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•

• 1

Niteobrachis elegans sp. nov., 9, lateral view. •

• SYSTEMATICS OF Nn-EOBRACHIS, A NEW IN THE TRIBE BRACHI8TINI (: : )

ANTONIO M. ARANGUREN

A thesis submitted to the Faculty of Graduate Studies and Research in partial fulfillment of the requirements • for the degree of Doctor of Philosophy

Department of Natural Resource Sciences Macdonald Campus of McGill University Ste. Anne de Bellevue, July, 1999

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Canadl •

Thedarkest days as a graduate student were lit by my belief in God, the support of my mother Carmen, my brother Luis and my friends Janet Forrester, Sharlene Sing, Stuart Brown, Duyen Gauthier, Miriam Barrios, Pilar Schneider and Nina Zitani. These people believed in me and assured me that the iII-intentioned are already locked in a dark hell of their own fabrication. Therefore, 1dedicate this dissertation to my supporters and detractors as a testimony ta the triumph of love and scholarship over malfeasance.

• • ABSTRACT Niteobrachis, a new genus in the tribe Brachistini (Hymenoptera: Braconidae: Helconinae), is described. The genus is endemic to the New World and contains 24 species, ail of which are newly described: amazonicus, attonitus, candeo, carmenae, clarus, delicatus, elegans, fulgeo, gracilis, gustus, ictus, insignis (type species), luceo, lucidus, luisi, lumen, luminosus, luna, mica, nitidus, serenus, splendidus, subtilis and venustus. Keys to the species of Niteobrachis and to the genera of Brachistini are given. Niteobrachis is diagnosed by a cambination of characters in the metasoma, including the absence of striations and/or sculpturing on the second and third median tergites with a glossylsmooth appearance; a shallow suture between the second and third median tergites; presence of a crease that separates the second median tergite from the second laterotergite; absence of a crease that separates the third median tergite fram the third laterotergite; fused second and third 1aterotergites; the lateral • longitudinal carinae of the tirst median tergite reaching the articulation between the tirst and second median tergites; and the tirst three median tergites covering most of the remainder of the metasoma. The monophyly of Niteobrachis is supported by two synapomorphies: a minute median longitudinal carina of the propodeum, and the lateral longitudinal carinae of the tirst median tergite reaching the articulation between the tirst and second median tergites. Recognition of this genus resulted from a phylogenetic analysis of six tribes of Helconinae, including most genera of Brachistini based on a data matrix of 49 characters and 18 terminal taxa. The phylogenetic affinities of the genus are examined, a sister group relationship with Foersteria Szépligeti is hypothesized, and the problematic placement of the subgenera of Nees is considered. • RÉSUMÉ SYSTÉMATIQUE DU NOUVEAU GENRE NITEOBRACHIS DE LA TRIBU BRACHISTINI (HYMENOPTERA: BRACONIDAE: HELCONINAE)

Niteobrachis, un nouveau genre de la tribu Brachistini est décrit. Le genre est endémique aux Amériques et constitué de vingt quatre espèces, toutes, nouvellement décrites: amazonicus, attonitus, candeo, carmenae, clarus, delicatus, e/egans, fu/geo, gracilis, gustus, ictus, insignis (espèce type), Iucea, lucidus, luis;' lumen, luminosus, luna, mico, nitidus, serenus, splendidus, subtifis and venustus. Les clefs d'identification des espèces du genre Niteobrachis et d'autres genres parmi les Brachistini est présentée. Le genre Niteobrachis se distingue par une combinaison de caractéristiques du métasoma incluant l'absence de strie et/ou d'ornementation de la cuticule sur le deuxième et le troisième tergites médians et un aspect lisse et brillant; une suture peu profonde • entre le deuxième et le troisième tergites médians; la présence d'un plis qui sépare le deuxième tergite médian du deuxième tergite latéral; l'absence d'un plis qui sépare le troisième tergite médian du troisième tergite latéral; les deuxième et troisième tergites latéraux fusionnés; la carène latérale longitudinale du premier tergite médian; et les premiers trois tergites médians couvrant la majeure partie du metasoma. L'origine monophylétique du genre Niteobrachis est supportée par deux synapomorphies: une minuscule carène longitudinale située au milieu du propodeum et une carène latérale longitudinale du premier tergite médian qui s' étend la articulation qui sépare le premier y la deuxième tergita médian. Ce genre a pu être reconnu suite à une analyse phylogénique de six tribus d'Helconinae, incluant la plupart des genres de Brachistini. Cette analyse a été réalisée à partir d'une base de données comprenant 49 caractères et 18 taxons finaux. Les affinités phylogéniques à • ii ('intérieur du genre ont été examinées. Le placement des sous-genres de • Eubazus Nees est évoqué.

•

• iii • SUGGESTED SHORT TITLE

Systematics of Niteobrachis, new genus.

•

•• iv • ACKNOWLEDGMENTS 1 wish to express my profound gratitude to the Fundaciôn Gran Mariscal de Ayacucho, a Venezuelan govemmental program, and the Ministère de l'Enseignement supérior et de la Science du Québec, which supported my studies in Canada. 1am extremely grateful ta Dr. David J. Lewis, my supervisor and Chairman, for granting me the opportunity to obtain my degree through the Department of Natural Resource Sciences. Dr. Manfred E. Rau was very kind to give me encouragement and advice as to continue with my dissertation and made valuable comments to improve il. 1would Iike to acknowledge Dr. C. van Achterberg for generously suggesting which outgroup would be useful for the phylogenetic analysis and for making valuable comments on my preliminary results. Dr. Victor H. Reynoso introduced me ta the computer programs MACCLADE and PAUP, spent time discussing my approach to the cladistic analysis and made suggestions to improve il. Equally, Alain Roy introduced me • to the programs CLADOS and HENNIG86. To them 1am very grateful. It is very important to me to acknowledge the prompt and wise support, understanding and experience of the professors who took charge of the unwanted situation in which 1 was targeted by an insecure and rootless instructor. Dr. Martha Crago, Associate vice-president of the Graduate Faculty and Research; Dr. Rodger Titman, member of the committee on Graduate Studies, Department of Natural Resource Sciences; Dr. David J. Lewis, Chairman, Department of Natural Resource Sciences, and Dr. Estelle Hopmeyer, Ombudsperson, School of Social Work. Without their advice 1would not have been able to complete this degree.

Appreciation is extended to the following curators and professors who generously loaned material for this study: M. Smethurst, American Museum of • v Natural History, New York, New York; D. Wahl, American Entomological Institute, Gainesville, Florida; D. Ubick, california Academy of Sciences, San • Francisco; M.J. Sharkey and J.T. Huber, Canadian National Collection, Ottawa; J.K. Lièbherr, Cornell University collections, Ithaca, New York; C.B. de Femândez, Fundaci6n Miguel Lillo, Tucuman, Argentina; J. Papp, Hungarian Natural History Museum, Budapest; J. Jimenez, Instituto Nacional de Biodiversidad, Santo Domingo, Costa Rica; C.C. Hsiung, Lyman Entomological Museum, Ste. Anne de Bellevue, Québec; T. Huddleston, Natural History Museum, London, England; G.F. Hevel, National Museum of Natural History, Washington D.C.; C. van Achterberg, Nationaal Natuurhistorisch Museum, Leiden, The Netherlands; S.P. Cover, Museum of Comparative Zoology, Harvard University, Cambridge; R.A. Wharton, Texas A& M University, College Station; R.W. Brooks, Snow Entomological Museum, University of Kansas, Lawrence; J. Pickering, University of Georgia Collection of , Athens; J.B. Whitfield, University of Arkansas, Fayetteville; S.R. Shaw, University of Wyoming, Laramie; S.L. Heydon, University of California, Davis; S. Triapitsyn, University of • California, Riverside; J.L. Garcia, Universidad Central de Venezuela, Maracay; A. Penteado, Universidad de Sao Carlos, Brazil.

1 would like ta extend my thanks and gratitude to a number of people at the Canadian National Collection, whose help and support expedited the completion of this work. Dr. Jean François Landry allowed me the use of his laboratory equipment, gave me advice and helped me capture the images of the wings of the taxa involved in the project. Dr. Alex Smetana identified a specimen. Klaus Balte made the scanning electro-micrographs. Jocelyn Denis and Jennifer Read provided me with advice on how to solve various computer problems. Eric Rickey kindly taught me how to prepare specimens by using the critical point drier method with chemicals, how ta use the Corel Photo Paint Program and, in general, was a valuable resource in the laboratory. • vi ORIGINAL CONTRIBUTIONS TO RESEARCH

• The major original contributions of the present study to the knowledge of Niteobrachis and Brachistini, in general, are as follows:

1. Construction of the first data matrix for the purpose of phylogenetic analysis, including character states for Niteobrachis species, genera and subgenera of Brachistini and tribes of Helconinae.

2. Phylogenetic inference of the relationships of the genera and subgenera of Brachistini and the species of Niteobrachis.

3. Elaboration of the first phylogenetic inference, using cladistic procedures, of the possible relationship of the genera of Brachistini and other tribes of Helconinae which permits a new understanding of the relationships of its taxa and phylogenetic assumption that the monophyly of Brachistini is • questionable. 4. Recognition of Foersteria Szépligeti as the sister group of Niteobrachis.

5. Production of keys and the identification of the species of Niteobrachis and the genera and subgenera of Brachistini.

6. Description of Niteobrachis, new genus and recognition of 24 species by their adult external morphology. The known New World distribution of the genus and of each species presented in map form. Figures are presented iIIustrating the character states of diagnostic and phylogenetic significance.

vii .. 7. This thesis provides data sets useful for future phylogenetic analyses of • Helconinae.

•

• viii TABLE OF CONTENTS

• ABSTRACT ,. RÉSUMÉ , , '" " _--.. . ii SUGGESTED SHORT TITLE iv ACKNOWLEDGMENTS ,... . v ORIGINAL CONTRIBUTION TO RESEARCH _ ,. vii LIST OF FIGURES ,. xii LIST OF MAPS _. _. _" . xvi INTRODUCTION AND LITERATURE REVIEW , -... 1 The Family Braconidae '" ., : _', 1 The genus Niteobrachis , ,, _. '" .. ,. 1 Biology _,.". 2 Biocontrol _, _ ,... . 3 '" _,,. 3 Organization of thesis _'" ,_,. 7 • Objectives of present study , '" _'_,. 7 MATERIALS AND METHODS , .. __ 8 Collection of specimens _. 12 Mounting of specimens '" . 12 Illustrations '" '" . 13 Morphological terminology - , '" . 14 Measurements ...... 14 Description of the species 15 Selection of taxa for cladistic analyses ...... 16 Coding of the character states 17 Matrix and analyses ...... 17 Phylogenetic analysis of the tribe Brachistini to establish monophyly and placement of the genus Niteobrachis , '" . 18

• ix Phylogenetic analysis of the species of Niteobrachis ta establish their Relationship within their genus ,.. 19 • RESULTS ., , ,. . 22 CHARACTER DESCRIPTIONS FOR PHYLOGENETIC ANALYSES 22 PHYLOGENETIC ANALYSES 26 KEY TO GENERA AND SUBGENERA OF BRACHI8TINI , 38 NITEOBRACH/S, GEN. NOV. 42 Generic diagnosis '" ", ,...... 42 General features , '" , 42 Biology '" , ,, '" , 43 Etymology , '" '" ," ' '" .",.. 45 Distribution , , ., '" , '" , '" . 45 Identification of the species _'...... 45 Remarks ., '" '" , 46 KEY TO THE KNOWN SPECIES OF NITEOBRACHIS 47 • SPECIES DESCRIPTIONS ,. ... . 53 Niteobrachis de/icatus sp. nov " ., .." 53 Niteobrachis serenus sp. nov. . ,. '" _, ,. . 54 Niteobrachis /ucidus sp. nov , '" ,. _.. 55 Niteobrachis fu/geo sp. nov , , ,. 57 Niteobrachis /uœo sp. nov. . '" _" '" _.. . 58 Niteobrachis insignis sp. nov _.. 59 Niteobrachis elegans sp. nov. . '" .' ,. 61 Niteobrachis venustus sp. nov. . , ,". 62 Niteobrachis clarus sp. nov. . '" ,,,.. . 64 Niteobrachis gustus sp. nov. . ,. ... . 65 Niteobrachis luna sp. nov , , 66 Niteobrachis luisi sp. nov , '" .. , '" 67 • x Niteobrachis luminosus sp. nov. . ' '" ...... 68 Niteobrachis subtilis sp. nov '" " 69 • Niteobrachis splendidus sp. nov ,. '" 71 Niteobrachis amazonicus sp. nov ,. 73 Niteobrachis lumen sp. nov , '" 74 Niteobrachis ictus sp. nov , '" 75 Niteobrachis carmenae sp. nov '" 76 Niteobrachis candeo sp. nov. 78 Niteobrachis mico sp. nov , '" 79 Niteobrachis attonitus sp. nov. . , ...... 80 Niteobrachis nitidus sp. nov , '" ., , ,... 81 Niteobrachis gracilis sp. nov , 83 DISCUSSION , . 85 Phylogenetic analyses '" '" ...... 85 Genus Niteobrachis , '" ., ,, '" . 87 Distribution of Niteobrachis 88 • Tribe Brachistini -. 89 REFERENCES , 93 FIGURES 104 DISTRIBUTION MAPS 165 APPENDIX 1. CHARACTER FILE OF DELTA 173 APPENDIX 2. TOKEY FILE OF DELTA '" 177 APPENDIX 3. DATA MATRIX '" . 178 APPENDIX 4. CHARACTER STATUS '" 182

• xi LIST OF FIGURES

• 1. Strict consensus tree. Hypothesis of the relationships of Niteobrachis and related genera .. , -.- . 104 2. PAUP most parsimonious tree No. 120 -.-.. _ . 104 2A. PAUP most parsimonious tree No. 120. Hypothesis of relationships ' '" _ 105 28. PAUP most parsimonious tree No. 120. Hypothesis of relationships _ _ . 106 3. Strict consensus tree. Hypothesis of the relationships of the species of Niteobrachis _ . 107 4. PAUP tree No. 522 _. 107 4A. PAUP tree No. 522. Hypothesis of relationships . 108 48. PAUP tree No. 522. Hypothesis of relationships . 109 4C. PAUP tree No. 522. Hypothesis of relationships _,. 110 5. Niteobrachis luna, 9, lateral view ._ . 111 • 6. Head, anterior view. Niteobrachis serenus . 112 7. Head, posterior view. Niteobrachis splendidus - . 112 8. Face. Niteobrachis serenus '. '" ",'. 113 9. Head, anterior view. Urosigalphus (Bruchiurosigalphus) bruchi .. ,., ," __ '.' . 113 10A-8. Head. Eubazus (Allodorus) crassigaster _ '" '" . 114 11-14. Head. 11. Niteobrachis venustus. 12. N. insignis. 13. N. attonitus. 14. N. graci/is ,, ,. 115 15-16. Clypeus " , '" . 115 17. Head, posterior view. Niteobrachis candeo . 116 18. Head and mesoscutum, posterior view. Niteobrachis /una 116 19. Head, posterior and lateroposterior view. Niteobrachis

splendidus. . . 117 • xii 20· Head, lateroposterior view. kurtogaster 117 • 21-21A. Head, lateroposterior view. Foersteria laeviuscula ." . 118 218. Head, dorsal view. He/con tardator _.. 119 22. Mesosoma, dorsal view. Niteobrachis venustus .. _.. . 119 23-24. Mesosoma, dorsal view. 23. Niteobrachis serenus. 24. N. candeo ,, - -. 120 25. Head and mesoscutum, dorsal view. Niteobrachis carmenae '" ,. '" '" ., '" 121 26. Mesoscutum. Niteobrachis amazonicus '" ". 121 27. Mesoscutum and metanotum, dorsal view. Niteobrachis luna '.' '" _.. . 122 28. Mesoscutum, dorsal view. Niteobrachis lucidus 122 29. Head, posterior view, and mesoscutum, dorsal view. Po/ydegmon 'oveo/atus '" , ,. . 123 29A. Metanotum. Urosigalphus (Bruchiurosigalphus) bruchi .. '" . 123 30. Mesosoma, partial lateral view. Niteobrachis serenus 124 • 31. Pronotum, partial lateral view, and mesopleuron. Niteobrachis insignis 124 32. Mesosoma, partiallateral view. Eubazus (Brachistes) salicicola " , '" .,. . 125 33. Mesosoma, partial lateral view. Eubazus (Eubazus) pallipes. 125 34. Pretarsus. Niteobrachis elegans '" '" ... . 126 35. Pretarsus and apical tarsomere. Schizoprymnus (Schizoprymnus) ambiguus _.. ,. 126 36. Pretarsus and apical tarsomere of Urosigalphus (Bruchiurosiga/phus) bruchi 126 37A-B. Hind coxa of Foersteria laeviuscula 127 38. Forewing of Niteobrachis species ,. 128 39. Hind wing of Niteobrachis species 128 • xiii 40A-B. Measurement taken in the wing venation of Niteobrachis species ',,," '._, ,. . 129 • 41-44. Forewings of Niteobrachis species ,,. 130 45-48. Forewings of Niteobrachis species _ '" _.. . 131 49-52. Forewings of Niteobrachis species ,,, '" , _.. 132 53-56. Forewings of Niteobrachis species ',, '" .,.. 133 57.()O. Forewings of Niteobrachis species ,_, 134 61'()4. Forewings of Niteobrachis species 135 65. Forewing of Helconini. He/con yukonensis .. , '" .-, '" .,. . 136 66'()7. Forewing of Brachistini. Triaspis kurtogaster and Schizoprymnus (Schizoprymnus) ambiguus 136 68. Forewing of Brachistini. Urosigalphus (Bruchiurosigalphus) bruchi ,,,_,_,, '" . 136 69. Forewing of Brachistini. Schizoprymnus (Muiriella) concisa 137 70-71 . Forewing of Brachistini. Eubazus (Ca/yptus) chinensis and Eubazus (Brachistes) lapponicus - '" . 137 • 72-75. Hind wings of Niteobrachis species ,. 138 76-79. Hind wings of Nifeobrachis species 139 80-83. Hind wings of Niteobrachis species , " '" '" '" . 140 84-87. Hind wings of Niteobrachis species 141 88-91. Hind wings of Niteobrachis species , -. .. . 142 92-95. Hind wings of Niteobrachis species "" '" . 143 96. Hind wings Brachistini. Urosigalphus (Bruchiurosigalphus) bruchi '-,, '" , '" , '" ,. 144 97-98. Propodeum. dorsal view. Niteobrachis serenus and N. insignis ,, '" ., '" .. ,,,.. _. '" . 144 99. Metanotum and propodeum. Niteobrachis venustus .. '" . 145 100. Mesosoma. dorsal view. Niteobrachis splendidus .. -. '" .,. . 145 101 . Mesosoma. dorsal view. Niteobrachis candeo 146 • xiv 102. Propodeum and metasomal first median tergite. Niteobrachis /uns " '" '" '" , 146 • 103-104. Propodeum. dorsal view. Niteobrachis clarus and He/con tardator ,-.-..,," '" -. '" " _. 147 105-106. Metasoma, dorsal view. Niteobrachis splendidus and N. candeo ---..,, '" '" '" . 148 107. Metasoma, dorsal view. Nea/io/us rufus -. '" 149 108-112. First median tergite. Niteobrachis serenus, N. lucidus, N. venustus, N. splendidus and N. insignis '" -...... 150 113A. Metasoma, lateral view. Female and Male. Niteobrachis sp/endidus '" '" , _...... 151 113B. Metasoma, dorsal view. Female and Male 152 113C-O. Metasoma, lateral view. Niteobrachis candeo 153 114A-B. Metasoma, lateral view. Nea/io/us rufus 154 115A-B. Metasoma, lateral view. Eubazus (A/iD/US) crabilli '" 155 116A-B. Metasoma, lateral view. Foersteria laeviuscula 156 • 117A. Metasoma, lateral view. Triaspis kurtogaster 157 1178. Laterotergites. Triaspis kurtogaster 157 118A-B. Metasoma, lateral view. Po/ydegmon (oveo/atus .. __ . '" . 158 119A-B. Metasoma, lateral view. Eubazus (Brachistes) salicicola '" 159 120A-B. Metasoma, lateral and dorsal view. Che/ostes vardzia ..... 160 121A-B. Metasoma, lateral and dorsal view. Schizoprymnus (Muirie/la) concisa ,,,,...... 160 122. Metasoma, lateral view. Eubazus (Allodorus) crassigaster. 161 123A-B. Metasoma, lateral view. Urosigalphus (Bruchiurosigalphus) bruchi , ,. 162 124A-B. Metasoma, lateral view. Schizoprymnus (Schizoprymnus) ambiguus " ,. '" .,. . 163 125. Ovipositor length compared to body length ," ...... 164 • xv LISTOFMAPS

• Map 1. Composite distribution of Niteobrachis species in

Costa Rica and Panama '" ,, 0 O" •••• 165 Map 2. Composite distribution of.Niteobrachis species in

the Neotropical region 0 •••••••••••• 165 Map 3. Known distribution of Niteobrachis delicatus ,. 166 Map 4. Known distribution of Niteobrachis serenus .. , . 166

Map 5. Known distribution of Niteobrachis lucidus 0 •••• 166

Map 6. Known distribution of Niteobrachis fulgeo . o ••• , ••• • •••• 0 ••••••• 166

Map 7. Known distribution of Niteobrachis luceo ., o •••••••• 167 Map 8. Known distribution of Niteobrachis insignis . 167 Map 9. Known distribution of Niteobrachis elegans .. , . 167

Map 10. Known distribution of Niteobrachis venustus 0 0 ••••• , •••••••• 167

Map 11. Known distribution of Niteobrachis clarus o ••••• , •••••• 0 168

Map 12. Known distribution of Niteobrachis gustus - 0 •••• 168 • Map 13. Known distribution of Niteobrachis luna , '" . 168

Map 14. Known distribution of Niteobrachis luisi , 0 ••••••• 168 Map 15. Known distribution of Niteobrachis luminosus . 169 Map 16. Known distribution of Niteobrachis subtilis " .. 169

Map 17. Known distribution of Niteobrachis splendidus '00' •••••••••• 169 Map 18. Known distribution of Niteobrachis amazonicus . 169 Map 19. Known distribution of Niteobrachis lumen ,. 170 Map 20. Known distribution of Niteobrachis ictus .. ,,, '" . 170 Map 21. Known distribution of Niteobrachis carmenae ,.. 170 Map 22. Known distribution of Niteobrachis candeo , '" '" . 170 Map 23. Known distribution of Niteobrachis mico , '" . 171

Map 24. Known distribution of Niteobrachis attonitus o ••••••••••••••••••• 171 Map 25. Known distribution of Niteobrachis nitidus , .. '.-,.. 171 • xvi Map 26. Known distribution of Niteobrachis gracilis . 171 • Map 27. Known distribution of Foersteria Szépligeti in the Palearctic Region compared to that of Niteobrachis Gen. Nov. 172

•

• xvii INTRODucnoN AND LITERATURE REVlEW • The Family Braconidae: Species of Braconidae (Hymenoptera) are of other . They are an important influence on the population dynamics of theïr hosts, many of which are economically significant herbivorouslphytophagous insects (Shaw and Huddleston, 1991; LaSalle and Gauld, 1993). Despite the importance of these parasitic in biocontrol and insect ecology, their biology and taxonomy remain poorly known (Huber, 1993).

The Braconidae is a large and diverse family with a wide range of hosts and life cycles, and include an estimated 40,000 species worldwide (Townes, 1969). Closely synchronized with their hosts' lite history, braconids are of primary importance in the biological control of insect pests (LaSalle, 1993; Wharton, 1993). Their success in biological control programs has been mainly achieved against Lepidopteran and Dipteran pests, and predominantly by the genera • Apante/es Foerster (Microgastrinae), Bracon Fabricius (Braconinae), and Opius Wesmael (Opiinae) (Matthews, 1974; Greathead, 1986).

The genus Niteobrachis: This study provides a comprehensive review of a new genus, Niteobrachis (Braconidae: Helconinae: Srachistini). The projeet began as a revision of the genus Nea/io/us Mason and in the course of the examination of the material it was discovered that sorne specimens, although very similar to Nea/io/us, could not be placed in that genus. This raised the question as to whether or not Nea/io/us was a monophyletic group and the possibility that the unreported specimens could represent undescribed species, though similar to species in Nea/io/us, possibly represent an otherwise as yet undescribed genus.

