior section mostly or entirely concealed inside the semiannular in live specimens, but usually at least partially exposed in dried ones; this part usually finely transversely alutaceous. Median sulcus (Fig. 96 nur). A centro-longitudinal row o f closely aligned fovea or punctures on the mesoscutum, i^diich is sometimes partially hidden among coarse surfece sculpture; often weakfy^ dififerentiated or absent in species o fFoenaiopus; otherwise usually conspicuous. 27 Notaulus (Fîg, 96 n t). The oblique row of closely aligned fovea or punctures laterad of median sulcus; sometimes partially hidden among coarse sur&ce sculpture; often weakly dK&rentiated or absent in species o f Foenaiopus; otherwise usually conspicuous. Median lobe (Rg. 96). Area o fthe mesoscutum in between the notauli, containmg the median sulcus. Lateral lobe ^ g . 9Q. Each of the areas of the mesoscutum external to the notauli. Parapside ^ g . 93 p p ). The small and more or less rectangular areas next to each tegula; partially fused to the lateral lobes. Transscutal articulation (Fig. 92). A straight, wide, and always distinct sulcus between mesoscutum and axillae. Axilla (Fig. 92 A x), Each of the two large and somewhat triangular areas in front of the scutellum, and immediately posterior to the transscutal articulation; separated from scutellum by a series of deep crenulae. Scutellum (Figs 102,104, S c), fri stephanids, a composite sderite behind the axillae, formed by a larg^ glabrous, shmnin^ suboval central area, followed, on each side, by a triangular, densely pilose area, which is in a distinctly lower level than the central area. In the literature descriptions of sculpture patterns on the stephanid scutellum are virtually always referring to the central area only. Mesepistemum (Figs 100,101). The largest sclerite on thonu^ situated laterally between pronotum and metapleuron; dorso-posterioriy always concave and elongate usually polished smooth or at least partially so. 2S Mesopleural pit (sense Masner 1979) C^gs 101 mp^ 102, 103), A small central pit on the mesepistemum, surrounded by a few short rugosities (Foenatopus spp.) Mesopseudostemum (sense Shcherbakov 1980, Gibson 1993) (not illustrated). The ventral part of the mesepistemum; larg^ flat or weakly concave glabrous. Discrimen (Rg. 110 ok). A centro-longitudinal line on the mesopseudostemum; sometimes formed by aligned foveolae. Extends from the paracoxal suture to the base of the pectus. Paracoxal suture (Gibson 1993), The narrow transverse cuticular invagination near and along the basis of the mid coxae. Pectus (not illustrated). The broad and nearly flat anterior free of the mesepistemum, which is m contact with the posterior frees of the front coxae. Metanotum. (Fig. 104 A6%). A narrow, small, transverse sclerite mostly hidden behind and under the scutellum. Crenulate sulcus (Rg. 7). A wide transverse depresrion formed by a series of aligned, very deep crenulations, situated between metanotum and propodeum, crossing the entire dorsal width, endmg at the meeting pomt of metapleuron, metanotum, and propodeum, in a large and deep fovea (the pleuropropodeal fovea). Pleuropropodeal fovea (Rg. 7). A usually large and deep fovea at %ch end of the craiulate sulcus, maridngthe meeting point of metapleuron, propodeum and metanotum. Propodeum (Rg. 7). The large weakly convex dorsal area posterior to the crenulate sulcus, containmg the spiracles, spiracular sulcus, spiracular carina, and the parapetiolar depressions. Referred to as "median segment” ozKûttelsegment in most of the eariier literature on Stephanidae. 29 Flank 7). Each of the antero-Iateral comers of the propodeum; sculpture usually distinct and finer than that on the main area of the propodeum. Spiracular groove (Fig. 7). An elongate longitudinal depression in fiont of the spiracle, of variable length, but sometimes reachmg the crenulate sulcus; externally mariced by the sphacular carina. Treated under mai^ different names in the earlier literature on stephanids (“marginal suld,” “crenulate sulcus,” “stigmal suld,” “lateral impression,” “longitudinal impression,” or simply “a sulcus between metapleuron and propodeum”). Spiracular carina (Fig. 7). A longitudinal carina external to the spiracular groove, originatmg anteriorly on the propodeal spiracle, and usually following the trajectory of the spiracular groove. Parapetiolar depression (Fig. 7). Each of the usually sculptured depressions on each side of the petiolar foramen; usually subren^rm or suboval m shape. Metapleuron ^ g . 7). The large lateral, more or less triangular, convec or pyramidal area behmd the mesepistemum and ventrad of propodeum. Metaplqiral fovea (Fig. 7). The usually large depression occurring ventro-apically on the metapleuron, maridng the end of the mterfoveolar area; rarely absent hiterfoveolar area ^ g . 7). The oblique, weakly concave elongated area behind the mesepistemum, corresponding to the anterior edge of the metapleuron; delimited antero- dorsally by the pleuropropodeal fovea, and ventro-posteriorly by the metapleural fovea. Post-foveolar area 7). The ventral edge of the metapleuron, behind the metapleural fovea; usually beatmg a few strong transverse strigations posteriorly. 30 Median tooth ^ g . 128 mf), A conspicuous and usually broad triangular tooth placed midway o r subbasaOy on the ventral edge o f hind femur. Apical tooth (Fig. 128 a t) . Usually an elongate» dagger-shaped or lanceolate tooth placed subapically on the ventral edge of hmd femur. Femoral denticles ^ g s 131,132). Small tubercles located between the median and apical teeth, and between the apical tooth and apex of hmd femur; more rarely also present basad of median tooth. Wing venation. Veins and cells as in Figs 161-162. All front wing veins illustrated on Fig. 161 are present in all species of Stephanidae but in apical taxa most o f them may vary in shape or become entirely nebulous or spectral, or show different proportions (see for example Rgs 173-178). Spurious veins are relatively common (e g., vein stub coming out of2ronFig. 167). Hamuli, ^ g . 149) Only distal hamuli occur, but two different types can be distinguished; T )p e I hamuli are apical, decumbent, sinuously curved, showing a somewhat flat apex ^ g . 149 tl)y 2}peZr hamuli are basal, erect, slightly curved, apec shaped as a hook (Rg. 149(0). Abdommal tergites and stemites (Rgs 143,144,147,148). The original sclerites of the abdomen, referred to as T2-T9 (unmodified abdommal tergites) and S2-S8 (unmodified abdominal stemites). These terms were preferred over “metasoma” or ‘^gaster” because th ^ allow fi)r objective discussions on morphological homolo^ between stephanids and 31 other Hymenoptera. This seems significant because of the apparently intermediate placement o f stephanids between Symphyta and ^ocrita» and because ^mpbytans, by definition, do not have a structure equivalent to a metasoma or gaster. Petiole (Rg. 145). The narrowed anterior stalk of the abdomen which articulates with the propodeum, whether composed of part o f a segment or the whole segment (definition slightly modified fi*om Richards 1957). It must be noted that for most stephanids the petiole is a single structure, resulting from the complete fiision of T2 and S2 into a long and narrow %ibe" (Rg. 145), but this is not the case for Schlettererius. In this genus, T2 and S2 are distinctly separated and much wider (comparatively to other abdominal segments) than in other stephanids (Fig. 143). It seems therefore debatable whether it can be described as a petiole. For this reason, I refer directly to T2/S2 in this work when commentmg about Schlettererius, Petiolar spiracles (Fig. 143, 145). The vestigial, sometimes inconspicuous tubercle visible on each side of the petiole (or T2), located slightly basad of the middle. Pvfridial impression (Hustrations in chapter 3). An U- or V-shaped depression apically on T9, at the base of the pygidium. Pvgidium illustrations m chapter 3). A small convM plate or elongate projection dorso- apically on T9, usually preceded by the pygidial impression. 32 MORPHOLOGICAL CHARACTERS AND OBSERVATIONS Charactw description is followed by a formal (üscussion, organized according to the following order (but not necessarily including all items): (I) polarization o f states for polymorphic characters, and whether th ^ were run as additive or non-additive; binary characters always considered additive; (2) optimization selected (Deltran or Acctran); (3) description o f major consequences or assumptions hnplied by the selected optimization, and (4) a justffîcation for the selected optimization; (S) for polymorphic characters only, a schematic description of major character changes as observed in the preferred cladogram (eg., “0-^1-^2-^r). Polarization hypotheses are provided for 126 characters; the remaining 50 were treated as unpolarized. Character states for chs. 44,45,49/50,94/95, 97,103, 112 were renumbered in the matrbc (see discussion for these characters for justification). Character optimization was based on cladogram depicted in Fig. 198 (see item ‘Thylogenetic Analysis’* for details about the optimization process). 000. Head, degree o f compressiom (0) globose or almost so (Fig. 27); (1) w«ddy but distmctfy compressed antero-posterioriy ^ig. 28); (2) strongly compressed antero posterior^ (Fig. 29). Non-additive. Ambiguous at node 36. Optimization is unimportant because correspondent clade is of reduced mterest for the present analysis. Changes as suggested m the preferred cladogram: 1MM2. 33 001. H e a d y c o lo n (0) mostly or entirely black or dark-brown; (1) red or ferrugineous; (2) yellowish brown, but vertex darkened (dark brown or black); (3) entirely yellowish brown or brown. Yellow stripes on rirons and gena treated as a distinct character (ch. 32). Non-additive. Ambiguous at node 38. Optimization is unimportant because correspondent clade is of reduced interest for the present analysis. Changes as suggested in the preferred cladogram: 1MM3, with several independent origms for state 2. 002. Flagellum, shape transition from basal to epical flagellomeresr. (0) all flageOomeres approximately of the same proportion, i.e., uniformly short, or unribrmly elongate (Rg. 11), or becoming proportionally and progressively less elongate towards apex. (Rg. 8); (1) first 8 basal flageOomeres distinctly, although slightly, more elongate than foOowing ones (at least some o f the basal 8 flageOomeres about 5x as long as wide; apical ones about 3x as long as wide) OFig. 9); (2) most of 2nd-8th basal flageOomeres much more elongate than apical ones; the change in shape from basal to apical ones often abrupt (basal ones 8-16x or more as long as wide versus usuaOy 3-4x for apical ones) (Rg. 26). There is a tenden^ in several species to have basal flageOomeres with a distinctly difTerent structure rirom the apical ones. Such tendenqr is even more evident in the pupal or pre-imago stage of some species (eg., Parastephanellus rufoorm tus, P . granulatus\ suggesting it may indeed have some biological relevance. The nature of this difference has prhnarily to do with how much longer and narrower each set o f flageOomeres is in relation to the other, bi state 4, the basal flageOomeres are sOghtly narrowed where t h ^ connect 34 with the next or previous ones, so that each flagellomere is easily recognized, but the apical ones are fully connected with each other, so that it is more difiScult to visualize individual 9%ellomeres. Non-addhîve. Optimization: DELTRAN. Makes state 0 the basal condition for stephanids, with states 1 and 2 arising several times m apical taxa; change (M2 provides additional support to the Parcatephanellus clade, where state 2 occurs for she o f its seven taxa (reversal 2-»0 in f. cctu d a tu s Szépligeti). Justification: states 1 and 2 are distinct, but subtle specializations; therefore, it seems more reasonable to assume that these states evolved more times in separate taxa, rather than representing fundamental and fixed evolutionary changes. Changes in the preferred cladogram after optimization: l*-0-»2. 003. F/age//um, proportions o f most elongate flagellom ere'. (0) short, not more than 5.5x as long as wide (Rg. 11); (1) very long and narrow, 9.5-l6.0x or more as long as wide (Rgs 26,9); (2) moderately elongate, 5.6-9.0x as long as wide (Fig. 8). The most elongate flagellomere m stephanids is normally the 4th fi-om base to ^ex; more rarely the 3rd or 5th. Measured in dorsal view. Additive. Opthnization: ACCTRAN. Makes state I the basal condition for non- Schlettererius stephanids. Justffîcation: it conforms better with the additive hypothesis by allowmg a l-»2 transformation for S tep h a n a s and a MO reversal at node 4, as opposed to the more assuming 0-*^2 transformation for S tep h a n a s which occur under altematwe opfimuafions. Changes as suggested in the preferred cladogram after optimization: 0 -^ M 2 . 35 004. FlageUomere 1, organization o f Type I sensillae: (0) absent ^ g . 15, 17,21,22); (1) several, distributed ventrally along entire length of flagellomere ^ g s 10,18,25); (2) a few or several, restricted to apical 0.4-0.7 (this portion usually more dilated than remaining of flagellomere) (Figs 13, 16, 19,20,23); (-) inapplicable ^ig. 12). Albeit with large sensillae absent on flagellomere 1, this character was coded as QU^plicable for "Diastephanus" sp. A because sensillae on flagellomere 2 indicate this species does not have the same generilized Type I sensillae. Ambiguous at nodes 3 and 21. InsufScient evidence to support optimization. Changes as suggested in the preferred cladogram: 1-^2 and 0-»2; relationship o f states 0 and 1 is uncertain. 005. Flagellomere 2, relative length: (0) about as long (0.9-1.0) as flagellomere 3 (Figs 11,8,13,18); (1) distinctly shorter than flagellomere 3 (Figs 15,24). Changes are unambiguous. 006. Flagellomere 2, organization o f Type I sensillae : (0) several, distributed ventrally along enthe length o f flagellomere which is approximately unbformly cylindric (Figs 10, 13,15,16,18,20,25); (1) a few or several, restricted to apical 0.4-0.7, portion of flagellomere with sensillae usually somewhat dilated (Figs 17, 19,21,22,23); (-) inapplicable (Figs 12,14). Optimization: DELTRAN. Makes state 0 the basal condition fer stephanids, and assumes that no reversals occurred; fevors restricted occurrences of changes 0-+1, 36 minimizing occurence of state 1 in major ancestral nodes; provides additional support to tb& Parast^fumellus clade. Justification: Deltran conforms better with the additive hypothesis by assuming state 0 for the ancestors o f Schlettererius and StephanuSy thus allowing to explain occurrence of state I m S tep h a n a s as a progression fi’om state 0, rather that considering the highly specialized state I as the basal condition. It also prevents the complicated assumption 0->I-^0 to explain the occurrence of state 0 m Xonthostephanus n. gen. Such sequence seems unlikely because o f the several difierences between these states: state 1 d^ers fit)m state 2 at least by havmg fewer sensOlae these located apically, and flagellomere swollen where they are present. 007. Flagellomere 3, organization o f Type I sensillae ', (0) several, distributed ventrally along enthe length of flagellomere (Figs 13, IS); (I) a few or several, restricted to apical 0.4-0.7 (portion with sensillae usually more dilated than where thqr are absent) (Figs 17, 19,21,18,20,22); (-) inqiplicable or absent (Rgs 14,23). Optimization: DELTRAN. Favors restricted occurrences o f changes 0-» 1, minimizing occurence of state 1 m major ancestral nodes; provides additional support to the ParastephaneUus clade. Justffîcation: it prevents the complicated assumption 0-»l-»0 to explain the occurrence of state 0 in several taxa (see also justification for ch. 6 r%ardmg Jûmthostephanus n. gen.) 37 008. Fkigellum, presence o f transverse, stripe-like Type I sensillae: (0) absent (Kgs. 10, 12,13, etc); (1) present ^ g . 14). Changes are unambiguous. 009. Flagellomeres 1-3, presence ofmicrodepressions on ventral face: (0) absent in all (Figs 10,12,18,); (1) present in at least one of them (Figs 16,19,21). Ambiguous at nodes 2 and 29. ^sufficient evidence to support optimization. 010. Apical flagellomeres, presence o f Type I sensillae: (0) present on all flagellomeres fi’om 4th on, ventrally only (Fig. 17), even ^only one sensilla per flagellomere; (I) present or absent on lst-2nd, absent fi'om 3rd-6th, sometimes also 7th and 8th, present on remaining apical flagellomeres (not illustrated); (2) present on all flagellomeres firom 4th to apical, mostly ventrally, but also laterally, with some reaching the dorsal side (Fig. 24); (-) mapplicable(Fig. 14). Non-additive. Changes as suggested m the preferred cladogram; 1^0-^2, unambiguous. 011. Flagelbm , frequence o f Type I sensillae: (0) numerous, distributed in a wide area (Kg. 12); (1) few or several, usually organized in a single or double line (Kgs 10,13,15, 16,17,18, etc). Changes are unambiguous. 38 012. Flagellomeres 1-2, presence o f Type II sensillae'. (0) absent ^ig. 12); (1) present (Rgs ID, 13,14, 20,17, 15.25). Changes are unambiguous. 013. Flagellomeres, pruinescence'. (0) all flagellomeres densely pruinos^ although basal ones sometimes pilose (e g., m S. tibiator) (not illustrated); (1) basal 6-8 flagellomeres glabrous or with sparse pQosQr or pruinosity; from 9th, 10th, orl 1th until the apical flagellomere, entirely pruinos^ (e g., M califomicus, M tex a n sts, M anomalipes, H. m acrurus) (not illustrated); (2) flagellomeres glabrous, except for pilosity associated to Type n sensillae, mostly on ventral side (not illustrated). Additive. Ambiguous at node 11. Optimization is unimportant because correspondent clade is o f reduced interest for the present analysis. Changes as suggested in the preferred cladogram: 0-*l-^2. 014. Frons, dorsal half, overall orientation o f macrosculpture (not Qlustrate Paraocular strigation treated as a separate character (ch. 18). Non-addhîve. Ambiguous between nodes 15 and 29. Possible opthnizations are of reduced hnportance for the analysis because character is exceedingly homoplasious. Changes as suggested in the preferred cladogram: lM)-»2. 39 015. Frons, dorsal half, type o f mcKrosculpture (not illustrated): (0) mixed (broken or discontinuous, or partly strigate, partly areofate-rugose); (I) parallel or concentric, unbroken strigae or rugae, even if very weak; (2) mostly or entirely areolate or areolate- rugose; (>) im^plicable (macrosculpture absent). Paraocular strigation treated as a separate character (ch. 18). Non-additive. Ambiguous between nodes 17 and 29. Possible optimizations are of reduced importance for the analysis because character is exceedingly homoplasious. Changes as suggested in the preferred cladogram: all changes possible. 016. Frons, ventral half, overall orientation o f macrosculpture (not illustrated): (0) mainly transverse, somewhat straight, even if rugose or areolate; (1) mixed directions, or non-directional (e.g., areolate); neither oblique nor transverse, or only inconspicuously so; (2) oblique or arcuate; (-) inapplicable (macrosculpture absent). Paraocular strigation treated as a separate character (ch. 18). Non-additive. Ambiguous at nodes 2 and 28. Possible optimizations are of reduced importance for the analysis because character is exceedingly homoplasious. Changes as suggested m the preferred cladogram: I <-0-^2. Optimization: ACCTRAN fer state 0. Assumes occurrence of state 0 in most ancestral nodes for Hemistephcams and Parastephanellus, remfercing its condition of groundplan state for the Stephanidae. Xustffîcatîon: this seems to be the case, because states 1 and 2 have highly scattered occurrences in the cladogram. 40 017. Fram^ ventral ha^, type o f macrosculptiare (not illustrated): (0) mnced: strigate, areolate-rugose, and irr%ular; (1) mostly areolate-rugose; (2) rugose or rugulose; (3) strigate or carinat^ regular (even) or weakty ondulate; (-) inapplicable (macrosculpture absent). Paraocular strigation treated as a separate character (ch. 18). Non-additive. Ambiguous at node II. Optimization is unimportant because correspondent clade is of reduced interest for the present analysis. Changes as suggested m the preferred cladogram: 0-*3->(l, 2). 018. Frons, paraocular strigation (not QlustratefQ: (0) absent; (1) present and distinct; (-) inapplicable (macrosculpture absent). Changes are unambiguous. 019. Frons, direction o f pilosityi (0) upwardly inclined hairs, abundant or sparse (Fig. 31); (1) downwardly inclined hairs (sparse) (not illustrated); (2) vùtually glabrous, hairs mmut^ about as long as height of ridges of sculpturmg, very scarce (size and scarcity make direction of inclination difBcult to define), or totally absent (not illustrated); (3) haks conspicuous, abundant, transversaHy inclined mwardly, so that hairs on each side offrons are mclmed towards the other half (not illustrated); (4) two supra antennal hair patches (one on each side), with ventrally mclmed hairs; remainmg glabrous or nearly so ^ ig . 32). Hairs along ^ e orbits and those at base of lateral and anterior tubercles were not considered, as th ^ have freed dfrection m stephanids. 41 Non-additive. Optimization: ACCTRAN. Makes state 2 the basal condition for Foenatopus^ lending extra support to this clade. Justffîcation: this is strongly suggested by the occurrence of state 2 in nearly all basal taxa o f Foenatopus, The only acception is a smgle reversal in the somewhat unusual species “Neostephanus" albomaculatus. Changes as suggested in the preferred cladogram after optimization: 0-+2. 020. Face, pilosity in between antennal foramina, until cfypeal suture: (0) absent, or as on frons (not illustrated); (1) abundant, hairs usually much shorter than on frons (Fig. 31). Changes are unambiguous. 021. Face, pilosity tn area between antennalforamen, eye margin, and base o f mandible (not illustrated): (0) present, although sometimes delicate, or closer to antennal fbrame than to mandible ^ many cases mostly between antennal ferame and ^ e margin); (1) absent, or, ^ present, occuring in a more or less linear distribution, as part o f the paraocular hafrs, never primarily in between the three reference points mentioned. Hairs in state 1 seem to be derived from the paraocular hairs, which possibly became larger or more numerous near and b^ond the level of the antennal forame; however, in some species such hairs are present while paraocular hairs dorsad of them are totally absent (e g., F . mgricaut&s, Parastephanellus sp. A). 42 Ambiguous at node 21. LisufScient evidence to support optimization. State I occursÎO T 931 ParastephaneUus spp. and i33LFoenatopus spp., but it cannot be assumed to have been present in the common ancestor of these taxa because o f Xonthostephanus n. gen., which is placed between them and shows state 0. 022. Coronal area and fions, presence o f granular sculpture (not illustrated): (0) absent, polished smooth m between macrosculpture; (1) present, powdered or granular aspect in between the macrosculpture, or entirely granular (macrosculpture absent); (2) vestigial. State I is somewhat different for “M adegqfoenus" occfputalis and “M adegqfoenus” bekifyanuSy where the granular microsculpture is considerably coarser and more shining than in other spp. showing this state. Additive. Changes as suggested in the preferred cladogram: 0-»l-»2, unambiguous. 023. Intreocellar carinae, position: (0) mostly posterior to, with a few in between, the posterior ocelli (Fig. 33); (I) mostly anterior to and m between posterior ocelli (Rgs 35, 36). Optimmation: ACCTRAN for state I. Makes origin of state I unique, and occurrence o f state 0 within foenatopinae as isolated reversals. Justffîcation: F. seyriff. is somewhat atypical m that it has a very small siz^ and this could have easily contributed to modify the shape of the present character (as often occurs with small specimens of stephanids). At the same thn^ if state I is assumed for this species (mstead of the 43 observed 0), the character becomes unambiguous with aO»I change at node 17, and a reversal to 0 îov Xcmthostephanus n. gen., that is the same result as obtained with the present optimization. This situation suggest that mdependent origins of state 0 for these two species is a more reasonable assumption than to consider state 0 as the basal condition for the foenatopinae. 024. Vertex sculpture, structurel (0) areolate-rugose latero-anterioriy, remaining transversely strigate, except for polished smooth temples CFig. 33); (1) sculpture partially transvase, except laterally, or at least antero-laterally, where any other orientation, except transverse or areolate-rugose, occur (not illustrated); (2) mostly or entirely transverse, even ^ondulate or if vertec polished smooth posteriorly (not illustrated); (3) mostly areolate-rugose or deep foveolate, which extends partially to temple; transverse sculpture only inside central impression (not illustrated); (4) mixed rugose and areolate-rugose, weakly transverse; interrupted by transversely carinate central impression (not illustrated); (-) inapplicable (several different autapomorphic types which do not fit any of the above descriptions) (not illustratetQ. Non-additive. Ambiguous at node 11. Optimmation is unimportant because correspondent clade is of reduced interest for the present analysis. Changes as suggested in the preferred cladogram: most changes occur. 025. Vertex sculpture, composMan ^erocellar carinae not considered): (0) two or more deferent patterns, occurring approximately in the same proportion (Rg. 33); (I) mostly or 44 entirely rugose lugulose, or areolate-rugose (not illustrated); (2) mostly or entirely parallel, ondulate-nigose or rugose in zig-zag (not illustrated); (3) mostly or entirely strigate or strigulate, transverse (Fig. 27). Non-additive. Ambiguous between nodes 29 and 36. Possible optimizations are of reduced importance for the analysis because character is exceedingly homoplasious. Changes as suggested in the preferred cladogram: most changes occur. 026. Vertex sculpture, intensity (not illustrated): (0) very coarse; (I) fine, or very fine. Changes are unambiguous. 027. Vertex, anterior hal^ microsculpture (not illustrated): (0) polished smooth in between macrosculpture; (1) granular microstculpture in between macrosculpture. Di small specimens, state 1 is usually present only (or better seen) laterally. Changes are unambiguous. 028. Vertex, emargimOion o f posterior margin (dorsal view): (0) deeply convex ^ig. 34); (1) weakly convex or straight (Rg. 33); (2) angled, as an inverted “V”, even ^base shortly transverse ^ g s 35,36); (-) inapplicable Qiead compressed, as m Fig. 29, or dorso- posterior part o f vert&c structurally unique in“Neostephanus" alluaudi Kiefifer). Adopted standard dorsal view position: start by looking at the occiput Q.e., looking at head posterioriy), then proceed tummg head towards vertex, until occiput disx^pears and only its dorsal countoum (= posterior margm o f vertex) can be seen. 45 Non-addîtive> Optûxiization: DELTRAN. Retains state 1 to ancestors at nodes 17 (ParasfepAane/Titf-^Foenatopinae) and 24 (Foenatopinae); reversals 1-^0 lend ectra support to Parastephcmellus zxAFoemtopus clades. Justification: the relatively atypical statusofXanthostephanus n. gen. in relation to other foenatopinae is due in part by the similarity o f several o f its character states with those ofM egischus, the presence o f a weakly convex to straight posterior margin o f vertex (state 1) being one of thena. Since this character seems to be in more direct association vnthM egischus, I tried to preserve h at the ancestral level (nodes 21 and 28) by selecting Deltran optimization. Acctran for state 2 produces a similar result, but it also implies state 2 as the ancestral condition at node 32. Nonetheless, state 2 is clearly the more specialized of the three observed, making this latter optimization more suspicious than Deltran. Changes as suggested in the preferred cladogram after optimization: 0-^1-» 0-»2. 029. Gemt and temple, sculpture (not illustrated): (0) entirely polished smooth; (1) with conspicuous sculpture (microreticulate, or rugulose, or foveolae), at least on basal half of gena {^Stephanus serrator, M anomalipes); (2) finely alutaceous. Additive. Ambiguous between nodes 30 and 38. Optimization is unimportant because correspondent clade is of reduced interest for the present analysis. Changes as suggested in the preferred cladogram: 0-*^l-^2. 46 030. Gem, shcpe: (0) centrally prominent, even if weakly so: From a small inconspicuous central swelling or callosity, to a strong and acuminate protrusion (Figs 34, 35,36); (I) normal, uniformly spherical or globose (Rg. 33). Prommence very weak or inconspicuousiaM . maculipennis and H. macrurus, and absent in some small specimens ofM anomalipes. Changes are unambiguous. 031. Gena, pilosity behind mcdar space: (0) glabrous, or with same pQosity as rest of gena (even &Tgena pilose only in its ventral halQ (not illustrated); (I) with a well defined patch of hairs, but hairs sparse and sometimes minute; remaining of gena glabrous (Fig. 37,27). Optimization: ACCTRAN. Makes change 0-+I a synapomorphy for Hemistepluams and for clade at node 41 (an apical clade o f Foenatopus). Justification: although dubious as a true synapomorphy fo r H em istephanus, it is certainly a well defined state for 8 o f 11 species which appear under a single clade (at node 41) within Foenatopus. Deltran optimization would imply several mdependent origins of state 1, which makes little sense when most such occurrences are within a moderately well defined clade. This apical clade is of little importance p e r se, but its evaluation allows the selection of an optûnûation which provides further support for the Hemistephanus clade. 032. Gena andfrons, presence o f yellow spots or stripes (not illustrated): (0) absent, no yellow colors on gena; (1) a yellow spot basally, at or near malar space (faint ^head has a light color, as red or brown); (2) difitise yellow basally, usually ectending near ^ e margin 47 from gena to free and frons, or gena mostly or entirely dffîise yellow; sometimes similar to a stripy but restricted to ventral halÇ (3) a well d ^ e d yellow or ivory stripe «(tending from base ofmandible to temple. Additive. Optimization: DELTRAN. Makes state 2 the ancestral condition at nodes 30 and 31. Justification: the preservation of state 2 in all ancestral nodes within frenatopinae (against implication o f a 2-»I-+2 shift in these nodes) seems a more reasonable option, since other states in this subfrunily, including state I, also appear sporadically in the cladogram, and Independent occurrences. Changes as suggested m the preferred cladogram: G-^I-^2-^3, with a MO reversal at node 14 (origin of the Hemistephams clade), and independent occurrences of states 0,1, and 3 in Foenatopus. 033. Occipital carina^ lateral shape: (0) more or less uniformly curvilineous (Figs 27,28); (I) bent midway; dorsal and ventral portions nearly straight (Fig. 30). Changes are unambiguous. 034. Occipital carina^ ventral d^erentiation: (0) ending very close to, at, or extending alongside hypostomal carina, reaching its apex ÇFîgs 38,39); (1) completely disappearing frr from hypostomal carina ^ g . 40), or fitding gradually and converging toward each other, or toward center of hypostomal carina, but not meetmg (Fig. 41). Ambiguous at node 32. Optimization is unimportant because correspondent clade is of reduced interest for the present analysis. 48 035. Occiput, maximum width: (0) much wider than apical width o f pronotum (Fig. 41); (1) slightly wider; pronotum + prostemum fitting tightly inside occiput (Fig. 40). Changes are unambiguous. 036. Postgenal b rit^, structure: (0) sinking ünmediatlely behmd hypostomal carina, even if doing so only centrally; postgenal lin^ consequently, angled in relation to sagital plane o f oral fossa ^ g . 38); (I) forming a flat plataform behmd hypostomal carina, which is parallel to sagital plane of oral fossa (even when sometimes each side of postgenal bridge slightly angled in relation to each other); then sinking towards forame magnum; postgenal Im^ consequently, initially straight, horizontal, then bending posteriorly towards fbramem (Figs 39,40); (2) postgenal bridge deepfy sunk, with gena folding partially over it, generating a deep shaft {some Hemistephanus spp.) (Fig. 41); (-) inapplicable (head strongly flattened; Fig. 39). Non-additive. Changes as suggested in the preferred cladogram; 0-*^I-^2, unambiguous. 037. MatiUarypcdpus, relative length: (0) from slightly shorter to slightly longer than head is laterally high (not illustrated); (1) distmctfy longer than head is laterally high; mdividual segments elongated ^ g s 29,50); (2) very small, several times shorter than head height and nearly hidden under the mandibles; at least apical segments not elongate CFig.42). 49 Non-addhîve. Ambiguous at node 5. Optimization is unimportant because correspondent clade is of reduced interest for the present analysis. 038. M axilkay palpus, pilosity o f 3rd segment: (0) hairs and/or prumosity abundant (Fig. 44,45,46, 47); (1) hairs long and stiflÇ sparse (Fig. 43). Changes are unambiguous. 039. M andible, e^erentiation o f dorso-mesal tooth: (0) absent; seen dorsally, mesal margin o f mandibles straight, touchmg folly when completely closed (Fig. 51); (1) present basally, small (Figs 48, 52) or stout, distinctly detached (Fig. 53), seen dorsally, mesal margm o f mandibles normally somewhat concave or in V, or U, etc, barelly touching when closed. Changes are unambiguous. 040. Labrum, color, (0) entire^ crean^-yeOow dorsally; brown or yellowish brown ventrally (Schlettereritts); (I) yellow, or sometimes ivory, or at least pale, ventrally only; dorsalfy distinctively darker (usually same color as firons/foce); (2) yellowish to whitish ventrally and dorso-apically, dorso-basally distinctively darker (usually same color as foce andfoons). The present character has been coded as inapplicable for M e^schus n. sp. in order to allow other states to be considered as additive without the necessity o f splitting them with m ked codmg. This would have to occur because the labrum color for the mentioned 50 species shows a distinct apomorphic state (entirely glossy pale-yellow, except at the very base dorsally) which, as well as state 2, derives from state 1 in a polarized transformation series 0.e., + Afeg/jcAor n. sp.]). Addhive. Opthnization: ACCTRAN. Places change 1-+2 at node 25 (origin of F o e m to p u sclade), considering state I of “Neostepfmms” albomaculatus as a reversal from state 2. Justification: even though a définitive 2-^1 reversal occurs at an apical clade (node 48), state 2 still seems to be a specialization typical o f Foenatopus, occurring only once among non-fr)enatopinae in the cladogram {fl. cylindricus). Occurrence of state 1 in the basal “Neostephanus" albomaculatus is not a strong indication of ancestrally, since this species is somewhat unusual within frienatopmae.Changes as suggested in the preferred cladogram after optimization: (Ml-*^2-»l. 041. Thorax (except legs), color. (0) mostly or entfrely dark brown or black; (1) lightly colored, mostly or entirely yellowish brown or reddish brown, or similar. Changes are unambiguous. State 1 occurs m isolated and mostly unrelated taxa only, suggesting the present character does not offer much higher-level phyletic information. Nonetheless, it helps uniting "Diastephanus” alutaceous and F o en a to p u s n a ta lic u s in a clade, which have an unique morpholo^ among stephanids, and might be more closely related than currently known. The present character was maintained in the analysis mainly because of its hnportance for these two species. 51 042. Pronotum, apical sensoricd hairs: (G) inconspicuous or absent ^ g , 54); (1) linearly organized, in a line or stripe along antero-lateral margin, even if only a few or very small (e g., F o en a to p u s sp. B) ^igs 63, 79); (2) organized in a tuft or small area (Figs 57,60, 63, 73, 66,76). Non-additive. Optnnization: DELTRAN. Restricts change 1^2 to the Parastephanellus clade. Justification; State 2 is a distinct specialization, while state 1 is the obvious groundplan state ft>r stephanids. Acctran optimization seems less likely because it makes state 2 the ancestral condition for Parastephanellus + Foenatopinae only because of the somewhat ap ical Xanthostephanus n. gen., assuming then a very early reversal which is constant for nearly all Foenatopus. It also disturbs the more natural assumption, suggested by character distribution in the cladogram, that state I persisted in all mam ancestral nodes, assuming instead that it had two origins (0-+1 and 2 -^ I). Changes as suggested in the preferred cladogram after optimization: 0-^1-»2. 043. Pronotum, apical margin: (0) a simple rim, slightly to moderately reflexed upwards (Figs 54, 55,56,65,68, 77,57, 58,60,61,63, 64,66,67,79); (1) strongly reflexed upwards, convoluted, generating a wide collar (Figs 59,62), which is alutaceous dorsally; (2) the entire antero-lateral part of the pronotum vertically flat and broad laterally, changing to a narrow central margin reflexed nearly 90“ upwards ÇFigs 73,72,78,69,70, 71,75,76,74,77). Non-addhîve. Changes as suggested in the preferred cladogram: I«-0->2, unambiguous. 52 044. & 045. Pronotum, apical emargination: (0) weakly angled ^ g . 54 ); (1) distinctly and strongfy emarginat^ C-shapped ^ g s 57, 59, 60,62, 63,65); (2) as deep as, or deeper than wide; more or less U-shaped, but margins of emargination diverging towards apex ^ g s 66); (3) c-shaped, but small, restricted to about 0.3-0.4 of the apical width, occupymg central part only (Fig. 56); (4) deep, U- or V-shaped (Rgs 69,71,76,73,79, 80); (-) inapplicable pronotum atrophied anteriorly. Fig. 81). The species ”Neostephanus" albomacukOus has state 4, but modified, and somewhat similar to Parastephanellus. The species “Diastephanus” sp. B shows an intermediate state between F o en a to p u s and that of the aberrant F o en a to p u s n. sp.. The original character was split in two in the matrbc, as follows. Character 44; transformation series 0->l->2->4 renumbered as 0-^I-^2-+3; original state 3 coded as inapplicable. Character 45: states 0,1,2,4 renumbered as 0; original state 3 as 1. Ch. 44: Additive. Optimization: ACCTRAN. Places change l-»^2 at node 21, providing extra support for the Parastepfumellus + Foenatopinae clade. Justification: deferences between states 1 and 2 are relatively mtens^ so that it seems easier to accept a smgle changel-*'2, with one isolated reversal of 2->l (^orXanthostephanus n. gen.), than to assume two mdependent origins of state 2 fi’om state 1, which is suggested by optional optimizations. Changes as suggested in the preferred cladogram after optimization: 0 ^ l “^2“^4 Çh the cladogram: 0-^l-^2“^3). Ch. 45: Additive. Changes as suggested in the preferred cladogram: l-*^3 (cladogram: 0-+I), unambiguous. 53 046. Colo, mdmducd lateral hairs: (0) absent ^ g s 54,57, 60,66); (1) present (Fig. 63). These are isolated, usualfy relatively long and strong hairs present on or in between the transverse carinae on colo; only one hair per carina on each side, often originating directly from posterior side of carina, or basally; more commonly associated with first three complete carinae. Changes as suggested in the preferred cladogram; 0-^l-+0, unambiguous. 047. C o lo ,sAqpe: (0) transverse, very narrow (about three times wider than long) (Figs 54,55); (1) normal, about 0.5-1.0 as long as wide (Figs 57, 60, 62,63, 73, 66, 65,76, 79); (2) elongate or very elongate, at least 1.5 as long as wide, but usually 2.0 or more (Rgs 59,69, 71,80); (-) inapplicable (colo fused to preannular). Length is measured from apex of pronotum to pronotal fold; width corresponds to average width. When the colo is fused to the preannular the section of the pronotum in front of the semiannular looks more elongate than normal. However, this situation cannot be assumed as being equivalent to state 2 because there are cases of very long pronotum with a short colo, as for example mFoenatopus sp. E. Additive. Changes as suggested in the preferred cladogram: 0-+l-^2, unambiguous. 54 048. Cobi, presence qfcarim e: (0) present distinct even Wien weak ^ig s 54-58,60-72, 69-82); (1) inconspicuous or absent ^ g . 59). Changes are unambiguous. 049. & 050. Colo, carinae structure', (0) transverse and strong often raised as a blade-like structure complete or mcomplete; in lateral view transverse or weakly oblique towards apex ^igs 54, 55,57,58,60,61,63, 64); (1) carinae mostly complete, organized in a Ü- pattem, so that th ^ are oblique towards pronotal fold when seen in lateral view (Figs 66, 67); (2) in dorsal view straight and oblique on each side, generating a V-pattem; carinae incomplete, not or barely meeting dorso-centrally (not illustrated); (3) transverse, weak, or at least neither too strong nor blade-like; often not meeting dorso-centrally on colo J.e., mcomplete); in lateral view transverse or weakly oblique towards apex (Kgs 69-80); (4) strong oblique longhudmal, straight carina^ converging to inside pronotal fold (Fig. 82); (5) one strong centro-longitudinal carina on colo, and none or only a few oblique laterally 0 g . 56); (-) inapplicable (carinae absent). A branched transformation series was suggested when this character was run as unpolarized, as follows: (4,5)MM l-»2-^3. For this reason, the character was split m two to allow a polarization hypothesis to be applied for each subset of transformations. Subset 4«-0-+5 (ch. 49) was renumbered as I<-0-^2, and ran non-additive because states 4 and 5 are independent derivations; states 1-3 were considered as inapplicable. Subset 0-^l-+2-*^3 (ch. 50) was ran as additive with states 4 and 5 considered mapplicable. 55 Ch. 49: Non-additive. Ambiguous at node 4. Optimization is unimportant because correspondent clade is of reduced interest for the present analysis. Changes as suggested in the preferred cladogram: 4MMS (cladogram: I<-0-+2). Ch. SO: Additive. Optimization: ACCTRAN. Assumes change l-*^2 at the F oenatopus clade. Justification: confi)rms with additive hypothesis; other optimizations assume a l-*^3 change at node 33. Changes as suggested in the preferred cladogram after optimkation: 0-^l-+2-»3. 051. Colo, carinae shape: (0) linear, as a carina or strigation, etc (Figs 54, 55, 57, 58, 66, 67, etc); (I) leaf-like, wide and flat carinae present (Figs 63,64,73, 72). Ambiguous at node 15. InsufiScient evidence to support optimization. 052. Colo dorsalfy, differentiation o f centro-lanfftudinal depression: (0) absent, or present inside pronotal fold only ^ g s 54,57,60,66,62,79, 82); (I) present, although sometimes very weak (Figs 59,63,73,69,71,65,76). Ambiguous between nodes 28 and 36. hisufScient evidence to support optimmation. 053. Colo dorsalfy, concavify o f centro-longitudinal depression: (0) distinctly, even ft' only slightly, concave ^ g s 59,63,73,69,71,65,76); (I) deeply concave inside pronotal fold only (not illustrated); (-) mapplicable (centro-longitudinal dqiression absent). Changes are unambiguous. 56 054. Pronotal fold, d^erentiatiom (0) distinct, from sunply raised at about 90° m relation to surfrice of colo, to moderately or deeply excavated (Rgs 58-59,62, 63-72,68-74, 76, 82); (I) indistmc^ not a cuticular fr)ld but simply a “slope” or weak inclination between colo and preannular, or absent (Rgs 60,61,69-71,75,77-81); (-) inapplicable (Rgs 54, 55). Ambiguous between nodes 30 and 47. Although possible optimizations are of reduced hnportance for the analysis, it might be of interest to stress that the absence of a pronotal fold is a definitive tendency io a o n g F o en a to p u s spp., where it is distinctly more fr'equent than in any other stephanid taxa. 055, Pronotal fold, shape'. (0) distinclty and often deeply concave anteriorly, never collapsed centrally (Rgs 58,63,64,73,72,65,68,82); (1) centrally collapsed, or at least closed, leaving two shallow or moderately deep openings on each side QFig. 66); (2) weak and transverse, indicated by posterior end of centro-longitudinal depression of colo ^ig s 69,71, 75); (-) inapplicable pronotal fold absent). Yot Xanthostephanus n. gen. pronotal fold is collapsed centrally, similar to (1), but not ft)rming the characteristic lateral openings ofthis state. For “M akgqfoenus” seyrigi and Foenatopus elliotti (Cd)allos) the present character is very unusual in that th ^ are the only foenatopinae with a deeply concave pronotal fold. Non-additive. Optimization: ACCTRAN for state 2. Makes state 2 the ancestral state at node 30. Justffîcatîon: although the mapplicable occurrences are not considered a true state for current character, the foot that inapplicable here is equivalent to absence o f 57 the character makes h more likely to have progressed from state 2, where the character is weak or Glint, than from states 0 or 1, where it is conspicuous and well developed. Note also that both state 2 and "inapplicable" are part of state 1 of ch. 54. Changes as suggested in the preferred cladogram after optimization: I<-0-^2. 056. Preanmlar, presence o ftransverse ruga aandgroove-, (0) absent ^ g s 57, 59,60, 63, 73,66,69,71,65,76, 79, 80); (1) present ^ g . 62); (-) inapplicable (preannular not ^eremiated; Figs 54, 55). Ambiguous at node 4. Optimization is unimportant because correspondent clade is of reduced intérêt for the present analysis. 057. Preannular and semiannular, degree o f isolation: (0) preannular absent ^ g s 54, 55); (1) perfectly continuous with semiannular (Figs 58,61,67,72,68); (2) in lateral view, isolated from semiannular by a step or angle, or simply by a more swollen and dif&rently sculptured semiannular (Figs 64, 70,74, 75, 77,78). Character state for “Diastephanus" sp. B, “Diastephanus” sp. C, and “Diastephanus” sp. D similar to state 1, but this is due to their strongly flattened body. Additive. Ambiguous between nodes 11 and 21. InsufScient evidence to support optimization. Changes as suggested m the preferred cladogram: mostly 2-^ 1, but all changes are possible. 58 058. Preannular, inclination o fhairs (not illustrated): (0) inclined backwards; (I) inclined forwards, Note-that prumosity was considered as a distinct character (see ch. 63). Changes are unambiguous. 059. Semiannular, inclination o fhairs (not illustrated): (0) inclined backwards; (1) indined sideways, or arranged more or less radially; (2) mcOned forwards. Note that pruinosi^ was treated as a distinct character (ch. 63). Additive. Changes as suggested m the preferred cladogram: 0-^I->^2, unambiguous. 060. Pre- and Semiannular, d^erentiation o fa dorsal impression centrally. (0) absent ^ g s 54, 57,59,60,62,63, 66,69, 71,65, 82); (1) present as a fovea or a shallow and narrow longitudmal depression or series of small depressions ^igs 73,79,80,81). Observations in Stephanus serrator, M. califomicus, and P. grarmlatus may be difficult, and should be performed with extra attention. Dorsal impression absent on smallest observed specrnien ofF. annulipes. Changes are unambiguous. 59 061. Pre~arul Semiannular, shcpe o f dorsal mpression: (0) longitudinal, extending from semiannular to preannular, narrow and linear (not illustrated) or broad (Fig. 73); (I) circular, fbveola-like, placed exactly where preannular ends and semiannular starts ^igs 79,80,81); (-) mapplicable (absent). See notes for previous character. Ambiguous between nodes 13 and 31. Possible opthnizations are of reduced significance for the analysis. 062. Pre- and Semiannular, microreticulation: (0) absent, cuticle polished (not illustrated); (I) present, microareolate or with strong alutaceous, cuticle mat (Fig. 61). Ambiguous at node 47. Optimization is unimportant because correspondent clade is of reduced interest for the present analysis. 063. Semiannular, presence o f fine dense short pilosity (prumosity) lateralfy: (0) absent (not fliustrated); (1) present basally on femoral suclus only; hairs longer than in nect state (not Olustrated); (2) dense fine pOoshy (almost pruinose) present all over (not illustrated). Non-addhrve. Ambiguous at node 10. Optimization is unimportant because correspondent clade is of reduced interest for the present analysis. 064. Femortd sulcus (promtum), e^erendation: (0) present as a distinct oblique- longitudinal hnpression, evmi ^ weak; ventral area of pronotum distmctly delimited, with different sculpture from the usually polished smooth surfiice o f the fomoral impression 60 (Rgs 58, 64); (1) inconspicuous or not differentiated; no distinct physical limit between ventral area of pronotum and remaining of pronotum, although ventral are sometimes with distinct sculpture (not HlustratefQ. The femoral sulcus mP. cknotatusvi very weak, but is definitely diflferentiated. Optimization: ACCTRAN. Places change 0-»^l at node 17 (origin of AnwApAone/W+ Foenatopinae clade), providing ectra support to it. Justification: occurrence of state I is widespread in species o fParastephanellus and feenatopinae, while absent in the sister-group clade {Hemistephanus), as well as rarely occurring in other stephanids. These fects seem to suggest that state 1 was already present in the commom ancestor o fParastephanettus + Foenatopinae, mstead of evolving twice (once for Parastephanettus and once fer F oem topus) as suggested under Deltran. 065. Ventral Area ^onoium ), pilosity (not illustrated): (0) with fine, uniform dense p 3 o sî^ (1) fine dense pilosity basally, becoming more sparse apicaHy until glabrous fi’om midway on; somethnes finely e densety pilose only on extreme base; (2) entirely glabrous, or almost so. Non-additive. Changes as suggested in the preferred cladogram: 0-^2-» 1, unambiguous. 066. Prepectus, degyee o fexposition'. (0) mostly exposed (Fig. 85); (1) mostly or completely covered by postero-ventral apex of pronotum (not illustrated). 61 Optnnization: DELTRAN. Suggests state 1 for nodes 17 and 21, and change MO at node 28, providing further support to the foenatopinae clade. Justification: Although state 0 seems to represent the basal condition, it is typical of the foenatopinae, for which only a few species show state 1. At the same time, state 1 is the groundplan state for non- foenatopinae stephanids. Ambiguity occurs only in function of a few isolated occurrences of state 0 in Hemistephanus and Parastephanellus. These focts strongly suggest that the reversal MO occurred in a more definitive form for the foenatopinae, and that the occurrences ï o t Hemistephanus exA Parastephanellus represent independent reversals. 067. Pronotal spiracle, exposition'. (0) concealed by postero-ventral angle of pronotum (absent?) (not illustrated); (1) distinctly exposed ^ g . 85). The Australian Parastephanellus denotatus has a distinct spiracle and prepectus, but these are not visible m intact specimens (observed m a broken specimen). Optnnization: DELTRAN. Places change 0-+1 at node 28, providing further support to the foenatopinae clade. Justification: essentially the same as discussed in ch. 66, except that in the present case state 1 is the one characteristic of foenatopinae. 068. Semiannular, shape o f posterior margin dorsally. (0) slightly transverse centrally (Rgs 92,93); (1) unfibrmly round or arched ^ g . 96); (2) acummate centrally, or hyperbolic ^ g . 91); (-) inapplicable (species with l^erbolic mesoscutum). 62 The s iu ^ of the mesoscutum and that of the posterior margm of the semiannular are relatively mdependent, except apparently in Hemistephanus^ for which, therefor^ the present character was considered as mapplicable. Non-additive. Changes as suggested in the preforred cladogram: 0-^l->2, unambiguous. 069. Prostemum, angle between epical margins ofhemistenritesi (0) nearly 90® or more ÇFîg. 83); (I) distinctly less than 90* (Mg. 84). Changes are unambiguous. 070. Front coxa, lateral sculpture (extemalfy) : (0) several small foveolae present (Fig. 85); (1) foveolae absent (not illustrated). The presence of foveolae is apparently related to large size, but Xanthostepharms n. gen., F. aurantiiceps, and F. annulipes do not have it and represent large species. Changes are unambiguous. 071. Spur ofantenna cleaner, shcpe o fvelum: (0) width of ventral membrane more or less constant, or increasing slightly apically ^ ig . 86); (I) ventral membrane narrowing apically, disappearing before reaching apex o f spur (Mgs 87,89); (2) ventral membrane distmctly wider apically than centrally, so that it is somewhat lobed; endmg abruptly, somethnes with a small emargination apically ^ g s 88,90). 63 For state 2, the margin of the ventral membrane at the point o f the emargination is sometimes partially sclerotized, giving it approximately the same color and texture of the free apex of the spur; in lower magnifications this structure may appear as a spur with a “bifiircated” apex, which is a good indirect indication of character state 2, Additive. Changes as suggested in the preferred cladogram: 0-»'l->-2, unambiguous (except fer the 2 species clade at node 51). 012, Mesoscutum, overall shape: (0) large and round, distinctly larger than combined area of axillae (Figs 92, 95), usually I.5-2.0x as wide as long; (I) slightly transverse; nearly as large as, or slightly larger than combined area o f axillae (Fig. 96), usually 2.0-3 .Ox as wide as long; (2) conspicuously transverse more than 3.Ox as wide as long, and distinctly smaller than combined area of axillae (Figs 94,97,99); (3) hyperbolic, large ^ig. 98). Only area with macrosculpture considered (i.e., anterior “exposable” area excluded). Non-additive. Optimization: DELTRAN. Explains origm o f character state for Xanthostephanus n. gen. with a l-»0 reversal, and îo t F oem topus with a l-*^2 progression (both at node 28). Justification: other optimizations are more assuming m that they result either in a 0->-2 or a 2-»0 transformation. Since state 2 is highly specialized and intensely deferent form state 0, those last two possibilities seem much more difficult to occur in a single step than the gradual transfermations suggested by Deltran. Mam changes as suggested in the preferred cladogram: 0-*^l-»2, and 3 origmating fi^om 0 and I. 64 073. Mesoscutum, direction o fpilosity: (0) all haïra mcimed or decumbent toward transscutal articulation, or mostly glabrous (not illustrated); (1) hairs neartransscutal articulation inclmed or decumbent anterioriy, or at least antero-laterally (Fig. 98). Ambiguous between nodes 3 and 21. DisufBcient evident to support optimization. 074- Mesoscutum, size offunrsr. (0) very small or minute (not illustrated); (1) very lon& strong ^ g . 98). Ambiguous between nodes 3 and 21. DisufScient evidence to support optimization. OnS. Median line (mesoscutum), differentiation: (0) differentiated, even ifâint (Fig. 96); (1) absent on m m area (distinguishable only on anterior “exposable” area) (Figs 97,99). Changes are unambiguous. 076. Notaulus, differentiation: (0) dff&rentiated, even if feint ^ig. 96); (I) not differentiated (Figs 97,99). This character is difficul to interpret in foenatopmae. Character coding here is therefore somewhat imprecise. Codmg is particularly doubttiil for "Diasteplumus" sp. B, due to its small sfee. Changes are unambiguous. 65 077. Costal cell on front wing, presence o f short setae: (0) with small and sparse but conspicuous setae, usually more Sequent centrally or apically ^igs 163-167); (1) setae completely absent ^ ig s 171-178), or few and microscopic, inconspicuous CFig. 170). Changes are unambiguous. 078. VeinM+Cu, setae: (0) reaching base o f IM and usually advancing over ICu in at least one of the wings ^ g s 155,158,163-165,167); (1) endmg near or a t, but not reaching, base of Ih^ from which the most apical seta is separated by a distance always shorter than length o f Icu -a ^ tg s 157, 159, 166, 170, 172, 174, 175-177); (2) placed far from IM, from which th^r are separated by a distance greater than length of Icu-a (Figs 156, 168, 169, 173). Additive. Opthnization: ACCTRAN. Places change 0-^1 at node 13, lending extra support to clade Hemistephanus+Parastephanellus + Foenatopinae; prevents occurrence of change 0-^2. Justffîcation: the considerable distance of the M+Cu setae from base of vem IM in all Hemistephanus spp. (clade at node 14; see also Aguiar 1998) makes it dffîcult to accept that the character state m the ancestors of this clade was unmodffîed, as assumed under Deltran. Acctran also prevents the assumption of two d^erent origins for state 2, that is, a 0-»2 change for Hemistephanus and a l-*2 change for Xcaithostephanus n. gen., which would occur with Deltran. Changes as suggested in the preferred cladogram after optimization: 0-»-l->2. 66 079 Vein I-^IA, setae (not illustrated); (0) distmctly longer, or much longer, than setae on hffCu; (1) approximate^ of the same size, even if thicker than M+Cu setae. There are usually 5-6 setae on basal half of vem 1-lA. These are usually translucent, stiff and straigh, always 90** erect Changes are unambiguous. 080. ParasSignuM, d ^eren tia tio n i (0) distinctly longer or as long as IRs, and distmctly longer than wide ÇRgs 163-166); (I) shorter than IRs, but still longer than wide (Rgs 168, 169,171); (2) much shorter than IRs, or absent; when developed usually as long as wide, or wider than longer (Figs 170, 172-178). For Schlettererius cmctipes, Stephanus senator, and M. brurmeus the parastigma is “fractured” at point of connection with pterostigma, so that the two are not contiguous. For “Stephanus” tibiator the parastigma shows an intermediate shape between states 1 and 2; because of this, it was coded as multistated (12), rather than interpreted as a distinct state. Non-additive. Optimization: ACCTRAN. Places change 0-+2 at node 24, lending extra support to the foenatopinae clade. Justffîcation: state 2 is a distinct specialization for the foenatopmae, with only two independent occurrences for other stephanids, placed fiir appart from each other and from the foenatopmae. The character is ambiguous only because o f the occurrence of state 0 in K s e y r iff, but the reduced size of this species 67 suggest that most of its states which seem **unusuar in relation to other foenatopinae should be considered with caution. Changes as suggested in the preferred cladogram after optimizationr 081. Pterostigma, shape-. (0) short and wide, even if sometimes more or less lanceolate; or at least much wider near or at 2r ^ g s 163-166,171, 172, 178); (1) long and narrow; width more or less un^rm , or unifbrmly decreasing towards apex (Figs 167,168-170, 173-177). The shape of the pterostigma in Stephanidae is highly variable, and may perhaps be interpreted as representing more than two states. Nonetheless, several different interpretations have been tried for this woric, but the high variabOi^ makes it nearly unpossible to clearly define more than two states. Character state somewhat intermediate for P. granulatus. Character state for F oenatopus n. sp. was interpreted fi’om examination o f a male specimen. Ambiguous between nodes 3 and 29. Insufficient evidence to support optimization. 082. Pterostigma, sclerotimzation: (0) uniformly opaque, brown or black, usually heavily sclerotized, except by pale marie basally OFigs 163-173); (1) lightly sclerotized, with a distmct centro-longitudinal area, or most of pterostigma, except margins, clear translucent (Rgs 175-177). Some light-brown pterostigmae may look like state 1, but if the margms are not distmctly more mtensely sclerotôed (or more dark colored) than the center, or if the 68 texture is homogeneous all over, then it must be considered as state 0. For example, “Neostephanus" sp. A. The pterostigma m stephanids is never entn-ely translucent Ambiguous at node 32. Optimization is unhnportant because correspondent clade is o f reduced mterest for the present analysis. 083. Veins IR s and IM, contiguity: (0) IM normally at nearly 90** with IRs, making cell R truncate apically ^ g s 163-166,168,169,171), more rarely forming an obtuse angle (Fig. 167); (1) IRs aligned with IM, or nearly so, making cell R apically acute, triangular (Figs 170,172-178). Changes are unambiguous. 084. Veins IR s and IM, relative size: (0) IM from distinctly longer to much longer than IRs (observed: 1.2-6.5x) (Figs 163-171); (I) IM nearly as long as, or distinctly shorter thanlRs (observed: 0.3-1. Ix) (Rgs 172-178). If veins perfectly contiguous, search for spectral indication of (Rs+M)a, which marks the limit between IRs and IM This character was originally coded subjectively Q.e., not measured), reevaluated several times, and then finally measured and interpreted as above. Taxa coded with a **?” indicate that vein IM is perfectly fused and aligned with vem IRs, and that no spectral mdication o f ^s^M )a was found, so that the limit between these vems cannot be distmguished. Changes are unambiguous. 69 085. Vein IM. origini (0) origmatmg at or basad Icu-a, so that nodus formed by fusion of M-fCu/lM with Icu-a/lCu is usually oblique toward wing base (Figs 155,163-170, 178); (1) origmating ^ical to Icu-a, so that mentioned nodus is forwardly oblique (Figs 156- 158,159, 171-177). Optimization: ACCTRAN. Makes state 1 a synapomorphy for the Pcarastephanellus + Foenatopinae clade; assumes all occurrences of state 0 within this clade as isolated reversals. Justffîcation: character distribution in the cladogram strongly suggests that state 1 is a typical specialization of the mentioned clad^ with only one isolated occurrence for other stephanids. 086. Vein IM, shape: (0) distinctly and conspicuously arched (Figs 163, 167); (I) slightly arched (Figs 164,165,169-176); (2) straight, even if slightly bent basally (Figs 166, 168, 177,178). Non-additive. Changes as suggested m the preferred cladogram: multiple mdependent occurrences of 1-+2 and 2-*>l, unambiguous. Change 0-*^l seems distinct, but 1-^2 occurs almost with the same fiequen polarized. 087. Vein Icu-a, shepe: (0) straight or nearly so (Rgs 163-173, 178); (1) slightly convex toward apex o f wing; sometimes slightly clavate (Figs 156-158,174); (2) apex distmctly bent toward base of wmg (Kgs 159,175,176,177); (3) slightly convex toward base of wing (Fig. 155). 70 Non-additive. Ambiguous at node 32. Optimization is unimportant because correspondent clade is of reduced interest for the present analysis. Changes as suggested in the preferred cladogram: 3*-0-»l-»2. 088. Vein Icu-a, 'mdûr. (0) uniferm^ or gaining little width progressively from base to apex (Kgs 156,158,159, 191); (1) narrowed basally, thickened apically, or suddently thick only apically; so that it is slightly clavate, or clubbed (Figs 155, 157,178). Ambiguous at node 39. Opthnization is unimportant because correspondent clade is of reduced mterest for the present analysis. 089. Vein Icu-a, mclination: (0) straight, at nearly 90® with both M+Cu and ICu (Figs 155, 163-167, 170); (1) inclined towards wing base, forming an angle of little less than 90® with M+Cu, and usually a correspondent slightly obtuse angle with ICu (Figs 156-158, 168,169, 171,175,177, 178); (2) strongly inclmed towards wing base, angled at much less than 90® (to a minimum of 45®) with M+Cu (Kgs 159,168, 172, 173,174, 176). The best way to evaluate this character is to «camine the wing in an upside-down position (dorsal sid% right wing), which allows for more accurate observations on the slant o f vein Icu-a. Additive. Optimization: DELTRAN. Restricts change 0-»l to node 17, mamtaining state 0 as the ancestral condition for taxa basal to it, thus assuming isolated origins for state 1 at this level. Justification: character distribution in the cladogram strongly suggests states 1 and 2 are a distinct and strong tendenty for more specialized 71 stephanids (Hemistephanus, Patastephanelbis^ and feenatopinae). At the same time, states 0 and 1 are weakly different, so that isolated origins of state I from 0 is not a difhcult possibility. Changes as suggested in the preferred cladogram after optimization: (M M 2. 090, Vein ICu^ differentiationi (0) entirely tubular ^ g s 163-172, 174); (1) tubular on basal 20% or more, but distinctly less than 100%, with apex nebulous or spectral (Figs 173, 175); (2) spectral or nebulous, even if with a short tubular stub basally (Figs 176- 178). F o rK form osanus the tubular portion of ICu varies from 30-80% of the total length o f this vein in different specimens. Additive. Optimization: ACCTRAN. Places change 0-*^l at node 25 (origin of Foenatopus clade). Justification: conforms with additive hypothesis. Character distribution in the cladogram clearly indicates that loss of sclerotinization of vein ICu is a generalized tendenqr aiFoenaiopus. At the same time, state 2 is a distinct specialization of state 1. Acctran supports this idea, but alternative optimizations place state 0 at the base of F oenatopus clade, thus assuming an independent 0-»2 change at the same level (for ‘‘Mackgafoenus’’seyrigt). Changes as suggested in the preferred cladogram after optimization: (M1-+2. 091. Vein 2-IA, connection witit 1-IAi (0) distinctly connected, even if as a very short stub (Figs 155-157); or, if 2-1A entirely spectral, apoc of 1-1A still showing as a 72 bifurcation (Rg. 158); (I) completely disconnected, vein 2-1A spectral, apex of 1-1A as an mdependent vein, distinctly curving anteriorly towards aper o f crossvein Icu-a OFig. 159). Changes are unambiguous. 092. Vein 2-lA, shape: (0) straight longitudinal (Figs 155, 163-167); (1) curving but not strongly, toward posterior margin o f win& reaching margin opposite to or a little before apex o f 2Cua (Figs 175, 177); (2) strongly curved, reaching wing margin before middle o f ceU2Cu^gs 173.170). Note that character states apply even if vein is entirely spectral. Coding for F oenatopus sp. E and "Diastephanus” sp. B is doubtfiil (specimens in bad condition). Additive. Optimization: ACCTRAN. Places change 0-*^l at origin of the foenatopmae clade (node 24), assuming state 0 mF. seyrigi asa l-»0 reversal. JusthScation: conforms with additive hypothesis. Character distribution in the cladogram clearly mdicates that a curved 2-1A is a generalized tendency in foenatopinae. At the same tim^ change 1-+2 is not only an mtuitive progression but in Act occurs, unambiguously, for the clade at node 49. These Acts suggests that state 2 îov Xanthostepfuams n. gen. probably also originated from a 1-^2 change, a possibQity^ supported only by Acctran. Changes as suggested m the preferred cladogram after optimization: 0-*l-*2. 093. Vein 2-IA, différentiation: (0) entire^ nebulous; when mtensely so, appearing xa be tubular (Kg. 163); (1) tubular on 90% or more ^ g s 155,167); (2) shortly tubular basally, 73 remammg mostly or entirely nebulous ^ g s 164-166,170,191); (3) a small tubular stub, which, sometimes is briefly nebulous apically, remaining absent or spectral, or 2-1A entirely spectral ^igs 168,169, 171, 172, 174-178). Additive. Main changes as suggested in the preferred cladogram: 0-*^l-*^2-*^3, unambiguous. 094. & 095. Vein 2Cna, (^erentiation: (0) entirely tubular (Figs 163, 167, 168, 169, 170); (1) entirely and distinctly nebulous (Figs 165, 166,172); (2) partially tubular (basal 50% or less); apically nebulous, even ^ strongly and appearing to be tubular (Hgs 164, 171, 174); (3) feintly nebulous or spectral ^ g s 173, 175-178). If considered as a single character, and run as non-additive, the tran^ormation series l*-0-^2-»^3 is strongly supported imder Acctran optimization in resulting cladograms. Deltran results in state 3 deriving twice from state 2, which is a more presumptuous assumption in the present case than a single origin plus a 3-^2 reversal in Acctran. Based on this result, the character was split in two, and run as additive, as fellows: (ch. 94) states 0 and 1 renumbered as 0; states 2 and 3 renumbered as 1 and 2; and (ch. 95) states 0,2,3 renumbered as 0; state 1 remaining the same. Ch. 94: Addhhre. Optimization: ACCTRAN^. Anticipates change 2-*^3 (cladogram: 1-^2) to node 25 (foenatopus clade). Justification: conforms with additive hypothesis. Ambiguity at node 25 results from occurrence of state 1 (cladogram: state 0) for “Nèostephanus" alèomacufatus; but this state has been hypothesized as a separate line of derivation (ch. 95), representing an mdependent origin fer this species. If the state fer this 74 taxon is disconsidered, change 1-^3 (cladogram: 1-^2) at node 25 becomes unambiguous, and this is an indication that Acctran is the most adequate optimization. Changes as suggested in the preferred cladogram after optimization: 0-^2-+3 (cladogram: 0-^l-^2). Ch. 95: Ambiguous between nodes 11 and 15. Optimization is unimportant because correspondent clade is o f reduced mterest for the present analysis. 096. Vein 2CvOy relative size: (0) longer than portion of vein Icu-a above CuP, but less than 1.5x as long ^ g s 163, 164, 166,171); (1) more than 1.5x as long as portion of vein Icu-a above CuP (Figs 167,170); (2) as long as (rare), or shorter than length of portion of vein Icu-a above CuP O^igs 165,168, 169, 172-178). Non-additive. Optimization: ACCTRAN. Makes state 0 the ancestral condition for ÛieParastephanellus clade (node 18). Justification: since state 0 is the groundplan condition, for Parasi^fianellus, Deltran optimization would imply multiple independent origins (reversals) for this state, which seems less likely than the occurrence of independent origms for the much less fi’equent state 2 miimParastephanellus. Changes as suggested in the preferred cladogram after optimization: l*-0-»^2. 097. VeinlCua, anglewith ICu: (0) nearly 90" (Figs 163, 165, 166, 169, 173, 174, 177); (1) distinctly acute ^ g s 167,168,170); (2) distmctly obtuse ^igs 164,172,1827,175, 176,178). 75 Ifnin as non*additiv^ the transfonnatioa lH)->2 is strongly supported in resulting cladograms. To conform with this hypothesis 0.e., two steps for transformations between derived states), character states were renumbered in the matrix as and run as additive. Additive. Optimization: ACCTRAN for state 2. Assumes independent origins for state 1 (cladogram: state 0) in the “Stephaams”pachylomerus clade; anticipates change (M2 (cladogram: M 2) to ihtFoenatc^nis dad^ lending extra support to it; delays reversal 2-^0 (cladogram: 2-+1) for “Neostephanus” albomaculatus. Justifications: 1. The scattered distribution of state I (cladogram: state 0) suggests that it is indeed prone to arise independently among stephanids 0t probably represents a highly specialized trait, important only at the species level). 2. State 2 is a typical specialization oîFoenatopus spp., occurring throughout the genus, while only four mdependent occurrences of this state occur for other stephanids in the cladogram. These foots suggest that state 2 represents a characteristic evolutionary change îor Foenatopus, and its occurrence was therefore prioritized for this clade. Changes as suggested in the preferred cladogram: I-HM 2 (cladogram: 0*-M2). 098. Vein 2Cub, eb^erentiation: (0) distmct; strongly nebulous, often tubular basally ^ g s 163,164,170); (I) nebulous (Figs 165, 167); (2) partially and weakly nebulous, rest spectral, or entirely spectral (Kgs 166,169,171-178). 76 Additive. Ambiguous between nodes 3 and 29. Possible optimizations are of re&tced significance for the analysis. Changes as suggested m the preferred cladogram: 099. Vein 2Cub, forking oftqtex: (0) forked, even if visible only in a few specimens of each species ^ g s 163,164); (1) simple, always visibly even in specimens with this vein spectral (Figs 165-178). Changes are unambiguous. 100. Vein 2cu-a, differentiation: (0) semi-tubular or nebulous (Figs 163, 164, 166, 167); (1) spectral or absent ^ g s 165, 168-178). State 1 originally interpreted as two separate states (2cu-a “spectral” and “absent”), but these are difficult to tell appart, and always appear together and randomly dispersed within clades apical to state 0, thus suggesting th^r represent a same condition. Optimization: DELTRAN. Restricts 0-+1 change to node 17, providing additional support fer the correspondait clade. Justificative: State 1 is uniformly present withm the mentioned clade (except only for “Neostephanus" allwxutS)^ suggestmg it is a typical specialization o f this group. Ambiguity occurs only in function of state 1 va M e^sch u s anomcdipes^ but ocamined specmiens o fthis species had a reduced body size, which could be related to the feet that vein 2cu-a was not observed in this species. These fects suggest 77 that state 1 is better mterpreted as a charactenstic evoliitionaiy change tor Hemistepfumus ■^PcarasfephaneUus + Foenatopmae and its occurrence as a synapomorphy was th er^ re prioritized for this clade. 101. Vein R I, differentiaUon: (0) shortly tubular or densely nebulous, very narrow (linear); not reachmg midway distance betwemi pterostigma apex and point where R l meets wing ma%in ^ g s 163-166,171, 172); (1) tubular or densefy nd)ulous b^ond midway pomt between pterostigma and wing margin at apex of Rl (Figs 167,168, 169, 173,174); (2) rl entirely absent; wing margin between pterostigma and apex of Rl weakly nebulous or completely clear (Figs 170, 175-178). Additive. Ambiguous at node 47. Optimization is unimportant because correspondent clade is o f reduced interest for the present analysis. Changes as suggested in the preferred cladogram: 0->l->-2, some 1-+0 and 2-^l reversals. 102. Vein (Rs-^M)b, connection with 2-^3Rs: (0) interrupted by a bulla (Figs 163-169); (1) connected and contiguous with one another (Figs 170,171); (2) both veins spectral or, more rarely, nebulous (Figs 172-178). Strictly, state 2 is “inapplicable,” but smce it mdeed represents an apical state, polarizing the progression 0-*^l-+2 is not inappropriate. Non-addhive. Changes as suggested in the preferred cladogram: 0-+l-^2, unambiguous. 78 103. Vein (Rs+A^b, connection with 2-^3M: (0) interrupted by a bulla ^igs 163-166, w); (I) connected and contiguous ^ g . 167); (2) both veins spectral, rarely nebulous (Figs 172-178). If run as non-addhhr^ the transformation lM)-+2 is strongly supported in resulting cladograms. To conform with this hypothesis ^e., two steps for transformations between derived states), character states were renumbered in the matrix as 0*-l-*^2 and run as additwe. Additive. Changes as suggested in the prrferred cladogram: l<-0-^2 (cladogram: 0<-l-^2.), unambiguous. 104. Vein (Rs+M)b, relative length: (0) distinctly short (Figs 163, 164, 168, 169,175, 177); (1) moderately long or intermediate (Figs 165-167, 170,172, 173,174, 178); (2) very long (Figs 171,176). This subjective delimitation o f the character states was due to the foct that measurements such as length/width are highly imprecise for vein (Rs^M)b, because this vem does not have clearly defined limits in all species. Non-addhive. Ambiguous between nodes 15 and 281 hisufficient evidence to support optimization. Changes as suggested in the preferred cladogram: 0-^l-*^2. 79 105. Spurious vein arisingfrom (Rs+M)b, towards base o fpterostigma. (0) absent (Rgs 163-171); (I) present, normally spectral, sometimes nebulous, never tubular ^ g s 172- 175,177,178). Ambiguous at node 38. Optimization is unimportant because correspondent clade is of reduced Interest for the present analysis. 106. Vein 2-^3Rs, shape: (0) straight or slightly curved, tubular QFigs 163-171); (1) strongly sinuate, spectral or, more rarely, nebulous ^igs 172-178). Changes are unambiguous. 107. Vein 4Rs, structure: (0) tubular, or at least intensely nebulous, for nearly all its length, becoming lightly nebulous, spectral, or disappearing completely, only when very close to wmg margin (Figs 163-171,174); (1) tubular part ending 6 r from wing margin, becoming apically spectral, weakly nebulous, or simply absent ^ ig s 172, 173, 175-178). Changes are unambiguous. 108. Vein 4Bs, connection with 2r. (0) distinctly angled, or at least distmclty imperfectly contiguous in some other way (Figs 163-164, 166, 167,170-178); (1) smooth transition from 2r to 4Rs, these vems perfectly contmuous, never forming an angle (Figs 168, 169). 80 Ahhough. the alignment of 2r with 4Rs in Fig. 165 is equivalent to present state I, it represents an exceptional individual variation ofM anomalipes, for which the present character was coded as state 0. Changes are unambiguous. 109. Vein 4Rs, shape: (0) longitudinal, but neither perfectly straight, nor unifermly curved; if straight, not parallel to pterostigma, and usually slightly inclined anteriorly ÇFigs 163- 166,168,193,1827,174,176); (I) entirely and uniformly curved towards anterior margin, even if slightly ^ig s 169,171); (2) nearly perfectly straight and parallel to pterostigma (Mgs 167,173, w); (3) with a short horizontal basal portion, or at least a knob, then forked, with anterior branch tubular, bent towards anterior wing margin, and ventral branch spectral (rarely nebulous) and extending from half as long to as long as anterior branch (Figs 175,177,178). Non-additive. Optimization; ACCTRAN. Anticipates change 0-^ 1 to node 22, providing further support to the Petrastephanettus clade. Justification: state 1 is typical for Parastephanelhis^ occurring fer all spp. of this genus in the cladogram except the smallest on^ P. pyffnaeuSy and this occurrence alone is the one responsible fer making the character ambiguous at this node. However, since the reduced size (rather than mheritance) could easily be related to the lack of curvature of vein 4Rs in this species, it seems more reasonable to assume only one origin o f state 1 for the Parastephanellus spp., instead o f two, as suggested with Deltran. Main changes as suggested in the preferred cladogram: 0->(l, 2,3). 81 110. Veins 4Rsand2r, relative sizez (0) 4Rs more thaa 3x as long as 2r (up to 1 Ix) (Rgs 163-166,169,171); (1) 4Rs from as long as, to not more than 3x as long as 2r (Figs 167, 168,170, 172, 173-178). Ambiguous between nodes 3 and 30. Possible optimizations offer multiple but ^parently equally plausible possibilities, and therefore could not be selected. 111. Veins2+3R5and4Bs, alignment: (0) angled ^igs 163, 165, 169); (1) aligned, even if 2+3Rs sinuous (Figs 164,166, 167,168,170-178); (-) inapplicable (forked; state 3 of character 109). Note that coded state for species illustrated in Rg 164 is 0. The alignment of 2+3Rs with 4Rs in the mentioned illustration represents an exceptional individual variation for Stephanas serrator. Changes are unambiguous. 112. Veins 4Rs and 2-^3M, relative distance at point where they meet wing margin: (0) 4Rs (or its projected trajectory) intercepting wmg margin nearer to wing apex than 2+3M (Figs 163,167, 168, 169,171); (1) interception point of each vem with wing margin equidistant (or nearly so) to wing apex (Figs 164,165,166,172, 174); (2) 4Rs (or its projected trajectory) reachmg wing margin at or very near wmg apex (Rgs 170,173); (-) inapplicable (Rs forked; considered separately in character 109). 82 In case vein ends before reachmg apex, consider a projection o f its trajectory. For observations wings were oriented in the following standart position: wing on vertical position, apex pointing anteriorly, costal margm to the right. Note that (1) although perfoct examples for states 1 and 2 exist (e.g., M brurmeus, Xanthostephanus n. gen.), th ^ are not entW y distinct in some foenatopinae; (2) character state for “Stephamis" tibiator is more or less intermediate between states 0 and I; and (3) coding for “Neostephanus" albomaculatus could be equivalent to state 3, but this could not be confirmed due to limitations on the preservation condition of the examined spechnens. l^run as non-additive, the transformation series l*-0->2 is strongly supported in resultmg cladograms. To conform with this hypothesis (i.e., two steps for transformations between derived states), character states were renumbered in the matrix as0^1-+2 and run as additive. Addhwe. Changes as suggested in the preferred cladogram: l<«-0->2 (cladogram: 0^1-^2), unambiguous. 113. Vein 2+5A^ differentiation', (0) 80% or more (but never entirely) tubular; apex nebulous or spectral ^ g s 164-166,167-169); (I) tubular approximately on basal 50%, or les% remammg nebulous or spectral (somethnes strongly nebulous, cppearfng to be tubular) (Figs 163,170,171); (2) entirely nebulous or spectral ^ g s 172-178). 83 State for “Siephanus" tibiator could corresponde to state 1 abov^ but it could not be confirmed because vem changes progressively fi*om tubular to nebulous to spectral. Additive. Changes as suggested m the preferred cladogram: 0-+M2, unambiguous. 114. Vein CuP, trajectory: (0) sinuous, from slightly convex at level of Icu-a, to concave under vein 2Cua (Rgs 165-171); (1) straight or nearly so, but apically usually curving slightly in anterior direction, along 2Cub; or at least mostly straight, concave under cell 2Cu only (Rgs 172-178). Changes are unambiguous. 115. Cells 1+2R1 and IM, relative size: (0) area of cell Rl up to 2. Ox as large as area of cell IM ^igs 163-170); (1) nearly or more than 3.Ox as large ÇFigs 171-173,174, 175- 178). Changes are unambiguous. 116. Width o fwing membranefrom its posterior margin to CuP, at level o f 2Cua: (0) narrower than distance from apex of vein ICu to CuP (Rgs 163-167,169,171, 172, 177, 178); (1) about as wide (±10%) or distmctly wider (Rgs 168, 170,173-176). The above mterpretation is the most reasonable result o f many different interpretations tried for this character, including comparing the mentioned width with the length of vein Icu-a, length of 2Cua, and width of wing membrane below Icu-a. I also 84 tried to consider the width bdow apex of 2-1A and below apex of 2Cua. Di all these trials, however, intermediate forms made coding too difhcult or too subjective, even with a reduced number of character-states. This character is partially correlated with a few others, especially the position of the retinaculum, which determines the origin o f a slight antero-ventral swelling of the wmg margm, the relative size of vein 2Cua, and maybe with the small transverse, usually spectral vein (and/or wing fold) which originates from the apex of vein 2-1A The degree of elongation o f the wing, and small size o f some foenatopinae may also influence the present character slightly. The current interpretation probably "attenuates" the phylogenetic significance of a very narrow width for this caracter in some species (e g., ScMettererius cinctipes^ “Stepfmais” tibiator), which could perhaps be considered as a separate state, but will help p rient many mismterpretations, as commented above. Furthermore, the characters mentioned above as partially correlated to the present character have all been considered separately, which will hopefidly help to compensate for the lack of a specific state accounting for extreme cases of the present character. Character 114, "trajectory of CuP," may also be correlated with all the characters discussed above, although it seems difficult that it may exert any influence on their mterpretation. Ambiguous at node 36. Opthnization is unimportant because correspondent clade is of reduced interest for the present analysis. 85 in . Reüfutculum, position: (0) mostly under cell 2Cu, hs apex nearest to level of vein 2Cua ^ g s 163-167,171,172); (1) midway between cell iCu and cell 2Cu, its apex approximately equidistant from veins Icu-a and 2Cua (Figs 168,169, 174-178); (2) mostly or entirely under cell ICu, its apex nearest to level of vrâi Icu-a ^ g s 170,173). In evaluating this character, the most important is to consider how much of the retinaculum is under which cell. The position of the apex is an auxiliary detail. Additive. Changes as suggested in the preferred cladogram: 0-+l-*^2, unambiguous. 118. Ciliar hairs between retinaculum and apex o fvein R I, degree o fdifferentiation: (0) hairs lon& originating nearly at and greatly surpassing wing margin (Fig. 1849); (I) few bans, or entirely absent; if present, very short and delicate, originating distinctly before, and not or barely extendmg b^ond wing margin (Fig. 1848). Changes are unambiguous. 119. Ciliar hairs between retinaculum and apex o f vein R l, distribution (not illustrated): (0) hairs ending between apec of 4Rs and 2+3H or at least near 2+3M; (1) hairs ending more or less opposite to middle o f vein 2Cub, far from wing apeq hairs usually minute, or absent Changes are unambiguous. 86 120. HamuH, typical number. (0) usualfy 3-4 (up to 7) I hamuli m at least half of the specimens (Rg. 149), large or very large specimens often with 2-3 Type H hamuli; (I) only 2 Type I, and I Type H hamuli, very large specimens with no more than 3 Type I and only rarely with 2 Type H (not illustrated). Stephanids do not have basal hamuli. The present character therefore refers to the distal hamuli only. There seems to be some correlation between the number of hamuli and size of the specimen (Riek 1970; Quicke 1981; Basibuyuk & Quicke 1997) and this could be readily observed in the specimens examined for the present work as well. Nonetheless, there is a very distmct tendency towards a reduced number of hamuli (state 1) in more apical stephanid species. Very large foenatopinae, for example, may have one extra Type I hamuli, but the total number (3) is often inferior to th ^ o f a specimen of the same size in a more basal species (e.g, 6-7 in «pially large ScMettererius cinctipes^ 4 in equally large M nuzculipermis^ etc.) The number o f each type of hamuli also varies frequently from one wing to another in a same specimen, especially in more basal species. Due to these variations, it is advisable to evaluate specimens of average size for its species, and, if the variation persists, select the most frequent value. Changes are unambiguous. 87 121. H ind wing, shape ofSc+R: (0) straight or neariy so, or uniformly curved toward anterior margin of wing (Rgs 179-184, 186, 187,189, 192, 194); (1) apically slightly bent towards anterior wing margin ^ g s 185,188,190, 191, 193). Changes are unambiguous. 122. Hind wing, differentiation o f veinM Ç^ M+Cu, IM and 2M): (0) distinct, oblique, usually nd)ulous, but sometimes spectral ^ig s 179-184); (1) mostly or entirely spectral, longitudinal, apparently (used to Rs ^ g . 189); (2) absent (Figs 185-188, 190-194). Non-additive. Changes as suggested in the preferred cladogram: 0->2-^l, unambiguous. The observed changes suggest the character could be run as additive, but since the transformation sequece it suggests is highly counterintuitive, I chose not to polarize it 123. Hind wing, differentiation o f vein Ry. (0) usually nebulous, sometimes spectral, long, straight, and longitudinal, its origin distinctly unrelated to Sc+R (Figs 179-182, 184); (1) nebulous or spectral, short, originating near or at apex o f Sc+R, usually curving towards anterior margin, more rarely straight or somewhat undulate (Figs 186-192,194); (2) absent ^ g s 183, 193). Additive. Changes as suggested in the preferred cladogram: (Ml-»2, unambiguous. 88 124. Hindymg, vein Rs, structure o fshort nebulous types. (0) arched towards anterior wing margin ^ g s 186-188, 191,192, 194); (I) weakly curved, or straight, extending towards wing apex (Rg. 189); (2) more or less ondulât^ and approximately horûontal, not bent anteriorly QFig. 190). The “short nd)uIous ^ e ^ correspond to state 1 of ch. 123. Non-additive. Ambiguous at node 48. Optimization is unimportant because correspondent clade is o f reduced interest for the present analysis. Changes as suggested in the preferred cladogram: 0-^ 1, origin o f state 2 uncertain. 125. Hind wing, vein Rs, origin o fshort nebulous types. (0) originating distinctly behind expanded portion of Sc+R which bears the hamuli (Rgs 187,188, 190-192, 194); (I) originating from apex of M+Cu (which is spectral, and runs along and near Sc+R) (Fig. 189); (2) originating at (and fiised to) expanded portion of Sc+R which bears the hamuli (Fig. 186). The “short nebulous types” correspond to state I of ch. 123. The following hypothesis is a tentative explanation for the origin of the present character states: M+Cu moved anteriorty and fused with Rs; in the process, r-m disappeared; later, M+Cu also disappeared. An intermediate stage can be observed on Parastephanellus n. sp. and P. nigricauehis, where a convex spectral structure (M+Cu?), runs along Sc+R and meets Rs. 89 Vem M4Cu cam also be seen in an intermediate stage of fusion with Rs on "Neostephanus" sp. A ^ g . 191), providing some evidence to the hypothesis that these vems are fiised in some species (state I). Non-additive. Changes as suggested in the preferred cladogram: 1MM2, unambiguous. 126. HindMmg, development o f vein Ir-m: (0) present, even if spectral or incomplete; better developed in large specimens (Rgs 180,179, 184); (1) absent (Rgs 181,182, 183, 185-194). Changes are unambiguous. 127. Kind wing, differentiation o f vein Cua: (0) present, nebulous or spectral (Rgs 179, 184); (I) indicated by a stub coming out of M+Cu (Rgs 160,180); (2) absent ^ g s 181- 183, 185-194). State 1 is definitely the case fo t Stephanas coronator, but while it is distmct in large specimens of this species, h can be inconspicuous in medium or small ones. Additive. Optimization: ACCTRAN fiir state 1. Assumes state 1 for ancestral nodes of Stephanas serrator and those of the "Stephanas” pachylomeras clade, preventing the occurrence o f 0-^2 changes. Justffication: conforms with additive hypothesis. Deltran assumes two origms for state 2 through a 0-*^2 change. Changes as suggested in the pr^rred cladogram: 0-+1-+2. 90 128. Hind wing, (^erentiadon o fanterior wing fold: (0) present, complete, although sometimes difficult to see in small specimens; Ir-m, if present, interrupted by dorsal fold OFigs 179,180,184,185); (I) absent; Ir-m, i f present, entire, not mterrupted (Rgs 181, 182,186-194). \ ^ g folds differ ffom spectral veins by being concave creases ^ relation to dorsal surfoce of wing), while spectral veins are convex, and distinctly wider than wing folds. Ambiguous at node 19 (a daie vnÛmHemist^hanus). Optimization is unimportant because correspondent clade is o f reduced interest for the present analysis. 129. Hind wing, differentiation o f posterior wingfold: (0) present, at least basally, although somethnes difficult to see in small specimens (Rgs 179,180, 181,184); (1) absent (Rgs 182, 185-194). Determination of character state forM anonudipes was possible only when examinhig a large specimen. See also comments for previous character. Optimization: DELTRAN. Restricts change 0-^1 to node 17 (Hemistephanus-^ Parastephanellus + Foenatopinae clade). Justification: the occurrence o f state I for all taxa within the mentioned clad^ after its derivation from (0), seems to suggest that reversals are very rar^ ff not untenable, for the current character. In unambiguous l-»0 changes do not occur in the cladogram, leaving no evidence that reversals can occur. These observations agree whh the selected optimization, but are contradicted by Acctran, which hnplies the occurrence of two independent reversals. 91 130 Hindwmg, hasaL shape andwidüt (basal 0 ^ approximately): (0) round, wide, posterior margin convex (Rgs 179,184); (1) normal or weakly concave posteriorly, wmg distmctly not pedunculate ^ g s 180-183, 186); (2) very narrow, for a short distance approximately as narrow as or narrower than width of Sc+R, making the wing distinctly pedunculate (Figs 185, 187-194). Additive. Optimization: DELTRAN. Prevents occurrence of 1-^0 reversals; assumes independent origin of state 1 for Stephanus jxrrator. Justificative: conforms with additive hypothesis. Acctran does not contradict the additive hypothesis, but disrupts it by assuming the occurrence of reversals. Changes as suggested in the preferred cladogram after optimization: 0-»-l->-2, unambiguous. 131. Hirulwmg, epical shape'. (0) round and broad (Figs 179, 180, 184); (1) fi-om narrowed to distinctly poin^ knife-shaped (as in Figs 187,191,192, etc), or ^mmetrically acute-triangular (Rg. 189). Ambiguous between nodes 6 and 15. Possible optimizations are of reduced significance for the analysis. 132. Mesepistemum, dorsal part, pruinosityi (0) mostly or entirely uniformly pilose or pruinose, but moderately sparse to very sparse, with nearly 50-95% of its sur&ce visible through pQoshy (not OlustratetQ; (1) glabrous (not illustrated); (2) pruinosity restricted to anterior G.3-0.5, sparse as in state 0; remainmg glabrous (Rg. 103); (3) a small, well 92 defined, more or less oval patch white pQoshy anteriorly, concealing practically 100% of cuticular suiûce (Rg. 104); (4) patch of dense white pOostQr on area in fi’ont o f dorso f^ical ruga (Rg. 106). The pruinosi^ or pOosity^ of the dorsal part o f mesepistemum is always vertically decumbent towards ventral side, thus distinct from pilosity or pruinosi^ o f main mesopleural area, which is horizontally decumbent in posterior direction. "^Stepfumu^ x yriff shows state 4, but it has also a patch o f dense white pilosity centrally on the dorsal part of its mesepistemum. This patch is virtually identical to the patches occurring anteriorly in some species, coded as state 3. The central patch in S. seyriff^ however, seem to be an autapomorphy for the species, and, fr>r this reason, was not used in the analysis. Non-additive. Optimization: DELTRAN. Delays change (M2 within Foenato/n^r to node 30, by assummg state 0 for nodes 21,28, and 29; suggests independent origin of state 2 for non-foenatopinae taxa. Justffîcation: ambiguity in the present character occurs m function of two possible changes to «cpiain origin of state 4 in “Neostephanus" s^rigi: 0-+4 or 2-^4. Change G-^4, however, is probably better supported by the &ct that it has a precedent at node 7, with state 4 supporting the correspondent clade. State 3, which is similar to state 4, derives unambiguously from state 2 for two other species, “Neostephanus" attuaudi and ‘Neostephanus" sp. A, which, together with “N " seyrigi, are all endemic to Madagascar. Nonetheless, it still seems simpler to accept independent but mcclusive 0-+4 and 2-*3 changes rather than assume different origms for state 4 (i.e., 0-+ 4 and 2-»4). Changes as suggested m the preferred cladogram: 0-»(l, 2,4) and 2->3. 93 \22.M expistemum (except pectus and dorsalpert), size o f hairs (not illustrated); (0) hairs very small or minute indistinct from pruinosity (if present), decumbent, non- ovedapping 0.e., each hair barely or not reaching base of next nearest hair); (1) hairs long and conspicuous, (üstinct fit>m pruinosity (if present), decumbent and distmctly overlapping, or erect This character refers to hairs associated with macrosculpture, especially fbveolation and areolation (usually one hair originating inside and anteriorly on each fijveola or areola), as opposed to prumosity, which, when present is normally much more abundant occurring all over the cuticular surface. Non-additive. Changes as suggested in the preferred cladogram: unambiguous. 134. Mesepistemum (except pectus and dorsal pert), structure o fhetirs (not illustrated): (0) hairs regular looking length from minute to very lon& horizontally decumbent (I) long trichobothria-Iike hafrs, uniform width, semi-erect or erect (30°-90°). Non-addirive. Changes as suggested in the preferred cladogram: 0-*^!, unambiguous. 135. Mesepistemum (exceptpectus anddorxdpert), pruinosity: (0) absent or nearly so, at most with only a small amount of sparse pruinosity antero-dorsally (Rg. 103); (I) occurrmg on 90-100% o f mesepistemum area, moderately to densely abundant 94 concealing 50-95% of its cuticular suiûce (not illustrated); (2) similar to state I, but restricted to antero-ventral part (Kg. 100); (3) sunilar to state I, but restricted to anterior G.3-0.5, often also only on dorsal 0.50-0.75 o f this part ^ g . 101). It is important to stress that in several foenatopinae the pectus is perfectly contiguous with the mesepistemum, that is, the pectus is not flat, and does not form an angle with the mesepistemum, as is the case with most basal genera, as M egischus and Bemstephanus for example. Consequently, pectus pruinosity in the mentioned foenatopinae may lead to the impression that pruinosity is present anteriorly on mesepistemum, resulting in a mistaken interpretation of the character state as 3, instead of the correct 0. Non-addhive. Optimization: ACCTRAN. Favors occurrence of state 1 in ancestral nodes ïac Hemistephanus and Parastephanellus^ implying 1-^3 change for the Parastpehanellus clade. Justification: state 1 is clearly more related to states 2 and 3 than is state 0, and the only optimization consistent whh this observation is Acctran. Optimization whh Acctran for state 1 produces a similar result, but also assumes state 3 fer most ancestral nodes whhin Foenoro/ms; hnplying numerous independent 3-*-0 reversals whhin this genus. Changes in the preferred cladogram: 0-^l-+(2, 3). 126, M esopleuralpit: (0) absent ^ g s 100,104,106); (1) present, often very small, but always accompanied by characteristic transverse or oblique rugoshies arround h (Figs 101, 102,103). Changes are unambiguous. 95 137. Cooca 2 andM esepistem um , p ilo sity in between-. (0) pilose between mid coxa and ventral part of mesepistemum mesally ^ g . 110); (1) glabrous or nearly so (not illustrated). Hairs o n M eawmcdfpes are present in la%e specimens only. Ambiguous at node 34. Optimization is unimportant because correspondent clade is o f reduced interest for the present analysis. 138. Tibia 2 anteriorface, presence ofspirKsr. (0) without spines on anterior fece (not Illustrated); (1) with a few spines or spine-like setae along anterior Ace, inclined toward apexOFigs 112,113). The anterior Ace of tibia is the one which bears the tibial spur. The spines in the mid tibia of stephanids are sometimes intermediate between a seta and a true spine (e.g., in P. rrigricaudus), or are seta-like (F.formosanus), but still recognizable as spines by being shorter and thicker than nearby hahs, and by showing a prominent cuticule around the basis, which does not occur with regular hahs. Spines occur usually in smaller number and are more frequent on apical half of tibia in some non-Aenatopinae. Changes are unambiguous. 139. Propodeum, pilosity related to areohtion/foveolation (not illustratetQ: (0) one mmute hair, usually decumbent, origmating anterioriy on at least a Aw of the areolae or Aveae; (1) completely glabrous (2) each Aveolae with one very small elevation (vestigial hair base?) centrally or centro-anteriorly. 96 la evaluatmg this character, the area of propodeum apical to propodeal spiracle was not considered, because this area has a distinct sculpture and structure in most stephanid species. Propodeal pilosi^ is often difiScult to see, as it is often minute and translucent. It is particularly difficult to see in cyHndricuSy bemg almost completely absent in some specimens. In “Diastephanus” alntaceous, the propodeal sculpture is ^ically non-foveolat^ but one large male has ffiveolae laterally, and hairs associated with i t Non-additive. Changes as suggested in the preferred cladogram: 2*-0-*^l-»^0, unambiguous. 140. Spiracular impression ^opodeum ), (Sfferentiation: (0) absent ^ g , 104); (I) ill- defined, shallow and fiat, if well delimited, then with numerous transverse carinae (Fig. 105); (2) distinctly differentiated, well delimited, concave without transverse carina^ or carinae incomplete and m small number (Fig. 106). Additive. Ambiguous at node 7. Optimization is unimportant because correspondent dade is of reduced mterest for the present analysis. Changes as suggested m the prefèrred cladogram: 0-»l-»2. 141. Spiracular impression and carina vs. line o ffusion o f metapleuron andpropodeum: (0) spiracular carma and impression not differentiated (not illustrated); (1) spiracular impression and carma mdependent of Ime of fiision of metapleuron and propodeum ^ g s 105,96); (2) spftacular carina a t Ime o f fiision o f metapleuron and propodeum, origmating 97 at the propodeal spiracle and ending below 0.e., externally to) flanl^ at pleuropropodeal fovea; spiracular impression usually inconspicuous or absent, or «(tending alongside sphacular carina (Fig. 104). When evaluating this character, it should be observed whether the structures originating at the spiracle are extending towards the flank (/.e., a straight trajectory), or if th ^ are curving down ^icaUy, thus ending at pleuropropodeal fovea. Interpretation difBcultie^ however, arise from the many different topologies o f the metapleuron and propodeum, and dK&rent degrees of fusion and development of the spiracular impression and carina, and of the line of fusion between metapleuron and propodeum (e.g., “Diastephanus" sp. A, “Diastephanus” sp. D). A compromising situation would be finding a straight spiracular impression, with a distinct spiracular sulcus, curving towards the pleuropropodeal fovea, but this situation, as far as investigated, does not occur in stephanids. For state 1 ^igs 105, 96), the overall shape of the spiracular impression and carina may vary. T h ^ may be both straight, ending on or pointing towards flank, or the spiracular impression somethnes curves inwards apically, as m some species of Hemistephanus (e g., H . tener, see Aguiar 1998). When the spiracular carina ends outside flank, it is still separated from the line of fusion of metapleuron and propodeum, ending above the pleuropropodeal fovea. The spiracular carina sometimes is not developed, giving place to an area or “plateau” between spfracular hnpression and line of fusion of metaploiron and propodeum. This area gets progressively wider towards crenulate sulcus. 98 Note that (1) state 2 is usually indistinct in P, nrfoormtus^ but in some specimens the spiracular carina is well developed, and extends to pleuropropodeal fovea; (2) the attribution of state 2îor Foenatopus sp. F and F oem ttopus sp. G was estimated, since the metapleuron+ propodeum in these species is highly modified; and (3) both metapleuron and propodeum are highly modified m '‘Diastephanus" sp. A, “Diastephanus" sp. B, “Diastephanus" sp. C, and “Diastephanus" sp. D, with a resquice o f spnacular carina ending at pleuropropodeal fovea in some specimens; it runs almost in a straight line, mostly as a consequence o f a strongly flattened and elongate metapleuron + propodeum. Additive. Ambiguous at node 26. Optimization is unimportant because correspondent clade is o f reduced interest for the present analysis. Changes as suggested m the preferred cladogram: 0-^l-^2. 142. Metapleuron, Jitsionwith propodeum: (0) metapleuron distinctly triangularly prominent, thus not continuous with propodeum ^ g s 104, IDS, 106); (I) perfectly continuous with propodeum, or only very weakly prominent, not particularly prominent or detached as a triangular structure ^ g . 109). The spiracular groove and carma should not be considered when evaluating this character. These structures have been treated under character 141. Changes are unambiguous. 143. Metapleuron, size o fhairs associatedwith areokttion: (0) firom minute (sometimes absoitinverysmall sperânens), upto about 2x as long as diameter of areola where it 99 originates; delicate and fragile, nonnally completely decumbent (Fig. 102); (I) much longer, usually than 3-5x as long as the diameter o f areola where it originates, or longer; aU hairs strong and conspicuous erected 30-45" (not illustrated); (2) individual hairs as in state 1, but pilority very abundant, concealing neariy 100% of the sur&ce, hiding the sculpture (M texanus only). Nearly all stephanids have at least some coarse areolate-rugose on the mett^Ieuron, with one hair originating mside and anteriorly in each areolation. Pruinosity, or other short, densely abundant pilosity, is a distinct character, considered separately. Non-additive. Ambiguous at node 3, and between nodes 15 and 28. State 1 is definitely closely associated withA/egûcAur spp. and Hemistephanus spp., and a Deltran optimization could probably represent the correct evolutionary path for the character (by assuming state 1 for the correspondent ancestral nodes of these genera in the cladogram). However, possible optimizations are of reduced importance for the analysis because either character 1 or 0 become highly homoplasious ^napomorphies under each optimization. Changes as suggested in the preferred cladogram: ambiguous. Metapleuron^ type o fhtirs associatedwith areolation: (0) hairs regular looking mmute or lon& decumbent or erect; (1) thm and delicate, width uniform throughout lenght, thus similar to a trichobothria (similar to those on Rg. 43), each hair about twice as long as diameter of areola from which it origmates, and 45-90" erect. See comments for character 143. Changes are unambiguous. 100 lAS.MetcpIeuron, sfujrt pilosity not associatedwith areolation: (0) absent (not illustrated); (I) sparse and sometimes present only partially; 80-90% of cuticle surface always visible through pilosity (not illustrated); (2) centrally very densely abundant, hiding 50-100% of the sur&ce where hairs occuir, but glabrous or almost so on dorsal area (not illustrated); (3) as state 2, but dorsal area completely taken by short pilosity, which reaches spiracular carina (not illustrated); (4) restricted to a more or less central oval patch of very dense, moderate-size pilosity (Fig. 106). hi addition to hairs associated with areolation (see chs. 142 and 143), stephanids may also have an mdependent, usually abundant and decumbent pilosity in the metapleuron, which normally correspond to the shortest hairs in this area. Non-additive. Changes as suggested in the preferred cladogram: 0-^(1,2, 3,4). Ambiguous at node 7. Optimization is unimportant because correspondent clade is of reduced interest for the present analysis. 146. Post-fomolar area, association with mterfoveolar area: (0) post-foveolar area more or less aligned with mterfoveolar area (Fig. 105); (1) not aligned with, and not as a contmuation o f mterfoveolar area, appearing more characteristically as the ventral part of the metapleuron; if aligned, or more or less so, then mterfoveolar area deep, contiguous with metapleural fovea, so that post-foveolar area stands in a higher plane than mterfoveolar area (Rgs 104,107); (2) metapleural Avea absent or inconspicuous, post- foveolar and mterfoveolar areas continuous (Rgs 102,109). 101 Addhîve. Optmdzadon: ACCTRAN. Suggests state I for the common ancestor of Parastephanellus + Foenatopinae; explains occurrence of state 0 in Xmthostephanus n. gen. as a l-*0 reversaL Justffication: state I, or its derived form (state 2) are almost mvariably present among species o f Parastephanellus and foenatopinae, and appearing among othe stephanids only in an unusual group of species endemic to Madagascar (node 7). This occurrence o f state 1 almost restricted to apical clades, coupled with the foct that Parastephanellus and foenatopinae are closely related taxa (as sister groups), suggest a common origin of state 1 for these groups. Changes as suggested in the preferred cladogram: 0-^l-»2. 147. Post-foveolar area, shape o f ventral margin: (0) basally produced ventrally into a weak or very prominent lobe of sharp edge, which covers a little of the base of mid coxa ÇRgs 104, 105,106,109); (I) no distinguishable lobe basally, straight throughout, edge not sharp (Fig. 107). State 1 is a distinct synapomorphy for Parastephanellus. No significant variation or mtwmediate forms were observed. Changes are unambiguous. 148. Intercaxal space between hind coxae, sclerotinization: (0) membranous, petiolar foramen nearly always open behind petiole ^ g . 108); (1) covered by a heavily sclerotized plate formed by the retention and connection of metastemum with the posterior angles of propodaun(Fig. 111). 102 ff^the high and generalized morphological plasticity of stephanids is taken into consideration, it becomes somewhat surprismg to note that no intermediate forms were observed for this character. The only variation which may deserve some comments is found in Hemistephanus limpidipenniSy in which the petiolar foramen is closed behmd petiole with, the posterior angles of propodeum meeting behind it to form a large, heavily sclerotized plate; a structure unique for this species (see Aguiar 1998:420, figure 167). Nonetheless, in spite of this plate, the widely membranous intercoxal space va.H limpieBpenms undoubtedly corresponds to the basal state of the present character. Changes are unambiguous. 149. Coxa 3, structure at base: (0) deeply concave on the external side only (Mgs 114, 118); (I) deeply concave both externally and mesally (Figs 115,119). The internal constriction or emarpnation of the hind coxa in state 1 probably helps preventing the interference of the intercoxal plate (ch. 148) with movements of the hind coxae. These two characters might therefore be linked. However, since th ^ represent distinct and extreme specializations, in d^erent parts of the body, I decided not to assume fiill correlation, and considered them separately, foct, these characters are not associated in a fow taxa (e g., Xanthostephanus n, gen. and Foenatopus arcuatuSy examined for this work). Changes are unambiguous. 103 150, Caca 3, relative length (not illustrated): (0) shorter than maximum lateral length of mesepistemum; (1) as long as, or slightly longer (up to l.lx as long); (2) distinclty longer (1.3-l.Sx as long). Small o r medium specimens show less variation than large ones. Large specimens, thœ fore, should be avoided during interpretation of this character. Non-additive. Opthnization: ACCTRAN for state 1. Anticipates change 0-*l to node 3, providing one fiirther synapomorphy isolatmg Schlettererius+Stephanas from other stephanids. Justification: as with any quantitative character, the current definition of character states is inevitably at least partially artffîcial. Nonetheless, a moderately to distmctly elongate hind coxa is typical o f most “modem” stephanids, and the current optimization was selected as a mean of emphasizing this foct, mainly by making state 1 as inclusive as possible. As this is a continuous variable, the many independent origins of state 2 within foenatopinae resulting from Acctran optimization do not seem surprising. Changes as suggested in the preferred cladogram: 0-»l-»2. 151. C oxa 3, shcpe: (0) from globular to moderately elonj^te, but stout (Figs 116,117, 118, 123, 126); (1) very long and slender, 4.25x or more as long as laterally high (Figs 119,120). Several qualitative Q e., non-biometric) codmg schemes were attempted for this character, but m aH cases taxa fix>m basal and apical groups ended up as intermediate forms between states, suggesting that the devised schemes were not accurate. W th the loigth/height ratio approach, however, a “break pomf* was more or less evident at the 104 value 4.0, isolating the foenatopinae from other stephanids. The value 4.25 was chosen to prevent the inclusion ofthe evidmtly intennediate (although well defined) genus ParastepharKÏÏus. Changes are unambiguous. 152. Cam 3, c^erentiation o fa fla t or concave central area latero-dorsalfy'. (0) absent 0 g s 117, 120); (1) present, with different sculpture fi-om the rest of coxa (usually same kind, but finer, or sometimes polished smooth) and usually also with more sparse pilosity, or glabrous (Fig. 116). In state 1 hairs are sometimes completely decumbent, laying into tiny impressions (e.g., ‘W'easftpAamty" sp. A, “Diastephanus" alutaceousY Note: Hemistephanus cylindricus shows a more or less fattened area, but sculpture and pilosity are not distinct from that on the rest of the coxa. This species was therefore interpreted as having state 0. Ambiguous between nodes 21 and 31. Possible optimizations are of reduced hnportance for the analysis because character is exceedingly homoplasious. 153. Cam 3, mesal shape: (0) fiat, sculpture distmct from that of the dorsal and ventral sides (usually foier or mostly polished smooth), pilosi^ denser and with shorter bans than pilosity on dorsal or ventral sides, or glabrous (Fig. 118); (1) slightly convex or distmctly convex, pilosity identical to, and sculptmre not or only slightfy difierent from that on dorsal and ventral sides (Hg. 119). 105 Optmiization: ACCTRAN. Suggests change (M l at node 28, providing further support to the foenatopmae clade. Justification: ambiguity for the current character occurs in function of the character state fbr *7feostepfumus" seyriff. Nonetheless, the observed state for this species is of dffîcult mterpretation, as it may be linked to its reduced size. In addition, state I is a widespread trait within foenatopinae while there are only two mdependent occurrences of state I fbr other stephanids in the cladogram. Combmed, these foots strongly suggest that state I was present in the ancestral foenatopine. 154. Coxa 3, sculpture: (0) non-transverse coarse sculpture, even if polished smooth in between (not illustrated); (I) transverse but irregular, coarsely rugose or rugulose; (2) regularly transverse, fine (e.g., strigate or strigulate) or very fine (e.g., alutaceous) (Fig. 120); (3) non-transverse microsculpture, or polished smooth. Non-additive. Ambiguous at nodes 8,12, and between nodes 29 and 30. Possible optimizations are of reduced importance for the analysis because involved character state at clade o f interest (node 29) is highly homoplasious. (Changes as suggested in the preferred cladogram: O-^ (1,2,3) and 2-*^3. 155. Coxa apico-darsal spine: (0) present, conspicuous ^ g . 123); (1) absent (Fig. 116,117). Changes are unambiguous. 106 156, Coxa 3, pilosity (not illustratecQ: (0) at least externally with long or very lon^ stif^ erected 30^-45**, often overlapping hahs (except non-overlapping in Schlettererius cinctipeSy but this species with at least some very conspicuous hahs, which are longer than femoral teeth); (1) hahs minute, much shorter than basal femoral tooth, decumbent, delicate placed fer away from each other. Pruinosity in between pilosity should not be considered when evaluating this character. Changes are unambiguous. 157. Femur 3. pilosity (not illustrated) (pruinosity in between pilosity not considered): (0) at least mesally with long or very long, stifl^ 30M5® erect, often overlapping hairs (except non-overlapping in SchlettereriuscinctipeSy but this species with at least some very conspicuous hah^ longer than femoral teeth); (1) as m state 0, but all hairs completely decumbent, and more delicate; (2) most or all hairs minute, much shorter than basal femoral tooth, decumbent, delicate placed fer away from each other. Xanthostephanus n. gen, is a distmct intermediate, representing the only foenatopinae with long hairs ventrally on hmd femur. “Stephanu^ seyriff has also a patch of short white pilosity sub-basally on external side, in between the long hairs, but this is an autapomorphy fer this species. Additive. Changes as suggested in the preferred cladogram: 0-+1-+2, unambiguous. 107 158. Femur 3, structure o f the mesal ^obabfy gkmdular) opening basalfy: (0) absent (Kgs 128, 129); (1) pore-lik^ circular, border slightly raised; usualfy placed inside a shallow depresaon; Hghtfy colored around it (yellowish, ambar, or reddish), this coloration sometimes extendmg to whole basis o f femur (Kg. 124); (2) pore-like, elliptical; only rarely associated with a small yellowish spot or area; not necessarily associated with a shallow depression (Fig. 122); (3) a linear, slit-like impression ^ig. 121); (4) only a small depression, apparent^ not a true orifice; not associated with light coloration (Fig. 125). These structures are always situated basally, near the line of connection of femur and trocantellus, on the mesal side. To simplify coding state 4 was renumbered as 0 in the matix. This is justified because state 4 does not correspond to a true opening and is also an unique autapomorphy for “Neostephanus’'albomacuiatus. Additive. Changes as suggested in the preferred cladogram; 0-»l-»2->3, unambiguous. 159. Femur 3, d^erentiation o fa mesal compression: (0) with a dorso-mesal compression, on basal half or centrally, fi-om small and short (as in "Stephanus" tibiator) to relatively large and elongate (most feenatopmae), subtly often dffîcult to visualize (Kgs 126,127); (1) absoit, femur unffî)rmly curved in cross section (not illustrated). Optimization: ACCTRAN fer state 1. Assumes a single origin of state 1 within M egischus+ Hemistephanus + Parastephanellus^ as well as its contmuation through nodes 28 and 29 (origm o f feenatopmae clade). Justification: current optimization is 108 strongly suggested by character state distribution ht the cladogram, which shows dominance o f state 1, with few scattered occurrences o f state 0, among the central and apical AfegMcAuf spp. in the cladogram, and exactly the opposite situation for the feenatopinae. 160. Femur 3, presence o fa tooth or tubercles basad o f median tooth: (0) with one tooth or tooth-like tubercle basad of median tooth (Figs 128, 129); (1) teeth and tubercles of any kind absent (not illustrated); (2) two or more teeth, or several small tooth-like tubercles, or with small regular tubercles, as the ones in between median and apical teeth (Figs 130, 131, 132). Some feenatopmae may seon to have state I, because they show a well developed 3rd tooth, which can be as stout as or stouter than the median and the apical teeth (e g., “Diastephanus” alutaceous^ F. natalicus)y as well as have the same color (e.g., whitish in F. navagumeensiSy Foenatopus sp. F). However, such tooth is accompanied by other small tubercles, which characterize state 2. The only feenatopinae coded with state 1 in the present work was “Madegafoenus” (^Foenatopus) seyrigiy but the lack o f small tubercles in this species may be related to its reduced body size. Non-additive. Optimization: ACCTRAN fer state 1. Restricts change 1-^2 to node 21, lending mctra support to Parastephanellus + Foenatopinae clad^ and considering state 2 in Hemistephanus macrurus as an isolated ^araHeQ occurrence. Justification: it is difficult to decide whether the large size of AT. m acrurus is related to the presence o f basal tubercles in its hind femur, but, as observed during this work, such a tendency mdeed 109 seems to occur among some stephanids and therefore cannot be ruled out for the mentioned species. This is significant because if state 1 inÆ m acrurus is size related, then its nature is diferent of state I vaParastephanettus and feenatopinae. ^ so , it is better interpreted as an independent occurrence as suggested by Actran. Additional independent occurrences o f state 2 On the “Stephanus”pachylomerus clade) confirm the possibility o f parallel evolution for this state. Changes as suggested in the preferred cladogram: 0-+1-+2. 161. Femur 3, structure o f setae arising on tubercles behind epical tooth: (0) setae not particularly thick and stifi^ neither whitish, and not organized in a distinct comblike feshion (Figs 128, 129); (1) setae thick and s t^ usually perfectly straight; all with the same length and more or less equidistant to each other, similar to a “comb”; usually whitish (or similar), sharply contrasting with dark femur (Figs 131,132); (2) setae absent, tubercles with very small, delicate curved hairs (not illustrated). There is normally one single seta or hair orignating apically (or almost so) on each tubercle. Non-additive. Changes as suggested in the preferred cladogram: 1^0-^2, unambiguous. 162. Tibia 3, structure centredfy Cursed view): (0) weakly laterally compressed on basal 0.1, then widening progressively and moderately towards apec (Hg. 133); (1) laterally compressed on basal 0.3-0.5, dilated apically O^g. 134); (2) laterally compressed on basal 0.3-O.S, then firom moderately (not illustrated) to intensely compressed dorsally, then 110 dilated O^ig, 135); (3) as m state 2, but with a deep and mtense depression dorso-mesally, generating an edge or carina extemo-dorsally, which crosses all the way to the ventro- mesal side ^ g . 137); (4) as in state 3, but depression entirely shüted to the mesal side, so that its bordering carina or edge extends from the ventral side to the dorsal side, where it advances obliquely over base of compressed part (Fig. 136); (5) as in state 4, but without a dorso-longitudinal edge or carina on dilated part (ventral one distinct) (Fig. 138); (6) as m state 2, but dilated part with a dorso-basal depression, and a strong transverse ventral depression, also basally (Fig. 139). To allow the character to be run as additive, state 6 (applicable only to M brunneus^ and deriving from state 2) was renumbered m the final matrbc, being recoded as state 2, with which it seems to be related. This character is highly modified for F. arcuatus, but general structure is more similar to state 5 than to other states. Additive. Ambiguous at node 4. Optimization is unhnportant because correspondent clade is of reduced interest for the present analysis. Changes as suggested m the prrferred cladogram: 0-+l-^2-»(3,4->5). 163. Ubia 3, ventro-longitudinal carina: (0) dK&rentiated along compressed part only, albeit often advancing a little over dilated part ^ig. 141); (I) dK&rentiated along entire length o f tibia, conspicuous from base to apex, albeit usually as a sharp edge on compressed part, and as a carina on dilated part; sometimes weak on dilated part, disappearing befr)re reaching apex (Fig. 142). Ill Carina o f state 1 is often difScolt to visualize in lightly colored specimens, especially along the dilated part. Opthnization: DELTRAN. Assumes independent OparaileQ origins for state 1. Justification: state 1 appears within three reasonably independent clades (all with maximum branch support), each one containing species from a well defined biogeographic area, and in a heterogeneous way within two of these clades Q.e., not for all spp. in the clade). These Acts seem to offer better support to the hypothesis of parallel origins for state 1. 164. Ubia 3, relative size o fapical spury. (0) both spurs of same length (not illustrated); (I) mesal spur longer, even only slightly so (Figs 141,142). Ambiguous at node 3. Possible optimizations are of reduced importance for the analysis because character is exceedingly homoplasious. 165. Tcysus 3, number o ftarsomeres on fem aley (0) five (Fig. 146); (1) three (not illustrated). Changes are unambiguous. 166. Basitarstts 5, raw ofspines ventralfy: (0) with 1-2 rows oflon& translucent spines or stout hair^ which are bigger basally, gettmg smaller and weaker toward apex; (as hi Fig. 146 or more delicate) (Hgs ); (1) with 1-2 rows of short, opaque, conical spines, which get smaller and less distinct toward ap«c (Aguiar 1998:385, figure 63). 112 Dffîcult to see in most specie^ because spines are usually located in a densely pilose basitarsus, and some spines or setae are not distinctly differentiated. The double row o f spines/seta^ however, is additionally characterized by bemg glabrous on the space between the two rows. Changes are unambiguous. 167. 72 andS2, fitsioni (0) distinctly isolated (Rg. 143); (1) fused into a cylindric and elongate petiole ÇRg. 145). Changes are unambiguous. 168. 72 (orpetiole), length: (0) about as long as dorsal length of T3 (Fig. 143); (1) approximately I.6x as long as dorsal length of T3 (not illustrated); (2) two times or more as long (not illustrated). Females of “Diastephanus" alutaceous have petiole nearly I.8x as long as T3, but males have this value higher than 2.0x. Because of this, this species was interpreted as hawngstatel. Non-additive. Changes as suggested in the preferred cladogram: 0-+2-»l, unambiguous. 169. Petiolar spiracle, differentiation: (0) closed, unfunctional, mdicated by small, usually mconspicuous lateral tubercles, located near, but not reaching the middle (usually more easily detected m dorsal view) ^ g s 143,145); (I) completely absent (not illustrated). 113 There are several additional and similar tubercles near the middle on petiole o fai limpidtpemis, making mterpretation o f state fbr this species somewhat more dffîcult than fbr other stephanids. Ambiguous at nodes 5 and 34. Optimization is unimportant because correspondent clades are of reduced interest for the present anafysis. 170. Petiole dorsally, types o ftransverse sculpture (not illustrated): (0) basal 0.3 or more transverse rugulose, this sculpture becoming finer or polished smooth apically, (I) transversely strigate; (2) transversely strigulate, very fine, and very regular; (-) inapplicable (mostly or entuely with non-directional microsculpture). Additive. Changes as suggested in the preferred cladogram: 0-»l-»2, unambiguous. 171. Petiole dorsalfy, sculpture orientation (not illustrated): (0) transverse (rugulose, strigate, or strigulate); (1) non-directional (except sometimes basally). Changes are unambiguous. 172. Tergite 3, sculpture (not illustrated): (0) polished smooth (basal rugulosities not considérer^; (1) enthrely or partially covered with some kind of microsculpture; which is usually also present on terghes 4-6. 114 IxlH. limpieSpemds T3 is polished smooth for some specimens. Interpretation of state 0 for P. rufoomatus is not defiihive, as T3 has a waxy texture which complicates character state interpretation. Ambiguous between nodes 6 and 30. Possible opthnizations are of reduced significance fi)r the analysis. 173. Paratergites (T3), devehpment: (0) not dif&rentiatied, or short, distmctly not wide enough either to meet or overlap ventrally ^ig . 143); (I) wide enough to meet or overlap ventrally; or indeed meeting or overlappm& so that 3rd abdominal segment is nearly completely encased by T3 (Rg. 144). Paraterghes of T3 may appear either detached, and therefore easily seen as a pair of hanging flanges, or closely applied under S3, in which case earful examination wiH be needed for correctly mterpretmg its degree o f development. Changes are unambiguous. 174. Stemite 7, prexnce o f specialized xiae apically. (0) all setae an hairs regular lookmg (not illustrated); (1) with stout, curved, ^ically flat, translucent setae latero- ventrally (sometimes only one on each side) ^ g s 147,148). These setae are highly specialized m “Diastephemus" sp. A, but can still be interpreted as belonging to the general type described for state 1. Changes are unambiguous. 115 175. Ovipositor sheaÉhs^ coloration: (0) white- or yellow-banded subapically, (I) unicolorous, or progressively dadcer towards i^iex; (2) almost entirely yellowish-brown, except suddenly black or dark brown on apex. Additive. Optimization: DELTRAN. Assumes independent origins for state I for Stephanus and basal AAgwcAny spp. Justification: the scattered distribution of state I for M egischus spp., as well as personal observations, suggest that an unicolorous ovipositor in this genus tend to occur more or less randomly at least within this taxon, but probably also geographically. Changes as suggested m the preferred cladogram: (M l-»(0,2). 116 111111111122222222223333333333444444.44 Taxon. 012345678901234567890123456789012345678901234567 Diastephanusflavomaeulatus 020220000001120123000111231100012010110020123002 Diastephanus alutaceous 2312000000011210020201114111-2-02-10110021123001 Diastephanusflavidentatus 020220000001120103700111231100012010110020123002 DiasUphanus n. sp. 0372010-0101720012020111101100000010110021------0 - Diastephanus a 0102-0—11-1120103000111231120002010110020023002 Diastephanus sp. b 23020110000112-----30201112311-0-12—0-10020103002 Diastephanus sp. c 2302010000011210030201111011-0-02-1011002010300- Diastephanus sp. d 03020100000112000302011123110000201011002010300- Foenatopusformosanus 020220000101120102700111231120012010110020123002 Foenatopus annulipes 000120000101120123000111231120013010110010023002 Foenatopus arcuatus 022200000701120121020111100121002100110020102001 Foenatopus aurantiiceps 010120000101120103000111231120012010110110023002 Foenatopus natalicus 2312000000011210120201114111-0-02000110021123001 Foenatopus novaguineensis 100200000101121012020111411120-02000110020123001 Foenatopus 000220000001120102000111231120001010110010023002 Foenatopus sp, b 031220000001120102000111231100002010110020123002 Foenatopus sp,c 000210000001122102120121231020012010110010123002 Foenatopus sp, d 020220000101120123020111231120002010110010123002 Foenatopus sp, c 031200110101122002020111-11102001010110020107701 Foenatopus sp. f 120201000101121002020121410100-02000110020223001 Foenatopus sp. g 112200100001122002720121411102001000110020222001 Foenatopus sp, h 100220000101121002020121411120003000110010123002 Foenatopus sp, t 1302000001011210030201212011-000201011002010300- Hemistephanus macrurus 010110000001110103001001121110010100010110101001 Hemistephanus cylindricus 012110010001110123001011-21110110101110120101001 Hemistephanus elimatus 010110000001110111001001-21110100111210110011002 Hemistephanus limpitttpennis 010110000001110103001001-00110100101110010011001 Matkgafoenus seyrigi 002100110001011012120100231100012010110020702001 Megischusforcatus 000111000001100103000000000010001000010110101011 Madegafoenus bekylianus 032220110001011212040010-01000001100010011201001 Madegqfoenus occiputalis 012121110001710123040010-01001001100010010201001 Megischus anomalipes 001120110101111202000000111011001000110010101011 Megischus brunneus 0021110-0111110212000000310011001000111010201011 Megischus califomicus 0021211-0111111211010000111011001000121010101011 Megischus maculipennis 010211000101100103000000000010001100010110101011 Meffschusn, sp. 0202210001211002210100010000101020001100-0101001 Continued on next page. Table 1.3. Data matrbc for SS species and 176 morphological characters used in the analysis. Symbols: 0-5, character states; a-d, polymorphisms, a = 01; b = 02; c = 12; d = 23. 117 Table 1.3 (Continued): 11111111112222222222333333333344444444 Taxon 012345678901234567890123456789012345678901234567 Megfsdms rufofemoratus 000211000001100202000000000010101000010110101011 MegixJtus texanus 0321cl010101111211010000101002001000101011101011 Neostephanus albomacutatus 002220110001111211000111-11102001010110010103001 Neostephemus alluautB 1101-1—10-11201230001112311-0012000110010123002 Neostephanus sp. a 030110000001120122000111101120010000110021123002 Parastephanellus pygmaeus 032101110101111003100101231700003000110001202001 Parastephanellus denotatus 01215111010111------00111231120003000110010202001 Parastephanellus granulatus 002121110101110211100101211000003000110110202001 Parastephanellus n. 002121110101112003100101201120003000110110202001 Parastephanellus nigrlcaudus 020121110101110012100101201100003000110110202001 Parastephanellus rufiomatus 002101110101111012100101231020003000110010202001 Parastephanellus sp. 002121110101112103100101201000003000110010202001 Schlettererius cinctipes 0000- 0— 0000000000000000000000000000000000000000 Schlettererius determinatoris 0300-0—0000000000000000000000000000000000000000 Stephanus gigas 01102000010107121103000000000100100000001010-171 Stephanus pachylomerus 011010000001000123000000230010001000010110000001 Stephanus serrator 001201100101001010000000101001001000010010001011 Stephanus seyrigi 030110010001000123020000-00000001100010111201001 Stephanus tibiator 01101000010100121103000031000100100002101000-101 Stephanus xanthocephalus 020111000101110123000000101010002000110010201001 445555555555666666666677777777778888888888999999 Taxon 890123456789012345678901234567890123456789012345 Diastephanusflavomaeulatus 0030101-0200111012011112200111102111112201211320 Diastephanus alutaceous 002010120100111002101112200001102001102011201320 Diastephanusflavidentatus 0030101-02000-1012011111200111112111111202211320 Diastephanus n. sp. 1-----0-1-0200111012011112200001112011712211201320 Diastephanus sp. a 1-----101-0200111012011112200111102111111212211320 Diastephanus sp. b 1-----001-0200111012011112200001112111112211211320 Diastephanus sp. c 0030001-0200111012011017200001112111111211211320 Diastephanus sp. d 0030001-0200111012011711200001112111112211211320 Foenatopusfbrmosanus 0030101-02000-1012011112100111102111111112101320 Foenatopus annulipes 003010121200100002012111100011202111111112102320 Foenatopus arcuatus 00100-0101000-1012101112200011112001102111101320 Foenatopus aurantiiceps 0030101212000-001201211C100011202101111102102320 Foenatopus natalicus 002110020100111002101112200011102111112111101320 Foenatopus novaguineensis 00201002020010101201111220010111211111121Ilal320 Foenatopus sp, A 0030101-02000-0012012112100101102111111112201320 Foenatopus sp.b 0030101-0200111012011112200101102111111012201320 Foenatopus sp,c ______0030101-02000-1012012112100111202111111212102320 Contmued on nect page. l is Table 1.3 (ContmuetQ: 445555555555666666666677777777778888888888999999 Taxon 890123456789012345678901234567890123456789012345 Foenatopus sp.d 0030101-02000-1012011112100111102101101001001210 Foenatopus sp .t 0—0101-0200111012011112200111102001102012101320 Foenatopus sp,f 002010020200101012011112200001112111712201101320 Foenatopus sp, g 002110020200101012011112200011112111712211101320 Foenatopus sp. h 002010020200101012011111200011112111102201101320 Foenatopus sp.i 0030001-0200111012011112200111112111101211211320 Hemistephanus macrurus 00010-0012000-0001001101111000201100002002000300 Hemistephanus cylindricus 0001100012000-020110-101311000201100001001000300 Hemistephanus elimatus 1-----100212000-100100-101311000202100012101000300 Hemistephanus limpitBpennis 00000-0012000-000110-101311000201100012001000300 Afadegajbenus seyrigi 01-00-0001000-0012011112200011100001011111200320 MegischusJUrcatus 0001100012110-0101011001011000000100002000000100 Madegafoenus bekylianus 01-1110002000-0012102101100000000001001310000100 Madegafoenus occiputalis 01-0110002000-1002101107370000000001001310000100 Megischus anom alipes 00000 - 1201120-0002101001100000000000001000000201 Megischus brunneus 00010 - 0012110-0101101001000000010000002000000100 Megixhus califomicus 00000 - 1-0112100012101001100000100000002000000201 Megischus maculipennis 00010 - 0002110-0201011001011000000100002001000100 Me^schustt. sp. 02-11000-0770-0001101001011000000100012001000201 Meffschus rufofemoratus 00010-0002110-0002101001011000000100002001000201 Megischus texanus 00000-1201120-1012101001100000100000002000000d01 Neostephanus albomaculatus 0770101-02000-1012011112200011102001111012001301 Neostephanus alluaudi 7000100202000-1012011112200001102101111111001210 Neostephanus sp. a 0000100202000-0012012112200011102101111112001310 Parastephanellus pygmaeus 00200-0101000-0012101111100001000000012111000310 Parastephanellus denotatus 00100-010100101002101101200001000100011111000310 Parastephanellus granulatus 001010010100110012101101200001000100001111000310 Parastephanellus n. sp. 0010100101000-0101101101100001100100012111000310 Parastephanellus nigrlcaudus 00100-0101000-0012011111100001001100011101000310 Parastephanellus rufoomatus 00200-0101000-0012101111200001001000011111000310 Parastephanellus sp. 00100-0101000-0012011111100001000000011101000300 Schlettererius cinctipes 00000------0000-0000000000000000010000000000000000 Schlettererius determinatoris 00000------0000-0100000000000000000000000000000000 Stephanus ^gas 00010- 0001110-0002101101001000000100001000000201 ^ephanuspachylomerus 02 - 00 - 0012000-0001101001011000112100001001000100 Stephanus serrator 00000-0001001000020100010000000000000010000002a0 Stephanus seyrigi 01-1110002000-0002101001311000000100001000000101 Stephanus tibiator 02-0100012020-000210100101111000cl00000000000100 Stephanus xanthocephalus 0010100001000-0101011111000001202101001000002210 Contmued on next page. 119 Table 1.3 (Contmued): 11111111111111111111111111111111111111111111 999900000000001111111111222222222233333333334444 Taxon 678901234567890123456789012345678901234567890123 Diastephanusflavomaeulatus 222112222111001102110111102100121121100311101210 Diastephanus alutaceous 222112221111031--2110111102100121121200311111210 Diastephanusflavidentatus 222112227011021112111101102100121121200011101210 Diastephanus n. sp. 222112221111031--2111101102100121121200311101210 Diastephanus sp. a 2221122270110211121111011022—121121200011101210 Diastephanus sp. b 722112227711031—2111101102100121121100017101210 Diastephanus sp. c 212112221011031—2110101102100121121200310101210 Diastephanus sp. d 222112221011031—21111011022—121121200310101210 Foenatopusfbrmosanus 212112222011021122111201002100121121200011701210 Foenatopus annulipes 212111221111021122111211012120121121200311101200 Foenatopus arcuatus 222712227771021712111111102100121121200317101210 Foenatopus aurantiiceps 212111221111021122111211112120121121200311101200 Foenatopus natalicus 222112221111031—2110111102100121121100011101210 Foenatopus novaguineensis 222112220011031—2111101102100121121200311101210 Foenatopus sp.u 212112221111001122111211012120121121200311101200 Foenatopus sp.b 222112221111001102111111102100121121022311101210 Foenatopus sp. c 222110222011021122111211101111121121100011101210 Foenatopus sp. d 211111200110001112101111112100121121200011101200 Foenatopus sp.t 722712227111031—2111111102100121121100001101200 Foenatopus sp .t 222112227011031—2110101102100121121200311101210 Foenatopus sp. g 222112227771031—2110111102100121121200311101210 Foenatopus sp. b 722112227011031—2110111172100121121100011101210 Foenatopus sp.i 222112221011031—2110101102100121121200311101210 Hemistephanus macrurus 201111010000101010001100012100121121010100002101 Hemistephanus cylindricus 212111010000110010000100112100120121010100002111 Hemistephanus elimatus 212111010000110010000100112100121121010100002111 Hemistephanus limpitBpennis 212111010000110010000100112100120121010100002111 Matkgafoenus seyrigi 2221122271110001121000001022—121121400010171210 MegischusfUrcatus 0101010100000011100000000000— 021010010200011101 Madegqfbenus befylianus 2120100100001101000000001000—020011410000012101 Matkgafoenus occiputalis 2121100100000101000000001000— 721111400000711201 M egischus anom aiipes 2110100110000000000000001000— 121011000000001100 Megischus brunneus 0100011100000010100000000000— 011010010000021101 Megischus caiifbmicus 0111000100000001000000001000— 021011100001001100 Megischus maculipennis 2101010100000011100000000000— 021010010100011101 M egischus XL sp. 2111010100001011100000000000— 120011010100001101 Megischus rufbfémomtus 2101010100000011100010000000— 120010010000021101 Megischus textnnis 0121000110000001000000001000— 121111100000002102 Neostephtntus albomaculatus 202110221111001102110011112100121121200010101100 Continued on n%t page. 1 2 0 Table 1.3 (Continued): iiiiiiiiiiiiiiiiiiiiiiiiiim iL iiiiiiiiiiiii 999900000000001111111111222222222233333333334444 Taxoa 678901234567890123456789012345678901234567890123 Neostephamis cdlumuU 222101221111001112011111112100121111300011110210 Neostephamis qi. a 212111221110001102111111112100121121300010101200 Parastephanellus f^gmaeus 20211011200000010101000110????121121210300102100 Parastephanellus denotatus 012110112000010111010001102100121121011300101200 Parastephanellus granulatus 222110111000010101000001112100121121111000101100 Parastephanellus n. 012111111000011111000000001111121121000100111101 Parastephanellus ntgricaudus 012110112000010111010000101111121121200300101200 Parastephanellus rujbomatus 022110112000010111010001102100121121111000101200 Parastephanellus sp. 022110112000010110010000101111121121200300102100 Schlettererius cinctipes 0100000100000000110000000000— 000000000000000000 Schlettererius determinatoris 0100000100000000110000000000— 000000000000000000 Stephanas gigas 0110000110000210100000000000— 021000111001011100 Stephanas pachylomerus 1001010110000011100000000000— 000000010000011101 Stephanas senator 0200000100000000000000000000— 010010000000100200 Stephanas seyrig} OlllOlOllOOOalOOOOOOOOOOOOOO— 021111410000012101 Stephanas tibiator 1011010010000211100000110000— 000000011001011100 Stephanas xanthocephalus 100112111000001121001200002102121111010100101201 11111111111111111111111111111111 44444455555555556666666666777777 Taxon 45678901234567890123456789012345 Diastephcmusflavomaculatas 00101111102112202150110121201110 Diastephanus alutaceous 00201100113112102150110120-11112 DiastephanusJlavidentatus 00101121102112202150110120201110 Diastephanus n. sp. 0010112101211210215011012020117? Diastephanus sp. a 00101121012112202150110120201111 Diastephanus sp. b 00201111012112102150110120201100 Diastephanus sp.c 00101121012112112150110120201101 Diastephanus sp. d 00101121012112112150110121201101 Foenatopusfimnosanus 00101121012112302150110120201110 Foenatopus annulipes 00101121012112302150110120201110 Foenatopus arcuatus 00101010117112112150110120201101 Foenatopus aurantiiceps 00101121012112302150110120201110 Foenatopus natalicus 00201110113112102150110120-11102 Foenatopus novaguineensis 00101120102112212150110120201101 Foenatopus sp.^ 00101121012112202150110120201110 Foenatopus sp.\i 00101110112112202150110120201110 Foenatopussp.c 00101121012112202150110120201110 Foenatopus sp .i 00101121012112312050110120201010 Foenatopus sp.e 00101110112112002150110120201001 Continued on next page. 1 2 1 Table 1.3 (Continued): lllllllllllH lllllllllllllllllll 44444455555555556666666666777777 Taxoa 45678901234567890123456789012345 Foenatopus sp.f 00101121002112102150110120201100 Foenatopus sp. g 00001110103112102150110120201101 Foenatopus sp.h 00201120112112102150110121201002 Foenatopus sp.i 00101121012112112150110121201101 Hemistepiianus macrurus 03000010002100002040011120100000 Hentistephanus cylintMcus 03000010002100011010111120101000 Hemistephanus ellmatus 03000010002100011040011120101000 ffemistephanus limpidipennis 0300001000al00011040111120101000 Madegafoenus seyrigf 00101110103112011250110120-10102 Megischusfircatus 0200001000al00011021110120100000 Madegajbenus bekylianus 00100010003100001021110120-1110? idadegajbenus occiputalis 04100010002100011021110120-11100 idegf^tus anomalipes 00000000000100001020110120001000 Megischus brunneus 00000000001100011020010120100001 Adegischus califomicus 00000000003100001020010120-11000 Adegischus maculipennis 02000010000100011021110120100000 Adegix/nts TLsp. 01000010101100011030110120101001 Adegischus rujbfomoratus 01000010000100011020110120100000 Megischus texanus 0-000000003100001020010110-11000 Neostephanus albomaculatus 00101110013112002050110120201110 Neostephanus alluaudi 00101111112112202150110120201110 Neostephanus sp. a 00101110112112202150110120201110 Parastephanellus pygmaeus 00110010102100012040110127100001 Parastephanellus denotatus 00110010003100012040110120101011 Paraste^anellus granulatus 00110000102100012040110120101001 Parastephanellus n. sp. 01110010012100011040110120101101 Parastephanellus nigricaudus 00110010002100012040110120101101 Parastephanellus rujbomatus 00110010102100012040110120100001 Parastephanellus sp. 00110010102100012040110120100101 Schlettererius cinctipes OOOOOOOOOOaOOOOOOOOOOOOOOOOOOOOO Schlettererius determinatoris 01000000000000000000000000000000 Stephanas gigas 1000000000110001122011072??????? Stephanaspacfylomerus 00000010000100012231010120100101 Stephanas serrator 00000000003100000010000100000001 Stephanas s^rigt 04100010000112001030110120101000 Stephanas tibiator 10000100010100002231110121100001 Stephanusxanthocephaltts 03001021012101002040010120201011 1 2 2 PHYLOGENETIC ANALYSIS The data matrix for the examined stephanid taxa is shown on Table 1.3. Tree searching was performed with heuristic analyses in PIWE v. 2.8 (GolobofiT 1997), and NONA V. 1.6 (Goloboff 1993b), including a separate search in NONA aided by Ratchet (Nbcon 1999). Searches in NONA were performed with the commands “hold 1000, mult 1000” and “hold 1000, mult*50.” Further swapping of the cladograms found by “mult” was recommended, and then performed with “max*.” For Ratchet, a sample number of 27 characters was selected (15% of the total), and 3000 iterations performed in three simultaneous searches of 1000 iterations each. The resulting shortest trees were then submitted to successive approximations in NONA. For searches in PIWE the options “hold 500, mult*50” were used. For comparative purposes, separate searches were performed for all possible values of Ü, the constant of concavity. In all cases, further swapping of the trees was “unnecessary” according to the program. However, for the search with K = 1 different runs of PIWE using the same settings and matrix generated different results each time, producing cladograms with different topologies and fits. For this reason, a more thorough investigation was performed with ^ set to 1, in distinct and increasingly exhaustive searches: (1) multiple independent searches were performed with hold 500/mult*50; (2) the best trees fiom these searches were submitted to further swapping with max*, a procedure which also certifies that the resulting trees will belong to a “complete island” (cited from PIWE manuaQ; (3) resulting trees were then submitted to the command jump*3 Qiigher values produced the same results), which searches for better trees in 123 dif&rent “island^ by generating slightly less optunal trees from the ones found before and further swappmg them; (4) resulting trees were then repeatedly submitted to further swapping by using sswap* and tswap*, which did not find better trees. Support for the branches in the preferred cladogram was assessed using Bremer support (Bremer 1994, KâUerqô e t al, 1992) as implemented in PIWE. Note that with implied weights the Bremer support for each branch is expressed as a function of the difiference in fit (not steps) necessary to dissolve the correspondent clade. Ten thousand trees (defined with “hold 10000”) up to 1.0 less fit than the optimal were generated and investigated. After the evaluation of the whole data set was completed, a separate search was conducted for the taxa of fisenatopinae. The objective was to generate additional comparative information to help support or refute clades within fbenatopinae, a group which, albeit monophyletic, showed highly unstable mtemal resolution during the analyses of the complete matrbc For this search, Schlettererius spp. and Stephanus serrator, the most basal taxa in the âmify, were used as outgroups, and aU other non-foenatopinae taxa removed from the matrix, rather than simply deactivated. Character state numbering affected by the removal of taxa were revised accordingly. Characters 17,30,31,35,38,49, 117, and 140, polymorphic and unordered in the mam matrbc, became bmary in the reduced matrbc, and were run as additive. With the reduction in number of taxa, characters 8, 10,20,30,32,35,38,39,45,46,49,53, 58,59,73,74,95,108, 125,133,134,143,144, 145,147,163, and 166 (1st character= ch. 0) became uninfbrmative (identified with the command “mopup” in Clados), and were 124 removed 6om the matrcc Tree search for the foenatopmae was performed with PIWE only, usmgJC= I and following the same search procedures described above for the complete matrix. Analyses m NONA and Ratchet each produced 36 most parsimonious cladograms ^fPCs), all with 1145 steps. Cl 24, and R I72. Nonetheless, the strict consensus trees for each set of 36 MFCs (Figs 195 and 196) are not identical, with a less resolved consensus for the MFCs found with Ratchet, which has 62 extra steps. This indicates that NONA alone foiled to discover some of the tree topologies found with the aid of Ratchet. The MFCs found with Ratchet represent stable explanations of the data with equal weights applied to all characters. However, this approach implies that all characters provide equally strong evidence of relationship, an idea which is immediately compromised by the foct that some characters are much more homoplasious than others. The most widely used method that deals with this problem is successive approxhnations weighting, first proposed by Farris (1969). It assigns weights to characters according to their rescaled consistenqr mdex (Farris 1989), which combines a measure of homoplasy of the character (cQ with a measure of its usefulness in supporting clades (rQ. hi the process, the weighted characters in the MFCs found by simple parsimony are reconsidered: clades supported by the most parsimonious solution for characters with higher weights are preferred, even ^ it means conmdering solutions which are less parsimonious for the low weight characters than those achieved for them in the 125 unweighted search. The resulting cladograms are reanalyzed to produce a new set of MFCs, which are then used to assign a new series o f weights. The procedure is repeated until a stable solution is achieved. Both Hennig86 and NONA can perform successive approximations, but NONA is more accurate in that it assigns weight values ranging from 1-1000, against 1-10 m Hennig86. Furthermore, Hennig86 cannot process polymorphic characters. For these reasons, only NONA was used for such calculations. Calculations were «cecuted with the commands “run swtju n mult*50,” with an arbitrary choice of five iterations. Successive approximations for each of the two sets of 36 MFCs (found by NONA and Ratchet) stabilized at the second iteration, producing the same single cladogram, with length 1146 (one step longer than original cladograms). Cl 24 and RI 72 (Fig. 197). Successwe weighting is adopted by many contemporary cladists, mostly because there is a wide consensus about the value of weighting and the validity of successive approximations as a method for implementing it Yet, the situation seems inconsistent. As pointed out by GolobofT(1993), “fr’the data are properly weighted, those results alw ays are to be preferred, regardless of the results under equal weights” (emphasis is mine). Then, if the phylogenetic mformation provided by each character is different, and if this dË&rence can be objectively assessed in any cladogram (e g., by using the rescaled consistency index), why should parsimony precede weighting instead o f proceed whh it? ^ character weightmg is embraced as a valid approach, then the only logically consistent procedure is to consider it for all cladograms generated during a search. 