Niteobrachis is morphologically similar to Nea/io/us and A/iolus Say in the tribe Brachistini. Specimens of this new genus have been identified as Nea/iD/us or Aliolus in various entomological museums because they will readily key out to those genera when existing keys are employed (Marsh et al., 1987; Sharkey, 1997). However, an uncharacteristically noticeable and consistent smoothness on the second and third median tergites remains unaccounted for. The present work deals with 24 newly described species known to be distributed throughout the New World from Costa Rica to Argentina. Niteobrachis species have been poorly collected perhaps because of their small size and the specifie types of traps required; as such, specimens of this genus and braconids in general are more commonly collected by Malaise traps. Most specimens of the genus have been collected from Costa Rica due to the massive efforts of Dr. Paul E. Hanson. Niteobrachis is conspicuously absent from braconid material collected in Chile, although other undescribed and closely related taxa of Brachistini have been collected trom that part of the Neotropical Region.

Biology: • The biology of this genus is largely unknown. However, four specimens belonging to the National Museum of Natural History (Smithsonian Institution, Washington D.C.) have labels identitying the host as Faustinus apicalis Faust, 1896 (Coleoptera: ), thus reconfirming that members of Helconinae parasitize . ~elconinae have been determined to be koinobiont (parasitoids that allow hosts to continue to grow in size after parasitism, Godfray, 1994) endoparasitoids (Gauld, 1988; Gauld and Bolton, 1988; Shaw and Huddleston, 1991; Wharton, 1993) of egg and larval stages of wood-boring beetles (Clausen, 1954; Haeselbarth, 1962; Parnell, 1964; Shenefelt, 1970; Capek, 1971; Shaw and Huddleston, 1991; Papp and Maeto, 1992).

• 2 Biocontrol: The most racent reports of biocontrol involving genera of Brachistini have found • Eubazus atriconis Ashmead, E. minutus Nees and Triaspis pal/ipes ta contribute to the mortality of the curculionids (Coleoptera) Pissodes notatus Fabricius, P. piniphilus (Herbs), P. stmbi (Pack) (Alauzet, 1987; Campadelli and Scaramozzino, 1994; Kenis and MiIIs, 1994; Luik, 1994; Kenis etal., 1996) and Rhynchaenus fagi (Priore et al., 1991; Tsankov and Stalev, 1992, 1995). Allodorus crassigaster (Provancher) was reported to be an enemy of the white pine weevil which causes severe damage ta spruce in Canada and the United States (Alfaro et al., 1994). Triaspis aciculatus (Ratzeburg) was found to be a of the raspberry and bramble curculionid Anthonomus rubi Herbst (Scanabissi and Arzone, 1992). Likewise, unidentified species related ta the genus Urosigalphus were confirmed ta parasitize gall-forming beetles belonging ta the genus Prionomerus (Curculionidae) (Suarez, 1992). Charlet (1994) investigated the life cycJe of Nea/io/us curcu/ionis (Fitch), which parasitizes the sunflower stem weevil Cylindrocopturus adspersus (LeConte). Further study conduded by Charlet and • Seiter (1994) concluded that N. curcu/ionis parasitized the curculionids Smicronyx fulvus LeConte and S. sordidus LeConte feeding on sunflower seeds. Finally, Hemandez et a/., (1994) reported an undescribed species of Nea/io/us emerging trom fruit flies of the genus Anastrepha (Diptera: Tephritidae).

Taxonomy: Because the present research is based on the description of the new genus Niteobrachis, there are no previous references ta the genus. The following historical review deals with other genera belonging to the tribe Brachistini Mason.

Muesebeck and Walkley (1951), and Martin (1956) assigned the genera B/acus, Ca/yptus, A/iolus, Triaspis and Urosigalphus ta the subfamily Blacinae, whereas Telenga (1952, in Dudarenko, 1974) assigned the aforementioned genera to

.~.'"1 3 Calyptinae. Later, Capek (1970) took a further step and assigned them to the • tribe Blaeini (Heleoninae) based on the cephalie structures of the final-stage . Subsequently, Masan (1974) proposed the tribe Braehistini on the basis of the shape ôf the elypeus and the presence of only two cubital ceUs in the forewing (Le., the lack of forewing vein r-m). Brachistini is currently comprised of eight described genera and the newly described genus herein, and is cosmopolitan in distribution: Che/ostes van Achterberg (Palearetic), Eubazus Nees (Holarctic), Foersteria SZépligeti (Palearetic), Nea/iolus Mason (Nearctic, Neotropical), Niteobrachis new genus (Neotropical), Polydegmon Foerster (Palearctic), Schizoprymnus Foerster (Neotropical, Holarctic, Oriental, Australian), Triaspis Haliday (Neotropical, Holarctic, Oriental, Australian), and Urosigalphus Ashmead (Nearctic, Neotropical).

The most comprehensive reference for generic synonymy and host records of Brachistini is Shenefelt (1970), supplemented by Marsh (1979). Few general revisions on the genera of this tribe have been undertaken, perhaps because of • the problematic generic division of BrachistinL This problem results from their morphological similarities and especially from the fact that they do not present a sole characteristic that defines each individual genus, but offer instead the combination of a few characteristics. In addition, there is a point of contention among sorne Braconidologists who do not agree on the placement of certain taxa such as Schizoprymnus (Muiriella) and the subgenera of Eubazus. Finally, limited research on the family Braconidae, including the vast number of its subfamilies and species, may have contributed to the lack of research on the tribe Brachistini itself.

One signifieant contribution has been made by Mason (1974) who resolved the nomenclature of Brachistini. Mason's research defined, named the tribe as such, and identified two synonyms: Calyptinae Marshall (1887) and Triaspinae Viereck, • 4 (1918). Additionally, he delimited and gave synonyms for the following genera: • Eubadizus and Eubadizon, which were cited as junior objective synonyms of Eubazus Nees; Brachistes, Calyptus and Eubazus, which were reeognized as subgenèra of Eubazus; Alio/us and AI/odoms which were delimited and Nealio/us whieh was described as a new genus. The publication gives an aceount of the speeies of the newly delimited taxa and includes a key to genera and subgenera of Brachistini.

Martin (1954) redescribed Foersteria and subsequently (1956) reviewed the Nearetie fauna of Alio/us, Triaspis, Schizoprymnus and Urosigalphus. Martin's final research on Braehistini was published in 1958 with the description of a new speeies of Triaspis.

Recently, van Aehterberg (1990) reviewed the genera Foersteria and Po/ydegmon and deseribed the new genus Che/ostes. He also synonymized Alio/us as a subgenus of Eubazus and relegated AI/odorus ta a subgenus of Eubazus. No • explanation was given for this deeision. Papp (1984, 1991, 1993) redescribed Triaspis and described new Inda-Australian species of bath Triaspis and Schizoprymnus.

Baltazar (1966) synonymized the genus Muiriella Fullaway with the genus Triaspis, however it was most reeently treated as a subgenus of Schizoprymnus by Papp (1984). The ambiguous taxonomie status of Muiriella Fullaway is due to its elose relationship ta bath Triaspis and Schizoprymnus.

Additional related citations include Snoflak's (1952) monograph contributing ta the knowledge of Triaspis from the Czeehoslovakian fauna. Papp and Maeto (1992) described a speeies from the Oriental region and gave a list of speeies of the eastern Palearctie region. Papp later (1995) synonymized Sigalphus brucivorus • 5 Rondani with Triaspis thoracicus Curtis, whereas Belokobylskij (1994) contributed to the knowledge of Brachistini by identifying new species of Eubazus, • Schizoprymnus and Triaspisfrom far eastem Russia. Crawford (1914) revised the genus Urosigalphus ta indude the description of eight new species. More recently, Gibson (1972a, 1972b, 1974, 1982) reviewed the Westem Hemisphere fauna of the genus, including newly described species, and the known biology and range of each species.

Cornelius van Achterberg (1976, 1993), Shaw and Huddleston (1991) and Sharkey (1993) published keys identifying members of Helconinae as Brachistini. Works that provide iIIustrated keys for ail genera and/or subgenera of Brachistini in the Westem Hemisphere are those of Marsh et al. (1987), van Achterberg (1990), Shaw (1995) and Sharkey (1997). Gibson (1972a) and Martin (1956) keyed the genera of Brachistini as members of Blacinae. Papp gave keys for the Indo-Australian fauna of SChizoprymnus (1993) and Triaspis (1984), while Tobias • (1986) keyed the Russian fauna of Foersteria. Comelius van Achterberg (1984), Quicke and van Achterberg (1990), van Achterberg and Quicke (1992) and Wharton et al. (1992) assessed and reassessed phylogenetically the subfamily Helconinae. The tirst attempt ta hypothesize the relationships of sorne of its genera was pursued by Capek (1964), in which he stated that B/acus, Ca/yptus, Urosiga/phus, A/io/us and Triaspis presented an "unnatural and heterogeneous group on a basis of bath theïr bïonomics and larval taxonomy". A hypothesis of relationships of Brachistini with other tribes of Helconinae was suggested by van Achterberg (1983) in which its members were distinguished by the absence of forewing vein r-m as an apomorphous character state, and in which the possible relationships of the tribas of Helconinae were alsa considered.

6 Oraanization of thesis: The combined results of this thesis are divided into a sequence by which the • reader may be able to follow the evants undertaken in this systematic study. The morphological characters used for the identification of taxa and phylogenetic analyses are described first, then followed by the cladistic procedures used to supports the monophyly and erection of the new genus. A key to related genera and subgenera of the tribe Brachistini is presented to distinguish specimens of Niteobrachis within this tribe. Then, the description ofthe new genus Niteobrachis is given, followed by a key to the identification of its species. After a specimen of this new genus has been keyed out, the morphological information on the specimen can be compared to the description of the species which follows.

Objectives of present study: It is the hope that the information given here will enable specialists to make comparisons of Niteobrachis with other genera of Brachistini and Helconinae so that they can form their own conclusions regarding the classifications and • phylogenetic relationships of this new genus.

Because the phylogenetic relationships of the genera of Brachistini have not been conclusively anaIyzed, this study presents a preliminary approach to the understanding of the relationships of Niteobrachis with other genera of Brachistini. The goals of this studyare: to describe the genus Niteobrachis, to delimit and describe the included species, and to provide a key to their identification. A phylogenetic analysis of Niteobrachis and related genera is also performed to support the monophyly of Niteobrachis, to establish the sister group relationships of the genus, and to provide a preliminary analysis of the relationships of genera within the Brachistini.

• 7 MATERIALS AND METHODS

• This study is the result of the examination of over 3000 adult specimens tram four

subfamilies of Braconidae. Of the above-mentioned number of specimens,

around 650 specimens belong ta relatad taxa such as ,

Macrocentrinae, Orgilinae, and Helconinae other than the genus Neatia/us. An

excess of 2350 specimens belong to the latter genus from which 159 specimens

were separated to erect a new genus, Niteobrachis.

This material inaudes the following subfamilies, tribes, genera, subgenera and

their species:

Agathidinae: Earinus limitaris (Say), E. nitidulus (Nees).

Helconinae: • Tribe : Blacus (Blacus) chiJensis van Achterberg (Holotype), B. (B/acus) humi/is (Nees), B. (Blacus) Iongicaudatus Tobias~

Tribe Brachistini: Chelostes: C. varclzia (Enukidze).

Eubazus: E. (A/io/us) crabilli Martin, E. (AJioIus) fiskei (Rowher), E.

(AlioIus) marginatus Martin, E. (A/ialus) tomoxoiae (Rohwer), E.

(AllOOorus) crassigaster (Provancher), E. (Allodorus) definitus

Muesebeck, E. (AI/odorus) emobii (Muesebeck), E. (A/IOOorus)

semirugosus (Nees), E. (AJtodorus) tibia/is (Haliday), E. (Brachistes)

major (Cresson), E. (Brachistes) rotundiceps (Cresson) (Hototype), E.

(Brachistes) salicicola (Muesebeck), E. (Ca/yptus) chinensis

8 (Watanabe), E. (Eubazus) pallipes Nees. • Foersteria: F. laevïuscula Szépligeti. Nea/ioIus: N. coIlaris (Srues), N. crassipes (Martin), N. curculionis

(Fitch) (Holotype), N. rufus (Riley).

Polydegmon: P. foveolatus (Henich-8chaffer), P. sinuatus Foerster,

SChizoprymnus: S. (SChizoprymnus) ambiguus (Nees), 5.

(SChizoprymnus) brevipennis (Mefander and Brues) (Holotype), S.

(Muiriella) concisa (Fultaway).

Triaspis: T. caudatus (Nees), T. kurtogaster Martin (paratype), T

laticarinata Martin, T. ace/lulata Martin, T. thoracica (Curtis), T.

virgin;ensis (Ashmead).

Urosigalphus: U. (Bruchiurosigalphus) bruchi Crawford, U. • (Neourosigalphus) punctifrons Crawford, U. (Microurosigalphus) diversus Gibson (Halotype), U. (Microurosigalphus) femoratus

Crawford, U. (Microurosigalphus) neomexïcanus Crawford, U.

(Neourosigalphus) flexus Gibson (paratype), U. (Neourosigalphus)

porteri Gibson (HaIGtype), U. (Neourosigalphus) robustus Ashmead,

U. (Urosigalphus) barbe'; crawford, U. (Urosigalphus) donnae

Gibson, U. (Urosigalphus) torbesi Martin.

Tribe Brulleini: Brulfeia latiannulata (Cameron).

Tribe Cenocoeliini: nigriventris (Cresson), C. saperdae

(Ashmead). • 9 Tribe Diospilili: fomitus Masan, Diospi/us oIeraceus • Haliday. Tribe Helconini: Helcon tarclator Nees, H. yukonensis (Ashmead), H.

nunciator (Fabricius).

Macrocentrinae: Macroœntrus canarsiae Muesebeck, M. crambivorus Viereck,

M. uniformis Provancher.

Orgilinae: Orgi/us fictus Muesebeck (Holotype), O. graci/is (Srues).

As it was mentioned above, only 159 specimens belong to the genus

Niteobrachis. The foflO'A'ing abbreviations indicate the collections from which

matarial was borrowed (t'ne author alsa reviewed specimens by directly visiting

insect collections including: the Sohart Museum, the California Academy of • Sciences, the Canadian National Collection, the Museum of Comparative Zoology, the Museum of Insects at the University of Wyoming, the Museum of

Insects at the Universidad Central de Venezuela and the Smithsonian Institution).

AEI = American Entomological lnstitute, Gainesville, Florida.

AMNH = American Museum of Natural History, New York, New York.

BMNH = The British Museum (Natural History), London, England.

CAS = California Academy of Sciences, San Francisco. CNC = Canadian National Collection, Ottawa.

CUI = Comell University Insect Collections, Ithaca, New York. • 10 IML = Fundaci6n e Instituto Miguel Lillo. San Miguel de Tucuman. • Argentina. tNBio = rnstituto Nacional de Biodiversidad, Santo Domingo. Costa Rica.

LEM = Lyman Entomological Museum. Ste. Anne de Bellevue. Québec.

MCZ = Museum of Comparative Zoology, Harvard University. Cambridge.

NNML = Nationaal Natuurhistorisch Museum, Leiden, The Netherlands.

SEM = Snow Entomological Museum. University of Kansas, Lawrence.

TAMU = Texas A & M University, College Station.

TMA = Természettudomanyi Museum Allatara, Budapest, Hungary. UMM = University ofArkansas Museum, Fayetteville. UCDC = Bohart Museum, University of Califomia, Davis. UCR = University ofCalifomia, Riverside. • UCV = Universidad Central de Venezuela. Maracay. UDSC = Universidad de Sao Cartos, Brazil.

UGCA = University of Georgia Collection ofArthropods, Athens.

USNM = United States National Museum, (National Museum of Natural

History) Smithsonian Institution, Washington O.C.

UWY = University of Wyoming. Laramie.

ln order to make collection data citations more accurate, the spelling of locality

names given on specimen labels has been modified in sorne cases to conform

with those in various gazetteers of the world (Bartholomew, 1957; The Earth and • 11 • Man, 1972; National Geographie Atlas ofthe Wood! 19901- Collection ofspecimens:

Members of Braconidae were collected either by sweeping the wasps from

vegetation or by trapping specimens with flight-interception traps (Masner and

Goulet, 1981), yellow-pan traps or Malaise traps (Townes, 1962; Gauld and

Bolton, 1988). Malaise traps are considered the most effective method for

callecting large numbers of species (Wharton, 1997).

Mounting of specimens:

Specimens preserved in a'cahol were treated with a specialized drying procedure

to maintain three-dimensional morphology affording maximum visibility of ail body • ragions (Gordh and Hall, 1979; Wharton, 1997). The specimens were removed from the vials where they had been preserved, then were dehydrated through a

series of increasing concentrations of ethyl alcahol, ending with a solution of 95%

(Gordh and Hall, 1979). Samples were subsequently immersed in amyl acetate,

purified grade (Fisher Scientific), for 2 to 4 hours under a fume hood. After this

treatment, the specimens were removed and allowecl to dry under the fume hood;

the dried specimens were then mounted on triangular points.

As mentioned on the previous page, of the 3000 specimens studied, an excess of

2350 specimens belonging to Nealiolus Mason (1974) were examined and divided

12 into groups according to striations and/or sculpturing on the second and third • median tergites. 159 specimens nct exhibiting striations and/or sculpturing on either their second and third median tergites. and thus having a shiny and smooth

appearance, were assigned ta the group to be studied. These specimens were

examined with a Wild Heerbrugg stereo-microscope with magnifieations of 375X,

500X and 750X, that were chosen as dictated by the size of the strudures being

evaluated. Charaders with reliability in defining the species (Appendix 1) were

chosen and a database was created based on these characters and their states

for further analysis in the computer program Oesaiption Language for Taxonomy

(DELTA) (Dallwitz et al., 1994).

Illusbalions: • A fore and a hind wing of a specimen of each taxon was eut off at the tegula and placed in warrn water ta dean, relax and flatten the specimen. Flattened wings

were then temporarily mounted on slides with ladic acid (Sigma) 98% (Cumming,

1992). Siides of wing venation were viewed with a Nikon optiphot microscope and

the images captured with a Sony OXC-950 color video camera, 18X magnification.

The resulting TIF files were saved to disk and the images manipulated with Corel

Photo Paint version 5.0 to remove air bubbles and/or impurities produced during

mounting of the slides. The wings were then placect in an entomological microvial

containing glycerol and pinned beneath the specimens. Scanning electro

micrographs were taken with a Zeiss OSM-940A electron microscope; mast • 13 specimens were not gold coated. Ail specimens were put in a relaxing jar • ovemight ta moisten them ta enhance condudivity before photographing. Drawings were made with the aid of a Wild camera-Iucida attachment on the

stereo-microscope.

Morphologlcal termlnoIogy:

Terms used to desaibe adult morphology are based on van Achterberg (1988),

Shaw and Huddleston (1991) and Huber and Sharkey (1993). The system

employed for naming wing veins has been taken tram Huber and Sharkey (1993)

and Sharkey and Wharton (1997) (Figs. 38-39). Similarly, names of propodeal

structures were taken from Townes (1969) and Sharkey (1988).

• .e.surements: Apart trom ratios, ail measurements are given in millimeters and were obtained

with a Wild Heerbrugg stereo microscope using a micrometer eyepiece. The

measurement of the total body length was taken tram the distance of the insect

median ocellus to the posterior margin of the metasomal third median tergite (Fig.

125). Ovipositor measurements were taken tram the tip ta the suture between

stema 5 and 6 (Fig. 125). Whenever the number of specimens allowed. an

average of the ratio of body Jength ta ovipositor was made, taking measurements

fram the largest and smallest insects. The angles of fore wing veins were

calculated with a protrador tram enlarged pidures of the wings with straight lines • 14 imposed on them (Fig.40A). Lengths (in millimeters) of hind wing veins R over • 1M were macle to create ratios and were eut off arbitrarily into three continuous groups (0.6 to 0.7, 0.4 to 0.5 and 0.2 to 0.3) (Appendix 1, character 26). Equally,

lengths (in millimeters) of hind wing veins 1r-m over 1M were made to create

ratios and were eut off arbitrarily into three continuous groups (0.7 to 0.8, 0.9 ta

1.1 and 1.2 to 1.5) (Appenctix 1, character 27). The measurements of the above

mentioned hind wing veins were made with a ruler on enlarged pidures of the

wings (Fig. 408).

Descriptions ofthe speeies:

The descriptions of species and the key were produced with the computer

program DELTA (Dallwitz et al., 1994), a general system for processing • taxonomic interadive identifications and information retrieval. The key for the species was constructed using the weights given ta the characters

under the directive "character reliabilities" of the TOKEY subroutine (Dallwitz et

al., 1993) of the DELTA program (Appendix 2), rated from 1 to 10 depending on

theïr reliability, with 10 being the most reliabJe. The diagnostic descriptions of

each taxon were initially generated automatically using the command "diagnose"

of the INTKEY (interactive key) subroutine (Oallwitz et al., 1993) of DELTA

(Dallwitz and Paine, 1994). Each description distinguishes the species tram ail

the rest in at least one attribute. The bold-face sentences found in the resulting

desaiption of the species were the diagnostic charaders used ta identity each • IS • selection oftaxa for cladlstlc analyses:

Earinus Westmael (Agathidinae), Macrocentrus Curtis (Macrocentrinae), Orgi/us

Nees (Orgilinae) and Helcon Nees (Helconinae) were chosen to be the outgroups

taxa for charader polarization. Earinus, Macrocentrus and Olflilus are considered

closely related to Helconinae (Wharton et a/., 1992) while He/con is regarded as

the most primitive Helconinae (Capek, 1969; Tobias and Belokobylskij, 1983).

Genera belonging to tribes of Helconinae were examined, including Cenocoe/ius

Haliday (Cenocoeliini); Brul/eia Szép1igeti (BrulJeiini); Diospi/us Haliday

(Diospilini); B/acus (Blacus) Nees (Bladni); Eubazus (Eubazus) Nees, E.

(Brachistes) Westmael, E. (Calyptus) Haliday, E. (Ailodorus) Foerster, E. (A/ia/us) • Say, Nea/io/us Mason, Triaspis Haliday, Urosiga/phus Ashmead, SChizoprymnus Foerster, Schizoprymnus (Muiriella) Fullaway, Polydegmon Foerster and

Foersteria Szépligeti (Brachistini).

Blacini is considered as a tribe of Helconinae for the purpose of this study (van

Achterberg, 1976), due to the similarity of its morphological attributes with

Brachistini (Capek, 1983), although it has been previously regarded as a

subfamily of Braconidae (Capek, 1970; van Achterberg, 1988, 1993). Likewise,

Cenocoeliini is treated here as a tribe of Helconinae (Saffer, 1982; Sharkey, 1993;

Shaw, 1995), even though van Achterberg (1994) considered it a subfamily of • 16 • Braconidae. Codlng ofthe characterstates:

The coding of the charader states was done following the outgroup analysis

method in which slate 0 is the ancestral state. This characteristic is possessed by

the outgroups, thereby allowing subsequent reversais to state o. (Maddison et al.,

1984, Maddison and Maddison, 1996).

Matrix and analyses:

The charaeter matrix (Appendix 3), composed of 42 taxa and 58 charaders

(characters and states listed under "Character desaiptions for Phylogenetic

Analysis" on page 22), was crealed by entering the data as a spreadsheet in the • computer program for phylogenetic analysis MACCLADE version 3.06 (Maddison and Maddison, 1996).

ln the data matrix, characters 5, 7. 8, 22, 23, 26 and 27 were coded with a

question mark for ail genera involved in the study, other than taxa Kancestor" and

the species of Niteobrachis. This was because it was difficult to confidently

determine the states at the species level due to polymorphisms and to the

substantiaf number of species.

The matrix was analyzed using the computer program Phylogenetic Analysis • 17 Using Parsimony (PAUP)1 version 3.1.1. (SwoffordI 1993). PAUP is a program for • infening phylogenies from character data uncler the principle of parsimony.

Phylogenetic analysis of the tribe Brachistini to establish monophyly and

placement ofthe genus Niteobrachis.

The following analysis conventions are stated below 50 that the results can be

replicated and ifpossible later improved upon: multistate taxa were interpreted as

polymorphie; bath ACCTRAN and DELTRAN optimization were used (Swofford

and Maddison, 1987). AJthough ACCTRAN and DELTRAN charader

optimizations present different distributions of synapomorphies, ACCTRAN was

selected for the character optimization. Characters were treated as weighted

equally with a weight of unity and unordered (Appendix 4); the maximum number • of trees (MAXTREES) saved was 1000;24 taxa were deieted, including ail species of Niteobrachis (Appendix 4); outgroup chosen was "ancestor- or first

taxa in the data matrix (Appendix 3 & 4); ingroup included Cenocoelius, Brul/eia,

Blacus (Blacus), Diospilus, Eubazus (Allodorus), E. (Brachistes), E. (Calyptus), E.

(Eubazus), E. (Aliolus), Nealiolus, Triasp;s, Urosigatphus, Schizoprymnus

(Schizoprymnus), Schizoprymnus (Muiriella), Polydegmon, Foersteria and

Niteobrachis.