126 Sûnpie parsimony and weighted parshnony are not only cfadistîc tools to help interpret data, but also represent different hypotheses about character evolution. As such, it seems contradictory to evaluate the same infi)rmation with a combination of both approaches. Obviously, if characters reflect evolution homogeneously, then any weightmg method should produce the same results as simple parsimony, because the same weight would be attributed to all characters. Nonetheless, the opposite is not true: if characters are differently accurate m how th ^ reflect phylogeny, then simple parsimony would produce results as inaccurate as the amount of ambiguous information (homoplasy) inherent in the characters used. Weighted parsimony therefore must be used as a self- sufficient method. Weighting based on homoplasy is necessarily an a posteriori method, since homoplasy can only be defined in relation to a cladogram. It may therefore seem appropriate to generate the initial cladogram(s) via simple parsimony. This procedure, however, will limit character weighting possibilities by implying an initial weight (or range of weights, if more than one MPC) for each character. All subsequent cladograms will then be a reflection o f the weights suggested by the initial cladogram(s). The equally weighted cladograms, therefore, do not woric simply as a “starting point,” but ultimately prevail over character weighting by constraining data interpretation to its influence. This is particularly true ffir successive approximations, where, as noted by Goloboff (1993), weights are assigned before a search starts and are not modified until it is completed. Another signfficant problem with successive approximation as currently implemented is that character weights are defoied in relation to several cladograms at 127 once, e.g^ by using the best fit for each character among all MFCs in Henmg86, or by using an average value in PAUP. It means that m the final solution the weights will be stable only in relation to all trees at the same tune, but not necessarily in relation to the weights implied by each tree itself. Some or all of these trees may therefore be contradictory in the sense that each tree would “resolve character conflict in favor of exactly those characters the tree is telling us not to trust” (Goloboff 1993). Farris (1969) and Goloboff (1993) already noted that the initial set o f weights will affect the final solution. The main problem is that defending the final trees would require defbiding the initial weighting schemes, and this presents some practical difficulties. As pointed out by Harbach & Khching (1998): “errors will be introduced if heuristic methods are used for cladogram construction and the MPCs for the data (and weights) are not found, and unless ambiguous optimization is suppressed, weights could be based upon a set o f MPCs that mcludes spurious, over-resolved topologies.” A more basic difficulty was stressed by Carpenter (1988): multistate characters must be converted to additive binary coding before th ^ can be submitted to successive weighting. In Carpenteris words: “[tjhis is because higher integers may result in greater weight being given to a particular character shnply as a function of scoring whereas additive binary coding avoids this by evaluating each state of each character indwidually.” Goloboff (1993), however, proposed a non-iterative method for weighting characters which solves these problems. Characters are weighted according to the amount of homplasy implied for each of them in a given cladogram. The sum of the weights o f all characters gives the f i t of the cladogram, and the greatest fit is then used to select among 128 the dif&rent cladograms generated m a heuristic search. Weights are calculated accordmg to a concave Qiyperbolic) function of homopla^, as W=K/(K-ESi), where ESI is the number of extra steps implied for a character on a cladogram and K is the constant of concavity (reasons for not using the rescaled consistent mdex are «cplained by Goloboff 1993). Differences in the number of extra steps in characters that show more homoplasy are considered less influential, and these characters are thus accorded lower weight Note that Goloboff s method offers an additional improvement over previous weighting schemes; the possibility of objectively choosing the best supported cladogram(s) among all self-consistent cladograms that may be found. This is possible because different trees imply different weights, allowing self-KX>nsistent trees to be selected on the basis of their individual fit. The higher the overall sum o f weights, the more informative is the distribution of the characters in the three, so that the approach is not only objective, but also helps to prevent acceptance of results where part ofthe information had to be dismissed. A current limiting foctor with implied weights analyses, however, is that different int%er values for AT can be applied (m PIWE them range is 1-6) but there is not yet an objective way for deciding which value should be preferred (Goloboff 1995). Note, however, that higher values of AT will weigh less strongly against characters with homoplasy, thus makmg results tend towards those obtained firom equal weightmg. Lower values will down weight bomoplastic characters more strongly. It becomes obvious, therefor^ that the higher the level of homopla^, the smaller should be the value adopted for a :. 129 Di this anafysis, most characters were highly homoplasious, with observed numbers of extra steps rangmg from one to 24 (ch. 1 ^ . fri fact, even for searches using a JC value of 2 (also adopted, for example by Ronquist e t oL 1999) a large proportion of the ^napomorphies supporting clades o f mterest in the resulting cladogram still corresponded to highly homoplasious character states, appearing as synapomorphies five times or more in different parts o f the dadogram. It seemed obvious that the analysis could greatly benefit if the influence of homoplasious characters was down-weighted as much as possible within the limits of the program. Based on this rationale, results obtained from searches with ^ set to 1 were preferred over all the others. For the complete matrix, four MPCs were obtained when K was set to I, with fit 508.0. Total length varied slightly, with one cladogram with 1239 steps, two with 1240 steps, and one with 1241 steps; all had Cl 22 and RI 69 (rounded values). Differences m topology and characters supporting each clade were minimal, and unimportant for the objectives of this work, as they do not affect decisions at generic level. The strict consensus tree o f these cladograms (Fig. 198) was selected as the preferred cladogram. For the matrix restricted to foenatopmae taxa, three MPCs were obtained, with fit 725.5. One o f them has length 602, Cl 36, RI 61, and the other two have length 600, Cl 36 and RI 62. The strict consensus o fthese three cladograms is shown in Fig. 200. Clados and PIWE will show or list only unambiguous synapomorphies for each clade. Ambiguous characters, therefore, need to be optimized in order to determine whether th ^ will provide extra support for the clade or clades of interest Goloboff (1997) proposed an objective method for optimmng character^ based on unplied 130 Node 2 Mode 3 Hemiatephanus ParaatephantUus Xanthostephanus Foenatopua c h .II:0 -* I ch. 15:0-1 ch. 1:0—1 ch. 18:0-1 ch. 1:0—2 ch. 5:1-0 ch. 32: (M l ch. 1 7 :0 -3 ch. 5 :1 —0 ch. 24:1-2 ch. 63:0-1 ch. 1 7 :3 -2 ch. 37:0-^1 ch. 57:1-2 ch. 9:1—0 ch. 32:2-3 ch. 65:2-1 ch. 2 7 :0 -1 ch. 4 0 :0-»l ■ ch. 6 6 :0 -1 ch. 20:0—1 ch. 55:0-1 ch. 7 8 :1 -2 ch. 34:0—1 ch. 47: (M I ch. 68:0—1 ch. 25:0—2 ch. 8 7 :0 -1 ch. 8 9 :1 —0 ch. 7 1 :1 -2 ch. 57: (M l ch. 99:0—I ch. 2 7 :0 -1 ch. 101:1-0 ch. 92:01-2 ch. 76:0—1 ch. 65: (M2 ch. 101:0-1 ch. 32:1-0 ch. 115:0-1 ch. 93:3-2 ch. 88:0—1 ch. 7 1 :0 -1 ch. 111:0-1 ch. 33:0-1 ch. 147:0-1 ch. 9 6 :2 -1 ch. 102:1-2 ch. 86:0—1 ch. 133:0-1 du 39:0—1 optimized: ch. 97:0-1 ® ch. 103:0-2® ch. 93:0—2 ch. 139: (M l ch. 5 6 :0 -1 ch. 2:0-2 ch. 98:12-0 ch. 105:0-1 ch. 113:1-0 ch. 140:0—1 ch. 6 5 :2 -1 ch. 6:0—1 ch. 112:0-2® ch. 106:0-1 ch. 141:0-1 ch. 162:1-2 ch. 78:01-2 ch. 7:0—1 ch. 116:0-1 ch. 107:0-1 ch. 155:0-1 ch. 165:0-1 ch. 80:0—1 ch. 2 8 :1 -0 ch. 117:0-2 ch. 113:1-2 ch. 162:0-1 ch. 168:0-2 ch. 108:0-1 ch. 42:1-2 ch. 120:1-0 ch. 114:0—1 ch. 167:0-1 ch. 170:0-1 ch. Ill: 1-0 ch. 78:1-0 ch. 125:0-2 ch. 133:1-0 optimized: optimmed: ch. 117:0-1 ch. 96:2—0 ch. 130:2-1 ch. 136:0-1 ch. 3 :0 —1 ch. 150:0-1 ch. 121:0-1 ch. 109:0—1 ch. 145:0-3 ch. 149:0-1 ch. 44:0—1 ch. 159:0-1 ch. 140:1-2 ch. 150:1-2 ch. 156:0-1 ch. 127:0-1 ch. 160:0—1 ch. 145:0-3 Node 28 ch. 151:0—1 ch. 157: 1 -2 ch. 166:0-1 ch. 8 3 :0 -1 ch. 164:1-0 ch. 162:4-5 Stephanus Node 17 optimized: ch. 101:1-2 ch. 174:0-1 ch. 173:0—I ch. 3 :0 1 -2 ch. 17:12-3 ch. 31:0-1 ch. 141:1-2 optimized: optimized: ch. 5:0 —1 ch. 4 6 :1 -0 ch. 148:0-1 ch. 23:1-0 ch. 6:0—1 ch. 14:0—1 ch. 5 8 :1 -0 Node 21 ch. 157:0-1 ch. 42:1-2 ch. 7:0—1 ch. 24:0—1 ch. 5 9 :1 -0 ch. 3 2 :1 -2 ch. 170:1-2 ch. 44:2—1 ch. 19:0—2 ch. 2 6 :0 -1 ch. 69:0-1 ch. 50:0—1 optimized: ch. 6 4 :1 -0 ch. 2 8 :1 -0 ch. 2 9 :0 -1 ch. 9 3 :2 -3 ch. 70:0—1 ch. 6 6 :1 -0 ch. 7 2 :1 -0 ch. 40:1—2 ch. 46:0—1 ch. 122:0—2 ch. 77:0-1 ch. 6 7 :0 -1 ch. 8 5 :1 -0 ch. 5 0 :1 -2 ch. 60:0—1 ch. 123:0-1 ch. 8 6 :2 -1 ch. 80:0—2 ch. 146:1-0 ch. 72:1-2 ch. 67:0—1 ch. 130:1-2 ch. 94:0-2® ch. 92:0—1 ch. 159:1-0 ch. 90:0-1 ch. 9 3 :1 -2 ch. 154:0-2 ch. 102:0-1 ch. 153:0-1 ch. 94:2-3 ® ch. 97:0-2® ch. 162:2-4 ch. 113:0-1 ch. 97:0-2® ch. 112:0-1® optimized: ch. 138: (M l ch. 135: 1 -0 ch. 138:0-1 ch. 2 3 :0 -1 ch. 175:0-1 ch. 141:1-2 ch. 8 9 :0 -1 optimized: ch. 154:0-3 ch. 100:0—1 ch. 44:1—2 optimized: ch. 129:0-1 ch. 64:0—1 ch. 2:0—1 ch. 135:0—1 ch. 85:0-1 ch. 3 :1 -2 ch. 146:0-1 ch. 6 :0 —1 ch. 160:1-2 ch. 130:0-1 ch. 175:0-1 Table 1.4. Character state changes on shortest cladograms obtamed with K, in PIWE, set to I. Nodes correspond to those in Fig. 198. Changes 6 r terminal taxa (except Xanthostephanus n. gen.) and for clades within Foenatopus (node 29) are not listed because th ^ are not o f direct interest. Renumbered m matrix: I-^O, ® 1-^2, ® 0-> I. 131 transformation costs. However, while essentially a derivation of the principles o f implied weights parshnony, this new method represents in foct a distmct approach for tree searching which I did not have the chance to use and evaluate until the present work was practically completed. Because of this, the method of implied costs was not considered for the present work. Optimizations were performed instead aposrerib/7, by means of argumentation based on the evaluation of each character separately. Character changes for the ^napomorphies supporting clades of mterest are listed in Table 1.4. Monophyty and Classification of Stephanid Taxa Monaphyty o f the Stephanidae As explained in the item “Selection of outgroups,” non-stephanid taxa were not included in the analyses o f the mtemal phylogei^ o fthe fomEy. The foUowmg discussion is therrfore based mostly, albeit not entirely, m literature data. The monophyly o f the fomily Stephanidae is currently supported by the followmg synapomorphies: 1. Ocellar corona with a ring of cuticular teeth encircling the median ocellus (Rasnitsyn 1988), 2. Presence of large apodemes contmuous with tormae O ^elm sen 1996), 3. Presence of an sclerite for the retractor of the epiphaiynx (VOhelmensen 1996), 4. Paraglossae reduced distally (Vilhelmsen 1996), S. Labial palp with four s%ments (Vilhelmsen 1996), 6. Ventral wall ofupper ovipositor valve strongly thickened medially (Quickee t aL 1994), 7. Presence of Garge) tubercles on the ventral 132 margin of the hind femur ^ônigsmann 1978), 8. The lateral displacement of the mandibular bases (\HheImsen 1997). All these features, accept number 8, are also found mdependently in other Hymenoptera, and therefore seem to represent parallelisms. O f the above synapomorphies, perhaps the most characteristic is the presence of the coronal area and tubercles. The fact that th ^ occur in orussids as well has long been noticed (e.g., Ashmead 1900b), and it has been commonly used to suggest that stephanids and orussids are closely related. However, it is important to notice that the structure of these tubercles is different in each femily: stephanids have an odd number of tubercles, with a posterior and a central pair, and a single anterior tubercle placed medially, or S total, bi orussids, there are 6 tubercles, organized in three aligned pairs. This difference helps to support the idea, implied by the cladistic analyses in Vilhelmsen (1997) and Ronquist e t aL (1999), that the presence of tubercles are independent (parallel) ^napomorphies for each fomily. The réévaluation of literature data on Hymenoptera phylogeny performed for the present work (see item “Relationships of Stephanidae with other Hymenoptera”), could, in principle suggest a different set of synapomorphies for stephanids. Nonetheless, this investigation focused on a comparative analysis o f several cladograms obtained under diffaent approches, and no attempts were made to select or defend a single preferred cladogram. Consequently, an objective list o f synapomorphies for stephanids could not be derived. It might be of interest to report, however, that in the mentioned analysis the presence o f coronal tubercles was the only character invariably retrieved as a synapomorphy for stephanids in cladograms obtained with any method or parameter. 133 The following characters also seem to be typical for stephanids. Thty were not thoroughly compared with other Hymenoptera, but are proposed here as possible tynapomorphies for thû fomily: (I) postgenal bridge folly developed (i.e., postgenae completely fosed); (2) a wide; projectm^ “blade-like” Itypostomal carina; (3) presence o f apical hairs on anterior margin of pronotum, laterally; (4) front fomur basally compressed and somewhat twisted; (S) costal cell slightly more sclerotized than rest of wing membrane, and rolled or folded upwards (perhaps to compensate structural problems due to the absence of a costal vein); (6) costal cell, and other basal cells of front win& without microtrichia; (7) vein 2Cua at 90° or less with ICu (apparently less than 90° in other Hymenoptera); (8) vein (Rs+MQb distinct; (9) a bulla between veins 2+3Rs and/or 2+3M and apex of ^sfh ^ b ; (10) propodeal spiracles placed near petiolar forame; (11) wing membrane distinctly concave at meeting pomt o f veins 2+3Rs, 2+3M, and (Rs+M)b; (12) line of fosion between T2/S2 (petiole) visible at least basally on segment, or T2/S2 completely isolated (Schlettererius^ (13) presence of specialized setae on M-K^u, I-I A, and S(H-R; (14) a sharp longitudinal edge ventrally on basal half of hind tibia; (IS) brush of paddle shaped hafrs on hind tibia (mostly on dilated part) and tarsus; (16) margins of abdominal stemites deeply emargmate centrally, as a (17) tubercles on coronal area organized m a pentagon; (18) exceptionally large vertex; (19) occipital carma folly and conspicuously developed (varies apîcally only); (20) a slightly twisted articulation between mid fomur and tibia: the tibia closes onto the ventro-Iateral side o f the fomur, rather than dfrectly on its ventral side; (21) presence o f a parastigma (also on Trigonalidae). 134 Classification in Subfamilies The consideratioii of subfiunilies for stephanids is an almost inevitable concept when the worid âuna is compared. This is because the âmify is composed by two very distinct morphological ^ e ^ each represented by numerous species, and both readily recognized by dramatic differences in wing venation (e.g.. Figs 163-178), as well as in many other characters. The division in subâmilies, however, can also be useful in providing a meaningful and reasonably stable reference for main evolutionary lineages within the femily, each of which are taxonomically fer less stable internally. The breadth and definition of Foenatopinae or Stephaninae, fer example, is essentially the same throughout the taxonomic literature, and seems unlikely to change dramatically in the future. Taxa within these groups, however, have been and continue to be interpreted in a variety o f different ways. Three subfemilies have been proposed fer the Stephanidae, but only two are adopted here. Thty are discussed below. Subfamity Stephaninae Endertein Di spite o f the feet that Parastephanellus was retrieved here as the sister group of Foenatopinae its mclusion in Stephanmae as originally suggested by Enderlein (1905), was mamtamed fer a number of practical reasons: (1) it contributes to maintain taxonomic stability, preventmg major changes m the mterpretation of Stephanmae and Foenatopinae; (2) albeit equivalence of Lirmean taxa with monophyletic groups is deshable, there is no consensus or adopted conventions about the necessity o f feUowing this strictly, nor is it 135 always practical (see n&ct point); (3) a rigidly monophyletic interpretation o f subfamilies for stephanids, if based in any of the cladograms obtained here, would force the proposition o f numerous subâmilies, or the disconsideration of them all; (4) the synapomorphies for Foenatopinae without faras&pAongAts are much more stable and easier to interpret and visualize than those defining a monophyletic subfomily including this genus (see also discussion for node 21); and (5) past connections between Australia and South America, the main areas of occurrence îqx Pcarastephanettus and HemistephanuSy suggest that interpretations for the phylogenetic relationships between these two genera can not yet be considered as “settled.” More accurate conclusions, however, would depend on biogeographic and molecular analyses, both of which are beyond the scope of the present work (see also discussion îor Hemistephanus). The considerable phylogenetic distance between Schlettererius and the remaining stephanids (see discussion for this genus) seems to fiivor the idea of a separate subfamily, Schlettereriinae, as proposed by Orfila (1949). Once more, however, such subdivision does not seem adequate at the moment First because such subfiun3y is of little practical value, since it would encompass a single genus and only two species; and second because its adoption implies that other highly distinct taxa, such as Xanthostephanus n. gen., would also have to be placed in its own subfomily, a decision which would agam contribute to decrease taxonomic stabili^ and thus the overall usefiilness of the classffîcation. Because o fthe above reasons, Stephanmae Enderiem is mterpreted here m its origmal sens^ that is, including Schlettererius Ashmead (by implication, since this genus 136 was proposed after Stephaninae) and PamsliepAane/Tus Enderiem, Not^ however, that because Stephanmae is a paraphyletic group, it can be defined phylogeneticaly only in terms of the absence o f derived states which support Foenatopmae (see discussion for this sub&mily). For a diagnosis o f the Stephaninae, see item to SubâmOies and Genera,” Genus Schlettererius Ashmead The identification of synapomorphies & r the genus Schlettererius would in 6ct depend on a &mily-level investigation, since this taxon seems to represent the most basal extant genus of Stephanidae, as already discussed. For this reason, Schlettererius will be discussed here only with respect to its phylogenetic distance fi-om other stephanids, A total o f 18 ^napomorphies separate all other stephanids firom Schlettererius (node 2), Of these, 15 were recovered as unambiguous; flageOomeres with sensillae d ire n t from Fig, 12 (ch, 11:0-^I, unique); gena with a yellow spot basally, at or near malar space (ch, 32:0-^1, one full reversal for Hemistepharms)', maxillary palpus distinctly longer than head is laterally high (ch, 37:0-»l, two independent reversals); labrum lightly colored ventrally, darkened dorsally (ch. 40:0-^1, one mdependent reversa^; colo shape normal, about 0,5-1,0 as long as wide (ch, 47:0-+1, unique); preannular perfectly continuous with semiannular (ch, 57:0-^1, one mdependent occurrence); ventral area of pronotum glabrous or almost so (ch, 65:0-+2, unique); vellum o f fi'ont tibial spur narrowing apically, disappearing before reaching the ap«c (ch, 71: 0-*^!, unique); vein IM slight^ arched (ch, 86:0-+1, one mdependent reversal); vein 2-1A shortly tubular basally, then mostly or entmely nebulous (ch, 93:0->2, unique); vem 2+3M 80% or more (but 137 never entirely) tubular, ^ e x nebulous or spectral (ch. 113: l-»0, one reversal); spiracular impression and carina independent of Ime of fiision between metapleuron and propodeum; both straight,- ending on or pomting to flank, or spnacular impression curving inwards apically; spiracular carina somethnes not developed (ch. 141:0-*^I, unique); hind coxa without an apical-dorsal spine (ch. 155:0->l, unique); hind tibia structure highly specialized, n o t as in Fig. 133 (ch. 162:0-+1-5, unique); T2 and S2 fused into a cylmdric and elongate petiole (ch. 167: (M l, unique). Three additional ^mapomorphies resulted fi'om character optimization: most elongate flagellomere very long and narrow, 9.5-l6.0x or more as long as wide (ch. 3 : 0-*^l, one reversal); apical emargination of pronotum distinctly and strongly emarginate, C-shapped (ch. 44:0-*^l, one independent reversal); vein Cua on hind wing indicated by a stub coming out of M+Cu (ch. 127:0-+1, two independent reversals). This large set of weakly homoplasious synapomorphies not only indicates that Schlettererius is very distinct fi'om other stephanids but also suggests that it is distantly related to other octant species o f the fiimily. Note also that this phylogenetic distance is reinforced by 18 synapomorphies leading to node 3, which make Schlettererius distinct form all other extant stephanid species, except only Stephanus serrator, by a total of 33 synapomorphies (those at nodes 2 and 3). There has been virtually no disagreement about the delimitation and validity o f Schlettererius smce it was proposed by Ashmead (1900b) for a single species, Stephanus cinctipes Cresson. Indeed, Schlettererius is morphologically very distmct fi'om all other 138 stephanid^ and the taxonomic consistent^ of its unique features can be confirmed by their occurrence in a second, recently discovered species, S. determmatoris Madl, fi'om Korea O ^dl 1991). GenusStephanus Jurine The monophyly of this genus is supported by 14 apomorphies. O f these, 9 were recovered as unambiguous: most elongate flagellomere moderately elongate (ch. 3; l-»2, feur other occurrences, including as a synapomorphy fer clade at node 30); flagellomere 2 distinctly shorter than flagellomere 3 (ch. 5 :0-+1, highly homoplasious); sculpture on dorsal half of fi'ons in mixed directions, or non-directional (eg., areolate) (ch. 14:0-*^!, highly homoplasious); vertex sculpture partially transverse, except laterally, or at least antero-laterally, where any other orientation, except transverse or areolate^rugose, occur (ch. 24:0-^l, highly homoplasious); intensity of sculpture on vertex normal or fine (ch. 26:0-^l, highly homoplasious); gena and temple with conspicuous sculpture (ch. 29: 0-»l, highly homoplasious); individual lateral hairs on colo present (Fig. 63 ) (ch. 46: 0-^1, one more occurrence); presence of a dorsal impression centrally between preannular and semiannular (ch. 60:0-*^l, three other occurrences, mcluding as a ^napomorphy for clade at node 31); pronotal spiracle «posed (ch. 67:0-+1, four other occurrences, inclusmg as a ^napomorphy fer Foenatopmae); vein 2-lA shortly tubular basally, remaming mostly or entirely nebulous (ch. 9 3 :1-+2, highly homoplasious); angle between veut 2Cua and ICu obtuse (ch. 97:0-+2 [renumbered], highly homoplasious); interception point of vems 4Rs and 2-^3M with wmg margm equidistant (or neariy so) to wing ap « 139 (ch. 112; O-^I [renumberecQ, highfy homoplasious); area in between mid coxa and mesepistemum glabrous or nearly so (ch. 138:0-»l, also a synapomorphy for clade at node 21); spiracular impression distinctly dK&rentiated, well delimited, concave; without transverse carina^ or carinae incomplete and m small number (ch. 141:1-+2, highly homoplasious); hind coxa with non-transverse microsculpture, or polished smooth (ch. 154; (M3, highly homoplasious). Five additional synapomorphies resulted from character optimization; first 8 basal fiagellomeres more elongate than following ones (some or all of basal ones about 5x as long as wide; apical ones about 3x as long as wide) (ch. 2; 0-^1, highly homoplasious); most elongate flagellomere moderately elongate, 5.6-9.0x as long as wide (ch. 3; 1-+2, highly homoplasious); flagellomere 2 with a few or several Type I sensillae restricted to apical 0.4-0.7, portion of flagellomere with sensillae usually somewhat dilated (ch. 6; O-^I, highly homoplasious); basal shape of hind wing normal or weakly concave posteriorly, wing distinctly not pedunculate (ch. 130; 0->l, highly homoplasious); ovipositor sheaths unicolorous, or progressively darker towards apex (ch. 175; 0-+1, highly homoplasious). Albeit most apomorphies listed above are highly homoplasious, their signffîcance and the monotypic interpretation of Stephanus can be further supported by the long series o f synapomorphies isolating Stephanus serrator from Schlettererius (node 2) and fi'om the remaining stephanids (node 3). The genus Stephanus was originally proposed to accommodate a single European species Qurine 1800). A few other taxa were later incorporated mit, but they were large tropical specie^ considerably distinct flom the genotype Stephanus serrator (P.). BruHe 140 (184Q used these differences to isolate S. serrator as the only valid species o f Stephanus, and to propose to accommodate all the other. Since then, opinions on the delimitation o fthe genus Stephanus varied enormously, ranging from the m ono^ic concept o f Jurine and Brullé, to a genus containing about one-third of the stephanid species (e g., Elliott 1922), to the only valid genus in Stephanidae (Schletterer 1889a). The lack o f a precise definition fi)r this genus lead to a situation where numerous and diverse stephanid species were described as, or later transferred to or fi'om Stephanus. This situation is well illustrated by the fiict that aU stephanid genera ever proposed have had or still have species with junior synonyms which were originally described as members ot Stephanus. The adoption o fStephanus s. 1. persisted in the literature until very recently (e.g., Medler 1980, De Santis 1980), but most contemporary authors recognize it as monotypic (eg., Oehlke 1984, Pagliano 1986, Madl 1991, Tobias 1993, Scaramozzino 1995), an opmion which is in agreement with the results obtained here. It must be noted, however, that the main features used to justify Stephanus s. s. were the occurrence o f three femoral teeth in the hind leg (ch. 160:0), and the presence o f five tarsomeres in the hind leg of females (ch. 165:0), which are both plesiomorphic with Schlettererius. Genus Megischus BniDé (Nodes 3-17) Monophyfy M ost authors which recognize AfegMcAus as a valid genus adopt a sensu lato interpretation, including m ît the genus Hemistepharms Enderlein, and considering 141 Stephanus Jurine, with which is often ^onymized, as monotypic (e g., Townes 1949, Oifila 1956 & I960, Gauld 1995). An aftemative interpretation isolates Hemisiepfumus as a valid genus, and includes the remaining species as part of Stephanus (e g., Enderiein 1906, Orfila 1956). And finaly there is a sensu stricto interpretation, which is approximately equivalent to a combination o f the two previous approaches: it excludes &omMeffschus the monotypic Stephanus and all Hemistephanus spp. Periiaps surprismgly, this last interpretation has not appeared in the literature except until very recently, m Aguiar (1998). The results obtamed in all stages o f the present study, however, invariably indicated that none of the above interpretations o fMegischus correspond to a monophyletic concept, but are instead equivalent to groups containing several independent clades. This can be easily observed in the cladograms illustrated in Figs 195-198. Results in the preferred cladogram (Rg. 198), however, show that all clades and species of M egischus sensu stricto appear adjacent to one another m the preferred cladogram, usually with a small number o f synapomorphies separating one clade fi’om another. Clade support also greatly fevors a sensu stricto interpretation, being maximum where M egischus s. s. connects with Stephanus and Hemistephanus^ but showmg low values mtemally. That i^ although paraphyletic, M egischus s. s. seem to represent a distinct lineage within Stephanidae, placed between nodes 3 and 17, apical to Stephanus and basal to Hemistephanus (Rg. 198). Albeit not ideal, the present consideration of a paraphyletic remains taxonomically sound, smce it represents a well delimited group o f species. A strict 142 monophyletic interpretation o fMegischus would greatly limit its definition, retricting it to the clade at node 10, ânce it is the one with the genotype species. Although not a problem p e r se, this would lead to drastic taxonomic changes, in order to accomodate the several clades which would be left unnamed. An additional problem with such decision is that many of the clades between nodes IS and 3 are highly unstable (see Bremer support values in Fig. 198). In fiict, most clades vntimMegischus as defined here proved to be highly variable along the development o f this study, often changing considerably after each of the many revisions and re-runs of the data matrix. This was not true for clades which define other genera. Possibility o f generic statusfo r M. anomalipes+M, califomicus-^M. texanus For searches with K set to 2 or more in PIWE, and in all searches with NONA (both alone and with aid of Ratchet), the PalearticM anomalipes, and the NearcticM califomicus and M texanus appear invariably together as a clade in the resulting cladograms (as m Figs 195 and 196). This fact is contradictory with the arrangement suggested by the preferred cladogram, which associates M brunneus with M anom alipes in one clade, andM texanusanomalipes m another, the two placed considerably appart ftom each other ^ ig . 198). Phylogenetic relationships for this group of species as suggested by the preferred cladogram are intuitively dffîcult to accept, because of strong morphological dissimilarities between the associated taxa. This is indicated, for example^ by the long branches leadmg to M, califomicus and to M texanus in the correspondent 143 clades. When combined, these 6 c ts seem to support the idea o f a clade w ithM anom alipes + M califomicus+ M texcams, thus suggesting they could be assigned separate generic status. Nonetheless, the Act that this group is not supported in the preferred cladogram cannot be ignored. Note, for example, that the clade withM califomicusbrunneus has maximum branch support (Fig. 198), indicating that the synapomorphies supporting it are characteristic enough 0.e., non-homoplasious enough) to prevent other combinations even in grossily sub-optimal cladograms. The disconsideration of the results &om the preferred caldogram would also, of course, require the refutation of the arguments discussed m the item “Phylogenetic Analysis.” Therefore, a conservative approach Q.e., maintaining these taxa as part aîMegischus) seems more reasonable at the moment. Delimitation As interpreted here, M egischus can be easily delimited morphologically, on the basis of inclusive and exclusive character states (eg., those at nodes 3 and 17). However, the group would be more strongly supported if its species shared unique character states. I searched fer such features by examining the synapomorphies at node 3 individually, in Clados. Even though character changes occurring at this node always reverse or progress to a more derived state somewhere else in the cladogram, it almost never happens precisely at node 17. There is a single exception, however character 162 (structure of the hind tibia), fer which there is a 1-^2 change at node 3, then a 2-*^3 change withm M egfxhuSy then a 3->4 change at node 17, with no other occurrences fer states 2 and 3 in the cladogram. States 2 and 3, therefore appear to be unique to M egischus. 144 There are 18 synapomorphies at node 3, isolatmg from Schlettererius and Stepfum us. Ofthese; 15 were recovered as unambiguous: as follows: frons dorsally mostly or entirely areolate orareolate-rugose (ch. 15:0-+2, four other occurrences, mcluding as ^napomorphy for Hemistephanus and for clade at node 42); ventral half of frons strigate or carinate, regular or weakly ondulate (ch, 17:0-^3, unique); preannular isolated from semiannular, by a step or angle, or simply by a more swollen and difrerently sculptured semiannular (ch. 57: l-»2, highly homoplasious); prepectus mostly or entirely covered by postero-ventral apex of pronotum (ch. 66:0-^1, highly homoplasious among non-foenatopmae); posterior margin o f semiannular in dorsal view arched (ch. 68:0-*^ 1, unique); vein 2Cub simple, not forked at apex (ch. 99:0-»l, four independent reversals); vein RI tubular, or at least densely nebulous b^ond its basal half (ch. 101:0-^1, many reversals); veins 2+3Rs and 4Rs aligned, even if 2+3Rs sinuous (ch. Ill: O-^l, many reversals); hairs on mesepistemum long and conspicuous, decumbent and distinctly overlapping or erect, always distinct from pruinosity (tf present) (ch. 133:0-+1, many reversals); pflosity^ related to areolation/foveolation on propodeum absent (ch. 139: 0-^1, highly homoplasious); sphacular impression ill-defined, or very shallow and with numerous transverse carinae (ch. 140:0-*^l, one independent reversal); hind tibia structure as in Figs 135 or 137 (ch. 162:1-+2, unique); female hmd tarsus with three tarsomeres (ch. 165:0-*^l, unique); petiole two times o r more as long as dorsal lenght o f T3 (ch. 168: 0-+2, unique); petiole dorsally transversely strigat^ but not very finely so (ch. 170: O-^l, one mdependent reversal). 145 Three additional synapomorpbies resulted from character optimization: hind coxa as long as, or slightly longer than maximum lateral length of mesepistemum (up to l.lx as long) (ch. ISO: 0-»l, several independent reversals); hind Amur without a mesal conq>ression, uniformly curved in cross section (ch. 159:0-+I, many reversals); bind femur without teeth or tubercles o f any kind basad o f median tooth (ch. 160:0-*^!, unique). For ^napomorphies which exclude A /egfsc^ from the remaining stephanids see below. Hemstephanus+Parastephanellus + Foenatopinae (Node 17) This group is supported by 16 synapomorpbies, of which the following 11 were recovered as unambiguous: ventral half of frons strigate or carinate, regular or weakly ondulate (ch. 17:12-+3, highly homoplasious); individual lateral hairs on colo absent (ch. 4 6 :1-+0, highly homoplasious); hairs on preannular and on semiannular inclined backwards (ch. 58: l-»0 and ch. 59: l-»0, both unique, but state 0 occurs for most basal species); angle between apical margins of hemistemites of prostemum nearly 90' or more (ch. 69:0-+1, two other occurrences); vein 2-1A a small tubular stub, which sometimes is briefly nebulous apically; remainmg absent or spectral; or entirely spectral (ch. 93:2-*^3, three independent reversals); vein 2cu-a spectral or absent (ch. 100:0-*^l, two other occurrence^ one independent reversal); vem M absent on hind wmg (ch. 122:0-+2, unique); hind wing vem Rs nd)ulous or spectral; short, originating near or at apex of Sc+R; usually curving towards anterior margin, more rarely straight or somewhat undulate 146 (ch. 123; unique); posterior wmg fold on hind wing absent (ch. 1 2 9 : three mdependent occurrences); hmd wing basally very narrow, for a short distance approximately as narrow as or narrower thaa width of Sc^R, making the wing distinctly pedunculate (ch. 130:1-^2, two independent reversals); hind coxa regulariy transverse, sculpture intensity fine or very fine (ch. 154:0->2, one independent occurrence); hind tibia structure as m Fig. 136 (ch. 162:2-»^4, unique). Five additional tymqx>morphies resulted fi’om character optimization: intreocellar carinae placed mostly anterior to or in between posterior ocelli (ch. 23:0-*-1, one independent occurrence, two independent reversals); vein Icu-a inclined towards wing bas^ forming an angle of little less than 90° with M+Cu, and usually a correspondent slightly obtuse angle with ICu (ch. 89:0-^1, three other occurrences, one independent reversal); vein 2cu-a spectral or absent (ch. 100:0-+1, two mdependent occurrences, one independent reversal); posterior wing fold on hind wing absent (ch. 129: 0-^1, three independent occurrences); pruinosity on mesepistemum (except pectus and dorsal part), occurring on 90-100% of its area, moderately to densely abundant, concealing 50-95% of its cuticular surfitce (ch. 135:0-*^l, two independent occurrences, many reversals). The mam importance of the present clade lies in that it distinctly isolates Hemistephanus ûomM effschus s. s., first by unequivocal^ placing Hemistephanus apical to all MegischuSy and second by suggesting that Hemistephanus is the sister group of Parastephanellus + Foenatopinae, instead o f any taxa of or closely related to M egischus. This is an interesting result because it contradicts the generalized assumption that Parastephanellus and Hemistephanus are closely related taxa. In foct, it is probably 147 because o f this idea that there has been very little speculation or discussions about the relationships between these two genera, and virtually no discussion between their relationships with Foenatopinae. Nonetheless, the Act that Hemistephanus is mostly restricted to South America (with a Aw spp. in Central America), that Parastephanellus is most speciose in Australia, and that these two contments have been connected in the past, suggest that the phylogenetic relationships between these two taxa need be investigated further. This, however, would requhe the use of biogeographic and molecular methods, which are b^ond the scope of the present work. For relationships between Parastephanellus and Foenatopinae see discussion for node 21. Hemistqfhanus Enderiein ^ o d e 18) The clade which de&ies this genus is supported by 21 synapomorpbies, of which the fbllowmg 20 were recovered as unambiguous: head red or ferrugineous (ch. 1:0-*^!, several independent occurrences, except also a synapomorphy for clade at node 4); flageOomere 2 about as long (0.9-1.0) as flagellomere 3 (ch. S: 1-^0, highly homoplasious); flagellomeres 1-3 without microdepressions on ventral Ace (ch. 9:1-^0, highly homoplasious); Ace pilosity m between antennal foramina, until clypeal suture very abundant, hairs usually much shorter than on firons (ch. 20:0-*^ I, unique); vertec sculpture mostly or enthely parallel ondulate-rugose or rugose m a zig-zag pattern (ch. 25:0->2, unique but with reversal to 0 inÆlimpidipemns)\ anterior half of vertec with granular 148 microstculpture m between macrosculpture (ch. 27:0-> 1, three other occurrences, ipcfuding as a synapomorphy forFoenatopus); no yellow colors on gena (ch. 32: l-^O, two mdependent 2-^0 reversals); occipital carina in lateral view bent midway; dorsal and ventral portions nearly straight (ch. 33: (H-1, three other occurrences); dorso-mesal tooth of mandible present, small or stout and distinctly detached (ch. 39: (M l, highly homoplasious); preannular with a transverse ruga and groove (ch. 56:0-+1, five independent occurrences); ventral area o f pronotum with &ie dense püosi^ basally, becoming more sparse apically until glabrous fiom midway on (ch. 65:2-^1, several independent occurrences); setae on M+Cu placed 6 r fiom IM, fiom which th ^ are separated by a distance greater than length of Icu-a (ch. 78: M 2, three independent occurrences); parastigma shorter than IRs, but still longer than wide (ch. 80: 0-» 1, three independent occurrences); smooth transition fiom 2r to 4Rs, these veins perfectly contmuous, never formmg an angle (ch. 108:0-+I, two independent occurrences); veins 2+3Rs and 4Rs angled (ch. Ill: MO, three independent occurrences); retmaculum placed midway between cell ICu and cell 2Cu; its apex approximately equidistant fiom veins Icu- a and 2Cua (ch. 117:0->I, also a qmapomorphy for clade at node 31); hind wing Sc+R apically slightly bent towards anterior wmg margm (ch. 121:0-^I, four other occurrences); sptracular impression distinctly dK&rentiated, well delimited, concave; without transverse carmae, or carinae incomplete and m small number (ch. 140: M 2, four mdependent occurrences); short pilosity on metapleuron very densely abundant, blocking 50-100% o f pilose surface dorsal area completely taken by short pQosity, which reaches 149 sptracular carina (ch. 145: (M3, one mdependent occurrence); hind basitarsus ventrally with 1-2 rows of short, opaque conical spines, which get smaller and less distinct toward apex (ch. 166: (M l, unique). One additional synapomorphy resulted from character optimization: gena behmd malar space with a well defined patch of hairs, but hairs sparse and sometimes minute; remamingofgenaglabrous (ch. 31:0->1, highly homoplasious, revaW to 0 m H . e lm a tu s and H. limpidipennisy The genus Hemistephanus has been most often interpreted either as a valid taxon or, neariy with the same fiequency, as a Junior synonym îo t M egischus. Albeit such opinions were repeatedly presented without formal justification, two problems are likely to have played a role in generating the lack o f consensus: the overall morphological similarity between these two genera, and doubts about the taxonomic value of the diagnostic characters traditionally used to separate them. The single most important feature generally used to characterize Hemistephanus is the great reduction or absence of vein 2-1A (ch. 93), which leaves cell ICu widely open posteriorly. This character was originally proposed by Enderiein (1906) to separate both Hemistephanus and Parastephanellus fi’om other stephanids, but until very recently it represented the only effective diagnostic difference between Hemistephanus and M egischus. The only discussion expressing concerns about using vein 2-1A as a diagnostic character was published by Roman (1917), who pomted out that its development shows some variation, and was completely developed in at least one (supposed) Hentistephamts 150 specimen he observed. Roman also noted ibstM effschtts texanus (Cresson) has a very weak, and sometimes almost entirety “absent” vein 2-1 A, Because o f this, Roman consideredM texanus as a member o f Hemistephanus^ but maintained that this species had “strong afBnities m th Stephanas s. I ” (which included Megùc/tns). The degree of sclerotinazation of vein 2-lA, however, varies highly among stephanids, often making the decision about “presence or absence” considerably subjective. This character was given particular attention and completely reevaluated in the present work (see ch. 93). According to this reinterpretation no ovelapping was observed; vaMegischus vem 2-lA is always complete, nearly reaching apex of 2Cua, albeit sometimes entftely weakly nebulous (or sometimes mostly spectral inM. texanus, but these are rare exceptions for the species), while in Hemistephanus it never reaches beyond the basal 0.3 of cell ICu, usually remaining restricted to basal 0.1, or is absent. Character 93 ftiOs to hold as a synapomorphy for Hemistephanus alone, but has been useftil in supporting the clade Hemistephanus+Parastephanellus + Foenatopinae, discussed above. Enderiein (1906) proposed three diagnostic characters to separate Hemistephanus from Parastepfumeilus, and these were considered here as chs. 81 (shape of pterostigma), 85 ^oint o f origin of vein IKQ and 115 (relative size of cells RI and IM). Character 115 wOl be discussed imdtr Parastephanellus, which has the derived state. An elongate and sharply pointed pterostigma, the condition observed for Henristephanus, was retrieved as apomorphic among stephanids, but it is widespread in the ftunEy, occurring in several Parastephanellus as welL It has little diagnosric valu^ and 151 does not represent a synapomorphy for Hemistephanus. The point of origin of vein IM was originally interpreted by Enderiein as cell IM (Enderiein’s Cubitalzelle) either appendiculate or bluntly fused to cell ICu(^ubmedianzeUe). As interpreted here, however, the derived state (ch. 81:1) is shared hyhoÙL Hemistephanus and Parastepfumeilus^ and by many other taxa as well, with some independent reversals for some species. Very little testable information was published to further support or refute the validity o îHemistephanus. Roman (1917), however, noted that the species in this genus seemed to have a smaller overall size HamMegischus spp., implying that the différence could be diagnostic between these two genera. Elliott (1922) cited Roman’s observation in his revision, but was unable to decide about its taxonomic value. It was also investigated by Aguiar (1998), who used it as an auxdiar character to separate stephanid genera occurring in South America. However, several species o f Megischus from areas other than South America match well the typical body size for Hernistepfumus^ and this measure also overlaps greatly among other genera. Since it offered little promise as an informative character m a comprehensive evaluation, it was not investigated further for this work. The occurrence o f a clear hyaline or evenly infüscate wing membrane, and of a red head contrasting with a daric body were mentioned by Elliott (1922) as the “general rule” far Hemistepfumus spp. While many species in this genus indeed show a unusually clear wing membrane, its degree ofinfuscation in stephanids is highly variable and very difScult to categorize. For this reason it was not considered for this work. A red head occurs with a much M ^er frequency and is usually more brightly colored m Hemistepfumus than 152 among other stephanid genera. It was retrieved here as a synapomorphy for this genus, but there are several independent occurrences o f a read head for other stephanids. There are also some Hemistephanus species which are entirely dark brown or brown (Aguiar 1998). Aguiar (1998) was the only other author to provide information for discussions about the taxonomic status o fHemistephanus, mainly by proposing new characters to support it as a valid taxon (considered in the present woric as chs. 56,78, 110, and 166). Of these, chs. 56,78, and 166 were confirmed to be synapomorpbies îor Henristephanus. Character 110 has some diagnostic value, but is not a synapomorphy for any clade containing Hemistepfumus spp. investigated here. Parastephanellus + Foenatopinae (Node 21) This group Is supported by 15 ^apomorphies, of which the following 10 were recovered as unambiguous: gena with yellow or ivory maks or a stripe (ch. 32: l->2, one mdependent occurrence; state 3 for all Parastepfumeilus)', carmae on colo mostly complete, organized in a U-pattem, so that th^r are oblique towards pronotal fold when seen in lateral view (ch. 50:0-+1, two isolated reversals, state 1 with several fiuther specializations); front coxa without foveolae laterally (ch. 70:0-*l, two reversals); cell C on front wing without or with a few, microscopic, inconspicuous setae (ch. 77:0-*^ 1, unique); vein IM slightly arched (ch. 86:2-» I, highly homoplasious); vein 2Cua tubular on basal 50% or less; apically nebulous, even ^ strongly and cppearing to be tubular (ch. 94:0-^2 [renumbered], unique, but with 2->3 change at node 31); veins ^W-M)b and 2+3Rs cormected and contiguous with one another (ch. 102:0-*^l, one independent 153 occurrence, state 2tor Foenatopus); vrâi 2+3M tubular approximately on basal 50%, or less, remaining nebulous or specttal (ch. 113:0->l, unique, with one independent reversal, state 2 for Foenatopus); mid tibia anterioly with a fow spmes or spine-like setae along anterior foce, inclined toward apex (ch. 138:0-^1, one mdependent occurrence); met^leuron hahs associated with areolation from minute up to about 2x as long as diameter of areola where it origmates; ovipositor sheaths unicolorous, or progressively daricer towards apex (ch. 175:0-» I, four other occurrences, many reversals). Five additional synapomorpbies resulted from character optimization: apical emargination of pronotum as deep as or deeper than wid^ more or less U-shaped, but margins of emargination divergmg towards apex (ch. 44: l-»>2, one mdependent reversal); femoral sulcus pronotum) inconspicuous or not differentiated (ch. 64:0-*l, three independent occurrences, several independent reversals); vein IM originating apical to Icu-a, so that nodus formed by fusion ofM+Cu/lM with Icu-a/lCu is oblique toward wmg apex (ch. 85:0-^1, two other occurrences, several reversals); post-foveolar area not aligned with, and not as a continuation o f interfoveolar area, or interfoveolar area deep, contiguous with mettqileural fovea, with post-foveolar area standing in a higher plane than interfoveolar area (ch. 146:0-*^l, one independent occurrence, two independent reversals); hmd femur with two or more teeth or several small tooth-like tubercles basad of median tooth, or with small regular tubercle^ as the ones in between median and apical teeth (ch. 160: l-*^2, three independent occurrences, two mdependent reversals). The traditional consideration o fwing venation as the mam diagnostic feature to define and recognize stephanid genera has historically compelled authors to believe that 154 Parasfephanelhts is more closely related to Hemistephemus, with which it shares a very similar wing venation, than with any of the foenatopinae sensu Enderiein (1905), which show a highly derived venation. This situation is periiaps best illustrated by the 6 c t that these two genera were first recognized as a smgle taxon, ParastephanuSy which, together with Sdtlettererius and Sfephanus s. L, formed the subfiunily Stephaninae (Enderiein 1905). It does not seem surprising therefore, that when Enderiem (1906) split Parastephanus into Parastepfwnellus and Hemistephanus he maintained these taxa within Stephaninae. Results obtained in the present work, however, suggest a different view; Hemistephanus appears as the sister group of Parastephanellus + Foenatopinae ^ig. 198). hi the preferred cladogram Parastephanellus is separated fi’om its hypothetical common ancestor with (node 17) by a total of 52 synapomorpbies (those for nodes 18,21,22), whOe a much smaller number of 27 synapomorpbies separates it fi’om a common ancestor with Foenatopinae (those for nodes 22,28). Albeit the strong deferences in wing venation still represents an intuitive difiScuIty to the suggested relationship, it is reasonably counterbalanced by the intermediate position of Xanthostephanus n. gen., which is decidedly related to Foenatopus but has a wing venation which is similar to that o îParastephanellus, Additionally, even though it was not retrieved as a ^napomorphy for the group, the presence of a cell 1-^2R1 much larger than cell lM (ch. 115:1) represents a derived state shared by most Parastephanellus and Foenatopinae but does not occur in aiy other stephanid. The high frequent and confinement ofthis character state into a well 155 supported clade suggest that it should not be diconsidered as additional evidence of relationship. Overall, this clade is particulaly well supported by chs. 21,44, SO, 64,77, 85, 138, and I6Q. Parast^handtus Enderiein ^ode 22) The clade which defines this genus is supported by 18 synapomorpbies, of which the following 8 were recovered as unambiguous: paraocular strigation on fi'ons present and distinct (ch. 18:0-+I, two independent occurrences); vertex sculpture mostly or entirely transverse, even if ondulate or if vertex polished smooth posteriorly (ch. 24 :1-+2, three other occurrences, including as a synapomorphy for clade at node 42); gena with a well defoied yellow or ivory stripe extending fi’om base of mandible to temple (ch. 32: 2-+3, two independent occurrences); pronotal fold centrally collapsed, or at least closed, leaving two shallow or moderately deep openings on each side (ch. 55:0-+1, one independent occurrence); vein Icu-a slightly convex toward apex o f wing; sometimes slightly clavate (ch. 87: (M l, five independent occurrences, also a ^apom orphy for clade at node 47); vem RI shortly tubular or densely nebulous, very narrow (linear); not reaching midway distance between pterostigma apex and point where RI meets wing margin (ch. 101: l-»0, highly homoplasious); area of cell RI nearly or more than 3.0x as large as area o f cell IM (ch. 115:0-*^l, also a synapomorphy for clade at node 30, three independent reversals); post-foveolar area without a distinguishable lobe basally, straight throughout, not sharp (ch. 147:0-^I, unique). 156 Tea additional synapomorpbies resulted from character optimization: most of 2nd- 8th basal flagellomeres much more elongate than apical ones, the change in shape from basal to apical ones often abrupt (basal ones 8-I6x or more as long as wide versus usually 3-4x fr)r apical ones) (ch. 2 :0-+2, highly homoplasious); flagellomeres 2 and 3 with a few or several Type I sensillae, restricted to apical 0.4-0.7, portion of flagellomere with sensfllae usually somewhat dilated (ch. 6:0-^1 and ch. 7 :0-+1, both with many independent occurrences, but this is the onty clade consistently supported by these characters); posterior margm of vertex deeply convex in dorsal view (ch. 28: l-»0, highly homoplasious); apical sensorial hairs on pronotum organized in a tufr or small area (ch. 42: l->2, four other occurrences for small clades); setae on vein M+Cu reaching base of IM and usually advancing over ICu in at least one of the wmgs (ch. 78: l-»0, reversed for PoarastepfumeUm n. sp.; state 0 is also the groundplan state îotMegischuSy Stephanas, and Schlettererius); vein 2Cua longer than portion of vein Icu-a above CuP, but less than l.Sx as long (ch. 96:2-»0, reversed for P. pygm aeus and P. granulatus, state 0 is also the groundplan state for Megischus, Stephanas, and Schlettererius ); vein 4Rs entirely and ui#)rm ^ curved towards anterior margm, even slightly (ch. 109:0-*-l, two other occurrences, reversed tor P. pygmaeusy, prumosi^ on dorsal part of mesepistemum restricted to anterior G.3-0.5, remaining o f dorsal part glabrous (ch. 132:0-+2, highly homoplasious); pruinosi^ on mesepistemum (except pectus and dorsal part) restricted to anterior 0.3-0.S, often also only on dorsal 0.50-0.7S o fthis part, and moderately to 157 densefy abundant, concealing 50-95% of cuticular suiâce (ch. 135: l-»^3, revawd to I or 0 in several Parastephanellus spp.; state 3 common among F oenatopus spp. but does not occur in other stephanids). This genus was first proposed zs Parastephanus (Enderiein 1905), which included species fi'om the Ihdo-Australian as well as the Neotropical regions. The name was preoccupied by Haekel, and was later replaced by Parastephanellus (Enderiein 1906), but the Neotropical species were then assigned to a new genus, Hemistephanus. Among all features proposed by Enderiein (190Q to characterize and separate Parastephanellus fi’om other stephanids, only the relative size of cells I+2RI (Enderlein’s Cubitalzelle) and IM (Submendianzelle) seems to be found in a derived state for this genus, interpreted here as cell 1+2R1 considerably larger than IM (ch. 115:1). It was recovered as a synapomorphy for Parastephanellus in the preferred cladogram, but the plesiomorphic condition fisrthis character also occurs among some of its species. The point at which vein 4Rs ends in relation to the wing margin was also used by Enderiein as a diagnostic character to separate Parastephanellus and Hemistephanus. This vein, however, never effectively reaches the wing margin in stephanids, but ends at varied distances from it, so that the character is not discrete as suggested by Enderiein, but continuous and subjective. Due to the difficulty in defining character states, it was not considered in the cladistic analysis. Other features proposed by Enderiein to characterize Parastephanellus are discussed vmAes Hemistephanus^ which shows the derived states. Elliott (1922) mentioned the uniffirmfy colored sheaths o f the ovipositor in Parastephanellus spp. (that is, the absence o f a pre-apical white band) as a diagnostic 158 character. As investigated here (see ch. 175) it in 6 ct proved to be entirely reliable as a diagnostic difference Parastephanellus and Hemistephanus^ but was not retrieved as a synapomorphy for any genera. Ovipositor sheaths of a single color or without a pre-apical white band also occur in many basal taxa of Foenatopinae and in several species o f Megischus. An incompletely developed vein 2+3M (ch. 113:1) is present in most Parastephanellus^ and very rare in other stephanids, but was not retrieved as a synapomorphy for the genus because of its occurrence in the closely related Xanthostepharms n. gen. The few known exceptions in Parastephanellus I have been able to observe are the following: vein 2+3M complete and tubular for an undescribed species from the Solomon Is. (included in the present analysis) and for most species indigenous to New Caledonia, and this vein entirely absent or spectral for one unidentified species from Mindanao (Philippines). Foenatopinae Enderiem ^ode 28), N ew S ta tus This group is supported by 11 synapomorpbies, of which the following 6 were recovered as unambiguous: vein IRs aligned with IM, or nearly so, making cell R apically acute, triangular (ch. 83:0-^l, one more occurrence, at node 8); vein RI entirely absent; wmg margin between pterostigma and apex of RI weakly nebulous or completely clear (ch. 101: \-^2 y unique but with a few 2-+1 and 2-^0 reversals); spuracular carmaon Ime of fiision of metaploiron and propodeum, that is, origmating on spiracle and endmg «eternally to flant^ at pleuropropodeal fovea; spiracular impression usually inconspicuous 159 or absent, or extendmg alongside sptracular carina (ciu 141:1-^2, five mdependent occurrences); intercoxal space between bind coxae c o v e rt by a heavily scierotized plate formed by the extention and connection of metastemum with the posterior angles of propodeum (ch. 148:0-»-l, unique); bind coxa mesaUy slightly convex, albeit still somewhat flattened in relation to external side; hind femur at least mesally with Ion& delicate and completely decumbent hairs, or hairs minute (ch. 157:0-*^12, unique); petiole dorsally transversely strigulate, very fine, and very regular (ch. 170: 1-+2, unique). Five additional synapomorpbies resulted fi'om character optimization: prepectus mostly exposed (ch. 66: l->>0, highly homoplasious); pronotal spiracle distinctly ®q)osed (ch. 67:0-^l, four independent occurrences, two reversals); parastigma much shorter than IRs, or absent; when developed usually as long as wide, or wider than longer (ch. 80: 0-+2, two independent occurrences, one independent reversal); vein 2-1A curving, but not strongly, toward posterior margin of win& reaching margin opposite to or a little before apmc of 2Cua (ch. 92:0-^1, one independent reversal); hind coxa mesally slightly convex or distinctly convex, pilosity identical to, and sculpture not or only slightly different fi'om that on dorsal and ventral sides (ch. 153:0-+1, two independent occurrences, several reversals). This group was origmally proposed to include only those stephanids with greatly reduced wing venation and weakly scierotized pterostigma, equivalent to the literature genera F oenatopus Smith, Neostephanus Kiefifer, and Diastephanus Enderiein (treated here together asFoemtopus s. L). The deference in wmg venation in relation to other 1 6 0 stephanids is rather dramatic, and smce there are no known intermediate form^ the origmal proposition for the sub&nily seems to be well supported. Indeed, the mentioned literature taxa were recovered as a monophyletic group m the current analysis (Kg. 198, node 30). Nonetheless, I propose that the sub&mily concept be «cpanded to include Xantfmstephanus n. gen. and Foenatopus (^Madegqfoenus) xyrigi ^en o h ). The inclusion of F. x y r ig i as part of Foenatopinae is strongly supported by numerous synapomorpbies at node 29 (the longest branch in the cladogram), the 6ct that F. seyrigi shows the same type o f wing venation as Foenatopus s. 1., and the fact that Madegqfoenus is a polyphyletic genus, erected on the basis of artificial taxonomic characters (see discussion \xt\àe,T F oenatopu^. The inclusion o f Xanthostepfumus n. gen. seems justffîable mainly in function of two major synapomorpbies at node 28; vein 2-1A distinctly curved towards posterior margin o f wing (ch. 92:1) and presence of an intercoxal plate (ch. 148:1), both representing remarkable specializations within Stephanida^ which are invariably present and exclusive for those taxa withm the clade at node 28. Additionally, the independent origm o f ParastephanelluSy suggested by its well supported position as the sister group of Foenatopinae, and the numerous small clades or independent origins for taxa between Xanthostephanus n. gen. and F. arcuatus in the preferred cladogram, contribute to make node 28 a very reasonable breakmg point for the delimitation o f a (Linnean) taxon. 161 Xanthostqphanus m. gen. This taxon is supported by 29 autapomorphie^ of which 21 were recovered as unambiguous-: head yellowish brown, but vertex daricened (dark brown or black) (ch. 1: 0-+2, highly homoplasious); semiannular with fine dense short pilosity basally on femoral suclus only; hairs long (ch. 63:0-^1, four independent occurrences); ventral area of pronotum with fine dense pilosity basally, becoming more sparse apically until glabrous fiom midway on; sometimes finely e densely pilose only on extreme base (ch. 65:2-^1, highly homoplasious); setae on vein M-H]u placed far fiom 1\^ fiom which th ^ are separated by a distance greater than length of Icu-a (ch. 78: l-^2, two other occurrences, also a synapomorphy forHemistepharms); vein Icu-a straight, at nearly 90“ with both M+Cu and ICu (ch. 89: MO, state 0 also present in most taxa basal to node 17); vein 2- lA strongly curved, reaching wing margin befi)re middle of cell 2Cu (ch. 92: 0M 2, two other occurrences); vem 2-1A shortly tubular basally, remaining mostly or entirely nebulous (ch. 93:3-^2, two mdependent occurrences, but state 2 also present in several taxa basal to node 17); vein 2Cua longer than portion of vein Icu-a above CuP, but less than l.Sx as long (ch. 96:2->l, two independent occurrences); vein 2Cua making a distictly acute angle with ICu (ch. 97:0-^I [renumbered], five independent occurrences); vein 2Cub distinct, strongly nebulous, often tubular basally (ch. 98: MO, state 0 also present in most basal taxa); 4Rs (or its projected trajectory) reaching wing margin at or very near wmg apmc (ch. 112:0-+2, one more occurrence, at node 48); width of wing membrane fiom its posterior margin to CuP, at level of 2Cua, about as wide or distinctly wider than distance fiom apex of vein ICu to CuP (ch. 116: Q-»-l, three other occurrences, 162 includmg as a synapomorphy for clade at node 31); retinaculum placed mostly or entirely under cell ICu; its nearest to level of vein Icu-a (ch. 117: 0-^2, one more occurrence; at node 48); hamuli typical number usua% 3-4 (up to 7) Type ^ large or very large specimens often with 2-3 Type II hamuli (ch. 120:1-+0, highly homoplasious); hind wing vein Rs originating at (and fiised to) expanded portion of Sc+R which bears the hamuli (ch. 125:0-+2, unique); hind wing basally normal or weakly concave posteriorly, wing distinctly not pedunculate (ch. 130:2-^1, one independent occurrence, but state I also present in most taxa basal to node 17); metapleuron pilosity very densely abundant, blocking 50-100% of pilose sur&ce, dorsal area completely taken by short pilosity, which reaches spiracular carina (ch. 145:0-*^3, also a synapormorphy îox Hemistephanus)^ hind coxa distinclty longer (1.3-1.5x as long) than mesepistemum (ch. 150:1-+2, highly homoplasious, but restricted to Foenatopinae); hind coxa very long and slender, 4.25x or more as long as laterally high (ch. 151:0-+1, four other occurrences); apical spurs of hind tibia of equal length (ch. 164: l->0, four other occurrences, state 0 present in several taxa basal to node 21); stemite 7 with flat and curved, translucent setae latero-ventrally, even if only one on each side (ch. 174:0-*I, four other occurrences, including as a synapomorphy for clade at node 42). Eight additional synapomorpbies resulted fi’om character optimization: intreocellar carinae placed mostly posterior to, with a few m between, the posterior ocelli (ch. 23: l-*0, one independent occurrence; state 0 also occur for vaosX.Meffschus and other basal stephanids); apical sensorial hahs of pronotum organized in a tuft or small area (ch. 42: 1-+2, four other occurrences); apical emargmation of pronotum distinctly and strongly 163 emarginat^ C-shapped (ch. 4 4 : unique among Parastephanellus-i-Foeaaiopiaaey but state 1 also occur formost Megischus and other basal stephanids); femoral sulcus (pronotum) present as a distinct oblique-longitudinal impression, even if weak; ventral area of pronotum distinctly delimited, with different sculpture from the usually polished smooth surfece of the femoral impression (ch. 64: l->0, highly homoplasious); mesoscutum large and round, distinctly larger than combined area of axillae (ch. 72: l->0, unique within Hemisiephcptus +■ Parastephanellus + Foenatopmae, but state 0 also occur for most M egischus and other basal stephanids); vein IM originating at or basad of Icu-a, so that nodus formed by fusion of M-fCu/lM with Icu-a/lCu is oblique toward wing base (ch. 8 5 :1-+0, highly homoplasious); post-foveolar area practically aligned with interfoveolar area (ch. 146:1-+0, one independent occurrence, but state 0 also occur for all Hemistephanus and most basal stephanids); hind femur with a dorso-mesal compression, on basal half or centrally, but subtle and often difGcult to visualize (ch. 159: 1-+0, highly homoplasious). There seems to be only one non-homoplasious apomorphy fat Xanthostephanus n. gen. (ch. 125:2), all other appearing at least once more somewhere else in the cladogram. Note, however, that the large number of homoplasious apomorphies represent a highly unique and somewhat surprising mix of plesiomorphic and specialized features. In feet, about 50% o f the «qmmorphies for this genus (chs. 23,44,64, 65,72, 85,89, 93,96,97, 98,120,130,146,164) represent states which occur essentially in members of the basal genus txdiile several of the remainmg (chs. 112,116,117,150,151,159,174) 164 are shared mostly or only with apical taxa o fFoenatopus. Hbmoplasy in this case can therefore be understood as an obvious result of the consideration of a large proportion of these features either as independent reversals or as independent parallel derivations. Additional support for the independent taxonomic status o f Xanthostephanus n. gen. derives from the feet that it (^peered m the same position in all cladograms, weighted or unweighted, never forming a clade with any other stephanid. Its distant position in relation to other stephanids, indicated by the lengths of the three branches associated with node 28 (Hg. 198) also seem to warrant its treatment as a distinct genus (note also that the computer program used to draw the cladogram considers only unambiguous characters; ^the additional ^mapomorphies inferred from character optimization could be included the length of the branches would increase). Finally, the overall shape of the front wing is also unique for Xanthostephanus n. gen. QFig. 170), but it has not been coded and included in the matrix because of the great difficulty in objectively categorizing wing shapes. Foenatopus Smith ^ode 29) This taxon is supported by 32 synapomorpbies, of which the following 21 were recovered as unambiguous: flagellomere 2 about as long (0.9-1.0) as flagellomere 3 (ch. S: 1-+0, highly homoplasious); frons ventralfy rugose or rugulose (ch. 17:3-*^2, highly homoplasious); anterior half of vertex with granular microstculpture in between macrosculpture (ch. 27:0-+I, three other occurrences); occipital carina completely disappearmg far from hypostomal carina, or feding gradually with both sides converging 165 toward each other, or toward center of hypostomal carina, without ever meeting (ch. 34: 0-*^!, one mdependent occurrence, several reversals); spur of antennal cleaner with vellum distinctly wider apically than centrally, so that it is somewhat lobed; ending abruptly, sometimes with a small emargination apically (ch. 7 1 :1-*^2, four mdependent reversals); notauli not differentiated (ch. 76:0-+1, one independent occurrence, several reversals); vein Icu-a narrowed basally, thickened apically, or suddently thick only apically; so that it is slightly clavat^ or clubbed (ch. 88:0-+1, three independent occurrences, four reversals); vein ^s+M )b, 2+3Rs, and 2+3M spectral or, more rarely, nebulous (ch. 102: 1-^2, unique; ch. 103:0-*2 [renumbered], unique); spurious vein arising from (Rs+M)b, towards base of pterostigma present, normally spectral, sometimes nebulous, never tubular (ch. 105:0-*^l, several reversals); vein 2+3Rs strongly sinuate, spectral or, more rarely, nebulous (ch. 106:0-»^l, unique); vein 4Rs tubular part ending fer from wing margin, becoming apically spectral, weakly nebulous, or simply absent (ch. 107: O-^I, unique, two independent reversals); vein 2+3M entirely nebulous or spectral (ch. 113: l-*2, unique); CuP straight or nearly so, but apically usually curving slightly in anterior direction, along 2Cub; or at least mostly straight, concave under cell 2Cu only (ch. 114:0-+1, unique one independent reversal); mesepistemal hafrs, except on pectus and dorsal part, very small or minute, indistinct from pruinosity (if present), decumbent, each hair barely or not reaching base o f next nearest hahr (ch. 133: l-»0, highly homoplasious, but no reversals m Foenatopus); mesopleural pit present, always accompanied by characteristic transverse or oblique rugosities arround it (ch. 136:0-»^l, one mdependent reversal); hind coxa basally deeply concave both externally and mesally (ch. 149:0-*^l, one independent occurrence 1 6 6 one independent reversal); hind coxa with minute hair^ ixdiich are much shorter than basal femoral tooth, decumbent, delicate placed fer away from each other (ch. 156; (M l, one independent occurrence); hind femur with most or all hafrs minute, much shorter than basal femoral tooth, decumbent, delicate, placed fer away from each other (ch. 157: M 2, one mdependent (M2 change); hind tibia structure as in Fig. 138 (ch. 162: M 5, unique); paratergites of T3 wide enough to meet or overlap ventrally; or meeting or overIappm& so that 3rd abdommal segment is neariy completely encased by T3 (ch. 173:0-> 1, three other occurrences, three mdependent reversals). Eleven additional ^apomorphies resulted from character optimization: flagellomeres 2 and 3 with a few or several Type I sensillae, restricted to apical 0.4-0.7, portion of flagellomere with sensillae usually somewhat dilated (ch. 6:0-*^! and ch. 7: (M l, both highly homoplasious); frons virtually glabrous, hairs minute, about as long as height of ridges of sculpturing, and very scarce, or totally absent (ch. 19:0-+2, one mdependent occurrence, nearly complete reversal to 0 at node 42); posterior margin of vertex (dorsal view) deeply convex in dorsal view (ch. 28: MO, highly homoplasious); labrum yellowish to whitish ventrally and dorso-apically, dorso-basaUy distinctively darker (usualfy same color as fece/frons) (ch. 40: M 2, one mdependent occurrence, several reversals); carinae on colo in dorsal view straight and oblique on each side, generating a V-pattem; carmae mcomplete, not or barely meetmg dorsally (ch. 50: M 2, one independent occurrence, one reversal); mesoscutum conspicuously transverse, more than 3.0x as wide as lon^ and distinctly smaller than combined area o f axillae (ch. 72: M 2, one mdependent occurrence one complete reversal at node 48); vein ICu tubular on basal 167 20% or more, but distinctly less than 100%, with apex n^ulous o r spectral (ch. 9 0 :0-^l, unique four independent reversals); vein 2Cua âintly nebulous or spectral (ch. 94:2-^3 [renumbered], unique four mdependent reversals, including one to state I [=0 m the dadogram]); vein 2Cua making a distinctly obtuse angle with ICu (ch. 97:0-+2 [renumbered], three other occurrences, three reversals, including one to state 0); pruinosity of mesepistamum (except pectus and dorsal part) absent or nearly so, at most with a small amount of sparse pruinosity antero-dorsally (ch. 135:1->0, highly homoplasious). The genus Foenatopus is immediately distinct from other stephanids by its extremely “reduced” wmg venation, that is, most veins on the apical half of the front wing are entfrely spectral. In fact, since proposed by Smith (1861a), there has been nearly complete consensus in the literature as to attributing separate taxonomic status for its species, even though generic concepts varied. The only exceptions were Schletterer (1889a) (see discussion îo t Stephcmus) and Benoit (1951a) (discussed below). There is a considerable variation in the degree of sclerotinazation of the vems ICu and 2Cua among Foenatopus s. 1., a 6ct which led to the proposition of the goiera Neostephanus (Kiefrèr 1904) for spedes with both these veins tubular or strongly nebulous, andDiastephanus (Enderiein 1906), for spedes in which such veins are entirely spectral. The genus F oenatopus Smith was then restricted to contain the intermediate forms, that is, taxa with vein ICu scierotized and 2Cua spectral (Enderiem 1906). In spite 168 o f the Act that no other character has ever been proposed to support these genera, Endedein’s interpretation persisted as the most widely adopted classification for this group o f stephanids. Elliott (1922), however, had already suggested that albeit helpful in aiding the separation of its numerous species, such division seemed considerably artificial. Even so, the only alternative to Enderiein’s scheme was proposed by Benoit (1951a, 1956). By attributing higher taxonomic value to the shape of the hind tibia than to the traditionally used wing venation characters, Benoit (1951a) proposed the ÿscmMeukgqfœnus for a group of species which included forms with wing venation o f M egischus type as well as forms with the F oenatopus type. However, while the shape o f the hind tibia as remterpreted here indeed seems to be of high taxonomic and phylogenetic significance (see ch. 162), it contributed instead to demonstrate that the genus Madegqfoenus is distinctly pofyphyletic: most species mcluded in it fidl ^MÜàsiMegischus, and the genoQrpe, M seyrigiy appears basal to all oHaer Foenatopus s. I. (Fig. 198). Albeit several apomorphies support M seyrigf, utilizing this species to redefine and maintain the validity of Madegafoenus does not seemjustffîable at the moment for at least two reasons: (I) as one o fthe smallest known foenatopmae spedes, it is difficult to define whether the derived or basal condition of many characters inM seyrigi have supra-specffic relevance or ^ thqr represent spedes-level modifications related to character reductions or fusions, which tend to occur in small spedmens of Hymenoptera; and (2) the large number of synapomorpbies which placesM seyrig i within the clade at node 29 not only nnply a close relationship with, but also make this spedes extremely similar to other spedes with reduced wing 169 venation, providing strong practical and intuitive difficulties for its justification as a separate taxon. A third problem derives fiom the observed instability of the clades within Foenatopinae, which will be discussed below. Benoit (1956), however, provided a more effective critique to Enderiem (1906) by noticing that vein ICu may show intermediate stages of development between species of Foenatopus wvi. Diastephanus, suggMting that a full range of forms fi’om ICu entirely spectral to entirely tubular may occur. Based on this foot, Benoit (1956) justifiably proposed the suppression oîDiastephanmEtiéeùeâxi, considering it as a junior synonym foTFoenatopus Smith. Similar variation was also observed intra-specifically by Benoit (1984a) who reported that while most F. macrocephalicus ^enoit) have an entirely spectral ICu, some specimens show a short basal portion of it distinctly tubular. The high variability of this character was confirmed in the present work, and even with the remterpretation of each character state as a particular “range” of possibilities (see ch. 90) they continued to prove highly homoplasious within Foenatopinae (nonetheless, note that an incompletely tubular vein ICu represents a synapomorphy for F oenatopus s. 1). In feet, none of the foenatopinae taxa as revised and re d d e d by Enderiein (1906) were recovered in the present work as monophyletic units, but appear instead scattered withm the large clade hymning at node 30 (see also discussion for Foenatopinae). At the same tune, clades with the genotype species (nodes 45 and 46) are highly exclusive, containing only a few species. If a generic concept is to be attributed to them, then other equivalent but unnamed clades would require the same approach, forcing the proposition o f several new genera. Most of these clades, however, are supported by a small number of 170 weak synapomorphies, and have been highly variable througout the development of the current study. Such variability is dramatically demonstrated by the very poor resolution in the consensus tree for the unweighted searches ^igs 195, 196), as well as the intensely different clades between the preferred cladogram (Fig. 198) and the cladogram obtamed with a separated analysis o f the foenatopinae (Fig. 200). Such results indicate that in spite o fthem enormous morphological variability, there is no convincing evidence at the moment to justify more than one genus among the stephanids with “reduced” wing venation. General Considerations The historical difSculty in recognizing and defining stephanid taxa in a convincing manner, and the ensuing confusion towards the recognition of the few described genera, are a reflect of the high morphological complexity of the group. Stephanids have a peculiar general structure, which makes them difficult to interpret taxonomicaHy: the general structure and morpholo^ are quite uniform throughout the femily, but the intra- and interespecific variation of taxonomicaHy important characters is very high. For example the veins appearing in the fi'ont wmg are nearly the same for all stephanid species, but dËfer greatly in their degree of sclerotinization, to the point that in Foenatopus spp. most vems are c^pa rentfy completely absent. Similar variation patterns occur with surfece sculpture, pHosity, and serveral other characters. Since only a few comparative investigations, of limited scope, have been conducted with the group, it seems e a^ to understand why definitions for its genera could but resist tenuously to any kmd o f reassessment. 