PAUP 3.1.1. has fundional limits regarding the number of taxa and the type of

search that can be used to test a matrix (Swofford and Segre, 1993); therefore, • 18 large matrixes must be analyzed using heuristic algorithms (Swofford and Begle, • 1993). Onlyexhaustive searches are guaranteed to find the most parsimonious trees, tharefore, a branch-and-bound search was performed. Branch-and-bound

searches require less than 20 taxa in the matrix in arder to be completed. Taxon

No. 18 or Niteobrachis (Appendix 3) represents the summary of ail data in the 24

species of that genus, 50 the number of taxa could be reduced and this type of

exhaustive search could be undertaken. As a result, only 18 taxa were included in

the search (one taxon in the outgroup and 17 in the ingroup) (Appendix 4); of the

58 charaders in the data matrix, 49 were informative and 9 were uninformative

(charaders 5, 7, 8, 19, 22,23,26,27 and 28) (Appendix 4).

The strict consensus tree (Fig. 2) was chosen ta summarize the most • parsimonious resolutions because it contains only monophyletic groups that appear in ail of the rival trees (Sakai and RohIf, 1981). One of the most

parsimonious resolutions, PAUP tree No. 120, was chosen arbitrarily in arder to

discuss the synapomorphies, since any of the most parsimonious resolution

developed in this analysis could have been a possible hypothesis to have

occurred.

Phyloaenetic analysis of the sPeCies of Niteobrachis to establish their

relationships within thaïr genus.

This analysis was alsa performed using the following conventions: multistate taxa • 19 were interpreted as polymorphie; bath ACCTRAN and DELTRAN optimization • were used (Swofford and Macldisan, 1987); and ACCTRAN was setectecl for character optimization. Characters were treated as weighted equally with a

weight of unity and unordered (Appendix 4); the number of maximum trees saved

was 1000 since by raising this number the effectiveness of the search was

diminished due ta tree-buffer overflow; six taxa were deleted including Ancestor,

Cenocoe/ius, Bru/leia, Blacus (Blacus), Diospilus and Niteobrachis; outgroups

chosen were Eubazus (Allodorus), E. (Brachistes), E. (Calyptus), E. (Eubazus), E.

(A/ioIus), SChizoprymnus (SChizoprymnus), S. (Aluine/la), Foersteria, Nea/io/us,

Polydegmon, Triaspis and Urosiga/phus (Appendix 4): ingroup included the 24

species of Niteobrachis.

• The analysis was performed by a heuristie search. Heuristie searches do not ensure the most parsimonious resolutions and trees obtained by using this type

of search are referred ta as the minimum length trees. As suggested by Swofford

and Begle (1993), ail combinations of heuristic search options were performed,

including general, starting trees, stepwise addition and branch swapping. The

branch swapping option that produced the minimum length trees was the

alternative tree-bisection-reconnection (TBR). To maximize the performance of

the heuristic search, changes in the input order of taxa into the data matrix were

performed because input arder can potentially influence the resull The input

arder of taxa into the data matrix that proved to produce the minimum length trees • 20 • was as it is in the data matrix (Appendix 3). Only 36 taxa in the data matrix were included in the heuristic search (12 taxa in

the ct.etgroup and 24 in the ingroup) (Appendjx 4); of the 58 charaders in the data

matrix only 42 were informative and counted in length calculations and 16 were

uninformative (charaders 1,3,4, S, 7, 8 10, 12, 15, 18,20,22,23,26,27 and 36

were either excluded or constantlignored) (Appendix 4). Characters 5, 7, 8, 22,

23, 26 and 27 were exduded from the analysis since the chosen outgroup taxa had

no data to polarize them. When there is a question mark in a character that belongs

to the outgroup or anœstor, by default the PAUP program d100ses the next available

datum of the next taxon in the data matrix and makes the assumption that il is the

primitive state. If those character had been left included in the analysis, they would • have alter the results. In tact, although they were exduded from the anaJysis and were net inclucled in length calculations, they appear to be informative (Appendix 4).

A stnd consensus tree (Fig. 3) was chosen to summarize the minimum length

trees because it contains only monophyletic groups that appear in ail ot the rival

trees (Sokal and Rohlf, 1981). One of the minimum length trees, PAUP tree No.

522 (Fig. 4), was arbitrarily chosen to discuss the synapomorphies since any of

the minimum length trees could have been a possible hypothesis to have

occurred.

• 21 RES ULTS

• CHARACTER DESCRIPnONS FOR PHYLOGENETIC ANALYSES

Head:

1. Frontal cavity: (0) deeply concave, (1) concave. 2. Frontal ridge: (0) absent, <,1) present. 3. Anterior oœllus: (0) within frontal cavity, (1) extemal to frontal cavity. 4. Medial projection of frons: (0) present, (1) absent or greatly reduced. 5. Texture of face: (0) rugose, (1) punctate, (2) smooth. 6. Inter-tentorial line compared to ocular-tentorial line: (0) equal or longer, (1) shorter. 7. Texture of clypeus: (0) rugose or heavily pitted, (1) punctate. • 8. Surface of clypeus: (0) flat to weakly convex, (1) very convex. 9. Width of clypeus: (0) 2.S to 2.7 limes wider than long, (1) 2.0 times wider than long. 10. Lower margin of clypeus: (0) slightly curved, (1) with medio-ventral tooth, (2) medially depressed, (3) straight. 11. Number of segments of maxillary palpi: (0) S, (1) 6. 12. Number of segments of labial palpi: (0) 4, (1) 3. 13. Temples: (0) evenly convex, (1) raised laterally. 14. Occipital carina: (0) ending ventrally on hypostomal carina, (1) ending ventrally on subgenal carina, (2) reduced ventrally.

22 Mesosoma:

• 15. Forewing vein 1RS: (0) present, (1) absent.

16. Forewing vain 3RS: (0) not deeply curved anteriorly, (1) deeply curved anteriorly. 17. Forewing vein 2RS: (0) straight or nearly so, (1) curved. 18. Forewing vein (RS+M)b: (0) present, (1) absent. 19. Forawing vein 1m-cu: (0) straight. (1) curvad. 20. Forewing vain r-m: (0) present. (1) absent. 21. Forewing vain 2cu-a: (0) present. (1) absent. 22. Forewing vein 2RS angle with respect to (Rs+M)b from: (0)1260 to 130°, (1) 121° to 125°, (2) 116° to 120°, (3) 111° to 115°, (4) 106° to 110°. 23. Anterior half of forewing vein 1cu-a angle with resped to A trom: (0) 121° to 125°, (1) 1160 to 120°, (2) 111 0 to 115°, (3) 1050 ta • 110°. 24. Forewing vain 1a: (0) present, (1) absent. 25. Forewing vein 2a: (0) present. (1) absent. 26. Ratio of length of hind wing vein R to 1M from: (0) 0.6 ta 0.7, (1) 0.4 to 0.5, (2) 0.2 to 0.3.

27. Ratio of length of hind wing vein 1r-m ta 1M from: (0) 0.7 to 0.8t (1) 0.9 to 1.1, (2) 1.2 to 1.5. 28. Hind wing vein cu-a: (0) curved at least in posterior 1/5, (1) straight. 29. Longitudinal carina on median lobe of mesoscutum: (0) absent. (1) present 1/3 to 1/4 posterioriy, (2) complete. 30. Crenulation of notauli: (0) medium, (1) large, (2) small. 31. Number of scutellar sulci: (0) 4 or more, (1) 2. • 23 32. Stemaulus: (0) rugose, (1) impressed, (2) not impressed. 33. Median longitudinal carina ofpropodeum: (0) long, (1) minute. • 34. Lateral longitudinal carinae of propodeum: (0) straight, (1) eurved. 35. Posterior transverse carinae of propodeum: (0) absent, (1) present. 36. Median areola ot propodeum: (0) present, (1) absent. 37. Median areola of propodeum: (0) divided, (1) undivided. 38. Claws: (0) unlobed basally, (1) lobed basally.

Metasoma:

39. First three median tergites: (0) not covering remaining segments, (1 ) covering remaining segments. 40. First three median tergites: (0) not densely setose, (1) densely setose. 41. First three median tergites: (0) without a flanged lateral margin, (1) with a flanged lateral margin. • 42. Lateral longitudinal carinae on tirst median tergite: (0) present, (1) absent. 43. Lateral longitudinal carinae on tirst median tergite: (0) not reaching posterior margin, (1) reaching posterior margin. 44. First median tergite posterior border between lateral longitudinal carinae: (0) shorter than posterior border lateral to lateral longitudinal carinae, (1) nearly equal to posterior border lateral ta lateral longitudinal carinae, (2) longer than posterior border lateral ta lateral longitudinal carinae. 45. Striations and/or sculpturing on tirst median tergite lateral to lateral longitudinal carinae: (0) present, (1) absent. 46. Second median tergite: (0) completely delimited trom laterotergite by lateral crease, (1) partially delimited trom laterotergite by lateral t :e.' 24 crease, (2) not delimited tram laterotergite by lateral crease. 47. Striations and/or sculpturing on second median tergite: (0) present, • (1) absent 48. Articulation between first and second median tergites: (0) present, (1) absent. 49. Articulation between first and second median tergites: (0) narrow, (1) wide. 50. Third median tergite: (0) not delimited from laterotergite by lateral crease, (1) delimited trom laterotergite by lateral crease. 51. Striations and/or seulpturing on third median tergite: (0) present, sometimes weakly, (1) absent. 52. Suture between second and third median tergites: (0) very weak and shallow, often almost completely absent, (1) weil defined, (2) absent medially. 53. Posterior and lateral margins of third median tergite: (0) without a tlanged margin, (1) with a flanged margin. • 54. Posterior margin of third median tergite: (0) curved (sometimes slightly), (1) notched, (2) with one or two projections. 55. Shape of second and third laterotergite: (0) rectangular, (1) trapezoidal, (2) nearly transversely ellipsoideal. 56. Suture between second and third laterotergites: (0) present, (1) absent. 57. Second and third laterotergites: (0) not covering sterna, (1) covering stema. 58. Ovipositor: (0) equal to or longer than body length, (1) shorter than body length.

~. 25 PHYLOGENETIC ANALYSES

• Phylogenetic analysis of the tribe Brachistini ta establish the monophyly and placement of the genus Niteobrachis: The branch-and-bound search found 120 most parsimonious trees of 212 steps, with a consistency index (CI) of 0.736, a retention index (RI) of 0.600 and a rescaled consistency index (RC) of 0.442. The strict consensus tree (Fig. 1) is unresolved due to ambiguity in the evolutionary steps and/or lack of character support (Swofford and Begle, 1993). The branches of the third most basal Iineage compose two unresolved polytomies, one consisting of Brul/eia, Eubazus (AI/odorus), E. (Brachistes), Blacus (Blacus), Eubazus (Calyptus), E. (Eubazus) and Diospilus, with the other branch consisting of Eubazus (Alio/us), Nealio/us, Triaspis, Urosigalphus, Schizoprymnus (Muiriella), S. (Schizoprymnus), Polydegmon, Foersteria and Niteobrachis. Within this latter polytomy Urosigalphus and Schizoprymnus (Muiriella) show a sister group relationship. • The subsequent results emphasize the diagnostic characters of specifie branches as reconstructed from the above analysis and are based on PAUP most parsimonious resolution tree No. 120 (Figs. 2, 2A-B) and are explained starting trom the most basal taxon of Brachistini, Eubazus (AI/odorus) towards more inclusive taxa. 70% of the most parsimonious resolutions had a character distribution in which Foersteria was the sister group of Niteobrachis. Tree No. 120 was arbitrarily chasen ta represent this sister relatianship.

Eubazus (Allodorus) is supported by three synapomorphies: 1(0), lower margin of clypeus with media-ventral tooth (Fig. 10B); 49(1), wide articulation between tirst and second median tergites (Fig. 122), a characteristic that has also evolved independently by parallelism in Eubazus (Atio/us); and 52(1), a suture between second and third median tergites which is weil defined (as Fig. 115A). A wide articulation between first and second median tergites is a character state also • 26 featured by E. (Alio/us), where this state is shown ta have evolved independently. • Although E. (Allodorus) figures more basally in the cladogram (Figs. 2 & 28) than the rest of Brachistini genera, the author expected it ta be closely related to E. (Alio1us) and Nealiolus because of the presence of a carapace-like tirst three median tergites. In reality, Eubazus (AI/odorus) does not have a carapace-like structure formed by its first three median tergites; rather, its third median tergite does not even caver the remaining median tergites as it equally occurs in Eubazus (Brachistes), E. (Calyptus) and E. (Eubazus).

Eubazus (Brachistes) (Fig. 2A) is supparted by one synapomorphy: 43(1), lateral longitudinal cannae on first median tergite reaching posterior margin (as Rg. 105). This derived character state also evolved independently in E. (Calyptus), Nealiolus and Niteobrachis (Figs. 2A & 28). The lengths of the carinae vary either reaching or not reaching the posterior margin of the first median tergite in Eubazus (Eubazus), E. (AHo/us) and Triaspis making the character dimorphic, • thus creating ambiguity.

Eubazus (Calyptus) (Fig. 2A) is supported by one synapomorphy: 43(1), lateral longitudinal carinae on first median tergite reaching posterior margin (as Fig. 105). As stated in a paragraph above, this is a derived character state that has evolved independently in lineages such as Eubazus (Brachistes), Nealiolus and Niteobrachis (Figs. 2A &28).

Eubazus (Ca/yptus) forms a monophyletic group with B/acus (B/acus) (Slacini) (Fig. 2A) supported by one synapomorphy: 21 (1), forewing vein 2cu-a absent (Fig. 70). The absence of this vein also •occurs in sorne species of Urosigalphus in the tribe Brachistini. The presence of forewing vein 2eu-a is of taxonomie • 27 importance because it is a charader that diagnoses Helconinae when combined • with other structures; even so, it is not a synapomorphy for this subfamily of Braconidae (Shaw and Huddleston, 1991). It is of interest to mention that B/acus (B/acuS) (Blacini) is taxonomically differentiated from the genera of Brachistini by a scutellar dise that is marginated by carinae and by a deep dorsope, character state that were not included in the phylogenetic analyses because are uniquely found in Blacini and as a result they are uninformative characterizations.

The immediately preceding common ancestors of Eubazus (AI/odorus), E. (Brachistes), E. (Calyptus) and E. (Eubazus) are hypothesized to be different with respect to the other genera and subgenera of Brachistini. As a result, this tribe is rendered paraphyletic (Figs. 2 &2A).

Eubazus (Eubazus) is hypothesized to form a monophyletic group (Fig. 2B) with • a branch that includes Eubazus (A/io/us), Nealiolus, Tnaspis, Schizoprymnus (Schizoprymnus), Urosigalphus, Schizoprymnus (Muiriella), Polydegmon, Foersteria and Niteobrachis (Fig. 2B). This relationship is supported by one synapomorphy: 37(1), an undivided median areola of propodeum (Fig. 97). The areola of propodeum is a character that is widely found in the subfamilies Ichneumonoidae and Braconidae; the carina that usually divides the areola appears and disappears throughout the species in these two families of Hymenoptera.

Eubazus (Eubazus) is shown not to be monophyletic since it is not supported by one synapomorphy (Fig. 28). the species of this subgenus are distinguished from other subgenera of Eubazus byan epicnimial carina that does not reach the mesostemum (Fig. 33), a unique characteristic within the tribe Brachistini. • 28 Apart from Eubazus (AI/odorus), E. (Brachistes) and E. (Calyptus) the following • genera and subgenera of Brachistini form a monophyletic group including E. (Aliolus), Nea/io/us, Triaspis, Schizoprymnus (Schizoprymnus), Urosigalphus, Schizoprymnus (Muiriella), Polydegmon, Foersteria and Niteobrachis, a derived group supported by two synapomorphies (Fig.2B): 39(1), the first three median tergites form a carapace that covers the remaining segments (Fig. 1130) and 55(1), the shape of the second and third laterotergites are trapezoidal (as Fig. 114B). These derived charaeter state are so outstanding that they separate this group trom four subgenera of Eubazus (Allodorus, Brachistes, Calyptus and Eubazus) and support the redefinition of Brachistini as a monophyletic group without the latter subgenera of Eubazus. Nevertheless, a phylogenetic analysis of the species of ail subgenera of Eubazus must be preceded by a reexamination of the relationships of the genera of Brachistini, a task beyond the scope of this dissertation. Excluding Foersteria and Po/ydegmon, the above monophyletic group (Fig. 2B) also contains genera and subgenera with a primitive character state shared with their ancestor: 46(0), a second median tergite completely • delimited trom laterotergite bya lateral crease (Fig. 1148).

Within the above branch there are two lineages (Fig. 2B), one which supports the sister group relationship of Foersteria and Niteobrachis. This will be discussed further below. The other branch contains Eubazus (Aliolus, Nealiolus, Triaspis, Schizoprymnus (Schizoprymnus), Urosigalphus, Schizoprymnus (Muiriella) and Po/ydegmon, none of which are supported by synapomorphies; instead, they share two plesiomorphic states: 47(0), striations and/or sculpturing on second median tergite (Fig. 115A); and 51 (0), striations and/or sculpturing on third median tergite (Fig. 115A).

• 29 As mentioned above, Eubazus (A/io/us) contains those species supported by a sole synapomorphy, 49(1), a wide articulation between first and second median • tergites (Fig. 115A). This articulation was also independentJy derived in Eubazus (AI/adoms) and it is not found in any other genera of Brachistini. The sister group of Eubazus (Alio/us) is a group of four genera and two subgenera: Nealio/us, Triaspis, Schizaprymnus (Schizoprymnus), Urosiga/phus, Schizoprymnus (Muiriella) and Po/ydegmon (Fig. 2B). The latter taxa form a monophyletic group based on three synapomorphies: 24(1) and 25(1), forewing veins 1a and 2a absent respectively (as Fig. 67); and 33(1), median longitudinal carina of propodeum minute (as Fig. 97), although this latter character state has reverted in Schizoprymnus (Muiriella).

Nealio/us is supported by the possession of two synapomorphies. The tirst, 38(1), are claws lobed basally (as Fig. 34) that have evolved independently in Triaspis, Foersteria and Niteobrachis; these lobes are morphological structures • that have not been discussed in Braconidae and their function is perhaps to give the support when grabbing the host during oviposition. The other derived character is 43(1), the lateral longitudinal carinae on first median tergite reaching posterior margin (Fig. 107), and it is a derived character state which evolved independently in Eubazus (Brachistes), E. (Calyptus) and Niteobrachis (Figs. 2A & 2B). As it is shawn in the most parsimonious tree (Fig. 28), Nealiolus possesses one symplesiomorphy, 11 (0), which is also present in Triaspis, Schizoprymnus (Schizoprymnus) and Urosigalphus. This primitive character state is the maxillary palpi with five segments (as Fig. 19).

Triaspis, Schizoprymnus (Schizoprymnus), Urosiga/phus and Schizoprymnus (Muiriella) form a monophyletic group (Fig. 2B) based on three synapomorphies: 6(1), an inter-tentorial line that is shorter than the ocular-tentoiral line (as Fig. • 30 108), a character state also found in three subgenera of Eubazus (Allodorus, Brachistes and A/io/us); 41(1), in which the first three median tergites have a • flanged lateral margin (fi8g. 121 A); and 48(1}, where the articulation between first and second median tergites is absent (Rg. 123A-B). The last two synapomorphies have only been derived in Triaspis, Schizoprymnus (Schizoprymnus), Urosigalphus and Schizoprymnus (Muiriella) in the tribe Brachistini, which indicates that this is a strongly supported monophyletic group. These four taxa do not have an articulation between their first and second median tergites; this absence is a taxonomie characteristic that only occurs in these genera within the Brachistini.

Triaspis and Schizoprymnus (Schizoprymnus) have a sister group relationship (Fig. 2B) based on one synapomorphy: 16(1}, the forewing vein 3RS, is deeply curved anteriorly (Fig. 66), a derived character state that has only evolved in these two taxa within Brachistini. The forewing vein 3RS is also found as curved • in other species of Braconidae, such as in Coelinius sp. (Alyssinae), Dinocampus sp. and Westmalia sp. (Euphorinae), implying that this character state evolved in eartier ancestors. Triaspis is supported by two synapomorphies: 11 (1), maxillary palpi that have six segments (cf. Fig. 19); and 38(1), claws lobed basally (as Fig. 34), a character state that has been discussed above.

Schizoprymnus (Schizoprymnus) is supported by one synapomorphy: 52(2), the suture between the second and third median tergites is absent medially (Fig. 124A). This is a character state found only in the spedes of this genus. The sister group of Schizoprymnus (Muirie/la) is hypothesized to be Urosigalphus, a phylogenetic relationship that is supported by two synapomorphies. The first one, is the 14(1}, occipital carina ending ventrally on subgenal carina (as Fig. 2a), which is a charaeter state occurring in these two taxa within the tribe • 31 Brachistini, but which is also found in Cenocoelius (Cenocoeliini). The other synapomorphy is 53(1), in which the posterior and lateral margins of third median • tergite have a flanged margin (Fig. 121). The latter synapomorphy evolved independently for the species of Polydegmon. The aforementioned sister groups possess two synapomorphies with their hypothesized ancestor, including: 31 (0), four or more scutellar sulci (Fig. 29A); and 52{O), a suture between second and third median tergites that are very weak and shallow, and which are often almost completely absent (Fig. 121 A). the phytogenetic sister relationship suggests that the subgenus Schizoprymnus (Muiriella) is more closely related to Urosigalphus than it is to the genus Schizoprymnus where it had been praced by Papp (1984).

Urosigalphus is supported by two synapomorphies; the first being 10(3), where the lower margin of the clypeus is straight (Fig. 9); this is a derived state that is only found in the species of this genus within the tribe, in contrast to most Brachistini that have a slightly curved lower margine The other synapomorphy is • 13(1), in which the temples are raised laterally (as Fig. 17), a character state that is also found in Nealiolus and Niteobrachis. The species of Urosigalphus also share a plesiomorphy: 3(0), an anterior ocelfus on the frontal cavity of the head (as Fig. 21 B).

Schizoprymnus (Muiriella) is supported by one synapomorphy: 34(1), a curved lateral longitudinal carina of the propodeum (as Fig. 102). The species of S. (Muiriella) also possess four symplesiomorphies: s 9(0), the width of the the clypeus is 2.5 ta 2.7 times wider than it is long (as Fig. 15); 30(0), the crenulation of the notauli is of a medium size (as Fig. 26); 33(0), the median longitudinal carina of the propodeum is long (as Fig. 104); and 56(0), there is the presence of a suture between the second and third laterotergites (as Fig. 1158).

• 32 The genus Po/ydegmon is supported by five distinct synapomorphies. Firstly, • 35(1), there is the presence of posterior transverse carinae on the propodeum (as Fig. 37). This attribute has evolved independently in Eubazus (AHo/us), NeaHo/Os, Foersteria and Niteobrachis. Next, 42(1), is the absence of lateral longitudinal carinae on first median tergite (as Fig. 1238). Thirdly, 46(1), the second median tergite is partially delimited trom its laterotergite by a lateral crease (Fig. 1188); also, 53(1), the posterior and lateral margins of the third median tergite have a flanged margin (Fig. 1188); and 1astiy, 57(1), the second and third laterotergites coverthe sterna (Fig. 118A).

Foersteria and Niteobrachis have a sister group relationship based on one synapomorphy, 38(1), in which the claws are basally lobed (as Fig. 34). These claws were independently derived in Nealio/us and Triaspis within Brachistini and in Cenocoelius (Cenocoeliini) within Helconinae; other braconids outside the subfamily Helconinae also possess claws basally lobed such as Alabagrus sp. • (Agathidinae) and Bracon (Braconinae), suggesting that this state appeared in earty Iineages as a primitive state. Foersleria and Niteobrachis also possess a symplesiomorphy, 30{O), in that the crenulation of notauli are of a medium size {Fig. 26).

Foersteria is supported by four synapomorphies, as follows: 14(2), the occipital carina reduced ventrally (Fig. 21), a derived character state that only occurs in the species of this genus; 25(1), the absence of forewing vein 2a (as Fig. 67) which is a widely spread character state in genera of Brachistini and one that supports a monophyletic group mentioned above; 46(1), where the second median tergite is partially delimited trom the laterotergite bya lateral crease (Fig. 116B); and 57(1), wherein the second and third laterotergites cover the stema

• 33 (Fig. 1168). The latter two synapomorphies have also evolved independently in • Polydegmon.

The erection of the new genus Niteobrachis is based on two synapomorphies: 33(1), a minute median longitudinal carina of the propodeum (Fig. 97), which is a derived character state that has evolved independently and supports a monophyletic group consisting of Nealiolus, Triaspis, Schizoprymnus (Schizoprymnus), UlOsigalphus and Polydegmon; and secondly 43(1), where the lateral longitudinal carinae on the first median tergite reach the posterior margin (Fig. 105). This final character state has evolved independently in Eubazus (Brachistes, E. (Calyptus) and Nealiolus.

Phylogenetic analysis of the soecies of Niteobrachis to establish their relationship • within their genus: The heuristic search branch swapping option, alternative tree-bisection­ reconnection, found 1000 minimum length trees of 175 steps, with a consistency index (CI) of 0.629, a retention index (RI) of 0.698 and a rescaled consistency index (RC) of 0.439.

The following discussion deals only with the steps placed on minimum length tree No. 522 (Fig. 4 & 4A-C) by the heuristic algorithm and starts from the most basal branch and continues ta more inclusive ones. The discussion is based on the 24 species in the genus Niteobrachis, since its relationship with other genera of Brachistini has already been discussed in the previous phytogenetic analysis. 70% of the minimum length trees had a character distribution that showed three monophyletic groups. Tree No. 522 was arbitrarily chosen to represent the latter clades. • 34 The strict consensus tree (Fig. 3) does not show resolution on the relationships • of the species; ail but three species are grouped in an unresolved polytomy. N. elegans, insignis and lucea are shawn ta have a close relationship. This polytomy implies that data are insufficient to determine a fully resolved tree; the majority of the evolutionary steps in the minimum length trees are ambiguous or lack support (Swofford and Begle, 1993).

The genus is hypothesized ta be supported by one synapomorphy, 47(1), an absence of striations and/or sculpturing on the second median tergite (Fig. 113C­ D). The most basal branch that supports Niteobrachis (Fig. 4A) also possesses two symplesiomorphies. 30(0) is the crenulation of notauli of medium size (Fig. 26); this primitive state is found in most of the species, except for N. serenus, N. carmenae and N. splendidus in which the crenulate are of a large size. The other symplesiomorphy is 52(0), where the suture between the second and third median tergites is very weak and shallow (Fig. 105); this primitive state is found • in ail species and is a diagnostic character for the genus.

The cladogram possesses three major monophyletic groups and many species that are not included in monophytetic groups including, N. delicatus, N. serenus, N. lucidus and N. fulgeo (Fig. 4 &4A).

Within a more inclusive branch stands a monophyletic group that is comprised of N. luceo and N. insignis, which is supported by one synapomorphy: 35(1), a presence of a posterior transverse carinae of the propodeum (Fig. 98). These carinae are also found in N. subtilis and N. carmenae, species where the carinae evolved independently. The sister group of this monophytetic group is hypothesized to be N. elegans. This phylogenetic relationship is not supported • 3S by a synapomorphy. N. lucea, N. insignis and N. elegans share a primitive • character state (fig. 4B), 44(0), where the posterior border of the first median tergite between the lateral longitudinal carinae is shorter than either of the two outer laterallongitudinal carinae (Fig. 112).

Within a more internai node N. clarus and N. venustus form a monophyletic group (Fig. 4 & 4A) based on a sole synapomorphy, 45(1), an absence of striations and/or sculpturing on the first median tergite lateral to lateral longitudinal cannae (Fig. 110). This derived state has evolved independently in N. luisi; it also evolved in a more inclusive branch that supports a monophyletic group formed by nine species. This group will be discussed later so that the logical progression in which the tree is being explained may continue without interruption.

• The monophyletic group formed by N. claros and N. venustus is shown to be the sister group of a paraphyletic ensemble formed by N. gustus, N. luna, N. luisi and N. luminosus (Fig. 48). Apart from N. luminosus, ail of the latter species shared a plesiomorphic state 9(0), which is a clypeus 2.5 to 2.7 times wider than it is long (Fig. 15); this state has also been hypothesized ta occur independently in N. lumen. Within a more inclusive branch, N. subtilis is not included in a clade (Fig. 4C).

A more inclusive monophyletic clade consists of nine species, including N. splendidus, N. amazonicus, N. lumen, N. ictus, N. carmenae, N. candeo, N. attonitus, N. nitidus and N. gracilis. These are ail supported by a sole synapomorphy: 45(1), an absence of striations and/or sculpturing on the first median tergite lateral to laterallongitudinal carinae (Fig. 110). Niteobrachis mica • 36 does not possess the latter derived state. Within the aforementioned described group, there is an unresolved polytomy (Fig. 4 & 4C) that consists of N.

• splendidusz N. gracilis and a clade supported by one synapomorphy: 29(1), a presence of a posterior longitudinal carina on the median lobe of mesoscutum

(Fig. 26). This posterior longitudinal carina has evolved in N. amazonicusz N. lumen, N. ictus, N. carrnenae and N. nitidus. Although, N. lumen and N. amazonicus are sister groups, this relationship is not supported by a synapomorphy (Fig. 4C).

The next inclusive branch possesses one synapomarphy: 19(1), a curved forewing vein 1m-cu (Fig. 43). This supports a manophyletic group consisting of

N. carmenaez N. candeo, N. mico, N. attonitus and N. nitidus, which are the only species in the genus Niteobrachis ta have this derived character state.

• A derived state, 29(2), has also evolved on the medium lobe of the mesoscutum in which the longitudinal carina is complete (Fig. 24) and is found in N. candeo, N. mica and N. attonitus. This complete longitudinal carina is a synapomorphy that supports the latter three species as a monophyletic group. The sister group of the latter monophyletic group is hypothesized to be N. nitidus.

• 37 KEY TO THE GENERA AND SUBGENERA OF BRACHISTINI

• (Key modified from Mason, 1974; Marsh et al., 1987; van Achterberg, 1990; Sharkey, 1997)

1 First three median tergites forming a rigid dorsal carapace covering remaining segments (Figs. 114A,8-117A,B); second and third laterotergites either subtrapezoidal or subellipsoidal, not always divided by a suture (Figs. 1148, 1158, 117B)...... 2 First three median tergites not forming a rigid dorsal carapace, not covering remaining segments, (Fig. 119A,B); second and third laterotergites subredangular, not delimited from second and third median tergites by a crease and a shallow suture between second and third laterotergites (Fig. 119B), if only second laterotergite delimited trom second median tergite by a crease, then second and • third laterotergites delimited by weil defined suture (Fig. 122) ...... ', ...... 11 2(1) First and second median tergites delimited by articulation (Figs. 105-107); first three median tergites without a flanged lateral margin (Fig. 1148); third median tergite posterior margin without projections or notches (Figs. 114A,B-115A,B). 3 First and second median tergites delimited by suture (Fig. 117A); tirst three median tergites with a flanged lateral margin (Fig. 118A); third median tergite posterior margin with projection(s) or a notch (Figs. 123B, 124B) .. 4 3(2) Lateral longitudinal carinae on first median tergite reaching articulation between tirst and second median tergites (Figs. 105­ 107); second median tergite completely delimited from second • 38 laterotergite by a crease (Figs. 1130, 1148, 1158)...... 7 Lateral longitudinal cannae on first median tergite not reaching • articulation between tirst and second median tergites or absent (Figs. 1168, 120A-B); second median tergite partially delimited trom second laterotergite by a crease (Figs. 116B, 118B)..... 9 4(2) Forewing vain 3RS deeply curved anteriorly (Figs. 66-67) 5 Forewing vein 3RS not deeply curved anteriorly (Figs. 41 ~)... . 6 5(4) First and second median tergites, and second and third median tergites delimited by a complete suture (Fig. 117A); claws lobed basally (as Fig. 34)...... 1r";éI~";~ tiélliciél1f First and second median tergites, and second and third median tergites laterally delimited by a suture (Fig. 124A); claws unlobed basally (Fig. 35) .. Schizo"rymnus (Schizo"rymnu~) Foerster 6(4) Flange of third median tergite with denticles (Fig. 121A); juncture of forewing veins C+Sc+R with pterostigma superficial (Fig. 69); hind wing œil R broad (as Fig. 95); posterior margin of third • median tergite with a notch (Fig. 121 B); outer claw of hind leg same size as inner claw (as Fig. 35) ...... Schizoprymnu~ (Afuiriel/éI) Fullélway Flange of third median tergite without denticles (Fig. 123A); juncture of forewing veins C+Sc+R with pterostigma deep (Fig. 68); hind wing œil R narrow (Fig. 96); posterior margin of third median tergite with one or two projections (Fig. 1238); outer claw of hind leg larger than inner claw (Fig. 36) ...... Uro~;gél/"hu~ Ashmeaci 7(3) Second and third laterotergites with suture (Fig. 1158); claws unlobed basally (as Fig. 35); tirst median tergites delimited from second median tergite by wide articulation (Fig. 115A) ...... Eubélzus (A/io/us) Sél1f • 39 Second and third laterotergite without suture (Figs. 1130. 1148); claws lobed basally (Fig. 34); tirst median tergites delimited trom • second median tergite by narrow articulation (Figs. 105. 114A)... 8 8(7) Second and third median tergites with striations and/or sculpturing (Fig. 114A); third median tergite delimited trom third laterotergite by a crease (Fig. 1148) Nea/io/us Mason Second and third median tergites without striations and/or sculpturing (Fig. 113C); third median tergite not delimited trom third laterotergite by a crease (Fig. 1130) ... Niteobrachis new genus 9(3) Third median tergite posterior and lateral margins with a flange (Fig. 1188); third laterotergite lower margin concave (Figs. 118A,B); hind coxa with a lateral tooth (Fig. 118B) '" " '" ,," Po/ydegmon Foerster Third median tergite posterior and lateral margins without a tlange (as Fig. 115B); third laterotergite lower margin not concave (as Fig. 115B); hind coxa without a lateral tooth 10 • 10(9) Occipital carina ending ventrally on hypostomal carina (as Figs. 6. 20); second and third laterotergites not covering sterna; hind coxae without longitudinal carina Che/ostes van Achterberg Occipital carina reduced ventrally not ending on hypostomal carina (Fig. 21); second and third laterotergites covering stema (Figs. 116A,B); hind coxa with longitudinal carina (Fig. 37ArB)...... Foersteria Szépligeti 11 (1) Second and third median tergites not delimited trom laterotergites by a crease (Fig. 119A); clypeus lower margin without a median tooth (as Fig. 9); second median tergite without striations and/or sculpturing (Fig. 119A); tirst and second median tergites delimited by narrow articulation (Fig. 119B); second and third median tergites delimited by shallow suture (Fig. 119B) '" 12 • 40 Third median tergite not delimited tram laterotergites by a crease (Fig. 122); clypeus lower margin usually with a median tooth (Fig. • 108); second median tergite with striations and/or sculpturing (as Figs. 114A, 122); tirst and second median tergites delimited by wide articulation (as Figs. 115A, 122); second and third median tergites delimited by weil defined suture (Fig. 122) ...... Eubazus (Allodorus) Foerster 12(11 ) Second and third median tergites smooth or nearly so (as Fig. 119A); epicnimial carina always reaching mesosternum (Fig. 32) ...... ' ,. '" ," '.' ,. 13 Second and third median tergites mostly sculptured (as Fig. 115A); epicnimial carina occasionally not reaching mesostemum (Fig. 33) ...... "...... Eubazus (Eubazus) Nees 13(12) Metasoma comparatively short; torewing vein 2cu-a present (Fig. 71 ), Eubazus (Brachistes) Wesmael Metasoma comparatively long; forewing vein 2cu-a present or • absent (Fig. 70) Eubazus (Calyptus) Haliday

r-'•. 41 • NrrEOBRACHIS gen. nov. Type-species: Niteobrachis insignis sp. nov. (by present designation).

Generic diagnosis: The genus Niteobrachis can be recognized by a combination of characters because no single charaeter is diagnostic for the genus. This combination includes: first three median tergites exposed completely, forming a carapace-Iike structure; first median tergite with or without subparallel longitudinal striations lateral to the longitudinal cannae; lateral longitudinal carinae of first median tergite reaching articulation between first and second median tergites; striations and/or sculpturing on the second and third median tergites absent; second and third median tergites with glossy/smooth appearance; second median tergite and second laterotergite separated by a crease; third median tergite not separated from laterotergite by a crease; second and third median tergites separated by a shallow suture; second and third laterotergites fused together.

General features: The following characters, although found in Niteobrachis, may or may not be found in other genera of Brachistini and are presented here for completeness. Head: quadrate to slightly triangular in anterior view; antennal f1agellomeres from 23 to 31, long and slender basally but with the more apical segments broader than long in ail species; first and second flagellomeres usually subequal; scape short and stout; scape and pedicel nearly always differing in colar from the flagellum; vertex, temples and genal regions generally smooth; vertex either raised laterally or evenly convex when viewed posteriorly; occipital carina completely reaching hypostomal carina; frons divided by a frontal carina; face either nitidus with fine setiferous punctures or with rugose to coarse

• 42 appearance, sometimes heavily setose, face usually with curved parallel sculpturing below the antennal sockets; clypeus elongate to almost round, either • punctate or rugose with a fiat to curved surface; inter-tentorial line longer than ocular-tentorial line; mandibles bidentate and overlapping, upper tooth longer than lower: maxillary and labial palpi with 6 and 3 segments respectively. Mesosoma: in general, sparsely pubescent. Pronotum with polymorphie sculpturing and carinae. Notaulus impressed with crenulations of variable size. crenulation sometimes absent in anterior region. Median lobe of mesoscutum usually with a flat median longitudinal carina. Propodeum: with a median areola sometimes divided transversely; propodeal median longitudinal carina minute or nearly absent; propodeal lateral longitudinal carinae either curved or straight. Stemaulus either impressed or unimpressed, when impressed the number of crenulations varies infraspecifically. Wings: forewing veins 1a and 2a present. although these veins tend to be spectral; forewing vein r-m absent (cf. Fig. 65); wing color is hyaline or dark with a smoky tinge in sorne species. Legs: as in other Helconinae, medial tibial spurs of hind leg usually less than or equal to 1/3 • length of basitarsus; upper region of tibiae usually lighter in color than the rest; lateral regions of legs sometimes darker. Metasoma: ovipositor length varies trom shorter than body length to equal or longer. Body length varies trom 1.75 mm to 5 mm.

Biology: Biological information is lacking for ail the species. however, as members of Helconinae. it is expected that the species of Niteobrachis may be koinobiont endoparasitoids of larvae of Curculionidae (Shaw and Huddleston, 1991), Cerambycidae, Bruchidae, Anobiidae and Mordellidae (Coleoptera) (Martin, 1956; Marsh, 1979; Quicke and van Achterberg, 1990; Charlet and Sei1er, 1994), Coleophoridae (Lepidoptera) (Gauld and Bolton, 1988) and Tephritidae (Diptera) (Hemandez et al., 1994). Sorne specimens of N. insignis were • 43 recorded to have emerged from Faustinus apicalis Faust 1896 (Curculionidae: Cryptorhynchinae) which is a plant feeding species and a serious pest of • cultivated plants (Borror and White. 1970). This suggests that species of Niteobrachis may have potential as bioconlrol agents of species related to F. apicalis. such as the plum curculio Conotrache/us nenuphar (Herbst). Specimens of Niteobrachis were collected at altitudes ranging from 200-2200 m; the majorities were caught between 900-1300 m.

Of the genera of Brachistini few biologiesl studies have been published only on Triaspis and Eubazus. In arder ta add understanding ta the life history of Niteobrachis, these accounts are mentioned as follows. Parnell (1964) described a fairly complete life cycle of the female parasitoid Triaspis sp. nr. obscure/la. The parasitoid locates an egg with its antennae and then oviposits into the developing egg of Bruchidius alter (Marsham) (Coleptera: Bruchidae), a procedure that takes 46 seconds. The females of T. sp. nr. obscurella were observed to oviposit into eggs in ail stages of development but no record was • made of a second oviposition into the same host egg. Pupal periods lasted 16 to 17 days; male adults were observed ta emerge from their hosts nine days later whereas females emerged a week later. Adults survived for only three weeks when fed on honey and sugar water. It is unlikely that this species overwinters in the adult stages. These braconids probably parasitize another unknown host that has an overwintering egg stage, a host that presumably develops early in the following year so that adults of Triaspis emerge in May ready ta oviposite on eggs of B. alter once again.

The bionomics of Eubazus semirugosus (Nees) have been described by Alauzet (1987). This species is an endophagous egg-Iarvae parasitoid of Pissodes notatus Fabricius (Coleoptera: Curculionidae). Eggs of P. notatus parasitized by E. semirugosus develop and hatch within nine to fourteen days of oviposition. • 44 The parasitoid has three larval instars, with the first one emerging from the host in its pupation chambar to feed extemally. Parasitized hosts are induced to • construct abnonnal pupal ceUs from which the adult parasitoid can easily eclose. The complete development of this wasp is conditional upon the speed of the host's development as mediated by climatic conditions. In laboratory settings (190 C) and in the forests of southem France (where the experiments took place during the summer months), full development required approximately two months but it was slower when the temperature was lower as in winter. E. semirugosus is known to be bivoltine (having two generations in a year; De La Torre-Bueno, 1989), the first generation emerging from April to the beginning of June and the second generation emerging trom the end of June up to mid-August.

Etymology: Niteo is Latin for glossy, referring to the second and third metasomal median tergites and Brachis is part of the tribe name Brachistini. The gender of the name Niteobrachis is masculine according to the International Code of • Zoological Nomenclature, Article 30 (b), and it is given by the noun IIBrachis".

Distribution: Ali species of Niteobrachis are known only from the New World. The specimens were collected in Costa Rica, Panama, Ecuador, Colombia, Venezuela, Peru, Bolivia and Brazil (Maps 1-2). The above distribution suggests that this genus may occur in other ragions of Central and South America from which specimens have not yet been collected.

Identification of the species: The key for the identification of Niteobrachis specimens is based on numerous well-defined morphological characters which are reliable in delimiting the species and not on such ambiguous charaders as coloration and size (Appendix •• 45 1). Male genitalia do not show variation that could be used as a delimiting character ta identify the species, however, contrary to that, the length of the • female ovipositor compared ta that of the length of body is the only genital characteristic that has enough variations ta use for species identification. Males can easily be separated trom the females by the presence of tapered fourth to seventh metasomal segments (Fig. 1138). The males are, in general, one flfth smaller in size than the females (Fig. 113A). The females can be easily separated from the males by the oval-shaped fourth to seventh metasomal segments (Fig. 1138), and by the presence of an ovipositor and its sheaths (frontispiece, Fig.113A). It is of importance to mention that the characters used for the identification of the species found in Appendix 1 involvecl the examination of specimens of bath genders.

Remarks: Because of the wing venation and/or metasomal carapace (Sharkey, 1993), species of Niteobrachis can be mistaken for other Helconinae and/or other • subfamilies of Braconidae. However, the genus differs from genera in other tribes of Helconinae such as Helconini, Cenocoeliini, Brulleini and Diospilini by the absence of forewing vein r-m (cf. Fig. 65). Additionally, it possesses a metasoma connected to the propodeum immediately above the hind cavities of the coxae, in contrast ta Cenocoeliini. Niteobrachis can be distinguished from Blacus (Blacus) (Blacini) by the absence of a margined scutellar dise and the possession of a shallow dorsope; it can be distinguished from Cheloninae by the absence of a deeply concave occiput, absence of forewing vein r-m and in general, a different forewing venation, absence of a suture between tirst and second median tergites and the smoothness of the second and third median tergites.

• 46 • KEY TO THE KNOWN SPECIES OF NITEOBRACHIS Females are distinguished by having the fourth to seventh metasomal stema oval in shape and by the presence of an ovipositor and its sheaths (Figs. 113A,B). In males, the fourth to seventh metasomal stema are tapered in shape (Fig. 1138).

1 longitudinal median carina of median lobe of mesoscutum complete (Figs. 22-23) 2 Longitudinal median carina of median lobe of mesoscutum present only posteriorly (Figs. 25-26)... 13 Longitudinal median carina of median lobe of mesoseutum absent (Figs. 27-28) ,. 19 2(1 ) Ratio of length of hind wing vein R ta 1M fram 0.2 to 0.3 (Figs. 83- 84)...... 3 Ratio of length of hind wing vein R ta 1M from 0.4 ta 0.5 (Figs. 78, • 88) 7 Ratio of length of hind wing vein R to 1M from 0.6 to 0.7 (Figs. 77, 92)...... 10 3(2) Ratio of length of hind wing vein 1r-m to 1M from 1.2 to 1.5 (Figs. 83, 95)...... 4 Ratio of length of hind wing vein 1r-m ta 1M from 0.9 to 1.1 (Figs. 78, 93)...... 6 4(3) Temple raised laterally (Fig. 17); lateral longitudinal carinae of

propodeum straight (Figs. 97r 101)...... 5 Temple evenly convex (as Fig. 18); lateral longitudinal carinae of propodeum curved (as Fig. 99)...... Niteobrachis luceo sp. nov. 5(4) Notaulus completely crenulate (as Fig. 26); clypeus very convex (as Fig. 8); posterior border of tirst median tergite between lateral • 47 longitudinal carinae shorter than posterior border lateral to lateral longitudinal carinae (Fig. 112); posterior transverse carinae of • propodeum present (Fig. 98).. Niteobrachis insignis sp. nov. Notaulus smooth anteriorly. crenulate posterioriy (as Fig. 28); clypeus fiat to weakly convex (as Figs. 9, 10A); posterior border of first median targite between lateral longitudinal carinae nearly equal to posterior border lateral to 1atera1 longitudinal carinae (as Figs. 110-111); posterior transverse carinae of propodeum absent (as Fig. 102)...... Nittte»i»l1Ichis ~l1Icilis sp. nov. 6(3) Face smooth (as Fig. 11); lateral longitudinal carinae of propodeum curved (as Fig. 99); tirst median tergite with striations external to the lateral longitudinal carinae (as Fig. 112) ...... Nittte»i»lëIclJis ~1I/~ttC) lap. nov. Face rugose (as Fig. 13); lateral longitudinal carinae of propodeum straight (as Fig. 97); first median tergite without striations external to 1atera1 longitudinal carinae (Fig. 105) . • ...... Nit~i»lëIclJis SIJlttllcJricJrlls sp. nov. 7(2) Ratio of length of hind wing vein 1r-m ta 1M from 1.2 to 1.5 (Fig. 94); posterior transverse carinae of propodeum present (as Fig. 98).... NiteobraclJis slIbtilis sp. nov. Ratio of length of hind wing vein 1r-m to 1M from 0.9 to 1.1 (Figs. 78, 93); posterior transverse carinae of propodeum absent (Fig. 97)...... a 8(7) Sternaulus with crenulae (as Fig. 31); forewing vein 1m-cu straight (Fig. 58); temple evenly convex (Fig. 18); clypeus punctate (Fig. 14)...... 9 Stemaulus without crenulae (as Fig. 30); forewing vein 1m-cu curved (Figs. 59-60); temple raised laterally (Fig. 17); clypeus

• 48 rugose or heavily pitted (as Fig. 13) ...... Niteobrachis candeo sp. nov. • 9(8) Clypeus 2.5 ta 2.7 times wider than long (as Fig. 15); posterior border of first median tergite between lateral longitudinal carinae shorter than posterior border lateral ta lateral longitudinal carinae (as Fig. 112); 1atera1 longitudinal carinae of propodeum curved (as Fig. 99). Niteobrachis e/egans 5p. nov. Clypeus 2.0 times wider than long (as Fig. 16); posterior border of first median tergite between lateral longitudinal carinae nearly equal to posterior border lateral to lateral longitudinal carinae (as Figs. 110-111); lateral longitudinal carinae of propodeum straight (as Fig. 97)...... Niteobrachis "lIsf&lS 513. nov. 10(2) Ratio of length of hind wing vein 1r-m ta 1M from 1.2 ta 1.5 (Figs. 83, 95); lateral longitudinal carinae of propodeum straight (Fig. 97)...... 11 Ratio of length of hind wing vein 1r-m to 1M from 0.7 to 0.8 (Fig. 76); • lateral longitudinal carinae of propodeum curved (as Fig. 99)...... Niteobrachis clams 513. nov. 11(10) Stemaulus with crenulae (as Fig. 31); forewing vein 2RS straight or nearly so (Fig. 63); clypeus 2.5 to 2.7 times wider than long (Fig. 15); notaulus completely crenulate (Fig. 26)...... 12 Stemaulus without crenulae (Fig. 30); forewing vein 2RS curved (Fig. 61); clypeus 2.0 times wider than long (as Fig. 16); notaulus smooth anteriorly, crenulate posterioriy (Fig. 23) ...... Niteobrachis sel1!nllS 5fJ• .,ov. 12(11 ) Forewing vein 1m-cu straight (Fig. 64); clypeus very convex and punctate (as Figs. 8, 14); temple evenly convex (as Fig. 18) ...... Niteobrachis "..nllstlls 5J)• .,ov.