171 The considerably uniform body structure and the occurrence of perfectly intermediate forms between different species and different character states throughout the âmEy, persuaded some authors to adopt (Schletterer 1889a), defend (Ceballos 1926) or at least show some complacenty C^Hiott 1922) towards the idea that all stephanids belong to a single genus. Obviously, even today the idea of how inclusive a genus concept should be is not entirely objective, but the nature of the concern implied by the above authors was dffferent: that among stephanids, perfectly well delimited (i.e., non-overlapping) groups of species did not exist or could not be defined. However, the present results indicate that such idea can be hnmediatly and efficiently refuted by the demonstration that at least two very different lineages of stephanids are retrieved under all known phylogenetic algorithms, virtually under any parameter the gtasxdiHemistephanus, Xanthostephanus^ and Foeruaopus s. 1. These taxa represent very distinct monophyletic units, the latter two supported by the longest series of synapomorphies in the cladogram. Effects of “long- branch attraction" in the definition o f Foenatopus and Xanthostephanus are disregarded here, in full agreement with the opinion of Siddall & Whiting (1999) that there is no logical justffication to interpret it as a problem. Furthermore, both branches have exactly the same “length,” Le., thqr are supporte by 21 synapomorphies each. Perhaps the main reason for the belief in the idea of a single genus was the fact that near^ all proposed stephanid genera were defined on the basis of wing venation characters (e.^. Ashmead 1900b, Kieffer 1904, Enderlein 1906), many of which showing a range of intermediate forms. Benoit (1951a) went as for as considered wing venation too variable to be used m recognizing stephanid genera; he proposed a new set o f (a few) characters to 172 support and define a new stephanid genus, Mad&gqfoenus^ which mciuded fenns with very different types of wing venation. Nonetheless, Benoit restricted this interpretation to the proposed genus, and continued to adopt stephanid taxa defined exclusively by wmg venation characters (e g., Benoit 1951b, 1953), and to synonymize (and redefine) stephanid genera based on these characters ^enoit 1956). The only character which proved to be informative among those proposed by Benoit (1951a) (ch. 162) ironically contributed to refiite Aforikgtybemts in fiivor o f previously established genera, as seen in the discussion îor Foenatopus. Interestingly, however, when data from the present analysis were reevaluated using wing venation characters only (chs. 80-129) results were essentially the same when compared with analyses of the entire data matrix, that is, all major monophyletic groups important for genus-level definition were recovered (under implied weights), accept only for S tep h a m s (Fig. 199). These results suggest that the long history of ambiguous interpretation of stephanid genera was probably more related with the lack of adequate methods for a comprehensive evaluation of the fomüy than with a supposedly “low” taxonomic value of the characters used. Key to Subfamilies and Genera 1 Intercoxal space between hind coxae membranous (Rg. 108); most or all veins on apical half o f front wing tubular or strongly nebulous (Figs 163-169, 171) ...... Stephaninae... 2 173 r bitercoxalspacebetweenhmdcoxaecoveredbyaheaviiysclerotizedpiate(Fîg. Ill); veins on apical half of fix>nt wing spectral or weakly nebulous C^gs 172-178) (except Jûmthostephanus n. gen.. Fig. 170) ...... Foenatopinae... 6 2(1) Second abdominal tergite not more than I.Sx as long as T3; hind tarsus of females withS tarsomeres ...... 3 2’ Second abdominal tergite or petiole distinctly more than 1.5x as long as T3; hind tarsus o f females with 3 tasom eres...... 4 3(2) Preannular absent, pronotum with a flat vertical wall between colo and semiannular (Fig. 55); with a conspicuous apico-dorsal spine on hind coxa (Fig. 123); T2 and S2 isolated, not forming a petiole (Fig. 143); gena without yellow marks; vein 2+3M tubular on basal half or less, remaining nebulous (sometimes strongty) or spectral (Hg. 163) ...... Schlettererius 3 ’ Preannular dKforentiated, although continuous with semiannular, pronotum elongate (Fig. 58); apico-dorsal spine on hmd coxa absent; T2 and S2 fosed into a petiole, albeit a lateral edge where both sclerites fuse together can be seen basally (similar to Fig. 145); gena with a yellow spot basally, on or near malar space; vein 2+3M tubular on basal 0.8 or more, but never entirely, with apex nebulous orspectral(Rg. 164)...... S tepha nus 4(2) Ventral margin of post-foveolar area basally produced mto a lobe, which covers a little the base of the mid coxa ^ g s 104,106,109); mid tibia without spmes on anterior fece; paraocular strigation absent; (Rs+hdQb normally (99%) separated flrom 2+3Rs owf 2+3M by distinct bullae (Figs 163-166,168,169), 174 or continuous only with 2+3M (Fig. 167), or all spectral; ovipositor sheaths color variable...... 5 4’ Ventral margin ofpost-foveplar area straight throughout ^ig. 107); mid tibia with a few, ventrally inclined spines or spine-like setae along anterior fece (Figs 112, 113); paraocular strigation usually (90%) distinct (Fig. 2); (Rs+M)b continuous with 2+3Rs and separated from 2+3M by a bulla (Fig. 171); ovipositor sheaths unicolorous ...... Parastq/handbis 5(4) Setae on M+Cu reaching IM and usually advancing over ICu in at least one of the wings, or at least ending near IM, separated from it by a distance never greater than length of Icu-a; retinaculum mostly under cell 2Cu, its apex nearest to level of vem 2Cua (Figs 163-167, 171, 172); apical margin of pronotum a simple rim, slightly to moderately reflexed upwards (Figs 57,60, 63); wings often partially or unftbrmly infuscated ...... M egischns 5’ Setae on M-K^u placed far from IM, from which th ^ are separated by a distance greaterthan length of Icu-a; retinaculumplaced midway between cell ICuand cell 2Cu; its apmc approxhnately equidistant from vems Icu-a and 2Cua ^ g s 168, 169); apical margin of pronotum usually convoluted, generating a wide collar OFigs 59,62), which is alutaceous dorsally; wings usually clear hyaline ...... Hendstephanus 6(1) Most vems on apical hafr^offront wmg spectral or weakly nebulous ^ g s 172-178); vein 2+3Rs distmct^ smuate (spectral); (Rs+M)b distmctly basad of point where vein 2r arises from parastigma; mesopleural pit present, always 175 accompanied by characteristic rugosities (Rgs 102, 103); mesoscutum transverse, neariy as large as, or smaller than combined area of axillae (Figs - 97,99) ...... F oenatopus 6’ Most veins on apical half of fiont wmg tubular or strongly nebulous (Kgs 170); vein 2+3Rs straight or slightly curved (tubular); (Rs+M)b distinctly opposite to point where vein 2r arises 6om parastigma; mesopleural pit and rugosities absent; mesoscutum large and round, distinctly larger than combined area of axillae (as inFigs 92,95) ...... Xanthostephanus n. gen. Descriptions GenusSchlettererius Ashmead Stephanus (psri): Cresson, 1887:52. Schlettererius A^ameeâ, 1900b: 150; Szépligeti, 1902: 528; Enderlein, 1905:474; 1906:290; Kieffer, 1908:3, 5; Elliott, 1922: 710, 713; Townes, 1949:362; Madl, 1991: 118-120; Belokobylskij, 1995; Gauld, 1995:184. Type species: Stephams dnctipes (Cresson), designated by Ashmead 1900b: 150, H ead.^ Head globose or almost so (as Hg. 27). Paraocular strigation absent. Face pQosity m between antennal foramina until clypeal suture absent, or as in frons. Granular sculpture on coronal area and &ons absent: polished smooth m between macrosculpture. Vertex sculpture areolate-rugose latero-anteriorly, transverse centrally, polished smooth on temples; two or more different sculpture patterns, approximately in the same proportion; sculpture very coarse. Posterior margin of head seen dorsally deeply convec 176 ^ g . 34). Gena without yellow or white colors. Occipital caiina ending very close to^ or extending alongsde hypostomal carina, reaching its apec (as Figs 38,39). Post-genal bridge sinkmg immediately behind hypostomal carina, even doing so only centrally; post-genal line, in consequence, entirely inclined m relation to saghal plane of oral fossa (as Fig. 38). Maxillary palpus approximately as long as head is laterally high. Prothorax.- Pronotum (Figs 54,55): ^ ic a l sensorial hairs of pronotum inconspicuous or absent. Apical m ar^ of pronotum a simple rim, slightly to moderately refleced upwards, so that it is L- or C-shaped in cross section. Apical margin of pronotum weakly angled. Colo transverse and very narrow, about three times wider than long; with transverse, complete or incomplete carinae, which are strong, often with at least one pair, or one complete carina, vertically raised as a blade-like and weakly translucent structure; centro longitudinal depression absent. Preannular absent Prepectus mostly exposed (as Fig. 85). Angle between apical margins of hemistemites of prostemum about 90" or more (Fig. 83). Front coxa laterally with several small foveolae (as Fig. 85). Meso andMetathorax.- Mesoscutum large and round, distinctly larger than combined area of axillae ^ig . 92). Dorsal part of mesepistemum mostly or entirely uniformly pilose or pruinose, but hairs moderately sparse to very sparse, so that 50-95% of cuticular surface is visible. Pruinosity on mesepistemum, secluding pectum and dorsal part, absent, or nearly so, at most with a small amount of sparse pruinosity antero-dorsaily (shnilarto Hg. 103). Mesopleural pit absent. Mid tibia without spines on anterior face. Spiracular impression and carina not differentiated. Ventral margin of post-foveolar area basally produced into a small or very prominent lobe with sharp edge, which covers a little 177 o fthe base ofthe mid coxa (as Hgs 105,106). Intercoxal space between hind coxae membranous (as R g. 108). Hmd coxa basally deeply concave on external side only (as Rgs 114, 118); shorter than maxhnum lateral length of mesepistemum; shape short and wid^ subconicai (Fig. 123). Apico-dorsal spine on hind coxa present and conspicuous ^ g . 123). Mesal glandular opening basally on hind femur absent. Ventral edge of hmd femur basad of central tooth with one single tooth or tooth-like tubercle (Fig. 129). Hmd tibia weakly laterally compressed on basal 0.1-0.2, then widening progressively and moderately towards apmc (Fig. 133). Ventral longitudinal carina on hind tibia restricted to compressed part, although often advancing a little over dilated part (as Fig. 141). Hnd tarsus o f female with five tarsomeres (Fig. 146). BEnd basitarsus ventrally with 1-2 rows of long translucent spmes, or at least seta^ which are bigger basally, getting smaller and weaker towards apex (as Fig. 146 or more delicate). Front Wing.^ (Fig. 163) Costal cell with very small and sparse but conspicuous setae, usually more firequent centrally or apically. Setae on M+Cu reaching base of IM and usually advancing over ICu in at least one of the wings (Figs 155, 158). Parastigma distmctly longer, or at least approxhnately as long as vein IRs, and distinctly longer than wide. Vein IM usually at about 90" with IRs, making cell R truncate apically; more rarely forming a distinctly obtuse angle; distinctly longer to much longer than IRs (observed: 1.2-6.5x); originating at or basad Icu-a, so that nodus formed by fusion ofM+Cu/lM with Icu-a/lCu is backwardly oblique; distmctly and enthely curved. Vein Icu-a straight or neariy so; at about 90" with both M+Cu and ICu. Vein ICu entirely tubular. Vein 2-1A straight; entirely nebulous; when intense^ so, appearing to be tubular. Vein 2Cua entirely 178 tubular, at about 90** with ICu. Vein 2Cub distinct; strongly nebulous, often tubular basally. Often forked {^>ically. Vein 2cu-a semi-tubular or ndiulous. Vein (Rs+M)b disconnected from 2+3Rs by presence of a bulla; disconnected firom 2+3M by presence of a bulla; distmctly short. Spurious vein arising from (Rs+hQb absent. Vein 2+3Rs straight or slightly curved, tubular. Veins 4Rs and 2r forming a distinct angle with each other, or at least notably discontinuous. Vem 4Rs longitudinal, but usually neither perfectly straight, nor uniformly curved, if straight, not parallel to pterostigma, and usually slightly inclmed anteriorly; more than 3x as long as 2r, up to 1 Ix. Vein 2+3M tubular on basal half or less, rest nebulous (sometimes strongly), or spectral. Area of cell R1 up to 2x as large as area of ceil IM. Retinaculum mostly under ceil 2Cu, its apex nearest to level of vein 2Cua. Hind Wing.- (Fig. 179) Hamuli usually 3-4 (as Fig. 149), but up to 7 Type I hamuli in at least half of the specimens, large or very large specimens often with 2-3 Type II hamuli. Vein M+Cu distmct, oblique usually nd>ulous, but sometimes spectral, often continued by, or indiscernible from IM. Vein Rs usually nebulous, but sometimes spectral; lon& straight, and longitudinal, its origin distinctly unrelated to Sc+R Vein Ir-m present, even very faint, better developed m large specimens. Vein Cua present, nebulous or spectral. Dorsal wing fold present, complete, although sometimes difficult to seem small specimens; Ir-m, if present, interrupted by dorsal fold. Ventral wing fold present at least basally, although somethnes dKGcult to see in small specimens. Wing membrane basally round, wide; posterior margin convex. Wing apically round and broad. 179 A bdom en -Second tergite and stennte distinctly isolated, not forming a petiole; about as long as dorsal length of T3 (Bg. 143). Ovipositor sheaths white- or yellow-banded subapically. GenusS tep h a n u s Jurine Aie/iAantts Jurine, m Panzer, 1800: tab 13; Jurine, 1807:91-3; Latreille, 1807:3-4; LatreOI^ 1810:298; Nees ab Esenbeck, 1818:301; LatreOle, 1825:445; Nees ab Esenbeck, 1834:8; Blanchard, 1840:329; BruUé, 1846: 536; Blanchard, 1849:23; Desmarest & Chenu, 1860:172; Sichel, 1865; BCirchner, 1867: 141; Westwood, 1874; Oehlke, 1984:163;Pagliano, 1986:13; Madl, 1991:120-1; Tobias, 1993: 391; Belokobylskji, 1995:16; Scaramozzino, 1995:3. .Xbrwfes (part): Lamarck, 1817:134-5; 1835:346-7. Stephanus (pdxi): Ashmead, 1900b: 150; Kieffer, 1904:1; 1905:478-9; 1908:3; Enderlein, 1905:474-5; 1906:290,306; Elliott, 1922: 710, 713, 715; CebaUos, 1926: 137; Dutt, 1926:1; Schmiedecknecht, 1930; Berland, 1951:928. Morphology, biolo^, figure; Orfila, 1956:6; DeSantis, 1980:7. Type species: Stephcams senator ÇBûmàas, 1798), by monotypy. H ead,- Head globose or almost so (as Fig. 27). Paraocular strigation absent Face pilosity in between antenna! foramma until clypeal suture absent, or as in firons. Granular sculpture on coronal area and ffons absent: polished smooth in between macrosculpture. Vertex sculpture partially transverse except laterally, or at least antero-Iaterally, where any other orientation (except areolate-rugose) may occur; two or more different sculpture 180 patterns, approximate^ ui the same proportion; sculpture intensity regular or very fine. Posterior margin of head seen dorsally deeply convex (as Fig, 34). Gena with a yellow spot basally, at or near malar spac^ which is fiunt if head has a light color. Occipital carina endmg very close to, at, or extending alongside hypostomal carina, reaching its apex (Figs 38,39). Post-genal bridge sinking immediately behind hypostomal carina, even if doing so only centrally; post-genal line, in consequence entirely inclined in relation to sagital plane of oral fossa (Fig. 38). Maxillary palpus distmctly longer than head is laterally high (as Kg. 29). Prothorax.- ^igs 57, 58) Apical sensorial hairs of pronotum organized linearly, in a single line or narrow stripe even if only a few or very small. Apical margm of pronotum a simple rim, slightly to moderately reflexed upwards, so that it is L- or C-shaped in cross section. Apical margin of pronotum distmctly and strongly emarginate C-shapped. Colo normal, about 0.5-1.0 as long as wide; with transverse, complete or incomplete carinae which are strong often wnth at least one pair, or one complete carine vertically raised as a blade-like and weakly translucent structure; centro longitudinal depression absent. Pronotal fold distinct; and distinclty, often deeply, concave anteriorly, never collapsed centrally. Transverse ruga and groove on prenannular absent Preannular perfoctly continuous with semiannular. Prepectus mostly exposed (as Fig. 85). Angle between apical margins o f hemistemites of prostemum about 90" or more (as Fig. 83). Front coxa laterally with several small foveolae (as Fig. 85). Meso and Metathorax.- Mesoscutum large and round, distinctly larger than combmed area of axillae (Kg. 93; see also Kg. 92). Dorsal part of mesepistemum mostly 181 or entirety uniformly pilose or pruinose, but hairs moderately sparse to very sparse so that 50-95% o f cuticular surAce is visible. Pruinosity on mesepistemum, excluding pectum and dorsal part, absent, or nearly so, at most with a small amount of sparse pruinosity antero-dorsalty (similar to Fig. 103). Mesopleural pit absent Mid tibia with a few spmes or spme-like setae along anterior Ace, inclined toward apex (as Fig. 112). Spiracular carina af line of Asion of metapleuron and propodeum, ending below (i.e., externally to) flanlt at pleuropropodeal Avea; spiracular impression usually absent ^ present extending alongside spiracular carina (as Fig. 104). Ventral margin of post-Aveolar area basally produced into a small or very prominent lobe with sharp edge, which covers a little of the base of the mid coxa (as Figs 105, 106). Intercoxal space between hind coxae membranous (as Fig. 108). Hind coxa basally deeply concave on external side only (as Figs 114,118); shorter than maximum lateral length of mesepistemum; shape similar to Figs 117 and 123. Apico-dorsal spine on hind coxa absent Mesal glandular opening basally on hind Amur absent Ventral edge of hind femur basad of central tooth with one single tooth or tooth-like tubercle (Fig. 128). Hind tibia laterally compressed on basal 0.3-0.5, dilated apically ^ g . 134). Ventral longitudinal carina on hind tibia restricted to compressed part although often advancing a little over dilated part (as Fig. 141). lAid tarsus o f Amale with Are tarsomeres (Hg. 146). Hmd basitarsus ventrally with 1-2 rows of Ion& translucent spines, or at least seAe, which are bigger basally, getting smaller and weaker towards apex (Hg. 146). Front Wmg.- (Hg. 164) Costal cell with very small and sparse but conspicuous seta^ usually more Sequent centrally or apically. Setae on M-KJu reaching base of IM and 182 usually advancing over ICu in at least one ofthe wings (Figs 155, 158). Parastigma distinctly longer, or at least approximately as long as vein IRs, and distinctly longer than wide. Vein IM usually at about 90" with IRs, makmg cell R truncate apically; more rarely forming a distinctly obtuse angle; distinctly longer to much longer than IRs (observed; I.2-6.5X); originating at or basad Icu-a, so that nodus formed by fosion of M-Küu/IM with Icu-a/ICu is backwardly oblique; slightly curved. Vem Icu-a straight or nearly so; at about 90" with both MKZu and ICu. Vein ICu entirely tubular. Vein 2-1A straight; tubular on 0.9 or more of its length. Vein 2Cua entirely tubular, or partially tubular (basal 50% or less); apically nebulous, even if strongly, appearing to be tubular; making a distmct, but never greatly obtuse angle with ICu. Vein 2Cub distinct; strongly nebulous, often tubular basally. Often forked apically. Vein 2cu-a semi-tubular or nebulous. Vein (Rs+M)b disconnected from 2+3Rs by presence of a bulla; disconnected from 2+3M by presence o f a bulla; distinctly short. Spurious vem arising from (Els+M)b absent. Vein 2+3Rs straight or slightly curved, tubular. Veins 4Rs and 2r forming a distmct angle with each other, or at least notably discontinuous. Vem 4Rs longitudinal, but usually neither perfoctly straight, nor uniformly curved, ft' straight, not parallel to pterostigma, and usually slightly inclined anterioriy; more than 3x as long as 2r, up to I Ix. Vein 2+3M tubular on basal 0.8 or more, but never entfrely, with apex nebulous or spectral. Area of cell RI up to 2x as large as area of cell IM. Retmaculum mostly under cell 2Cu, its apex nearest to level ofvein2Cua. Hind Wing.- (Fig. 180) Hamuli usually 3-4 (as Fig. 149), but up to 7 Type I hamuli in at least half of the specimens, large or very large specimens often with 2-3 Type II 183 hamuli. Vem M-K^u distinct» oblique usually nebulous, but sometimes spectral, often continued by, or indiscernible from IM. Vein Rs usually nebulous, but sometimes spectral; lon^ straight, and longtudinal, its origin distinctly unrelated to ScfR. Vein Ir-m present, even ifvery frtint, better developed in large specunens. Vem Cua mdicated by a nebulous stub commg out ofMK^u. Dorsal wing fold present, complet^ although sometimes difScult to seem small specimens; Ir-m, if present, interrupted by dorsal fold. Ventral wing fold present at least basally, although sometimes difiScult to see in small specimens. V ^g membrane basally normal or weakly concave postenoriy, wing distinctly not pedunculate. Wing apically round and broad. A bdom en.- Second tergite and stemite fused into a petiole, which is about as long as dorsal length of T3. Ovipositor sheaths unicolorous, sometimes progressively darker towards apex. Genus Afegischus BruUé Stepham s of authors nec Panzer, 1800, nec Jurine, 1807. Pim pla (part): Fabricius, 1804:112. MegischusBvjM^ 1846:537; Westwood, 1851:227; Smith, 1861a: 59; Sichel, 1865:467; Cresson, 1887:52; Townes, 1949: 362-3; Baltazar, 1962:744-5; 1966:15; Madl, 1991:118,122-5; Tobias, 1993:391; Belokobylskij, 1995:18; Scaramozzino, 1995: I; Aguiar, 1998:365. M egischus (part): Gauld, 1995. 184 Stephanus: Desmarest & Chenu, I860:172; Szépligeti, 1902:528; Westwood, 1874; Ashmead, 1900b: ISO; Enderiein, 1905:474-5; Kie&r, 1908:3; Viereck, I9I4:22; CebaUos, 1926:137;EUiott, 1922: 710,713,715-46; 1926:517; 1927a: 211;Brues& Melander, 1932:476; Medler, 1980:443. Boihrioceros Sichel, 1860:759. Madegqfoenusipzit): Benoit, 1956:205. Stephanus (Megixhus): DeSantis, 1980:7. Type species: Stephanusjiircatus LeBelcder & ServiUe, 1825 (^Megischus onntttoor BruUé). Designated by Viereck, 1914:91. H ead,- (Figs 33,39) Head globose or almost so (as Rgs 27,33). Paraocular strigation absent. Face pilostty in between antennal foramina until clypeal suture absent, or as in frons. Granular sculpture on coronal area and frons absent: polished smooth in between macrosculpture, or present: powdered or granular aspect in between macrosculpture, or enthely granular with macrosculpture absent. Vertex sculpture often areolate-rugose latero-anteriorly, transverse centralfy, polished smooth on temples, but many different patterns occur; sculpture normaUy very coarse, but sometimes regular or very fine. Posterior m ar^ of head seen dorsaUy deeply convex (as Fig. 34), or weakly convex or straight (Fig. 33). Gena with a yeUow spot basaUy, at or near malar space, which is fidht if head has a Ught color, or with diSUse yeUow basaUy, usuaUy extending near m ai#i fi'om gena to firce and ffons, or gena mostly or entirely difiuse yeUow, sometimes similar to a strips but restricted to its ventral half Occipital carina ending very close to, at, or «(tending alongside hypostomal carina, reaching its apex (Rgs 38,39). 185 Post-genal bridge sinkmg immediately behind hypostomal carina, even if doing so only centrally; post-genal line, in consequence, entirely inclined in relation to sagital plane o f oral fossa (as Fig. 38), or forming a flat plataform behind hypostomal carina, which is paralld to sagital plane of oral fossa, then sinkmg towards forame magnum; post-genal line; in consequence, initially straight, horizontal, then bent towards foramen (as Figs 39, 40). Maxillary palpus distinctly longer than head is laterally high ^ig s 29, SO); rarely approximately as long or much shorter than head is laterally high. Prothorax.- Pronotum (Figs 56,60,61, 63,64): Apical sensorial hairs of pronotum organized lineariy, m a single line or narrow stripe, even d* only a few or very small (also Fig. 79), or organized in a tuft or small area (Figs 60, 63), or inconspicuous or absent. Apical margm of pronotum a simple rim, slightly to moderately refleced upwards, so that it is L- or C-shaped m cross section. Apical margin of pronotum distinctly and strongly emarginate, C-shapped, although sometimes small, restricted to about 0.3-0.4 of the apical width, occupying central part only (Fig. 56), more rarely weakly angled (as in Fig. 54). Colo normal, about 0.5-1.0 as long as wide; with transverse, complete or mcomplete carinae, which are strong often with at least one pair, or one complete carina, vertically raised as a blade-like and weakly translucent structure (as those m Figs 63,73); rarely with oblique, longitudinal, straight carinae, converging to inside pronotal fold (as in Fig. 82), or with one strong centro-longitudinal carma (Fig. 56); centro longitudinal depression absent or present. Pronotal fold normally distmct and distinclty, often deeply, concave anteriorly, never collapsed centrally; but sometimes shallow and transverse, or absent (as Fig. 60). Transverse ruga and groove on prenannular absent, rarely distinct. 186 Preannularm lateral view isolated from semiannular by a small step, or simply by a more swollen and differently sculptured semiannular, rarely perfectly continuous with semiannular or absent Prepectus exposed ^ig. 85), or mostly or complete^ covered by postero-ventral apex of pronotunt Angle between apical margins of hemistemites of prostemum about 90" or more (as Fig. 83), or distinctly less than 90" (as Hg. 84). Front coxa laterally with several small foveolae ^ig. 85). Meso ondMetathorax.- Mesoscutum large and round, from smaller to distinctly larger than combined area of axillae OFig- 95; see also Figs 93, 96), rarely hyperbolic (as Fig. 98). Dorsal part of mesepistemum mostly or entirely unifbrmly pilose or pruinose, but hairs moderately sparse to very sparse, so that 50-95% of cuticular surfece is visible; rarely glabrous, or with a patch of dense white piloshy on area in front of dorso apical mga (Fig. 106). Pminosity on mesepistemum, excluding pectum and dorsal part, absent or nearly so, at most with a small amount of sparse prumosity antero-dorsally (as Fig. 103), or occurring on 90-100% of its area, and moderately to densely abundant, concealing 50-95% of surfece, or restricted to antero-ventral part (Fig. 100; M fu rc a tu s only). Mesopleural pit absent Mid tibia without spm% on anterior fece. Spiracular impression and carma independent of line of fusion between metapleuron and propodeum; both straight ending on, or pointing to, flank, or spfracular impression curving inwards apically; spiracular carina sometimes not developed, givmg place to an area or plateau between spiracular impression and line of fusion of metapleuron and propodeum, this area progressively wider towards crenulate sulcus ^ g , 105), or carina or Ime of fusion of metapleuron and propodeunt endmg below (t.e., externally to) flank, at pleuropropodeal 187 fovea; spôacular impression usually absent, if present, extending alongside spiracular carina (as Fig. 104). Ventral margm of post-foveolar area basally produced into a small or very prominent lobe with sharp edge, which covers a little of the base of the mid coxa (Kgs 105, 106). fotercoxal space between hind coxae membranous (Fig. 108). Hind coxa basally deeply concave on external side only (as Figs 114,118), rarely deeply concave externally and mesally (as Figs 115,119); shorter than maximum lateral length of mesepistemum, or as long as, or slightly longer than maximum lateral length of mesepistemum (up to l.Ix as long); shape globular, barrel-like, subconic, or moderately elongate (as Figs 116,117). Apico-dorsal spine on hind coxa absent. Mesal glandular opening basally on hind fomur absent. Ventral edge of hind femur basad of central tooth without teeth or tubercles of any kind or with two or more teeth, or several small tooth-like tubercles; o r with small regular tubercles, as the ones in between central and apical teeth (as Figs 130,131,132). Hind tibia laterally compressed on basal G.3-0.5, then laterally from moderately to intensely compressed or “pinched,” then dilated ^ig . 135); more rarely as in Fig. 93. Ventral longitudinal oirina. on hind tibia restricted to compressed part, although often advancmg a little over dilated part (as Fig. 141), or extending along entire length o ftibia, conspicuous from base to apex, albeit usually as a sharp edge on compressed part, and as a carina on dilated part; sometimes weak on dilated part, disappearing before reaching apec (Fig. 142). Hnd tarsus of female with three tasomeres. Hind basitarsus ventrally with 1-2 rows of lon^ translucent spines, or at least setae, which are bigger basally, getting smaller and weaker towards apex (as Kg. 146 or more delicate). 188 Front Wtng.^ (Figs 165-167) Costal cell with very small and sparse but conspicuous setae, usually more frequent centrally or apically. Setae on M+Cu reaching base o f IM and usually advancing over ICu m at least one of the wings (Figs 155, 158), or ending near or at, but not reaching, base of IM, from which the most apical seta is separated by a distance always shorter than length of Icu-a ^ g s 157, 159). Parastigma distinctly longer, or at least approximately as long as vein IRs, and distinctly longer than wide, or shorter than vrâi IRs, and longer than wid^ or much shorter than IRs, or absent; when developed, usually as long as wide, or wider than longer. Vein IM usually at about 90° with IRs, making cell R truncate apically; more rarely forming a distinctly obtuse angle, or aligned with IRs, or nearly so, making cell R apically acutely triangular, distinctly longer to much longer than IRs (observed; 1.2-6.5x); originating at or basad Icu-a, so that nodus formed by fusion of M+Cu/IM with Icu-a/lCu is backwardly oblique, or originating apical to Icu-a, so that it is forwardly oblique; straight, or slightly curved, or distinctly and entirely curved. Vein Icu-a straight or nearly so, or slightly convex toward base of wing; at about 90° with both M+Cu and ICu, or somewhat inclined towards wing base, forming an angle of little less than 90° with M+Cu, and a correspondent slightly obtuse angle with ICu. Vein ICu entirely tubular. Vein 2-1A straight; tubular on 0.9 or more o f its length, or shortly tubular basally, remaming mostly or entirely nebulous. Vein 2Cua entirely tubular, or entirely and distinctly nebulous; at about 90° with ICu, or making a distinct, sometimes intensely acute angle with ICu. Vein 2Cub distinct; strongly nebulous often tubular basally, or nebulous, or partially and weakly nebulous rest spectral, or entirely spectral. Often forked apically, or apically not 189 foriced. Vem 2cu-a semi-tubular o r nebulous, or spectral or absent. Vein ^s+M )b disconnected from 2+3Rs by presence of a bulla, or connected and contiguous with 2+3Rs; disconnected from 2+3M by presence of a bulla, or connected and contiguous with 2+3M; distinctly short, or moderately long. Spurious vein arising from (Rs+M)b absent Vein 2+3Rs straight or slightly curved, tubular. Veins 4Rs and 2r fbrming a distinct angle with each other, or at least notably discontinuous, or perfectly continuous with one another, not forming an angle. Vem 4Rs longitudinal, but usually neither perfectly straight, nor unifbrmly curved, if straight, not parallel to pterostigma, and usually slightly inclined anteriorly, or entirely and unhbrmly curved towards anterior margin, even if slightly, or nearly perfectly straight and parallel to pterostigma; more than 3x as long as 2r, up to 1 Ix, or from as long as, to not more than 3x as long as 2r. Vein 2+3M tubular on basal 0.8 or more, but never entirely, with apex nebulous or spectral. Area of cell R l up to 2x as large as area of cell IM. Retinaculum mostly under cell 2Cu, its apex nearest to level of vein 2Cua. Hind Wing.- ^ g s 181-184) Hamuli usually 3-4, but up to 7 Type I hamuli m at least half of the specimens (Fig. 149), large or very large specimens often with 2-3 Type H hamuli, or only 2 Type I, and 1 Type H hamuli, very large specimens with no more than 3 Type I or, more rarety, 2 Type XL Vein MKZu distinct, oblique, usually nebulous, but sometimes spectral, often continued by, or indiscernible from IM. Vein Rs usually nebulous but sometimes spectral; long, straight, and longitudinal, its origin distinctly unrelated to Sc+R. Vein Ir-m present, even if very 6int, better developed in large specimens, or absent Vein Cua absent, or present, nebulous or spectral, or indicated by a 190 nd)ulous stub coining out of M-K ^ Dorsal wing fold present complete although somethnes difiScult to seem small specunens; Ir-m, if present, interrupted by dorsal fold, or absent; Ir>m, if present, entire not interrupted. Ventral wing fold present at least basally, although sometimes difiScult to see in small specimens, or absent. Wing membrane basally normal or weakly concave posteriorly, wing distinctly not pedunculate, or round, wid^ posterior margin convex. Wing apically round and broad, or narrowed or distinctly pointy; knüè-shaped, or shnmetrically acute-triangular. Abdom en.^ Second tergite and stemite fused mto a qrlindric and elongate petiole (Fig. 145); 2x or more as long as dorsal length of T3. Ovipositor sheaths white- or yellow-banded subapically; or unicolorous, sometimes progressively darker towards apex. Genus Hendst^hanus Enderiein (part) Enderlein, 1905:474-5. HemistephamsEiiàisA&m^ 1906:291,301-6; BCiefifer, 1908:3,4; Elliott, 1917b: 130; 1922:710,713,760-777; Bmes & Melander, 1932:476; Orfila, 1956:6; DeSantis, 1980:8; Aguiar, 1998. Stephcams(HemistepfKams)-i^taaxiy 1917. M egischus Towne% 1949: 363; Carlson, 1979:741; Gauld, 1995: 181-4; Belokobylskij, 1995:22. Parastephanellus(Hemestephams) [sicl]: Narendran & Sheela, 1995:45. Type species: StephemusmcuanarusSciAsXXgxeCr by original designation. 191 H ead.- Head globose or almost so (as Fig. 27). Paraocular strigation absent. Face pOoshy in between antennal foramina until clypeal suture very abundant, hairs usually much shorter than on firons (Fig. 31). CSranular sculpture on coronal area and firons absent: polished smooth in between macrosculptur^ or present: powdered or granular aspect in between macrosculpture, or entirely granular with macrosculpture absent. Vertex sculpture varied, but usually partially transverse, except laterally, or at least antero-Iaterally, where any other orientation (accept areolate-rugose) may occur, mostly or entirely parallel ondulate-rugose, or zig-zag rugose, or two or more different sculpture patterns, approximately in the same proportion (Fig. 33); sculpture intensity regular or very fine, or sculpture very coarse. Posterior margin of head seen dorsally weakly convex or straight (as Fig. 33). Gena without yellow or white colors. Occipital carina ending very close to, at, or actending alongside hypostomal carina, reaching its apac (as Figs 38,39), or disappearing for fi’om hypostomal carina (as Fig. 40), or narrowing gradually and disappearing, with both sides converging without ever meeting ^ g . 41). Post-genal bridge sinking Immediately behind hypostomal carina, even if doing so only centrally; post-genal line, in consequence, entirely inclined in relation to sagital plane of oral fossa (as Rg. 38), or forming a fiat plataform behmd hypostomal carma, which is parallel to sagital plane of oral fossa, then sinking towards forame magnum; post-genal line, in consequence initially straight, horizontal, then bent towards foramen (as Figs 39,40), or sunlq with gena folding partially over It, generating a deep shaft ^ g . 41). Maxillary palpus distinctly longer than head is laterally high (as Fig. 29). 192 Prothorar.- Pronotum ^ g s 59,62); Apical sensorial hairs of pronotum inconspicuous or absent, or organized Imeariy, in a single line or narrow stripe, even if only a few or very smaü (as Rg. 63). ^ ic a l margin of pronotum strongly reflexed upward^ generatmg a wide collar; more rarely a simple rim, slightly to moderately reflexed upwards, so that it is L- or C-shaped in cross section. Apical margin of pronotum distinctly and strong^ emarginate, C-shapped. Colo normal, about 0.5-I.0 as long as wid^ or very long usually 2.0x as long as wide or more, but at least l.Sx; with transverse complete or incomplete carinae, which are strong often with at least one pair, or one complete carina, vertically raised as a blade-like and weakly translucent structure (as those on Figs 63 and 73), or carinae mostly complete and at least anteriorly organized In an U-pattem, so that th ^ are backwardly oblique when seen in lateral view; dorsally somethnes straight, transverse; centro longitudinal depression absent or present. Pronotal fold distmct and often deeply concave anteriorly, never collapsed centrally; or shallow and transverse, indicated by posterior end of centro-longitudinal depression on colo, not concave anteriorly. Transverse ruga and groove on prenannular usually distinct. Preannular m lateral view isolated ftom semiannular by a small step, or simply by a more swollen and dKferently sculptured semiannular. Prepectus mostly «cposed (as Fig. 85), or mostly or completely covered by postero-ventral apex of pronotum. Angle between apical margins of hemistemites o f prostemum distinctly less than 90” (as Fig. 84). Front coxa laterally with several small foveolae (as Fig 85). Meso andMelathorax,^ Mesoscutum hyperbolic, large (Fig. 98), or slightly transverse; neariy as large as, or slightly larger than combmed area o f axillae (as H g 96). 193 Dorsal part of mesepistemum mostly or entirely uniformly pilose or pruinose, but hairs moderately sparse to very sparse; so that 50-95% of cuticular surâce is visible. Pruinosity on mesepistemum, mccludmg pectum and dorsal part, occurring on 90-100% of its area, and moderately to densely abundant, concealing 50-95% of surAce. Mesopleural pit absent. Mid tibia without spines on anterior Ace. Spiracular impression and carina mdependent of Ime of fusion between metapleuron and propodeum; both straight, ending on, or pointing to, flanl^ or spiracular impression curving inwards apically; spiracular caiina sometimes not developed, giving place to an area or plateau between spiracular impression and line of fusion of meApleuron and propodeum, this area progressively wider towards crenulate sulcus (Fig. 105). Ventral margin of post-foveolar area basally produced into a small or very prominent lobe with sharp edge, which covers a little of the base of the mid coxa (as Figs 105,106). Intercoxal space between hind coxae membranous (as Fig. 108). Hind coxa basally deeply concave on «eternal side only QFigs 114, IIS); as long as, or slightly longer than maximum lateral length o f mesepistemum (up to I.Ix as long); shape from barrel-like to moderately elongate (Figs 116, 118). ^ico-dorsal spine on hind coxa absent. Mesal glandular opening basally on hind femur absent. Ventral edge of hind Amur basad of central tooth without teeth or tubercles of any kind; more rarely with some small tooth-like tubercles CFig. 130). Hind tibia laterally compressed on basal G.3-0.5, then with a weak depression «eternally, and a correspondent deep semicircular depression mesally, its bordering carina or edge extending from the ventro-mesal side to the dorsal side, where it is oblique (Fig. 136); more rarely as in Fig. 134. Ventral longitudinal carina on hmd tibia restricted to compressed part, although often 194 advancmg a Ihtle over dilated part ÇFig. 141). (find tarsus of female with three tasomeres. Hind basitarsus ventrally with 1-2 rows of short, opaque, conical spines, which get smaller and less distinct towards apac. Front Wmg.- (Hgs 168,169) Costal cell with very small and sparse but conspicuous seta^ usually more frequent centrally or apically. Setae on M+Cu placed frr from IM, from which they are separated by a distance greater than length of Icu-a (Fig. 156). Parastigma shorta- than vein IRs, and longer than wide, or much shorter than IRs, or absent; when developed, usually as long as wide, or wider than longer. Vein IM usually at about 90° with IRs, making cell R truncate apically; more rarely forming a distinctly obtuse angle; distinctly longer to much longer than IRs (observed: 1.2-6.5x); originating at or basad Icu-a, so that nodus formed by fusion ofM+Cu/lM with Icu-a/lCu is backwardly oblique, or originating apical to Icu-a, so that it is forwardly oblique; straight, or slightly curved. Vein Icu-a straight or nearly so, or slightly convoc toward apex of wing somewhat inclmed towards wing base, forming an angle o f little less than 90° with M+Cu, and a correspondent slightly obtuse angle with ICu, or strongly inclined towards wing base, angled at much less than 90° (to a minimum o f 45°) with M+Cu. Vein ICu entirely tubular. Vein 2-1A straight; very shortly tubular basally, forming a “stub,” which sometimes is briefly nebulous apically; remaining absent of spectral; or entirely spectral. Vem 2Cua enthely tubular; makmg a distmct, sometimes intensely acute angle with ICu, or at about 90° with ICu. Vein 2Cub partially and weakly nebulous, rest spectral, or enthely spectral, or nebulous. Apically not forked. Vein 2cu-a spectral or absent. Vein ^IsfNQb disconnected from 2+3Rs by presence of a bulla; disconnected from 2+3M by 195 presence o f a bulla; distinctly sh o rt Spurious vein arising fi-om (Rs+NQb absent Vein 2+3Rs straight or slightly curved, tubular. Vems 4Rs and 2r perfectly continuous with one another, not forming an angle. Vein 4Rs entirely and uniformly curved towards anterior margin, even if slightly; more than 3x as long as 2r, up to I Ix, or from as long as, to not more than 3x as long as 2r. Vein 2+3M tubular on basal 0.8 or more, but never entirely, with apex nebulous or spectral. Area of cell RI up to 2x as large as area of cell IM. Retinaculum midway between cell ICu and cell 2Cu, its apex nearly equidistant from Icu-a and 2Cua. Hind Wing.- (Figs 185) Hamuli only 2 Type I, and I Type H hamuli, very large specimens with no more than 3 Type I or, more rarely, 2 Type H, or usually 3-4, but up to 7 Type I hamuli in at least half of the specimens OFig. 149), large or very large specimens often with 2-3 Type H hamuli. Vein M-hCu absent Vein Rs nebulous or spectral; short originating near or at apex of Sc^R, usually curving towards anterior margin, more rarely straight or somewhat undulate. Vein Ir-m absent. Vein Cua absent. Dorsal wing fold present complete, although sometimes difficult to see in small specimens; Ir-m, if present mterrupted by dorsal fold, or absent Ir-m, if present entire, not interrupted. Ventral wing fold absent Wing membrane basally very narrow, for a short distance approximately as narrow as width of vein Sc+R, making the wing distinctly pedunculate. Wing apically narrowed or distinctly pointy; knifb-siuq>ed, or shnmetrically acute-triangular. A b dom en- Second tergite and stemite fused into a cylmdric and elongate petiole (Fig. 145); 2x or more as long as dorsal length of T3. Ovipositor sheaths white- or yellow-banded subapically. 196 Genus Parasfephanettus Endcriein Stephanus (p 2o i): Westwood, 1874:126. Pco’ostephanusEixdeA&xi (preoccupied by Hâckel forProtozooa), 1905:474. PcarastephaneUusBDàeA&Oy 1906:191,301; Kiefkr, 1908:3,5; Elliott, 1922: 710,713, 746-60; 1926:521; 1928a; CebaÜos, 1926:142; Dutt, 1926:5; Orfila, 1956:6; Baltazar, 1962:744-5; 1966:17; Beiokobylskij, 1995:18, 22. (misidentifed): Moore, 1961:93. Parastephanellus (Parastephanellus): Narendran & Sheela, 1995:45. Type: Stephanaspyg^eusEni&ûàxi, designated by Enderlein (1906). H ead ,- Head globose or almost so (as Fig. 27). Paraocular strigation distinct (Fig. 2). Face püosity in between antennal foramina until clypeal suture absent, or as in foons. Granular sculpture on coronal area and frons absent: polished smooth in between macrosculpture, or present: powdered or granular aspect in between macrosculpture, or entirely granular with macrosculpture absent. Vertex sculpture mostly or entirely transverse even if intensely ondulate or if vertex polished smooth posteriorly; two or more dKforent sculpture patterns, approximately in the same proportion, or mostly or entirely transversely strigate or strigulate, or mostly rugose, rugulose, or areolate-rugose; sculpture mtenshy regular or very fine. Posterior margm o f head seen dorsally deeply convex (as Fig. 34), or weakly convex or straight (as Fig. 33). Gena with a well defined, yellow or ivory stripe near ^ e margm, extending foom base of mandible to temple. Occipital Carina ending very close to, at, or extending alongside hypostomal carina, reaching its ap&c (as Figs 38,39). Post-genal bridge forming a flat plataform behind 197 hypostomal caruia, is parallel to sagitai plane of oral fossa, then sinking towards ferame magnum; post-genal line, m consequence^ mitiany straight, horizontal, then bent towards foramen (as Figs 39,40). Maxillary palpus distinctly longer than head is laterally high(asRgs29). Prothorax.- Pronotum (Figs d66, 67): Apical sensorial hairs o f pronotum organized in a tuft or small area, ^ ic a l margm o f pronotum a simple rim, slightly to moderately reflexed upwards, so that it is L- or C-sh^ed in cross section, ^ ic a l margin of pronotum as deep as, or deeper than wide; more or less U-shaped, but margins of emargination diverging towards apex. Colo normal, about 0.5-1.0 as long as wide; carinae mostly complete, and at least anteriorly organized in an U-pattem, so that th ^ are backwardly oblique when seen in lateral view; dorsally sometimes straight transverse, or carinae straight, laterally distinctly oblique towards pronotal fold, dorsally interrupted, or at least distinctly weakened, rarely blade-like; centro longitudinal depression absent or present. Pronotal fold distinct, centrally collapsed, or at least closed, leaving two shallow or moderately deep openings on each side. Transverse ruga and groove on prenannular absent Preannular perfectly commuons with semiannular. Prepectus mostly or completely covered by postero-ventral apex of pronotum, or mostly exposed (as Fig. 85). Angle between apical margins ofhemistemites ofprostemum distmctly less than 90° (as Fig. 84). From coxa laterally without foveola^ or with several small fbveolae (as Fig. 85). M eso cm dM etathorax,- Mesoscutum slightly transverse; nearly as large as, or slightly larger than combined area of axillae (Rg. 96), or conspicuously transverse, and distmctly smaller than combmed area of axillae (as Figs 94,97,99). Dorsal part of 198 mesepistemum mostly or entirely uniformly pilose or pruinose, but hairs moderately sparse to very sparse so that 50-95% o f cuticular surfece is visible; or with pruinosity restricted to anterior G.3-0.5, showing moderately sparse to very sparse hairs, so that 50-95% of cuticular surfoce is visible, rest glabrous (as Fig. 103); or entirely glabrous. Pruinosity on mesepistemum, excluding pectum and dorsal part, restricted to anterior 0.3-0.5 (Fig. 101), often also restricted to dorsal 0.5-0.7 of this part; or nearly absent, at most with a small amount o f sparse pruinosity antero-dorsaUy (as Fig. 103); more rarely occurring on 90-100% o f its area, and moderately to densely abundant, concealing 50-95% of surfoce. Mesopleural pit absent. N£d tibia with a few spines or spine-like setae along anterior face, mclined toward apex (Fig. 112). Spiracular impression and carina independent of line of fusion between metapleuron and propodeum; both straight, endmg on, or pointing to, flank, or spiracular impression curving inwards apically; spiracular carina somethnes not developed, ghdng place to an area or plateau between spiracular impression and line of fosion of metapleuron and propodeum, this area progressively wider towards crenulate sulcus (as Figs 105,106); or carina or line of fusion o f metapleuron and propodeum, ending below Q.e., externally to) flank, at pleuropropodeal fovea, and spiracular impression usually absent, if present, extending alongside spiracular carina (as Fig. 104). Ventral margin of post-foveolar area straight, edge usually longitudinally carinate, not produced into a lobe basally (Bg. 107). Intercoxal space between hind coxae membranous (as Bg. 108). Hnd coxa basally deeply concave on external side only (as Bgs 114,118); as long a^ or slightly longer than maximum lateral length of mesepistemum (up to 1. Ix as long), or shorter than maximum lateral length o f mesepistemum; shape as in Bg. 116 or 199 moderate^ elongate. Apico-dorsal spine on hind coxa absent Mesal glandular opening basally on hind femur absent Ventral edge of hind femur basad o f central tooth with two or more teeth, or several small tooth-like tubercles; or with small regular tubercles, as the ones m between central and apical teeth (as Figs 130,131), or without teeth or tubercles of any kind. Hind tibia laterally compressed on basal 0.3-0.S, then with a weak depression externally, and a correspondent deep semicircular depression mesally, its bordering carma or edge extending from the ventro-mesal side to the dorsal side, where it is oblique (as Fig. 136). Ventral longitudinal carina on hind tibia restricted to compressed part, although often advancing a little over dilated part (as Fig. 141). Hind tarsus of female with three tasomeres. Hind basitarsus ventrahy with 1-2 rows of lon^ translucent spines, or at least setae, which are bigger basally, getting smaller and weaker towards apex (as Fig. 146 or more delicate). Front Wing.- ÇFig. 171) Costal cell setae completely absent, or few and microscopic, inconspicuous. Setae on Mh-Cu reaching base of IM and usually advancing over ICu in at least one of the wings (Figs 155, 158), or ending near or at, but not reachm& base of IM, from which the most apical seta is separated by a distance always shorter than length of Icu-a (Hgs 157,159). Parastigma distmctly longer, or at least ^proximately as long as vein IRs, and distmctly longer than wide, or shorter than vein IRs, and longer than wide. Vem IM usually at about 90* with IRs, making cell R truncate apically; more rarely ft)rming a distinctly obtuse angle; distmctly longer to much longer than IRs (observed: I.2-6.5x); originating ^ical to Icu-a, so that nodus formed by fusion of M-Khi/IM with Icu-a/ICu is ferwardly oblique, or originatmg at or basad Icu-a, so 200 that It is backwardly oblique; slight^ curved, or straight Vein Icu-a slightly convex toward apec of wing; somewhat inclmed towards wing base, forming an angle o f little less than 90“ with M+Cu, and a correspondent slightly obtuse angle with ICu. Vein ICu entirely tubular. Vein 2-1A straight; very shortly tubular basally, forming a “stub,” which sometimes is briefly nebulous apically; remaining absent o f spectral; or entirely spectral. Vein 2Cua partially tubular (basal 50% or less); apirally nd)ulous, even if strongly, appearmg to be tubular, or entirely tubular; making a distinct, but never greatly obtuse angle with ICu, or at about 90“ with ICu, or making a distinct, sometimes intensely acute angle with ICu. Vein 2Cub partially and weakly nebulous, rest spectral, or entirely spectral. Apically not forked. Vein 2cu-a spectral or absent. Vein (Rs+M)b connected and contiguous with 2+3Rs; disconnected fi-om 2+3M by presence of a bulla; very long, or moderately long. Spurious vein arising from (Rs+NQb absent. Vein 2+3Rs straight or slightfy curved, tubular. Veins 4Rs and 2r forming a distinct angle with each other, or at least notably discontinuous. Vein 4Rs entirely and uniformly curved towards anterior margin, even if slightly, or longitudinal, but usually neither perfectly straight, nor uniformly curved, ^ straight, not parallel to pterostigma, and usually slightly inclined anteriorly; more than 3x as long as 2r, up to 11?^ or from as long as, to not more than 3x as long as 2r. Vem 2+3M tubular on basal half or less, rest nebulous (somethnes strongly), or spectral, or tubular on basal 0.8 or more, but never entirely, with apoc nebulous or spectral. Area of cell RI nearly or more than 3x as large as area of cell IM, or up to 2x as large as area of cell IM. Retinaculum mostly under cell 2Cu, its apex nearest to level of vehi2Cua. 201 Hind Wing.- (Rg. 187) Hamuli only 2 Type ^ and 1 Type II hamuli, very large specimens whh no more than 3 Type I or, more rarely, 2 Type U, or usually 3-4, but up to 7 Type I hamuli in at least half ofthe specimens ^ g . 149), large or very large specimens often whh 2-3 Type H hamuli. Vein MfCu spectral, longitudinal, apparently fused to Rs; IM indistinct, or absent Vein Rs nebulous or spectral; short, originating near or at apex of ScfR, usually curving towards anterior margin, more rarely straight or somewhat undulate. Vein Ir-m absent Vein Cua absent Dorsal wing fold absent; Ir-m, present entire, not interrupted. Ventral wmg fold absent. Wing membrane basally very narrow, ft>r a short distance approximately as narrow as width of vein Sc+R, making the wing distinctly pedunculate. Wing apically narrowed or distinctly pointy; knife-shaped, or simmetrically acute-triangular. A bdom en.- Second terghe and stemite fused into a cylindric and elongate petiole ^ig. 145); 2x or more as long as dorsal length of T3. Oviposhor sheaths unicolorous, somethnes progressively darker towards apex. GenusJùmthostephanus n. gen. Stephanusiÿdx^'i Cameron 1912:358; Elliott 1922:716,724; Ceballos 1926:138; Benoh 1949:286,1950:266, 1984a: 216. Type specie: Stephams xanthocephanus Cameron, by monotypy and present designatioiL H ead.- Head globose or almost so (as Fig. 27). Paraocular strigation absent. Face pOoshy in between antennal foramina until clypeal suture absent, or as in ftons. Granular 202 sculpture on coronal area and frons absent: polished smooth in between macrosculpture. Vertex sculpture partially transverse, except laterally, or at least antero-Iaterally, where ary other orientation (except areolate-rugose) may occur; two or more diffèrent sculpture patterns, approximately m the same proportion ^ig . 33); sculpture intensity regular or very fine. Posterior margin of head seen dorsally weakly convoc or straight (Fig. 33). Gena with diffuse yellow basally, usually extending near ^ e margin firom gena to face and ffon^ or gena mostly or entirely diffuse yellow, sometimes similar to a stripe, but restricted to its ventral half. Occipital carina ending very close to, at, or extending alongside hypostomal carina, reaching its apex (as Figs 38,39). Post-genal bridge forming a fiat plataform behind hypostomal carina, which is parallel to sagitai plane of oral fossa, then sinking towards forame magnum; post-genal line, in consequence, initially straight, horizontal, then bent towards foramen (as Figs 39,40). Maxillary palpus distinctly longer than head is laterally high (as Rg. 29). ProthorcK.- (Fig. 65,68) Apical sensorial hairs of pronotum organized in a tuff or small area (see also Figs 76, 66). Apical margin of pronotum a simple rim, slightly to moderately reflexed upwards, so that it is L- or C-shaped in cross section. Apical margin of pronotum distinctly and strongly emargmate, C-shapped. Colo normal, about 0.5-1.0 as long as wide; carinae mostly complet^ and at least anteriorly organized in an U-pattem, so that they are backwardly oblique when seen m lateral view (as Rg. 67); dorsally sometimes straight transverse; centro longitudinal depression present, sometimes very weak. Pronotal Aid distinct; distinclty, offen deeply, concave anteriorly, never collapsed centrally. Transverse ruga and groove on prenannular absent. Preannular perfectly 203 continuous with semiannular. Prepectus mostly exposed (as Fig. 85). Angie between apical margins ofhemistemites of prostemum distinctly less than 90" (as Fig. 84). Front coxa laterally without foveolae. Meso andMetatfmrax.- Mesoscutum large and round, distinctly larger than combined area of axillae (as Figs 92,95). Dorsal part o f mesepistemum mostly or entirely unffî)rmfy pilose or pruinose, but hairs moderately sparse to very sparse, so that 50-95% o f cuticular sur&ce is visible. Pruinosity on mesepistemum, excluding pectum and dorsal part, occurring on 90-100% of its area, and moderately to densely abundant, concealing 50-95% o f surtiice. Mesopleural pit absent. Mid tibia with a few spines or spine-like setae along anterior fece, mclined toward apex (as Fig. 112). Spiracular carina a t line of fesion of metapleuron and propodeum, ending below Q.e., externally to) flank, at pleuropropodeal fovea; spiracular impression usually absent, if present, extending alongside sphacular carina (as Fig. 104). Ventral margin of post-fbveolar area basally produced into a small or very prominent lobe with sharp edge, which covers a little of the base of the mid coxa (as Figs 104,105,106). Intercoxal space between hind coxae covered by a heavily sclerotized plate, formed by the extension and connection of metastemum with the posterior angles of propodeum (as Fig. 111). Hmd coxa basally deeply concave on external side only (as Hgs 114, 118); distinclty longer than maximum lateral length o f mesepistemum (1.3-1.5x as long); shape very long and slender, 4.25x or more as long as laterally high (as Figs 119,120). Apico-dorsal spine on hind coxa absent. Mesal glandular opening basally on hind femur absent. Ventral edge of hind femur basad of central tooth whh two or more teeth, or several small tooth-like tubercles; or with small 204 regular tubercles, as the ones m between central and apical teeth (as Figs 131 ,132), I£nd tibia laterally compressed on basal 0.3-O.S, then with a weak depression externally, and a correspondent deep semichcular depression mesally, its bordermg carina or edge extending 6om the ventro-mesal side to the dorsal side^ where it is oblique (as Hg. 136), Ventral longitudinal carina on hind tibia restricted to compressed part, although often advancmg a little over dilated part (as Fig, 141), Hind tarsus of female with three tasomeres, (End basitarsus ventrally with 1-2 rows oflon& translucent spines, or at least setae, which are bigger basally, getting smaller and weaker towards apex (as Fig, 146, or more delicate). Front Wmg.~ (Fig, 170) Costal cell setae completely absent, or few and microscopic, inconspicuous. Setae on M+Cu placed 6 r from IM, from which th ^ are separated by a distance greater than length of Icu-a (Fig. 156), Parastigma much shorter than IRs, or absent; when developed, usually as long as wide, or wider than longer. Vein IM aligned with IRs, or nearly so, makmg cell R, apically acutely triangular; distinctly longer to much longer than IRs (observed: l,2-6,5x); originating at or basad Icu-a, so that nodus fr)rmed by fusion of M-fCu/lM with Icu-a/lCu is backwardly oblique; slightly curved. Vein Icu-a straight or nearly so; at about 90“ with both M+Cu and ICu. Vein ICu entirely tubular. Vein 2-1A strongly curved, reaching wing margin before middle of cell 2Cu; shortly tubular basally, remaining mostly or entfrely nebulous. Vein 2Cua partially tubular (basal 50% or less); apically nebulous, even ^ strong^, appearmg to be tubular; making a distinct sometimes intensely acute angle with ICu, Vein 2Cub distinct; strongly nebulous, often tubular basally, ^ically not forited. Vein 2cu-a spectral or absent. Vein 205 connected and contiguous with 2+3Rs; disconnected from 2+3M by presence of abulia; moderately long. Spurious vem arising from (Rs+N^b absent Vein 2+3Rs straight or slightly curved, tubular. Veins 4Rs and 2r forming a distinct angle with each other, or at least notably discontinuous. Vein 4Rs longitudmal, but usually nehher perfectly straight, nor unfrbrmly curved, ^ straight, not parallel to pterostigma, and usually slightly inclmed anteriorly; from as long as, to not more than 3x as long as 2r. Vein 2+3M tubular on basal half or less, rest nebulous (sometimes strongly), or spectral. Area of cell RI up to 2x as large as area of cell IM. Retinaculum mostly or entirely under cell ICu, its apex nearest to level of vein Icu-a. H in d Wmg.~ ^ig. 186) Hamuli usually 3-4, but up to 7 Type I hamuli in at least half of the specimens (Fig. 149), large or very large specimens often with 2-3 Type II hamuli. Vein M+Cu absent. Vein Rs nebulous or spectral; short, originating near or at apex ofSc+R, usually curving towards anterior margin, more rarely straight or somewhat undulate. Vein Ir-m absent Vein Cua absent. Dorsal wing fold absent; Ir-m, if present, entire, not interrupted. Ventral wmg fold absent. Wmg membrane basally normal or weakly concave posteriorly, wing distinctly not pedunculate. Wing apically narrowed or distmctly pointy; knife-shaped, or simmetrically acute-triangular. Abdomen.~ Second tergite and stemite fused into a ^lindric and elongate petiole (as Mg. 145); 2x or more as long as dorsal length of T3. Ovipositor sheaths unicolorous, sometimes progressively darker towards ap«c. 206 GenusF oenatopus Smith Stephams (pact): Westwood, 1841:274; 1874:127; Schletterer, 1889a: 121. M egiscfm s (pait): Westwood, 1851:230. F oenatopus Smith, 1861a: 58; Szépligeti, 1902:528; Kiefifer, 1904:1; 1905:478, 488; Enderlein, 1905:474,476; 1906:291,298; Kiefifer, 1908:6; Elliott, 1922:710,713, 782-800; CefaaUos, 1926:144; Elliott, 1926: 525; 1928b: 79; Orfila, 1956:6; Benoit, 1956:205; Baltazar, 1962:744-5; 1966: 19; Mateu, 1972:601; Medler, 1980:443; DeSantis, 1980: 10;Madl, 1991:118, 123-4; Beiokobylskij, 1995: 18; Aguiar, 1998:364. Foenatopus \ûc\\: Tobias, 1993:391. Neostephanus'KieSxx, 1904:1-4; 1905:478; Enderlein, 1905: 474,476; 1906: 291, 299-301; Kieffer, 1908:6-7; Elliott, 1922: 713, 777-81; Ceballos, 1926: 143; Orfila, 1956: 6; 1956:6; Medler, 1980:443. DiastephanusEndedtiSiy 1905:474,476, New Synonymy; 1906: 291-8; Kiefier, 1908:6-7; Viereck, 1914:44; Eüiott, 1922: 710,713,800-827; Ceballos, 1926:146; Elliott, 1926:525; 1927b: 349; Brues & Melander, 1932:476; Orfila, 1956:6; Medler, 1980:442. MaetegcrfoenusBmoxt, 1951a(part): 269,278, New Synonymy; 1956:205. Type species: Stephanus indicus Westwood, by monotypy. In the original description. Smith (1861a:58) proposes a new specie^ F. n ificep s, and the generic transfer o fA feg tsch u s indicus (Westwood) to F oem topus, These two species, however, are ^onym s (Schletterer 1889). 207 H ea d ,- Head globose or almost so ^îgs 27,37), or weakly but distmctly compressed antero-posteriorly 2&), or strongly compressed antero-posteriorly (Hg. 29). Paraocular strigation normally absent, rarely distinct Face pilosity m between antennal foramina until clypeal suture absent or asm frons. Granular sculpture on coronal area and foons present: powdered or granular aspect in between macrosculpture, or entire^ granular whh macrosculpture absent or vestigial, or absent: polished smooth in between macrosculpture. Vertex sculpture mostly or entirely transverse, even if intensely ondulate or if vertex polished smooth posteriorly, or partially transverse, except laterally, or at least antero-laterally, where any other orientation (except areolate-rugose) may occur, or mked rugose and areolate-rugose, hnperfoctly transverse, interrupted by transversely carinate central impression; mostly or enthely transversely strigate or strigulate (Hg. 27), or mostly or entirely rugose, rugulose, or areolate-rugose, or two or more different sculpture patterns, approximately in the same proportion; sculpture intensity regular or very fine; sculpture rarely very coarse. Posterior margin of head seen dorsally angled, as an inverted V (Fig. 36), even if base is truncate (Hg. 35), or deeply and regulariy convex (Hg. 34). Gena whh diffese yellow basally, usually extendmg near tye margin from gena to fece and fi’ons, or gena mostly or entirely dififese yellow, sometimes similar to a stripe, but restricted to hs ventral balÇ or with a yellow spot basally, at or near malar space^ which is feint ^head has a light color, or whhout yellow or whhe colors, or whh a well defined, yellow or ivory stripe near tye margin, extending fiom base of mandible to temple. Occipital carma ending very close to, at, or extending alongside hypostomal carina, reachmg hs ap«c (as Hgs 3S, 39), or disappearing fer from hypostomal 2 0 8 carma 40), or fading gradually with both sides converging toward each other, or toward center of hypostomal carina, without ever meeting (similar to Fig. 41). Post-genal bridge forming a flat plataform behind hypostomal carina, which is parallel to sagitai plane of oral fossa, then smking towards forame magnum; post-genal line, in consequence, initially straight, horûrontal, then bent towards foramen ^ g s 39,40). Maxillary palpus distinctly longer than head is laterally high (Fig. 29). Prolhorax.- Apical sensorial hairs of pronotum inconspicuous or absent, or organized Imearly, in a single line or narrow stripe, even if only a few or very small (Fig. 79), or organized in a tuft or small area O^igs 73,76). Apical margin of pronotum laterally broadly flat, vertical, changing to a narrow central margin reflexed upwards (Figs 73,72, 78,69, 70,71, 75,76,74, 77), or a simple rim, slightly to moderately reflexed upwards, so that it is L- or C-shaped in cross section (Figs 77,79). Apical margin of pronotum deep, U- or V-shaped (Figs 69,71,76,73,79,80), or as deep as, or deeper than wide; more or less U-shaped, but margins of emargination diverging towards apec (as Kg. 66). Colo very lon% usually 2.0x as long as wide or more, but at least l.Sx ^ g s 69, 71, 80), or normal, about 0.5-1.0 as long as wide (Figs 73,76, 79); entirely transversely strigate, never oblique, most or all strigae complete, often weak, never with too strong or blade-like (Figs 69-71,74-80); or carinae laterally distinctly oblique towards pronotal fold, dorsally mterrupted or weakened ^ g s 78,82); rarely with carina vertically raised as a blade-like and weakly translucent structure ^ g s 73,72); centro longitudinal depression present, sometimes very weak ^ g s 73,69,71,76), or absent (Kgs 79,80,81, 82). Pronotal fold absent (Kgs 69-71,75,77-81), or distinct shallow or deep (Figs 73,72,74, 209 76,82), rarely centrally coll^sed or closed (as Fig, 66), leaving two shallow or moderately deep openings on each side. Transverse ruga and groove on prenannular normalfy absent; distinct in South American species only. Preannular in lateral view isolated fix)m semiannular by a small step, or simply by a more swollen and diflTerently sculptured semiannular (Figs 70, 74,75,77,78), or perfectly continuous with semiannular ^ g . 72). Prepectus mostly exposed (as Fig. 85), or mostly or completely covered by postero-ventral apex of pronotum. Angle between apical margins ofhemistemites of prostemum distinctly less than 90* ^ig . 84), rarely about 90" or more (as Fig. 83). Front coxa laterally without fbveolae. Meso and Metathorax.- Mesoscutum conspicuously transverse, and distinctly smaller than combmed area of axillae (Figs 94, 97, 99), or slightly transverse; nearly as large as, or slightly larger than combined area of axillae (as Kg. 96). Dorsal part of mesepistemum with pruinosity restricted to anterior 0.3-0.5, showing moderately sparse to very sparse hairs, so that 50-95% of cuticular surface is visible, rest glabrous (Rg. 103), or mostly or entirely uniformly pilose or prumose, but hairs moderately sparse to very sparse, so that 50-95% of cuticular surâce is visibly or glabrous; more rarely as in Figs 104,106. Pminosityon mesepistemum, excluding pectum and dorsal part, restricted to anterior 0.3-0.5 ^ g . 101), often also restricted to dorsal 0.5-0.7 of this part; or nearly absent, at most with a small amount of sparse pruinosity antero-dorsally (Fig. 103). Mesopleural pit present, very small, but always accompanied by characteristic transverse or oblique rugosities ^ g s 101,102,103). Nfid tibia with a &w spines or spine-like setae along anterior 6ce, inclmed toward apex (as Kg. 112). Spiracular carina a t line o f fusion 210 of metapleuron and propodeum, endmg below 0.e., externally to) flank, at pleuropropodeal fovea; spiracular impression usually absent, if present, exten<&ig alongside spiracular carina ^ig . 104). Ventral margin of post-fbveolar area basally produced into a small or very prominent lobe with sharp edge, which covers a little o f the base o fthe mid coxa ^ ig s 104, 105, 106, 109). Intercoxal space between hind coxae covered by a heavily sclerotized plate, formed by the extension and connection of metastemum with the posterior angles of propodeum (Fig. 111). Hind coxa basally deeply concave externally and mesally (Figs US, 119); rarely deeply concave on external side only (Foenatopus arcuatus); as long as or distinclty longer than maximum lateral length of mesepistemum (1.0-1 5x as long), rarely shorter, shape very long and slender, 4.25x or more as long as laterally high (Rgs 119,120), or moderately elongate (as Fig. 118). Apico-dorsal spine on hind coxa absent. Mesal glandular opening basally on hind femur normally present and variedly developped (Figs 124,122,121); more rarely as a punctual depression (t.e., not a tm e orifice) (Fig 125), or absent Ventral edge of hind femur basad of central tooth with several small tubercles; or with small regular tubercles, as the ones m between central and apical teeth ^ g s 130, 131, 132); sometimes absent in very small specimens. Hind tibia laterally compressed on basal 0.3-0.5, compressed also mesally and ventrally where dilated part meets compressed part generating a ventro-longtudinal carma or edge; not compressed «eternally; dorso-central carina absent (Fig. 138). Ventral longitudinal carina on bind tibia restricted to compressed part although often advancing a 211 little over dilated part (as Rg. 141). Hind tarsus of female with three tasomeres. Hind basitarsus ventrally with 1-2 rows of lon^ translucent spines, or at least seta^ which are bigger basally, gettmg smaller and weaker towards apex (as Fig. 146 or more delicate). Front Wing:- (Figs 172-178): Costal cell setae completely absent, or few and microscopic^ inconspicuous. Setae on M+Cu ending near o r at, but not reaching base of IM &om which the most apical seta is separated by a distance always shorter than length of Icu-a (Figs 157, 159), or placed 6 r &om IM, from which thqr are separated by a distance greater than length of Icu-a (Fig. 156). Parastigma much shorter than IRs, or absent; when developed, usually as long as wide, or wider than longer, or distinctly longer, or at least approximately as long as vein IRs, and distinctly longer than wide. Vein IM aligned with IRs, or nearly so, making cell R apically acutely triangular, nearly as long as, or distinctly shorter than IRs (observed: 0.3-l.lx), or distinctly longer to much longer than IRs (observed: 1.2-6.5x); originating apical to Icu-a, so that nodus formed by fusion of M+Cu/IM with Icu-a/lCu is forwardly oblique, or originating at or basad Icu-a, so that it is backwardly oblique; slightly curved, or straight. Vein Icu-a apex distinctly bent towards the base o f the win& or slightly convex toward apex of win& or straight or nearly so; somewhat inclined towards wing base, forming an angle of little less than 90" with M+Cu, and a correspondent slightly obtuse angle with ICu, or strongly mclined towards wmg base, angled at much less than 90" (to a minimum o f 45°) with M+Cu. Vein ICu spectral or nebulous, but sometimes with a short tubular stub basally, or tubular on basal 20% or more, but distmctly less than 100%, with apex nd)ulous or spectral, or entirely tubular. Vein 2-1A curving but not strongly, toward posterior margm of wm^ reaching 212 margin opposite to, or a little before, 2Cua, or strongly curved, reaching wing margin before middle of cell 2Cu, or straight; very shortly tubular basally, forming a ''stub," which somethnes is briefly nebulous apically; remaining absent o f spectral; or entirely spectral, or shortly tubular basally, remainmg mostly or entirely nebulous. Vein 2Cua &intly nebulous or spectral, or partially tubular (basal 50% or less); apically nebulous, even if strongly, appearmg to be tubular; making a distinct, but never greatly obtuse angle with ICu, or at about 90" with ICu, or makmg a distinct, sometimes intensely acute angle whh ICu. Vem 2Cub partially and weakly nebulous, rest spectral, or entirely spectral, or nebulous. Apically not forked. Vein 2cu-a spectral or absent, or semi-tubular or nebulous. Vein (Rs+M)b and 2+3Rs spectral or nebulous; 2+3M spectral, or connected and contiguous whh 2+3M; moderately long, or very Ion& or distinctly short. Spurious vein arising from ^LyfM)b present, normally spectral, but sometimes nebulous, never tubular, or absent Vein 2+3Rs strongly sinuate, spectral, more rarely nebulous. Veins 4Rs and 2r forming a distinct angle whh each other, or at least notably discontinuous. Vein 4Rs with a short horizontal basal portion, or at least represented by a knob, then forked, with anterior branch tubular, bent towards anterior wing margin, and ventral branch spectral (rarely nebulous) and half as long or as long as anterior branch, or longitudinal, but usually neither perfectly straight, nor unffî)rmly curved, if straight, not parallel to pterostigma, and usually slightly inclined anteriorly, or nearly perfectly straight and parallel to pterostigma; from as long as, to not more than 3x as long as 2r, or more than 3x as long as 2r, up to I Ix. Vem 2+3M entirely nebulous or spectral. Area of cell RI up to 2x as large as area of cell 1\^ or neariy or more than 3x as large as area of cell IM Retinaculum midway 213 between cell ICu and cell 2Cu, its apex nearly equidistant &om Icu-a and 2Cua, or mostly or entirely under cell ICu, its apex nearest to level of vein Icu-a, or mostly under cell 2Cu, its apex nearest to level o f vein 2Cua. Hirui Wmg,- ÇFîgs 188,190-194): Hamuli only 2 Type I, and 1 Type II hamuli, very large specimens with no more than 3 Type I or, more rarely, 2 Type H, or usually 3-4, but up to 7 Type I hamuli in at least half of the specimens ^ig . 149), large or very large specimens often with 2-3 Type II hamuli. Vein M+Cu absent. Vein Rs nebulous or spectral; short, originating near or at apex of Sc^R, usually curving towards anterior margin, more rarely straight or somewhat undulate, or absent. Vein Ir-m absent. Vein Cua absent Dorsal wing fold absent; Ir-m, if present entire, not interrupted. Ventral wing fold absent. Wing membrane basally very narrow, for a short distance approximately as narrow as width of vein Sc+R, making the wing distinctly pedunculate, or normal or weakly concave posteriorly, wing distmctly not pedunculate. Wing apically narrowed or distmctly pointy; knife-shaped, or simmetrically acute-triangular. Abdomen,- Second tergite and stemite fused into a cylmdric and elongate petiole (Rg. 145); 2x or more as long as dorsal length of T3. Ovipositor sheaths white- or yellow-banded subapically, or unicolorous, sometimes progressively darker towards apex; more rarely amost entirely yellowish-brown then suddenly black or dark brown at apex. Relationships of Stephanidae with other Hymenoptera Hstoricaly, stephanids have been unplicitly or etplicitly associated with Ichneumonoidea (most authors), Evaniidae (Westwood 1951, Smith I86Ia/b, Cameron 214 1903), Megalyridae (Riek 1973), and Orussidae (Ashmead 1900, Rasnitsyn 1980, Gibson 1985:1412, Whitfield 1992, Gauld 1995), in many different ways. But the most discussed hypothesis about the phylogenetic relashionships o f stephanids is that of Rasnitsyn (1988), which considers this fiunily as a member of the Evaniomorpha, a group proposed earlier (Elasnitsyn 1980) to include the Megalyridae, Trigonaiidae, Evanioidea, and Ceraphronoidea. Most authors disagree with this hypothesis, and Whitfield (1992), based also on mfonnation fi-om Mason (unpublished), Gibson (1985), Johnson (1988) and Whitfield e t al. (1989), concluded that stephanids “are not closely related to the Evaniomorpha.” This is also implied in Neumann's (1992) conclusion that “Stephanoidea do not appear to be closely related to any other Apocrita.” Johnson (1988) studied the midcoxal articulation in Hymenoptera and confirmed that the Evaniomorpha sensu Rasnitsyn (1980), i.e., excluding Stephanidae, is supported by a derived condition in the mesal articulation o f the mid coxa. Ronquist e t al. (1999), however, insisted that the derived condition is also present in stephanids, but apparently missed the 6ct that Johnson’s interpretation o f the character is distinct fi-om that of Rasnitsyn (1980 and 1988). Jonhson (1988) states that “the unique aspect of the mesal articulations pn Evaniomorpha] is in the degree o f elongation of the mesal lobes and not [as proposed by Rasnitsyn 1980] in the Act that the articulation is formed with the surfime of the coxa.” I checked this character in several stephanid species, and the presence of a short mesal lobe seems to be constant throughout the family, which agrees and lends further support to the observation of Johnson (1988). 