". 49 Forewing vein 1m-cu curved (Fig. 42); clypeus fiat to weakly convex and rugose to heavily pitted (as Figs. 9-10A, 13); temple raised • laterally (as Fig. 17)...... Niteobrachis atlonitus sp. nov. 13(1) Median lobe of mesoseutum without raised and carina-like margin (Fig. 22); notaulus completely crenulate (Fig. 26); crenulae of notaulus medium (Fig. 24); posterior transverse carinae of propodeum absent (Fig.97) ,, 14 Median lobe of rnesoseutum with raised and carina-like margin (Fig. 25); notaulus smooth anteriorly, crenulate posteriorly; crenulae of notaulus large (Fig. 25); posterior transverse carinae of propodeum present (as Fig. 98)...... Niteobraehis earmenae sp. noy. 14(13) Ratio of length of hind wing vein 1r-m to 1M from 1.2 to 1.5 (Figs. 83, 95) 15 Ratio of length of hind wing vein 1r-m to 1M from 0.9 to 1.1 (Figs. 78, 93) 16 Ratio of length of hind wing vein 1r-m to 1M from 0.7 to 0.8 • (Figs.76-77) 18 15(14) Stemaulus with crenulae (as Fig. 31); forewing vein 1m-cu straight (Fig. 41); ratio of length of hind wing vein R to 1M from 0.2 to 0.3 (Fig. 72); ovipositor shorter than body length (as Fig. 125) .

...... Nit~lJlëlehis alJJa~olJic:L1s tsp. 110Y. Sternaulus without crenulae (as Fig. 30); forewing vein 1rn-cu curved (Fig. 60); ratio of length of hind wing vein R to 1M trom 0.4 ta 0.5 (Fig. 91); ovipositor equal ta or longer than body length (Fig. 125) . Niteobraehis nitidLls sp. noy. 16(14) Stemaulus with crenulae (as Fig. 31); hind wing vein cu-a curved at least in posterior 1/5 (Figs. 81, 86) 17 Sternaulus without crenulae (as Fig. 30); hind wing vain cu-a straight (Fig. 82)...... Niteobraehis ietLls sp. noy.

50 17(16) Clypeus 2.5 ta 2.7 times wider than long (as Fig. 15); ratio of length • of hind wing vein R ta 1M from 0.4 ta 0.5 (Fig. 88); temple evenly convex (as Fig. 18); clypeus punctate (as Fig. 14) .

...... l\tit~iJlëlc:"i!f ICllJ7i"()~&I~ !SI). 110". Clypeus 2.0 times wider than long (as Fig. 16); ratio of length of hind wing vein R to 1M from 0.2 to 0.3 (Fig. 87); temple raised laterally (as Fig. 19); clypeus rugose or heavily pitted (as Fig. 13)...

.. NitecJiJrac:his I&lmen SI). 110". 18(14) Stemaulus with crenulae (as Fig. 31); ratio of length of hind wing vein R to 1M from 0.2 to 0.3 (Fig. 86); posterior border of first median tergite between lateral longitudinal carinae nearly equal to posterior border of tirst median tergite lateral to lateral longitudinal carinae (as Figs. 110-111); lateral longitudinal carinae of

propodeum curved (as Fig. 99) ...... N;t~iJrachi~ I&li~i !sp. 110V• Stemaulus without crenulae (Fig. 30); ratio of length of hind wing • vein R to 1M from 0.6 to 0.7 (Fig. 77); posterior border of tirst median tergite between lateral longitudinal carinae shorter than posterior border of tirst median tergite lateral to lateral longitudinal carinae (as Fig. 112); lateral longitudinal carinae of propodeum

straight (as Fig. 101 ) '. Nitee»iJlëlc:hi!f delic:at&l!f Spa 110". 19(1 ) Ratio of length of hind wing vein R to 1M from 0.2 to 0.3 (Fig. 89); temple evenly convex (as Fig. 18); face punctate (as Fig. 12) .

...... Nit~brach;~I&lna Spa 110V. Ratio of length of hind wing vein R to 1M from 0.4 ta 0.5; temple raised laterally (Fig. 17); face rugose (Fig. 13)..... 20 20(19) Forewing vein 1m-cu straight (Fig. 54); ratio of length of hind wing vein 1r-m to 1M tram 0.9 to 1.1 (Fig. 85); clypeus 2.0 limes wider than long (as Fig. 16); notaulus smooth anteriorly, crenulate • 51 posterioriy (Fig. 28)...... Niteobl'8chis lucidus sp. nov. Forewing vein 1m-cu curved (Fig. 59); ratio of length of hind wing • vein 1r-m to 1M from 1.2 to 1.5 (Fig. 90); clypeus 2.5 to 2.7 times wider than long (as Fig. 15); notaulus completely crenulate (as Fig. 26) _...... Niteobrachis mico sp. nov.

•

•• 52 SPECIES DESCRIPnONS

• Note: The bold face phrases or sentences found in the following desaiptions of the species are either the sole charader or the combination of characters that identity each species. The descriptions involved the examination of specimens belonging to males and females when bath genders were available. The order of the species desaiptions that follows is based on the phylogenetic analysis.

Niteobrachis delieatus sp. nov. (Figs. 46, 77. Map 3) Color. Dark and light brown, and yellow; dark brown as follows: hind tibia and tarsus, and ail ctaws; light brown as follows: mandible, scape. pedicel. lower pronotum and mesopleuron, middle tibia, fore and middle tarsi; yellow as follows: maxillary and labial palpi, ail coxae, trochanters and trochantelli, and femur. Head. Antenna flagellomeres with unknown number. Temple evenly convex (as Fig. 18). Clypeus 2.0 times wider than long (as Fig. 16), punctate (as Fig. 14), • very convex (as Fig. 6). Face smooth (as Fig. 11). Mesosoma. Median lobe of mesoscutum without carina on margin (as Fig. 24), longitudinal median carina present only posterioriy (as Fig. 26). Notaulus completely crenulate, crenulae medium (as Fig. 26). Stemaulus without crenulae (as Fig. 30). Posterior transverse carinae of propodeum absent, lateral longitudinal carinae straight, median areola undivided (as Fig. 97). Forewing veins: (Fig. 46) 1m-cu straight, 2RS straight or nearly so, 2cu-a with an angle of 121-125 degrees with respect to 2CUa, anterior hall of 1cu-a with an angle of 105-110 degrees with respect to A, 2RS with an angle of 121-125 degrees with respect to (Rs+M)b. Hind wing veins: (Fig. 77) cu-a curved at least in posterior 1.5. ratio of length of 1r-m to 1M trom 0.7 to 0.8, ratio of length of R to 1M trom 0.6 toO.7. Metasoma. First median tergite with striations extemal to lateral longitudinal • 53 carinae, posterior border between lateral longitudinal carinae shorter than posterior border lateral to lateral longitudinal cannae (as Fig. 108). Length of • ovipositor equal to body (as Fig. 125), with a length of 3.3 mm. Body length 3.3 mm. Male. Unknown. Host. Unknown. Distribution and material examined: Panama (Map 3). Holotype 9: PANAMA: Chirique, 2 km W Cerro Punta, 1700 m, 19.V- 8.V1.1977 (Pack & Howden) (CNe). Comments. This description is based on one specimen, which has the antennae broken off and missing a few segments, as weil as the metasoma broken off from the rest of the body, but bath pieces are attached to paper point; otherwise, the specimen is in good condition. Etymology. From the Latin delicatus meaning "delicate" due to its fragility.

Niteobrachis seTenus Spa nov. (Figs. 6,8,23,30,61,92,97, 108. Map 4) • Color. Black, dark and light brown, and yellow; black as follows: head, mesosoma; dark brown as follows: tegula, tirst to third median tergites, coxae, hind femur and tibia; light brown as follows: mandible, fore and middle femora; yellow as follows: ail trochanter and trochantelli, fore and middle tibiae and tarsi, ail claws. Head. Antenna with 29-31 flagellomeres. Temple raised laterally (Fig. 7). Clypeus 2.0 times wider than long, pundate, very convex (Fig. 6). Face punctate (Fig. 6). Mesosoma. Median lobe of mesoscutum without carina on margin, longitudinal median carina complete (Fig. 23). Notaulus smooth anteriorly crenulate posterioriy, crenulae large (Fig. 23). Stemaulus without crenulae (Fig. 30). Posterior transverse carinae of propodeum absent, lateral longitudinal carinae straight, median areola undivided (Fig. 97). Forewing veins: (Fig. 61) 1m-cu straight, 2RS curved, 2cu-a with an angle of 115-120 degrees with respect 10 2CUa, anterior hait of 1cu-a with an angle of 111-115 degrees with resped ta A, • 54 2RS with an angle of 116-120 degrees with respect to (Rs+M)b. Hind wing veins: (Fig. 92) cu-a straight, ratio of length of 1r-m to 1M trom 1.2 to 1.5. ratio of length • of R to 1M from 0.6 to 0.7. Metasoma. First median tergite with striations extemal to lateral longitudinal carinae. posterior border between lateral longitudinal carinae longer than posterior border 1atera1 to 1atera1 longitudinal carinae (Fig. 108). Length of ovipositor shorter than body (as Fig. 125), with a ratio to body length trom 0.71 to 0.80. Body length trom 3 mm ta 3.3 mm. Male. Fourth to seventh metasomal stema tapered in shape (as Fig. 113B). Host. Unknown. Distribution and material examined: Brazil (Map 4). Holotype 9: BRAZIL: Guanabara, Foresta de Tijuca. 31.1.1969 (L.& J. Stange) (AEI). Paratypes: BRAZIL: Amazonas, 71°38'W, 4°33'8, 29, 1d', IX.1979 (Alvarenga); Guanabara, Floresta da Tijuca, 1d. IV.1966 (Alvarenga & Seabra); Guanabara, Floresta da Tijuca, 19,31.1.1969 (L. & J. Stange) (AEI). Comments. The border of the median lobe of mesoscutum has a raised edge in • the "Floresta da Tijuca" specimens. First median tergite varying from completely smooth to having few striations on lateral and mesal regions of lateral longitudinal carinae. Etymology. From the Latin serenus meaning "clear, fair and bright" in reference to its carapace.

Niteobrachis lucidus sp. nov. (Figs. 28. 54, 85. 109. Map 5) Color. Dark brown. light brown and yeIlow; dark brown as follows: head. mesosoma. hind tibia. hind tarsus. ail claws. median tergites, ovipositor sheath; light brown as follows: clypeus, mandible, antenna. tegula. hind distal lateral femur, laterotergites, stema. ovipositor; yellow as follows: maxillary and labial palpi. fore and middle legs. hind coxa to femur. • 55 Head. Antenna with 25-30 flagellomeres. Temple raised laterally (as Fig. 17). Clypeus 2.0 times wider than long (as Fig. 16), rugose or heavily pitted (as Fig. • 13), very convex (as Fig. 8). Face rugose (as Fig. 13). Mesosoma. Median lobe of mesoseutum without carina on margin (cf. Fig. 25), longitudinal median carina absent (Fig. 28). Notaulus srnooth anteriorly, erenulate posterioriy, erenulae medium (Fig. 28). Sternaulus with crenulae (as Fig. 30). Posterior transverse carinae of propodeum absent (as Fig. 97), lateral longitudinal carinae curved (as Fig. 99), median areola undivided (as Fig. 97). Forewing veins: (Fig. 54) 1m-cu straight, 2RS straight or nearly so, 2cu-a with an angle of 136-140 degrees with resped to 2CUa, anterior half of 1cu-a with an angle of 121-125 degrees with respect to A, 2RS with an angle of 116-120 degrees with respect to (Rs+M)b. Hind wing veins: (Fig. 85) cu-a curved at least in posterior 1.5, ratio of length of 1r-m to 1M trom 0.9 to 1.1, ratio of length of R to 1M trom 0.4 ta 0.5. Metasoma. First median tergite with striations external to lateral longitudinal carinae (as Fig. 108), posterior border between laterallongitudinal carinae longer • than posterior border lateral to laterallongitudinal carinae (as Fig. 108). Length ot ovipositor equal to or longer than body (as Fig. 125), with a ratio to body length trom 1.0 to 1.2. Body length tram 3mm to 4mm. Male. Unknowo. Host. Unknown. Distribution and material examined: Peru (Map 5). Holotype 9: PERU: Quincemil, IX.1962 (R.D. Shenefelt) (AEI). Paratypes: PERU: Cuincemil, 19, 2C-30.X.1962, 10-15.XI.1962 (R.D. Shenefelt) (AEI). Comments. N. Lucidus is the only known species of this genus that presents the above combination of charaders written in bold letters. Etymology. From the Latin lucidus meaning "luminous" due to its shining metasomal median tergites.

• 56 Niteobrachls fulgeo sp. nov. (Figs. 48, 79. Map 6) • Color. Black, dark and light brown, and yellow; black as follows: head, mesosoma, median tergites; dark brown as follows: antenna, distal lateral hind femur, ail claws, laterotergites, stema, and ovipositor sheath; light brown as follows: clypeus, mandible, scapa, pedicel and ovipositor; yellow as follows: maxillary and labial palpi, tegula, fore leg, middle leg, hind coxa, trochanter and trochantellus, femur and proximal tibia. Head. Antenna with 29 flagellomeres. Temple raised laterally (as Fig. 17). Clypeus 2.5 to 2.7 times wider than long, punctate (as Fig. 14), very convex (as Fig. 8). Face smooth (as Fig. 11). Mesosoma. Median lobe of mesoscutum without carina on margin (as Fig. 24), longitudinal median carina complete (as Fig. 26). Notaulus completely crenulate, crenulae medium (as Fig. 26). Stemaulus with crenulae (as Fig. 31). Posterior transverse carinae of propodeum absent (as Fig. 97), 1atera1 longitudinal carinae curved (as Fig. 99), median areola undivided (as Fig. 97). Forewing veins: (Fig. • 48) 1m-cu straight, 2RS straight or nearly sa, 2cu-a with an angle of 131-135 degrees with respect to 2CUa, anterior half of 1cu-a with an angle of 111-115 degrees with respect to A, 2RS with an angle of 116-120 degrees with respect to (Rs+M)b. Hind wing veins: (Fig. 79) cu-a curved at least in posterior 1.5, ratio of length of 1r-m to 1M from 0.9 to 1.1, ratio of length of R to 1M from 0.2 to 0.3. Metasoma. First median tergite with striations external ta lateral longitudinal carinae (as Fig. 112), posterior border between lateral longitudinal carinae nearly equal to posterior border lateral to lateral longitudinal carinae (as Fig. 110). Length of ovipositor shorter than body (as Fig. 125), with a length of 3.00 mm. Body length 3.2 mm. Male. Unknown. Host. Unknown.

• 57 Distribution and matarial examined: Brazil (Map 6). Holotype 9: BRAZIL: Serra da Bocaina, Barreiros, 1600 m, 4-7.XI.1967 (Alvarenga and Seabra) (AEI). • Comments. Niteobrachis fulgeo resembles N. /uceo. Etymology. From the Latin fulgeo meaning ''to glow, to gleam" in reference to the glossiness of ils metasomal median tergites.

Niteobrachis luceo sp. nov. (Figs. 53, 84. Map 7) Color. Dark reddish brown and light brown; dark reddish brown as follows: scape, pedicel, head, mesosoma; light brown as follows: antenna, maxillary and labial palpi, ail legs, metasoma. Head. Antenna with 31 flagellomeres. Temple evenly convex (as Fig. 18). Clypeus 2.5 ta 2.7 times wider than long, pundate (as Fig. 14), fiat to weakly convex (as Fig. 10A). Face punctate (as Fig. 12). Mesosoma. Median lobe of mesoseutum without carina on margin, longitudinal median carina complete (as Fig. 24). Notaulus completely crenulate, crenulae • medium (as Fig. 26). Stemaulus with crenulae (as Fig. 31). Posterior transverse carinae of propodeum present (as Fig. 98), laterallongitudinal carinae curved (as Fig. 99), median areola divided (as Fig. 99). Forewing veins: (Fig. 53) 1m -cu

straight, 2RS straight or nearly SO, 2cu-a with an angle of 126-130 degrees with respect to 2CUa, anterior half of 1cu-a with an angle of 116-120 degrees with respect to A, 2RS with an angle of 116-120 degrees with respect to (Rs+M)b. Hind wing veins: (Fig. 84) cu-a curved at least in posterior 1.5, ratio of length of 1r-m to 1M from 1.2 to 1.5, ratio of length of R ta 1M from 0.2 to 0.3. Metasoma. First median tergite with striations external to lateral longitudinal carinae (as Fig. 108), posterior border between laterallongitudinal carinae shorter than posterior border lateral to lateral longitudinal carinae (as Fig. 112). Length of ovipositor longer than body (as Fig. 125), with a length of 4.7 mm. Body length 3.7 mm. • 58 Male. Unknown. Host. Unknown. • Distribution and matarial examined: Brazil (Map 7). Holotype 9: BRAZIL: Serra da Bocaina, Jose Barreiros, 1600 m, Il.1971 (F.M. Oliveira) (CNC). Comments. Niteobrachis /uceo resembles N. fu/geo. Etymology. From the Latin /uceo meaning nto be bright" due to the brightness of its carapace.

Niteobrachis insignis sp. nov. (Figs. 12,31,52,83. Map 8) Color. Black, dark and light brown; black as follows: head, mesosoma, tirst median tergite; dari< brown as follows: antenna, hind legs, second and third median tergites; light brown as follows: labrum, maxillary and labial palpi, fore and middle tarsi, ail tibial spurs, ovipositor, laterotergites, stema. Head. Antenna with 29-30 flagellomeres. Temple raised laterally (as Fig. 17). Clypeus 2.5 to 2.7 times wider than long, pundate (as Fig. 14), very convex (as • Fig. 8). Face punctate to rugose (as Figs. 13-14). Mesosoma. Median lobe of mesoscutum without carina on margin, longitudinal median carina complete (as Fig. 24). Notaulus completely crenulate, crenulae medium (as Fig. 26). Stemaulus with crenulae (Fig. 31). Posterior transverse carinae of propodeum present, lateral longitudinal carinae nearly straight, median areola divided (as Fig. 98). Forewing veins: (Fig. 52) 1m-cu straight, 2RS straight or nearly so, 2cu-a with an angle of 156...160 degrees with respect to 2CUa, anterior hait of 1cu-a with an angle of 116-120 degrees with respect to Ar 2RS with an angle of 111-115 degrees with respect to (Rs+M)b. Hind wing veins: (Fig. 83) cu-a curved at least in posterior 1.5, ratio of length of 1r-m to 1M frem 1.2 ta 1.5, ratio of length of R to 1M trom 0.2 ta 0.3. Metasoma. First median tergite with striations extemal to lateral longitudinal carinae, posterier border between lateral longitudinal carinae shorter than • 59 posterior border lateral to lateral longitudinal cannae (Fig. 112). Length of ovipositor longer than body (as Fig. 125), with a ratio ta body length trom 1.21 to • 1.5. Body length trom 4.33 mm to 5.33 mm. Male. Fourth ta seventh metasomal stema tapered in shape (as Fig. 113B). Host. Larvae of Faustinus apicalis Faust, (Curculionidae: Cryptorrhynchinae). Distribution and material examined: Costa Rica, Ecuador, Colombia, Venezuela, Bolivia (Map 8). Holotype 9: ECUADOR: Nape, Baeza, 2000 m, Il.1979 (W.R.M. Mason) (CNC). Paratypes: COSTA RICA: Puntarenas, San Vito, Las Cruces, 29, S.VII.1983 (B. Gill); Puntarenas, Goito Oulce, 3 km SW Rincon, 10 m, 19, VI.1991 (P. Hanson); Parque Nacional Braulio Carrillo, 500 m, 84°07'W, 100 10'N, 19, 10.lV.1985 (H. Goulet & L. Masner) (CNC). ECUADOR: Nape, Huahua Sumaco, km 45 on Honin-Loreto road, 19 2<1, Malaise trap, 14-20.XII.1989, 20.XII.1989 (M.J. Wasbauer & H. Real). COLOMBIA: Cundinamarca, Anolaima 19 2

Niteobrachis e/egans sp. nov. Caver plate. (Figs. 34, 47, 78. Map 9) Color. Black, dark and light brown; black as follows: head, mesosoma and tirst median tergite; dark brown as follows: antenna, ail coxae, femur, tibia, second and third median tergites; light brown as follows: labrum, mandible, maxillary and labial palpi, fore and middle tarsi, ail tibial spurs, ail claws, laterotergites, stema, and ovipositor. Head. Antenna with 27-30 flagellomeres. Temple evenly convex (as Fig. 18). Clypeus 2.5 ta 2.7 times wider than long. pundate (as Fig. 14), very convex (as Fig. 8). Face smooth (as Fig. 11). Mesosoma. Median lobe of mesoscutum without carina on margin, longitudinal median carina complete (as Fig. 24). Notaulus completely crenulate, crenulae • medium (as Fig. 26). Stemaulus with crenulae (as Fig. 31). Posterior transverse carinae of propodeum absent (as Fig. 97), lateral longitudinal carinae curved, median areola divided (as Fig. 99). Forewing veins: (Fig. 47) 1m-cu straight, 2RS straight or nearly so, 2cu-a with an angle of 136·140 degrees with respect to 2CUa, anterior half of 1cu-a with an angle of 121-125 degrees with respect to A, 2RS with an angle of 116-120 degrees with resped to (Rs+M)b. Hind wing veins: (Fig. 78) cu-a curved at least in posterior 1.5, ratio of length of 1r-m to 1M from 0.9 to 1.1, ratio of length of R to 1M tram 0.4 ta 0.5. Metasoma. First median tergite with striations external to lateral longitudinal carinae, posterior border between lateral longitudinal carinae shorter than posterior border lateral to lateral longitudinal carinae (as Fig. 108). Length ot ovipositor equal to or longer than body (as Fig. 125), with a ratio ta body length tram 1.0 to 1.36. Body length trom 3.0 mm ta 4.33 mm. • 61 Male. Color pattem very similar ta that of female but differing in having either lighter or stronger light brown hues in the laterotergites. Fourth to seventh • metasomal stema tapered in shape (as Fig. 1138). Host. Unknown. Distribution and matarial examined: Costa Rica, Venezuela (Map 9). Holotype 9: VENEZUELA: Lara, Yacambu, 1200 m, 13.V.1981 (H.K Townes) (AEI). Paratypes: COSTA RICA: Alajuela, Penas Blancas, 700m, 6d\ Malaise trap, IV.1987, V.1987 (E. Cruz). VENEZUELA: Lara, Yacambu, 1200m, 4d' 9d', 7.V.1981, 10.V.1981, 13.V.1981 (H.K Townes); VENEZUELA: Aragua, Parque Nacional Rancho Grande, 29 1er, 22.11.1970 (H. & A. Howden) (AEI) (CNC). Comments. Reticulations found in the propodeum and the length of the lateral longitudinal carinae are variable. Specimens from Rancho Grande generally have yellow legs and paler coloration. Males from Costa Rica present lighter forelegs. Etymology. From the latin elegans meaning "elegant" due to ils exquisite beauty.

NiteobTachis venustus Spa nov. • (Figs. 11, 22, 64, 95, 99, 110. Map 10) Color. Black, dari< and light brown, and yellow; black as follows: head, mesosoma; dark brown as follows: antenna, mandible, laterotergites and stema, and ovipositor sheaths; light brown as follows: scapa and pedicel, ail tibiae, tarsi and claws, and ovipositor; yellow as follows: ail coxae, trochanter and trochantelli, and femora. Head. Antenna with 25-27 flagellomeres. Temple evenly convex (as Fig. 18). Clypeus 2.5 to 2.7 times wider than long, pundate (as Fig. 14), very convex (as Fig. a). Face smooth (Fig. 11). Mesosoma. Median lobe of mesoscutum without carina on margin, longitudinal median carina complete (Fig. 22). Notaulus completely crenulate, crenulae medium (as Fig. 22). Stemaulus with crenulae (as Fig. 30). Posterior transverse carinae of propodeum absent, lateral longitudinal carinae curved, median areola • 62 divided (as Fig. 99). Forewing veins: (Fig. 64) 1m-cu straight, 2RS straight or

nearly SOI 2cu-a with an angle of 121-125 degrees with respect to 2CU., • anterior hait of 1cu-a with an angle of 121-125 degrees with respect to A. 2RS with an angle of 116-120 degrees with respect to (Rs+M)b. Hind wing veins: (Fig. 95) cu-a curved at least in posterior 1.S, ratio of length of 1r-m to 1M trom 1.2 to 1.5, ratio of length of R to 1M trom 0.6 to 0.7. Metasoma. First median tergite without striations external ta 1atera1 longitudinal carinae (as Fig. 106), posterior border between lateral longitudinal carinae nearly equal to posterior border lateral to lateral longitudinal carinae (Fig. 110). Length of ovipositor longer than body (as Fig. 125), with a ratio to body length frcm 1.1 to 1.2. Body length tram 3.3 mm to 4.0 mm. Male. Fourth to seventh metasomal stema tapered in shape (as Fig. 1138). Host. Unknown. Distribution and material examined: Costa Rica, Ecuador (Map 10). Holotype 9: COSTA RICA: Puntarenas, San Vito, Estacion Biologica Las Alturas, 1750 m, IX-XI.1992 (P. Hanson) (UWY). Paratypes: COSTA RICA: Puntarenas, San Vito, • Estacion Biologica Las Alturas, forest border, 1500 m, 69, Malaise trap, IX.1991, V.1992, VI.1992 (P. Hanson); Puntarenas, San Vito, Las Cruces, 1200 m,59, Malaise trap, 9.VII-7.VII1.1982, 17.VIII- 12.1X.1982, 1.VII.1983, VIII-IX.1988, (P. Hanson; B. Gill); Puntarenas, Peninsula de Osa, Cerro Rincon, Sierra dei Hilo, 200 m & 745 m, 19 4d', IX. 1990 (E. Ouiros); San Jose, Zurqui de Moravia, 1600 m, 29, 111.1992, VII.1992 (P. Hanson); Guanacaste, Volcan Cacao, Cerro Pedregal, 1000 m, 19, 1I-IV.1989 (1. Gauld & O. Janzen); Guanacaste, Parque Nacional Guanacaste, Estacion Pitilla, 9 km S Santa Cecilia, 700m, 19, 22.1X-14.X.1992 (C. Moraga). ECUADOR: Pichincha, near Tinalandia, 1150 m, 19, Malaisetrap, 9-13.V.1987 (B. Brown & L. Coote) (CNC, INBia, UWY). Comments. Anterior carinae of the propodeum may or may not be present. First median tergite occasionally has striations internai to and external to the lateral longitudinal carinae. The specimen trom Ecuador has a rather coarse textured face • 63 Etymology. From the Latin venustus meaning "graceful" due to the elegance of its • specimens. Niteobnlchis clams sp. nov. (Figs. 45, 76, 103. Map 11) Color. Dari< and light brown, and yellow; dark brown as follows: head, scape and pedicel slightly lighter, flagellum, hind tibia and tarsus, ail claws, mesosoma and metasoma with hues of lighter brown, ovipositor sheath; light brown as follows: middle tibial spurs and ovipositor; yellow as follows: maxillary and labial palpi, tegula, fore and middle leg, hind coxa ta tibia. Head. Antenna with 24-25 flagellomeres. Temple evenly convex (as Fig. 18). Clypeus 2.5 to 2.7 times wider than long, pundate (as Fig. 14), very convex (as Fig. 8). Face smooth (as Fig. 11). Mesosoma. Median lobe of mesoscutum without carina on margin, longitudinal median carina complete (as Fig. 24). Notaulus completely crenulate, crenulae medium (as Fig. 26). Stemaulus with crenulae (as Fig. 31). Posterior transverse • carinae of propodeum absent, lateral longitudinal cannae curved, median areola divided (Fig. 101). Forewing veins: (Fig. 45) 1m-cu straight, 2RS straight or nearly so, 2cu-a with an angle of 126-130 degrees with respect to 2CUa, anterior hait of 1cu-a with an angle of 121-125 degrees with respect to A, 2R5 with an angle of 116-120 degrees with respect to (Rs+M)b. Hind wing veins: (Fig. 76) cu-a curved at least in posterior 1.5, ratio of length of 1r-m to 1M trom 0.7 ta 0.8, ratio of length of R to 1M trom 0.6 ta 0.7. Metasoma. First median tergite without striations extemal to lateral longitudinal carinae (as Fig. 1(6), posterior border between laterallongitudinal carinae shorter than posterior border lateral to laterallongitudinal carinae (as Fig. 112). Length of ovipositor longer than body (as Fig. 125), with a ratio to body length from 1.12 to 1.25. Body length from 3.20 mm ta 3.27 mm. Male. Unknown. • 64 Host Unknown. Distribution and maleria' examined: Costa Rica (Map 11). Holotype 9: COSTA • R'CA: San Jose, Zurqui de Moravia, 1600 m, V.1992 (P. Hansen) (UWY). Paratype: COSTA RICA: Puntarenas, San Vito, Estacion Biologica Las Alturas, 1750m, 19, IX-XI.1992 (P. Hanson) (UWY). Comments. Niteobrachis clarus resembles N. gustus. Etymology. From the Latin clarus meaning "clear, bright" due ta the glossiness of its carapace.

Niteobrachis gustus sp. nov. (Figs. 50, 81. Map 12) Color. Dark and light brown, and yellow; dark brown as follows: head, mesosoma, hind tibia to tarsus, ail claws, distal lateral hind femur and metasoma; light brown as follows: clypeus, mandible, maxillary and labial palpi, ovipositor; yellow as follows: fore and middle legs, hind coxa to femur. Head. Antenna with 25 flagellomeres. Temple evenly convex (as Fig. 18). • Clypeus 2.0 times wider than long (as Fig. 16), punctate (as Fig. 14), very convex (as Fig. 8). Face smooth (as Fig. 11). Mesosoma. Median lobe of mesoscutum without carina on margin, longitudinal median carina complete (as Fig. 24). Notaulus completely crenulate, crenulae medium (as Fig. 26). Stemaulus with crenulae (as Fig. 31). Posterior transverse carinae of propodeum absent, lateral longitudinal carinae straight (as Fig. 97), median areola divided (as Fig. 99). Forewing veins: (Fig. 50) 1m-cu straight, 2RS straight or nearly so, 2cu-a with an angle of 136-140 degrees with respect to 2CUa, anterior half of 1cu-a with an angle of 116-120 degrees with respect ta A, 2RS with an angle of 126-130 degrees with respect to (Rs+M)b. Hind wing veins: (Fig. 81) cu-a curved at least in posterior 1.5, ratio of length of 1r-m to 1M from 0.9 to 1.1, ratio of length of R to 1M trom 0.4 ta 0.5. Metasoma. First median tergite with striations extemal to lateral longitudinal

.- 65 carinae (as Fig. 108), posterior border between lateral longitudinal cannae nearly equal to posterior border 1atera1 to lateral longitudinal carinae (as Fig. 110). • Length of ovipositor longer than body (as Fig. 125), with a length of 3.3 mm. Body length 3.0 mm. Male. Unknown. Host Unknown. Distribution and material examined: Costa Rica (Map 12). Holotype 9: COSTA RICA: Escazu, 23.V.1987 (H. & M. Townes) (AEI). Comments. Niteobrachis gustus resembles N. elegans. Etymology. From the Latin gustus meaning "to be fond of' due the grace of its specimens.

Niteobrachis luna sp. nov. (Figs. 5, 18, 27, 58, 89, 102. Map 13) Color. Black, dark and reddish brown. Black as foIlows: head; dark brown as follows: antenna, mesoscutum, ail claws, second and third median tergites, • 1aterotergites, stema, ovipositor sheath; reddish brown as follows: rest of body. Head. Antenna with 27 flagellomeres. Temple evenly convex (Fig. 18). Clypeus 2.0 times wider than long (as Fig. 16), rugose or heavily pitted (as Fig. 13), very convex (as Fig. 8). Face pundate (as Fig. 12). Mesosoma. Median lobe of mesoscutum without carina on margin, longitudinal median carina absent. Notaulus completely crenulate, crenulae medium (Fig. 27). Stemaulus with crenulae (as Fig. 31). Posterior transverse carinae of propodeum absent, lateral longitudinal carinae curved and sigmoidal-shaped, median areola divided (Fig. 102). Forewing vains: (Fig. 58) 1m-cu straight, 2RS straight or nearly so, 2cu-a with an angle of 136-140 degrees with respect to 2CUa, anterior hait of 1cu-a with an angle of 121-125 degrees with respect to A, 2RS with an angle of 116-120 degrees with respect to (Rs+M)b. Hind wing veins: (Fig. 89) cu-a curved at least in posterior 1.5, ratio of length of 1r-m to 1M from • 66 0.9 to 1.1, ratio oflength of R to 1Il tram 0.2 to 0.3. Metasoma. First median tergite with striations extemal to lateral longitudinal • carinae (as Fig. 108), posterior border between lateral longitudinal carinae nearly equal ta posterior border lateral ta lateral longitudinal carinae (Fig. as Fig. 110). Length of ovipositor shorter than body (as Fig. 125), with a length of 2_67 mm. Body length 4.0 mm. Male. Unknown. Host. Unknown. Distribution and matarial examined: Brazil (Map 13). Holotype 9: BRAZIL: Sao Carlos, Parque Ecologico, 29.X.1983 (A.S. Soares) (UDSC). Comments. This species has been described from the one specimen, which is distinctive when compared to other species of Niteobrachis. Etymology. From the latin luna meaning "moon" due to brightness of its carapace.

Niteobtachis lu;s; sp. nov. (Figs. 55, 86. Map 14) • Color. Dark and light brown, and yellow; dark brown as follows: head, mesosoma, middle distal femur, hind lower lateral femur, tibia and tarsi, ail claws, ovipositor sheath; light brown as foIlows: metasoma (Iaterotergites and stema of lighter hues); yellow as follows: maxillary and labial palpi, mandibles, fore leg and middle leg. Head. Antenna with 23 flagellomeres. Temple evenly convex (as Fig. 18). Clypeus 2.0 times wider than long (as Fig. 16), punctate (as Fig. 12), very convex (as Fig. 8). Face smooth (as Fig. 11). Mesosoma. Median lobe of mesoscutum without carina on margin (as Fig. 24), longitudinal median carina present only posterioriy (as Fig. 25). Notaulus completely crenulate, crenulae medium (as Fig. 26). Stemaulus with crenulae (as Fig. 31). Posterior transverse carinae of propodeum absent (as Fig. 97), lateral longitudinal carinae curved, median areola divided (as Fig. 99). Forewing veins: • 67 (Fig. 55) 1m-cu straight, 2RS straight or nearly so, 2cu-a with an angle of 121-125 degrees with respect to 2CUa, anterior haitof 1cu-a with an angle of • 116-120 degrees with respect to A, 2RS with an angle of 121-125 degrees with resped ta (Rs+M)b. Hind wing veins: (Fig. 86) cu-a curved at least in posterior 1.5, ratio of length of 1r-m to 1M trom 0.7 to 0.8, ratio of length of R to 1M from 0.2 ta 0.3. Metasoma. First median tergite without striations extemal to lateral longitudinal carinae (as Fig. 1(6), posterior border between lateral longitudinal carinae nearly equal ta posterior border lateral to lateral longitudinal carinae (Fig. 110). Length of ovipositor longer than body (as Fig. 125), with a length of 3.47 mm. Body length trom 2.93 mm. Male. Unknown. Host. Unknown. Distribution and material examined: Ecuador (Map 14). Holotype 9: ECUADOR: Napo, Provincia Huahua, Sumaco km 45 on Hollin-Loreto road, Malaise trap, 15-22.XII.1989 (M. & J. Wasbauer, & H. Real) (UCOC). • Comments. This species resembles N. delicatus. Etymology. This species is named in honor of my brother, Luis, for his moral support throughout my studies.

Niteobrachis luminosus sp. nov. (Figs. 57, 88. Map 15) Color. Brown, light brown, yeIlow. and light yellow; brown as follows: head, mesosoma, median tergites; light brown as follows: antenna, mandible, hind tibia, tarsus and claw, ovipositor and sheath, laterotergites, and stema; yellow as follows: scape and pedicel, and tegula; light yellow as follows: maxillary and labial palpi, fore and middle legs, hind coxa, trochantellus and femur. Head. Antenna with 27-28 flagellomeres. Temple evenly convex (as Fig. 18). Clypeus 2.5 to 2.7 times widerthan long, pundate (as Fig. 14), very convex (as • 68 Fig. 8). Face smooth (as Fig. 11). Mesosoma. Median lobe of mesoscutum ~thout carina on margin (as Fig. 24), • longitudinal median carina present only posteriorly (as Fig. 25). Notaulus completely crenulate, crenulae medium (as Fig. 26). Stemaulus with crenulae (as Fig. 31). Postenor transverse carinae of propodeum absent (as Fig. 97), laterallongitudinal carinae curved, median areola divided (as Fig. 99). Forewing veins: (Fig. 57) 1m-cu straight, 2RS straight or nearly so, 2cu-a with an angle of 136-140 degrees with respect ta 2CUa, anterior hait of 1cu-a with an angle of 105·110 degrees with respect to A, 2RS with an angle of 116-120 degrees with respect to (Rs+M)b. Hind wing veins: (Fig. 88) cu-a curved at least in posterior 1.5, ratio of length of 1r-m to 1M from 0.9 to 1.1, ratio of length of R to 1M from 0.4 to 0.5. Metasoma. First median tergite with striations external to lateral longitudinal cannae (as Fig. 10S), posterior border between lateral longitudinal carinae nearly equal to posterior border lateral ta lateral longitudinal carinae (as Fig. 110). Length of ovipositor equal ta or longer than body (as Fig. 125), with a ratio to body length from 1.12 to 1.27. Body length from 3.27 mm to 3.40 mm. Male. Unknown. Host. Unknown. Distribution and material examined: Venezuela (Map. 15). Holotype 9: VENEZUELA: Aragua, Tiara, 1200 m, 2-3.VII.1994 (V. Sabrini & C. Rosales) (UCV). Paratype: VENEZUELA: Aragua, Tiara, 1200 m, 19, 2-3.VII.1994 (V. Sabrini & C. Rosales) (UCV). Comments. Niteobrachis luminosus resembles N. ictus. Etymology. From the Latin luminosus meaning "full of light" due to luster of its metasomal median tergites.

Niteobrachis subtilis sp. nov. (Figs. 63, 94. Map 16) Color. Black, dark and light brown, and yellow; black as follows: head; dark brown

• 69 as follows: antenna, mesosoma, hind tibia, first and flfth tarsomeres, ail daws first median tergite, ovipositor sheath; light brown as follows: clypeus, mandible, hind • tibial spurs, hind basitarsus, second and third median tergites, hind second to fourth tarsomeres, stema, ovipositor; yellow as follows: maxillary and labial palpi, fore and middle legs. Head. Antenna with 22-23 flagellomeres. Temple raised laterally (as Fig. 17). Clypeus 2.5 to 2.7 times wider than long, punctate (as Fig. 14), very convex (as Fig. 8). Face rugose (as Fig. 13). Mesosoma. Median lobe of mesoscutum without carina on margin (as Fig. 24), longitudinal median carina complete (as Fig. 22). Notaulus completely crenulate, crenulae medium (as Fig. 26). Stemaulus with crenulae (as Fig. 31). Posterior transverse carinae of propodeum present (as Fig. 98), lateral longitudinal carinae straight (as Fig. 97), median areola undivided (as Fig. 97). Forewing veins: (Fig. 63) 1m-eu straight, 2RS straight or nearly so, 2cu--a with an angle of 138-140 degrees with respect to 2CUa, anterior hait of 1cu-a with an angle of 121-125 degrees with respect to A, 2RS with an angle of 121-125 degrees with respect • to (Rs+M)b. Hind wing veins: (Fig. 94) cu-a curved at least in posterior 1.5, ratio of length of 1r-m to 1M from 1.2 to 1.5, ratio of length of R to 1M from 0.4 to 0.5. Metasoma. First median tergite with striations external to 1atera1 longitudinal carinae (as Fig. 1OS), posterior border between lateral longitudinal carinae nearly equal ta posterior border lateral ta lateral longitudinal carinae (Fig. 110). Length of ovipositor equal to or longer than body (as Fig. 125) with a length of 2.4 mm. Body length 2.4 mm. Male. Unknown Host. Unknown. Distribution and material examined: Costa Rica (Map 16). Holotype Q: COSTA RICA: Puntarenas, San Vito, Las eruces, 2o-26.V.1988 (P. Hanson) (TAMU). Comments. Niteobrachis subtilis resembles N. splendidus .

• 70 Etymology. From the Latin subtilis meaning "delieate and slender" in reference to • the fragility of its specimens. Niteobrachis splendidus sp. nov. (Figs. 7, 19, 62, 93, 100, 105, 111. Map 17) Color. Black, dark and light brown, and yellow; black as follows: flagellum, head, median tergites; dark brown as follows: scape and padicel, mandible, clypeus, 1aterotergites, distal femur, hind tibia, ovipositor sheaths, ail claws; light brown as follows: mesosoma, fore legs, hind leg tarsus; yellow as follows: maxillary and labial palpi, ail coxae, trochanter and trochantelli. and hind femur. Head. Antenna with 26-28 flagellomeres. Temple raised laterally (Fig. 7). Clypeus 2.5 to 2.7 times wider than long, punctate (as Fig. 14) to rugose or heavily pitted (as Fig. 13), very convex (as Fig. 7). Face rugose (as Fig. 13). Mesosoma. Median lobe of mesoscutum without carina on margin (as Fig. 24), longitudinal median carina complete (as Fig. 22). Notaulus completely crenulate (as Fig. 26), crenulae medium (as Fig. 26) to large (as Fig. 26). Stemaulus with • crenulae to without crenulae (as Figs. 30-31). Posterior transverse carinae of Propodeum absent (Fig. 100), lateral longitudinal carinae straight, median areola undivided (Fig. 100). Forewing veins: (Fig. 62) 1m-eu straight, 2RS straight or nearly so to curved, 2cu-a with an angle of 161-165 degrees with respect to 2CUa, anterior half of 1cu-a with an angle of 121-125 degrees with respect to A, 2RS with an angle of 111-115 degrees with respect to (Rs+M)b. Hind wing veins: (Fig. 93) cu-a curved at least in posterior 1.5, ratio of length of 1r-m to 1M from 0.9 to 1.1, ratio of length of R to 1M tram 0.2 to 0.3. Metasoma. First median tergite without striations extemal to lateral longitudinal carinae (Fig. 105), posterior border between lateral longitudinal carinae nearly equal to posterior border lateral ta lateral longitudinal carinae (Fig. 111). Length of ovipositor either equal to or longer than body, or shorter than body (as Fig. 125); with a ratio to body length trom 1.0 to 1.1. Body length trom 3.06 to 3.8 mm.

71 Male. Fourth ta seventh metasomal stema tapered in shape (as Fig. 113B). Host. Unknown. • Distribution and matarial examined: Costa Rica, Ecuador, Colombia, Peru (Map 17). Holotype 9, COSTA RICA: San Jose, Zurqui de Moravia, 1600 m, IX-X.1993 (P. Hanson) (TAMU). Paratypes: COSTA RICA: Limon, 15, 16 km W Guapiles, 400 m, 194<3', 1V.1989 III-V.1990 (P. Hanson); Limon 4 km NE Bribri, 50 m, 19 1d', IX.XI.1989, XII.1989-1I1.1990 (P. Hansen); Puntarenas, Golfo Dulce, 3 km, SW Rincen, 10 m, 19, X.XII.1990 (P. Hanson); Puntarenas, Golfo Dulce, 10 & 24 km W Piedras Blancas, 100 & 200 m, 29 1d', VI.VII.1989, VI.VIII.1989 (P. Hanson); Puntarenas, Estacion Carara, Bijagoal, 500 m, 29, X.1989. (P. Hanson); Puntarenas, San Vito, Jardin Botanico, Las Cruces, 1200 m, 19, VIl-VIII.1988, (P. Hanson); Puntarenas, San Vito, Las Cruces, 1200 m, 19 1d', 9.VII-7.VII1.1982, 1.VII.1983 (B. Gill); Puntarenas, Parque Nacional Corcovado, Estacion Sirena, 29, Malaise trap, 21.111-21.1V-1991, III-IV.1991 , (Z. Fuentes); Peninsula de Osa, Sirena, 29, VII.1977. (D.H. Jansen); San Jose, Zurqui de Moravia, 1600 m, 29 1d', V.1992, IX-X.1993 (P. Hanson); San Jose, Parque Nacional Braulio Carrillo, 0.5 • km E tunel, 1000 m, 19, VI.1989 (P. Hansen); Parque Nacional Braulio Carrillo, 84°07' W, 100 10'N, 500 m, 1d', 10.lV.1985 (H. Goulet & L. Masner); Guanacaste, Parque Nacional Guanacaste, Estacion Pitilla, 9 km S Santa Cecilia, 700 m, 19, 22-14.X.1992 (C. Moraga); La Selva, 19, X1.1993 (J. Longine). PANAMA: Chirique, 15 km NW Hato dei Velcan 1200 m, 19, 24- 31.V.1977 (Peck & Howden); Reserva San Bias Nusagandi 9°20'N, 79°0'W, 29 19, 25-30.VII.1994, 6-13.11.1995 (J. Pickering); ECUADOR: Pichincha 47 km S, Santo Domingo, Estacion Rio Palenque, 19, 22.VII.1976 (S. & J. Pack); Pichincha. 16 km SE Santo Domingo, Tinalandia, 680 m, 2d', 15-30.V1.1975, 4.V-25.VII.1975 (S. & J. Pack); Pichincha, Tinalandia, 800 m, 1d', 2.11.1983 (Masner & Sharkey); Sucumbios, Rio Nape, Sacha Lodge, 0°05'5, 76°OS'W, 270 m, 29, Malaise trap, 20- 30.lX.1994 (P. Hibbs); COLOMBIA: Valle, Anchicaya, 19, 27.VII.1970 (H. & A. Howden); PERU: Loreto, Iquitos, Ouistococha, 1d', 8-10.11.1984 (L. Huggert); Madre de Dias, Puerto Maldonado, 19, 6-11.1.1984 (L. Huggert). (AEI, CNC, • 72 INBio, TAMU, UGCA, UWY). Comments. The following are some infrequent variations that occur within the • species: presence of pits on temples and striations on the first median tergite lateral ta lateral longitudinal carinae; color changes trom dark to light tones in most body parts; clypeus ranges trom dark to light brown and propodeum from light brown to black. Etymology. From the Latin splendidus meaning "brilliant" in reference to the smoothness and glossiness of ils carapace.

Niteobrachis amazonicus sp. nov. (Figs. 26, 41, 72. Map 18) Color. Black, dark and light brown, and yellow; black as follows: head, mesosoma, median tergites; dark brown as follows: antenna, clypeus, tegula, middle and hind coxae and femora, hind tibia, ovipositor sheath; light brown as follows: fore coxa and femur, middle and hind tarsus, ovipositor; yellow as follows: ail trochanter and trocantelli, fore tibia and tarsi. • Head. Antenna with 27-29 flagellomeres. Temple raised laterally (as Fig. 17). Clypeus 2.5 ta 2.7 times wider than long (as Fig. 15), rugose or heavily pitted (as Fig. 13), fiat ta weakly convex (as Fig. 10A). Face rugose (as Fig. 13). Mesosoma. Median lobe of mesoscutum without carina on margin (Fig. 26, cf. Fig. 25), longitudinal median carina present only posterioriy (Fig. 26). Notaulus completely crenulate, crenulae medium (Fig. 26). Stemaulus with crenulae (as Fig. 31). Posterior transverse carinae of propodeum absent, lateral longitudinal carinae straight, median ar80la undivided (as Fig. 97). Forewing veins: (Fig 41) 1m-cu straight, 2RS straight or nearly so, 2cu-a with an angle of 141-145 degrees with resped to 2CUa, anterior half of 1cu-a with an angle of 116-120 degrees with respect to A, 2RS with an angle of 106-110 degrees with respect to (Rs+M)b. Hind wing veins: (Fig. 72) cu-a curved at least in posterior 1.5, ratio of length of 1r-m to 1M trom 1.2 to 1.5, ratio of length of R to 1M trom 0.2 to 0.3. ~. 73 Metasoma. First median tergite without striations extemal to lateral longitudinal carinae (as Fig. 1(6), posterior border between laterallongitudinal carinae nearly • equal to posterior border lateral to lateral longitudinal cannae (as Fig. 110). Length of ovipositor shorter than body (Fig. 1258), with a ratio to body length trom 0.9 to 0.91. Body length trom 3.3 mm to 3.67 mm. Male. Unknown. Host. Unknown. Distribution and material examined: Brazil (Map 18). Holotype 9: BRAZIL: Estacion Rio de Janeiro, Silva Jardim, 5.111.1974 (F.M. Oliveira) (CNC). Paratype: BRAZIL: Guanabara, represa Rio Grande, 19, XII.1969 (M. Alvarenga) (AEI). Comments. Niteobrachis amazonicus resembles N. nitidus. Etymology. This species is named after the Amazon jungle where the specimens were colleded.