215 At the same time, and in a more or less mdependent way, the idea that stephanids represent a basal group within Hymenoptera has also been discussed by several authors: most phylogenetic studies on Hymenoptera which include Stephanidae (Kônigsmann 1978, R a s n it^ 1980, Gibson 1985, Whitfield 1992, Dowton & Austin 1997, VOhelmsen 1997) agree m that this âmily has a very basal position within the Apocrita, and could perh^s represent its sister group, showing a close relationship with orussids. On the other hand, however, extensive data from Rasnitsyn (1988), both as originally interpreted and as reanalysed by Ronquist etcd, (1999), suggest a different position, with stephanids appearing in a distinct clade within Apocrita. Yet, in spite of the above efforts, Ronquist (1999) concluded that very little is known about Apocritan phylogeny, and summarized the state of knowledge for this group with a highly unresolved cladogram, which neither support nor refute current or early ideas on the phylogenetic relationships o f stephanids with other Apocritan taxa. The above situation seem to suggest that complementary data about the phylogeny of Hymenoptera might be necessary to support more accurate conclusions. However, in spite o f all published opinions, the available data on the phylogenetic position of stephanids has not yet been put together and formally analysed with cladistic techniques. Of particular interest are the data matrices in Vilhelmsen (1997), which compliles most of the available data from the literature for basal taxa of Hymenoptera, as well as provides origmal data, and Ronquist e t aL (1999), which present a revised matrnc for the extensive 2 1 6 orignal data set provided by R asnh^ (1988). Together, these two works compile nearly all o fthe morphological characters ever coded for stephanids in investigations about Hymenoptera phylogeny. Vilhelmsen (1997) and Ronquist e t aL (1999) used almost entirely different character sets, but their data are fully compatible as characters were drawn mostly feom the same source i.e., skeletal morphology. Taxa selected are also similar. These data sets can therefore be easily combined and used to generate more complete phylogenetic mfonnation for stephanids. Combmed analysis o fliterature data I combined data from Vilhelmsen (1997) and Ronquist e t aL (1999) into a single matrbc, and analysed it following the same principles and procedures explained before for investigations on the internal phylogeny of stephanids. As already discussed, however Qtem Thylogenetic Analysis”), I agree with Goloboff (1993) in that results from implied weights should be preferred over those obtained with equal weights or successsive approximations. For this reason, results from the last two methods will be mentioned below only for the sake of completeness, and discussions will focus on results from hnplied weights. In the present cas^ however, justifymg the selection of a smgle K value (for the implied weights analyses) seems madequate, because it would depend on detailed knowledge about the homoplasious nature o f characters throughout the order Hymenoptera, and this was not closely investigated here. To follow the original authors would be equally problematic because th ^ adopted very different approaches (K was set 217 to 6 m Vilhelmsen 1997, and to 2 in Ronquist e t al. 1999) and did not provide a justification fi>r their choice. For these reasons, all allowed values for K in PIWE (i.e., 1-6) were investigated for the present andysis. Results were evaluated comparatively. I decided not to mclude or discuss molecular data based on two reasons: first because results from molecular analyses are still very preliminar for the Hymenoptera, and consequently not sufSciently reliable; and second because the nature of molecular data, at least in studies about Hymenoptera, is not yet as well undestood as that of morphological characters, so that differences are dffîcult to compare and conclusions tend to fitvor morphological characters (see for example discussion on the monophyly of Orussidae + Apocrita in Ronquist era/. 1999). fri combming the original matrices two basic problems appeared. First, a total of 78 taxa were investigated by Ronquist e t aL (1999), but only 20 by Vilhelmsen (1997). All taxa from the latter were considered by the former, but it becomes obvious that characters from Vilhelmsen (1997) had to be considered as "unknown" for most of the taxa in Ronquist e t aL (1999). Second, data for extmct taxa in Ronquist e t aL (1999) are, as expected, highly incomplete. In order to compare the effect of these limitations, the combined data were evaluated under three separate approaches; (1) with all taxa considered, (2) extinct taxa excluded, and (3) only taxa appearing in both works considered. Taxonomic concepts are equwalent in the mentioned works, but data for Tenthredhudae and Diprionidae from Vühefansen (1997) were fused in the combined matrices because these fiunilies are treated together as Tenthredinidae sensu lato in Ronquist e t aL (1999). The few characters which Vilhelmsen (1997) used from Rasnitsyn 2 1 8 (1988) were ignored m Avor o f the same characters considered by Ronquist e t aL (1999) because the latter revised and complemented Rasnitsyn’s observations. The only exception is the midcoxal articulation, used in the combined matrix as in Vilhelmsen (1997) because this author adopted Johnson’s (1988) reinterpretation o f this character, which seems to be more accurate, as discussed. Character number 51 in \Hlhelmsen (1997) was ignored in &vor o f character 64 in Ronquist e t aL (1999) because t h ^ are the same, and coding is more complete in the latter work. For analyses following approaches (1) and (2) above, taxa appearing only in Ronquist e t aL (1999) where assigned a for characters used by Vilhelmsen (1997). The question marie symbol is interpreted as “missing information’’ in the computer programs adopted here. For the analysis following approach (3), 20 taxa were considered. With the reduction in number of taxa many of the characters considered by Ronquist e t aL (1999) became unin&rmative, and therefore were removed from the combined matrix. Character states were renumbered accordingly. Character additiveness follows Vilhelmsen (1997) and Ronquist e t aL (1999) strictly. The following characters, numbered as in the original works, have been used. The plus sign indicate it was considered as additive. From Vilhelmsen (1997): 2 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21+ 22 23 24 25 26 27 28 30 31 34 36 39 40+ 41 42 43 44 45 46 47+ 48 49 50 52 53 54 55 56+ 58+ 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76+ 77+ 78 79 80 81 82 83 84 85 85 87+ 89 90 91+ 92 93 95 96 97 98; from Ronquist era/. (1999): 2 3 4 5 6 7+ 8 9 10 14+ 15 16 17+ 18+ 19 23 24 28 29 219 3(R 32 34 35 36f 37 39 40+ 41 42 43 47+ 48 49+ 50 52 53+ 54 55 57 58 59+ 61+ 64 67 69 74 75 76+ 77 78 80 81 82+ 84 86 88+ 91 92 94+ 97 98 99 100+ 101 102 103 104 108 109 110 113 120+ 124 125+ 126 127+ 133+ 137 138 139 141 142 143 148 149 150 151 152+ 153 154 155+ 156 157 159 160 161 162 163 164 165 167 168 169. Overall, 188 characters were evaluated, 110 (of 169) from Ronquist (1999) and 78 (of 98) from \^elm sen (1997). Results caul Discussion Whh all taxa considered, the analysis in NONA/Ratchet found 411 MFCs of 960 steps. Cl 38, RI 80, for which the strict consensus tree shows low resolution for Apocritan taxa (Rg. 201). When these cladograms were submitted to successive weighting in NONA, one MPC resulted (972 steps. Cl 37, RI 79) (Rg. 202), in which Apocrita appears distinctly divided in two main monophyletic groups, and stephanids at the base of the Microhymenoptera clade (sensu Ronquist e t ai. 1999; = Proctotrupomorpha + Evaniomorpha + Stephanidae). Implied weights under all allowed values for AT produced essentially the same result^ although whh some variation for Evaniomorpha taxa ^ g s 203-207; steps 997+18, aU whh Cl 36 and RI 78). Analyses with data restricted to extant taxa only produced results which essentially match the ones reported above, but the overall resolution of cladograms was, expectedely, better than in the analysis with all taxa considered (since the matrix for restricted taxa is considerably more complete). This is reflected in three differences in relation to the analysis whh all taxa: (1) only 107 MFCs Wiere found by NONA/Ratched (907 steps. Cl 40, RI 78) (strict consensus in Fig. 208); (2) a byphyletic Apocrha was retrieved in all 220 cladograms ^ g s 208-210); and (3) implied weight analyses for different values o îK produced more un%nn results ^ g . 210; steps 944+19, Cl 38, R I76+1). A final relevant difiference is that in results from NONA/Ratchet and successive weighting ^ig s 208, 209) the Ichneumonoidea do not appear as part o f the Aculeata clade, but are placed instead in between evaniomorpha taxa and stephanids. For shared taxa only (approach 3) the analysis in NONA/Ratchet generated 6 MFCs of nearly identical topologr, with 497 steps. Cl 52, RI 77 (strict consensus in Fig. 211). These resulted in one MPC under successive weighting in NONA (Fig. 212), same topolo^ and statistics. Implied weights analyses produced different results for Apocrita only, but always retrieving the same two MFCs for all allowed values of K, with stephanids m the most apical position in both of them (Fig. 213). Although the cladograms derived fi'om the combined matrices were generally similar to those of Ronquist e t aL (1999), two notable differences emerged for analyses with all taxa and with extant taxa only: (1) the strict consensus trees for most parsimonious cladograms found under equal weights are considerably less resolved than the correspondent cladograms in Ronquist e t cd. (1999), but (2) cladograms derived firom implied weights analyses were much less ambiguous for the Apocrita than those found with the same method by the same authors. The first difference is hardly surprising considering that the proportion of “unknown” character states increased considerably in the combined matrix, for reasons ecplamed before. Such entries are particularly damaging for the resolution of cladograms found under equal weights analyses, because th ^ affect how homoplasious a character 221 win be in the matrbc the more taxa coded with a “?” for a particular character, the more homoplasious this character wUI tend to behave in the matrbc. With implied weights, however, increasmg homopla^ (apparent or real) is compensated by the proportional down weighting of the homoplasious characters, which allows the unambiguous informational content to be efficiently utilized even in a highly incomplete matrix. This not only explains the increase in resolution obtained for results with the combined matrices, but also provides an additional reason for preferring results from implied weights. It is also interesting to note that results from implied weights obtained for all values o fK were extremely similar, a fret which makes the much higher resolution of these results particulariy significant. Such resolution and uniformity in the results from unplied weights indicates two important frets: I. if the influence of homoplasy is even minimally down-weighted (by setting ÜT to 6), then a dramatically different hypothesis is supported by the same data, showing that in the current case homoplasy is a highly significant frctor, and 2. the fret that similar cladograms are obtained for all values of ^ indicates that differences in homoplasy between characters are distinct enough to produce approximately the same results no matter how much influence is attributed to the weighting scheme. Both these fiicts, as well as the discussion m the previous paragraph, strongly suggest that results from weighted parsimony should be given preference in the present analysis. Overall, results from the combined matrices suggest the following hypotheses: (1) that stephanids are more related to taxa from Evaniomorpha (except ceraphronids and megaspilids) than to any other Hymenoptera, (2) that stephanids always fidl within one of 222 the clades of a biphyletic Apocrita, and not basal to them 0 e , stephanids do not appear as the sister group of Onissidae), and (3) that stephanids and the non-ceraphronoid evaniomorphs are the most basal taxa within Mîcrol^menoptera. Note that hypothesis 3 also implies that, at this point, evidence suggesting that stephanids are the most basal taxon of Microhymenoptera seems no better or worse than similar evidence for members of Evanioidea, Megalyroidea, or Trygonalidae. Although these points are partially evident in the cladograms found by Ronquist et aL (1999), th ^ were not discussed by these authors. Ronquist (1999), citing Ronquist et aL (1999), stated that “the Evaniomorpha form a paraphyletic grade of basal apocritan lineages,” which is similar to hypothesis 2 above. This view, however, is not (and could not be) clearly defended by Ronquist e t aL (1999) because these authors attributed equal importance to the somewhat incompatible results obtained with equal weights and with implied weights. Support for the hypotheses formulated above, as interpreted from the results with the combined matrices, are discussed below. Hypothesis (I): Stephanids are closely related to non-ceraphronoid evaniomorphs. This possibility is suggested mainly by the fret that in the present analysis stephanids always appeared as the sister group of one or more families or clades of Evanioidea, Megalyroidea, and Trigonalyoidea. It is also mdfrectly supported by the frict that ceraphronids and megaspilids appear as a very apical group within the Proctotrupomorpha in all cladograms, several nodes apart from other evaniomorphs. As mentioned above, this relationship can be defended at the moment only as a “paraphyletic grade” Ronquist 1999). Phylogenetic relationships, however, can be criticized but not 223 refilled only because a group of taxa is not unambiguous^ monophyletic: a weakly supported hypothesis remains the best explanation if data interpretation can not provide a better supported alternative. In the second approach, however, stephanids also appeared near the Ichneumonoidea in the consensus tree fiir the MFCs fiiund under equal weights, and in the cladogram firom successive weighting (Figs 208-209). Even though such results are considered here of secondary importance (based on methodological grounds), it is interesting to note that (I) most Evaniomorpha in the mentioned cladograms still appear near Stephanidae, inunediately basal to the Ichneumonoidea and (2) extensive research about the phylogeny of Ichneumonoidea suggest that its placement in the Microhymenoptera is a less well supported hypothesis than considering it as part of the Aculeata (Ronquist 1999, Carpenter & Wheeler 1999). Hypothesis (2): Stephanids are members o f a clade wtthin a biphyletic Apocrita^ and therefore are not the sister group o fOrussidae, Nearly all results obtained with and without «ctmct taxa, including PIWE searches for all allowed values fbr^ suggest that the Apocrita are biphyletic, with one origin fi)r the Nficrohymenoptera, and a separate origin for the remanmg ^ocrita. The only exception was the strict consensus tree fi)r cladograms derived from equal weights parshnony with all taxa considered, which shows very low resolution fisr the Apocritan clade. This, however, simply indicates that results from equal weights are highly ambiguous, and therefore do not strictly contradict the hypothesis supported by unplied weights. 224 Although the matrbc restricted to taxa appearing in both VOhelmsen (1997) and Ronquist e t aL (1999) had only 20 taxa, results derived from its evaluation can be considered equhralent to those of the most complete matrices: in all cladograms obtained via implied weights the Microhymenopteran groups appear in one branch, and vespids (representing Aculeata) in another. Equal weights parsimony and successive approxunations fr)r this matrbc placed stephanids basally, apical only to Orussidae, but the significance of this observation is surpassed by the Act that with implied weights stephanids appear completely apical under all possible values îo tlL It is mteresting to note that results obtained by Carpenter & Wheeler (1999), combining morphological and molecular data, corroborate with the idea of two major clades within Apocrita. This possibility is significant in the present context because if stephanids are indeed part of one of such clades, as seems to be the case, then it becomes obvious that thebr basal or apical position cannot be defined in relation to Apocrita as a whole, as proposed by some authors (see introductory considerations for current section). It may seem surprising that stephanids form a “clade” with Ibaliidae in the implied weights cladograms for the restricted matrix. Note, however, that because of the low representativeness of Apocritan taxa, the mentioned clade does not necessarily imply a dose relationship o f stephanids with ibaliids proper, but with the lineage ibaliids represent. Le., the parashoid taxa which appear apical to stephanids in the cladograms fr>und with more complete matrices. 225 Hypothesis (3): stephanids and non-cercphronoid evaniomorphs are the most basal Microhymenoptera. With more complete matrices (approaches I and 2), close phylogenetic relationships between stephanids and other non-ceraphronoid Evaniomorpha are supported by all results obtamed with PIWE, regardless of the value selected for f ^ g s 203-207,210), and by the cladogram obtained under successive approximations when all taxa are considered (Fig. 202). Note that to evaluate these results based on strict consensus trees only is not a fiilly appropriate procedure in the present case. This is because strict consensuses preserve only the clades all trees have in common, while the current idea is to ddSne with which individual taxon or taxa stephanids appear closely associated in all possibilities. Note, &r example, that the apical placement of evaniids in the cladogram of Fig. 204 does not compromise the fact that stephanids and megalyrids or gasteruptiids appear near each other in all other PIWE cladograms. However, since evaniids also appear near stephanids in other trees (e g.. Figs 205-206), the total variation m then relative position will force all the mentioned taxa, as well as several other, to appear collapsed together into a strict consensus tree, thus eliminating the information which can be learned by examining the cladograms individually. The analysis restricted to shared taxa (approach 3) is limited in allowing further discussion on the relationships of stephanids with non-ceraphronoid evaniomorphs. The dffîculty lies in the high proportion of evaniomorphs, which will lead to their “close reWonship" with stephanids independent of the final results, thus compromising the objectiveness o f results fi>r discussions about this issue. 226 C onclusion ^results from implied weights are adopted as the best hypotheses, as defended her^ then it is much more reasonable to say that current knowledge does not support the sister group relationship of Stephanidae and Orussida^ but suggests instead a basal position o f stephanids, together with non-ceraphronoid evaniomorphs, within the Microhymenoptera clade of a biphyletic Apocrita. 227 FIGURE 1 Habitus ofParastephemellus rufoomatus (Cameron). 228 229 HGÜRES 2-7 General morpholo^. ParastephanellusdamenicusQNeStwooi): 2, head, frontal; 3, head, dorsal; 4, pronotum, dorsal; 5, head, posterior view; 6, prothorax, lateral; 7, propodeum and metaploiron, lateral. Scale line: 1.0 mm. 230 Coronal area Posterior tubeicle Anterior tubercle Coronal ocellus. Lateral tubercle Lateral tubercle CoronaloccHux Intemcellar carinn Posterior tubercle Anterior tubercle Lateral ocellus Frontal Carina Genol angle Gerrnl angle Antenrurllbrame Teniple Occipital Carina Centco-longilutlinal impression Apical Carina Vertex Occiput Occipital Carina C n io -j Rimarounil Foramen magnum foramen magnum Pol Preannular Eye Eye Gena Gena Setniimnular Hypostomal Carina Apex of Occipital carina Pronoml lobe - — Proporleumv Spiracular groove Spiracular carina Parapetiolor depression Semiaiuiular Pmpodcal spiracle: Pnmotal Anterior plate fobt and Posterior plate Occipital - Pronotai lobe Dorsal Pleutopmpodeal fbven Intcrfovcolor area (( Ventral 'l Ventral a Metapleural fovea Femoral impression Pnstfbveolararen 231 FIGURES 8-26 Antennae and basal flagellomeres, ventral (except otherwise indicated). 8, St^hanus senator, right, dorsal; 9, Hemistephanus cylindricus, right, dorsal; 10, H, cylindricus, fis. 1-2; 11, Schlettererius cinctipes, nght, dorsal; 12, Megischus tibiator, fis. 1-2; 13, Foenatopus aurantiiceps, pedicell and fis. 1-3; 14, Foenatopus (j=Diastephanus) sp. A, scape, pedicell, fis. 1-3; 15, Foenatopus sp., fis. 1-3; 16, M egischus n. sp., scape, pediceU, fis. 1-2; \l,Parastëphamttusrufoornaîus, scape, pedicell, fis. 1-5; 18, Æ cylmdricus, pedicell and fis. 1-3; 19, P. nigricaudus, pedicell and fis. 1-3; 20, A/, texanus, scape, pedicell, fis. 1-5; 21, P. gycarukttus, fis. 1-3; 22, Parastephanellussp,, pedicell and fis. 1-4; 23, M calÿomicus, scape, pedicell, fis. 1-4; lA,Megtschusti. sp., pedicell and fis. 1-6, dorsal; 25, H. limpidipennis, scape, pedicell, fis. 1-2; 26, H. cylmdricus, right, dorsaL S«des: 0.5 mm. 232 î S â ^ Lc>O O o C > O _jjOc»oc»maa °®^^^^3B2=tn3CîB2Cl^^ Tl^Ç^O^OL^g ?.QOSuLa.Sa?i5* R ^ g jQ S HGURES 27-39 Head. 27, Foenatopusflavi (=Diastephanus) sp. B, lateral; 30, Hemistephanus otfilanus, posterior hal^ lateral; 31, H. cylineticus, &ontal (sculpture and mandibles not shown); 32, M egischus (^Meuiegqfoenus) occiputaliSy frontal (sculpture not shown); 33, M. maculipermiSy dorsal; 34, Schlettererius cinctipeSy dorsal; 35, F. annulipeSy dorsal; 36, F. aurantiiceps, dorsal; 37, F. armulipeSy lateral; 38, Schlettererius cinctipeSy ventro-lateral, right; 39, M texanus, ventro-lateral, right Scale lines: 0.5 mm. 234 /fi 37 235 FIGURES 40-53 Head. 40, Foenatopus sp. A, ventro-lateral; 41, Hemistephanus elimatuSy ventral; A9,Me0schus n. sp., latero-ventral. Maxillar palpus: 42, Megischus tibiator, articles 3-5, lateral; 43, M tibiator, article 3, dorsal; 44, M egischus n. sp., articles 2-3, dorsal; 47, M. texanus, articles 2-3, dorsal; 45, Schkttererius cinctipes, articles 2-3, dorsal; 46, M maculipennis, articles 2-3, dorsal; 50, Megischustl sp., articles 2-6, lateral. Mandible and clypeus, dorsal: 48, H validus; 51, Stephanus serrator, 52, M. maculipennis, 53, H, macrurus. Legends: he, hypostomal carina. Scale lines: 42-46,0.1 mm; all others 0.5 mm. 236 57 52 53 237 FIGURES 54-68 Pronotum. 54, Schlettererius cinctipes^ dorsal; 55, S. cinctipes, lateral; 56, Megischus tibiator, colo, dorsal; 57, Stephanus serrator, dorsal; 58, S. serrator, lateral; 59, Hemistephanus erugatus, dorsal; 60, M texanus, dorsal; 61, M texanus, lateral; 62, K limpidipennis, dorsal; 63, M maculipennis, dorsal; 64, M maculipennis, lateral; 65, XarOhostephanusxanthocephalus, dorsal; 66, Parastephaneüus isoglyptos, dorsal; 67, P, isogfyptos, lateral; 68, Xanthostephanus xanthocephalus, lateral. Scale lines; 0.5 mm. 238 239 HGURES 69-82 PronotunL 69, Foenatopus annulipes, dorsal; 70, F. annulipes, lateral; 71, F. sp., dorsal; 72, F. alutacem^ lateral; 73, F, cdutaceus^ dorsal; 74, F. novagumeensis, lateral; 75, F. sp., lateral; 76, F. novagumeensis^ dorsal; 77, F pentheri, lateral; 78, F. (rNeostepharms) sp. A, lateral; 79, F. pentheri, dorsal; 80, F (=Diastephanus) sp. D; 81, F oenatopus n. sp.; 82, F. s ^ r ig i, dorso-lateral. Scale lines: 0.5 nun. 240 f s 241 FIGURES 83-101 Pro- and mesothorax: 83, Schlettererius cinctipes, apex of prostemum; 84, F. am m lipes, apex of prostemum; 85, Megischus maculipennis, connection of pro- and mesothorax, lateral; 91, F. annulipes, posterior margin of pronotum, dorsal. Front tibial spur and vellum, mesal: 86, Schlettererius cinctipes, 87, Stepharms serrator, 88, Foenatopus SÇ. D; 89, M maculipennis, 90, F. formosanus. Mesothorax, dorsal; 92, Schlettererius cinctipes (sculpture not shown); 93, Stepharms serrator (sculpture not shown); 94, M texanus (sculpture not shown); 95, M. maculipennis (sculpture not shown); 96, Parastephaneüus n. sp.; 97, F, albomaculatus, 98, Hemistephanus cylhuiricus, including posterior magin o f pronotum (sculpture not shown); 99, F. aurantiiceps. Mesepistemum: 100, M Jurcatus, 101, F, aurantiiceps, arrows indicate direction of prumosity. Legends: Ax, axilla; C x i, front coxa; M e, mesoscutum;mp, mesopleural pit; M t, mesepistemum; ms, median sulcus;ra, notaulus;p , prepectus; Pn, pronotum; p p , parapside; P s, prostemum. Scale lines: 0.5 mm. 242 cP 243 HGURES 102-113 Thorax (sculpture not complete in most drawings). 102, Foenatopus (^Diastephanus) sp. B, dorso-lateral; 103, Foenatopus sp. A , mesepistemum; 104, F. (^Neostephanus) sp. A, thorax (except legs and apoc of pronotum); 105, M egischus fiacatuSy propodeum + metapleuron; 106, M (;=Mackgqfoemts) seyri^, thorax (except legs and apex o f pronotum), lateral; 107, ParastephaneUus n. sp., posterior half of mesepistemum, mid coxa, propodeum + metapleuron; 108, M coronator, intercoxal space between hind legs, posterior view; 109, F oenatopus sp. H, propodeum + metapleuron, lateral; 110, M ntfofemoratuSy bases of mid coxa^ ventral view; 111, F. (-Neostephanus) sp. A, intercoxal space between hind legs, posterior view; 112, ParastephaneUus n. sp., mid tibia, mesal side OpHosity not shown); 113, detail of spines. Legends: <6, discrimen; M r, metanotum; Scy scutellum. Scale lines: 0.5 mm. 244 / «yyatiCfc KJ ri2 245 FIGURES 114-129 Hind coxa and Amur. 114, Hemistephams elimatus, articulation o f hind coxa with thonu^ dorsal view; 1 IS, Foenatopm (^Neostephanus) sp. A, articulation of hind coxa wiht thorax, dorsal view; 116, Parastephanellus granukttus, bind coxa, dorso-lateral; 117, Megischus texcams, hind coxa, lateral; 118, H. elimatuSy hind coxa, dorsal; 119, F. cmrantiicepSy hind coxa, dorsal; 120, F. aurartiicepSy hind coxa, lateral; 121, F. aurantiicepSy base of hind femur, mesal; 122, Foenatopus sp. A, base o f hind femur, mesal; 123, Schlettererhts cmctipeSy hind coxa, lateral; 124, Foenatopus sp. F, base of hind femur, mesal; 125, F. albomaculatus, base of hind femur, mesal; 126, F. arnmlipes, hmd femur, dorsal; 127, F. annulipes, mid-transversal section of hind femur (arrow shows dorso-mesal depression); 128, Stephanus senator, hind femur, lateral; 129, Schlettererius (kterminatoriSy hind femur, lateral. Legends; a t, apical tooth; mt, median tooth. Scale lines: 0.5 mm. 246 c 723 727 722 724 725 726 727 728 129 a t 247 FIGURES 130-142 ISnd femur, lateral: 130, Hemistephanus macrurtts\ 131, Foenatopus aurantiiceps; 132, Foenatopus sp. L Hind tibia, dorsal: 133, Schlettererius cmctipes; 134, Stephanus serratOTy I35y Megischusmaculipermis; 136, Parastephanellus n. sp.; 137, M tibiator, 138, F, annulipes. Hind tibia, lateral: 139, M bnameusr, 140, M tibiator. Hind tibia, ventro-Iateral; 141, Æ cylineùicus; 142, M Jurcatus. Scale lines: 132, 140, 141,0.5 mm; aH others 1.0 mm. 248 130 T3I 732 733 734 735 736 739 Î40 747 742 737 738 249 FIGURES 143-160 Abdomen: 143, Schlettererius cmctipes, basai segments, lateral; 144, Foenatopus sp. A, apex of petiole and segments3-5, iatero-ventral; 145, sam^ petiole, lateral; 147, F. (=Diastephanus) sp. A, apex, latero-posterior; 148, F. annulipes; apex, latero-posterior. ICnd tarsus: 146, Stephanus serrator 9, lateral 0>dosity and setae of apical tarsomeres not shown). Wmgs: 149, Megischus crassicauda, hamuli; 150, M fitrca tu s, front wing, posterior margin apical to retinaculum; 151, detail of short hairs; 152, detail of long hairs; 153, M tibiator, front wing, posterior margin apical to retinaculum, with detail of short hairs; 154, detail of long hairs. Front wm^ four-vein nodus and setae on M+Cu: 155, M Ç^^adegafoenus)hétàlyanusr, 156, Hemistepfumus elimatus; \51,ParastephanellusTi. sp.; 158, P. nigricaudus; 159, F. flavidentatus. Hind wing: 160, Stephanus serrator, ventro-apical portion. Scale lines: 143,145,160, 1.0 mm; 144,147-146, 155-63,0.5 mm; 149 0.25 mm; 150, 153, 1.0 mm; 151-c, 154,0.1 mm. 250 TS, 143 144 146 ...... 151 'X 154-^r\ \ \ 1 U*Cu 251 HGURES 161-162 Vein an ceQ nomenclature. \6 l, MegischuscrassicaudaMatief, 162, ScMettererim cirwtipes Original lengths, in nun: 161,10.9; 162,7.9. 252 Parasrigma 1+2R1 I+Z+3R, (Ki+M)a ^ — ^ 2+3W iM ryt**^a s ScfR- JRsb 2M Cub 162 FIGURES 163-167 Front wing. 162, Schletteritis cmctipes-, \6A, Stephanus serrator, 165, Megischus anomalipesr, 166, A/, tacanus; 167, M tibiator. 254 163 164 165 166 167 255 FIGURES 168-172 Front wing, 168, Hemistepharms macrurus (membrane of costal area folded upwards); 169, H. cylindricus (membrane o f costal area folded upwards); 170, Xanihostephanusxanthocephalus; 171, Parastephanellusrufoomatus; 172, Foenatopus pentheri. 256 Î68 169 257 FIGURES 173-178 Front wing. 173, Foenatopus catrantticeps; 174, F. (=Neostepfumus) sp. A; 175, F. navaguineensis; 176, Foenatopus sp.; 177, Foenatopus sp.; 178, F. alutaceus. 258 259 HGURES 179-194 i& dw îng. 179, Schlettererius cmctipes; 180, Stephanus serratory ISl, M egischus anomalipes; 182, M texanusTy 183, M crassicaudOy 184, M tibiatory liSyHemistephanus limpidtpermiSy \ZSy Xanihostephanus xanthocephalus; 1%1, Parastephanellus rufoomcttusTy 188, Foeruttopus 189, Parastephanellus tl sp.; 190, F. aurantHcepSy 191, F. ^Neostephams) sp. A; 192, F. rwnfaguineensis; 193, Foenatopus sp.; 194, F alutaceus. 260 .... - /y . 184 185 186 187 188 189 190 191 _ 192 198 194 2 6 1 HGURE 195 Strict consensus tree o f 36 MFCs of 1145 steps (Cl 24, R I72) found by NONA (characters all accorded equal weight). 262 SCHLEnSSERlOS cÙ K t^ iuséetembuuoris VHANÜS serrator ie g i^ m anomaUpes -Mègisdius texanus fegfsdaa ea^fbndcus ■egüdaa bnumeus r~Megfsdua nfq/èmoratus jilEŒSCHüSftiraaia XUegtsdmmaoê^pennis ~ ' 'ytomerus rtibktor ia^M ^rm ocdputaUs faaegttfom a ber^Banus r— Megischus lusp. — HEMI^TEPHANÜS macrurus r n w ^ e im ta h a im cyUndricus ^ " •\r-^ a m a h a n u s eBmatus ^-^enBae^uamUmptdtaennis Stephanus xanthocepnaba 195 ^Parastepfunàbis n.sp. ^mParastepihaneBus rigrkauim ^^Parastephandba m, —Parastepfumelluin^rmaus —^Parastephaneüus granuUaus ^ P arastephanettm denotatus *ABÀSIEPHdNELLüSpygmaeus r-^JdADEGAFOEmJS seyrigt ^^eostephanus albomaculatus ^-^oenatopva jpJ? PoauMpus sp.C mPoenatopus sp J ) -foenatopus sp.E FOENATOPUSindittotr m^jytastephanusn.^, mmmmDiast^usnus s p ^ Dîastep/uaua spM D iasttpbanus m J ) ——Diastephanusflavideniatus loenatopus navaguineensis foenatopus spJF •Foenatopus sp .G —FoenatopusFoenatopus sp.H ’iUSTEPuAmJSfisvornaadi]atus fo e n a to p u s s p ^ ——I rFoenatopus auraïuiiceps ^foenatopus annulipes ^mmNeostephanus spui ""^l— ilEOSIEPBAmS oBuaudi i-Diastephanus sp.C "•foenatopus sp.i . - Foenatopus arcuatus 1 r-Oiastepnanus alutaceous ^"Voenatopus nataBcus 263 HGURE 196 Strict consensus tree o f 36 MFCs o f 1145 steps (Cl 24, RI 72) found by NONA with aid o f Ratchet (characters all accorded equal weight). 264 SCHLETTERERIUS dnctipes ^ d d e ttm rtuxdetemüfKUOTîs I rnm^STEFHAHUSsemur I mMegbdutt anomales I i-H Y ^^^bdmtexam ^"^M em d a a a d ^ n d a a mmma^hanuTgjlgas — mjfeguchus brwuieus mmmUetfxnus n^bfimoraaa ^t^Tuam sesfiÿ^ - I w^adezafienusocclpmaBs ^"^y^Jdadegqfiam bel^Bam >aluampa^bmena übbttor — Afegischusn. HANUSmaenaus qfUndrtaa eSmatus x U n ^^e m is 196 — iSipppytow •Panatalumdba m, 'arastepnaneUus n ^ m a tu s rnPanat^umellus granulatus ^arastqmaneOusaauaaais mmmPARASIEPHANELLUSmmaats ~ ~MADEGAEO^S seyrtgji ^oenatopussp^ loautopussp.C J) 'oeiuaopussp., -Foemiopus arcmius )ENAIOPÜSinTaaa — d astephanus iLsp. THiatephama sp ^ ■ ^dasuphanusspM I—I ^Dtastepfuuaa sp.C —JHastephanus spJ> Diastepfuomflavidauattu Eoaiatopia novapmeauis ~^oenatopus sp.F Foenatopus sp.G —Foenatopus spM loenate^spJ iostephanus albonwadatus ^MSTEPHANÜSflavomaculatus _ _ w — N eoste^ am spJL “ '''n-^.lŒOSIEPHANaS aUtmtU ^ ^ w —Dfast^fumus alutaceous ^-Foenatopus natalicus ■oenatopussp.A '^oenatapus aurantiiceps foenatopus annuEpes 265 HGURE 197 Smgle MPC cladogram found by NONA following the application of successive approximations character weighting to the 36 MFCs found in the unweighted analysis with aid of Ratchet. 266 SÇHLEnERERIüS cbialpes iusdetémùiatoris PHAtiüS serrator egjbdm anomalies " 'sdaistexanus tc a ^ m k u s egis^ûrirunneus fM eàsdm ndbfimoratus jèiÈGlSCÈiuSfiuarais iMe^sdmmaail^eiuiîs ~ ihanuspadryUmerus Stephanus tibiator s ^ g t '(^benus ocdputalis •gq/betm bdgUatm w-MegfvAtajum, mmHEMlSIEPEUNÜS macrurus fH fH èm ist^hatm ÇjfUtubkus *“\r^emistmhatmeWtuitus 'tepfuinusxaiuhocepnàlus 197 Parastepàanelbis rusp, Tarastephanelm tügriamdus •Parastephanelha sp. rnParaste^umeUus dauruuus ParastephatteOus granulatus Parastephanellus n^rruttus PARASTEPHANELLUS mgmaeus OEGAFOE^Sseyrtgi feostepfumus albomaculatus fmFoertatt^usqiÆ III Foenatopus arcuatus J)tastepharm alutaceous foenatopus tuttaEcus foenatopus sp.G Foenatopus spM foenatopus novaguineensis foenatopm sp.F r^ ia step n a tw s n ap. f\fDiast^hanm spM Hm^laste^tanus sp.B \rDiast^hanm sp.C t-mFoenatopus sp.I foenatopm spJ llASJEPttANÜSJUsvomaculatm ')iastephanmflavfdentatm Hastephanus spA fOElUTOPÜS ituBcm foenatopm sp.C foenatopm spJ) ''feostephanm spA m on E P H A N üS albiaudi foenatopmspA Foenatopm auran^ceps foenatopm anmiUpes 267 FIGURE 198 A, Strict consensus of feur MPCs obtained with implied weighting implemented using PIWE with set to 1. All trees have fit S08.0, Cl 22 and RI 69. Number o f steps varied slightly, with one cladogram with 1239, two with 1240, and one with 1241. Values on branches indicate Bremer support in terms of fit (trees up to 1.0 less fit were calculatecQ.B, node numbers for the same tree. 268 ^CBLEnERERIOSefncàpes ' ^•SdikttembadaemOnmris . Schlettererius I w-mmmSIEPHANUSsenrnr Stephanus i.a n.Ê—^ahampaaÿlomerus T lJ Æ riMator mJifoMq/benus bdfyaaniu ùSfàÊEGISCBmfiacatus Megischus ^i^egfsduanuai^ienils 798A fm ca^fbnüaa \bnmneus )w-èügbdatt Tvfifimoratus 1 •U egbdm lu p . aai JBregfsdmtexaaus ^•m Uegfa^ ano f-BEUlSlEPHANUSnucnina 'T j i i 269 FIGURE 199 Cladogram obtained using wing venation characters only (strict consensus of 76 MPCs obtained with implied weighting implemented using PIWE with AT set to 1. Individual trees with fit I66J2, steps 308+8, Cl 24+1 and RI 76+1). 270 ÎCEŒEZTERERZro céldÿa îdœrmtnaioris ybm ena or îserrator rnmmUègbdm bnuBieia mlÎEŒSCHüSfiaaittis y — pufaSs iq /b a m l ylùam calÿbrTiicus îeÿsdmtexam 1—Megfsdua n^femoratus “‘^-mUegüdua 11^. i-tâe^rnmmommpes •mmm^EUlSIEPBANUSmacrunu I— I fmrnSmtstq^ham eWnanis *“Hmmaemîst^hamqf&itMau ^OmpamUanu *jŒASIBPBANELLUS pygfnaeus n/foomatus 199 iptianelbadaotaau 'Jbttiusp. tdgricaudus nûtephaïuOus^. I — •Stephanui xantkocephalus I w^oauaopmspJ} I I mmNèoaephanus spA I "I- ■ ifeosuphanus albomaculam L— j \^—NEOSl&mmiSaauaudl j^oauuoDUs aunattüceDS oenau^sp.i -•mFoenatopus arauuus ^oenatopui nataücus loauuopussp.G loenàopusspM ''oenatopia sp.F ^oenatopus novaguineaais mFoenatopus sp.C VENAXOPUSfadlcus mmFoauiu^us ’’œtuttopus spM - Dtastephama tusp, ^iastepfuuua atutaceous JdADBGAFOENUSsejrigi r-Diasuphanus sp.3 •mOUatephanus sp,C —^iastephaimiflavidauatus pu sspJ __ISÏI_____ mmmDiastephanux s p ^ *-mDlastephanus sp.D 271 FIGURE 200 Restricted analysis of Foenatopinae taxa. Strict consensus of 3 MFCs obtained with implied weighting implemented using PIWE with AT set to 1. All trees have fit 725.5; one has 602 steps. Cl 36, R I61, and the other two have 600 steps. Cl 36, R I62. 272 jSCHU^riERERIOS dnctipts mScMettenrius determimtaris f^mSJEPBAHUS senator 'upfuamxan^ux^habis j— JiiADl^jÀFOESÜS seyrigi m-^^eostephanus albomaculatus —Foenatopus s p £ ^^mJiUaupfianusn.sp. I Foenatopus arcuatus n ■ «^J)iastephamts abaaceus ImFoenautpia nataScus 200 m-Foenatopus navagumeensis j ^—Foenatopus sp.F tmm^oenatopus sp.G 'oenatopus sp.H feostephama sp^ OSTEFHANUS aUuaudi FOENATOPUS indicus 'oenatopus spJ} foenatopus sp^ -j I Foenatopus sp.C ^ foenatopus aurantiiceps imFoenatopus annulipes foenatopus spJ F^iastephanus sp.A F)iastepfumusjlavidentatus ^lASTEPHANUSflavom, ^Foenatopus spJ mF)tastepluinus sp.B iastepfumussp.C -Diastephanus sp.D 273 FIGURE 201 Réévaluation of literature data for Hymenoptera. Ali taxa (approach 1; see text for details). Strict consensus tree of 411 MFCs o f960 steps (Cl 38, RI 80) found by NONA with aid of Ratchet (characters all accorded equal weight). 274 -Jünhydnidm» rKuibmluka ■Ephulfitiriift ■—Onun^ ^ MegdyiUw Slyphaniditt "I I -TnMMlyidiB Gutguptadie ■■■PrMMilindiri -MeaoMmhtdae ■^Tunsniaae PiaMclmrmiiOBfdie i-i-Bethyloiiyimdac ScelioQiitM» U g a f e . _»-PeIecimdae ^^-Pradoliupi^ •PtuiDKudas gubidae la lemidae liidaa Mntillidaa imofphii£e 275 FIGURE 202 Réévaluation of literature data for Hymenoptera. AU taxa (approach 1; see text for details). Single MPC cladogram found by NONA foUowing the application o f successive approximations character weighting to the 411 MFCs found in the unweighted analysis with aid o f Ratchet. Steps 972, Cl 37, RI 79. 276 Cfmhidd geBdme jJPphfmlhhX»* 202 racoBidae "™%,ydudme ibbidae ^jboSmjidaft ■ . Rhoomlomommhdme VwpdM iSieroloiiinfpnuMft onmct^ mlmfbimddme Bnutynobaenidae tBphmiw Evaniidie dme —TrisomdyidiB Mimiwisluae dae —Kopraqiidw Helondie . unpriidaB 277 FIGURES 203-207 Réévaluation of literature data for Hymenoptera. All taxa (approach I; see text for details). Strict consensus of MFCs obtained with implied weighting implemented using PIWE under all possible values for ^ (1-6). All cladograms showed Cl 36, RI 78. 203, Strict consensus of 4 MFCs obtained with AT=1 (fit 1436.9, steps 1011+4); 204, relationships among non-ceraphronoid evaniomorpha in the strict consensus tree of 3 MFCs obtained with .A3=2 (fit 1682.2, all with 1013 steps); 205-12, same, 5 MFCs obtained with/T=3 (fit 1838.3, steps 1005+3), identical to single MFC obtained with A=4 (fit 1942.6, steps 1000); 206, sam^ 4 MFC obtained with ^=5 (fit 2018.0, steps 998+4); 207, sam^ one MFC obtained with .^=6 (fit 2078.3, steps 997). 278 cidae i rPneanlacidae pBdfaylMjnw -Tngooalyidie* ■I I KfiimfffiihWM ennbbidM —I p H elm i^ Siemlomoq^iidme -Fonnuridae -FmW^n^Wdme __ÜSJto 2ÎSS?;.-. 205 Mgnnmarhidifl -Meanidae -Pxoclotnvidae vaniuiift* .pwuidie*" ssar odae rAiutrooiidaB 206 I p J u ia p ^ ^lapmdM upidme ^idae kArmammmalidaa 207 Higmaptomidae 279 FIGURE 208 Réévaluation of literature data for Hymenoptera. Extant taxa only (approach 2; see text for details). Strict consensus tree of 107 MFCs of907 steps (Cl 40, RI 80) found by NONA with aid of Ratchet (characters all accorded equal weight). 280 [Outgroup JAamqrelmme ilesticotomUi» •TenHuedûiidae Cephidw F—AaeqreUd* -SiiiddM -]%hydrüdme f-Scolebythidas —Bethylidme etogibbufae 208 Dryûùdme -Smpyguhe —MuliUi&e — BndyiiobMHiirltn ia^ierolomorphidu —Fbrmicidme -wVe^idme i-Tîpfaiïdie i-Scoliidae •Sphecidae —LApidme Lpon^ilidae ■Megatyridw [viniidae Tngoomlyidae Idme liepfaamdae Ropraoiidae pHebnihe pPdecmidae **L>Pfoctotn9idae rAuatroaudae ‘t-Mbnomarhidaff [Diaprüdae pKgilmdae HfÔiupida» -Chalcidoiilee (ygBaïonuiiatîdaB -ifcalinnidiaw ygaatridae 281 HGURE209 Réévaluation of literature data for Hymenoptera. Extant taxa only (approach 2; see text for details). Single MPC cladogram found by NONA following the application of successive approximations character weighting to the 107 MFCs found in the unweighted analysis with aid of Ratchet Steps 914, Cl 40, RI 80. 282 gop&onihe iTeelhiediiiidie Cephidw ■^Antiydidae P«Sffiriitn ■X^ydriklae pPhmiariidae I |i5colebytfai(Iie I pBeihylidie Li rh^nirifilir p -Scleropbbiifae ^"(rlkymidme 209 I Bmhotomiriift ■J pPonpiliidH HfSpIiectdae ^ A p id a e j-V e^idm e | j —Stenbmorphtdie iponmcidae ■Tiphiidie ■Scolii MbaonachidaB [ymannmiiatidae 283 FIGURE 210 Réévaluation of literature data for Hymenoptera. Extant taxa only (approach 2; see text for details). Strict consensus of all MFCs obtained with hnplied weighting implemented using PIWE under all possible values for ^(1-6) (steps 946+17, Cl 37+1, RI 76+1). 284 I—XydàiaB -BlutüBotoinkfae rTnlnwlmida» I r ~L CünbkMmn f— PMBphiliîrfaw Meg>lndnnrf 210 igibbufae ■PmhnlmiiîH»» Bel&ylWme mSphocidme LrniApidmm " h PbmpilHM L mi Rhopmlommmlidmm J —VeipüiiH jpHfiieroldiiiofpliiibe TPonmddme -Sapygidae J rMutfllidw T r t—Bimdynobmemdie Tj-ïïphüdme nLscoliidae -Megalyridae «SlrphmmdmR — Tdgonalyidw tvaniidae ■Ropfoniidae p—Halondae —AuatfonndaB I KfoBomirhidiB mDi^i&Xaa —I fFîgitüdae *—-UÔrnipid» njÛüdae ebcûudae - c 'PioctotnipidaB ^ MegaapiltiiMt Cuaphmiûii [audae •Plalygaatridae 285 HGURES 211-213 Réévaluation of literature data for Hymenoptera. Shared taxa only (approach 3; see text for details); 211, strict consensus tree of 6 MFCs o f497 steps (Cl 52, RI 77) found by NONA with aid of Ratchet (characters all accorded equal weight); 212, single MFC cladogram found by NONA following the application of successive approximations character weighting to the 6 MFCs found in the unweighted analysis with aid of Ratchet. Steps 497, Cl 52, RI77; 213, strict consensus of two MFCs o f502 steps. Cl 51, RI 77, obtained with implied weighting implemented using FIWE under all possible values for AT (1-6). QEach cladogram with fit 1164.5 for^= l). 286 Oat(toup •Ouiciaap pMtctoocyd&BB I jAamqtlWe —Xyt&me L xvc &hb ■WmfaotoniHMt pAiikhe pAigtte '“H . ‘itedBBflBite .1.TMhrnfiniiitB » . LonUddK I-Oaimbkübe -PnnpUlidie -CephirtiB -CephMme —-AonysUte —WrfcidiB l-JQphydifite I— O iu n ite — Trigoomlidme .Vopidie pMepl: lytidie ■llwHiifae 211 272 tI dstetaptSdM^ G« 213 287 CHAPTER! THE FOSSIL STEPHANIDAE (HYMENOPTERA), WITH DESCRIPTION OF TWO NEW TAXA FROM BALTIC AMBER, AND KEY TO SPECIES OF ELECTROSTEPHANUS BRUES INTRODUCTION Two genera of fossil Stephanidae have been proposed, Protostephanus Cockerell, 1906, for one species from the Tertiary shales o f Florissant, Colorado, and ElectrostephanusBmes, 1933, originally including three species from Baltic amber. Brues suggested Hat Protostephanus was not a stephanid, because it seemed to lack a toothed hind fomur, a synapomorphy for the fomfly, but based his opinion exclusively on the description provided by Cockerell (1906). fri a recent publication. Carpenter (1992) still mentioned the Gundy level placement o f Protostephanus as doubtfiil This problem is investigated in the present work. The ÿeaasEIectrostephanusdiso presents some taxonomic problems. All species originaUy included m the genus dK&r considerably from each other in characters o f genus- level significance, and Brues himself suggested that one ofthem (£1 brevicomis) could possibfy represent a distmct genus. There ar^ however, some impediments to solving this 288 problem, particularly because all original type specimens oîEIectrostephanus are now mîMîng, and because all known specimens are males, which are much more problonatic to define taxoncmically than are females (Aguiar 1998). The aim of this chapter is to review the classification of fessil Stephanidae, comparmg published infisrmation with two recently discovered fessil species, as well as extant taxa, and to provide information relevant fer a cladistic revision of the family. MATERIAL AND METHODS This study is part of an ongoing revision o f the world Stephanidae (Aguiar & Nfiisetti 1998), and is based mainly on the examination o f the holotype o fProtostephanus ashm eadi Cockerell, deposited in the Museum of Comparative Zoology, Harvard University, and two recently discovered specimens fi’om Baltic amber assigned to Electrostephanus. The amber containing E sulcatus sp. n. is only partially clear, yellow brownish, and was cut several times, at different angles, to allow more accurate observations and drawings. The piece contaming E neavenatus sp. n. is of clear pale yellow amber, and was satisfectorily observed by submersion in pure glycerin, covered with a glass slide. Details of the cuticular sculpture were independently observed and confirmed by the author and by Jens-WOhelm Janzen, who also prepared the drawings. None ofthe types fer Bruy's species oîEIectrostephanus were feund, and may have been destroyed during the World War II (see efferts by Johnson 1998). In dealing with this problem, the fellowing procedures were adopted: (I) the distinction between E petiokttusBmssmiE Brues m the kty to species was based ecclusively on 289 characters provided by Brues (1933); and (2) several taxonomicaily significant characters not discussed by Brues (op. c.) for £. p etio ko u sand E. brevicomis Brues were interpreted dhrectly fi’om habitus illustrations o f these species provided m his paper. The character set and illustrations used here to describe bothJSL sulcatus sp. n. and K neavenatus sp. n. correspond to what was possible to observe rather than the most adequate for describing either fossil or extant species. Terminology follows Aguiar (1998). RESULTS AND DISCUSSION GenusProtostq^hanus Cockerell Protostephanus CocVsxeVi, 1906: 57. TyT^spedes: Protostephanus ashmeadi Cockerell, by original designation. Monotypic. Protostephanus. Carpenter, 1992:466. Familial statuso f Protostephanus ashmeadi Cockerell and sex of its holo^e. The examination o f the holotype o f this species confirmed that it is correctly classified in the fiunily Stephanidae. The overall condition o f the specimen is poor, but the following structures, characteristic of stephanids, could be observed (Figs 214-217): anterior and lateral tubercles of the head, all segments of the typically specialized hind leg, including an apically dilated tibia and a 3-segmented tarsus, with elongate basitarsus, and fi-ont wmg venation and shape neariy identical to that of basal extant species o f the âmily, as Schlettererius spp. and M egischus spp. fi)r example. The presence of three tarsomeres on the hind le& the basal one elongate and o f uniform width (suggesting the original. 290 tridimensional shap^ was (^Imdric), and the distmct absence of male genitalia, with an indication of what appears to be an ovipositor, unquestionably indicates the specimen is a &male. Recognition and taxonomic status. See genus Electrostephanus and item “Taxonomic status and relationships o f Protostephanus and Electrostephanus” GenusElectrostephanus Brues Electrostephanus ^roes, 1933: 12; Carpenter, 1992:466. Type species: Electrostephanus brevicomis Brues, by original designation D iagnosis. Vein IM arched, and distmctly longer than m-cu; veins Rs+M and ICu not parallel. Parastigma long and tubular, as long as, or usually distinctly longer than IRs. Vein 2r much shorter than 2+3Rs. Cell 1+2R1 relatively short, twice as long as its maximum width. Comparative notes. Separated from Schlettererius Ashmead by lacking a pronotal wall, colo dififerentiated, and dorso-apical tooth on hind coxa apparently absent. Separated from Stephanus Jurine by having a short petiole, with tergite and stemite not fused, o r without bullae between vem and veins 2+3Rs and 2+3M. Probably polyphyletic. All known species from Eocene Baltic amber. Females unknown. The most important features shared by Protostephanus and Electrostephanus are the Mowing: petiole short; vems 2+3Rs and 2+3M contiguous with RgfM (i.e., bullae absent); vein IM distmctly arched; vem IM distinctly longer than m-cu; and vein 2r 291 distmctly shorter than vein 2+3Rs. Nfoin dffîerences include: vein C ^M )b differentiated; veins 2-1A and 2Cua entirely tubular; vein 2Cub distinct (apparently tubular); and hmd coxa long, cylindric, and transversely strigate for Protostephanus. Kqr to Species of Electrostephanus Brues I True petiole not developed, second abdominal tergite and stemite distinctly separated, with a narrow membranous area between them (as in Fig. 227), or at least tergite 2 shorter than tergte 3 ...... 2 1 * True petiole developed, second abdominal tergite and stemite fused into an elongate cylindric structure that is distmctly longer than abdominal tergite 3 (asm Fig. 218) ...... 3 2(1) Vein ^s+M)b absent, and no bullae at origin of veins 2+3Rs and 2+3M (as in Fig. 219); vein 2Cua apparently absent ^ig . 4 ofBmes 1933:13) ...... K brevfconrisBmos 2’ Vem (Rs+M)b distinct, with two bullae between it and veins 2+3Rs and 2+3M (Fig. 225); vein 2Cua present, tubular...... K neavenatus sç. n. 3(1*) Hind femur with 3 teeth, one near base, and one near middle and apex ...... 4 3* Hind femur with 2 teeth, one near middle and one near apical fourth (Fig 220) ...... ...... E.sulcatussp.xL 4(3) % id femur with apical tooth near apex; basal tooth long acute ....EL petiolatusBmes 4* Hind femur with apical tooth at apical fourth; basal tooth short, not acute ...... ...... E . tridenta tusB ruts 292 Electrost^hanus sulcatus sp. n. (Figs 218-222,228) E tym ology. Ladnsir/cus, finrow, groove. Di reference to femoral impression. Type material. Holotype male m the collection o f the Geological-Paleontological histhute and N^iseum of the University of Hamburg Germany, Type Number SGPIH 3892. Included in Baltic amber (Eocene), from Samland peninsula, Baltic coast. Description. Male ÇEIolotype): Habitus as in Rgure I (chapter 1). Total length 7.8 mm. Scape about I.7x as long as wide; pedicel subglobular; first three flagellomeres subtylindric, somewhat flattened, distinctly wider centrally than basally and apically. Maxillary palpus S-segmented; basal segment the shortest and widest; 2nd segment elongate-fitsdbrm, nearly three times as long as first; 3rd to 5th segments elongate 3rd and 4th o f similar length, 5th the longest; 3rd and 4th s%ments thicker apically than basally. Coronal area areolate-rugose behind central ocellus; mostly polished smooth anteriorty. Distance between lateral tubercles slightly greater than distance between posterior tubercles, which are contiguous with each other mesaHy. Gena round, apparently polished smooth. Occipital carina very narrow. Pronotum finely transversely strigose dorsally; colo differentiated; femoral impression deep and wide (Fig. 218). Median sulcus and notauli distinct, formed by series of aligned foveola. Front tibia normal, not compressed basally; a series of strong short, stKf setae aligned on anterior fitce, inclined about 45* towards apmc, progressively stouter 293 and more numerous apically. Mesopleuron apparently smooth. Discrimen strongly foveolate. Metanotum and propodeum separated by very wide crenulate sulcus formed by several, deep; subrectangular crenulations (Fig. 222). Propodeum strongly foveolate, with some punctures (Fig. 222). Metapleuron strongly areolate-rugose; inter- and post-foveolar areas transversely strigate ^ g . 221). All tarsi S-segmented; ventral side of hind pre-apical tarsomere prolonged anteriorly, almost reaching apex of apical tarsomere. Hind leg with coxa stout, twice as long as its maximum height; ventral side of femur without teeth basally, central tooth broadly triangular, apical tooth more acute (Fig. 220); tibia with basal 0.3 slightly compressed, apically dilated; coxa and femur apparently smooth; tibia strongly obliquely alutaceous. Pronotum, mesonotum, and hind legs with many long erect hans; with hairs also on front and mid legs, but shorter. Front wing (Rg. 219) with cell 1+2R1 relatively short, only about twice as long as high; vein IRs at nearly 90" with Sc+R, and about half as long as 2r; IM about I.8x as long as m-cu; ICu 1.4x as long as Rs+M; 2r 0.4x as long as 2+3Rs; (Rs+h^b absent; bullae absent; 2-1A mostly developed, tubular basally to nebulous apically; 2Cua tubular on basal 0.2, remaining nebulous; 2Cub nebulous. Parastigma very long. Pterostigma more or less pointed. Chaetota^qr of veins (Figs 219, 228): 6 short and stout setae apically on M+Cu; 10 long setae basally on Sc+R, 3 basally on M+Cu, and S on basal half of vein I- lA H nd wmg not visible. Petiole 7.4x as long as lateral width at apex, much longer than abdommal tergite 3, and 0.5Sx as long as gaster. Fifth abdominal tergite with very fine microareolation. 294 Entirely black; wing membrane hyaline, veins black or brown, clear basally on pterostigma. General condition o f the holotype. Complete; most of microsculpture obscured by a series of small air bubbles attached to the body, and in some parts, by a film of pyrite, (ktster partially decomposed, mostly enveloped by a milky substance so that its sculpture is visible only for abdommal tergite 5. Amber piece yellow-brownish, only relatively clear; cut and polished in a polyedral shape, measuring 3.5 x 1.0 x 0.4 cm. It shows a laminated structure characteristic of Baltic amber. One unpolished surface shows small cracks of weatherm& indicating the amber piece was unearthed for some time before it was found. Body parts of a specimen of Pteromalidae (Hymenoptera), a sprrngtad (CoUemboIa), and several stellate plant hairs are embedded together with the stephanid. Comparative notes. Immediately separated from E. brevicomis and E neavenatus sp. n. by the presence of a long, qrlmdric petiole, and from E petioiatus and E tridentatus by the presence of only two teeth ventrally on the hind femur and by a longer petiole Qengtfa/Iateral width at apex 7.4 versus 5.0 fb r& petioiatus and 4.0 forE tridentatus). Further distinguished from E petioiatus by having a short hind coxa Oength/maxhnum height 2.00 versus 2.75 in E. petioiatus) and a deep femoral hnpression, and from E tridentatusby its uniformly black body. 295 EJectros^hanus neavenaùts sp. n. (Figs 223-227,229) Etymology. Greekneo, new; Latin vena, vem, venation. In reference to vein (Rs+M)b and bullae on front wing features apparently rare in fossil Stephanidae. "type material. Holotype male in the collection of the Geological-Paleontological Institute and Nfiiseum o f the University o f Hamburg Germany, Type Number SGPIH 3893. fricluded in Baltic amber (Eocene), from Samland peninsula, Baltic coast. Description. Male ^olotype): Total length 6.7 mm. Head (Fig. 223): scape 1.4x as long as wide; pedicel subglobular; first three fi%ellomeres weakly thicker apically than basally, somewdiat fiattened, first one more distinctly, remaming flageOomeres cylindric. Maxillary palpus 5-segmented; basal segment the shortest and widest; 2nd segment elongate-fiisifbrm, twice as long as 1st; 3rd to Sth segments elongate, all of similar size and slightly longer than 2nd; 3rd and 4th segments thicker apically than basally. Frons transversely arcuate-rugose, with some areolae near right lateral tubercle Qefr side not visible). Vertex with scattered erect hair% which are about as long as length of first flagellomere, sculpture not visible. Gena round, polished smooth. Occipital carma very narrow. Frostemum obliquely allutaceous, with a few fbveolae. Colo differentiated; pronotal fold apparently dKferentiated, weak. Pronotum strigate as follows: transversely dorsally, includmg colo; obliquely laterally, including shallow and broad femoral 296 impression; more or less longitudinally on ventral area; strigation fine dorsally on semiannular, strong otherwise. Semiannular with a few feveolae and several long and stout erect hairs. Pronotal lobe and depression in front of lobe distinct. Median sulcus and notauli distinct, formed by aligned fbveolae; notauli contiguous posteriorly, at transscutal articulation. Front tibia weakly compressed posteriorly, at basal 0.3; a series of strong short, stiff setae aligned on anterior fece, inclined about 45” towards apex. Mesopleuron rugulose and foveolate near pronotum, bearmg several scattered long erect hairs. Discrimen foveolate. Propodeum macrosculptured (details not visible). Metapleuron areolate-rugose; each areola with one erect hair anteriorly, with each hair about as long as diameter of corresponding areola. Inter- and post-foveolar areas strongly transversely strigate, including inside large metapleural fbvea (Fig. 227). All tarsi 5-segmented, ventral side of pre-apical tarsomeres prolonged anteriorly to a distance as long as lenght of apical tarsomere (e.g.. Fig. 224). (find coxa short and stout, almost twice as long as high, more or less “bullet-shapetT in lateral view (Rg. 224); mostly polished smooth; covered with sparse, medium-sized hairs, which are inclined posteriorly at about 45”. Hind femur tansversely alutaceous; with sparse, long erect hairs; without teeth basally, central tooth broadly triangular; apical tooth dagger-shaped ^ g . 224). Hind tibia transversely to obliquely alutaceous; basal half strongly compressed, apical half strongly dilated; pflosity^ as fer femur. Front wing (Fig. 225) with cell 1+2R1 relatively short, only about twice as long as high; vein IRs at distmctly acute angle with Sc+R, and slightly longer than 2r; IM about I.4x as long as m-cu; ICu slightly longer than (Rs^NQa; 2r 0.4x as long as 2+3Rs; 2 9 7 distinct, about as long as 2r; distinct bullae between 2r and vems 2+3Rs and 2+3N^ which is complete ending at short distance &om wing margin. All veins tubular, except apex of 4Rs and 2+3M, and entire 2-lA and 2Cub, nebulous; 2cu-a developed, ^parently spectral. Parastigma long. Pterostigma blunt. Chaetotaj^ of veins ^ g s 225, 229): Sc^R with 11 long, light colored setae along its entire length, the setae more or less equidistant from each other; M-K^u with same ^ e of setae on basal 0.8, apically with 7 short and stout setae on left win& two of them beyond IM; vein 1-lA with 5 long setae within its basal haK and 2 more setae apically, these about half as long as basal setae. Hnd wing ^ g . 226): Sc+R tubular, stout, as thick as or thicker than I-IA on front wing. Anterior wing fold developed. Four hamuli on left wing, which are straight, but bent basally, thus decumbent [not visible on illustrated right wing]. Veins IRsb and 2M visible on apical half of win& nebulous. True petiole absent, abdominal T2 and S2 separated; T2 distinctly longitudinally rugulose, stronger basally; about 0.85x as long as T3, and 0.3Ox as long as gaster. T3-9 polished smooth (observed laterally). T9 apical margin simple, round. Stemhes with centro-longitudinal crest, or keel, which is much stronger on S5-7 (as in males of Stephanas senator). Black; whitish spot on malar space. Front tibia basally discolored (white?). ) ^ g s clear hyaline. Vems brown; clear basally on pterostigma. General condition o fthe holotype. Right mid le^ flageHomeres after 4th on right antenna, and after 9th on left, missing. All fine sculpture of mesothorax obscured by unifonn film of air. Gaster with whitish substance covering all tergites centrally. 298 Amber piece clear pate yellow, polisiied m a rounded sliape, measuring 3.5 x 1.5 x 0.5 cm. ADolicliopodidae (Diptera) and a mite nymph (Erythraeidae, probabfyZ^mr sp.) are anbedded together with the stephanid. Comparative notes. The absence of a true petiole, with T2/S2 short, wid^ and cleariy separated, malces tliis species similar only to K brevicomis, from wliich it can be distinguished by its more specialized wmg venation (key characters) and differently structured maxillary palpus (see description). Taxonomic status and relationships of Protost^hanus and Electrostqthanus The genus Protostephanus Cockerell (1906) is very similar to, and could be a senior synonym of Electrostephanus Brues (1933), based on its type species K brevicom is Brues. The establishment of this synonymy, however, is premature because of the following problems. (1) Protostephanus is based on a female, making it very dffîcult to compare with EkarosApAomw, for which only male specimens are known. As discussed in Aguiar (1998), wing venation, biometric ratios, and other characters of taxonomic importance are usually not readily comparable between males and females of Stephanidae. (2) Several characters of taxonomic importance could not be observed or confirmed for the two taxa. This was due to the poor condition and weakly informative fossilized position (only ventral side visible) of the single specimen of Protostephanus, and because several characters o f the wing venation o îElectrostephanus could only be 299 determined from an illustration inBrues’s paper, which does not seem to be entirely accurate (eg., shape o f right and left wing veins are not exactly the same), rendering the drawing of questionable value fr)r taxonomic discussions. In the origmal description o fElectrostephanus, only two characters were explicitly mentioned by Brues as being different from other stephanid genera: “Venation of the complete type, as in Stephanus, but differing in that the cubitus [Icu-a] forms an angle with the lower part o f the basal vein [IM], not a continuous line,” and “ .. differs from aU living forms of Stephanidae in the small number of joints in the antennae [17-23] . . . other recent genera have at least 32 joints”. The taxonomic value of both these features, as well as all other characters Brues used in defining the genus, is not corroborated by what one of us (APA) has observed along extensive taxonomic work with the Stephanidae (Aguiar & Musetti 1998): Veins Icu-a and IM do form an angle in the genus Stephanus, as well as inMegischus, a related genus. It may appear aligned in more apical groups (e.g., some Foenatopus), but not without showmg intraspecific variation. The number of antennal segments also varies widely within the family, appearing to be correlated with the size o f the specimeiL In &ct, a reduced niunber of antennal segments appears to be a general trend among males of Stephanidae, which are generally much smaller than the females. This is especially relevant, considering that all known spechnens o f Electrostephanus are males. Males of the Australian ParastephaneUus rufoomatus (Cameron), for «cample, are among the smallest stephanids known, often with overall length under 5 mm, and may have as fow as 12 antennal segments, below the known range fa t Electrosteplumus, 300 Nonetheless the following character states described for the genom e species, E. brevicomiSy by Brues (1933) are either unique or highly unusual among Stephanidae: (1) structure o f maxQlary palpus C^three basal segments much thickened, short; apical ones very long and slender^); (2) vein Sc+R, and apparently also 1-1 A, with setae along all their length; (3) vein (Rs+NQb absent; (4) vein 2r very small, much shorter than 2+3Rs; (5) no bullae at origin of vein 2+3Rs and/or 2+3M; (6) structure of basal flagellar segments (“first three flagellar joints widened apically [ ...] flattened, as when viewed fi*om the side they are not thickened apically”); (7) Ion& well developed parastigma; (8) vein IM arched; (9) vein IM distinctively longer than m-cu; (10) cell 1+2R1 about twice as long as wide; (11) veins Rs+M and ICu not parallel; and (12) petiole short and stout. Only characters 6-12 occur in extant stephanids, namely Schlettererius spp. and Stephanus serrator (Fabricius). The apomorphic condition of the states listed above cannot be accurately hypothesized without a cladistic analysis. Nonetheless, th ^ form a unique combination, not yet observed for any living species in the fomily, which suggests that E brevicomis probably cannot be properly classified in any of the extant genera. Character states 1,2,6, and 11 abov^ plus the apparent absence of veins 2Cua (suggested in Brues’s illustration) m ake& brevicom is also considerably different fi*om all other known Baltic amber species, ht fitct, Brues had already noted that “Possibly this species should be placed in a distmct genus,” but declmed doing so, reasoning it would be a premature decision. The genus Electrostephanus, therefore, may indeed represent a valid taxon, but its precise definition and classffication, includmg the possibility of monotypy, or synonymy 301 'v^Prostostephanus, are still problematic. The major difficulty is that most taxonomically important characters at genus level vary widely among the known Baltic amber specie^ mcluding the two new taxa described in this publiortion. From the character states listed abov^ for example only numbers 4 and 8-11 seem to be present mall Baltic amber species. All other characters taxonomically important at the genus-level either could not be properly observed for all species, or lead to conflicting conclusions. For example, K brevicom is and K neovemtus sp. n. share a distinctively short and stout “petiole,” but while the petiole is undoubtedly formed by an isolated terghe and stemite in K neavenatus sp. n., its exact structure was not described and remams unknown for& brevicomis. Additionally, the structure of the basal antennal segments and of the maxfllary palpus, and some characters o f the wing venation for K neovenatus sp. n., are also all considerably different from those ofE. brevicomis. On the other hand, the wing venation for £1 sulcatus sp. n. is almost entirely the same as that oîE. brevicomis^ but the petiole of the former is elongate and cylindric, with terghe and stemite fused. Similar differences occur among aiy two of the five currently known Baltic amber species. The level of morphological variation discussed in the previous paragraph is usually sufficient to ju s t^ recognizing genus-level taxa in extant Stephanidae, but the fiict that such differences were observed in several individual specimens, renders taxonomic decisions based on them too costly, that is, too many insufficiently understood genera would have to be created to accomodate them. 302 A further problem is the similarity^ and, at the same tim^ the taxonomic incompatibility between the genders for the available specimens of Electrostephanus and Protostephanus, discussed previously. Finally, none o f the characters described by Brues could be confirmed, since all his types are now missing. A more stable solution fi)r most o f the problems discussed above can probably be achieved by a detailed cladistic analysis o f the stephanid genera, but this is beyond the scope of the present publication. For this reason, and to prevent generation of further taxonomic problems by erecting new genera based on taxonomic characters which are not well understood, the revised diagnosis of the genus Electrostephanus Brues presented here is only provisory. It was a necessary step, however, because Electrosteplumus was originally defined by features of no taxonomic relevance, as already discussed. Relationships with extant species. Both Electrostephanus and Protostephanus lack important synapomorphies of most of the apical groups of extant Stephanidae, from which they can, therefore, be safely occluded. Some plesiomorphic states for fossil species are (1) veins m-cu, R, and M, tubular (spectral in Foenatopinae) and (2) short petiole or setae on M-K^u placed apically (setae on M+Cu placed centrally in Hemistephanus; petiole long and tylindric in all extant Stephanidae accept Schlettererius). Additionally, vem IM is arched and distinclty longer than m-cu m all fossil species ^ g s 217,220,225), two plesiomorphic conditions which exclude the fossil species from most extant stephanid genera, where IM is straight and as long as, or shorter, than m-cu. The only acceptions are Schlettererius Ashmead, and Stephanus Jurm^ which are also, by fer, the most similar genera (morphologically) to Protostephanus and Electrostephanus. 303 CONCLUSIONS (1) Contrary to the opinion of Brues (1933), Protostephanus ashmeadi Cockerell is a species of Stephanidae. Its holotype is a female. (2) The genus Electrostephanus Brues, when interpreted solely on the basis of its genotype, K brevicomis^oxes^ shares many important genus-level features with Protostephanus Cockerell. However, the final decision on whether or not these two genera should be synonymized depends on further discovery o f males o fProtostephanus and on a more precise understanding of the systematics of male stephanids. (3) Establishing the synonymy would also require a redefinition o f Electrosteplumus relative to the remaining species. However, the five known species in this genus show together a greater variation in most genus-level characters than is observed among the species o f extant genera of the family, which suggests that more than one genus would have to be recognized. (4) As feras investigated here, fossil species of Stephanidae lack important tynapomorphies of most of the recent genera, firom which, therefore, fossil species can «q)parently be safely occluded. This feet suggests a basal placement for all fessU species in the family, probably near the extant genera Schlettererius and Stephanus. (5) I suggest that the next most adequate step m mvetigating these problems is a cladistic analysis o f the Stephanidae including male specimens. 304 FIGURES 214-217 Protostephanus ameadi Cockerell; 214, head; 215, body ventrally, from prostemum to near apex of abdomen; 216, front left wing, ventral side; 217, detail of structure o f cell IM and related venation. Scale bars: 214,217,0.25 mm; 215, 0.50 mm; 216, 1.00 mm. 305 306 FIGURES 218-222 Electrostephanus sulcatus sp. n.: 218, habttus; 219, front wing; 220, hind trochantellus, trochanter, and femur; 221, metapleuron and propodeum, lateral view; 222, propodeum, dorsal view. 307 ma 222 308 FIGURES 223-229 Electrostephanus neovenatus sp. n.: 223, head, fronto-Iateral; 224, lefb hind Ie& external side; 225, left fi-ont wing; 226, apical 0.75 of right hind wing; 227, metapleuron and basal section of “petiole” in latero-ventral view, left side; 229, setae on apex of vein M-K^u, left wing inverted, dorsal side. Electrostephanus sulcatus sp. n.: 228, setae on apex o f vein M-K^u, right wing dorsal side. 309 01£ ) ”D+Vi % PHYLOCæNETICS AND SYSTEMAnCS OF THE WORLD STEPHANIDAE (HYMENOPTERA) Volume H DISSERTATION Presented in Partial Fulfillment of the Requirements for the Degree o f Doctor of Philosophy in the Graduate School o f The Ohio State University By Alexandre Pires Aguiar, M.S. ***** The Ohio State University 2000 Dissertation Committee; Approved by Norman F. Johnson, Adviser Woodbridge A. Foster HansKIompen EntomologrSraauate Program JohnW. Wenzel TABLE OF CONTENTS ListofTables...... xxx ListofRgures...... xxxi Chapters: 3.Revisionofthe Australian Stephanidae ...... 311 Introduction...... 311 Material and methods ...... 313 General...... 313 Morphometricanalysis ...... 315 Spheridtymdex ...... 316 Identffîcation ...... 317 Sex récognition ...... 317 Morphometricratios ...... 318 Body size and interpretation ofcharacter states ...... 319 Key to AustrahangeneraofStephanidae ...... 320 Genus Schlettererius...... 321 GemxsMegischus...... 323 Key to AustraliansgeàssoîMeffschus ...... 323 M crassicaudaÇ^oûe^) ...... 324 M cenetmatrvcxL sp ...... 328 GenusParastephanellus ...... 331 Groundplan ...... 333 Kqrto species ...... 338 Descriptions ...... 349 P. maculatusn. sp ...... 349 P. brevisti. sp ...... 354 P. laevigatustL sp ...... 358 P. isog^tosïL sp ...... 363 P.dbme///ct£s(Westwood) ______368 P. albigenuisTL sp ...... 376 P. aeffazetosTL sp ...... 380 xxvui P, callidictyon n. sp ...... 3 84 P. tarytarsusvL sp ...... 388 P.granulatus(JE3!lio\£)...... 393 P. platyceratusTL sp ...... 397 P.strigosusrirSÇ...... 401 P, trifasciatusvLsp ...... 406 P. squamatusTL sp ...... 411 P. H{/bo/7iaA£s(Cameron)...... 417 P. vertipictusiL sp...... 423 P. denotatusvL sp ...... 427 P, caudatusW iotX...... 431 Species mquffendum...... 436 Stephanid taxamîstakenlyreportedûiAustralia...... 438 Plates ofOIustrations...... 440 List ofReferences...... 496 APPENDIX Ranges of morphometricratios for species ofAxx^XxdlimParastephanellus ...... 506 XXIX U SX OF TABLES Table Page 3.1 Selected morphometric ratios evaluated for taxonomic usefulness with species of Australian ...... 318 3 Morphometric ratios to be avoided in taxonomic decisions with species of AustralianParastephaneUus ...... 318 3.3 Number of specimens studied (n) for each species of Australian ParastephaneUus^ correspondent percentage (%) in relation to total number of specimens, and indication o f species for which males are unknown...... 332 3.4 Main differences between morphotypes off. squam atus n. sp ...... 416 3.5 Intermediate characters in the holotype of P, ptcticeps Roman...... 437 3.6 Minimum and maximum values of taxonomically important morphometric ratios for Australian species o fParastephaneUus, Females...... 506 3.7 Minimum and maximum values of taxonomically important morphometric ratios for Australian species o îParastephaneUus, Females...... 507 3.8 \finimum and maximum values of taxonomically important morphometric ratios for Australian species o îParastephaneUus, Females...... 508 3.9 Minimum and maximum values of taxonomically important morphometric ratios for Australian species o îParastephaneUus, Females...... 509 3.10 Minimum and maximum values of taxonomically important morphometric ratios for Australian species o î ParastephaneUus, Females...... 510 XXX USTOFnGÜRES Figure Page M ep sch u s crassicauda Morley: 230 Vertex ...... 441 231 Pronotum...... 441 232 Propodeum ...... 441 233 Metapleuron...... 441 234 Pronotum, mesoscutum, and axillae...... 441 235 Hind leg view oftrochanters trough tibia ...... 441 236 Axillae, scutellum, and crenulatesulcus...... 441 M epschus ceneonatrix n. sp.: 237 Vertex ...... 443 238 Pronotum...... 443 239 Propodeum ...... 443 240 Metapleuron and propodeum ...... 443 241 Pronotum, mesoscutum, ax21a^ andpartofscutellum...... 443 242 Hind 1% outer side view offèmur and tibia ...... 443 243 Detail ofhindfèmur, outerside ...... 443 244 Locality records fbrM crassicaudaMoxie^ andM ceneonatrix n. sp ...... 445 245 Localrtyrecordsfor&Afe/ferermyancftpe^CCameron) ...... 445 246 SeazonaIdistnl)utionoO Measurements used to calculate morphometric ratios, ParastephaneUus vertipictus n. sp.: 248 Body in lateral view ...... 447 249 H ead ...... 447 250 Pronotum...... 447 251 Tergites 3-4 ...... 447 Values o f morphometric ratios for all examined female specimens o f all species: 252 Ratio Ov/Ptl...... 449 253 Ratio Ov/Tt...... 449 XXXI 254 Ratio Ov/Hdl ...... 449 255 Ratio Ov/Saw...... 449 256 RatioCxI/h ...... 449 257 Ratio CI/Fml...... 449 258 RatioFmSI ...... 449 259 Ratio tw^EE ...... 449 260 RatiobtI/w...... 451 261 Ratio Pnl/Saw...... 451 262 Ratio twAo...... 451 263 Ratio Pnl/Ew...... 451 264 Ratio Gsl/Ptl...... 451 265 Ratio Ptl/T3l...... 451 266 Ratio Hdl/Ew ...... 451 267 Ratio Ew/tw...... 451 268 Ratio îEE/eEE ...... 453 269 Ratio iEE^o ...... 453 270 Ratio Ppl/Ew ...... 453 271 Ratio io/oo...... 453 ParastephaneUus maculatus n. sp.: 272 Head sculpture...... 455 273 Pronotum and mesoscutum...... 455 274 Axillae, scutellum, basai section of propodeum...... 455 275 Pronotum...... 455 276 AxOIae, scutellum, propodeum...... 455 277 Posterior half ofmesepistemum, metapleuron, propodeum ...... 455 278 Hind fémur and tibia, outerside ...... 455 279 Tergites 6-9 and pygidium ...... 455 ParastephaneUus brevis n. sp.: 280 Head sculpture ...... 457 281 Pronotumand mesoscutum...... 457 282 Tergites 7-9 and pygidium ...... 457 283 Mesoscutum sculpture...... 457 284 Pronotum...... 457 285 Apec of scutellum, propodeum ...... 457 286 Propodeum and metapleuron ...... 457 287 Right front win^ venation centrally...... 457 288 Hind femurand tibia, detail ofsculpture, outerside ...... 457 ParastephaneUus laemgatus n. sp.: 289 Headsculptureand colour, morphotype 1 ...... 459 290 Same, morphotypeü ...... 459 xxxu 291 Pronotum...... 459 292 Axillaeandscutellumsculpture...... 459 293 Same, difièrent specimen ...... 459 294 Pronotum...... 459 295 Propodeum ...... 459 296 Thorax, except pronotum ...... 459 297 Propodeum, detail ofsculpture ...... 459 298 Mesepistemum ...... 