Niteobrachis lumen sp. nov. (Figs. 56, 87. Map 19) • Color. Dark and light brown, and yellow; dark brown as follows: head, antenna, mesosoma, ail claws, hind tibia, median tergites, ovipositor sheath; light brown as follows: scape, pedicel, clypeus, mandibles, maxillary and labial palpi, ail coxae, mid and hind femora, terga, ovipositor; yellow as follows: fore leg (tibia with light brown hues), mid and hind trochanter and trochantelli, median and hind tarsus with yellowish hues. Head. Antenna with 25 flagellomeres. Temple raised laterally (as Fig. 17). Clypeus 2.0 times wider than long (as Fig. 16), rugose or heavily pitted (as Fig. 13), fiat to weakly convex (as Fig. 10A). Face rugose (as Fig. 13). Mesosoma. Median lobe of mesoscutum without carina on margin, longitudinal median carina present only posteriorly (as Fig. 26). Notaulus completely crenulate, crenulae medium (as Fig. 26). Stemaulus with crenulae (as Fig. 31). Posterior transverse carinae of propodeum absent, lateral longitudinal carinae • 74 straight, median areola undivided (as Fig. 97). Forewing veins: (Fig. 56) 1m-cu

straight, 2RS straight or nearly SOI 2cu-a with an angle of 141-145 degrees with • respect to 2CUa, anterior hait of 1cu-a with an angle of 116-120 degrees with respect to A, 2RS with an angle of 116-120 degrees with respect ta (Rs+M)b. Hind wing veins: (Fig. 87) cu-a curved at least in posterior 1.5, ratio of length of 1r-m ta 1M tram 0.9 to 1.1, ratio of length of R to 1M from 0.2 to 0.3. Metasoma. First median tergite without striations external to lateral cannae (as Fig. 106), posterior border between lateral longitudinal cannae nearly equal to posterior border lateral to lateral longitudinal carinae (Fig. 110). Length of ovipositor shorter than body (as Fig. 125), with a length of 3.13 mm. Body length 3.67 mm. Male. Unknown. Host. Unknown. Distribution and material examined: Brazil (Map 19). Holotype 9: BRAZIL: Minas Gerais, Santa Barbara, Serra do Caraca, 1600 m, Il.1969 (F.M. Oliveira) (AEI). Comments. Niteobrachis lumen resembles N. attonitus. • Etymology. From the Latin lumen meaning "Iight" in reference to the glossiness of its metasomal median tergites.

Niteobtachis ictus sp. nov. (Figs. 51, 82. Map 20) Color. Dark and light brown, and yellow; dark brown as follows: head, mesosoma, lower hind femur, tibia, median tergites and ovipositor sheaths; light brown as follows: antenna, clypeus, mandibles, hind tarsus, laterotergites, stema, ovipositor; yellow as follows: scape and pediœl, maxillary and labial palpi, fore and middle legs, hind coxa, trochanter, trocantellus and femur. Head. Antenna with 24-26 flagellomeres. Temple raised laterally (as Fig. 17). Clypeus 2.5 ta 2.7 times wider than long, pundate (as Fig. 14), very convex (as Fig. 8). Face rugose (as Fig. 13). . • 75 Mesosoma. Median lobe of mesoscutum without carina on margin (as Fig. 24), longitudinal median carina present only posteriorly (as Fig. 26). Notaulus • completely crenulate, aenulae medium (as Fig. 26). Stemaulus without crenulae (as Fig. 30). Posterior transverse carinae of propodeum absent, lateral longitudinal carinae straight, median areola undivided (as Fig. 97). Forewing veins: (Fig. 51) 1m-eu straight, 2RS straight or nearly so, 2cu-a with an angle of 148-150 degrees with respect to 2CUa, anterior hait of 1cu-a with an angle of 121-125 degrees with respect ta A, 2RS with an angle of 106-110 degrees with respect ta (Rs+M)b. Hind wing veins: (Fig. 82) cu-a straight, ratio of length of 1r-m to 1M from 0.9 ta 1.1, ratio of length of R to 1M trom 0.4 ta 0.5. Metasoma. First median tergite without striations extemal to lateral longitudinal carinae (as Fig. 106), posterior border between laterallongitudinal carinae nearly equal to posterior border lateral ta lateral longitudinal carinae (as Fig. 110). Length of ovipositor shorter than body (as Fig. 125), with a ratio to body length from 0.7 to 0.89. Body length trom 2.87 ta 3.0 mm. Male. Unknown. • Host. Unknown. Distribution and malerial examined: Brazil (Map 20). Holotype 9: BRAZIL: Caruaru, Pemambuco, IV.1972 (M. Alvarenga) (CNe). Paratypes: BRAZIL: Jose Barreiros, Serra da Bocaina, 1650 m, 19, X1.1968 (Alvarenga & Seabra) (AEI). Comments. This species resembles N. nitidus. Etymology. From the Latin ictus meaning "sunbeam" in reference to the brilliancy of ils carapace.

Niteobrachis cannenae Spa nov. (Figs. 25,44,75. Map 21) Color. Black, yellow, reddish and dark brown; black as follows: head, propodeum, and middle terga of melasoma; yellow as follows: tarsi and tibia offore and middle

76 legs, distal fore femur, pronotum and notaulus; reddish as follows: mesopleuron; dark brown as follows: remainder of body. • Head. Antenna with 24 flagellomeres. Temple raised laterally (as Fig. 17). Clypeus 2.5 to 2.7 limes wider than long (as Fig. 15), punctate (as Fig. 14), very convex (as Fig. 8). Face rugose (as Fig. 13). Mesosoma. Median lobe of mesoscutum with raised carina-like rnargin, longitudinal median carina present only posteriorly (Fig. 25). Notaulus smooth anteriorly crenulate posteriorly, crenulae large (Fig. 25). Stemaulus without crenulae (as Fig. 30). Posterior transverse carinae of propodeum present, lateral longitudinal carinae straight, median areola divided (as Fig. 98). Forewing veins: (Fig. 44) 1m-cu curved, 2RS straight or nearly so, 2cu-a with an angle of 141-145 degrees with respect ta 2CUa, anterior hait of 1cu-a with an angle of 116-120 degrees with respect ta A, 2RS with an angle of 116-120 degrees with respect to (Rs+M)b. Hind wing vains: (Fig. 75) cu-a curved at least in posterior 1.5, ratio of length of 1r-m to 1M tram 0.9 ta 1.1, ratio of length of R to 1M trom 0.2 to 0.3. Metasoma. First median tergite without striations extemal to lateral longitudinal • carinae (as Fig. 1(6), posterior border between laterallongitudinal carinae nearly equal to posterior border lateral to lateral longitudinal carinae (as Fig. 110). Length of ovipositor shorter than body (as Fig. 125), with a length of 2.66 mm. Body length from 3.2 mm. to 3.26 mm. Male. Fourth to seventh metasomal stema tapered in shape (as Fig. 1138). Host. Unknown. Distribution and material examined: Ecuador (Map 21). Holotype 9. ECUADOR, Napo, Huahua, Sumaco km 45 on Hollin-Loreto road, Malaise trap, 2o-XII-1989, (M.J. Wasbauer & H. Real) (CNC). Paratype: ECUADOR: Napo, Puerto Misahualli, 350 m, 1d', 18-22.11.1983 (M. Sharkey) (CNC). Comments. Niteobrachis carmenae has a conspicuously broad crenulation on the notaulus.

• 77 Etymology. This species in named in honor of my mother, Carmen, for her • insightful generosity and love, for the constancy of her support and the benefit of her experience.

Niteobrachis candeo sp. nov. (Figs. 17,24,43,74,101,106. Map 22) Color. Dark and light brown; dark brown as follows: head, mesosoma, middle and hind femora, tibiae and tarsi, ail claws, metasoma, ovipositor sheath; Iight brown as follows: maxillary and labial palpi, fore leg, mid and hind trochanter and trochantelli, ovipositor. Head. Antenna with 25-27 flagellomeres. Temple raised laterally (Fig. 17). Clypeus 2.5 to 2.7 times wider than long, rugose or heavily pitted (as Fig. 13), very convex (as Fig. 8). Face rugose (as Fig. 13). Mesosoma. Median lobe of mesoscutum without carina on margin (as Fig. 26), longitudinal median carina complete. Notaulus completely crenulate, crenulae medium (Fig. 24). Sternaulus without crenulae (as Fig. 30). Posterior transverse • carinae of propodeum absent, lateral longitudinal carinae straight, median areola undivided (Fig. 101). Forewing veins: (Fig. 43) 1m-eu curved, 2RS straight or nearly so, 2cu-a with an angle of 141-145 degrees with respect to 2CUa, anterior half of 1cu-a with an angle of 121-125 degrees with respect to A, 2RS with an angle of 121·125 degrees with respect to (Rs+M)b. Hind wing veins: (Fig. 74) cu-a straight to curved at least in posterior 1.5, ratio of length of 1r-m to 1M from 0.9 to 1.1, ratio of length of R to 1M from 0.4 to 0.5. Metasoma. First median tergite without striations external ta lateral longitudinal carinae (Fig. 106), posterior border between lateral longitudinal carinae longer than posterior border lateral ta lateral longitudinal carinae (as Fig. 108). Length of ovipositor equal ta or longer than body (Fig. 125), with a ratio to body length from 1.02 to 1.2. Body length from 2.67 mm to 3.24 mm. Male. Fourth ta seventh metasomal stema tapered in shape (as Fig. 1138). • 78 Host. Unknown. Distribution and matarial examined: Ecuador, Peru (Map 22). Holotype 9: PERU: • Quincemil, lX.1962 (R.D. Shenefelt) (AEI). Paratypes: ECUADOR: Nape, Durene, 200 m, 19, 23-28.1x.19n (L. Penal. PERU, (unplaced) Avispas, 39, 20-30-lX.1962, 1-15.X.1962 (R.D. Shenefelt); Quincemil, 1er, IX.1962 (R.D. Shenefelt) (AEI, CNC). Comments. The locality IIAvispas· in Peru where sorne specimens were collected, was not found in any Gazetteer. The ward means "wasp" in Spanish, giving the impression that there may have been a misunderstanding when the rocality of the colleding site was written on the labels. Etymology. From the Latin candeo meaning "glow" due ta the luminescence of its metasomal median tergites.

Niteobnlchis mica sp. nov. (Figs. 59, 90. Map 23) Color. Dark and light brown; dark brown as follows: head, mesosoma, metasoma; • Iight brown as follows: clypeus, mandible, ail trochanter and trochantelli, fore distal femur, fore tibia, fore tarsus, mid tarsus, hind proximal tibia. Head. Antenna with 28 flagellomeres. Temple raised laterally (as Fig. 17). Clypeus 2.5 to 2.7 times wider than long (as Fig. 15), rugose or heavily pitted (as Fig. 13), flat toweakly convex (as Fig. 10A). Face rugose (as Fig. 13). Mesosoma. Median lobe of mesoscutum without carina on margin (as Fig. 24), longitudinal median carina absent (as Fig. 28). NotauJus completely crenulate, crenulae medium (as Fig. 24). Stemaulus with crenulae (as Fig. 31). Posterier transverse carinae of propodeum absent, lateral longitudinal carinae straight, median areola undivided (as Fig. 97). Forewing vains: (Fig. 59) 1m-cu curved, 2RS straight or nearly so, 2cu-a with an angle of 136-140 degrees with respect to 2CUa, anterior half of 1cu-a with an angle of 121-125 degrees with respect ta A, 2RS with an angle of 111-115 degrees with resped to (Rs+M)b. Hind wing veins: • 79 (Fig. 90) cu-a straight. ratio of length of 1r-m to 1M from 1.2 to 1.5, ratio of length of R to 1M tram 0.4 to 0.5. • Metasoma. First median tergite with striations extemal to lateral longitudinal carinae (as Fig. 108), posterior border between lateral longitudinal carinae nearly equal to posterior border lateral to laterallongitudinal carinae (as Fig. 110). Body length 4.06 mm. Female. Unknown. Host. Unknown. Distribution and material examined: Brazil (Map 23). Holotype d: BRAZIL: Nova Teutonia, 27°11'8 52°23I W, 4.X.1937 (F. Plaumann) (BMNH). Comments. This is the only species in the genus Niteobrachis to have been described trom a male specimen because it has enough attributes to distinguish it from the other species. Etymology. From the Latin mica meaning "brilliancy" because of the luster of its carapace.

• Niteobrachis attonitus sp. nov. (Figs. 13, 42, 73. Map 24) Color. Dark and light brown, light yellow, and black; dari< brown as follows: maxillary and labial palpi, ail trochanter and trochantelli; light brown as follows: ail tibial spurs, fore and middle tarsi, basitarsus of hind leg, and ovipositor; a combination of dari< and light brown as follows: hind tarsus; light yellow as follows: proximal tibia of middle and hind legs; rest of the body black. Head. Antenna with 28 flagellomeres. Temple raised laterally (as Fig. 17). Clypeus 2.5 to 2.7 times wider than long, rugose or heavily pitted (Figs. 13, 15), flat to weakly convex (as Fig. 10A). Face rugose (Fig. 13). Mesosoma. Median lobe of mesoscutum without carina on margin, longitudinal median carina complete (as Fig. 22). Notaulus completely crenulate, crenulae medium (as Fig. 26). Stemaulus with crenulae (as Fig. 31). Posterior transverse • 80 carinae of propodeum absent (as Fig. 97), laterallongitudinal carinae straight (as Fig. 101), median areola undivided (as Fig. 97). Forewing veins: (Fig. 42) 1m-cu • curved, 2RS straight or nearly so, 2cu-a with an angle of 161-165 degrees with respect to 2CUa, anterior hait of 1cu-a with an angle of 121-125 degrees with respect ta A, 2RS with an angle of 116-120 degrees with respect to (Rs+M)b. Hind wing veins: (Fig. 73) cu-a straight, ratio of length of 1r-m to 1M from 1.2 to 1.5, ratio of length of R to 1M trom 0.6 to 0.7. Metasoma. First median tergite wïthout striations external to 1atera1 longitudinal carinae (as Fig. 1(6), posterior border between laterallongitudinal carinae nearly equal to posterior border lateral ta lateral longitudinal carinae (as Fig. 110). length of ovipositor equal to or longer than body (Fig. 125A), with a ratio to body length trom 1.0 to 1.25. Body length from 5.6mm to 6.0 mm. Male. Unknown. Host. Unknown. Distribution and material examined: Brazil (Map 24). Holotype 9: BRAZll: Faz, Canchim Sao Carlos, 26.X.1989 (l.A. Joaquim) (UDSC). Paratypes: BRAZll: Faz, Canchim Sao Carlos, 29 12.XII.1985, 5.11.1987 (l.A. Joaquim ) (UDSC). • 1 Comments. Niteobrachis attonitus resembles N. venustus. Etymology. From the Latin attonitus meaning "seized with inspiration" because of the attractiveness of its specimens.

Niteobrachis nitidus Spa nov. (Figs. 60, 91. Map 25) Color. Dark brown, light brown and yellow; dark brown as follows: head, mesosoma except propodeum, hind middle to distal tibia, median tergites, metasoma; light brown as follows: clypeus, mandible, hind tarsus, propodeum; yellowas foIlows: maxillary and labial palpi, fore and middle legs, hind coxa to proximal tibia. Head. Antenna with 26 flagellomeres. Temple raised laterally (as Fig. 17) Clypeus • 81 2.5 to 2.7 times wider than long (as Fig. 15), rugose or heavily pitted (as Fig. 13), fiat to weakly convex (as Fig. 10A). Face rugose (as Fig. 13). • Mesosoma. Median lobe of mesoscutum without carina on margin (as Fig. 24), longitudinal median carina present only posteriorly (as Fig. 26). Notaulus completely crenulate, crenulae medium (as Fig. 26). Stemaulus without crenulae (as Fig. 30). Posterior transverse cannae of propodeum absent, 1atera1 longitudinal carinae straight, median areola undivided (as Fig. 97). Forewing veins: (Fig. 60) 1m-cu curved, 2RS straight or nearly so, 2cu-a with an angle of 146-150 degrees with respect to 2CUa, anterior hait of 1cu-a with an angle of 116-120 degrees with respect to A, 2RS with an angle of 116-120 degrees with respect to (Rs+M)b. Hind wing veins: (Fig. 91) cu-a straight, ratio of length of 1r-m ta 1M tram 1.2 to 1.5, ratio of length of R ta 1M trom 0.4 to 0.5. Metasoma. First median tergite without striations extemal to lateral longitudinal carinae (as Fig. 106), posterior border between laterallongitudinal carinae nearly equal ta posterior border lateral to lateral longitudinal carinae (as Fig. 110). Length of ovipositor equal to or longer than body (as Fig. 125), with a length of • 3.0 mm. Body length 3.0 mm. Male. Fourth to seventh metasomal stema tapered in shape (as Fig. 113B). Host. Unknown. Distribution and material examined: Brazil (Map 25). Holotype 9, BRAZIL: Jose Barreiros, Serra da Bocaina, 1650 m, XI.1968 (Alvarenga & Seabra) (AEI). Paratype. ARGENTINA: Salta, Tartagal, 1d', 11.V1I1.1973 (C. Porter) (IML). Comments. In this species the conspicuosly rugose clypeus which separates it easily from N. ictus, in addition to the combination of diagnostic charaeters above written in bold letters. Etymology. Derived trom the latin nitidus meaning "bright, gleaming" in reference to the reflection of light that its carapace produces.

• 82 Nlteobl'llchls gTaci/is sp. nov. (Figs. 14. 49, 80. Map 26) • Color. Dark brown. light brown, and yellow; dark brown as follows: head, propodeum. fore and hind coxae, distal hind femur, hind tibia, hind tarsus, ail claws. median tergites, ovipositor sheath; light brown as follows: clypeus, mandible, mesosoma except propodeum, middle coxa, middle tibia, middle tarsus, laterotergites, tergal ovipositor; yellow as follows: maxillary and labial palpi, fore leg except coxal middle trochanter and trochantellus. hind trochanter and trochantellus. proximal hind femur. Head. Antenna with 27 flagellomeres. Temple raised laterally (as Fig. 17). Clypeus 2.5 ta 2.7 times wider than long, pundate (as Fig. 14), flat to weakly convex (as Fig. 10A). Face rugose (as Fig. 13). Mesosoma. Median lobe of mesoscutum without carina on margin (as Fig. 24), longitudinal median carina complete (as Figs. 22-23). Notaulus smooth anteriorly crenulate posteriorly (as Fig. 28), crenulae medium (as Fig. 26). Stemaulus with crenulae (as Fig. 31). Posterior transverse carinae of • propodeum absent. lateral longitudinal carinae straight. median areola undivided (as Fig. 97). Forewing veins: (Fig. 49) 1m-cu straight, 2RS straight or nearly so, 2cu-a with an angle of 141·145 degrees with respect to 2CUa, anterior hait of 1cu-a with an angle of 121-125 degrees with respect to A. 2RS with an angle of 116-120 degrees with respect to (Rs+M)b. Hind wing veins: (Fig. 80) cu-a curved at least in posterior 1.5, ratio of length of 1r-m to 1M from 1.2 to 1.5, ratio of length of R ta 1M from 0.2 to 0.3. Metasoma. First median tergite without striations external to lateral longitudinal carinae (as Fig. 106), posterior border between laterallongitudinal carinae nearly equal to posterior border lateral to lateral longitudinal carinae (as Fig. 110). Length of ovipositor equal to body (as Fig. 125). with a length of 4.0 mm. Body length 4.0 mm. Male. Unknown. • 83 - Host Unknown.

Distribution and malerial examined: Peru (Map 26). Holotype Q. PERU: (unplaced) • Avispas, 1-15.X1962 (R.D. Shenetelt) (AEI). Comments. Althaugh the species can be distinguished by the highlighted charaderistics above, at tirst il might be confused with N. candeo. Holotype with right antenna broken. The locality IIAvispas· in Peru was nat found in any Gazetteer. This is a ward that means "waspJl in Spanish, giving the idea that there may have been a confusion when the locality of the collecting site was written on the label. Etymology. Derived trom the Latin gracilis meaning "slim and graceful" in reference ta the graciausness that mast hymenapterans possess. •

• 84 • DISCUSSION Ph~ogenetic analyses: The phylogenetic analyses presented here must be considered preliminary; the data matrix was made of adult morphological characters only, ten species were described trom a single specimen and other species are represented by only a few specimens. However, as a systematist the author is only estimating the diversity of species in the group or fauna investigated. Answers will never be known with certainty (Wiley, 1981). The resolution of phylogenetic relationships of Niteobrachis species and the relationship of that genus with other genera of Brachistini presented in this study may be improved by including more adult morphological characters that may have been overlooked, larval morphology, behavioral characters as to host preferences, and molecular data in order to generate a more comprehensive hypothesis. In this respect, the elucidation of biological aspects of the species involved in the study could improved the hypothesis given here. Nevertheless, the results of the analyses present a • preliminary pattern of relationships among the species of Niteobrachis and other genera of Brachistini which can be tested with more data.

The most important result of the branch-and-bound cladistic analysis is the recognition of Niteobrachis as a monophyletic group. The synapomorphies that support the monophyly of this genus are as follows: a median longitudinal carina of propodeum. The length of of this carina is not only found in other genera of Brachistini such as Nealio/us, Triaspis, Schizoprymnus (Schizoprymnus), Urosiga/phus, Po/ydegmon but also in Diospilini, genus Diospilus and in Brulleini, genus Brulleia. The carina is minute compared to that of, for instance, He/con or Eubazus (AHo/us). Since the minute size varies within the species of Niteobrachis, it could be a character state to be included in a future analysis of the genus. The other synapomorphy that defines the genus Niteobrachis is the • 85 lateral longitudinal carinae on tirst median tergite that reach the posterior • margine This is a feature which serves as a diagnostic characteristic of the genus. Although the branch-and-bound search placed this derived character state as synapomorphies that evolved independently only in Nea/io/us and Eubazus (Ca/yptus) and Eubazus (Brachistes), it also occurs in Eubazus (AI/odorus), E. (Eubazus), E. (AHo/us) and Triaspis. The lateral longitudinal carinae on first median tergite are missing in Po/ydegmon, perhaps due to a convergent loss trom a distant ancestor such as Cremnops sp. or A/abagrus texanus (Agathidinae).

The genus Niteobrachis resembles Nealio/us from which its species where removed. Although both genera have the lateral longitudinal carinae on tirst median tergites reaching posterior margin and claws lobed basally, the species of Niteobrachis are shown to be more closely related to those of Foersteria, by the possession of one derived character state, a six-segmented. The species of Niteobrachis are in fact more closely related to those of Triaspis, Schizoprymnus • (Schizoprymnus) and Urosiga/phus, ail of which possess a tive-segmented palpus.

Not ail the species of Niteobrachis were shown, by the heuristic search, branch swapping option, to be ensembled in monophyletic groups. The groups of species supported by synapomorphies where those made by Niteobrachis insignis and N. /ucea which share a presence of posterior transverse carinae on the propodeum. Two other monophyletic groups, the tirst formed by Niteobrachis venustus and N. clarus, and the second formed by Niteobrachis sp/endidus, N. amazonicus, N. lumen, N. ictus, N. carmenae, N. candeo, N. attonitus, N. nitidus and N. gracilis, are supported by the same synapomorphy: the absence of striations and/or sculpturing on the first median tergite lateral to lateral longitudinal cannae, a derived character state which is shown to have

• 86 evolved independently for both groups. Niteobrachis mico is not part of this • monophyletic group since it does not share the aforementioned synapomorphy, however it is the sister group of N. attonitus relationship based on a derived characfer state, an impressed stemaulus. The rest of the species were shown not ta be a part of a monophyletic group including Niteobrachis delicatus, N. serenus, N. lucidus, N. fu/gea, N.e/egans, N. gustus, N. /una, N. luisi, N. /uminosus and N. subtilis.

Genus Niteobrachis: The new genus Niteobrachis has been taxonomically defined on the basis of metasomal characteristics and its monophyly is phylogenetically supported on the basis of synapomorphies including a minute median longitudinal carina of propodeum and the lateral longitudinal carinae on tirst median tergite which reach the posterior margin.

Due ta the surface uniformity of the carapace-like metasoma of Niteobrachis • species, their morphological divergence is subtle compared ta that of the species of Nealiolus, which are distinguished by the striations/microsculpturing of the carapace. Wing venation does not provide sufficient attributes to differentiate Niteobrachis from other genera of Brachistini. This study presents useful identifying characters associated with the forewing veins including 2RS, 1m-cu, (Rs+M)b, 2CUa, 1cu-a, 2cu-a, A, and hind wing veins including R, 1M, 1r-m and cu-a. These were found to reveal differences that identify the species.

The most important combination of taxonomie characters for the delimitation of the genus are associated with the metasoma, including: tirst three median tergites only exposed completely forming a carapace-like structure, tirst median tergite with or without subparallel longitudinal striations lateral to longitudinal • 87 carinae, lateral longitudinal carinae of first median tergite reaching articulation • between first and second median tergites; absence of striations and/or sculpturing on the second and third median tergites; second and third median tergites with glossylsmooth appearance; presence of a crease that separates the second median tergite from the second laterotergite; third median tergite without a crease separating third laterotergite; shallow suture between second and third median tergites; second and third laterotergites fused together.