459 ParastephaneUus isogïyptosn. sp.: 299 Head sculpture ...... 461 300 Pronotum...... 461 301 Scutellum and propodeum...... 461 302 Metapleuron and propodeum ...... 461 303 Pronotumand mesoscutum...... 461 ParastephaneUus damellicus (Westwood): 304 Head sculpture...... 463 305 Pronotum...... 463 306 Pronotum, detail ofsculpture...... 463 307 Axillae and scutellum...... 463 308 Pronotum...... 463 309 Axillae and scutellum, second specimen ...... 463 310 Scutellum, detail ofsculpture, third specimen...... 463 311 Apex of scutellum and propodeum ...... 463 312 Mesepistemum...... 463 313 Propodeum and metapleuron ...... 463 314 Propodeum, detaü ofsculpture...... 463 ParastephaneUus albigenuisn. sp.: 315 Head sculpture...... 465 316 Pronotum...... 465 317 Mesepistemum posterior half and metapleuron ...... 465 318 Head posteriorly and pronotum ...... 465 319 Thorax, except anterior portion of pronotum ...... 465 320 Axillae, scutellum, basal halfofpropodeum ...... 465 321 G aster...... 465 ParastephaneUus aeUazetosTL sp.: 322 Head sculpture...... 467 323 Pronotumand mesoscutum...... 467 324 Scutellum and basal halfofpropodeum ...... 467 325 Pronotum...... 467 xxxm 326 Thorax ...... 467 327 Mesepistemum posteriorhalÇ propodeum and metapleuron ...... 467 ParastephaneUus œllidîctyon n. sp.: 328 Head sculpture ...... 469 329 Vertex sculpture, détail ...... 469 330 Frons...... 469 331 Scutellum...... 469 332 Pronotum and mesoscutum...... 469 333 Pro^emum...... 469 334 Pronotum, detail ofsculpture...... 469 335 Scutellum apex and propodeum ...... 469 336 Mesepistemum posterior halÇ propodeum and metapleuron...... 469 337 Pronotum...... 469 ParastephaneUus tarytarsus Ti. sp.: 338 Head sculpture ...... 471 339 Head sculpture anteriorview ...... 471 340 Pronotum and mesoscutum...... 471 341 Right front wing, venation centrally ...... 471 342 Pronotum...... 471 343 Scutellum apex and propodeum ...... 471 344 Axillae andscuteHum...... 471 345 Prostemum, ventral...... 471 346 Mesopseudosteraum, discrimen...... 471 347 Hind femur and tibia, outer side...... 471 ParastephaneUus granulatus (Elliott): 348 Head sculpture ...... 473 349 Pronotum and mesoscutum...... 473 350 Rightfront wing, venation centrally ...... 473 351 Pronotumandanteriorhalfofmesepistemum ...... 473 352 Prostemum, ventral...... 473 353 Mesepistemum...... 473 354 Scutellum and propodeum...... 473 ParastephaneUusplatyceratusn. sp.: 355 Head sculpture...... 475 356 Head posteriorly and pronotum ...... 475 357 Pterothoraxand propodeum ...... 475 358 Thorax, except pronotum ...... 475 XXXIV ParastephaneUusstrigosus n. sp.: 359 Head sculpture ...... 477 360 Pronotum, detail...... 477 361 Propodeum andmetapleuron ...... 477 362 Head sculpture, detail...... 477 363 Pronotum...... 477 364 Axillaeand scutellum...... 477 365 Rightfrontwing ...... 477 ParastephaneUus trtfasciatus n. sp.: 366 Head sculpture...... 479 367 Pronotum...... 479 368 Head, anteriorview ...... 479 369 Pronotum...... 479 370 Mesoscutum, axillae, scutellum...... 479 371 Same, different specimen ...... 479 372 Mesepistemum...... 479 373 Mesepistemum posterior hall^ propodeum and metapleuron ...... 479 ParastephaneUus squamatus n. sp.: 374 Head sculpture...... 481 375 Pronotum and mesoscutum, morphotype 1...... 481 376 Head sculpture, posteriorview ...... 481 377 Pronotum, morphotypell ...... 481 378 Pronotum posteriorly, mesoscutum, axillae, scutellum...... 481 379 Pronotum...... 481 380 Thorax, exceptpronotum ...... 481 381 Head and thorax ...... 481 ParastephaneUus rufoomatus (Cameron): 382 Head sculpture ...... 483 383 Pronotum and mesoscutum...... 483 384 Thorax ...... 483 385 Pronotum...... 483 386 Mesepistemum ...... 483 387 Mesothorax ...... 483 388 Mesothorax ...... 483 389 Mesepistemum posteriorhaK propodeum and metapleuron ...... 483 390 ScuteUumandpropodeum ...... 483 XXXV Parasiephanellus vertipictus n. sp.: 391 Head sculpture ...... 485 392 Temples, pronotum, mesoscutum...... 485 393 Scutellum...... 485 394 Pronotum and mesothorax ...... 485 395 Prostemum, ventral...... 485 396 Thorax, except pronotum ...... 485 397 MesepîstemumpostenorhalÇ propodeum and metapleuron ...... 485 Parastephanelfus denoiatus n. sp. ; 398 Head sculpture ...... 487 399 Pronotum and mesoscutum...... 487 400 Thorax ...... 487 401 Scutellum and propodeum...... 487 402 Hind femurand tibia, outer side...... 487 403 Tergites 7-9 and pygidium, detail ...... 487 Parastephanellvs caudatus n. sp.: 404 Head sculpture...... 489 405 Pronotum and mesoscutum...... 489 406 Axillae and scutellum...... 489 407 Pronotum...... 489 408 Propodeum ...... 489 409 Propodeum and metapleuron ...... 489 410 Tergites 6-9 and pygidium ...... 489 Locality records for species oîParastephanelhts occurring in Australia: 411 AUspp ...... 491 412 P. isoglyptosn. sp ...... 491 413 P, maculatusn, sp ...... 491 414 P. brevistL sp ...... 491 415 P. laevigatusTL sp ...... 491 416 P. albigemnsn. sp ...... 491 417 P^aeüæetosTLsp...... 491 418 P. callidictyon n. sp ...... 491 419 P. dbwe///ctt^(Westwood) ...... 491 420 P. tm^ctrsusn. sp ...... 491 421 P. granulatus (Elliott)...... 491 422 P . platyceratusn. sp ...... 491 423 P.strigosusTLsp...... 491 XXXVI Locality records for species o f Parastephanellus occurring in Australia: 424 P. trifasdatusxL sp ...... 493 425 P,vertipictusTLSÇ...... 493 426 P,squamatusn,sç ...... 493 427 P,denotatusTLSÇ ...... 493 428 P. n^oornahis(ÇamtcovC)...... 493 429 P, cawrfofttf Elliott...... 493 430 Seasonal distributionofAustralianParartepAonc/fttf...... 495 Seasonal distribution for species of Australian Parastephanellus known foom seven or more specimens: 431 P.rufoomatus...... 495 432 P.damelücus...... 495 433 P.isogfyptosn.sg...... 495 434 P. trifasciatusvL sp ...... 495 435 P.strigosusn.sp...... 495 436 P. platyceratusn. sp ...... 495 437 P,laevigatusn,sç ...... 495 438 P.squamatusTLSÇ ...... 495 xxxvu CHAPTERS REVISION OF THE AUSTRALIAN STEPHANDDAE (HYMENOPTERA) INTRODUCTION Current knowledge about native Australian stephanids is very limited, consisting mainly of sbc isolated species descriptions (Westwood 1874, Cameron 1905, Roman 1915, Morlqr 1917, Elliott 1919a, Girault 1926), and a description of the oviposition behaviour and early developmental stages for an ‘Hinidentified” species (Rodd 1951) {^Pearastephmellus rufoomatus, see this woric). More recent publications include a series of papers by Taylor (1967a, 1967b, 1976, 1981) that deal with the introduction, biolo^, and status ofS, cinctipes, imported into Tasmania from California (USA) to help control the pine wasp, Sirex noctilio Fabrichis. Hosts of the native stephanid species are virtually unknown, but Rodd (1951), mentioned above, and Moore (1962), reporting on a ‘^ffemistephanusT (very likely a ParasIî^Aane/Ttts species), observed oviposition and emergence on logs infosted with bostrichid beetles. Native stephanids occur in most of Australia, except apparently in Tasmania. This, however, could be a consequence o f sampling. Furthermore, the apparent madequa(y of 311 Nfolaise traps (Aguiar 1998) and yellow pan traps (Aguiar & Sharkov 1997) in collecting stephanids is probably an important 6ctor Ibniting the representation o f these insects in most collections. The body plan o f stephanids is rather uniform throughout the fomQy, but intra- and mterspedfic variation o f taxonoroically significant characters is usually high, making species difiScult to recognize and characterize. Evidently, such dffîèrences can only be detected through comprehensive morphological comparisons, but the foct that stephanids are almost invariably considered “rare” (which is not always the case; see Aguiar & Sharkov 1997) has been fi'equently used by authors to justify the proposition of new stephanid taxa based on single specimens. Because of this, most of the taxonomic literature on stephanids has, ironically, very little value for identffîcation purposes, as th ^ provide no perspective on the large and complex intra-specific variability o f the group. Another consequence is that character sets used by difièrent authors vary greatly, making most available descriptions not, or only marginally comparable. Considermg that morphological variabilify is particularly intense among the Australian stephanids, it seems fortunate that casual studies on this group are few, as much taxonomic confiision was undoubtedly prevented. On the other hand, knowledge about these species is o f considerable importance for the systematics o f stephanids in general. This is mostly because of the country’s unique and diverse fauna of Parastephanellus^ a taxon which phylogenetically, as well as geographically, occupies an intermediate position between Western and Oriental taxa of the fiunily (Aguiar, in 312 preparation). This becomes of further interest in 6ce of past connections between Australia and South America, and of close morphological similarities between some stephanids from these areas, especially some Parastephanellus and Hemisiepharms. The main objective of this work is to provide the first revision ofthe Australian stephanids, but it is also aimed at contributing to a better understanding of the genus Parastephanellus, mainly by investigating the morphological variabili^ of its species in Australia, providing mfôrmation which will be useful m establishing homologies between this and other stephanid genera. The focus of this work, however, is at the species level. A fiiU taxonomic revision and phylogeny of the stephanid genera will be published elsewhere. MATERIAL AND METHODS A total o f347 spechnens were examined, including all relevant type material, except the holotype of Stephanas (^Parastephanellus) damellicus Westwood, originally deposited in the Oxford Museum, England, but not found in this institution in a recent search. Specimens are deposited in the institutions listed below, which are referred in the text with abbreviations from Arnett e t aL (1993). AEIC American Entomological Institut^ Gainesville, Florida AMSA Australian Museum, Sydn^ ANIC Australian National Insect Collection, Camberra BMNH The Natural History Museum, London BPBM Bernice P . Bishop Museum, Honolulu, Hawai 313 FCNI Forestry Comission of New South Wales Insect Collection, Beecroft MVMA Museum ofVictoria, Melbourne NHRS Natudiistoriska Riksmuseet, Stockholm NSWA Orange Agricultural Institute, New South Wales QDPC Queensland Department of Primary Industries, Brisbane QMB A Queensland Museum, Brisbane SAMA South Australian Museum, Adelaide UCRC University of California, Riverside UQIC University of Queensland, Brisbane USNM United States National Museum, Washington, DC WAMP Western Australian Museum, Path ZMAN histhuut voor Taxonomische Zoologie, Amsterdam ZMUC Zoologisk Museum, University of Copenhagen, Denmark All specimens were barcoded and databased. Label data for all specimens and species are available at http://iris.biosci.ohio-state.edu/projects/stephanids, and, except for holotype^ will not be repeated here. Morphological terminology and general methodolo^ follows chapter 1. Codens for lodging mstitutions follows Arnett e t cd. (1993). Observations on color and sculpture were always made with fluorescent illumination. Incandescent illumination without a light dffîisor will not allow observation of color and structures as described and illustrated in the present work. All illustrations and photographs were prepared &om female specimens. 314 The term “mîcroacîculate” as used here refers to an especialized kind of strigation, which is very fin^ with a somewhat “vetvef* tecture. It occurs on different abdominal tergites of many species. Morphometric Analysis Selection of morphometric ratios and investigation on their relative accuracy and usefulness in separating species were based on an evaluation of the Parastepfumeüus spp. only, and performed only after all species had been “qualitatively” defined. Only females were measured, following recommendations by Aguiar (1998). Morphometric ratios throughout this work are therefore inappropriate for the identification of male specimens. Measurements are illustrated on Figs 248-251. Ratios are represented by using their original measurements, e.g., Ov/Tt, Pnl/Hdw. Letters‘T , “w”, and “h” at the end stand for length, width, and height respectively. Lettos “v” and “c” in the begining stand for vein and cell respectivelly (wing venation ratios), ^tw o dimensions of the same structure are used in a ratio, only the letter for the dunension of the dividing measurement is used in representing the ratio, e.g., Cxl/h instead of Cxl/Cxh. The only exception is ratio FmSI, explained below. A total o f 45 morphometric ratios were investigated (Tables 3.1 and 3.2). Prelimmaty data were organized in graphs as those in Kgs 252-271, and selection of the best ratios was performed m two step^ based on the evaluation of these graphs: (1) an elunmatory evaluation, using five species, taxonomicaHy related to each other in the form (a((bc)(dc))), where a= P . ntfoorruOus^ b = f . kuvigatusn. sp., c =/*. damellicus, d = f . 315 trifasciatus n. sp., e=A strigosus n. sp. This approach allowed each ratio to be evaluated for its power in separatmg, as well as grouping closely related (b vs. c, d vs. e, and be vs. de) or more distantly related (a vs. bcde) species. And (2) an overall subjective evaluation o f the selected ratios, considering all species, case by cas^ to determine more accurately the relative importance of each ratio. The greater the non-overlapping range (proportionally to the overall range o f the ratio for all species), the better and more accurate was considered the ratio. For step two, all species were measured, and ratios were selected and ranked according to then usefulness in separating species or groups of species found to be closely related morphologically in the qualitative analysis. Minimum and maximum values for the selected ratios, for all species, are presented in tables 3.6 to 3.10 (Appendix). Individual values of all ratios for all specimens and species were organûed graphically, in Figs 252-271. Species numbers in these graphs (Y axis) correspond to those in tables 3.6 to 3.10. This numeration reflects the overall simflarity between species, so that species in close proximity along the Y axis are more similar morphologically to each other than species for apart To mitigate problems with allometry, the relative size of a given structure is sometimes illustrated by more tlian one ratio in the descriptions. For example, the ratios Ov/Ptl, Ov/Tt, and Ov/Hdl can be used all at once to support a given statement about the relative length of the ovipositor. SphericiQr îndmt (SI). The overall shape of the hind fomur is important for dif&rentiating many stephanid species, but femoral shape is difficult to interpret objectively because of significant intra- and mterspedfic variability. Simple ratios, as 316 length/height or length/width, can be used, but then at least two different values are generated to evaluate the three-dhnensional structure of a femur. Instead, the following formula was used, based on the division of the ratio length/height by the ratio width/height, thus taking into consideration all three axial dimensions of the hind femun FmSI = ^m length. Fm width)/(Fm height)* The formula returns a value of 1.00 for a perfect sphere, and increasingly greater values for more elongate or flattened/compressed shapes. Extreme values found in the present work are 1.33 and 2.63. Identification. Species identification for most stephanids is done in great part by interpretation of the cuticular sculpture, which is inherently difScult to interpret. Males and fomales of a same stephanid species are generally similar to each other, but variability in colour, morphometric proportions, and sculpture patterns are usually much greater and more irr^ular for males. and descriptions, therefore, may be less efScient when used with male specimens. Sex recognition. Secondary sexual dimorphism for males of Australian AfogMcAnr and Parastephanellus include a S-segmented hind tarsus (3-segmented on females), abdomen flattened dorso-ventrally (tylindric or laterally compressed on females), and size usually small or very small. Males ofSchlettererius have the last two features only; the number o f segments in the hind tarsus is five for both soces. 317 Morphometric Ratios. Twenty ratios were selected as taxonomically relevant in separating and characterizing species o f Australian ParastiepAane/7as^ (Table 3.1). Ratios listed on Table 3.2 have been used in descriptions of stephanid species by some authors, but th ^ have low or no taxonomic value for Parastephanellus^ and shold be avoided in descriptions and species interpretation. Differences in importance for each of the ratios selected m this work were much less pronouced than for those selected îor Hemistephanus (Aguiar 1998). Because of this, I ranked the present ratios in “groups” of importance (Table 3.1), rather than discussing them one by one, as in the mentioned work. Ov/Pü CxI/h btl/w GslÆÜ iER/eHR Ov/Tt CxI/Fml Pnl/Saw Ptimi iEE/io Ov/Hdl FmSI twAo Hdl/Ew Ppl/Ew Ov/Saw tw/iEE Pnl/Ew Ew/tw io/oo Table 3.1. Selected morphometric ratios evaluated for taxonomic usefulness with species of AustralianParastephanellus. Ratios in each column nearly equally accurate but progressively less accurate from first to last column. Measurements as in figures Figs 248- 251. See text for explanation of FmSL fl3/fll Gw/Oc PnI/T4I vMl/cMIi Ppl/h io/bl Oc/cw Sal/w vMI/cRl Ptl/w lo/Oc Hdl/Gw Msl/h cRl/h Ptl/MsI lEE/Gw Pnl/cw R/M cMl/h T31/T4I tw/Oc Pnl/Sal vMI/vRsl cRl/cIM T4I/W Table 3.2. Morphometric ratios to be avoided in taxonomic decisions with species of AsxsMsSasLParastephanellus. Mrasurements as in Figs 248-251. R and M (3rd column) correspond to area of cells 1+2R1 and IM. 318 No special investigation was conducted for iSa&Megischus spp., but all morphometric ratios tested for Parasiephanellus spp. were calculated for the two Australian species ofMegischus, and ratios showing non-overlapping values included in the species description^ in decreasmg order of accura^. Body Size and Interpretation of Character States.Variation in body length o f Australian stephanids is extreme, «(tending from 2 mm in some males of Parastephanellus rufoom atus (Cameron), the smallest known stephanid, to a maximum of 33 mm, ovipositor excluded, for some females ofM crassicauda (Moriey), one o f the largest species o f the fomily. IntraspecrGc variation for fomales is moderate, and most species have specimens which may diffor l.S-2.0x in length; but the disproportion can be as much as 4.Sx in females of P. rufoomatus. Extreme values between males and fomales are similar, but males are consistently smaller than fomales (exact values not assessed). As previously observed for Hemistephanus (Aguiar 1998), morphological variation for small specimens is more intense than in large ones. Consequently, it is often much more difficult to interpret characters, and thus characterize species, for small or very small sperànens On relation to the mean for each species). This problem is particularly relevant for identifying males, which are generally much smaller than females. The major tendency is a lessening o f intensify and densify of micro sculpture patterns in smaller spechnens. Stronger sculpture for small specimens also occurs, but is less common. 319 RESULTS Key to Australian Genera ofStephanidae The following is intended to be used for the Australian âuna only, and may not work properly with stephanids from other regions. 1 Vein 2+3M with apical half spectral, or at most weakly nebulous. Cell 2Cu entirely open posteriorly (vein 2-1A absent) (Fig. wI6). Vein 2+3Rs contiguous with (Rs+^Qb (Hg. wI6). Vein 2Cub spectral or weakly nebulous. Ovipositor sheaths unicolorous...... ParastephaneUtis^éexiàn. r Vein 2+3M almost entirely tubular, becoming nebulous only at apex, near wing margin. Cell2Cu closed posteriorly (vein 2-1 Acomplete, tubular) (Fig. wml). Veins 2+3Rs and (Rs+NQb not connected (Fig. wml), or at least appearing as broken, rarely contiguous. Vein2Cub complete, tubular orat least strongly nebulous. Ovipositor sheaths with a distmct uiute pre-apical band 2 2 Second abdommal segment comprized ofvery long and slender, closely transversely strigate petiole. Pronotum long, subconical. ESnd wing venation mostly restricted to ScfR (Fig. wl I). Abdominal tergites 3-9 dark-brown or black ...... MegischusBrvM T Second abdominal segment with tergite and stemite separate, short and wide (only slightly longer than T3), mostly polished smooth. Pronotum blunt, with a deep, 90** step nnmediately in front o f mesonotum, fr>rming a flat, polished 320 wall. Hmd wing with well developed venation, although veins mostly nd)uIous (Fig. w42). Abdominal tergites 3-4 yellowish-brown ...... ...... &MettereriusP%^asaud Genus Schlettererius Ashmead For a ^onym ic list, see chapter I. Comm ents. This genus contains two known species, S. cmctipes (Cameron), native to eastern United States, and S. determinatoris Madl, &om North Korea. Of these, only S. cm ctipes occurs in Australia. This species is easily recognized by the distinctive yellowish-brown T4-5, and the white marks apically on the femora and basally on the tibiae o f all legs. Females measure mostly between 1.5-2.0 cm in lenght, reaching up to 3 cm, excluding the ovipositor, but and males are smaller, normally between I.G-1.5 cm in length. Schlettererius cmctipes was introduced into Tasmania between 1967 and 1973, together with 23 other species of parashoid Hymenoptera and one nematode species, to help control Sirex noctilio F. (Hymenoptera, Siricidae), a pest of pine plantations (Taylor 1976). It was first selected as a potential biological control agent after the collection of a male and female in the vicinity o f Pinus Jefteyi Murr. and Abies spp. trees infested by sitîcid^ m Califemia (USA). The specimens were sent to Australia, and the female exposed to a log infested with S. noctilio larvae fium which a progeny of three males and feur females o fS. cinctipes emerged (Taylor 1967a). A detailed description of the species biological tycle is also given by this author. The fact that S. cinctipes was later observed 321 to be one o f the dominant species emerging &om siricid-in&sted logs collected in Arizona and New Modco (USA) ^Cirk 1975) suggested that this species could be using siricid larvae as its primary host. More S. cmctipes specimens were then imported from the United States, and a laboratory culture was established m Tasmania and Victoria (Taylor 1976). Successive releases in areas infested by & noctilio followed in Tasmania, with regular assessment of the status of both host and parasitoid populations. According to Taylor (1981), control was no doubt effective in all study sites in Tasmania, with 60-80% of the S. noctilio population destroyed by the parasitoid complex. The overall level o f attack by & cinctipes is low (Taylor 1981; percentage of parasitism not provided), but it does not seem to be much inferior than that of several other species in the complex. It attacks the host at different times than other introduced species, thus helping to balance the action of the parasitoid complex, making control more effective. Known distribution is mapped on Fig. 245; seasonal records are tabulated m Fig. 247. Although distribution data is scanty. Fig. 245 clearly shows the species has spread to mainland Australia. There seems to be one generation a year in Tasmania, with species emergm^ searching for hosts, and ovipositing throughout, but restricted to, the period from the begmning o f December to the end of March (Taylor 1981). 322 GenusA fe ^ sc k u s BruDé For a ^onym ic list, see chapter 1. Comments. The AustnüanMegischus spp. are very characteristic, not resembling any other species in this genus. The two known Australian species are keyed below. Although based on a single specimen, M ceneoncttrix n. sp. is very distinct from the more frequent^ collectedM crassicauda ^ o d q r ). Key to the Australian Species of M egischus (females) 1 Metapleuron with short, very dense pilosity centrally and dorsally, and with long sparse hairs laterally (Fig. 233). Basal halfof hind coxa and apical half of hind femur, at least, with short dense pilosity in between sparse long hairs ^ g . 235). Colo usually with three carinae of unequal size, 2nd and 3rd carinae very strong 3rd largest, blade-like, somewhat translucent (Figs 231, 234). Pre- and semiannular finely sculptured, with dense short pilosity, except for a narrow dorso-longitudinal glabrous area (Fig. 234). Genal angle prominent, somewhat detached as a callosity (Fig. 230). Hnd femur slender ^ g . 235). K ndtarsi reddishoryellowish. Ov/PtI 1.02-1.22. Eastern Australia ...... ...... Ad. crassiccmdaQAotiey) 1 ' Metapleuron with long sparse hairs centrally only, dorsally mostly polished smooth ^ig . 240). Hind coxa and femur without short dense pilosity, polished and glabrous in between sparse long bans (Figs 242,243). Colo with five pairs of strong carmae, all carinae more or less the same size, none blade-like or 323 translucent (pgs 238,241), Pre- and semiannular strongly sculptured, short pilosity sparse, occurring laterally only (Fig. 241). Genal angle round, not prominent (Fig. 237). Hind femur stout (Fig. 242). B&id tarsi dark-brown (same coloras tibia). Ov/PtI 1.42. Southwestern Australia...... ...... M ceneonatrixAgaiar, n. sp. Megischus crassicauda (MoHqr), New Combination (Figs 230-236, 246,244) StephanuscrassicaudaMot\e^, 1917: 106;Elliott 1922:716, 728. Hemistephanus giganteus CnxdxlX, 1926: \1. New !^nonymy E tym ology, L. crassus, thick, stout; L. cauda, tail. In reference to the relatively stout ovipositor sheaths. Material Examined, Holotypes: ? Stephanus crassicaudalAotley, Australia, Jan/1917; OSUC20SS6 (BMNH). $ Hemistephanus giganteus Giiwiity no label data [collected at Tingalpa Creek, Queensland, according to Girault 1926]; OSUC20S60 (QMBA). Other specimens examined. 29 9$ and 3 F em ale L ength, Mean 25.1 (range 18.5-33.1). General structure, Frons strigate-rugose in zig-zag pattern, this sculpture from dorso-laterally oblique to ventrally transverse^ surface in between macrosculpture polished 324 smooth or weakly microreticulate (SOx). hiteroceOar carmae (Fig, 230) often broken mto three short arcuate carmae laterally and centro-IateraSy on each side, and 2-3 larger carinae centro-posteriorly; sometimes simply wfth several arcuate and concentric carinae, becoming progressively smaller posteriorly. Vertex (Kg. 230) centrally irregularly rugose or transversely rugose, abruptly changing to laterally areolate-rugose, posteriorly, including temple, densely transversely strigate. Gena prominent densely pilose-punctate with long inclined haire but polished smooth on and behmd genal prominence to occipital carina; in fi-ont o^ and dorsal to genal prominence, alongside e y t margin, with stronger punctures and short sparse hairs. Prostemum apex alutaceous, rugulose and finely punctate sub-apically, including inside and near prosternai depression, otherwise densely micropunctulate and very sparsely punctate; with dense, short pilosity. Pronotum (Figs 231,234) with dense and short pilosity and sparse long hairs, except for a narrow dorso-longitudinal glabrous area; colo usually with three pairs of carinae on each side, 1st small, 3rd the strongest, leaf-lik^ somewhat translucent, extending inside o f pronotal fold; sometimes an extra pair of weak carinae anteriorly or posteriorly; pronotal fold very stout and deep, its anterior margin undulate approximately shaped as a “}” when seen anteriorly; preannular strongly transversely strigate anterioriy, this sculpture contmuing centro-dorsally on semiannular, becoming weaker towards posterior margin; pronotum otherwise dull, very finely rugulose-punctulate; femoral impression weak, very shallow; ventral area rugulose-punctate. Mesepistemum finely but distinctly rugulose, coarser and subtransverse anteriorly, also with dense short pilosity and punctulate. Propodeum ^ g . 232): centro-Iongitudihally transversely strigate-rugose^ laterally, ihclunding fiank and 325 spiracular groove, changing progressively to finely and strongly areolate-rugose, with both sculptural patterns usuatty organùed somewhat radially near petiolar foramen; a small antero-lateral area (mesal to flank) also finely alutaceous; antero-laterally often also with a few fi)veae. Spiracular groove with same sculpture as propodeum laterally; spiracular carina distinct, in zig-zag or almost straight, fused anteriorly with flank sculpture. Metapleuron (Fig. 233): dorsally glabrous and polished smooth; centrally approxhnately areolate-rugose, with sparse long hairs and very dense short pilosity; ventrally with sparse short pilosity only. Post-foveolar area polished smooth, sometimes with a few faint transverse strigations or ventral margin with punctures; interfbveolar area strongly scrobiculate. Hind femur (Fig. 235) finely and weakly alutaceous, with sparse long hairs; laterally on apical 0.65, and on mesal side dorso-apically, covered with dense short pilosity. Bfind tibia ^ g . 235): compressed on basal 0.35, dilated part dull, strongly and uniformly microreticulate and sparsely covered with long hairs, compressed part only with sparse punctate and associated pilosity, otherwise polished smooth, Pygidium large and convex, basally with same sculpture as T9, apically polished smooth, with a fiinge of long hahs where the two sculpture patterns meet C olour. Dark-brown, with some reddish-brown on legs. Antermae daric-brown; sometimes at least the first o f the three basal flageOomeres, and scape and pedicel more rarefy, with yellowish or reddish hue. Yellowish spot near or on malar space. Reddish-brown or yellowish-brown: trochanters, firont and mid femora basally, most of fi’ont and mid tibiae, hind tibia apically, all tarsi. All tibiae basally yellowish, usually 326 followed by a brief blackish rmg on front and mid tibiae. Sometimes reddish-brown: pronotum dorso-posterior margin* petiole apeq some tergites posterior margin. Wings infiiscate. Ovipositor with a white subapical band. All hairs whitish. Morphometries, Ov/Pt 1.02-1.22, FmSI 2.79-3.70, Pnl/Ew 0.47-1.70, Cxl/h 2.17-2.79, lEE/eEE 0.53-0.57, Ov/Tt 0.96-1.34, twriEE 0.49-0.55. Seven specimens measured. M ale Length, Mean 22.1 mm (range 18.1-25.7 mm). Most important difrbrences from the females as follows. Without short dense pilosity among long hairs on hind coxa and femur; long pilosity much more abundant than in females, especially on legs; three to five carinae on colo, the anterior ones sometimes complet^ central or posterior ones sometimes wide, leaf-like, posterior ones extending mside pronotal fold; post-feveolar area entirely punctate and foveolate, rarely mostly smooth (i.e., as fer females); hind coxa and tibia much slender (CxI/h 2.70-3.82, FmSI 3.81-4.53). Comments, Easily distinguished fromM ceneonatrix by having dense short pilosity on metapleuron, hind coxa, and femur, relatively longer ovipositor (Ov/Ptl 1.02-1.22), longer hind coxa (Cxl/h 2.17-2.79), slender hind femur (FmSI 2.79-3.70), distinct pronotal structure (Figs 231,234), and lightly colored hmd tarsi. Geographic distribution also distmct (Fig. 244). hi male specimens, the lack o f dense short pilosity among the long hairs on hind coxa and femur is somewhat similar to the condition observed in the single female ofM 327 (xneoruxûix n. sp., but maies ofM crassicauda have dense short püosity dorsally on metapleuron, a much more slender hind coxa and fèmur, yellow or reddish yellow hind basharsi and hind tibia basally and apically, gena ventrally densefy punctate with dense long pilosity, and pronotum nearly identical to that of the corresponding females, all these features being characteristic of the species. Eastern Australia, except Tasmania (Fig. 244). Adults apparently more abundant in the Summer ^ig. 246). Megischus ceneonatrix Aguiar, n. sp. (Figs 237-243, 244) Etym ology, Gr. keneon, flank, side; Gr. athrix, without hair. In reference to pilosity on dorsal part o f metapleuron. Material Examined, Holotype: 9, Drummonds Cove, nr. Geraldton, WA, Australia, Dec/1972. N. McFarland; OSUC20846 (ANIC). Fem ale Length, 21.4 mm. General structure, Frons transversely porcate and subareolate, this sculpture dorsally more oblique and m zig-za^ sur&ce m between macrosculpture polished smooth. One short, arcuate interocellar carina behind each posterior ocellus, followed by series of concentric, progressively smaller undulate rugosities, which are contiguous with 328 areolate-rugose laterally; latero-posteriorly, near eye, more irregularly or longitudinally rugose. Vertex, mduding temples and post-verter, transversely substrigate, stronger centrally ^ g . 237). Gena round, polished smooth centrally and posteriorly, with strong punctures and foveae anterioriy and dorsally, along ^ e margin, latero-ventrally with long sparse hairs, distinct punctures, and short, denser pilosity and small punctures. Prostemum apex finely alutaceous, rugulose subapically, otherwise punctate with short pilosity and sparsely strong-foveolate with hairs. Pronotum (Fig. 238) with weak short pilosity and sparse long hairs, except glabrous dorso-longitudinally; colo dorsally deeply concave, with five paks of strong but regular looking carinae approximately of the same size; pronotal fold very stout and deep, its anterior margin hregular; preaimular coarsely transversely rugose, changing to transversely strigate on semiannular centro-longitudinally; preannular and semiannular laterally obliquely rugulose with punctations or foveolae; anteriorly and on ventral area with much stronger sculpture; femoral impression distinct. Mesepistemum with deep punctations and short pilosity, and sparse foveolae and areolae. Propodeum ^ g s 239,240); strongly areolate-rugose m a longitudinal pattern, converging to petiolar foramen, except for a few transverse areolae centro-anteriorl)^ flank with same overall sculpture. Spiracular groove weakly concave, polished smooth, ending at flank; spiracular carina distinct, straight on its basal 0.7, ending at flank. Metapleuron (Hg. 240); dorsally and ventrally glabrous and polished smooth; centrally areolate-rugose and with very long pilosity; dorso-laterally with a few weak short hairs and some 6int microreticulate. Post-fi)veolar area polished smooth, with a line of punctures along its ventral margin; 329 interfôveolar area strongly scrobiculate. Hind femur ^igs 242,243) weakly transversely alutaceous, with sparse long hairs. Hind tibia ^ g . 242) compressed on basal 0.3; otherwise as inM crassicauda. Pygidium as inM crassicaueh. Colour. Dark-brown. Yellowish spot right above malar space. Daric-brown with reddish hue: pronotum dorso-posterior margin, front and mid tarsi, post-abdomen. Reddish: hind 2nd trochanter, petiole apex; much lighter on pygidium and apical hind tarsomere. Wngs in&scate. With a white subapical band on ovipositor valves. All hairs whitish or yellowish. Morphometries. Ov/Pt 1.42, FmSI 2.58, Pnl/Ew 1.78, Cxl/h 2.03, lEE/eEE 0.60, Ov/Tt 0.92, tw/iEE 0.48 (Holotype). M ale Unknown. Comments. Easily distinguished fromM crassicawki by the absence of dense short pilosity on metapleuron, hind coxa, and femur, relatively short ovipositor and hind coxae, more stout hind femur (see FmSI); distinct pronotal structure (Fig. 238,241), and concolorous hind tibia and tarsi. Geographic distribution also distinct (Fig. 244). 330 GenusParastepkanèUus Enderiein For a synonymic list, see chapter L Com m ents, An efiScient way to recognize Parastepfumelbis in Australia is to search for front wing vem 2+3M tubular on its basal half only, as opposed to entirely tubular or entirely spectral in other genera ofStephanidae. The features used by Enderiein (1906) to characterize f arasApAaneZ&s, and later partially adopted by Elliott (1922) in his monograph of the frmily, are usually too variable to be diagnostic for the genus. For example, the large size of cell 1+2R1 in relation to cell IM, the most commonly cited diagnostic character for recognising Parastephanellus^ varies considerably within this taxon, being sometunes very similar to the condition observed in Hemistephamts. As a result, some Australian specieso fParastephanellus have been described (Elliott 1919a) or identified ^ e f f e r 1908, Moriey 1917, Elliott 1922, Moore 1962) as H em isteplum us (see also item “stephanid species mistakenly reported in Australia”). The genus Parastephanellus occurs all over Australia, except apparently in Tasmania (Fig. 411). There is a notably greater number o f Parastephanellus species in the eastern Australian States (16) than in the central ones (3) or in Western Australia (S). The only widespread species is P, rufoorruOus (Cameron), albeit it is distinctly more abundant in the eastern side. Parastephanellus tanytarsus n. sp. and P. granulatus (Elliott) occur only in Western Australia (Figs 420-421), and P. squam atus n. sp. occur in both eastern and central states (Fig. 426). The other 13 species seem to occur ecclusively along the eastern coast, with 11 of them apparently restricted to eastern Queensland. 331 Although avaflable data is still notably insufficient for some species ^igs 416-418, 425,429), it is nonetheless apparent that within the western or the eastern groups there is little or no geographic isolation among dK&rent species, at least in terms of the geographic scale considered. Also somewhat unusual are the western records for P. trtfasciatus n. sp, (Fig, 424) and P, rufoom atus ^ g , 428), Although based on apparently reliable label data, these occurrences were inferred from a single specimen for each species, and must therefore be considered with caution. The most commonly collected species in Australia are P. rufoom atus and P. damellicus, with all other species apparently much less abundant (Table 3.3). Adults are generally more numerous during the Summer, especially December and January (Fig. 430). species n % species n % species n % P, rufoomatus 131 46.6 *P. squam atus 8 2.8 *P. vertipictus 3 1.0 P. dameilicus 51 18.1 P. laevigatus 7 2.5 *P. brevis 2 0.7 P. trifasciatus 16 5.7 *P. granulatus 7 2.5 •P. adiazetos 2 0.7 P, isoglyptos 14 5.0 P. denotatus 7 2J *P. albigenuis 1 0.4 P. strigosus 13 4.5 P. m aculatus 4 1.4 ♦P. callidictyon 1 0.4 P. piatyceratus 9 3.2 P. tatnytarsus 4 1.4 *P. cauebtus 1 0.4 Table 3.3. Number of specimens studied (n) for each species of Australian ParastephanelluSy correspondent percentage (%) m relation to total number of spechnens, and indication o f species fr>r which males are unknown (*). 332 Groundplan Most AustralianParastephanelbis can be treated as a group with a characteristic morphology. The only exceptions are P. c a u d a tu s Elliott, described from, and apparently more closely related to New Guinean species, and P. maculatus n. sp., which is more similar to forms occurring in Borneo and New Guinea. Two other species, /*. dam ellicus (Westw.) and P. denotatus n. sp., occur also in New Guinea, but morphologically are distinctly more closely related to the typical Australian forms. The characteristic morphology of most Australian Parastephanellus seem significant, as results from a phylogenetic analysis of the Stephanidae (Aguiar, in preparation) suggest they may form a monophyletic group. Although a comprehensive analysis would be necessary to confirm this, the possibility of monophyly is suggested by the fiict that P. rufoomeOus, P. granulatus, and P. denotatus, representing most of the morphological variability of the typical Australian forms, were invariably recovered as a clade in the mentioned analysis, which also included Parastephanellus species from New Gumea, Salomon Islands, and New Caledonia. The groudplan morpholo^ for these forms is described below. It is important that this general structure be well understood before any taxonomic work with this group is attempted, as it «(plains multiple morphological variations which are of generalized occurrence, and therefore have little taxonomic importance at the species level. Note, however, that it is neither a general description for the genus, nor a list of plesiomorpbies. H ead. Frons subareolate-rugos^ frontal carina developed, formed by coalescence o f neighboring wrinkles, sculpture more regulariy transverse near ^ e . Coronal area 333 transversely or subradîally strigate around ocellus, more irregular anterioriy behind anterior tubercle. A few distinct, straight interocellar carinae. Vertex with weak and transversely strigate centro-Iongitudinal hnpression, sculpture behind interocellar carinae somewhat irregular rugose or concentrically undulate-strigate, then transverse or concentric, becoming finer posteriorly. Temple polished smooth or with same general sculpture o f vertex. Genal angle prominent, gena polished smooth. Occipital carina narrow, but distinct, widest latero-dorsally. Occiput polished smooth. Hypostomal carina polished smooth, alutaceous, or rugulose. Post-genal bridge alutaceous. Thorax. Prostemum apicaUy alutaceous, subapically on lateral margin of each hemistemite obliquely rugulose, centrally polished smooth, prosternai depression distinct Colo anterior margin slightly upturned and deeply emarginate, anteriorly with a few oblique lateral carinae, latero-posteriorly with oblique carinae, which enter pronotal fold dorsally, centro-posteriorly somewhat concave, including within pronotal fold, which is distinct, somewhat flattened, distinctly but not deeply concave anterioriy, centro-dorsally collapsed. Preannular transversely strigate or transversely rugose, anterioriy finely transversely alutaceous. Semiannular dorso-lateralfy with stronger or more irregular sculpture, often also with shallow foveolae, dorso-posterior margin polished smooth or weakly alutaceous. Pronotal lobe and depression in fi*ont of it longitudinally carinate or strigate. Femoral impression distinct, polished smooth or with same general sculpture as pronotum; ventral area sculpture coarser anteriorly, finer posteriorly. Mesoscutum anteriorly finely and distinctly transverse^ alutaceous, this part variably exposed, sometimes covered by posterior margin o f semiannular; median sulcus and notauli 334 extending through entire length of mesoscutum, and formed by deep aligned foveolae or lacuna^ partially fused with areolate-rugose or polished rugose sculpture of median and lateral lobes. Axilla sculpture in somewhat concentric pattern; with a larg^ deep, polished smooth fovea where scutellum and axillae meet. Scutellum with a narrow elongate sulcus along postero-lateral margin, ending midway towards scutellum apex; deeply crenulate between scutellum and axillae. Mesepistemum anteriorly usually coarsely rugose or rugulose and densely pilose, sculpture finer ventrally or posteriory; its dorsal part mostly polished smooth, glabrous. Mesopseudostemum finely transversely alutaceous, accept narrowly polished smooth near foveolate discrimen. Propodeum sculpture often combining three or four different patterns, usually coarser centro-longitudinally, well delimited polished smooth fbveae never present; flanks with finer and fewer sculpture patterns than main area of propodeum, or often only finely alutaceous. Parapetiolar fovea not distinctly delhnited, partially polished smooth or carinate, partially merged with general propodeal sculpture. Metapleuron centrally subtriangularly prominent, areolate-rugose, ventrally and post-foveolar area transversely carinate, polished smooth between carinae, which are contiguous with central areolate pattern, ventro-anterior margin finely alutaceous. Literfoveolar area with a few weak transverse carina^ polished smooth between carmae. Metapleural fevea distinct, subcircular. Hind coxa mostly transversely strigate^ but laterally rugose; or rugulose basally; a somewhat fiat or weakly concave centro-dorsal area with much finer sculpture; mesally also partially punctulate or microreticulate. Hind femur feiely and longitudinally alutaceous, this pattern more elongate laterally or ventrally, denser and more regular dorsally, from weak to polished smooth basally, sculpture finer 335 i^ically; basal tooth subtriangular; median tooth longer; 2-7 small tubercles between base o f &mur and basal tooth, 3-7 between the two teeth, 4-5 apically, which are the strongest; each tubercle bearmg at least one subaptcal seta or hair; a broad semicircular notch ^ically, b^ond tubercles. Hind tibia: compressed part very finely longitudinally or obliquely alutaceous, ventrally sharp. Dilated part ventrally obliquely carinate on each side, forming an inverted V-pattem. F ore w ing. Setae on M*H3u more or less equally spaced, those on basal 0.6 (often broken ofO longer and more delicate than apical ones; setae on ScfR. and 1-1A stouter and longer than basal setae of M-fCu, absent on apical 0.3-0.1 of Sc+R. and apical 0.6-0.4 of 1-1 A. Veins IM and ICu basally isolated or fused for a short distance (leaving cell IM basally “appendiculate”). Crossvein Icu-a weakly sinuous. IRs oblique, its angle with IM greater than angle with (Rs+h^a. Vein 2-1A entirely spectral, or a very short, tubular stub, usually becoming shortly nebulous apically. Vein 2Cua usually tubular on basal 0.2, otherwise nebulous, apically dilated and bent towards wing apex, continued by 2Cub, which is spectral or weakly nebulous, ending near wing margin. Vein (Rs+h^b distinct, 6-IOx as long as wid^ contiguous with 2+3Rs, which is slightly arched. Vein 2+3M originating at, but not fiised to, junction of (RsfM)b with 2+3Rs, tubular on basal 0.3-0.6, otherwise spectral or weakly nebulous. Crossvein 2r slightly sinuous, with 0.2-0.3 length of ^Is, and 0.7-0.8 length o f 2+3Rs. Vem Cup distmct, crossmg the entire wmg length, but more evident between apical part of 1-1A and 2Cua. Pterostigma sublanceolat^ pale basalfy, otherwise unftbrmly sclerotized. Cell I+2R1 usually distmctly larger than cell 336 although sometimes almost as large (areas R/M 1.24-2,55, n = 22, six species evaluated). Cell IM diamond shaped, veins Arming its opposite sides [IM/m-cu and (Rs+h^a/ICu] practically parallel to each other and approximately of the same length. H in d wing. Venation very reduced, no closed cells. Sc+R. distinct, tubular, other vans absent, distmctly curved IRsb spectral, sometimes nebulous. Mostly 3-5 hamuli. Abdomen. Petiole line of fusion between terghe and stemite weak but visibly terghe semicircular in cross-section, from finely rugulose to transversely strigate, basally with sculpture coarser or irregular, apex polished smooth; stemite somewhat V-shaped in cross-section, with a centro-Iongitudinal undulate carina extendmg from base to apex, otherwise sparsely coarsely rugulose, sometimes also punctulae or microreticulation basally; petiolar spiracles nonfunctional, inconspicuous, indicated by small lateral tubercle on each side, located slightly basad of middle. Tergite 3 rugulose basally, otherwise polished smooth or apically faintly and transversely microaciculate, T4-8 transversely microaciculate, T8 apically sometimes transversely alutaceous, T9 distinctly alutaceous. Pygidiai impression triangular, V-shaped, pygidium narrow, convex, emargioate. Colour. Mostly dark brown, with legs partially brown, white on base and apex of fit)nt and mid tibia, and base o f mid basitarsomere. Genal stripe mctendmg adjacent to ^ e margm ventrally, separated from it dorsally. Wing veins brown or dark brown. Ovipositor entirely dark brown, except very briefly white or reddish brown on tip, pre-apical white band absent. 337 Key to Species o f Australian ParastqphaneUus Enderlein The Mowing was built mainly firom the analysis of female spechnens» but will also woric for males o f most species (except morphometric ratios, calculated for fomales only). Males of some species, however, are considerably difficult to recognize, and were treated separately into the kqr. The sculpture of the scutellum is of primary importance in recognising males of P. damellicus and P. isogfyptos, but not a decisive character per se for P. laevigatus (couplets 23-24). Ditraspecffic variation for males o ff. dam ellicus is very high, and some specimens cannot be recognized by the key alone. Species for which males are unknown are indicated in Table 3.3. Notes. (1) Specimens which have been in alcohol for some years may show severely faded or modified colours (e g., fi’om red or dark brown to yellow, pale yellow, or white); this was observed in pin-mounted specimens off. denotatus n. sp., and f . nrfoomatus; (2) values between square brackets are expanded limits, within which overiapping with those of closely related species would still not occur; (3) note that number 19 in the kqr is a triplet. 1 Body mostly shining (Figs 272-397), strong sculpture at least on pronotum, metapleuron, propodeum. Head neverferrugmeous, usually dark brown, genal stripe well defined, ivory or yellow. Hind femur variable, but usually not slender (FmSI usually <2.30)...... 2 r Body dull ^igs 398-401), frons, thorax, including hind legs, and propodeum with very fine microreticulation. Head usually light forrugineous, genal stripe 338 dîfiiis^ creamy yellowish, on light-brown gena. Hind femur slender (FmSI 2.26-2.63; Rg.402) ...... P. denotatusn, sp. 2(1) Pygidium simple ^igs 279, 282, 321,403). % id coxa always shorter than femur (Cxl/Fml < 1.00). Semiannular with strigations or coarser sculpture (e.g., Rgs 308,334,363), sometime with polished smooth spaces, but never entirely alutaceous and shinning (e.g.. Figs 275,356). Ov/Ptl usually <5.00. Ov/Hdl <12.00 (accept P. strigosus). Interocular space normal or wide QEE/eEE usually larger than 0.45)...... 3 2" Pygidium ‘bifid’ (Rg. 410). Hind coxa longer than correspondent femur (Cxl/Fml 1.03-1.12). Semiannular laterally weakly alutaceous, shining (Rg. 407). Ovipositor relatively long (Ov/Ptl 4.40-6.09, Ov/Hdl 12.58-19.32). Interocular space very narrow ^E/eEE 0.40-0.41) P. caudatusEMaotL 3(2) Vertoc with three colours: broadly black centrally, reddish anteriorly, narrowly along eye margin, more widely on temple, and broadly yellow on lateral part of temple (fiom genal stripe). Scutellum sculpture as in Rg. 363. Mesepistemum microreticulate (Rgs 394, 397). Front and mid legs b^ond coxa, and hind tibia and tarsus, yellowish brown. Petiole distinctly more lightly coloured than thorax and abdomen, reddishbrown ...... P. vertipictusvusç^ 3’ Vertex mostly unicolourous, except by genal stripe, and sometimes base of interocellar carmae reddish. Scutellum sculpture not as m Rg. 363. Mesepistemum sculpture variable...... 4 339 4(3) Hther tonple relatively broad, gena round (twÆEE 0.67-0.80; Figs 1 [chapter I], 382) and ovipositor short (Ov/Hdl 5.82-8.46), or head and thorax strongly - flattened (tw^EE 0.18-0.35; Figs 374,381). Mesepistemum microreticulate, at least centrally (as in Figs 384, 386). Pygidiai impression usually deeply notched basally. Ov/Saw <10.31. Length normally <10 mm ...... 5 4’ Temple usually normal, genal angle sometimes conspicuously prominent laterally (as in Figs 359, 366, 391), head and thorax never flattened (tw/iEE always greatw than 0.45 and usually smaller than 0.67, e.g., Rgs 289-290,304,348, 366), and Ov/Hdl usually greater than 8.46. Mesepistemum and pygidiai impression variable, but if mesepistemum microreticulate and pygidiai impression notched, then frons has yellow stripes along ^ e margin (females; Fig. 368), or flice is entirely yellow (males). Ov/Saw usually >10.31. Length commonly >10 mm...... 6 5(4) Temple broad (twfiEE 0.67-0.80; Rg. 382), body normal (Fig. 1 [chapter 1]). Broadly yellow or brownish yellow around each antennal forame. Pronotum short ^nl/Saw 0.60-0.80, Pnl/Ew 1.04-1.33; Fig. 383). Vertex without scale like projections laterally. Posterior margin of head concave in dorsal view (Rg.2696). Pronotalfold stout, distinct (Rg. 383) ...... ______P.m/bomornf (Cameron) 5’ Head and thorax strongly flattened (tw/iEE 0.18-0.35; Figs 374-375, 381). Frons entirely black or weakly reddish along ^ e margin or near antennal forame, never with yellow. Pronotum longer (Pnl/Saw 0.85-1.00; Pnl/Ew 1.46-2.07; 340 Figs 375,377). Vertex with scale-Iike projections dorso-Iaterally on flattened part (Fig. 376). Posterior margin of head straight in dorsal view (Fig. 374). Pronotal fold flat and shallow ^ g s 377,379), or absent (Fig. 375) ...... ...... P.squanatusvLsg. 6(4) Males. Frons entirely lightly coloured, yellowish or light-brown ...... 7 6’ Males and fomales. Frons variable, but never entirely lightly coloured ...... 9 7(6) Frons entirely bright sulflir yellow or pale yellow. Temple with strigation from vertex (as in Figs 362, 366). Metapleuron dorsally transversely strigate (as in Figs 361,373), albeit sometimes weakly...... 8 T Frons brown or yellowish brown. Temple polished smooth (as in Fig. 355). Metapleuron dorsally variable, but ofren polished smooth (as in Fig. 358) ...... ...... P.platyceratusn. sp. (seealso couplet 11) 8(7) Semiannular laterally distinctly strigate ^ g . 369). Femoral impression longitudinally strigate (Rg. 369). Scutellum distinctly alutaceous (Rgs 370-371). Body colour usually light, yellowish brown or shnilar ...... P. trifasciatusn, sp. 8* Semiannular laterally finely alutaceous (as in Rg. 363), but distinctly more coarsely sculptured latero-dorsally. Femoral impression ofren transversely carinate ^ g . 363). Scutellum coarsely sculptured, strongly alutaceous to strongly substrigate or areolate-rugose (Fig. 364). Body colour only partially lightly coloured, meso- and metathorax, incluging propodeum, and petiole usually darkbrown...... P. strigosusxL sp. 341 9(Q Neaiiy always with vertical, yellowish stripes on frons (e g., Rg. 368), at least along ^ e margin, although sometimes frint, difficult to see. Genal angle usually promment, somewhat acuminate (Figs 355,359,366). Ovipositor long (Ov/Tt 1.13-1.69). Hind coxaslender(Cxl/h>3.30) ...... 10 9* Without longhudmal yellow stripes on frons (e g.. Fig. 339), although sometimes yellowish around antennal foramen. Genal angle usually less prominent (e g.. Figs 299, 348). Ovipositor shorter (Ov/Tt often below 1.15). Hind coxa normal (CxI/h usually < 3.30)...... 12 10(9) Frons with two yellow stripes along e y t margin, and absolutely no central stripe. Ptl/T3I 2.50-3.09. Basal 0.2 of hind tibia whitish (except a few atypiod specimens) ...... 11 10’ Frons with three vertical stripes, two along eye margin, one centrally, stripes always present, but sometimes faint, inconspicuous, especially central one OFig. 368). Ptl/T312.09-2.46. Compressed part of hind tibia entirely yellowish brown ..... ...... P. trifasciatusiL sp. 11(10) Temple with same general strigation o f vertex, which reaches broadly occipital carina and genal stripe (Rg. 362). Femoral impression distinctly transversely carinate ^ g . 363). Semiannular laterally distinctly alutaceous (Fig. 363), dorsally with much coarser sculpture (Fig. 360). Scutellum coarsely sculptured, strongly alutaceous to strongly substrigateorareolate-rugose(Fig. 364). Flagellomeres on apical halfof antenna ^lindric ...... P.strigosusTL sp. 342 i r Temple broadly polished smooth, vertex sculpture not or only barely reaching occipital carina, ending 6 r from genal stripe (Fig. 355). Femoral impression smooth (Fig. 356). Semiannular, mcluding dorsally, mostly polished smooth (Fig. 356). Scutellum centrally weakly alutaceous ^ g . 357) or polished smooth. Flagellomeres on apical half of antenna somewhat dattenned (females)...... P. platyceratusn, sp. 12(9) Semiannular dorsally strong!^ rugose OFigs 323,340), or at least coarsely transversely carinate ^igs 332, 349). Prostemum strongly rugulose or rugose (Figs 333, 345,352), or at least strongly alutaceous, or both. Frontal carina absent (e.g., Figs330,339), except males sometimes ...... 13 12’ Semiannular usually ordinarily strigate (as in Fig. 308); if transversely strongly carinate, or rugose (one specimen off. damellicits\ then prostemum weakly alutaceous orpolished smooth. Frontal carina present or absent ...... 16 13(12) Scutellum areolate-rugose and punctate (Figs 344, 354). Prostemum strongly rugulose (Figs 345,352). Basal 0.4 or more o f mid basitarsomere distinctly more lightly coloured fellow or white) than apical part. Ov/Ptl < 4.25 [3.14- 4.01].WestemAustralia^gs420-421)...... 14 13’ Scutellum centrally mostly polished smooth (Fig. 331), or densely and finely transversely aIutaceous^g.326).Prostermim strongly alutaceous orcoarsely transversely strigate (Rg. 333). Mid basitarsomere lightly coloured on basal 0.2 or less, or unicolourous. Ov/Ptl > 4.25 [4.56-5.10]. Eastern Australia (Figs4I7-4I8)...... 15 343 14(13) Hmdbasharsusshort(btIAv4.82-5.05),yelIow.PetioIedistmctlytransversefystrigate. Ventral area of pronotum strongly longitudinally carinate ^ig. 351), Semiannular coarsely transversely strigate or transversely carinate, usually also rugulosedorso-Iaterally (Figs 349,351) ...... P .ff’camlatusWioVL 14’ Hind basitarsus long (btl/w 6,41-6.44), reddish brown. Petiole finely and densely rugulose ('granular’ texture) orfinely areolate-rugulose, coarser basally, finer apically. Ventral area of pronotum rugose or sub-rugose, never with long, unbroken carinae (except males) (Fig, 342), Semiannular rugose, with few transverse carinae (Figs340,342) ...... P. tanytarsusvL sp, 15(13) Scutellum centrally mostly polished smooth, laterally foveolate, with a few foveolae reaching central part ^ g . 331), Prostemum very coarsely transversely strigate (Rg, 333), Mid basitarsomere uniformly unicolorous. Hind basitarsomere short (btl/w 4,78), Semiannular coarsely transversely strigate dorso-anteriorly, becoming longitudinally strongly strigate laterally, including femoral impression and ventral area (Figs 332, 334, 337), Genal stripe complete. Metaplmiron dorsally mostly polished smooth (Fig, 336), Frons homogeneously areolate-rugose^g, 330), Large(15 mm) ...... ...... f , callidictyonvL sp, 15’ Scutellum centrally dull, densefy and strongly alutaceous ^ g s 324,326). Prostemum strongly alutaceous, NCd basitarsomere on basal 0.2 whitish. Hind basitarsomere long (btl/w 8,80), Semiannular entirely rugose, including femoral hnpression and ventral area ^ g s 323, 325), Genal stripe broadly 344 interrupted centrally, leaving one spot near mandible and another on temple (temporal spot absent on single male specimen observed). Metapleuron dorsally rugulose and transversely carmate ^ g . 327). Frons near ^ e s and ventrally with strong and sparse transverse strigation. Small (8 nun) ...... ...... P. aeUazetostL sp. 16(12) Pronotum general^ more elongate (Pnl/Saw 0.79-1.10; Pnl/Ew 1.37-1.79) (Figs 291, 303, 305). Ovipositor moderatly longer than petiole (Ov/Pt <4.1) and hind Amur slender (FmSI almost always>1.81). Temple width normal (tw/io 0.84- 1.21; tw/iEE 0.53-0.67). Vein 2Cua tubular on basal 0.3 or less (as in Fig. 350)...... 17 16’ Pronotum generally short (Pnl/Saw 0.57-0.83; Pnl/Ew 0.99-1.39) ^ g s 273, 281). Ovipositor much longer than petiole (Ov/Pt 4.19-5.90); hind femur more globular ^m SI usually <1.6; total range 1.42-1.81). Temple slightly wider than average (tw/io mostly 1.26-1.33, rarely more; twAEE 0.67-0.76). Vem 2Cua variable, but sometimes entirely tu b u la r^ g . 287)...... 22 17(16) T3 polished smooth or almost so, T4-7 transversely microaciculate (dull in relation to T3; clean specimens). Temple and prostemum variable. Scutellum centrally mostly polished smooth (Figs 292-293), or heavily sculptured, with 2-4 d ire n t patterns on same specimen ^ g s 301,309-310). Hind tibia basally concolourous with remaining of compressed part, never whitish. Mid tibia basally and apically rarefy whitish. Gsl/Ptl usually smaller than 1.40. Ov/Saw >9.50 ...... 18 345 17’ T3-6 polished smooth, T7 ftintly alutaceous (Fig. 321). Temple ^ g s 315,318) and prostemum polished smooth. Scutellum centrally strongly transversely alutaceous (Figs 319-320). Hind femur strongly alutaceous on both sides (as strong as on dilated part of correspondent tibia). Hind tibia on basal 0.3, and mid tibia basally and apically, whitish. Gsl/PtI 1.43. Ov/Saw 8.96 ...... ...... f . albigenuisTL sp. 18(17) Females; hind tarsus3-segmented ...... 19 18’ Males; hmd tarsus 5-segmented ...... 23 19(18) Vertex sculpture, including temples until genal stripe, uniformly transversely strigate (Fig. 299). Scutellum with fine transverse sculpture (coarse alutaceous, or substrigulate, or strigate) ^ ig . 301). Pronotum and mesothorax dark brown or black. Metapleuron dorsally always finely transversely strigate (Fig. 302) ortransversely alutaceous...... P. isogfyptosxL sp. 19’ Vertex transversely strigate, but this pattern arcuate centrally, or wavy at least laterally near posterior ocellus, temple sculpture variable (Figs 289-290,304). Scutellum varying fi’om coarsely areolate-rugose and foveolate (Figs 307, 309-310), to mostly polished smooth or very weakly alutaceous, with a few foveolae centrally (Figs 292-293). Pronotum sometimes dark brown, but usually with reddish hu^ or yellowish, contrasting with distmctly darker mesothorax. Metapleuron dorsally somethnes polished smooth (Figs 296, 313)...... 21 346 19’ Litennediate, or mostly as for P, isogfyplos but with some characters from second couplet...... 20 20(19) Prostdmum weakly or very weakly alutaceous (40x magnificatioii; sometimes apparently polished smooth). Compressed part of hind tibia distinct^ lightly coloured, reddish or yellowish, Interocellar space narrow OEE^o 1.90-1.94, tw^o 1.14-1.19) ...... P. isogtyptosxL sp. 20’ Prostemum polished smooth; sometimes with short strigations or microreticulation apico-laterally or basally, but never alutaceous. ICnd tibia, except sometimes apm^ usually dark brown or black, compressed part rarely lightly coloured. hiteroceOar space wide (iEE/Io 1.49-1.84, tw^o mostly between 0.84-1.10) (see also couplet21)...... P. dbmg/Bcits (Westwood) 21(19) Overall sculpture coarse: Scutellum usually coarsely areolate-rugose and foveolate (Figs 307, 309-311), somethnes also arcuate-strigate, rarefy only with a few foveolae centrally. Temple with same scoltpture as vertex centro-posteriorly ^ g . 304), rarely polished smooth; ^so, vertex sculpture endmg not very for from genal stripe. Pronotum usually shorter than in P. laevigatus (Pnl/Saw mostly < 1.02) (see also comments for morphofype H of P. laevigatus) ...... ...... P. dbme/7/cus(Westwood) 21’ Overall sculpture weak: Scutellum centrally mostly polished smooth, with sparse foveolae ^ g s 292-293). Temple broadly polished smooth, vertmc sculpture endmgfor away from genal stripe (Fig 289). Pronotum usually more elongate than mAdüme/KcMS (Pnl/Saw mostly > 1.02) ...... P . laevigatustLsp. 347 22(IQ Vem 2Cua entirely tubular ^ g . 287). Ovipositor much longer than petiole (Ov/PtI 5.52-5.90). Pronotum very short (Pnl/Saw 0.57-0.61) (Fig. 281). Front wing membrane amber-infiiscate. Pterostigma somewhat acute, elongate. Semiannular laterally stro n g strigate (Fig. 284) ...... P. brevisn, sp. 22’ Vein 2Cua tubular on basal 0.25 only (similar to Rg. 365). Ovipositor moderately longer than petiole (Ov/PtI 3.89-4.20). Pronotum elongated (Pnl/Saw 0.70- 0.83). Cell 2Cu strongly mliiscate, wing membrane otherwise hyaline. Pterostigmablunt. Semiarmularlatero-centrally polished smooth (Fig. 275).... ...... P. macuhtusTL sp. 23(18) ScuteOum conspicuously alutaceous ormicroreticuiate, apical 0.3 sometimes polished smooth. Vertoc, including temple until genal stripe, uniformly transversely strigate (similar to Fig. 299). Mesepistemum sculpture coarse. Semiannular distmctly, sometimes strongly, transversely strigate. Pronotum and mesothorax daric brown or black (See also comments for males of P. ntfoomatus)...... P. iso g ^ to siL sp. 23 ’ Scutellum with foveolae or mostly polished smooth. Vertex transversely strigate, but strigation is arcuate centrally or undulate laterally (shnilar to Figs 289-290, 304). Temple often polished smooth. Mesepistemum and semiannular variable. Pronotum very rarely dark brown, normally reddish black or yellowish, sharply contrasting with much darker colour o f mesothorax 24 24(23) Overall sculpture strong: Scutellum sculpture usually coarse, due to large foveolae and punctures, or at least with some foveolae hi partially polished smooth 348 background. Semiannular distinctly, sometimes strongly, strigate. Mesepistemum sculpture coarse somethnes partially strong alutaceous (see also comments for males o fP. rufoomatus)...... P, dbme/Kicitr (Westwood) 24* Overall sculpture weak: Scutellum mostly polished smooth or weakly alutaceous, usually with a few small, shallow foveolae laterally. Semiannular weakly or very weakly strigate partially polished smooth. Mesepistemum sculpture relatively sparse with polished smoothspaces between mgae/areolae ...... ...... P. laevigatusTLsp, Descriptions Note: values for morphometric ratios are presented comparatively for all species on Figs 252-271. Minimum and maximum limits listed on tables 3.6 to 3.10 (Appendix). Parastqrhanellus maculatus Aguiar n. sp. (Figs 252-271 (sp. 1), 272-279,413) E tym ology. L. macula, spot, stain, mark, hi reference to infuscation of cell 2Cu. M aterial Examined. Holotype: ?, Coastal Range, near Biggenden, Qld., 28 Nov 1975, H. Frauca. OSUC21041 (ANIC). Paratypes: 2 ? ?, 1 (f (QMBA, UQIC). F em ale L ength. Mean 6.7 mm (range 3.8-8.1 mm). 349 Hecui. Flagellomeres normal. Frons strongly areolate-rugose; frontal carina distinct. Vertex ^ g . 272) mostly transversely carinate or transversely strigate; posterior 0.20-0.25 transversely strigulate; more frregulariiy behind interocellar carina; temple polished smooth. Centro-longitudinal hnpression very weak or absent. Genal angle moderately prominent, with no distinct protuberance. Gena polished smooth. Occipital carina very narrow. Posterior margin of head concave m dorsal view. Occiput polished smooth. Post-genal bridge weakly alutaceous. Hypostomal carina apex, below mandible base, finely but weakly alutaceous. Thorax^ Prostemum with a few oblique luguIosMes anteriorly on prostemal depression, otherwise polished smooth. Pronotum (Figs 273,275); colo carinate, carinae incomplete, strongly oblique or almost longitudinal, running towards, posterior ones entering, pronotal fold; polished smooth centro-dorsally and between carinae; pronotal fold very stout, its dorsal margin deeply and broadly emargmated; preannular very short, almost absent centrally; semiarmular in dorsal view polished smooth or weakly alutaceous along posterior margm, especially centrally, then transversely carinate-rugose; apically dense-alutaceous on each side o f preannular; in lateral view broadly polished smooth centrally, dorsally with 2-3 small punctures, transversely carinate-rugose anteriorly, longitudinally dorsally and posteriorly ^eluding pronotal lobe); femoral impression distinct, or almost so, narrow, polished smooth; ventral area from transversely carinate apically to longtudmally alutaceous posteriorly. Mesoscutum areolate or areolate-rugose; lateral lobe weakly microareolate anteriorly. Axilla (Rgs 274,276) areolate-rugose in semicircular pattern; strigulate or alutaceous transversely on anterior margin. Scutellum 350 (Figs 274,276) laterally rug^Iose-Iacunose or areolate, or with some isolate lacunae; centrally weakly alutaceous or polished smooth. Mesepistemum (Fig. 277, part) anteriorly strongly rugose and densely pilose then punctate or foveolate on shallowly rugose- lacunose, punctulate on weakly alutaceous background posteriorly; dorsal part polished smooth; changing gradually to alutaceous on ventral part; discrimen simple or foveolate. Propodeum (Figs 276-277) laterally foveolate and distinctly alutaceous; centrally from foveate anteriorly to strongly areolate-rugose near petiolar foramen; flank alutaceous, finer anteriorly, coarser laterally. Parapetiolar fovea polished smooth or strongly transversely carinate. Spiracular groove defined by irregular and closely aligned lacunae; spiracular carina well mariced, endmg at some distance from flank. Metapleuron (Fig. 277) dorsally mostly polished smooth, short transverse carinae along spiracular carina, and transversely carinate dorso-posteriorly; centrally areolate-rugose; ventrally weakly or distinctly microreticulate; anteriorly and dorsally to metapleural fovea polished smooth. Post- foveolar area transversely carinate; transversely strigulate in small specimens. Interfoveolar area scrobiculate, deeper anteriorly. Ventral side o f fourth tarsomere of front and mid legs normal, not prolonged anteriorly. Hind coxa centrally and posteriorly on dorsal 0.6 of lateral side somewhat depressed and microreticulate, alutaceous, and some rugulose, otherwise transversely strigate, except microreticulate-punctate basally mesally. (find femur entirely and conspicuously alutaceous ^ g . 278). Hind tibia longhudmaOy alutaceous, finer than on hind fomur (Fig. 278); dilated part obliquely carinate ventrally. Front wmg: vein 2Cua tubular on basal 0.2-0.5. Pterostigma blunt, short. 351 Abdom en. (Fig, 279) Petiole transversely strigulate or finely transversely rugulose» coarser basally, apex polished smooth; ventrally transversely rugose or finely transversely rugulose. Tergite 3 on extreme base with about six short longitudinal carinae, otherwise polished smooth, T4-6 polished smooth, T7-9 subtransversely alutaceous, fi'om weakly on XT to strongly on T9 (Fig. 279). Pygidiai impression wide subtriangular or broadly semicircular. Pygidium narrow, deeply emarginate, not elevated (Fig. 279). C olour. Black, brownish legs. Genal stripe uniformly narrow, creamy-yellow, ending near mid of temple, for firom occipital carina. Ferrugineous base of coronal tubercles and interocellar carinae, apex o f petiole, extreme base of T3; usually also fi’om and mid coxae, trochanters, and femora (which sometimes are blackened towards apex), bind leg on coxa dorsally and ventro-apically, trochanter, femur «ctreme base, and tibia laterally; sometimes slightly ferrugineous on whole head and prostemum, mid and hind legs b^ond coxa. Yellowish brown or brownish antennae basally, mandible except apex, and all tibiae and tarsi except: black apex for each apical tarsomere, whitish apical 0.2 of mid tibia and basal O.S-0.6 o f mid basitarsus, and centro-Iongitudinal dark stripe mesally on hind tibia. Wing membrane amber, little more intense subapically on cells l+2+3Rs and 2 + 3 much stronger on cell 2Cu; veins brown, pterostigma and ovipositor valves dark brown. D iagnosis. Pronotum short ^ ig . 273). Semiannular broadly polished smooth laterally ^ g . 275). Scutellum centrally and apically polished smooth ^igs 274, 276). Mesepistemum punctate or foveolate on shallowly rugose^lacunose ^ g . 277). Dorsal area o f metapleuron mostly polished smooth (Fig. 277). Pterostigma blunt. Cells 2Cu and 352 1+2+3RS distinctly infiiscate. Basal 0.1 o f front and bind wings, including veins, creamy white. Front and lateral tubercles ferrugineous to reddish brown, with dark brown tips, sharply contrasting with blackish frons. Rem arks. Similar to P . brevis^ but immediately distinguished from it by havmg vein 2Cua tubular basally only and cell 2Cu infiiscate; also distinguished by diagnosis above, vertex sculpture, and colour pattern ofI%s. Similar to P. rufoomatus, from which it can be distinguished, in addition to the diagnosis provided above, by having transverse carinae on anterior half of vertex much stronger, preannular more well defined (more distinctly isolated from semiannular), propodeal sculpture stronger, with deeper punctures, and yellow around antennal foramina weak and contiguous, or absent. One «ctremely small specimen observed for this species (3.8 mm) similar to P. rufoomatus, especially in function of a widened gena and temple, presence of yellowish around antennal foramen (although diffiise and pale), and similar sculpture on vertex and propodeum. Also with octreme values for morphometric ratios (note extreme values for sp. I on Figs 252-271). M ale Unknown. Distributional and Seasonal Records. Four specimens from the northeast (Fig. 413), collected 23 April, 17-18 September, 28 November. 353 Parast^haneilus brevis Aguiar il sp. (Figs 252-271 (sp. 2), 280-288,414) E tym ology. L. brevis, short In reference to length of colo (pronotum). Material Examined. Holo^e: Gap Cr., 5 Km ESE Mt. Finningan, QId.,I5.50S I45.20E, 13-16 May 1981, at light; from alcohol collection; I. D. Naumann. OSUC21025 (ANIQ. Paratype: 1 ? (ANIC). Female Length. Mean 10.8 mm (range 10.3-11.3 mm). H ead. Flagellomeres normal. Frons areolate-rugose, more transverse near eyes; frontal carina obscure or absent. Vertex (Fig. 280) around posterior ocelli irregularly or subradially rugose; behind interocellar carina and centrally irregularly or transversely strigate; posteriorly transversely strigate, reaching occipital carina centrally only; temple mostly polished smooth, strigations ending 6 r from genal stripe; posteriorly somewhat irregularly rugose near ^ e s on parade. Centro-Iongitudinal impression present posterioriy, weak, visible with tangential light only. Genal angle weakly or moderately promment. Gena polished smooth. Occipital carina relatively wide dorso-Iaterady, very narrow or almost absent dorsally and ventrally. Posterior margin of head concave in dorsal view. Occiput polished smooth. Post-genal bridge anteriorly with rugulose from hypostomal carina, otherwise weakly alutaceous or polished smooth. Hypostomal carina apec, below mandible base, strongly rugulose or rugose. 354 Thorax. Prostemum mostly polished smooth; some oblique weak carinae subapically inside prostemal depression, and mesally on base. Pronotum (Figs 281,284): colo very short; 4-S oblique carinae laterally, which are weaker dorsally, absent dorso- centrally, where colo is weakly alutaceous; pronotal fold very high, stout, shallowly excavated, centrally collapsed; preannular very short, almost indistinct from semiannular, wich is broadly polished smooth on dorso-posterior margin; transversely strigate dorsally and antero-lateraUy; latero-posteriorly, at least more dorsally, longitudinally strigate or carinate-rugose including pronotal lobe; latero-dorsally also with some foveolae between carinae; alutaceous laterally on apical margin of preannular; femoral impression broad and shallow, same transverse sculpture as semiannular, contrasting with longitudinally carinate on ventral area. Mesoscutum ^igs 281,283) centrally foveolate and alutaceous; more distinctly alutaceous anteriorly; lateral lobe taken by large lacunation, which is punctate or rugulose inside; median sulcus narrow, formed by small, partially fused punctations; notaulus formed by partially fused lacunae. Axilla ^ g . 283) anteriorly finely transversely alutaceous, otherwise covered with small lacunae, punctations and rugosities, subconcentricaQy. Scutellum (Figs 283,285) punctate-punctulate laterally; polished smooth or arcuately transversely strigulate centrally and anteriorly; posterior margin broadly polished smooth. Mesepistemum finely areolate-rugose, or finely mgulose, and punctate; finely rugulose and sparsely shallow-foveolate anteriorly; dorsal part polished smooth ÇFig. 286); ventral part finely transversely alutaceous, polished smooth near discrimen, which is foveolate. Proppdonn ^ g s 285-286) foveolate to areolate-rugos^ and punctate; alutaceous between; sculpture denser centro-longitudinally m the paratype; 355 flank finely alutaceous, partially or entirely with same general sculpture o f propodeum. Parapetiolar fovea transversely carmate, polished smooth between, boundaries indistinct. Spiracular groove indicated by strong, aligned depressions; spiracular carina distinct; spiracular groove and carina partially blended with propodeum sculpture, ending at flank or further, near crenulate sulcus. Metapleuron (Fig. 286) centrally and dorso-posteriorly areolate-rugose; polished smooth dorso-anteriorly; dorsally scrobiculate along spiracular carma; ventrally polished smooth. Post-foveolar area with a few weak transverse carinae or polished smooth. Interfoveolar area scrobiculate on anterior 0.75 or more. Ventral side of fourth tarsomere of fi'om and mid legs normal, not prolonged anteriorly. Hind coxa densely transversely strigate, coarser and more regular basally; lateral side dorso-centrally with weakly concave area finely transversely strigate, almost alutaceous. Hind femur entirely strongly alutaceous (Fig. 288). Hind tibia finely and longitudinally alutaceous laterally and mesally on constricted part; dilated part weakly alutaceous to polished smooth, obliquely carinate ventrally (Fig. 288). Front wing: vein 2Cua completely tubular (Fig. 287). Pterostigma sub-elongate, apex more or less acute. Abdom en. Petiole dorsally finely rugose basally, otherwise normally or finely transversely strigate; apmc broadly polished smooth; ventrally finely areolate-rugose and punctate. Tergite 3 on extreme base longitudinally rugose, otherwise polished smooth, T4- 7 apparently polished smooth (feintly transversely microalutaceous), T8-9 alutaceous, strongest on T9 (Fig. 282). Pygidiai impression perfectly semicircular, very broad. Pygidium a narrow C-shaped stripe ^ g . 282). 356 C olour. Black. Genal stripe yellowish or ivory, almost reaching occipital carina. Caramel coloured basal 0.25 antennae, around antennal foramen, mandible (except black apex), front and mid legs on coxa, trochanter and femur, tip of hind coxa, hind tibia at least on dilated part, apical segments of all tarsi, except black tip of apical tarsomere, most of front basitarsus basally; also front and mid tibia, but lighter. Yellowish 2nd and 3rd front tarsomeres. Whitish or pale-yellow mid tibia apex, correspondent basitarsomere, and 2nd tarsomere; yellowish hind basitarsus. Hind trochanter partially reddish in holo^e. Wmg membrane lightly or conspicuously yellowish brown infiiscate, stronger near 2Cua knob; veins brown. Ovipositor valves brown. D iagnosis. Pronotum, especially colo, very short (Fig. 281). Semiannular strongly strigate (Fig. 284). Vein 2Cua entirely tubular (Fig. 287). Pterostigma elongate, somewhat pointed. Wings unfrbrmly light amber-infbscate. Anterior and lateral tubercles concolourous with frons. All flagellomeres, except first and last three, with numerous, relatively large placoid sensilae underneath (difficult to see if specimens even lightly grea^). Rem arks. Similar to P. maculatus n. sp., but immediately distinguished by having vein 2Cua entirely tubular and wmg membrane unffi)rmly weakly infiiscate; also distinguished by diagnosis above, vertmc sculpture and colour pattern of legs. General structure and microsculpture more similar to some New Guinean and Bornean species than to Australian species, suggesting this species could be a recent introduction from one of those areas. 357 M ale Unknown. Distributiaml and Seasonal Records, Two specimens from the North-northeast (Rg. 414), collected 13-16 May, 14 March 14. ParastephaneUus laevigatus Aguiar n. sp. (Rgs 252-271 (sp. 3), 289-298,415,437) E tym ology. L. laevigatus^ smooth. In reference to sculpture of scutellum. M aterial Examined. Holotype: 9,24 Km N by W of Mareeba, 16.47S, 145.22E, Qld., 24Nov 1981, at Hght, D. H. CoUess. OSUC21030 (ANIC). Paratypes: 5 99,1 (ANIC, UQIC). Fem ale Length. Mean 10.1 mm (range 6.8-13.6 mm). H ead. Basal flagellomeres moderately inflated apico-ventrally in small specimens. Frons subareolate-rugose or transversely areolate-rugose; firontal carina obscurely or distinctly defined. Vertex (Figs 289-290) fi-om concentrically undulate-rugose anteriorly to transversely undulate-rugose or transversely strigate posteriorly, rugosities reaching occipital carina centrally; more irregularly rugose near ^ e s in large specimens; temple polished smooth. Centro-Iongitudinal impression absent. Genal angle weakly prominent, (kna polished smooth. Occipital carina very narrow, slightly wider laterally. Posterior 358 margin o f head concave in dorsal view. Occiput polished smooth. Post-genal bridge and occiput polished smooth. Hypostomal carinaapex, below mandible base, polished smooth. Thorax. Prostemum polished smooth; usually a few rugosities or fbveolae on antero-lateral margins; prosternai depression deep. Pronotum (Figs 291-294): colo arcuate-strigate; strigation sometimes stronger postero-laterally, sometimes well spaced and polished smooth between; deep and wide depression posteriorly, ending within pronotal fold, which is normal, moderately excavated; preannular and semiannular dorsally transversely strigate, changing to obliquely or longitudinally strigate laterally; semiannular, especially laterally, with sparse or numerous, small or large, shallow foveolae; sometimes mostly polished smooth in small specimens; femoral impression indistinct; ventral area longitudinally strigate. Mesoscutum (Fig. 291): Median lobe posteriorly, median sulcus, and notaulus, lacunose or densely foveate; lateral lobe usually almost entirely taken by one large and shallow lacune; median and lateral lobes anteriorly alutaceous. Axilla (Figs 292- 293) punctate to lacunose and alutaceous, or rugulose-lacunose to rugose-Iacunose. Scutellum (Figs 292-293) laterally foveate or with moderate size lacunae, otherwise usually weakly alutaceous, almost polished smooth, or sometimes distinctly alutaceous; also usually sparsely punctate or puncticulate. Mesepistemum (Figs 296-298) anteriorly punctate- or foveolate-rugulose, otherwise &om foveate-punctate (small specünens) to lacunose-punctate, with small polished smooth areas in between centro-dorsaUy; dorsal part polished smooth; ventral part foom polished smooth to transversely alutaceous; discrimen foveolate. Propodeum (Figs 295,297) foveate, punctate, and finely alutaceous; foveolation sometimes very dense centrally, almost rugose; flank sometimes «cclusively 359 alutaceous. Parapetiolar fovea polished smooth or transversely carinate. Spiracular groove somewhat inconspicuous, defined by aligned fbveolae and spiracular carina, which is complete, reaching flank. Metapleuron (Fig. 296) polished smooth dorsally, small fbveolae aligned along spnacular carina; areolate-nigose centrally; transversely carinate ventrally and on post-foveolar area. Interfoveolar area polished smooth or transversely carinate. Ventral side o f fourth tarsomere of front and mid legs normal, not prolonged anteriorly. Hmd coxa from weakly to strongly transversely strigate more frregularly or subareolate- rugose basally; strigations stronger on base of hairs; flattened centro-dorsal area on lateral side not differentiated, or small, fidntly roicroreticulate/alutaceous. Hind femur laterally from polished smooth basally to distinctly alutaceous on apex; mesally weakly alutaceous. Hind tibia finely longitudinally to obliquely alutaceous; dilated part obliquely carinate ventrally. A bdom en. Petiole rugose basally, otherwise transversely strigate; ap«c broadly polished smooth; ventral sculpture similar, but coarser. Tergite 3 polished smooth, T4-8 weakly microaciculate transversely, T9 alutaceous. Pygidial impression large, triangular. Pygidhim narrow, slightly elevated. C olour. Black; some brown or reddish on appendices and extremities; more brownish/reddish in small specimens. Base of coronal tubercles and mteroceOar carinae ferrugineous. Genal stripe narrow, white to pale-yellow, adjacent to ^ e margin, ending on temple &r from occipital carina ^ g . 289), or broader dorsally, ending at occipital carma (Rg. 290). Pale yellow or white on 0.2 basal and 0.1 apical of mid tibia, and basal 0.3-0.8 mid basitarsomere. Apical tarsomeres blackish. Yellowish brown or reddish basal G.3-0.7 360 flagelum, apex colo, posterior margm semiannular (sometimes), front and mid tibiae, trochanters, compressed part and apec of dilated part bind tibia, tarsi (eccepetd as noted above), propodeum near petiolar foramen (sometimes), and apex of petiole and «ctreme base of T3; darker on front and mid femora than in correspondent tibiae. Wing membrane weakfy^ infüscate or hyaline; veins reddish brown or pale-brown. Ovipositor valves dark brown. Front wing: vein 2Cua tubular on basal 0.3-0.4. Pterostigma weakly pointed. D iagnosis, Absolutely no yellow stripes on frons. Temple polished smooth, vertex strigation not reaching genal stripe ^igs 289-290). Scutellum shining, mostly polished smooth or weakly alutaceous between macrosculpture ^igs 292-293). Mesepistemum foveate-punctate or lacunose-punctate, polished smooth spaces in between (Figs 296, 298). Metapleuron dorsally, from pleuropropodeal fovea to posterior magin, always polished smooth (Fig. 296). Medium and small specimens partially polished smooth or weakly sculptured on semiannular and mesally on hind femur. Compressed part of hind tibia, apex o f hind tibia sometimes, and hind tarsi usually yellowish brown; hind tarsi always lightly coloured. Rem arks. Very shnflar to P. (kmellicus, but scutellum sculpture sparse, especially centFO-posteriorly; alutaceous on scutellum and propodem much weaker or absent, resulting in polished smooth or almost so between punctures/lacunae. Overall sculpture on propodeum similar, but never with the typical 'areolate-rugose laterally and densely punctate/puncticulate centrally* o f P. damellicus. A few small P. damellicus with similar scutellum sculpture^ but sculpture on temple, pronotum, mesepistemum, and hind femur mesally dififerent. 361 Also similar to P. isogfyptos, from which it can be difrbrentiated by having more irregular sculpture on vertex, temple and dorsal part of metapleuron polished smooth, semiannular, scutellum, and bind femur mesally partially polished smooth, base o f mid tibia sometimes pale-yellow, and mesepistemum with polished smooth spaces between general sculpture. Two morphotypes, keyed below. I Genal stripe narrow, not getting wider dorsally, ending as soon as it reaches temple, far from occipital carina ^ g . 289). Compressed part of hind tibia and basitarsus reddish or yellowish brown ...... Morphotype I r Genal stripe getting wider dorsally; broadly crossing the temple, reaching occipital carina (Fig. 290). Compressed part of hind tibia brownish. Hmd basitarsus light brownish yellow. [Males with colour similar to females, but genal stripe not reaching occipital carina] ...... Morphotype H Morphotype I is apparently the most common, corresponding to typical specimens for the species. Morphotype H is based on two fomale specimens (OSUC21038 and OSUC21039, both at ANIC) collected on the northernmost limit of the known geographical range for the species (Fig. 415); it is very similar to, and could be conspecific with some P. damellicus. fri spite of intensive mvestigation, results are still inconclusive, and further studies will be necessary to define whether a subspecffîc status for morphotype E is the most correct solution. Morphotype E is also similar to typical P. laevigatus, except for the following. Genal stripe gettmg wider dorsally, crossing temple^ ending at occipital carina. Mid 362 basitarsus white or whitish on basal 0.6 or more. Apex and basal 0.2 o f mid tibia whitish or pale-yellow. Compressed part of hind tibia brownish; apoc of dilated part yellowish or orange. Hmd basitarsus clear brownish yellow. Legs and antennae mostly brownish or yellowish, with very little reddish. Scutellum sculpture as for P. damellicus. M ale Similar to female. Rem arks ^dale). Similar to some males ofP. platyceratus, but with head uniformly brown (except for genal stripe), as opposed to lightly coloured yellow stripes along eye margin or frons in P. platyceratus. Similar to P. isogfyptos and some P. damellicus, from which it can be distinguished by having weak overall sculpture, polished smooth areas on temple, pronotum, scutellum, mesepistemum, and hind femur. Foveolae on scutellum aligned laterally, differing from more irregularly distributed and centrally present sparse or dense foveolation of P. damellicus. Distributional am i Seasonal Records. Northeast (Fig. 415); probably flying all year(Rg. 437). Parastephanellus isogfyptos Aguiar n. sp. (Figs 252-271 (sp. 4), 299-303,412,433) Etym ology. Gr. isos, equal, like; Gr.gfyptos, carved. En reference to sculpture of vertex. 363 Material Examined, Holotype: Î, NSW, Sydn^, Pennant Hills, Cumberland S. P., Acacia longffolia tree (ca. 4m) dead ca. 12 months; timber collected 8 Apr 1986, adult emerged 23 Nov 1986, G. A. Webb. OSUC21077 (FCNI). Paratypes: 4 ? ?, 9 cfd* (ANIC, FCNI, QMBA, UQIC). Fem ale Length, Mean 9.0 mm (range 7.3-11.1 mm). H ead, Flagellomeres normal. Frons areolate-rugose, sometimes subtransversely; transversely rugose along ^ e margin; frontal carina distinct. Vertex (Fig. 299) from transversely rugose behind hind ocelli to transversely strigate posteriorly, reaching occipital carina; temple strigate, strigation ending at genal stripe. Centro-longitudinal impression 6int or absent Genal angle moderately prominent. Gena polished smooth. Occipital carina very narrow laterally, marginated or absent dorsally. Posterior margin of head concave in dorsal view. Occiput polished smooth. Post-genal bridge usually polished smooth, sometimes weakly alutaceous anteriorly. Hypostomal carina apex, below mandible base, from alutaceous m small specunens, to finely longitudinally rugose in large ones. Thorax, Prostemum mostly polished smooth, apically alutaceous, basally with a few oblique rugosities; sometimes entfrely alutaceous, obscurely centrally. Pronotum (Figs 300-303): colo with several carinae on each side, some complete; polished smooth near and within pronotal feld; pronotal feld normal, moderately excavated; preannular and semiannular transversely strigate, more densely on semiannular; femoral impression feint 364 or indistinct; ventral area longitudinally strigate. Mesoscutum broadly microreticulate anterioriy, otherwise strongly rugose-punctate. Axilla rugose-Iacunose. with alutaceous between, which is more conspicuous anteriorly. Scutellum (Mg. 301) rugose-Iacunose laterally, from densely micropunctate anteriorly to arcuately transversely strigulate centrally; strigulate sometimes on almost entire scutellum; sculpture frding apically. ap«c almost polished smooth. Mesepistemum finely areolate-rugulose. especially anteriorly, with dense microreticulation or punctulation, which is more abundant centro-ventrally and posteriorly; sometimes strongly alutaceous and microreticulate in small specimens; dorsal part fidntly obliquely alutaceous/microreticulate, or polished smooth; ventral part transversely alutaceous, sometimes faint, almost polished smooth; discrimen foveolate. Propodeum (figs 301-302) finely areolate-rugose with microreticulate between; areolae bigger centrally and posterioriy; flank transversely alutaceous. Parapetiolar fovea weak, boundaries indistinct, posteriorly formed by large polished smooth areolae. Spiracular groove ill-defined, indicated by aligned areolae, or indistinct; spiracular carina distinct, but weak or irregular, usually partially blended with propodeum sculpture, ending near or at flank. Metapleuron (Fig. 302) dorsally polished smooth or transversely carinate, with shallow small punctures, always shining; centrally areolate-rugose. much coarser than on propodeum; ventrally transversely carinate. contiguous with areolate of lateral part, extending to post-fi>veolar area. Interfoveolar area polished smooth. Ventral side of fourth tarsomere o f front and mid legs normal, not prolonged anteriorly. Hind coxa laterally on basal 0.2 sub areolate-rugos^ with dorso-central finely sculptured on somewhat depressed area, otherwise transversely strigate, including mesally, where it is also densely punctulate 365 ventro-basally. Hind femur conspicuously alutaceous on both sides, weakly basalfy. Hind tibia alutaceous much finer than on bind femur, obliquely on constricted part; dilated part obliquely strigate ventrally. Front wing: vein 2Cua tubular on basal 0.1-0.3. Pterostigma more or less pointed. Abdom en, Petiole from rugulose basaly to transversely strigulate subapically, sometimes finely; apex broadly polished smooth; ventral sculpture similar, also densely punctate or microreticulate, at least in part Tergite 3 on extreme base rugose, otherwise polished smooth, T4-8 dull and distinctly transversely microaciculate, T9 alutaceous. Pygidial impression triangular, sometimes concave sides, forming a collapsing triangle. Pygidium narrow, convex, emarginate. Colour. Deep black to brownish black, with a few yellowish brown or reddish parts. Genal stripe narrow, creamy whitish or yellowish, ending fer or very near occipital carina. Yellowish posterior margin of semiannular, including pronotal lobes, front tibia, except blackish anteriorly, mid tibia and mid tarsomeres. Yellowish or reddish yellow antennae basally, hind leg on trochanter, base of femur, sometimes constricted part of tibia, apmc of tibia, and tarsomeres, except blackish apex of apical tarsomere. Whitish basal 0.25-0.33 mid basitarsus. Reddish base of coronal tubercles and interocellar carmae. Wing membrane hyaline or light-yellow infiiscate; veins light or dark brown. Ovipositor valves brown o r dark brown. D iagnosis. Vertex microsculpture, mcluging temple until genal stripe, regularly transverse (pg. 299). Scutellum distinctly transversely alutaceous or finely transversely strigate (Fig. 301). Pronotum and mesothorax concolourous. Mesepistemum rugulos^ 366 with tendency to microreticulate centro-ventrally. Metapleuron dorsally with fine transverse microsculpture (Fig. 302). Prostemum always alutaceous, although usually weakly or inconspicuously. Body mostly black or dark brown, mid tibia basally not yellowish; compressed part and tip of hind tibia reddish. R em arks, Similar to P. laevigatus n. sp., fi’om which it can be distinguished by having a distinctly regular vertex and temple scultpture, dorsal part of metapleuron always with fine transverse scultpture (fi’om alutaceous to strigate), semiannular completely strigate, scutellum finely transversely sculptured, without polished smooth areas or fbveolae centrally, hind femur distinctly alutaceous mesally, mid tibia more or less uniformly coloured, not pale-yellow basally, and mesepistemum mgulose. Similar to P. damellicus^ fi'om which it can be separated, in addition to the characters in the key, by showmg the following: Propodeum sculpture finer, transverse (compare Figs 301.and 311). T3 polished smooth (accept sometimes fointly and finely alutaceous on apical margin). Scutellum dull, finely and densely transversely arcuate-strigose. Pygidial impression with weakly or strongly concave sides, forming a collapsing triangle, with a notch or channel anteriorly. Mid tibia uniformly coloured or with darker tones basally, never with yellow. Compressed part and tip of hind tibia usually distinctly light to dark ferrugineous or yellowish brown, contrasting with black on dilated part. Mesepistemum rugulos^ posterior half with tendency to microreticulate. In small specunens, most rugose patterns tend to alutaceous, and delicate impressions (e.g., centro-longitudinal Impression on vertoc) tend to disappear. 367 M ale Similar to the female. R em arks O^ale). Sunilar to males o ff. laevigatus n. sp. and P. damellicus^ from which it can be distinguished especially by its dull, finely microsculptured scutellum, and uniformly transversely strigate vertex and temples. Sculpture of vertex, gena often wide, overall sculpture of thoraiq and small size sometimes make this species very similar to males o f P. ntfoomatus, from which it can be distinguished by having a more elongate hind coxa, no yellow marks ventrally on frons, and pronotum comparatively longer. The smallest specimen examined is atypical, with vertex sculpture somewhat irregular, polished smooth temple, and weak overall sculpture. Distributional and Seasoned Records. East coast ^ig. 412); apparently more abundant from November through March (Fig, 433). Associations. Several specimens emerged from a branch oîAcacia longifolia (Andr.) Willd. (Leguminosae), removed from a tree about 4 m tall, dead for about a year. The branch was collected in April S, and specimens of P. isogfyptos emerged as follows: one 9 11 November; one d* 23 November; one 9 and two «fcf 25 March o f the following year. Parastephandlus damellicus (Westwood) (Figs 252-271 (sp. 5), 304-314,419,432) Stephemus Damellicus Wescwoody 1S74 (nec Enderlein, 1901. 1906): 127; Schletterer 1889:116. 368 Stephanus damellicus: Dalla Torre 1902:6. Foenatopus damellicus: Szépligeti 1902:529. Parastephanus dameUicusr. Enderlein 1905:475. Parastepfumellus damellicus: Kieffer 1908:5; Elliott 1917b: 129; Brues 1918: 100;Elfiott, 1919a: 29;ElHott, 1922:747, 753 HemisteplumusD Stephanus (JParastephanellus) damellicus.^ Elliott 1917c: 266. ParastepfumellusalbicepsWiiott, 1919b: 130; Elliott 1922: 747, 751. N ew S y n o n y n ^. E tym ology. Dedication to the collector, cited simply as 'Darnell' by Westwood (1874). Material Examined. Holotype: 9 Parastephanellus albicepsEVàoVt, Victoria, 'c. 7.9.1901.’ OSUC20558 (BMNH). Other specimens examined. 32 ? ?, 18 d‘ AMSA, ANIC, BMNH, FCNI, MVMA, NSWA, QDPC, QMBA, UQIC). Fem ale Lengjth. Mean 10.6 mm (range 5.3-14.3 mm). H ead. Eagellomeres normal. Frons areolate-rugose; rugose in small specimens; fiontal carma distmct. Vertex (Fig. 304) sculpture transverse; usually narrow centro- longttudinal area regularly transversely rugose; laterally from transversely undulate-rugose anteriorly to strigate or substrigulate posterioriy, readüng occipital carma and spreading 369 on most of temple. Centro-Iongitudmal impression weak, but usually distinct, at least in part. Genal angle weakly or moderately prominent. Gena polished smooth. Occipital «arina very narrow, wide dorso-laterally in large specimens. Posterior margin o f head concave in dorsal view. Occiput polished smooth. Post-genal bridge usually polished smooth; weakly alutaceous on colour type 3; substrigate narrowly near and along hypostomal carina. Hypostomal carina apex, below mandible base, polished smooth or weakly rugulose. Thorax. Prostemum polished smooth, with apex weakly alutaceous; sometimes subapically rugulose or subpunctate. Pronotum (Figs 305-306,308): colo varying from homogmieously arcuate-strigulate to sometimes with Just a few well marked arcuate carinae; polished smooth or weakly alutaceous between carinae; anterior strigulation or (arinae contiguous dorso-anteriorly, interrupted and entering pronotal fold posteriorly; usually a centro-posterior polished smooth depression, also entering pronotal fold, which is stout, deeply excavated; dorsally emarginated; sometimes centrally collapsed; preannular transversely strigate, sometimes strongly; apical margin finely transversely alutaceous; semiannular latero-posteriorly usually more irregularly rugose, substrigate or subareolate- lugose, sometimes with shallow fbveolae or fine rugosity, semiannular otherwise finely strigat^ from transversely arcuate dorsally to obliquely arcuate laterally; femoral hnpression indistinct; its equivalent area, ventral area, and pronotal lobe longitudinal^ strigate or sometimes longitudinally carinate. Mesoscutum: Median sulcus and notaulus foveolate to scrobîculate; median lobe posteriorly rugose-lacunose-foveate or foveolate; anteriorly alutaceous; half or little more of lateral lobe taken by large shallow lacune 370 posteriorly; anterioriy alutaceous. Axilla ^ g s 307.309) from strongly subareolate-rugose. and partialy microreticulate or punctate (large specimens), to rugulose- or alutaceous- punctate (small specimens); sculpture sometimes concentrically arcuate. Scutellum ^ g s 307.309-311) areolate-rugose and densely punctate or lacunose; areolae much larger laterally, finer or absent centrally; punctation/punctulation much denser centrally, usually dispersing laterally between and inside areolae; sometimes also with puncticulate or shining alutaceous intermixed; in a few specimens central sculpture arcuately arranged; posterior margin polished smooth or alutaceous. Mesepistemum ^ g s 312-313) areolate- rugulose. sometimes weaker or absent centrally or centro-dorsally, and punctulate, with some microreticulate or alutaceous, especially ventrally; dorsal part polished smooth; ventral part transversely alutaceous; discrimen foveolate. Propodeum (Figs 311,314) centro-Iongitudinally areolate-rugose, laterally strongly alutaceous and puncticulate/punctate or lacunose/variculose; sometimes areolate-rugose also laterally, but weaker than centrally, with alutaceous on flanks only, or subareolate-rugulose centro- longitudinally or centro-posterioriy; flank as on lateral propodeum. but sometimes alutaceous or finely alutaceous. Parapetiolar fovea ill-delimited, anterioriy inconspicuous, hidden by propodeal sculpture posterioriy as small polished smooth area. Spiracular groove mdicated by weakly defined areolae or frjveolae; together with spiracular carina usually ending at some distance or at flank; rarely both indistinct; always ill-defined or irregular on colour type 3 specimens. Metapleuron (Fig. 313) centrally prominent, subtriangular, areolate-rugose; dorsally polished smooth or weakly alutaceous. transversely rugulose or with punctulation next to and along spiracular carina; antero- 371 dorsally sometimes microreticulate. Ventrally and on post-foveolar area transversely rugose, usually contiguous with central areolae. Interfoveolar area with about two transverse carinae anterioriy, otherwise polished smooth or 6intly transversely carinate. Ventral side of fourth tarsomere of front and mid legs normal, not prolonged anteriorly. Hind coxa mostly transversely strigate, subareolate-rugose basally; usually with slightly concave or flattened area centro-dorsally, which is very finely transversely rugose or microreticulate. Hind fomur strongly and densely alutaceous; sometimes polished smooth basally. Hind tibia finely, but distinctly alutaceous, which is more oblique or longitudinal on constricted part; dilated part strongly obliquely carinate ventrally. Front wing: vein 2Cua tubular on basal 0.2-0.4. Pterostigma elongate more or less pointed. Abdom en. Petiole transversely strigate, coarser basaly, polished smooth apex; ventral sculpture similar, but coarser and more irregularly transversely strigate. Tergite 3 on basal 0.5-0.8 weakly alutaceous, or sometimes apically transversely and weakly microaciculate rarely polished smooth, T4-8 transversely microaciculate (distinctly stronger than in T3), T8 apically and all T9 strongly alutaceous. Pygidial impression triangular, deep. Pygidium narrow, emarginate. Colour. Dark brown or black; some yellowish brown on legs. Whitish genal stripe and hind leg colour in three common patterns: (1) genal stripe narrow, endmg at some distance from occipital carina; bind tarsi completely, hind tibia apex and laterally up to basal 0.4, reddish; (2) genal stripe narrow ventrally, much wider on temple, almost reaching occipital carina; hind tibia usually reddish only apically, sometimes also yellowish brown on compressed part, or almost entirely of this colour, although always darkened 372 centrally; hind tarsi yellow, yellowish brown, or orange; (3) genal stripe as in colour type 1; bind tibia and tarsus as in colour ^ e 2. Intermediate colour forms and variations do occur. Pronotum usually with reddish hue, or sometimes brownish; front legs reddish and black or brown and yellowish brown; mid femur brownish or reddish; mid tibia white (or ivory) for a short distance basally, then shortly black; apical 0.7-0.8 brownish; sometimes whitish apex; sometimes entirely brown; hind basitarsus basal half whitish. Propodeum sometimes reddish near petiolar foramen. Wing membrane slightly infiiscate, sometimes stronger around vein 2Cua, near crossvein Icu-a, and cell l+2+3Rs; veins brown to dark brown (sometimes with reddish hue), usually lighter pterostigma. Ovipositor valves dark brown. D iagnosis. Absolutely no yellow stripes on frons. Vertex transversely strigate centrally, changing to more irregularly laterally; postero-Iateral sculpture taking most of temple (Fig. 304). Scutellum usually densely sculptured, with following general structure (Figs 307,309-310): areolate-rugose and densely punctate/puncticulate; areolae much larger laterally, finer or absent centrally; punctation much denser centrally, usually dispersing laterally between and inside areolae. Mesepistemum finely sculptured, mgulose or with some microreticulate, without polished smooth spaces (Fig. 312). Interfoveolar area with about two transverse carinae anteriorly, otherwise polished smooth or frintly carinate. Rem arks. This species has a high intraspecific morphological variability, and this problem is fiirther aggravated by its similarity^ with some equally variable species, m particular PL platyceratus n. sp., P. isogfyptos n. sp., and P. laevigatus n. sp., and by the 373 fîict that the range of morphological variability of these new taxa is likely to be much higher than what could be observed. The morphological differences discussed below must, therefore, be considered critically. Very similar to f . laevigatus n. sp. and f . isogfyptos n. sp.; somewhat similar to P. strigosus n. sp. and P. trifasciatus n. sp. From P. laevigatus it can be distinguished by having areolate-rugose and centrally densely punctate scutellum, axila punctate with polished smooth or alutaceous between, and propodeum areolate-rugose centrally to punctate/lacunose laterally, with alutaceous or punctulate between and inside patterns. From P. isogfyptos it can be distinguished, in addition to characters in the key, by showing the following: Propodeum sculpture coarse, with large areolae, especially centro- longitudinally, this sculpture less or not transverse (compare Figs 311 and 301). T3 apparently polished smooth, but almost always with fine transverse aciculation or weak alutaceous, at least partially. Scutellum punctate, or sparsely punctate with some foveolae, polished smooth or weakly alutaceous between, or densely arcuate-rugose with puctures and areolae. Pygidial impression usually with sides conv«c (dome-shaped) or straigth (triangular), rarely weakly concave; never forming a channel or notch basally. Base of mid tibia often tricoloured yellow-black-brown. Compressed part of hind tibia dark brown or black, more rarely reddish, never sharply contrasting with black of dilated part. Mesepistemum without or with very little microreticulate. T3 microsculpture must be evaluated with great care, since it is usually very weak and only slightly (although consistently) different in these species. 374 From P, strigosus and A trifasciatus tt can be dtfi&rentiated by lacking totally yellow maries on frons, and by its distinct scutellum sculpture. M ale Similar to female. R em arks (Male). Similar to males of P. laevigatus n. sp. and to P. isogfyptos n. sp., from which it can be distinguished chiefly by presence of foveolae centrally on scutellum (similar to Fig, 307), and pronotum usually reddish black. Vertex sculpture, gena width, and very weak yellowish ventrally on frons (rare) sometimes similar with males o f P. rufoomatus, from which it can be distinguished by having a more elongate hmd coxa and foveolae always present on scutellum. The presence of foveolae centrally on scutellum is diagnostic for males of P. damellicus, but intraspecifrc variation is high. The male scutellum is usually densely covered by very strong foveolation and some punctation, producing a coarse texture, but in rare occasions only a few foveolae are present, with spaces between polished smooth or almost This condition is shnilar to that observed in the single known male of P. laevigatus, but the foveolae on this specimen are small and somewhat aligned laterally, while large, more irregularly distributed, including centrally, in males of P. damellicus, Synor^nny, The last report about the location of the h o lo ^e of the present species appears in Elliott (1917c), which mentions OXLM as the lodging museum. Nonetheless, this h o lo ^e could not be found there. The Hymenoptera collection at OXUM could not 375 yet be completely reorganized since its evacuation to a more protected place during the Worid War H (C. O’Toole, pers. comm.). Therefore, it is possible that the holotype ofP. dam ellicus may be still deposited there, although with its precise location unknown. fo spite of the unavailability of the holotype; Westwood’s original description of P. damellicuSy the redescription o f the holotype ^Uiott 1922), and a series of specimens of this species studied by Elliott (1922) match well with the holotype of P. albiceps Elliott, clearly mdicating its synoitymy. Parastephanellus picticeps Roman is also similar to P. damellicus, but the holotype ofP. picticeps is somewhat intermediate between the present species and P. rufoom atus (see discussion for P. p icticep s under species inquirenebtm), so that its tynonymy with P. dam ellicus cannot be reliably determined at this time. Pcarastephanellus damellicus sensu Enderlein (1901, 1906), from New Guinea, is a distinct species, renamed as P. rubripictus by Elliott (1917b). Distributional arul Seasonal Records. Throughout the eastern coast (Fig. 419); flying all year, but more abundant from January through March (Fig. 432). Associations. Female OSUC2101S emerged from ‘C. australis’ (label data; genus and classification not found) on September 29. Parastephanellus albigenuis Aguiar n. sp. (Figs 252-271 (sp. 6), 315-316,416) E tym ology. L. albus, white; L. genuis, knee, joint. In reference to basal colour of hmd tibia. 376 MaterialEsummed. Holotype: Î, 6 KmNTaroom, Qld., 25®36’E, 149®46’S, 7 Fd) 1981, mercury vapor light, G. Daniels, A. Daniels, C. J. Burwell. OSUC20965 (UQIC). Fem ale Length, IS.2 mm. H ead. Flagellomerra normal. Frons areolate-rugose; frontal carina absent. Vertex ^ g . 3 IS) arcuate-rugose anteriorly, especially near interocellar carinae, otherwise from transversely rugose anteriorly to transversely strigulate posteriorly, reaching occipital carina broadly, not extending to temple, which is polished smooth. Centro-longitudinal impression absent. Genal angle weakly prominent Gena polished smooth. Occipital carina narrow, dorsally almost absent. Posterior margin of head concave in dorsal view. Occiput polished smooth. Post-genal bridge mostly polished smooth, weakly alutaceous apically. Hypostomal carina apex, below mandible base, weakly alutaceous. Thorax. Prostemum anterioriy alutaceous, otherwise polished smooth. Pronotum (Figs 316,318): colo arcuate-carinate; posterior carmae dorsally interrupted, entering pronotal Aid; a polished smooth depression, almost entnely within pronotal Aid, which is stout, deep, emarginated dorsally; preannular arcuate-rugose; semiaimular dorsally obliquely rugose laterally; centrally partially covered by arcuate pattern from pre-annular area; laterally, preannular, and semiannular anterioriy, transversely rugose and carinate, semiannular otherwise obliquely sculpturate; from strongly strigate ventrally and on pronotal lob^ to weakly strigulate or alutaceous centro-dorsally; femoral impression 377 distmct; long narrow strigation between it and ventral area, which is strongly longitudinal alutaceous. [Mesoscutum broken}. Axilla (Figs 319-320) arcuate-rugose or arcuate- rugulose. Scutellum ^ g s 319-320) strongly transversely alutaceous; also punctate and foveolate^ except centrally and centro-posterioriy. Mesepistemum (Fig. 317, part) rugulose and white-pHose dorsally and anteriorly; centro-Iongitudinally transversely alutaceous; centro-ventrally fium densely punctate posteriorly to punctate and rugulose anteriorly; dorsal part polished smooth; ventral part obliquely alutaceous; discrimen sub- foveolate anteriorly, otherwise simple. Propodeum ^ig s 319-320) strongly alutaceous, puncticulate, and sparsely shallowly lacunose; flank finely alutaceous. Parapetiolar fovea distinct, polished smooth, relatively small, open anteriorly, blending with overall sculpture of propodeum. Spiracular groove inconspicuous, indicated by weak spiracular carina developed on basal G.4-0.5 ÇFig. 317). Metapleuron (Fig. 317) strongly areolate-rugose centro-posterioriy; anteriorly and dorso-anteriorly sparsely punctulate, polished smooth between; sub-scrobiculate along spiracular carina and dorso-anterior margin, reaching pleuropropodeal fovea; dorso-posteriorly and antero-ventrally very densely punctulate; ventro-posteriorly central areolate-rugose fusing with strong transversely carinate of post- foveolar area. Diterfoveolar area contiguous with very lon& eliptical metapleural fovea, wich is polished smooth inside. Ventral side o f fiiurth tarsomere of front and mid legs normal, not prolonged anteriorly. Hind coxa ventro-basally, especially mesally, micro- areolate or microreticulate; a large flattened to slightly concave dorso-central area very finely strigulate almost alutaceoue otherwise transversely strigate. Hmd femur strongly longitudinally alutaceous, except almost polished smooth basally on lateral side. Hind tibia 37& strongly longitudinally alutaceous on dilated part (weaker than on hind &mur); Biely obliquely alutaceous on compressed part; dilated part obliquely carinate ventrally. Front wing: vein 2Cua basal 0.3 tubular. Pterostigma sub-elongate, more or less pointed. Abdomen. Petiole finely transverse!^ rugulose dorsally; microreticulate or microareolate ventrally; apex dorsally polished smooth, laterally microreticulate. Tergite 3 basally with a few rugae, otherwise polished smooth, T4-6 polished smooth, T7 very weakly alutaceous, T8 transversely alutaceous, weak basally, distinct apically, T9 strongly alutaceous (Fig. 321). Pygidial impression large, very deep, shape similar to a Pygidium convex, weakly emarginate (Fig. 321). C olour. Dark or reddish brown; pronotum lighter, reddish or brownish; no yellow marks on fi^ons; black gaster and ovipositor. Genal stripe narrow, yellowish, wider on temple, almost reaching occipital carina. Yellowish brown basal anteimal segments, mandible (except black apex), posterior margin of scutellum, base of fi’om femur, apex of fi-ont tibia, fiont tarsi, mid femur basally, mid tarsi except basitarsomere basally, hypopygium. White or pale yellow mid tibia base and apex, mid basitarsomere basal half scale-like apex of hind coxa, compressed part of hind tibia basal 0.4. Reddish brown base of head tubercles and base o f interocellar carinae, hind leg on trochanters, extreme base of femur, tibia apex lateral^, most of dilated part mesally, basitarsomere, and second tarsomere, petiole apex, and «ctreme base of T3. Wing membrane hyaline, somewhat infiiscate near vein 2Cua; veins brown. Ovipositor valves dark brown. Yellowish brown pygidium. 379 D iagnosis, Mid and hind tibia basally creamy white. T3-6 polished smooth (Fig. 321). Metapleuron ventrally and dorso-posteriorly densely punctulate (Fig. 317). Hind femur strongly alutaceous. Absolutely no yellow on feons. Petiole ventrally microareolate. A long straight strigation between femoral impression and ventral area ^ ig . 3 IS). R em arks. Similar to P. rufoomatus, from which it can be distinguished by the diagnose above, and by having sparser vertex microsculpture, temples mostly polished smooth, without anteriorly curving strigulation, gena normally developed, mesepistemum sculpture more complex, without microreticulation (although dense punctulation may be similar to it), and cell 2Cua, and area below it, hyaline. Also similar to P. platyceratus n. sp., from which it can be distinguished, in addition to the diagnosis above, by having vertex arcuate-strigose centrally, changing to transversely strigulate laterally and broadly transversely strigulate posteriorly, and genal angle not prominent. M ale Unknown. Distributional and Seasonal Records, Holotype from southeastern Queensland ^ g . 416), collected 7 February. Parastepkanelltts mSazetos Aguiar n. sp. (Figs 252-271 (sp- 7), 322-327,417) E tym ology, Gr. adiazetos^ not polished. In reference to the sculpture of prostemum. 380 M aterial Examined. Holotype: ?, Brisbane, Qld., 20 Sep 1961, R.G. Winks. OSUC21023 (UQIQ.Paratype: I ? (MVMA). F em ale L ength. 8.4 mm. H ead. Antennae lost. Frons transversely rugose centro-ventrally to obliquely rugose near coronal area; transversely carinate in broad area along eye margin, polished smooth between transversely carinae. Frontal carina not defined, but partial confluence of wrinkles centrally. Vertex OFig. 322) transversely strigate; strigation undulated centro- anteriorly, arcuate posteriorly inside centro-longitudinal impression; strigate reaching tyes radially. Temples transversely strigulate; strigulations not reaching gena, perpendicular to radial striction around eyes. Centro-longitudinal impression weak; wider and more conspicuous posteriorly. Genal angle prominent. Gena polished smooth. Occipital carina very narrow laterally, centro-dorsally marginated, almost absent. Posterior margin of head concave in dorsal view. Occiput polished smooth. Post-genal bridge longitudinally rugulose. Hypostomal carina apex, below mandible base, longitudinally rugulose. Thorax. Prostemum alutaceous apically to microreticulate centrally to weakly microreticulate basally; some weak transverse carmae subapically and on posterior margm. Pronotum QFigs 323,325-326): colo weakly obliquely strigulate laterally, strigulations contiguous dorsally, producing an arcuate-strigulate pattern; stout carina posteriorly curving dorso-laterally within pronotal fold, which is not excavated, indicated by transverse carina; preannular and semiannular strongly rugose; some diamond-shaped 381 microreticulation centro-posterioriy; posterior margin narrowly weakly alutaceous; weakly microreticulate apically on each side. Femoral impression rugose as in semiannular, with polished smooth spaces among rugae; sometimes well marked longitudinal carinae between femoral impression and ventral area, which changes from alutaceous posteriorly to strongly rugose apically. Mesoscutum (Figs 323,326) areolate-rugose, anterior margin narrowly finely alutaceous. Axilla (Fig. 326) mostly areolate-rugose, finely alutaceous on anterior margin. Scutellum ^igs 324,326) densely and microscopically areolate-rugose centrally, laterally more coarsely, as on axdia; posterior margin weakly alutaceous. Mesepistemum (Fig. 327, part) laterally and dorso-anteriorly strongly rugulose and microreticulate; rugulose stronger dorsally and anteriorly, microreticulation more conspicuous centrally, dorsal margin of dorsal part strongly transversely carinate (weakly so on male); posteriorly M itly microreticulate (apparently alutaceous); ventral part strongly transversely alutaceous, except polished smooth near discrimen, which is foveolate. Propodeum (Figs 324,326-327) strongly and transversely subareolate-rugose; flank strongly alutaceous. Parapetiolar fovea not well delimited, indicated by elongate concave area; same sculpture as propodeum. Spiracular groove weak, indicated by spiracular carina, which does not reach flank. Metapleuron (Fig. 327) dorsally strongly transversely carinate; centrally and sub-ventrally strongly subareolate-rugose. Post- foveolar area transverse^ carinate polished smooth between. Interfoveolar area with a fow transverse carmae. Ventral side o f fourth tarsomere of front and mid legs prolonged anteriorly in a process about as long as apical tarsomere. Hind coxa laterally rugulose to rugose on basal 0.25; transversely strigate on apical 0.2; dorso-centrally weakly depressed, 382 very finely rugulose or alutaceous or microreticulate; centro-ventrally, ventrally, and mesally on apical 0.5-0.6 transversely strigulate; basal 0.4-0.S mesally rugulose and microreticulate. Hind femur totally alutaceous, weaker basally on lateral side. EBnd tibia longitudinally alutaceous, finer than on hind femur; dilated part obliquely carmate ventrally. Front wmg: vein 2Cua tubular on basal 0.1. Pterostigma relatively blunt. Abdom en, Petiole transversely substrigate, coarser basally; apex polished smooth; ventral sculpture similar, coarser. Tergite 3 on «ctreme base rugulose, centrally weakly longitudinally alutaceous, which changes to inconsptcuoulsly micro-aciculate subapically, T4-8 transversely microaciculate, T9 distinctly alutaceous (as on hind femur). Pygidial impression triangular. Pygidium basally flat, alutaceous, apically convec, pilose on irregular surfece. C olour. Black, with brownish or yellowish legs and head. Genal stripe not completely developed; large pale-yeQow spot fi’om base of mandible to eye margin, small yellowish spot on genal angle, ferfiom ^ e . Yellowish brown clypeus, mandible except apex, palpi, octreme base of fiont and mid femora, basally and apically on fiont and mid tibiae, all tarsomeres, except apical ones and white basal 0.2 of mid basitarsus, and compressed part of hind tibia. Rufous base o f coronal tubercles and interocellar carina^ apex of petiole extreme base of 13, hind trochanter laterally. Reddish black gena, occiput, post-genal bridge, most of pro- and midlegs. Wing membrane lightly infiiscate stronger on cells ICu and 2Cu; veins and sclerites brown. Ovipositor valves dark brown. D ia ffw sis. Pronotal feld low, not eccavated, differentiated as a transverse carma (Hgs 323,325). Compressed part of hind tibia twice as long as dilated part; scutellum 383 microareolate (Figs 324,326); pronotum strongly rugose CFigs 323, 325); prostemum microreticulate, with subapical transverse carinae; dorsal part of mesepistemum almost totally rugulose and carinate ÇFig, 327); cell IM narrow. Remarks. Somewhat similar to P. tanytarsus n. sp.; distinguished by having prostemum sculpture finer (mostly microreticulate), hind coxa and femur more elongate (sp. 7 on Figs 256,258, and table 3.7), hind femur alutaceous as in dilated part of correspondent tibia, metapleuron centrally not or weakly prominent (Fig. 326) and entirely coarsely sculptured, compressed part of hind tibia distinctly longer than dilated part, vertex sculpture more or less un%rm, colour pattern dififerent. M ale Unknown. Distributional and Seasonal Records. Holotype firom southeastern Queensland ^ g . 417), collected 20 September. Parastephandlus cidlidictyon Aguiar n. sp. (Figs 252-271 (sp. 8), 328-337,418) Etymology. Gr. kallion^ beautdW; Gr. (tiktyon, net In reference to reticulate sculpture of fi’ons. Material Examined. H olo^e: 9,20 Km South Lotus Ck., MEQ, Brigalow forest 26 Apr 1979, E. Dahms. OSUC20971 (QMBA). 384 F em ale Length. 15.2 mm. H ead. Flagellomeres normal. Frons strongly areolate-rugose; frontal carma absent ^ g . 330). Vertec (Figs 328-329) rugose (weakly oblique); posteriorly transversely strigate until occipital carina, frding laterally when it reaches temple, which is mostly polished smooth. Centro-longitudinal hnpression weak, visible centrally and posteriorly under tangential light ^ g . 329). Genai angle weakly, but distinctly prominent. Gena polished smooth. Occipital carina very narrow. Posterior margin of head concave in dorsal view. Occiput polished smooth. Post-genal bridge weakly longitudinally alutaceous. Coarsely longitudinally strigate near and along hypostomal carina; basally this pattern weaker and more fragmented. Thorax. Prostemum (Fig. 333) coarsely obliquely strigate on anterior half and basally; much more sparsely centro-posteriorly. Pronotum (Figs 332, 334,337): colo densely arcuate-rugose anteriorly (along apical emargination); transversely strigate lateral^ frintfy alutaceous or polished smooth centro-posteriorly; pronotal fold flat, but distinct, shallowly excavate centrally collapsed, so that a narrow central portion of preannular is contiguous with colo; preannular and semiannular very strongly transversely strigate dorsally (frner and more irregular centro-dorsally); this sculpture longitudinal on ventral area, including area equivalent to femoral impression, which is indistinct, and on postero-lateral margin, including pronotal lobe and depression in front of it; some weakly defined foveolae dorso-posteriorly. Mesoscutum ^ g . 332): anterior part alutaceous; median lobe densely fr>veate; lateral lobe strigate-alutaceous on anterior margin, otherwise 385 deeply transversely lacunose. Axilla strongly sculptured» approximatelly areolate-rugose with concave areolae, which have irregular, polished smooth surface; a few very small areolae between, Scutellum ^ig . 331) mostly polished smooth, with stout foveae, which are more abundant laterally, sparse centrally. Mesepistemum (Fig. 336, part) confüsed- rugulose, also with microreticulate or alutaceous ventraHy; dorsal part polished smooth; ventral part weakly transversely alutaceous; discrimen fbveolate. Propodeum (Figs 335- 336) sculpture very complex, firom densely foveate to areolate-rugose centro- longitudinally; then with large and small foveae more laterally, with strong microreticulate between; near and along spiracular groove strongly more or less microareolate-rugose, with some microreticulation between; all sculpture patterns reaching flank, which is finely transversely alutaceous. Parapetiolar fovea strongly and broadly transversely strigate. Spiracular groove inconspicuous; position o f spiracular carina indicated by contrasting dorsal sculpture of metapleuron. Metapleuron ^ig . 336) delimited dorsally by line of sub- fbveolae fi’om pleuropropodeal fovea to posterior margin, alongside spiracular carina, dorsal part otherwise broadly polished smooth or fiuntly alutaceous; centrally strongly concave-areolate-mgose, with one short pale hair inside each areolation; ventral part and post-foveolar area very strongly transversely carinate (almost porcate). Interfoveolar area polished smooth, with weak transverse carinae. Ventral side of fourth tarsomere o f fi’ont and mid legs normal, not prolonged anteriorly. Hind coxa latero-ventrally on basal half rugulose and areolate-rugulose, otherwise transversely strigate weaker centro-mesally, coarser dorso-basally; a centro-dorsal flattened area laterally, with finer overall sculpture and weakly transversely alutaceous centrally. Hmd femur distinctly alutaceous on apical 386 0.3 only, âding to practically polished smooth basally; alutaceous much weaker mesally. Hind tibia finely, but distinctly obliquely alutaceous; dilated part strongly obliquely carinate ventrally. Front wing: vein 2Cua tubular on basal 0.15. Pterostigma sub-elongate, apex more or less acute. Abdomen. Petiole transversely substrigate, basally coarser dorsally, finer ventrally. Tergite 3 with short strong rugulosities basally, otherwise polished smooth, T4-7 and T8 basally very finely and transversely microaciculate, T8 apically weakly transversely alutaceous, T9 strongly alutaceous. Pygidial impression distinct, V-shaped, its sides sligthly concave, forming a collapsing triangle. Pygidhim basally fiat-triangular, apically convec, weakly emarginate. Colour. Dark brown, with reddish legs and petiole. Genal stripe yellowish, narrow, width uniform; detached fi’om ^ e margin dorsally only, barely reaching temple; ending very 6 r fi’om occipital carina. Reddish brown basal half of the antennae, &ons ventrally (at level of antennal foramina), mandibles, except black tips, palpi, basally on coronal tubncles and interoceOar carinae, most of legs, except dark brown on front and mid coxae and basal 0.5 of hmd coxa, hind fomur centrally on dorsum and sides, petiole; lighter on antennae and front and mid tarsi. Wing veins and ovipositor valves brown. D ia ^io sis. Scutellum sparsely foveolate, polished smooth between (Fig. 331). Frons strongly and distinctly areolate (Fig. 330). Anterior half of prostemum strongly obliquely strigate (Fig. 333). Pre- and semiannular strongly strigate (Figs 334,337). Vein 2Cua basal 0.2 tubular, otherwise nebulous, apical half bent towards apex of wmg. ^ e x 387 of tubular part o f vein 2+3M truncate changing suddenly to nebulous or spectraL Hind basharsomere short (btl/w 4.78). No whitish or yellowish basally on mid basharsus and mid tibia. Legs reddish. Rem arks. Similar to P. ff^anulatus, from which it can be distinguished by having scutellum polished smooth between sparse foveolation, mid basitarsus and correspondent tibia not pale-coloured basally, and distinct structure for veins 2Cua and 2+3M. Overall colour pattern and geographical distribution O^gs 418,421) also different M ale Unknown. Distributional and Seasonal Records. H olo^e from mid-eastern Queensland (Rg. 418), collected 26 April. Parastephandlus tanytarsus Aguiar n. sp. (Figs 252-271 (sp. 9), 338-347, 420) E tym ology. Gr. Uaiyo, stretched out long Gr. tarsis, tarsus. In reference to length o f hmd basitarsus. M aterial Examined. Holotype: ?, 57 Km S Norseman, WA, 32“38’S, I2M 2'E, 31 Dec 1985, G. Daniels, A. Daniels. OSUC2I027 (UQIC). Paratypes: 2 9 ?, I cf ÇklVMA, WAMP). 388 Fem ale iM ïg th , Mean 12.2 mm (range 11.8-12.7 mm). H ead, Flagellomeres normal. Frons areolate-rugose, transversely ventrally; frontal carina absent ORg- 339). Vertex (Fig. 338) transversely subrugose, more distinctly transverse along centro-longitudinal impression; posteriorly, including temple, transversely strigate, ending anteriorly on temple at genal stripe, continuing down posteriorly on gena as weak strigulation along occipital carina, until almost ventrally. Centro-longitudinal impression weak, distinct under tangential light. Genal angle moderately prominent. Gena polished smooth, except centrally near occipital carina. Occipital carina very narrow, centro-dorsally marginated. Posterior margin of head concave in dorsal view. Occiput polished smooth. Post-genal bridge distinctly alutaceous. Hypostomal carina apex, below mandible base, weakly rugulose. Thorax. Prostemum rugose, except conspicuously alutaceous on apec (Fig. 345). Pronotum ^ g s 340,342,345): colo with 4-5 carinae, which are oblique anteriorly, transverse posteriorly, contiguous dorsally; pronotal fold collapsed and fused with colo medially, originating two lateral depressions instead o f single large excavated area; preannular and semiannular strongly rugose; longitudinally carinate on pronotal lobe and depression m front of it; femoral impression shallow, more or less hidden by general pronotal sculpture; ventral area as for pre- and semiannular. Mesoscutum (Fig. 340) areolate-mgose; weakly alutaceous narrowly on anterior margin. Axilla (Fig. 344) areolate-rugose, finer than on mesonotum, much coarser than on scutellum. Scutellum (Figs 343-344) areolate-rugose centrally; some large foveolae laterally; lateral ap«c 389 punctate; sulcus along postero-lateral margin crenulate; posterior margin polished, weakly alutaceous. Mesepistemum areolate-rugose finer than on axilla, and punctate or fbveolate, producing densely and richly sculptured sur&ce; dorsal part polished smooth on posterior 0.6; overall sculpture getting finer ventrally, to densely punctate on ventral part (Fig. 346); polished smooth in narrow area around discrimen, which is foveolate (Fig. 346). Propodeum ^ g . 343) strongly areolate-rugose, with some sub-circular areolae near flank, which is distinctly alutaceous. Parapetiolar fovea indicated by larger and more transverse areolae, otherwise inconspicuous. Spiracular groove ill-defined, indicated by distinct spiracular carina, ending near, but not at flank. Metapleuron polished smooth on dorsal part; centrally strongly areolate-rugose anteriorly, weakly posteriorly; ventrally areolate- rugose as in antero-central region; post-foveolar area transversely carinate; interfoveolar area transversely carinate, firom strongly anteriorly to very weakly posteriorly. Ventral side of fourth tarsomere o f front and mid legs prolonged anterioiiy in a process about as long as apical tarsomere. Hind coxa areolate-rugose, punctate in some areas (similar to mesepistemum), dorso-centrally on lateral side a small, slightly concave area of much finer and more transverse sculpture; apically transversely strigate. Hind femur completely and finely alutaceous, shnflarto alutaceous on lateral side of dilated part of tibia (Fig. 347). Hind tibia very finely alutaceous longitudinally on constricted part; dilated part finely alutaceous (but coarser than on constricted part) ^ g . 347), obliquely carinate ventrally. Front wing: vein 2Cua tubular on basal 0.4-0.5 ^ g . 341). Pterostigma elongate, more or less pointed (Fig. 341). 390 Abdomen. Petiole transversely areolate-rugose, coarse basally to very finely apicalfy; apex polished smooth; ventral sculpture similar, but finer. Tergite 3 on extreme base rugulose and microreticulate, centrally microcoUicuIate (visible at SOx), its apical 0.3 and all T4-8 transversely microaciculate T9 strongly alutaceous. Pygidial impression subtriangular. Pygidium flat basally, with a short centro-longitudinal elevation, changing to convex and slightly emargmate apically. Colour. Rufous with reddish black areas, bind femur and abdomen black. Brownish basally on antennae and mandibles. Genal stripe yellowish, much narrower to almost absent centrally, reaching no flirther than genal angle, ending far from occipital carina. Head otherwise reddish black. Rufous on pronotum, prostemum, mesepistemum, including ventral part, mesonotum and axila centrally, scutellum laterally, propodeum, metapleuron, petiole basally and apically, most of front and mid femora and tibiae, hind coxae ventrally, hind trochanter, hind tarsi, and hind tibia except for black centro-Iaterally and narrow longitudinal stripe mesally. Front tarsi reddish brown. Mid basitarsus white on 0.6 basal. Otherwise thorax and abdomen black or reddish black. Wing membrane slightly in&scat^ stronger on cells R, ICu, IM, and 2Cu; veins brown, pterostigma and subcostal area dark brown. Ovipositor valves from brown basally to black on apex. D iagnosis. Pronotum (Rgs 340, 342), prostemum (Fig. 345), and propodeum (Fig. 343) strongly rugose. Ventral side o f fourth tarsomere of front and mid legs strongly projected anteriorly, almost reaching apec o f apical tarsomere. Vein 2Cua straight, at 90" with ICu; tubular on basal 0.4, then nebulous, forming large spot (Rg. 341). Knd femur 391 very stout, robust, very finely and conspicuously alutaceous, similar to alutaceous on dilated part of hind tibia ^ g . 347). Ventral area of mesepistemum densely punctate and rugulose (Fig. 34S). Frontal carina absent (Fig. 339). Remarks, Similar to P. granukOus, fi’om which it can be distinguished especially by its fine, not transversely strigate microsculpture on petiole, longer hind basitarsus, distinct colour pattern, and structure of vein 2Cua. Also somewhat similar toP, acUazetos n. sp., but distinguished by having prostemum sculpture coarse, hind coxa and femur relatively shorter (sp. 9 on Figs 256,258, and table 3.7), hmd femur more finely alutaceous than on dilated part of correspondent tibia (Fig. 347), metapleuron centrally triangularly prominent, polished smooth dorsally, compressed part of hind tibia shorter or about as long as dilated part, vertex sculpture centrally in two distinct patterns (tranverse centrally, irregular antero-laterally), and différent colour pattem. M ale Similar to female, except fi>r the fisllowing. Frontal carina developed, distinct Daric ferrogineous to reddish dorsally on posterior margin of semiannular, scutellum apically, metanotum, petiole, accept dorso-centrally, hind coxa and femur apically, all tibia (except dark brown centrally on dilated part) and tarsi. Prostemum basally weakly sculptured to polished smooth. Ventral part of mesepistemum alutaceous. Vein 2Cua mostly nd)ulous and linear, not forming a spot 392 R em arks ^dale). Microsculpture on the prostemum, mesostemum, hind femur, pronotum and 13 similar to females o f P. granuIatuSy but sculpture on petiole and scutellum, and overall colour patem, which are more complex and more stable characters, match better the condition observed fer females of the present species. Distributional and Seasonal Records. Four specimens from the Southwest (Fig. 420), collected m January (day unknown), 14 September and 31 December. Associations. Adult OSUC2I029 and pre-adult OSCU2I02S females collected fiom a buprestid tunnel in Casuarim campestris (Casuarinaceae), in September 14. Parast^kanellus granulatus (Elliott) ^ g s 252-271 (sp. 10), 348-354, 421) HemistepfumusgranulatusEliiotty 1919b: 130; 1922:762,771. Parastephanellus granulatus.- Aguiar, 1998:428. Etym ology. L. granum^ gram. In reference to overall body sculpture. Material Examined. Holotype: 9 HemistephamtsffranulatusPXlioVL, Australia: SwanR., no date. OSUC20559 (BMNIQ. Other specnnensexamined. 6 99 (ANIC, MVMA, ÜCRC, WAMP). Fem ale Length. Mean 15.6 mm (range 12.9-18.4 mm). 393 H ead. Flagellomeres suddenly shorter and thicker beyond 7th. Frons areolate- rugose, finer near ^ e s in one specimen; fi-ontal carina absent Vertex ^ ig . 348) coarsely rugose finer posteriorly; transversely strigulate on temples, sometimes also centrally near occipital carina. Centro-longitudinal impression absent Genal angle moderately prominent Gena polished smooth. Occipital carina very narrow. Posterior margin of head concave in dorsal view. Ocdput polished smooth. Post-genal bridge distinctly alutaceous. Hypostomal carina apex, below mandible bas^ longitudinally strigate. Thorax. Prostemum (Fig. 352) entirely rugulose or rugose, except alutaceous apically and sometimes sub-basaUy. Pronotum (Figs 349, 351): colo arcuate-strigate; strigation dorsally interrupted, running towards, some entering, pronotal fold, which is collapsed and fused with colo medially; preannular transversely strigate, sometimes finely; sometimes transversely alutaceous apically; semiaimular strongly transversely strigate, or transversely carinate; laterally rugose at least dorsally; pronotal lobe obliquely strigate; dorso-posterior margin polished smooth; femoral impression indistinct; ventral area longitudinally strigate. Mesoscutum areolate-rugose with some foveolae anteriorly, anterior margin finely alutaceous. Axilla (Fig. 354, part) areolate-rugose, stronger basally (finer than on mesoscutum), finer apico-laterally. Scutellum (Fig. 354) areolate-rugose and punctate. Mesepistemum (Fig. 353) punctate-reticulate postero-ventrally, otherwise finely areolate-rugose; surfirce inside and between rugosities irregular, producing very mtrmoite overall pattern; this microsculpture extending partially to dorsal part, which is polished smooth dorsally ventral part transversely alutaceous, some punctulations; rarely oblique strigate-punctate (one specimen); discrimen simple or finely foveolate. Propodeum (Fig. 394 354) densely areolate-rugose» coarser centro-Iongitudinally, finer laterally (as mesepistemum); flank alutaceous. Parapetiolar fovea deep posteriorly, disappearing anteriorly, with a few transverse carinae. Spiracular groove weak, narrow, weakly crenulat^ spiracular carina disappearing midway between spiracle and crenulate sulcus. Meü^leuron areolate-rugose in a narrow, triangular-elongate central area; dorsally partialfy polished smooth, some transverse strigation and microreticulation posteriorly; transversely strigate along spiracular carina. Post-foveolar area transversely strigate. Interfoveolar area with some crenulae anteriorly, otherwise polished smooth. Ventral side o f fourth tarsomere of fi’ont and mid legs normal, not prolonged anteriorly. Hind coxa laterally on basal 0.2-0.3 areolate-rugose, sometimes strongly; dorso-centrally finely transversely rugulose on small, slightly concave area; ventrally on center and anteriorly transversely strigate, more sparsely anteriorly, hind coxa otherwise transversely strigate, strigations stronger on base of hairs. Hind femur densely and distinctly alutaceous, weaker basally. Hind tibia alutaceous finer than on bind femur; dilated part obliquely strigate ventrally. Front wing; vein 2Cua tubular on basal 0.1-0.2 (Fig. 350). Pterostigma elongate, more or less pointed. Abdomen. Petiole transversely strigate; rarely finely transversely rugulose; sculpture more irregular basally, broadly polished smooth on apex; ventral sculpture similar, but coarser. Tergite 3 on extreme base rugulose, then polished smooth until middle, its apical half and all T4-8 very finely transversely microaciculate, T9 foiely and conspicuously alutaceous. Pygidial impression acute-triangular. Pygidium large, convec. 395 Colour. Black. Front and mid legs rufescent, tarsi brownish or yellowish. Genal stripe yellowish, much narrower to almost absent centrally, reaching no further than genal angl^ ending far from occipital carina. Extreme base of mid tibia, basal half of mid basharsus, apex of hind coxa, and hind basitarsus testaceous or yellowish. Hind trochanter and tibia mesally rufescent to ferrugineous. Posterior margin o f semiannular and basal wing sclerites brownish or yellowish. Wing membrane slightly brownish infhscate; veins daric brown. Ovipositor valves daric brown. D iagnosis. Scutellum duO, areolate-rugose and punctate (Fig. 354). Prostemum rugulose (Fig. 352). Hind basitarsus short (btl/w 4.82-5.05). Petiole distinclty transversely strigate. Vein 2Cua straight all over, at 90” with ICu, tubular on basal 0.2 only, then nebulous, but narrow, not forming a large spot apically (Fig. 350). Vein 2+3M gradually tappering and becoming nebulous apically. Frontal carina absent. Remarks. Similar to P. tanytarsus n. sp., from which it can be distinguished especially by showing transversely strigate petiole, short basitarsus, and distinct colour pattern. Also similar to P. callidictyon n. sp., but distinguished by having scutellum dull, areolate-rugose and punctate and mid basharsus (sometimes also mid tibia) pale-coloured basaHy; overall colour pattem and geographical distribution (Figs 418,421) also different. Distinguished from both species by the structure of veins 2Cua and 2+3M. Microsculpture o f the prostemum, mesosteraum, hind femur, pronotum, and T3 on males ofP. tanytarsus n. sp. similar to females o f the present species, but males of P. tanytarsus have the same typical petiole and scutellum sculpture of the females. 396 M ale Unknown. Distributional and Seasonal Records. Soc specnnens from the Southwest ^ g . 421), four collected 7 March, 12 October, 22 November; other specimens without date or locality. Parastephanellus platyceratus Aguiar n. sp. (Figs 252-271 (sp. 11), 355-358,422,436) Etymology. Gt.platys, broad, wide, flat; Gr. keratos, horn (antenna). In reference to the apical shape of antenna. Material Examined. H olo^e; ?, KalmgaPk., ‘12-9-1957’, C. P. McPhee. OSUC21049 (UQIC). Paratypes: 3 ??, 5 Female Length. Mean 6.9 mm (range 6.2-8.1 mm). H ead. Apical flagellomeres dorso-ventrally flattenned. Frons finely or strongly areolate-rugose, transversely carmate along ^ e margin; fluntal carina absent or distinct Vertex QRg. 355) behmd interocellar carinae and centrally irregularly rugose; laterally transversely carinate-rugose; posteriory transversely strigate centrally only; temple polished smooth. Centro-longitudmal impression absent Genal angle strongly prominent, but with no distinct protuberance. Gena polished smooth. Occipital carina very narrow, 397 wider laterally. Posterior margm of head concave in dorsal view. Occiput polished smooth. Post-genal bridge very weakly alutaceous or polished smooth. Hypostomal carina apex, below mandible base, very weakly alutaceous. Thorax. Prostemum with a few transverse rugulosities on posterior margin, apex alutaceous, otherwise polished smooth. Pronotum (Fig. 356): colo with about three complete carina^ which are oblique laterally, arcuate dorsally, polished smooth or alutaceous between; pronotal fold stout, distinctly excavated, not collapsed; preannular transversely substrigate; alutaceous apico-laterally; semiannular mostly polished smooth; some weak longitudinal carinae and punctules laterally on pre- and semiannular, polished smooth between; lateral posterior margin longitudinally carinate-rugose, including pronotal lobe; femoral impression shallow and faint, or indistinct; ventral area sub- alutaceous. Mesoscutum from rugose or fr)veate posteriorly to weakly alutaceous apically. Axilla (Fig. 357) from strongly rugose or sub-areolate-rugose near scutellum, to alutaceous on anterior margin. Scutellum (Fig. 357) laterally with some shallow lacunae, foveolae, and punctures, otherwise weakly alutaceous or from distinctly and finely alutaceous anteriorly to polished smooth posteriorly. Mesepistemum (Fig. 358, part) from rugulose anteriorly to rugulose-lacunose centrally; dorsal part broadly polished smooth; pectus densely and ventral part weakly and sparsely alutaceous; discrimen foveolate. Propodeum (Fig. 358, part) areolate-rugose; alutaceous narrowly along anterior margin, including anterioriy on flank. Parapetiolar fovea indicated by large areolae. Spiracular groove weak or indistmct; spiracular carma distinct, disappearing midway between spiracle and crenulate sulcus, or more complete almost reaching flank. Metapleuron (Fig. 398 358) mostfy polished smooth dorsally, accept somethnes some transverse carinae posteriorly; centrally areolate-rugos^ transversely carinate ventrally and on post-foveolar area. Interfoveolar area transversely carinate. Ventral side of fourth tarsomere offrent and mid legs normal, not prolonged anteriorly. Hind coxa transversely strigate; centro-dorsal area small, inconspicuous, very weakly depressed and weakly sculptured, or indistinct (probably more distinct in large specimens). Hind femur alutaceous, stronger dorso- posteriorly; almost polished smooth antero-ventraUy mesally. Hind tibia obliquely to longitudinally strong-alutaceous (similar to hind femur dorso-posteriorly) on both sides; dilated part obliquely strigate ventrally. Front wing: vein 2Cua tubular on basal 0.3-0.5. Pterostigma more or less blunt. Abdomen. Petiole transversely strigulate, coarser basally, broadly polished smooth apically; ventral sculpture similar, but finer and with some microreticulate between. Terghes 3-7 polished smooth, T8 polished smooth or weakly alutaceous, T9 alutaceous. Pygidial impression wide-triangular. Pygidium convex, emarginate. Colour. Brownish or dark brown. Genal stripe pale-yellow, wider dorsally, usually ending far from occipital carina, sometimes ending very close to h. Pale-yellow mandibles dorsally, wing sclerites, apoc of fiont tibia, base and apex of mid tibia, tip of hind coxa, basal 0.2 of hind tibia. Yellow on narrow longitudinal area on basal half of inner ^ e margin, around antennal foramen, clypeus. Femigmeous base of coronal tubercles. Head, prothorax, extreme base 13, petiole apex, antennae basally, most of front and mid legs, with tendency to brown or reddish brown; hind legs similar, but darker. Hind tibia with 399 some blackish marks, especially mesally on dilated part. Wing membrane hyaline, cells IM and 2Cu very lightly brownish; veins light-browiu Ovipositor valves yellowish brown or brown. D iagnosis, ^ ic a l flagellomeres slightly flattenned. Hnd tibia often with distinct light creamy colour on basal 0.2. Pronotum mostly polished smooth QFig. 356). Temple (Fig. 355) and T3-7 polished smooth. Scutellum centrally polished smooth or shinnmg alutaceous. Hind femur, at least mesally, polished smooth or distinctly more weakly alutaceous than dilated part of correspondent tibia. Vein 2+3M slightly longer than 2+3Rs. Small size. Rem arks. Similar to P. albigenuis n. sp. and P. strigosus n. sp. From P. albigenuis it can be distinguished by the diagnosis above and by having the following: Vextex irregularly rugose centrally, changing from transversely strigose laterally to posteriorly strigulose centro-Iongttudinally, this sculpture barely or not reaching occipital carina (Fig. 355); genal angle usually strongly prominent; frons usually narrowly yellow along eye margin and around antennal ferame. From P. strigosus it can be easily distinguished by having temple broadly polished smooth and hind tibia basal 0.2 often lightly creamy- coloured, sharply contrasting with remaining dark brown. M ale Sunilar to female, but morphological variability higher. Frons usually entirely yellow and/br reddish yellow; more rarely two yellow stripes along ^ e margin, as for females. Scutellum polished smooth or almost so. Semiannular, mesepistemum, and hind coxa laterally with weak sculpture or polished smooth areas. Petiole relatively lon^ 400 usualfy as long as, or longer than gaster. Temple polished smooth, vertec sculpture not reaching genal stripe, ending near or barely reaching occipital carina. Overall colour mostly dark brown, except gaster; one specimen from alcohol nearly entirely yellow; ^ ster apex always dark brown. Rem arks ^ ale). Very similar to males o ff. laevigatas n. sp., from which it can be distinguished basically by showing frons with vertical yellow stripe along each ^ e margin, or at least distinctly more lightly colored than in other areas of the head. Similar to some P. dameilicus; distinguished by absence of strong foveolae centrally on scutellum (similar to Figs 307 vs. 357), pronotum not reddish black, usually lightly coloured, and frons with yellow. Distributional and Seasonal Records. Mid- and North-northeast (Fig. 422); flying at least from April to October, but no specimens recorded from hottest months (Fig. 436). Associations. Holotype reared from ‘Tea-Tree Bark' [probably A/efo/eirca altemifolia (Maiden & Betche) Cheel, Myrtaceae]. Parasteplumenus strigosus Aguiar n. sp. (Figs 252-271 (sp. 12). 359-365.423,435) E tym ology. L. striga^ ridge. Di reference to strigation on femoral impression. Material Examined. Holotype: Hectra S. F., c. 25 Km S Bundaber^ Qld., 1 Dec 1976, H Frauca. OSUC21051 (ANIC). Paratypes: 7 ? Î , 5 cfd* (AMSA, ANIC, BMNH, SAMA. UQIC). 401 Female L e n g th . Mean 12.4 ram (range 7.4-16.4 ram). H ead. Flagellomeres normaL Frons transversely or obliquely scabriculous; frontal carina distract, but weak and usually only partially developed. Vertex (Rgs 359,362) from transvosely undulate-rugose anteriorly to transversely strigate posteriorly; sculpture extending to temple and distinctly reaching genal stripe; straight transversely rugose on centro-longitudinal impression. Centro-longitudinal impression weak, distinct under tangential light ^ g . 362). Genal angle strongly prominent, with distinct protuberance. Gena usually polished smooth, sometimes with oblique rugulosities ventrally. Occipital carrâa very narrow, wider laterally. Posterior margm of head concave in dorsal view. Occiput polished smooth. Post-genal bridge weakly alutaceous, stronger apically. Hypostomal carina apex, below mandible base, weakly longitudinally rugulose. Thorax. Prostemum transversely rugulose on apical 0.3 and basal 0.1, otherwise sparsely pQose or glabrous and polished smooth. Pronotum (Figs 359-360,363): colo from arcuate-strigulate (small specnnens) to strongly arcuate-carinate (large specimens); posterior carinae entering pronotal fold, which is stout, deeply excavated, not or only weakly collapsed or emarginate; semiannular dorsally transversely carinate centrally, frner iq)ically and basally; some frne alutaceous on apical margin; preannular and semiannular laterally obliquely rugulose dorsally; centrally frnely alutaceous to foiely obliquely rugulose; pronotal lobe, and concave area in front of it, strongly carinate-rugose longhudmally; femoral hnpression deep, narrow, strongly transversely carinate along all its extension; ventral area frnely longitudinally alutaceous, with a fow well marked 402 longitudinal carinae. Mesoscutum mostly strongly transversely rugose, narrow anterior margin finely alutaceous. Axilla ^ g . 364) strongly rugose or areolate-rugose. Scutellum (Fig. 364) transversely rugose centrally, much finer apically; laterally areolate-rugose; posterior margin finely alutaceous. Mesepistemum rugulose and microreticulate, extending to ventral 0.5 of dorsal part, otherwise polished smooth; ventral part obliquely finely alutaceous, except polished smooth near discrünen, which is simple. Propodeum strongly transversely rugose or areolate-rugose; flank finely alutaceous, sometimes also with a few punctures. Parapetiolar fovea weak, shallow, densely transversely carinate, carinae contiguous with propodeum sculpture. Spiracular groove indicated by series o f aligned areolae or, in smaller specimens, weakly defined areolae, not reaching flank; spiracular carina appearing as part of the areolation. Metapleuron (Fig. 361) strongly and densely transversely carmate dorsally, centrally areolate-rugose, ventrally weakly transversely carinate. Post-fbveolar and interfoveolar areas transversely carinate-rugose. Ventral side of fourth tarsomere of fi'ont and mid legs normal, not prolonged anteriorly. Hind coxa transversely strigate, more irregularly or finely dorso-centrally on lateral side; mesally transversely strigate (stronger basally) and punctulate (sparse apically); dorso-central area mdistmct Hind fomur finely alutaceous, which laterally and dorsally is composed of sub- circular units, thus similar to microreticulate; mesally with elongate units, thus more tipically alutaceous. Hind tibia finely obliquely alutaceous laterally, finely alutaceous mesally (finer than on bind fomur); dilated part obliquely strigate ventrally. Front wing OHg. 365):Vem 2Cua tubular on basal 0.3. Pterostigma apex more or less acute. 403 Abdom en, Petiole transversely strigate, more rarely finely transversely strigulate; apex polished smooth; ventral sculpture similar, sparser or weaker. Tergite 3 on extreme base rugulose and finely alutaceous,. otherwise polished smooth, T4-8 dull and transversely microaciculate, T9 alutaceous. Pygidial impression shallow, widely triangular. Pygidium laig^ convex. Colour, Dark brown or yellowish brown. Genal stripe pale-yellow, crossing templ^ where it is wider, reaching occipital carina (Figs 359,362). Yellow or yellowish on anterior and lateral tubercles (except black tips), fixons along ^ e margin, until level of lateral tubercles, and basally at least around antennal forame, and mandible (accept black tip); yellow sometimes replaced by orange or dark reddish brown in greasy or not well preserved specimens. Pronotum (I) brown, light-brown or light-reddish brown, with dark brown dorsally, anteriorly on preannular and posteriorly on colo (dark brown at least on anterior margin o f preannular), or (2) totally dark brown, with some reddish brown on semiannular. Scutellum, hind coxa mesally, axilla (sometimes), and posterior half of propodeum (1) reddish brown or (2) dark brown or black. Petiole basally or apically and T3 basally reddish brown. Pale-yellow or yellowish on apical O.I and basal 0.2 of mid tibia, basal 0.5 o f mid basitarsus, apex of hind coxa, and tegula. Hind tibia (I) pale-yellow on basal 0.3-0.4 of compressed part, then shortly black or blackish, otherwise hind tibia ^eluding dilated part) and tarsi light brown, or (2) hind tibia and tarsi reddish brown, lighter basally on compressed part, mostly black laterally on dilated part. Front legs brown, sometimes darker on femur. Nfid legs brown to reddish brown, except as noted 404 above. Wing membrane lightly yellow infiiscate; veins yellowish brown or brown, pterostigma brown marginally, light- or pale-brown centrally, never entirely opaque. Ovipositor valves from yellowish brown or brown basally to daric brown apically. D iagnosis. Anterior and lateral tubercles yellow anteriorly and/or basally; frons with yellow stripe along eye margin, which becomes wider ventrally, originating a subtriangular area on each side above and around antennal foramina, or stripe interrupted ventrally, with yellow spot dorsad and mesad of antennal foramina. Femoral impression conspicuously transversely carinate QFig. 363). Semiannular finely sculptured laterally, coarser latero-dorsally (Fig. 363). Pygidial impression shallow, pygidium large, convex. R em arks. Similar to P. platyceratus n. sp., from which it can be separated, in addition to the diagnosis above, by having stronger sculpture on semiannular, alutaceous of hmd femur as strong as on dilated part of correspondent tibia, transverse strigation on vertex covering temple and reaching genal stripe, and antennae apically normal, not flattened apically. Also similar to P. trifasciatus n. sp., from which it can be immediately distinguished by lacking a centro-longitudinal yellow stripe on frons, and by showing smaller values for Pt/T31. M ale Vertec, includihg temple until genal stripe, transversely strigate, more undulate or irregular anteriorly; finer and more straight posteriorly. Pronotal fold deeply «ccavated, not collapsed. Pronotum and scutellum sculpture Aiding rapidly with decrease m size of specimen: large specimens as fr)r females; small ones with semiannular and scutellum weakly alutaceous, and femoral impression polished smooth. Metapleuron dorsally always 405 transversefy substrigate usually conspicuous^. Colour pattern characteristic: Frons entirely sulfur yellow to pale yellow; genal stripe wide dorsally, reaching occipital carma, same colour as frons. Head, prothorax, legs (except hind coxa), and gaster partially, ligfatfy brownish or yellow, sharply contrasting with dark brown or blackish meso- and metathorax, propodeum, petiole hind coxe and sometimes also hind femur and tibia. Mesothorax dorsally, propodeum near petiolar fr)rame petiole basally, and sometimes coxa dorsally, dark reddish brown. Hind tibia and tarsi yellowish brown/brownish yellow; somethnes dilated part dark brown. Small specimens tend to mostly lightly coloured; large ones with dark brown predominating. Rem arks ^ale). Similar to P. trifasciatus n. sp.; distinguished by overall body colour pattern, coarse microsculpture of scutellum, fine microsculpture laterally on pronotum, and femoral impression usually at least anteriorly with transverse carinae. Distributional and Seasonal Records, North-northeast (Fig. 423); flying from October through January, but apparently more abundant in December (Fig. 435). Parastephanellus irifascitOus Aguiar n. sp. (Figs 252-271 (sp. 13), 366-373,424,434) Etym ology. L. tres^ three; L . fascia^ stripe. In reference to stripes of colour on front Material Examined. Holotype: ¥, Bundaberg Cr., Bundaberg, Qld., 13 Jan 1975, H. Frauca. OSUC21051 (ANIC). Paratypes: 9 ? ?, 6 406 Fem ale L e n g th . Mean 10.8 mm (range 9.1-14.4 mm). H ead. Flagellomeres slightly clavate. Frons (Fig. 368) areolate-rugose; parallel oblique rugose near eye margin; Aontal carina absent^ rarely very weakly developed. Vertex ^ g . 366) from finely transversely rugose anterioriy to transversely strigulate posterioriy and on temples, reaching occipital carina; laterally some strigations curving anteriorly towards ^ e margin; the ones near occipital carina curving postero-ventrally; temple strigulate; all strigation ending at genal stripe. Centro-longitudinal impression absent. Genal angle strongly prominent, sometimes with distinct protuberance. Gena polished smooth. Occipital carina very narrow laterally, marginated or absent dorsally. Posterior margin of head concave in dorsal view. Occiput polished smooth. Post-genal bridge weakly longitudinally alutaceous, stronger apically. Hypostomal carina apex, below mandible base, finely alutaceous in small specimens, finely rugose in large ones. Thorax. Prostemum alutaceous apically, transversely or obliquely rugulose basally, otherwise polished smooth. Pronotum (Figs 367,369); colo usually strongly arcuate- carinate, with alutaceous^ragulose between; rarely only arcuate-strigulate; pronotal fold stout, partially collapsed and moderately emarginate; preannular and semiannular densely transversely rugulose or transversely strigate dorsally, finer near apex of pronotal fold, more irregular laterally; transversely carinate on pronotal lobe and concave area in front of it; femoral impression sometimes distinct, polished smooth or longitudinally carinate- rugose; ventral area varying from entirely alutaceous to densely longitudinally rugulose. Mesoscutum conspicuously alutaceous on anterior 0.3, otherwise subtransversely rugose; 407 median sulcus and notaulus indicated by punctures. Axilla ^ g s 370-371) microreticulate with rugulosities. Scutellum ^igs 370-371) transversely alutaceous with microreticulations, basally transversely nigulose, punctate laterally, punctures sometimes weak. Mesepistemum (Figs 372-373) densely microreticulate, strongly rugulose anteriorly and dorsally; dorsal part polished smooth, centrally partially with same general sculpture o f mesepistemum; ventral part finely transversely alutaceous; discrimen distinctly microfoveolate. Propodeum (Fig. 373, part) areolate-rugose, sometimes coarser posteriorly or anteriorly; usually also densely microreticulate or alutaceous, especially laterally; flank very finely transversely alutaceous. Parapetiolar fovea indicated by large polished smooth, weakly defined depressions. Spiracular groove indistinct; spiracular carina distinct, receiving several transverse carinae or rugae firom both sides, ending near or at flank. Metapleuron ^ g . 373) densely shallow puncticulate or microreticulate ^ess abundant in small specimens); centrally also areolate-rugose; dorsally densely transversely carinate along spiracular carina, meeting areolate pattem anteriorly, polished smooth between it and central part, or totally transversely carinate. Post-fbveolar area transversely carinate or mgose. Interfoveolar area transversely carinate, sometimes weakly. Ventral side of fourth tarsomere of front and mid legs normal, not prolonged anteriorly. Hind coxa laterally rugose on basal 0.2; microreticulate in somewhat depressed area centro-dorsally; transversely strigate ventro-apically; mesally transversely strigate, also densely punctulate basally to sparsely apically. Hind femur strongly alutaceous on both sides, weakly on 408 extreme base. Hmd tibia obliquely alutaceous on constricted part; dilated part alutaceous, obliquely carinate ventrally. Front wing: vein 2Cua tubular on basal 0.2. Pterostigma elongate apex more or less acute. Abdom en. Petiole transversely substrigulate, rugose basally, broadly polished smooth apex; ventral sculpture similar. Tergite 3 polished smooth, T4-7 conspicuously transversely microadculate (dull), T8-9 alutaceous, strongest on T9. Pygidial impression triangular, usually deepfy notched basally (notch sometimes covered by posterior margin of T8). Pygidium narrow, convex, emarginate. C olour. Brownish or black. Frons yellow basally, with three vertical yellow stripes, two lateral, along eye margin, ending at lateral tubercles, one central, ending at frontal tubercle; stripes narrow, or very wide and almost fusing with one another, in which case frons is almost entirely yellow. Genal stripe crossing temple, ending near occipital carina. Yellow mandible (except black apex), wing sclerites, mid tibia basally and apically, basal half of mid basitarsus, hind coxa tip. Brownish basal antenna! segments, dypeus, palpi, posterior margin of semiaimular, front and mid legs (except black tip of apical tarsomeres), hind leg on trochanter, apex to most of femur, tibia, except sometimes dark brown centrally, and tarsi, accept black tip of apical tarsomere. Reddish to brownish base of coronal tubercles and interoceOar carinae, semiannular laterally, prostemum somethnes. Ocelli sometimes pink or pinkish. Wing membrane lightly yellow înfûscate; veins yellowish brown or brown, pterostigma brown marginally, light- or pale-brown centrally, never entfrely opaque; apex of vein Sc+R. and pterostigma basally sometimes black. Ovipositor valves brown or dark brown. 409 D iagnosis. Three vertical stripes and basal part of fions yellow ÇFig. 368). Vertex sculpture transverse never concentric, undulate anteriorly, straight strigulate posteriorly CFig. 366). Prostemum alutaceous apically and oblique strigations basally. Hind coxa long and slender (sp. 13 on Figs 256>2S7 and table 3.7). Scutellum transversely alutaceous centrally OFig. 371). Genal angle prominent ^ig . 366), usually forming weak protuberance. Ocelli sometimes pink or pinkish. Metapleuron with dense shallow punctulation or microreticulation all over, inclusive on and among coarser sculpture (Fig. 373). Rem arks. Close to P. strigatus n. sp. and P. platyceratus n. sp., from which it can be easily distinguished by having central yellow stripe on frons (although sometimes 6int) and distinct semiannular sculpture. Some specimens entirely light-brown, making yellow stripes on frons difScult to recognize. Because of this problem, the current species can be mistaken for someP. damellicus, but there are some important differences; P. trifasciatus n. sp. scutellum finely alutaceous, mesepistemum more strongly and characteristicaly microreticulate (compare Figs 66,126), vertec sculpture regularly transverse, and ventral area of pronotum with unbroken longitudinal strigation. M ale Similar to female, except following. Frons almost always entirely yellow, rarely with three longitudmal yellow stripes. Genal stripe wider on temple, ending very close to or at occipital carina. Body mostly lightly coloured: head, pronotum and legs yellowish brown; sometimes partially dark brown centrally on hind tibia; thorax yellowish or reddish brown; petiole and gaster often of same colour as thorax; apical 2-3 segments of abdomen 410 much darker than the others. Vertex finely transversely undulate-strigat^ strigation rarely as straight and uniform as in P. isogfyptos n. sp. Pronotal fold deeply excavated, not collapsed. Scutellum distinctly, sometimes coarsely, alutaceous. Dorsal part of metapleuron always with fine, usually transverse microsculpture. R em arks (Male). Similar to males ofP. strigosus n. sp.; distmguished by overall body colour pattern, fine microsculpture of scutellum, strigation on pronotum, femoral impression polished smooth or with longitudinal sculpture. Distributional and Seasonal Records. Mostly North-northeast, but one record fi’om western Australia (Pig. 424); flying activity recorded for all months from September though April (Fig. 434). Parastephanellus squamaius Aguiar n. sp. (Figs 252-271 (sp. 14), 374-381,426,438) E tym ology. L. squama^ scale. In reference to scale-like projections on back of vertex. Material Examined. Holotype: $, 10 Km SW Musselbrook Mining Camp, Qld., I8.39S, 138.12E GPS, 12 May 1995, at light, T. Weir. OSUC20959 (ANIC). Paratypes: 7 Î Ç (AMSA, ANIC, NSWA, UQIQ. Female Length. Mean 6.3 mm (range 2.6-7 6 mm). 411 H ead, Head antero-posterîoily, thorax dorso-ventralfy, strongly flattened (Fig. 381). Basal flageilomeres weakly clavate, otherwise normal. Frons areolate-rugulose; Êontal carina usually absent (distinct m one specimen). Vertex ^ g s 374, 376) dorsally transversely or arcuately rugos^ more arcuate laterally; posteriorly (flattened part) becoming transverse^ strigate or transversely strigulat^ usually some fine alutaceous between; this sculpture reaching occipital carina broadly; pair of scale-like projections laterally, on dorso-posterior angle; second pair, stronger, ventro-laterally. Centro- longitudinal impression distinct, but weak. Genal angle not prominent. Gena polished smooth. Occipital carina dorsally straight, transverse, weak; scale-like protuberance on each side, at begmning of lateral portion, where it is stouter, arcuate. Posterior margin of head straight in dorsal view. Occiput alutaceous; sometimes also rugulose dorsally. Post- genal bridge longitudinally alutaceous on specimens without pronotal fold (could not be observed on others). Hypostomal carina apex, below mandible base, finely longitudinally alutaceous; sometimes also rugulose. Thorax. Prostemum transversely alutaceous, stronger apically. Femoral impression mdistmct. Pronotum dimorphic: (Q Specimens with distinct pronotal fold ^ig s 376-378, 379): colo more sparsely arcuate-strigate, polished smooth or weakly alutaceous between; strigulation contiguous dorso-anteriorly; interrupted and entering pronotal fold posteriorly; a centro-posterior polished smooth depression, also entering pronotal fold, which is strongly flattened, but distinct; preannular sub-longitudinally, irregulariy, or transversely finely rugose; semiannular finety rugose, transverse on posterior margin (sub- concentric laterally in one specimen; pronotum rugose dorsally and laterally in one 412 spedmen). (H) Specimens without pronotal fold (Fig. 375): colo from arcuate-strigate anteriorly to transversely rugose centrally; preannular and semiannular dorsally arcuate- rugose; preannular laterally transversely rugose longitudinally rugose on ventral area, especially below semiannular; semiarmular laterally and latero-dorsally with some alutaceous and rugulose, overall sculpture arcuate sub-concentric. Both types: pronotal lobe lonptudinally carinate. Mesoscutum CFig. 378) coarsely rugose; median sulcus and notaulus defined by foveolae; sometimes anteriorly alutaceous. Axilla ^ g s 378,380) sub- arcuately and strongly alutaceous or microreticulate; lacunose antero-mesally on type I specimens ^ ig . 378). Scutellum ^ g s 378, 134) concentrically and strongly alutaceous or microreticulate; faintly alutaceous on posterior margin; punctate anteriorly and laterally on type I specimens (Fig. 378). Mesepistemum microreticulate, usually with some strong alutaceous and punctations; anteriorly rugulose; centro-dorsally weakly alutaceous; dorsal part alutaceous anteriorly, polished smooth posteriorly; ventral part finely transversely alutaceous; discrimen foveolate. Propodeum (Fig. 380) alutaceous; centro-Iongitudinally strongly rugose or areolate-rugose and punctate; areolae usually transverse anteriorly, elongate posterioriy; laterally puncticulate; flank sculpture finer. Parapetiolar fovea transversely carinate. Spiracular groove merging with overall propodeal sculpture; indicated by weak spiracular carma, which is distinct on basal G.5-0.7, and usually also by series of small aligned areolae or punctures. Metapleuron dimorphic: (1) Centrally subtriangularly prominent, areolate-rugose; dorsally weakly transversely rugulose or transversely alutaceous; ventrally strongly microreticulate; post-foveolar area microreticulate or transversely strigulate; (IQ Centrally not prominent (Fig. 380), changmg 413 from weakly areolate-rugose posteriorly to punctate anteriorly; dorsally polished smooth, scrobiculate along spiracular carina; ventrally microreticulate or as for post-foveolar area, which is weakly or strongly transversely carinate. Both ^ e s : foterfoveolar area very narrow, from usually polished smooth to sometimes transversely carinate. Ventral side of fourth tarsomere offrent (usually) and mid (always) legs prolonged anteriorly, almost reaching apmc of apical tarsomere. (End coxa rugulose basally; weakly concave dorso- central area with much finer sculpture; mesaHy microreticulate and finely rugose or alutaceous centro-basally, coxa otherwise transversely strigate. Hind femur mostly strongly transversely alutaceous, finer basally. Hind tibia longitudinally alutaceous, much finer than alutaceous on hind femur; dilated part obliquely carinate ventrally. Front wing: vein 2Cua tubular basal 0 J-0.6. Pterostigma sub-elongate, more or less blunt. Abdom en. Petiole basally rugulose, otherwise from weakly or finely rugose to finely transversely rugulose; ventral sculpture similar. Tergite 3 basally weakly rugulose, otherwise polished smooth or faintly longitudinally alutaceous, T4-7 weakly alutaceous, even weaker on apical 0.3-0.S, T8-9 alutaceous, strongest on T9. Pygidial impression very deep, V-shaped. Pygidium convex. C olour. Black, with abundant reddish or brownish on legs. Genal stripe narrow, wory, ending on latero-ventral scale-like projection, far from occipital carina. Reddish antennae basally, mandibles, except black tip, base o f head tubercles and ocellar carmae, gena sometimes, pronotum dorsally ^art, expecially apex and posterior margin), scutellum sometimes laterally or on posterior margin, propodeum near propodeal foramen, petiole, at l«tst apically, T3 basally. Legs reddish or brownish, «ccept dark brown or black on 414 most of hmd leg, coxa, femur, and laterally on dilated part of tibia, except apex. Brownish yellow apices of fi*ont and mid tibiae, basal half of mid basitarsus, apec of hind coxa. Posterior margin o f tergites sometimes narrowly yellowsOi or reddish brown, much clearer than, and sharply contrasting with their predominantly daric colouration. Wing membrane hyaline or lightly infbscate; strongly infiiscate cell 2Cu, especially near 'knob' of vein 2Cua; infuscate partial and less strong on cell Rs; veins daric brown. Ovipositor sheats dark brown. D iagnosis. Temple very narrow, head flattened antero-posteriorly ÇFigs 374, 381). Thorax conspicuously flattened dorso-ventrally (Fig. 381). Pronotum distinctly rugose (Fig. 377). Vertex with scale-like projections laterally (Fig. 376). Occipital carina dorsally straight, transverse, bearing scale-like projection dorso-laterally on each side (Hg. 376). Scutellum alutaceous in circular pattern QFigs 378,380); axilla alutaceous in semi-circular pattern (Figs 378,380). Pronotal fold present, flat, shallow (Figs 377, 379), or absent ^ g . 375). Petiole very short (Gsl/Ptl 1.51-1.98, usually >1.60; see sp. 14 on Fig. 264 and table 3.9). Rem arks. Distinguished from all other Australian species by the strongly flattened head and thorax. General sculpture, size, and colour, somethnes simflar to P. rufoomatus, from which it can be distinguished by the diagnosis above. The flattened body, however, nuiy perhaps represent a post-embrionic feature. The soft body of the larval/pupal stages could acquire a dorso-ventrally flatenned body by being 'molded' hi a tranversely oval gaiety o f a larval host, as is the cas^ for example, with the Buprestidae. 415 A fit>nto-posterioriy flattenned head could result if it is kept in an opisthognathous position inside the gallery. Ifthis possibility is true, species such as A ntfoomatus, to AA^ichf. squam atus n. sp. is similar in some ways, would be a likely candidate for conspec^ status. Type n specimens (OSUC20962, at NSWA, OSUC20963, at UQIC, and OSUC20964, at ANIC) have distinctive characters, and may perhaps belong to a different species (Table 3.4). The presence or absence of pronotal fold, for example, is usually enough for distinguishing species in most Stephanidae, but it is not known to which extent the strongly flattened body of the current species may have influenced the development of this structure. Nonetheless, the current state of knowledge of this species is still insufficient to allow a definitive conclusion. MbiphoftT»! Mbrphotype II 1. Head flattened posterior^ only 1. Head completely flat antero-posterioriy 2. Pronotal fold present 2. Pronotal fold absent 3. Pronotum sometims rugose 3. Pronotum strigate 4. Metapleuron central^ areolate-rugose 4. Met^Ieuron omitralty weakly areolate-rugose, and partly punctate 5. Met^leuronventralfy strongly punctate-areolate S. Metapleuron ventrally partially microreticulate or transversely carinate 6. Axilla lacunose antero-mesally 6. Axilla not lacunose 7. Scutellum partially punctate orfoveate______7. Scutellum without punctations or fbveae______ Table 3.4. Main deferences between morphotypes of P. squam atus n. sp. 416 M ale Unknown. Distributional and Seasonal Records. North central, and Southeast O^g. 426); probably flying all year (Fig. 438). Associations. Female OSUC20962 (morphotype II) collected in eucalypt forest, Decem bers. ParastephandJus rufoomatus (Cameron) (Figs I [chapter I], 252-271 (sp. IS), 382-390,428,431) Stephanus rufo-om atus CametoTiy 1905:45. Hemistephanusrufo-ornatus.‘ KieSer, 1908:5. Parastephanellus rufid)matus.~ Motley, 1917:35,112 [Invalid emendation according to ICZN article 3 Id] Parastephanellus rufo-omatus.-Eüiott, 1917a: 92; 1922: 748, 758. ‘Undetermined species’.-Rodd, 1951:341-343. Etym ology. L. rufus, red; L. om atus, decorate. In reference to the reddish areas on the body. MaterialBaanined. H olo^e: ? Burnett River, Qld., R. Semon, ‘1891/92’. OSUC20969 (ZMAN). Other specimens etamined. 74 ??, 56 cfd* (AHC, AMS A, ANIC, BMNH. BPBM, FCNI, MBMA, NSWA, QDPC, QMBA, SAMA, UQIC, USNM). 417 Fem ale L e n g th . Mean 7.0 nun (range 2.4-10.8 nun). Hecui. Flageilomeres subclavate in small specimens, otherwise normal. Frons areolate-rugose; frontal carina distmct. Vertex (pg. 382) transversely sculptured, anteriorly stronger, carinate or strigate, posteriorly strigulate or finely strigulate, sometimes arcuate in an inverted U-pattem. Centro-longitudinal impression absent in small specimens, frint in large ones. Genal angle not prominent, gena round. Gena sculpture dimorphic: (I) Entirely polished smooth, genal stripe complete, running along ^ e margin; or (2) Finely rugulose or strigulate between eye and genal stripe, which is constricted or interrupted sub-basally by same strigulation, otherwise polished smooth. Occipital carina very narrow. Posterior margm of head concave in dorsal view. Occiput polished smooth. Post-genal bridge weakly or distinctly longitudinally or obliquely alutaceous, usually stronger centro-apically. Hypostomal carina apex, below mandible base, from polished smooth in small specimens to weakly rugose in large ones. Thorax. Prostemum apical 0.2S alutaceous, otherwise polished smooth. Pronotum ^ g s 383,384-385); colo anteriorly arcuate-strigate (arcuate-carinate in large specimens); strigations interrupted dorso-posteriorly, entermg pronotal fold; alutaceous dorso- centrally and within pronotal fold, which is laterally well marked, centrally broadly collapsed Qess distinct m small specimens); preannular and semiannular densely transversely strigulate dorsally, laterally more irregular; rarely weakly alutaceous (almost polished smooth); preannular sometimes transversely alutaceous apically; frmoral hnpression sometimes distinct, long and narrow, or broad and indicated anteriorly only; 418 ventral area longitudinally rugulose. Mesoscutum apical 0.3 transversely alutaceous, otherwise rugose-punctate. Axilla ^igs 387-388) concentrically alutaceous-rugulose. Scutellum ^ig s 387-388,390) strongly alutaceous on 0.3 centro-longtudmal; laterally areolate-rugose or punctate. Mesepistemum (Figs 384, 386) microreticulate, some rugulose antero-dorsally; dorsal part alutaceous or polished smooth; ventral part transversely alutaceous, weakly in small specunens; discrimen microfbveolate, foveolae larger anteriorly. Propodeum (Figs 389-390) finely areolate-rugose centro-longitudinally with densely microreticulate or strongly alutaceous intermbced; sometimes a few weak foveae anteriorly (large specimens mainly); areolate-rugose sometimes weak or absent in small specimens. Parapetiolar fovea weakly indicated by large, polished smooth, weakly defined depressions, or indistinct. Spiracular groove indistinct; spiracular carina often weak, irregular, or indistinct; when distinct, ending near flank. Metapleuron ^ g . 389) dorsally fi’om polished smooth with some faint transversely strigulate, to densely transversely strigate, polished smooth between; centrally areolate-rugose, with some microreticulate between; microreticulate anteriorly and ventrally. Post-foveolar area microreticulate, 0-10 transverse carinae. Interfoveolar area transversely carinate, rarely weakly so. Ventral side of fourth tarsomere of front and mid legs normal, not prolonged anterioriy. Hind coxa laterally with a &w areolae basally; basal 0.6-0.7 microreticulate or microareolat^ otherwise transversely strigate; mesally densely punctulate or microreticulate on basal 0.7, apically transv^ely strigate; dorso-central area indistinct, or subdistmct, weakly flattened, finer sculpture, or most of basal half of hind femur flattened m a large subtriangular area. Hind fomur on both sides strongly alutaceous, weaker 419 basally. Hind tibia constricted paît alutaceous; dilated part obliquely alutaceous, ventrally obliquely or longitudinally strigate. Front wing: vein 2Cua tubular on basal 0.3. Pterostigma blunt in small specimens, more elongate in large ones. Abdom en. Petiole finely transversely rugulose, sometimes transversely strigulate coarser basally, broadly polished smooth apex; ventral sculpture similar, finer or microreticulate. Tergite 3 on extreme base rugose, otherwise polished smooth, T4-5 polished smooth, T6-7 often fi’om basally transversely microadculate to apically transversely alutaceous (faintly, apparently polished smooth), T8 sculpture as for T6-7, but much stronger, T9 alutaceous. Pygidial impression triangular, usually deeply notched basally. Pygidium narrow, convoc, emarginate. C olour. Deep dark brown with some rufous to yellowish brown. Genal stripe crossing temple to almost reach ocdpital carina. Yellowish brown, brown, or reddish base of coronal tubercles and interocellar carinae, basal antennal segments, fi’ont and mid legs (mccept white apically on mid tibia and basal 0.75 o f mid basitarsus), bind legs on coxa apically, ventrally and dorsally, trochanter, extreme base and apex of femur, tibia (except yellow basally and sometimes some black laterally), and petiole at least on apical 0.2 (often more). Weakly reddish on posterior margin of semiannular, colo anteriorly or entirely, mesonotum centrally, axillae mesally, scutellum centro-apically, and around petiolar foramen. Yellow mandibles (except black apex), and around antennal foramen. Wing membrane lightly yellowish, more rarely hyalme; often brown or yellowish brown mfiiscate cell 2Cu and area ventrad of it; variable on cells R, IM, 3R1, l+2+3Rs; veins brown, pterostigma brown or dark brown. Ovipositor valves brown or dark brown. 420 D iagnosis. Small size. Always broadly yellow independently around each antennal foramen. Pronotum strigate (Figs 383,385). Mesepistemum microreticulate OFigs 384, 386). Hmd femur stout (FmSI 1.33-1.99; sp. IS on Fig. 258 and table 3.7). Propodeum alutaceous microreticulate, areolate-rugoss finely (Fig. 390). Dorsal part of metapleuron with fins distinct, transverse sculpture (barely visible on Hg. 389 because o f reflexing light). Scutellum mostly strongly alutaceous ^ g s 387-388). Cell 2Cu usually at least partially daricened. Petiole densely rugulose. T8 dull, transversely weakly microalutaceous. Pygidial impression deeply notched. Basal half of hind coxa laterally microreticulate. R em arks. Somewhat similar to P. squamatus n. sp. and P. maculatus n. sp. From the first species it can be immediately distinguished by not having compressed head and thorax; from P. maculatus it can be distinguished by having preannular more or less indistinct fi’om semiannular, finer transverse carinae on vertex, scutellum strongly alutaceous, mesepistemum mostly microreticulate, semiannular laterally strigulate, never polished smooth (although very rarely fidntly alutaceous), dorsal area of metapleuron densely transversely strigate (rarely faint), yellow around antennal forame, finer propodeum sculpture. M ale Very similar to the female, but somethnes very small, showing some character reduction. 421 R em arks ^(fale). Sometimes very sW Iarto small males o f P. isogfyptoso. sp. and P, damellicus^ from which it can be distinguished by having shorter, stouter hind coxae, frons with yellow ventrally, and gena usually wider. The absence o f foveolae on scutellum also distinguishes it from P. damellicus. Distributional and Seamnal Records. This is mostly a eastern species, but there are a fow records for central and southwestern Australia (Fig. 428). Adults collected throughout the year, but apparently more abundant in December (Fig. 431). Associations. Male OSUC2II87 with label inscription ‘^Spotted Gum’ {Eucalyptus cypellocarpa Johnson) without further details. Males OSUC2I097/21098 with label inscription W. decurren^ {pxohtiAy Acacia decurrens WHldenow, MBmosaceae), without further details. Three malM and three females reared from Xylobosca bispinosa (Bostrichidae). hfrde OSUC20922 emerged in ‘December’ from dead A cacia twigs collected October 12. Male OSUC20929 collected aa. Acacia sp. (Leguminosae). Females OSUC20899/209I8 and males OSUC20902/20903 emerged from AngophoraflaribundaL. (Myrtaceae). Male OSUC20936 collected by N. W. Rodd at Lane Cove in 1948 matches local and date o f observations made by Rodd (1951) on the oviposition habits of an ‘undetermined species’ of Stephanidae an. Angophora lanceolata infested with Bostrychidae larvae. This author also provides illustrations for oviposition sequence, mature larva, and male and fomale pupae. 422 Parastephandlus verüpictus Aguiar n. sp. (Figs 248-5,252-271 (sp. 16), 391-397,425) E tym ology. L. vertex, top; h .p ic tu s, painted, coloured. Di reference to colours of vertex. Material Examined. Holotype: ?, Cairns, N. Qld., A, P. Dodd, no date. OSUC20968 (QMBA). Paratypes: 2 ? ? (pMNH, QMBA). Fem ale Length. Mean 12.9 mm (range 10.5-14.4 mm). H ead. Flageilomeres normal. Frons finely areolate-rugose to rugulose; fi^ontal carina weaK but disctincL Vertex (Fig. 391) antero-Iaterally, and shortly behind interocellar carinae, irregularly rugose; centro-Iongtudlnally and posteriorly until occipital carina transversely strigate, stronger anteriorly, finer posteriorly, fading laterally on temple, which is mostly polished smooth. Centro-longitudinal impression weak, distmct under tangential light. Genal angle moderately prominent. Gena polished smooth. Ocdpital carina very narrow. Posterior margin of head concave in dorsal view. Ocdput polished smooth. Post-genal bridge weakfy alutaceous. Stongly strigate near and along hypostomal carina. Thorax. Prostemum (Hg. 395) distinctly alutaceous, weaker centrally; a few oblique carinae anteriorly in larger spedmens. Pronotum ^ g s 392,394): colo fiom arcuate-strigate anterioriy, sometimes stron^y, to transversely strigate centro-laterally; 423 polished smooth or hdntly alutaceous centrally; pronotal fold moderately stout, somewhat flattened, dorsal margin distmctly emarginated; distmctly excavated; centrally weakly collapsed Qess evident in small specimens); preannular anteriorly finely transversely alutaceous, posteriorly, and semiannular dorsally, finely or normally strigate with transversely alutaceous or microreticulate, transversely dorsally, oblique laterally; latter two patterns more evident laterally on semiannular; pronotal lobe and anterior depression in front of it strongly longitudinally strigate. Femoral impression indistinct; ventral area, and sometimes area equivalent to femoral impression, with long and well marked longitudinal strigations. Mesoscutum distinctly microareolate on anterior part, stronger laterally; median lobe strongly areolate and foveolate-rugose. Axilla from strongly subareolate-rugose and alutaceous mesally and posteriorly, to very finely transversely strigulate-alutaceous anteriorly. Scutellum (Figs 393, 150) finely, but distinctly areolate- rugose laterally, otherwise very finely and distinctly alutaceous; in large specimens sparsely transversely strigate centro-anterioriy, contiguos with lateral areolation. Mesepistemum ÇFigs 394,398, part) densely rugulose and microreticulate, coarser anteriorly, where it is also whhe-pilose; microreticulate latero-ventrally, changing to very foiely transversely alutaceous on ventral part; dorsal part weakly or strongly alutaceous, glabrous; discrunen closely foveolate. Propodeum (Kgs 396-397) areolate-rugose centro- longitudinally; flank very finely alutaceous; these two patterns blending gradually laterally, or propodeum mostly areolate-rugose (large specimens); midway laterally also with a few shallow foveolae. Parapetiolar fovea formed by 1-3 large polished smooth areolae. Spiracular groove and carina mdistinct. Metapleuron ^ g . 397) centrally weakly 424 prominent, isolated fiom propodeum by distinct, subcrenulate sulcus originating on pieuropropodeal fovea, ending at propodeal spiracle; centrally strongly areolate-rugose, one single long hair inside each areolation, otherwise glabrous; a few weak rugosities extending anteriorly to origmate 2-3 transvese carinae dorsally on inteifi>veolar area; densely punctate dorsally and anteriorly, weakly so inside more lateral areolae (SOx, fluorescent illumination, steady specimen), Ventrally and on post-fbveolar area finely punctate-microreticulate, with some weak transverse carmae. fiiterfoveolar area âintly alutaceous (apparently polished smooth), 2-3 transverse carinae dorsally, from metapleuron. Ventral side of fourth tarsomere of fi’ont and mid legs prolonged anteriorly, reaching about 0.6-0.8 of length of apical tarsomere. Hind coxa basally rugose, especially mesally, otherwise weakly or ordinarily transversely strigate; a weakly flattened centro- dorsal area. Iflnd femur transversely alutaceous, weaker basally; alutaceous weak, basally sometimes fidnt, in small specimens. Hind tibia very finely obliquely alutaceous; dilated part weakly obliquely carinate ventrally. Front wing; Vein 2Cua tubular on basal 0.25. Pterostigma sub-elongate, apex more or less acute. Abdom en, Petiole dorsally strongly transversely strigate, irregulariy rugose basally; ventrally from finely rugulose or rugulose and transversely strigate; apex broadly polished smooth. Tergite 3 6intly longitudinally alutaceous (apparently polished smooth) in small specunens, more distinct in larger ones, its apical margin and T4-8 transversely microadculate (weak on T3 of small specimens), 18 apically sometimes transversely alutaceous, T9 strongly alutaceous. Pygidial impression deep, V-shaped, shortly notched basally. Pygidium basally flat-triangular, apically narrowly conver, emarginate. 425 C olour. Dark brown, reddish or orange areas on head, thorax, petiole, hind legs. Genal stripe wider oa temple (much wider, somethnes taking all temple, in large specimens), gradually diluted on dorsally reddish or orange and posteriory yellowish brown gena. Antenna yellovdsh or reddish brown on basal halÇ brown apically. Frons entirely reddish brown in small specimens, reddish only along ^ e margin and on triangular area between antennal foramina and base of anterior coronal tubercle (otherwise dark brown) in large specimens. Coronal area in part or entirely, and tips of coronal tubercles, dark brown. Reddish base of coronal tubercles and base of interocellar carinae. Vertex distinctly tricoloured, broadly black centrally, surrounded by reddish anteriorly, narrowly around ^ es, more widely on temple; temple dorsally broadly yellow due to very wide genal stripe. Reddish areas on semiannular laterally, mesothorax and propodeum centro- longitudinally, metapleuron posteriorly or ventrally. Front and mid legs reddish brown in larger specimens, yellowish brown in smaller ones; lighter basal 0.3 or less of mid tibia and basally on correspondent basharsomere. Reddish brown hind coxa ventrally, hind trochanters, apex of correspondent femur. Hind tibia and tarsi lightly coloured, yellowish brown, orange, or reddish brown. Petiole reddish brown or dark ferrugineous, conspicuously more lightly coloured than thorax and gaster, but darker tone than hind tibia and tarsi. Gaster and ovipositor dark brown. Wing veins yellowish brown to brown. Ovipositor valves brown to daric brown. D iagnosis. Head in dorsal view black centrally, reddish laterally, creamy yellow temple (see description for details). Metapleuron dorsally and anteriorly densefy punctate, centrally strongly areolate-rugose, ventrally finely ptmctate and microreticulate; very 426 sparse pttosity (Fig. 397). Scutellum finely, but distmctly areolate-rugose laterally, otherwise very finely and distinctly alutaceous (Fig. 393). Prostemum distinctly alutaceous, this sculpture weaker centrally (Fig. 395). Ventral area of pronotum, and sometimes area of femoral impression, with long and well marked longitudinal strigations (Fig. 394). Pronotum and mesepistemum sculpture also somewhat characteristic (see description). Remarks, Similar and probably related to P, trtfasciatus n. sp., firom which it can be readly distinguished by the diagnosis above and lack of yellow on fi’ons. Overall size also considerably bigger, but this must be considered with caution, since intraspecific size variation in Parastephanelltts is usually high. N ote. Head colours of paratypes was accidentally affected during cleaning process, especially the genal stripe, which almost disappeared, and reddish tones, which got darker. M ale Unknown. Distributiorud and Seasonal Records. Three specimens from northeastem Queensland ^ g . 425), without collecting date. Parastephanellus dénotants Aguiar n. sp. ^ g s 252-271 (sp. 17), 398-403,427) Etym ology. L. denotatus, marked out, conspicuous. In reference to the characteristic sculpture of the body. 427 Material Examined. Holotype; 9, Gayndah, Queensland, no date. OSUC21042 (AMSA). Paratypes: 4 ? Î , 2 cTcf Q^VMA, QMBA, UQIC). Fem ale Length. Mean 12.0 mm (range 8.0-15.1 mm). H ead. Flageilomeres normal. Frons finely, densely microreticulate, rugulose dorsally, sometimes also laterally; a few weak oblique or transverse carinae near antennal feramen; fi’ontal carina absent. Vertex (Fig. 398) rugose in small area behind interocellar carinae; fi"om transversely strigate centrally to transversely substrigulate posteriorly, reaching occipital carina and temples, disappearing laterally at genal stripe; strigation coarser near eyes. Centro-longitudmal impression very weak. Genal angle moderately prominent. Gena polished smooth. Occipital carina very narrow. Posterior margin of head concave in dorsal view. Occiput very finely transversely alutaceous. Post-genal bridge distinctly alutaceous. Hypostomal carina apex, below mandible base, finely longitudinally alutaceous. Thorax. Prostemum densely and conspicuously microreticulate. Pronotum CFigs 399,400): colo arcuately alutaceous or substrigulate; in large specimens, antero-laterally sometimes with a few oblique carinae; pronotal fold flat, transverse, excavated but centrally collapsed; not excavated m small specunens; preannular and semiannular densely and conspicuously microreticulate; dorsally also densely transversely strigulate; posterior margin o f semiannular alutaceous; pronotal lobe, and somethnes preannular apico- laterally, with some longitudinal carinae; femoral impression indistinct. Mesoscutum ^ g . 428 399) completely microreticulate, finer than on pronotum; posteriorly also partially transversely rugulose or transversely rugose. Axilla (Fig. 401) densely microreticulate, with small weak punctures. Scutellum (Fig. 401) densely microreticulate. Mesepistemum (Hg. 400, part) densely microreticulate, including dorsal part; ventral part more weakly, almost alutaceous; discrimen foveolate. Propodeum (Figs 400-401) densely microreticulate with small shallow variolations centrally; flank alutaceous anteriorly. Parapetiolar fovea areolate-rugose. Spiracular groove and carina mdistinct Metapleuron (Fig. 400) densely microreticulate with a few small wrinkles centrally; transversely carinate along line of fusion with propodeum. Post-fbveolar area densely microreticulate. Interfoveolar area weakly crenulate anteriorly, otherwise polished smooth. Ventral side of fourth tarsomere of fi’ont and mid legs normal, not prolonged anteriorly. Ifind coxa at least dorso-centrally and dorso-posteriorly on lateral side, and centro-longitudinally mesally, microreticulate, otherwise microreticulate and transversely strigulate; centro- dorsal flat area indistinct. Hind femur ^ g . 402) very finely transversely alutaceous. Ifoid tibia (Fig. 402) longitudinally alutaceous, as strong as on hind femur; dilated part obliquely strigate ventrally. Front wing; Vem 2Cua tubular on basal O.I. Pterostigma long and blunt Abdomen. Petiole transversely strigate, except transversely rugose basally and broadly polished smooth apeq ventral sculpture similar, sometimes finer; frontal carina distinct basally only. Tergite 3 on extreme base rugulose otherwise very weakly longhudmally alutaceous, its apical 0.1-0.2 and entire T4-8 frintly and finely transversely 429 microadculate (almost polished smooth), T9 alutaceous (Fig. 403). Pygidial impression V- shaped ^ g . 403). Pygidium basally flat-triangular, apical margin upturned, deeply emargmate (Fig. 403). C olour. Ferrugineous and yellow head, black body, partially ferrugineous legs and petiole. Genal stripe wid^ yellow or pale yellow, ending on temple, almost reaching ocdpital carina; gradually diluted on reddish brown gena (head unifbrmily light-brown m one specimen). Yellow broadly around antennal foramen, dypeus apically, mandible, except black apex, posterior margin of semiannular, and wing sclerites. Yellow or yellowish on basal 0.2 of mid tibia and hind basitarsus. Ferrugineous on most of head, semiannular at least latero-centrally, apical 0.2-0.6 of petiole, fl-ont and mid trochanters and femora, hind leg on coxa dorsally and apically, trochanter, apex of femur, and tibia, except narrow black stripe centro-mesally. Legs brownish otherwise. Weakly ferrugineous centro-transversely on tergites, mesally on axilla, basally on scutellum, and propodeum around petiolar foramen.Wing membrane very lightly yellowish; veins yellowish brown. Ovipositor valves browiL D iagnosis. Body dull; firons and thorax, mcludmg hind legs, with very fine and distinct microreticulation (Rgs 400-401). Head usually light ferrugineous, with a wide, fiiz^, creamy genal stripe (Fig. 398) on light-brown or reddish gena. Hind femur slender (FmSI 2.26-2.63; sp. 17 on Fig. 402 and table 3.7). Extremely fine vertex sculpture (Fig. 398) and overall colour pattern also characteristic (see description). 430 Rem arks. This is a very characteristic species, not resembling any other Australian Stephanidae. The diagnosis above, especially the extensive microreticulation, is sufSdent to ident^ both males and females of this spedes. M i/e Very similar to the female. Same diagnosis and comments apply. Distributional and Seasonal Records. Seven spedmens &om southMstem Queensland (Fig. 427), four collected 3 March, 21 November, 25 December; others without collecting date. ParastepkaneUus caudatus Elliott (Figs 252-271 (sp. 18), 404-410,429) ParastephanelluscaudatusEVàoiVi, 1926: 521; 1928:441. E tym ology. L. cauda, tail. In reference to the relative length of the ovipositor. Material Exmined. Holotype: 9 Parasteplumellus caudatus ESào% Borneo, Sandakan, Baker col., no date. 0SUC21195 (BMNH). Other specimens examined: 4 ? ? (Specimen from Australia in UQIC; others m BPBM, USNM) Fem ale Length. Mean 12.2 nun (range 8.2-14.5 nun). H ead. Flageilomeres 2-6 ^lindric, width very uniform. Frons weakly, but distinctly transversely strigate on narrow area along ^ e margin, becoming broadfy 431 subtransversely rugulose or subareolate-rugulose centrally; frontal carina absent Vertex (Fig. 404) from sub-concentrically undulate^strigate behind interocellar carinae to transversely strigate centrally; broadly polished smooth posterioriy and temples. Centro- longitudinal impression weak or absent Genal angle not or weakly prominent. Gena polished smooth. Occipital carina uniformly narrow. Posterior margin of head concave in dorsal view. Occiput polished smooth. Post-genal bridge polished smooth. A distinct carma along anterior portion of hypostomal carina, contiguous posteriorly with occipital carina. Thorax. Prostemum polished smooth. Pronotum ÇFigs 405,407): colo from weakly or fointly arcuate-srigate dorso-anteriorly to weakly alutaceous or polished smooth dorso-posteriorly; strigate lat«ally much stronger, oblique; pronotal fold stout, transverse, distinctly «ccavated, dorsal margin weakly or distinctly collapsed; preannular weakly transversely alutaceous, finer on anterior margin; semiannular widely polished smooth or fomtly alutaceous dorsally, sometimes distinctly alutaceous dorso-anteriorly and laterally; pronotal lobe and depression in front o f it strongly transversely strigate. Femoral hnpression weak, shallow, more or less distmct, polished smooth or almost so; ventral area longitudinally strongly alutaceous. Mesoscutum and axilla ^ ig . 406, axilla only) transversely alutaceous on weakly irregular sur&ce; notauli well marked, deep, more or less contiguous with one another posterioriy, forming a broad ‘U’; median sulcus weak, formed by small closely aligned ptmctations. Scutellum (Fig. 406) uniformly alutaceous induding apical m a r ^ some weak, very shallow foveolae near lateral margin. Mesepistemum distmctly alutaceous, including transversely on ventral part, and more or 432 less longitudinally, weaker, on dorsal part; also sparse very small and shallow punctures, which are coarser and accompanied by a few rugulosities in between anteriorly; discrimen simple. Propodeum ^ g s 408-409) finely and conspicuously transversely alutaceous, includmg mside fbveae, which are larg^ circular, with tendency to absent centrally, somethnes organised m roughly circular fitshion; flank more finely transversely alutaceous. Parapetiolar fovea polished smooth, sub-elongate. Spiracular groove defined by weak areolae or foveolae disappearing midway between spiracle and crenulate sulcus; sphacular carina distinct. Metapleuron (Fig. 409) centrally strongly areolate-rugose; otherwise entirely alutaceous, including inside areolae, faintly or polished smooth on dorso-posterior angl^ below and behind propodeal spiracle; isolated fi'om propodeum by line o f very small areolae or punctations, fi’om pieuropropodeal fovea to base of spiracle, alongside spiracular carina; practically glabrous (one short weak ham inside each areolation). Interfoveolar area fidntly alutaceous or polished smooth. Ventral side of fourth tarsomere of fi'ont and mid legs normal, not prolonged anteriorly. Hind coxa rugulose to subareolate- rugulose basal 0.2, finer or much finer on basal 0.3-0.S ventrally, otherwise transversely strigate, weaker mesally and weaker and finer on flattened centro-dorsal area on lateral side; this area elongate about half as long as coxa. Hmd femur weakly alutaceous; even weaker or polished smooth basally on lateral side. Hind tibia finely and strongly obliquely alutaceous; dilated part obliquely strigate ventrally. Front wing; Vein 2Cua almost entirely nebulous (tubular on less than its O.I basal). Pterostigma sub-lanceolate. Abdom en. Petiole strongly transversely strigate; more irregular basally, finer and weaker ventrally. Tergite 3 varying from fiuntly longitudinally alutaceous or polished 433 smooth m small specimens, to basaUy âintly microaciculate and apical 0.25 alutaceous in large specimens, T4-7 varying from weakly and finely transversely alutaceous in small specimens to strongly and very finely transversely alutaceous in large ones, T8-9 alutaceous, stongest on T9 C^g. 410). Pygidlal impression indistmct (Fig. 410). Pygidhun differentiated into two conspicuous tubular projections, which are apically perforated (Rg. 410). C olour. Dark brown, sometimes with distinct reddish hue. Reddish brown or yellow frons; reddish brown semiannular and prostemum. Genal stripe ivory, fading and disappearmg on temple midway between eye margin and occipital carina. Frons and basal 4-5 antennal segments reddish to yellowish brown. Mandibles, except black tips, and clypeus yellowish brown lighter than on frons. Prothorax dorsally dark brown, changing to reddish laterally and entire prostemum. Reddish between interocellar carinae, petiole partially sometimes, T3 basally. Vertex posterioriy and gena brown or reddish brown; fi%)nt and mid femora brown; front and mid tibiae and tarsi brown with yellowish or reddish hue; pale-yellow basally and apically on mid tibia, and basal 0.4 of correspondent basitarsomere. Hind frmur and tibia dark brown slightly lighter than that of coxa, with weak reddish hue; hind tarsi brown. Terghes yellowish (golden) centrally on apical margin. Wmgs hyaline or weakly infUscat^ vems brown. Ovipositor valves brown, with weak reddish hue. D iagnosis. Pygidium dfferentiated into two conspicuous tubular projections ^ g . 410). Vetter broadly polished smooth posteriorly (Fig. 404). Semiannular shmnin^ alutaceous at least laterally ^ g s 405,407). Frons yellowish or reddish; semiannular 434 laterally and prostemum entnely, reddish. Propodeum and metapleuron entirely and strongly alutaceous, including inside areolae and fbveae (Figs 408-409). Post-genal bridge polished smooth. Vein 2+3M short, only sliglthy longer than 2+3Rs. Vein 2Cua almost entirely nebulous (stability of this character for the present species uncertain). Hmd coxa as long as or longm^ than correspondent femur (Cxl/Fml 1.03-1.12); ^e-ocellus space very narrow Qo/oo 11.38-40.28; sp. 18 on Fig. 404 and table 3.10). Rem arks. Although original^ described 6om Borneo, P. caudatus Elliott is cleady related to the New Guinean species of Parastëphamllus^ which are considerably distmct from the Australian species. In fact, there is at least one undescribed species from New Guinea which is very similar, and probably closely related to P. caudatus. ^ th e collection record is accurate, and P. caudatus is really established in Australia, it can certainly be considered as a recent introduction from the Bomean or New Guinean fruna. It is readily distinguished from other Australian species by the diagnosis provided above. Most relevant differences between the Australian specimen and the Bomean holotype are described next. Australian specimen with yellowish frons, ventral area of pronotum very strongly alutaceous (almost strigate), mesepistemum posteriorly almost polished smooth, propodeum lackmg punctures centrally, and body colour darker, with reddish hue less evident Distinguished from P. m alayanus Cameron (from New Guinea), to which it is similar, by having a distinct pygidium, much narrower interocellar space, lightly coloured frons, well defined genal strips and T4-5 basally transversely microaciculate, contrasting with almost polished smooth T3 of P. malayanus Qiolotype). 435 Similar to description o f P. pygmaetts Enderiein ^ Elliott 1922); distinguished by having different pygidium; frons from laterally transversely strigate to more nregulariy sculptured centrally; scutellum weakly, but distinct^, alutaceous; propodeum strongly alutaceous and sparsely foveolate; petiole as long as or longer than gaster; Ov/TT I S. According to Elliott, P. pygmaeus has a simple pygidium, frons extremely finely and densely punctate, scutellum polished smooth, propodeum densely punctate, petiole shorter than gaster, and Ov/TT 0.7. M ale Unknown. Distributional and Seasonal Records. Essentially from Borneo and New Guinea, but one record in Australia, in the extreme Northeast (Fig. 429). The Australian specimen collected 22 March. Species Laquirendum Parastephanellus picticeps Roman ParastephanelluspicticepsVüonaUy I9 I5 :1 . Parasteplumelluspict^s IsiclJ.-Eliiott 1922:747,754. M aterial Examined^ Holotype: Î, Derby, N. V. Austr., Oct tyear 1910 according to Roman 1915), E. I^ôberg. OSUC20844 (NHRS). 436 Comments. None of the specimens exammed for this work match the holotype of P . picticepSy which, however, seems to be an intermediate between P. damellicus (Westwood)-and P. rufoom atus (Cameron) (Table 3.5). Its hmd leg proportions are mcompatible (FmSQ or peripheral (CxI/h) with those o fP. damellicus^ but Qrpical for P. rufoomcOus. The long gaster, similar vertex/temple sculpture, and small size, are also characteristic ofP. rttfoomatus. However, crucial diagnostic characters ofP. rufoom atus are ‘replaced’ by typical characters for P. damellicus, as the coarse sculpture of the scutellum, unicolourous frons, and a narrow temple {i.e., low tw^EE). Additionally, T3-8 microsculpture is much stronger than that of P. rufoomatus, and the relative length of the pronotum (Pnl/Saw 0.85) is mtermediate between the two recognised species. As in P . rufoomatus (Cameron) As in P. damellicus (Westwood) EBnd femur short and wide ^mSI 1.76) No yellow maAs on fions Hmd coxa relatively short (CxI/h 2.34) Temple relatively short (tw/iEE 0 J6) ^ g id ia l impression deeply notched Scutellum punctate-foveolate Fterostigma apically blunt Propodeum with large areolae Gaster relatively Ions (GsI/PtI 144) Overall size smian Table 3.5. Intermediate characters in the holotype of P. picticepsKovcan. 437 Elliott (1922:754) was able to compare the holotypes o f P. dam ellicus and P, picticeps^ and concluded that these species are closely related, and could be conspecffî<^ but ultimately (on page 156) declined of proposing the synonymy. In the provided in this work the h o lo ^e of P. p icticep s runs easily to couplet 14, and then it can, with a few problems, match P, damellicus. However, the presence of characters mentioned above as characteristic ferP, rufoom atus are considerably beyond the known morphological variability^ for P. damellicus. These mismatches, plus the fact that the morphological variability for P. dam ellicus could be thouroughly investigated, and is now reasonably well understood, make the synonymy unlikely. One possibility is that the holotype of P. picticeps is a hybrid of P. damellicus and P. rufoomatus, showing more dommant characters from the first. This is at least geographically and morphologically possible, as these species are sympatric and share structurally identical genitalia. Note that P. p icticep s was not included in the key to species. Stephanid Taxa Mistakenty Reported in Australia HemistepheamsEnAtAéxL This genus is a monophyletic group restricted to South America, Panama, and Costa Rica (Aguiar 1998). The '^Hemistephanud' species described or identified from Australia ^ e fie r 1908, M ori^ 1917, Elliott I9I9aand 1922, Girault 1926, Moore 1962) are either species o fParastephanellus or Megischus. Megischus coronator(FabriduSy 1804). Holotype 9 ZMHB, not exammed. This species was reported in Australia by many authors, including Schletterer (1889), Kieffrr 438 (1908), Elliott (1922), Ceballos (1926), Dutt (1926), and Baltazar (1966), but this is clearly erroneous. Both Australian species o fMegischus are very distinct from the available descriptions ofM caronator. Megischus duadis (Westwood, 1851). Holotype 9 BMNH, examined. Elliott (1922) mentioned this species as '^widely distributed in the Indo-Australian Region,” but I found no specimens collected in Australia. The two Australian species ofMegischus are very distinct from the holotype ofM ducalis, ParastephanettusscitusEVao% 1919. Holotype ? (BMNH?), not examined. This species is known only from the holotype, collected in Bengal, India. In the original description, Elliott (1919b) mentioned that all other specimens he knew of this “subgenus” (referring to Parastephanellus) were fium Cq^lon, Ding-Ding Island, Borneo and Australia. Dutt (1926) mistakenly mentioned all the localities, including Australia, as actual geographical records for P. scitus. 439 FIGURES 230-236 Megischus crassicauda Morley (holotype): 230, vertec, dorsal; 231, pronotum, dorsal; 232, propodeum, dorso-lateral; 233, metapleuron, lateral; 234, pronotum, mesoscutum, and axillae, dorsal; 235, hind leg, lateral view of trochanters trough tibia; 236, axillae, scutellum, and crenulate sulcus, dorsal. Scale line: 230-233,236, 1.O mm; 234-235, 2.0 mm. 440 441 FIGURES 237-243 Megischus ceneonairix n. sp. Oiolotype): 237, vertex, dorsal; 238, pronotum, dorsal; 239, propodeum, dorsal; 240, metapleuron and propodeum, latero-dorsal; 241, pronotum, mesoscutum, axQlae, and part of scutellum, dorsal; 242, hind le& outer side view of fémur and tibia; 243, detail of hind fémur, outer side. Scale line: 237-240,243, 1.0 mm; 241-242,2.0 mm. 442 443 FIGURES 244-247 Locality^ records for species ofMegischus and Schlettererius: 244; # , M crassicauda Moriey; ■, M. ceneonairix n. sp.; 245, Schlettererius cinctipes (Cameron). Seasonal distribution: 246, Megischus crassicauda Morley; 247, Schlettererius cinctipes (Cameron). 444 9 244 2 4 5 Sep Oct Nov Dec Jan Feb Mar Apr May Jul Aug Sep Oct Nov Dec Jan Feb Mar Month Month 246 2 4 7 445 FIGURES 248-251 Parastephanellus vertipictus n. sp., measurements used to calculate morphometric ratios: 248, body in lateral view; 249, head, dorsal; 250, pronotum, dorsal; 251, tergites 3- 4, dorsal. 446 Txl Gsl Hdl h E w Cxi Fml Ms! Ov 248 TbI I cEE cw f — iP R ---- Pnl T3I K Sal T4I Saw T3w 250 251 FIGURES 252-259 Values o f morphometric ratios for all examined female specimens o f all species. Species numbers as in table 3.8. 448 I I I II I 111 a I I I ! I I I III iiin I II» I If III III niiiiii II 111 ililii! I I illIII III 252 i 253 3 3.5 4 4.5 5 5.5 6 6. 0.75 0.95 1.15 1.35 1.55 1.75 Ov/PÜ Ov/Tt I I I Ill I \mm II II illlBli II ill t I III II II II II I : It I 8 11 I I I . 1 , I I B I 1 »! I I I I I I i n I III! i l l l l l l t l I II I l l I : ' II I ! i l l ' I I I i I i ! 254 255 8 to 12 14 16 18 8 to 12 14 16 18 20 Ov/Hdl Ov/Saw 1 1 1 18 L 1 lit ! ;i 1 16Vt [ \ illliti 1 1 'Ml1 illliI !| 1 |5 14 I. 1 III nil 1 II ram> 1 11 II 1 1 13 ^ nil III II I? f Ml! la 11 1 S I “ : 1 1 9 I Sr ; t 1 6 , !i III IllliI >1 IIII >1111 1 5 lit II III! Ill t ' 1 i i H 43 ! 1 :|l' IIII 1 1 1 11 256 257 1.5 2.5 3 3.5 4 4.5 0.6 0.7 0.8 0.9 I 1.1 1.2 Cxl/h Cxl/Fml 18 \ : I I I I 1 1 1 I 17 , I t , I I I 16 >11 I I I II MHIII 5 Bl 1 1 1 8 1 1 1 8 111 u 14 \ n \ \ \ Ill 111 13 i l l ! ; I i I I I I 1 12 t I I I ' . 1 i I I I I 1 1 1 1 I i ! » I I 1 1 I I I'l 8 \ ' I III iiiimiii 1 < 1 1 8 0 1 II 1 It I 4 M l II i III I t 3 1 1 1 I 258 259 ! Î 1.2 1.4 1.6 1.8 2 2.2 2.4 Z 6 2 8 0.1 02 02 0.4 0.5 0.6 0.7 0.8 0.9 FmSI tw /iE E 449 nOURES 260-267 Values o f morphometric ratios for all examined female specimens o f ail species. Species numbers as in table 3.8. 450 species species species species Q O—hJWAViOtMW^O^WWÎ^VlSi^W o o —K»w#kVi^*N»oe‘OO^h>WAvids^ob 0-'NW4vV%»»WOO'o5ZpW^ui5(Qoo K» bo W e _ I K a; Ov s I species species species species o^h>WAU)0>'^ee^o^Nv»Av>p^M99 o —h>W4i»ViO»'-400^0 0 ^^3 wKÎÂoî^4 oê Q O —NWfkV*(^%400»00—WW$kV)O\'U00 Q 0-»h»WAVlO>'«400'©0^^w!^uî5î^5 00 v% — ~ - - _ # _ : - - » 5 ----_ — U : - _ ^ p 5 W M - w - - 1 - ^ = ^ V, r - i s % S g . - :: - in K> " \ - K k) K N OÎ CO FIGURES 268-271 Values of morphometric ratios for ail exammed female specimens of all species. Species numbers as in table 3.8. 452 I 1 i l l 11 n i M i l I i l I t I Miiiiiiini 11 III II r lllli I I ; ( I I il i ! !1 II I I 1 i l 1 I l I II I I n I I I < I I I I I i II t till I I i I I I I i t i I Il I I M l I i I t i 1 il ! lUi III IBUI II I Ill 11mm 1 tiiiti I I I 1 1 i I i I I I I ! I ! Il ni 268 ‘! i 269 0.35 0.40 0.45 0.50 0.55 0.60 1.2 1.4 1.6 1.8 2 22 24 iEE/cEE ÎEE/Îo X 7 ; tt i i llli < fi il II 5 i I .lit 111 M i l linI I 'Iln 1 :!1 t ilÜ I II f I l I I '.\ II II o II A I II Si 9 1 a. X l in 7 l fi r 5 I I : | m i l l i III 1 1 0 0 i . I t r III i\ ' I a i ; i î\ i 11 u 270 277 i : h L ^ ',. - II 1.3 1.5 1.7 1.9 2.1 2.3 10 20 30 40 Ppl/Ew io/oo 453 FIGURES 272-279 Parastephanellus maculahis n. sp.; 272, head sculpture, dorsal; 273, pronotum and mesoscutum, dorsal; 274, axillae, scutellum, basai section of propodeum, dorsal; 275, pronotum, left; 276, axiHae, scutellum, propodeum, dorsal; 277, posterior half of mesepistemum, metapleuron, propodeum, left; 278, left hind femur and tibia, outer side; 279, tergites 6-9 and pygidium, dorsal. Scale ban 0.25 mm. 454 455 FIGURES 280-288 Parastephanellus brevis XL sp.: 280, head sculpture, dorsal; 281, pronotum and mesoscutum, dorsal; 282, tergites 7-9 and pygidium, dorsal; 283, mesoscutum sculpture, dorsal; 284, pronotum, left; 285, apex of scutellum, propodeum, dorsal; 286, propodeum and metapleuron, left; 287, right &ont win^ venation centrally, dorsal; 288, left hind femur and tibia, detail of sculpture, outer side. Scale bars: 0.25 mm. 456 457 FIGURES 289-298 Parastephanellus laevigatus n. sp.: 289, head sculpture and colour, morphotype I, dorsal; 290, same, morphotype U; 291, pronotum, dorsal; 292, axillae and scutellum sculpture, dorsal; 293, sam^ dififerent specimen; 294, pronotum, left; 295, propodeum, dorsal; 296, thorso^ except pronotum, left; 297, propodoun, detail of sculpture dorso lateral, left; 298, mesepistemum, left. Scale bars: 0.25 mm. 458 459 FIGURES 299-303 Parastephanellusisogfyptosn. sp.: 299, head sculpture, dorsal; 300, pronotum, left; 301, scutellum and propodeum, dorsal; 302, metapleuron and propodeum, left; 303, pronotum and mesoscutum, dorsal. Scale bars: 0.25 mm. 460 461 FIGURES 304-314 Parastephanellus damellicus (Westwood), dorsal view (except where noted otherwise); 304, head sculpture; 305, pronotum; 306, pronotum, detail of sculpture; 307, axillae and scutellum; 308, pronotum, left; 309, axillae and scutellum, second specimen; 310, scutellum, detail of sculpture, third specimen; 311, apex of scutellum and propodeum; 312, mesepistemum, left; 313, propodeum and metapleuron, left; 314, propodeum, detail of sculpture, dorso-lateral, left Scale bars: 0.25 mm. 462 £9P FIGURES 315-321 Parastephanellus albigenuis x\. sp.; 315, head sculpture, dorsal; 316, pronotum, dorsal; 317, mesepistemum posterior half and metapleuron, left; 318, head posteriorly and pronotum, left; 319, thorax, except anterior portion of pronotum, dorsal; 320, axillae, scutellum, basal half of propodeum, dorsal; 321, gaster, dorsal. Scale bars: 315-317,320, 0.1 mm; 318-319,0.2 mm; 321,0.25 mm. 464 465 FIGURES 322-327 Parastephanellus adieaetosn, sp.: 322, head sculpture, dorsal; 323, pronotum and mesoscutum, dorsal; 324, pronotum, left; 325, scutellum and basal half of propodeum, dorsal; 326, thorax, dorsal; 327, mesepistemum posterior halC propodeum and metapleuron, left. Scale bars: 322-324,326-327,0.25 mm; 325, 0.1 mm. 466 g 467 FIGURES 328-337 Parastephanellus callidictyon n. sp.: 328, head sculpture, dorsal; 329, vertex sculpture, detail, dorsal; 330, &ons; 331, scutellum; 332, pronotum and mesoscutum, dorsal; 333, prostemum; 334, pronotum, detail of sculpture, left; 335, scutellum apex and propodeum; 336, mesepistemum posterior hal^ propodeum and metapleuron, left; 337, pronotum, left. Scale bars: 0.25 mm. 468 469 FIGURES 338-347 Parastephanellus Uatytarsus n. sp.: 338, head sculpture, dorsal; 339, head sculpture, anterior view; 340, pronotum and mesoscutum, dorsal; 341, right front winfe venation centrally, dorsal; 342, pronotum, left; 343, scutellum apex and propodeum; 344, axillae and scutellum; 34S, prostemum, ventral; 346, ventral part of mesepistemum, discrhnen; 347, left hind femur and tibia, outer side. Scale bars: 0.25 mm. 470 471 HGURES 348-354 ParastephcmenusgranuIatus(^\&o\i): 348, head sculpture, dorsal; 349, pronotum and mesoscutum, dorsal; 350, right front win& venation centrally, dorsal; 351, pronotum, left; 352, prostemum, ventral; 353, mesepistemum, left; 354, scutellum and propodeum, dorsal. Scale bars: 0.25 mm. 472 473 HGURES 355-358 Pcarastephanellusplatyceratusn. sp.: 355, head sculpture, dorsal; 356, head posteriorly and pronotum, left; 357, pterothorax and propodeum, dorsal; 358, thorax, except pronotum, left. Scale bars: 0.2 mm. 474 475 FIGURES 359-365 ParastephaneilussùigosustLsp.: 359, head sculpture, dorsal; 360, pronotum, detail, dorsal; 361, propodeum and metapleuron, left; 362, head sculpture, detail, dorsal, left; 363, pronotum, left; 364, axillae and scutellum, dorsal; 365, right front wing. Scale bars: 359-364, 0.2 mm; 365, I mm. 476 477 FIGURES 366-373 Parastephanelltis trifasdatus n. sp.: 366, head sculpture, dorsal; 367, pronotum, dorsal; 368, head, anterior view; 369, pronotum, left; 370, mesoscutum, axillae, scutellum, dorsal; 371, same, different specimen, detail; 372, mesepistemum, left; 373, mwepistemum posterior haK propodeum and metapleuron, left. Scale bars: 0.2 mm. 478 479 FIGURES 374-381 Parastephanenussquamatus n. sp.: 374, head sculpture, dorsal; 375, pronotum and mesoscutum, morphotype I, dorsal; 376, head sculpture, posterior view; 377, pronotum, morphotype II, dorsal; 378, pronotum posteriorly, mesoscutum, axillae, scutellum, dorsal; 379, pronotum, left; 380, thorax, except pronotum, dorsal; 381, head and thorax, left Scale bars: 374-380,0.2 mm; 381,0.5 mm. 480 481 FIGURES 382-390 Parastephanellusrufoomatus (Cameron): 382, head sculpture, dorsal; 383, pronotum and mesoscutum, dorsal; 384, thorax, left; 385, pronotum, left; 386, mesepistemum, left; 387-388, mesothorax, dorsal; 389, mesepistemum posterior hal^ propodeum and metapleuron, left; 390, scutellum and propodeum, dorsaL Scale bars: OJZS mm. 482 483 FIGURES 391-397 Parastephanellusvertipictusn. sp.: 391, head sculpture, dorsal; 392, temples, pronotum, mesoscutum, dorsal; 393, scutellum, dorsal; 394, pronotum and mesothorax, left; 395, prostemum, ventral; 396, thorax, except pronotum, dorsal; 397, mesepistemum posterior hal^ propodeum and metapleuron, left. Scale bars: 0.25 mm. 4S4 485 FIGURES 398-403 Parastephanellmdenotatus n. sp.: 398, head sculpture, dorsal; 399, pronotum and mesoscutum, dorsal; 400, thorax, left; 401, scutellum and propodeum, dorsal; 402, left hind femur and tibia, outer side; 403, terghes 7-9 and pygidium, detail, dorsal. Scale bars; 0.25 mm. 486 487 FIGURES 404-410 Parastephanellus caudatus n. sp.: 404, head sculpture, dorsal; 405, pronotum and mesoscutum, dorsal; 406, axillae and scutellum, dorsal; 407, pronotum, left; 408, propodeum, dorsal; 409, propodeum and metapleuron, left; 410, tergites 6-9 and pygidium, dorsal. Scale ban 0.25 mm. 488 489 HGURES 411-423 Locality records for species o îParastephemellus occmnxvg in Australia: 411, all spp.; 412, P. isogfyptos n. sp.; 413, P. maculatus n. sp.; 414, P . brevis n. sp.; 415, P. laevigatus n. sp.; 416, P. albigermis n. sp.; 417, P. adiazetos n. sp.; 418, P. callidictyon n. sp.; 419, P. dcanelUcus (Westwood); 420, P. tanytarsus n. sp.; 421, P. granulatus (Elliott); 422, P. platyceratus n. sp.; 423, P. strigosus n. sp. 490 472 473 474 475 479 476 4 7 7 478 4 2 0 4 2 7 4 2 2 4 2 3 491 FIGURES 424-429 Locality records for species oîParastephanellus occurring in Australia: 424, P. trifasdatus n. sp.; 425, P. vertipictus n. sp.; 426, P. squamatus n. sp.; 427, f . deno tatus n. sp.; 428, P. rufoomatus (Cameron); 429, P. caudatus Elliott. 492 424 425 426 427 428 429 493 HGURES 430-438 Seasonal distribution for AustralianParastephanellus known from seven or more specimens; 430, all species; 431, P. rufoomatus; 432, P. damellicus, 433, P. isogfyptos n. sp.; 434, P. trifasdatus n. sp.; 435, P, strigosus n. sp.; 436, P. platyceratus n. sp.; 437, P. laevigatus n. sp.; 438, P. squam atus xt. sp. 494 I J JASOMDJ FMAM J JASONDJ FMAM J JASONDJ FMAM All specimens 4 3 0 P. rufoomatus 431 P. damellicus 43 2 10 10 8 a I 6Ï 6 I 4i 4 w ! 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XXTV + 205 pp. 505 APPENDIX RANGES OF MORPHOMETRIC RATIOS FOR SPECIES OF AUSTRALIAN PARASTEPHANELLUS NoTSpedes O v/Ptt Ov/Tt Ov/Hdl Ov/Saw n 01 P . m aculatus 3.89-4.20 0.86-I.I6 6.08-8.77 7 5 9 -1 0 5 9 3 02 P . brevis 4.90-5.90 I.0 7 -I.4 I 9.14-11.01 957-12.69 2 03 P . laevigatus 3.22-3.82 0.89-1.07 8.00-10.84 9.77-12.68 8 04 P . isoglyptos 3.60-3.99 0.97-1.02 8.55-9.75 10.32-1152 5 05 P . dam ellicus 3.404.08 0.92-1.16 8.56-11.80 952-13.73 26 06 P . albigerm is 3.95 0.92 7.41 8.96 1 07 P . adiazetos 5.10 1.31 10.81 11.98 I 08 P . callidictyon 4.56 1.06 9.53 10.58 I .09 P . tanytarsus 3.14-3.46 0.84-0.88 6.68-7.11 7.68-8.30 2 10 P . grarm latus 3 .4 I4 .0 I 0.89-1.12 8.35-10.34 9.08-11.37 7 I I P . platyceratus 3 5 5 4 .4 6 1.16-1.38 10.00-1150 12.26-14.05 4 I2 P . strigosu s 458-5.10 1.41-1.69 1251-16.89 1557-1857 8 13 P . trifasdatus 3 .7 7 4 5 7 1.13-155 1055-12.00 11.84-14.63 9 lA P. squamatus 455457 0.80-1.06 754-11.64 6.45-8.79 7 15 P, rufoomatus 3 5 4 4 .7 7 0.83-1.21 5.82-8.46 7.42-10.31 23 16 P . vertipictus 3.48-3.86 0.83-052 6.72-7.46 7.68-8.46 3 17 P . denotatus 3 .8 1 4 5 1 0.98-1.08 8.57-10.40 10.49-11.26 5 18 P . caudatus 4.40-6.09 154-1.77 12.58-19.32 14.91-2154 5 Table 3.6. Mmimum and maxmnim values o f taxonomically important morphometric ratios fi)r Australian species ofParastephanellus, Females. 506 APPENDIX (continued) MbTSpedes CxUh CxI/Fml FmSI tw ^E E n QlP.meuMlatus 2.24-2.53 0.75-0.79 1.42-1.51 0.67-0.75 3 02 P. b revis 2 .I0 -2 J 5 0.73-0.76 1.50-1.81 0.65-0.76 2 03 P, laevigatus 2.59-3.29 0.83-0.90 1.92-2.23 0.55-0.67 8 04 f . isoglyptos 2.81-3.04 0.81-0.88 1.82-2.03 0.60-0.62 5 05 P. damellicus 1-3.58 0.77-0.89 1.97-2.43 0.53-0.65 26 06 P. albigermis 2.48 0.88 1.84 0.58 1 07 P. atSazetos 2.69 0.78 2.03 0.60 1 08 P . callidictyon 1.87 0.70 1.71 0.52 1 09 P. tattytarsus 2.07-2.13 0.74-0.75 1.91-2.07 0.52-0.56 2 10 P . grarm latus 2.22-2.46 0.76-0.79 1.86-2.27 0.49-0.55 7 I I P. platyceratus 3.44-3.73 0.95-0.98 1.94-2.10 0.64-0.70 4 12 P. strigosu s 3.45-4.33 0.89-0.92 1.92-2.51 0.61-0.76 8 13 P. trijbsdatus 3 J3 -3 .9 7 0.87-0.91 2.12-2J0 0.59-0.67 9 14 P . squam atus 1.85-2.63 0.71-0.79 1.63-2.12 0.18-0.35 7 15 P. rujbom atus 2.08-2.83 0.69-0.81 1.33-1.99 0.67-0.80 23 16 P . vertipictus 2.53-2.71 0.73-0.76 2.24-2.46 0.62-0.64 3 17 P. denotatus 2.81-3.04 0.73-0.78 2.26-2.63 0.62-0.72 5 18 P. caudatus 3.67-3.89 1.03-1.12 1.43-1.77 0.72-0.81 5 Table 3.7. N&innum and maximum values o f taxonomically important morphometric ratios for Australian species o fParastephanellus, Females. 507 APPENDIX (contûuied) N oTSpedes btlAv Pnl/Saw twAo Pnl/Ew n 01 P . m aculatus 4.75-5J2 0.70-0.83 U6-1.45 1.07-U9 3 02 P . b revis 4.89-5.17 0.57-0.70 1 .16-U 3 0 .9 9 -U 3 2 03 P . laevigatu s 4.28-6.43 0.97-1.10 0.85-1.10 1.48-1.66 8 04 P . isogfyptos 5.97-6.46 0.84-0.96 1.14-1.19 1.37-1.60 5 05 P . dam ellicus 3.93-6.59 0.79-1.09 0.84-1.21 1.39-1.79 26 06 P . albigenuis 5.46 0.96 1.02 1.52 1 07 P . adiazetos 8.80 0.81 0i>9 1.43 1 08 P . callidictyon 4.78 0.64 0.80 1.10 1 09 P . tanytarsus 6.41-6.44 0.80-0.82 0.75-0.91 1.31-1.46 2 10 P . granulatus 4.82-5.05 0.73-0.95 0.76-0.80 1.26-1.54 7 IIP. platyceratus 4.39-5.15 0.61-0.84 1.29-1.51 1.00-1.34 4 11 P . strigosu s 4 .8 1 -6 J4 0.62-0.93 1.14-1.54 1.01-1.41 8 13 P . trifasdatus 6.92-8.64 0.78-0.95 1.11-1.35 1.23-1.47 9 14 P . squam atus 5.17-7.36 0.85-1.00 0.26-0.56 1.46-2.07 7 15 P . rufoom atus 4.78-7.71 0.60-0.80 1.07-1.46 1.04-1.33 23 16 P . vertipictu s 6.47-6.66 0.83-0.87 1.00-1.05 1.43-1.53 3 17 P . denotatus 6.41-8.16 0.73-0.89 0.94-1.14 1.21-1.59 5 18 P . caudatus 5.35-6.45 0.73-0.88 1.19-1.43 1.12-1.41 5 Table 3.8. Mmônum and maximum values o f taxonomically important morphometric ratios for Australian specie o î Parastephanellus. Females. 508 APPENDIX (contihued) NoTSpedes GsI/PU ptimi Hdl/Ew Ew/tw n 01 P . m aculatus 1.09-1.54 1.94-234 1.88-2.04 1.18-1.41 3 02 P. brevis 1.44-1.69 138-1.75 134-2.01 1.36-1.42 2 03 P . laevigatus 0.95-1.17 234-2.68 1.67-1.93 1.61-2.17 8 04 P. isogfyptos 1 .2 1 -1 J9 1.82-3.67 1.92-2.06 137-134 5 OS P . dam eilicus 1.01-1.42 1.95-235 1.73-2.19 132-2.06 26 06 P. albigenuis 1.43 1.89 132 1.64 1 0 7 PL adiazetos 1 J 2 2 3 6 1.95 1.46 1 08 P . callidictyon 1.51 2.18 1.90 1.75 1 09 P . tanytearsus 1.06-1.35 2.01-2.63 1.87-2.11 1.34-1.81 2 10 P . granulatus 1.06-1.47 2.10-238 1.75-1.98 1.52-1.86 7 IIP . platyceratus 0.88-1.08 2.70-2.98 1.88-2.08 1.26-1.55 4 12 P . strigosu s 0.87-IJll 2.50-3.09 I.72-I.91 1.41-1.69 8 13 P . trifasciatus 0.89-1.37 2.09-2.46 1.81-2.01 1.45-1.67 9 14 P . squamatus 1.51-1.98 1.42-1.92 1.32-1.74 230-4.71 7 15 P . rufoom atus 1.03-1.61 1.73-2.43 136-238 1.10-1.43 23 16 P . vertipictus 1.27-134 1.90-1.96 1.93-1.99 1.41-1.52 3 17 P . denotatus 133-1.43 1.94-2.27 1.85-2.00 137-1.52 5 18 P . caudatus 0.99-1.14 232-2.76 1.69-1.79 1.57-1.67 5 Table 3.9. Nfinimum and maximum values of taxonomically important morphometric ratios for Australian species ofParastephemellus, Females. 509 APPENDIX (continued) N b iS p ed es iEEleEE iEE/io M t/Ew io/oo n 01 P . m aculatus 0.50-0.54 1.88-1.94 1 2 0 -1 3 6 3.69-4.16 3 02 P . b revis 0.45-0.48 1.74-1.78 1.53-1.80 5.69-658 2 03 P . laevigatus 0.43-0.51 1.52-1.75 1.61-152 4.03-7.98 8 04 P . isogfyptos 0.48-0.53 1.90-1.94 1.69-158 3.22-4.50 5 05 P . dam eilicus 0.45-0.60 1.49-2.01 1.57-2.18 2.42-9.76 26 06 P . albigenuis 0.52 1.75 1.66 5.72 1 07 P . a ^ a zeto s 0.49 1.65 1.66 4.63 1 08 P . callidicfytm 0.50 1.53 1.97 7.32 1 09 P . tanytarsus 0.5I-OJ8 1.44-1.63 1.50-1.74 4.90-11.82 2 10 P , gyanulatus 0.49-0.53 1.41-1.57 1.85-2.04 6.99-10.95 7 11 P . platyceratus 0.48-0.52 2.02-2.22 1.44-1.71 2.54-3.17 4 12 P . strigosu s 0.42-0.50 1.82-2.17 1.59-1.77 3.09-5.44 8 13 P . trifasciatus 0.46-0.51 1.82-2.01 1.55-1.78 3.41-5.89 9 14 P . squamatus 0.47-0.54 1.37-1.78 1.59-2.06 5.47-26.46 7 15 P . rujbom atus 0.48-0.55 1.48-1.92 1.37-1.87 3.56-9.68 23 16 P . vertipictus 0.49 (w=3) 1.56-1.70 1.70-1.85 7.03-7.73 3 17 P . denotatus 0.48-0.50 1.45-1.57 1.65-1.82 7.69-1053 5 18 P . caudatus 0.40-0.41 UO-1.78 1.80-2.10 11.38-4058 5 Table 3.10. Mmmium and maximum values of taxonomically important morphometric ratios for Australian species o fParastephanellus. Females. 510