The biology of the species is virtually unknown. Although Helcaninae,

particularly Brachistinir are known ta be specialized hosts of curculionids, Hernandez et al. (1994) found an unidentified specimen of Nealio/us emerging trom a dipteran concluding that other orders of insects may serve as hosts for Brachistini. An expanded collection of adults and immature stages is desirable, particularly in central South America, where additional members of the genus Niteobrachis may occur.

• Distribution of Niteobrachis: Data obtained from the observed specimens show that Niteobrachis has a broad distribution in the Neotropical Region. The genus is not known to occur in the Nearctic Region and in the narthem part ot the Neotropical Region. Material belonging to Brachistini from both the central and southem United States and northem and central Mexico were evaluated and no specimens were found to belang to Niteobrachis. However, material from Guatemala and Nicaragua was not available. As a consequence, the distribution of the genus in those areas of Central America remains unknown. Similarly, material from insect collections of the Caribbean islands was not found in the specimens reviewed. The genus is not known trom Chile but it may weil occur there. The limiting influences on the

distribution of this genus in the Neotropics are thought to include raintaU r temperature, humidity (Brown and Gibson, 1983) and the availability of hosts • 88 • (Godfray, 1994). Based on the information available from the specimens studied, the species of Niteobrâchis are known to occur at altitudes ranging from 200-2200 m., with the majority collected at 900-1300 m. They are found in Costa Rica, Panama, Colombia, Venezuela, Brazil, Ecuador and Peru (Maps 1-2) in tropical rain forests of abundant evergreen broadleaf trees where annual rainfal! can exceed 200 cm and is never less than 130 cm. This level of rainfal! coincides with high relative humidity and an annual mean temperature of 25°C. (Starr and Taggart, 1992). Sorne species of Niteobrachis are also found in deciduous broadleaf forest in the eastem regions of Ecuador and in the eastem costal regions of Brazil (Map 2) where the temperatures remain mild but rainfal! decreases during a pronounced dry season.

Based on the Neotropical distribution of Niteobrachis and Nealiolus, it is possible that Foersteria, the sister group of Niteobrachis as hypothesized by the • branch-and-bound phylogenetic analysis, once existed in present-day South America prior to the continental breakup of Pangea over 200 million years aga (Brown and Gibson, 1983; Scherer, 1988), and became isolated in what is today the Palearctic Region (Map 27).

If this is accurate, the present distribution is possibly due to the cosmopolitan presence of hosts, beetles belonging ta the family Curculionidae. Had Niteobrachis originated before the continental breakup, it may still be dispersing ta northern areas of the Neotropica1 Region and has not yet reached the Nearctic Region.

Tribe Brachistini: Evidence that the tribe Brachistini is not a monophyletic group is provided by the • 89 position of Eubazus (Allodorus), E. (Brachistes), E. (Ca/yptus) and Eubazus (Eubazus) in the most parsimonious tree in which they are shown to have • different most recent eommon ancestors.

Nine genera and subgenera of the tribe Brachistini ineluding, Eubazus (Alio/us), Nealiolus, Triaspis, Schizoprymnus (Schizoprymnus), Urosigalphus, Schizoprymnus (Muiriella), Po/ydegmon, Foersteria and Niteobrachis are elosely related and form a monophyletic group based on two synapomorphies: their tirst three median tergites cover the remaining segments and their second and third laterotergites are trapezoidal in shape. These two characteristics separate those species from the subgenera of Eubazus excluding Eubazus (Alio/us) and eorroborates that the tribe may not be monophyletie.

The genera Triaspis, Schizoprymnus (Schizoprymnus), Urosiga/phus and Schizoprymnus (Muiriella) are shawn ta be elosely related within the tribe. They possess three synapomorphies including a shorter inter-tentorial line when • eompared to the ocular-tentorial line, their first three median tergites have a flanged lateral margin, and their tirst and second median tergites are fused together.

The genus Alio/us was synonymized by van Achterberg (1990) as a subgenus of Eubazus taking into consideration one morphological character: the degree of separation of the third epipleuron (Iaterotergite) fram its tergite. The morphologieal evidenee obtained from the present research suggests that Eubazus (Atio/us) is a separate Iineage from other Eubazus subgenera which is supported by one synapomorphy, a wide articulation between tirst and second median tergites. The results suggest that the current classification of Eubazus (Alio/us) is questionable and that a complete phylogenetie analysis of the subgenera of Eubazus is required to verity its monophyly and limits for it to be • 90 • transferred to genus status again. The classification of Schizoprymnus (Muiriella) is also open to interpretation. The gènus Muiriella was synonymized by Papp (1984) as a subgenus of Schizoprymnus. However, Schizoprymnus (Muiriella) was shown to be more closely related to Urosigalphus than to Schizoprymnus (Schizoprymnus). In fact. Schizoprymnus (Muiriella) is supported as an independent lineage by a synapomorphy: a curved lateral longitudinal carinae of the propodeum. In addition, the following morphological attributes diagnose the genus taxonomically and are uniquely found in the tribe Brachistini: carapace sculpturing uniformely crenulate. lateral longitudinal carinae on first median tergite polygonally-shaped, denticulate flange of the third median tergite and, both sutures of carapace almost completely absent. Due to the discrepancies of these results with Papp's decision (1984) to synonymize Muiriella as a subgenus of Schizoprymnus, it is recommended that a thorough phylogenetic analysis be • undertaken in arder ta establish its accurate placement. Foersteria is defined ta be the sister group of Niteobrachis based on one synapomorphy: basally lobed claws. Although Triaspis and Nealiolus also possess this characteristic, these two genera are shown to be more closely related ta one another than to Niteobrachis. Foersteria is known to occur only within the Palearctic Region where its species have been collected in temperate deciduous forests. Cornelius van Achterberg (1990) cites two hosts of F. puber (Haliday); the curculionids Rhynchites spp. and Furcipes rectirostris (Linnaeus) from a paper written by Rudow (1918). There appear to be no other records on the biology of this genus. However, if this genus is indeed the sister group of Niteobrachis, as hypothesized by the branch-and-bound cladistic analysis, it may be inferred that closely related species of beetles belonging to the genus Faustinus apicalis (Curculionidae: Chryptorhynchinae) which occur in that

• 91 biogeographic realm may serve as hosts of Foersteria species.

• Because this research deals primarily with the phylogenetic revision of the genus Niteobrachis, and because other genera of Brachistini were not revised at the species level, the author is not in a position to propose a new classification for the tribe Brachistini. Revisions of additional genera will be required before revised tribal classification is feasible.

•

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Seasonal abundance and impact of the sunflower stem W88vil parasitoid, Nes/iolus curculionis (Hymenoptera: Braconidae), in the northem Great Plains. Biologica' Control, 4 (1): 26-31. CHARLET, L.O. and G.J. SElLER. 1994. Sunflower seed weevils (Coleoptera: Curculionidae) and their parasitoids trom native sunflowers (Helianthus) in the northem Great Plains. Annals of the Entomological Society of America, 87: 831­ 835. CLAUSEN, C.P. 1954. The egg-Iarval host relationship among the parasitic Hymenoptera_ Bolletino dei laboratorio di Zoologia generale e agraria della facolta agraria in Portici, 33: 119-133. • CRAWFORD, J.C. 1914. A revision of the braconid genus Urosiga/phus (Hymenoptera, Braconidae). Insecutor inscitiae menstruus, 2: 22-27. CUMMING, J. 1992. Lactie acid as an agent for macerating Diptera specimens. Fly times, No. 8, pp. 7. DALLWITZ, M.J. and T.A PAINE. 1994. INTKEY version 3.13. CSIRO, Division of Entomology. Canberra, Australia. DALLWITZ, M.J, T.A PAINE, and E.J. ZUCHER. 1993. User's guide to the DELTA system. A general system for processing taxonomic descriptions. Edition 4.01. CSIRO, Australia. 142 pp. DALLWITZ, M.J, T.A PAINE and E.J. ZURCHER. 1994. DELTA PROGRAMS. CSIRO, Division of Entomology. Canberra, Australia. DE LA TORRE-BUENO, J.R. 1989. The Torre-Bueno Glossary of Entomology. The New York Entomological Society, 1989. 840 pp. '. 95 DUDARENKO, G.P. 1974. Formation of the abdominal carapace in braconids (Hymenoptera, Braconidae) and sorne aspects of the classification of the family. • Entomologiesl Revisw, 53 (1): 80-90. THE EARTH AND MAN, A RAND MCNALLY WORLD ATlAS. 1972. Rand McNally and Company, N.Y. 359 pp. FITCH, W.M. 1971. Toward defining the course of evolution: minimum change for a specifie tree topology. Systematic Zoology, 20: 406-416. FOERSTER, A. 1862~ Synopsis der Familien und Gattungen der Braconen. Verhandlungen frtr den Naturhistorischen Verein, Preussen Rheinland Westfahlen, 19: 225-228. GAULO, 1.0. 1988. Evolutionary patterns of host utilization by ichneumonoid parasitoids (Hymenoptera: Ichneumonidae and Braconidae). Biological Journal of the Linnaean Society, 35: 351-3n. GAULD, 1. and B. BOLTON. eds. 1988. The Hymenoptera. British Museum (Natural History), Oxford University Press, Oxford. 332 pp. GIBSON, L.P. 1972a. Revision of the Genus Urosigalphus of the United States • and Canada (Hymenoptera: Braconidae). Miscellaneous Publications. Entomological Society ofAmerica, 8 (3): 85-134.

GIBSONr L.P. 1972b. Urosigalphus of Mexico and Central America (Hymenoptera: Braconidae). Miscellaneous Publications. Entomological Society ofAmerica, 8 (4): 138-157. GIBSON, L.P. 1974. South American Urosigalphus (Hymenoptera: Braconidae). Miscellaneous Publications. Entomological Society ofAmerica, 9: 201-226. GIBSON, L.P. 1982. New species of Urosigalphus (Hymenoptera: Braconidae) tram South America. Proceedings of the Entomological Society of Washington! 84 (1): 167-176.

GOOFRAYr H.C.J. 1994. Parasitoidsr behavioral and evolutionary ecology. Princeton University Press, Princeton. 473 pp.

• 96 GORDH, G. and J.C. HALL. 1979. A critical point drier used as a method of mounting insects from alcahot Entomological News, 90 (1): 57-59. • GREATHEAD, D.J. 1986. Parasitoids in classical biological control. Pages 289­ 318 in Waage. J. and O. Greathead, eds. Insect parasitoids. Academie Press, London. 389 pp. HAESELBARTH, E. 1962. Zur Biologie, entwicklungsgeschichte und Okologie von Brachistes atricomis Ratz. (Hymenoptera: Braconidae) ais eines Parasiten von Pissocles piceae (III.) (Coleoptera: Curculionidae). Zeitschrift tuer angewandte Entomologie 49: 233-289. HERNANDEZ, V., R PEREZ and R.A. WHARTON. 1994. Native parasitoids associated with the genus Anatrepha (Diptera: Tephritidae) in Los Tuxtlas, Ve~ Mexico. Entomophaga. 39(2): 171-178. HUBER, J.T. 1993. Introduction. Page 1 in Goulet, H and J.T. Huber, ads. Hymenoptera of the world: an identification guide to families. Center for Land and Biologiesl Resources Research, Agriculture Canada, Ottawa. Publication 1894/E. 668 pp. • HUBER, J.T. and M.J. SHARKEY. 1993. Structure. Pages 13-59 in Goulet, H. and J.T. Huber, eds. Hyrnenoptera of the world: An identification guide to families. Center for Land and Biologiesl Resources Research, Agriculture Canada. Ottawa. Publication 1894/E. 668 pp. INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE. 1985. International Code of Zoological Nomenclature, third edition. International Trust for Zoological Nomenclature, London. KENIS, M.. M.A. HULME and N.J. MILLS. 1996. Comparative developmental biology of populations of three European and one North American Eubazus spp. (Hymenoptera: Braconidae), parasitoids of Pissodes spp. weevils (Coleoptera: Curculionidae). Bulletin of Entomologiesl Research, 86(2): 143-153. KENIS, M. and N.J. MILLS. 1994. Parasitoids of European species of the genus Pissodes (Col: Curculionidae) a potential for the biologiesl control of Pissodes

97 strobi (Peck) in canada. Biological Control, 4 (1): 14-21 . • LASALLE, J. 1993. Parasitic Hymenoptera, biological control and biodiversity. Pages 197-215 in laSalle, J. and 1. Gauld, eds. Hymenoptera and biodiversity. CAB International, Wallingford, UK. LASALLE, J. and 1.0. GAULD. 1993. Hymenoptera: their diversidy, and their impad on the diversity of other organisms. Pages 1-26 in LaSalle, J. and 1. Gauld, eds. Hymenoptera and biodiversity. CAB International, Wallingford, UK. LUIK, A. 1994. Sorne aspects of life history and population dynamics of Pissodes piniphilus Hrbst. (Coleoptera, Cureulionidae). Eesti Teaduste Akadeemia Toimetised Bioloogia, 43 (1): 27-36. MADDISON W.P., M.J. DONOGHUE and D.R. MADDISON. 1984. Outgroup analysis and parsimony. Systematie Zoology, 33: 83-103. MADDISON W.P. and D.R. MADDISON. 1996. Maeelade. Sinauer Assoeiates, Inc. Sunderland MA. MARSH, P.M. 1979. Braeonidae. Pages 144-313 (voI.1) in Krombein, K.V., Hurd, P.D., Smith, D.R. and Burks, B.D., eds. Catalog of Hymenoptera in America North • of Mexico. Smithsonian Institution Press, Washington, D.C. 1198 pp. MARSH, P.M., S.R. SHAW and R.A. WHARTON. 1987. An identification manual for the North American genera of the Family Braconidae (Hyrnenoptera). Mernoir of the Entomologieal Society of Washington, No. 13 MARSHALL, T.A. 1887. A monograph of British Braeonidae. Part Il. Transactions of the Entomologieal Society of London, pp. 51-131. MARTIN, J.C. 1954. Notes on the genus Foersteria Szépl. and a redeseription of Foersteria laeviuscula Szépl. (Hymenoptera: Braeonidae). Canadian Entomologist, 86 (3): 116-121. MARTIN, J.C. 1956. A taxonomie revision of the Triaspidine braconid wasps of Neardie America (Hymenoptera). Canada Department of Agriculture, Ottawa. No. 965.157 pp. MARTIN, J.C. 1958. A new species of Triaspis Haliday (Hymenoptera: • 98 Braconidae) tram Ontario. Canadian Entomologist. 90 (3): 191-192. • MASNER, L. and H. GOULET. 1981. A new model of flight-interception trap for sorne hymenopterous insects. Entomological News. 92 (5): 199-202. MASOf.J, W.R.M. 1974. A generic synopsis of Brachistini (Hymenoptera: Braconidae) and recognition of the name Charmon Haliday. Proceedings of the Entomological Society of Washington, 76 (3): 235-246. MATTHEWS. R.W. 1974. Biology of Braconidae. Annual Review of Entomology. 19: 15-32. MUESEBECK, C.F.W. and L.M. WALKLEY. 1951. Hymenoptera of America north of Mexico. Synoptic catalogue. U.S.D.A. monographs. 2: 90-184 NATIONAL GEOGRAPHIC ATLAS OF THE WORLD. 1990. VI edition. National Geographie Society, Washington D.C. NEES, von Esenbeck, C.G. 1812. Ichneumonides adsciti, in genera et familias divisi. Magazin der Gesellschaft für Naturffreunde Freude Berlin, 5: 3-37. PAPP. J. 1984. First survey of the Triaspidini species of the Indo-Australian region (Hymenoptera: Braeonidae: Calyptinae). 1. The genus Triaspis Haliday. • Acta Zoologica Hungarica, 30 (1-2): 137-158. PAPP. J. 1991. First survey of the Triaspidini species of the Indo-Australian region (Hymenoptera: Braconidae: Calyptinae). II. The genus Schizoprymnus Foerster. 1. Acta Zoologiea Hungarica, 37 (1-2): 75-99. PAPP, J. 1993. First survey of the Triaspidini species of the Indo-Australian region (Hymenoptera: Braconidae: Calyptinae). III. The genus Schizoprymnus Foerster.2. Acta Zoologiea Hungarica, 39 (1-4): 129-173. PAPP, J. 1995. Taxonomic revision of C. Rondani's braconid species (Hymenoptera: Braconidae). Redia, 78 (2): 303-309. PAPP. J.• and K. MAETO. 1992. Triaspis curculiovorus sp. n. (Hymenoptera: Braconidae) from Japan, parasitizing acom weevils. Japanese Journal of Entomology, 60 (4): 797-804.

• 99 PARNELL, J.R. 1964. The parasite complex of the two seed beetles Bruchidius • ater (Marsham) (Coleoptera: Bruchidae) and Apion fuscirostre Fabrieius (Coleoptera: Curculionidae). Transactions of the Royal Entomologieal Society of London·, 116: 73-88. PRIORE, R., E. TREMBLAY and A. RACANA. 1991. Sorne parasitoids (Hymenoptera, Braeonidae) of Rhynchaenus fagi L. (Coleoptera, Cureulionidae). BoUetino dei laboratorio di Entomologia Agraria "Filippo Silvestri", 48: 35-40. QUICKE, D.L.J. and C. VAN ACHTERBERG. 1990. Phylogeny of the subfamilies of the family Braeonidae (Hymenoptera: lehneumonoidea). Zoologische Verhandelingen (Leiden), 258: 1-95. RUDOW, F. 1918. Braeoniden und ihre Wirte. EnI. Z., Frankf. a M. 32: 7-8. SAFFER, B. 1982. A systematie revision of the genus Cenocoelius (Hymenoptera: Braeonidae) in North America, including Mexico. Poiskie Pismo Entomologiczne, 52: 73-167. SCANABISSI, G. and A. ARZONE. 1992. Epidemiological research on Anthonomus rub; Herbst (Coleoptera, Curculionidae). Redia, 75 (2): 537. • SCHERER, G. 1988. The ongins of Altieinae. Pages 115-130 in Jolivet, P., E. Petitpierre and T.H. Hsiao, eds. Biology of Chrysomelidae. Kluwer Academie Publishers. 615 pp. SHARKEY, M.J. 1988. A taxonomie revision of Alabagrus (Hymenoptera: Braconidae). Bulletin of the British Museum (Natural History) Entomology series, 57 (2): 311-437. SHARKEY, M.J. 1993. Family Braconidae. Pages 362-395 in Goulet, H. and J.T. Huber, eds. Hymenoptera of the world: an identification guide to families. Center for Land and Biologieal Resources Research, Agriculture Canada, Ottawa. Publication 1894/E. 668 pp. SHARKEY, M.J. 1997. Subfamily Helconinae. Pages 261-271 in Wharton, R.A., P.M. Marsh and M.J. Sharkey, eds. Manual of the New World genera of the Family Braconidae (Hymenoptera). Journal of Hymenoptera Research (special • 100 publications. No. 1).439 pp_ SHARKEY. M.J. and RA.. WHARTON. 1997. Morphology and terminology. • Pages 19-37 in Whartonr RA, P.M. Marsh and M.J. SharkeYr eds. Manual of the New Wood genera of the Family Braconidae (Hyrnenoptera). Journal of Hymenoptera Research (special publications. No. 1).439 pp. SHAW. M.R. and HUDOLESTON, T. 1991. Classification and biology of braconid wasps (Hymenoptera: Braconidae). Vol. 7, part 11. Royal Entomological Society of London. SHAW, S.R. 1995. Braconidae. Pages 431-463 in Hanson, P.E. and I.D. Gauld, eds. The Hymenoptera of Costa Rica. Oxford University Press. Oxford. 893 pp. SHENEFELT, RD. 1970. Braconidae 2. Heliconinae, Calyptinae, Mimagathidinae, Triaspinae. Hymenopterorum Catalogus, Pars 5: 177-306. SNOFLAK, J. 1952. La monographie de Triasp;s Hal. (Sigalphus Lair.) (Hymenoptera: Braconidae) de la Tchecoslovaquie. Acta Entomologica Musei Nationalis Pragae, 28: (1950): 285-395. SOKAl, R.R. and F.J. ROHlF. 1981. Taxonomie congruence in the • Leptopodomorpha reexamined. Systematie Zoo1ogy, 30: 309-325. STARR, C. and R. TAGGART. 1992. Biology. the unity and diversity of life. Sixth edition. Wadsworth Publishing Company. 921 pp. SUAREZ, E. 1992. Galls tram the young branches of Cinnamomum cinnamomifolium (Lauraceae) inteslect with Prionomerus (Curculionidae) and its relationships with parasitoid wasps and ants. Brenesia. 38: 137-145. SWOFFORDr D.L. 1993. PAUP, Phylogenetie Analysis Using Parsimony. Laboratory of Molecular Systematics. Smithsonian Institution, Washington, D.C. SWOFFORD, D.L and D.P. BEGLE. 1993. PAUP, Phylogenetie Analysis Using Parsimony. User's manual version 3.1. Laboratory of Molecular Systernatics. Smithsonian Institution, Washington, D.C. SWOFFORD, D.L. and W.P. MADDISON. 1987. Reconstructing ancestral character states under Wagner parsimony. Mathematical Biosciences, 87: 199-

.'. 101 229. TELEN~ H.A. 1952- The OOgen and evolution of parasitism in the Parasitica • and the formation oftheir fauna in the USSR Kiev Academy of SCience, Ukranian SSR Press, 1-139. TOBIAS, V.I. 1986. Opredelitel nasekomuch evropeiskoi tsasmi SSSR vol 3. Hymenoptera, pL 4. Opred. Faune SSSR, 145: 1-501. TOBIAS, V.I. and S.A BELOKOBYLSKlY. 1983. Aberrant venation in braconids (Hymenoptera, Braconidae) and ils significance for the study of the phylogeny of thefamily. Entomological Review, 62 (2): 107-114. TOWNES~ H. 1962. Design for a malaise trap. Proœedings of the Entomological Society of Washington, 64 (4): 253-262. TOWNES, H. 1969. The genera of Ichneumonidae, part 1. Memoirs of the American Entomological Institute, No. 11. TSANKOV, G. and Z. STAlEV. 1992. Studies on parasitoids of the larvae and pupae of the curculionid Rhynchaenus fagi in the Strandzha mountains. Nauka za Gorata, 29 (3): 45-49. • TSANKOV, G. and Z. STALEV. 1995. Certain features of the biology and ecology of Eubazus minutus (Braconidae, Hym.), the commonest parasitoid of larvae and pupae of the curculionid Rhynchaenus fagi in the Strandzha ragion. Nauka za Gorata, 32 (2): 46-50. VIERECK, H.L. 1918. A Jist of families and subfamilies of Ichneumon-flies of the super-family Ichneumonoidea (Hymenoptera). Proceedings of the Biologiesl Society of Washington, 31: 69-74 WHARTON, R. A 1993. Bionomics of the Braconidae. Annual Review of Entomology, 38:121-143. WHARTON, R.A. 1997. Introdudion. Pages 1-15 in Wharton, R.A, P.M. Marsh and M.J. Sharkey, eds. Manual of the New World genera of the Family Braconidae (Hymenoptera). Journal of Hymenoptera Research (special publications. No. 1). 439 pp. • 102 WHARTON, R.A., S.R SHAW, M.J. SHARKEY, 0.8. WAHL, J.B. WOOly~ J.B. WHITFIELD, P.M. MAR8H and W. JOHNSON. 1992. Phylogeny of the • subfamilies of the family Braconidae (Hymenoptera: Ichneurnonoidea): a reassessment. Cladistics, 8 (3): 199-235. WILEY, E.O. 1981. Phytogenetics. The theory and pradice of phylogenetic systematics. John Wileyand Sons, New York. pp. 78-79.

•

• 103 •

Fig. 1. Strict consensus tree showing a phylogenetic hypothesis of the relationships of Niteobrachis and related genera. The consensus tree was derived from 120 mast parsimonious trees of 212 steps, using the branch­ and-baund search with a 1000 maximum number of trees saved. • Fig. 2.

PAUP most parsimonious tree No. 120 (cf. Figs. 3A-B) showing a phylogenetic hypothesis of the relationships of Niteobrachis and related genera. The tree of 212 steps and of a CI 0.736, was obtained using the branch-and-bound search with a 1000 maximum number of trees saved.

• • • •

N ~

ANCESTOA Cenocoellus

Cenocoelius ~ Brullela

Brullela Eubazus(Allodorus) Eubazu.(Allodorusl 0 EubazU9(Brachlslssl Eubazus(Brachlstes) Slacus(Slacus)

SIacus(BIacus) Eubazus(Calypfus)

Eubazu8(Eubazus) Eubazus(CaJyp1us) Olospllus Dlospllus Eubazus(Aliotus) /EUbaZUS(EUbaZUS) ~o '//////"( . Neatlolus Eubazus(Aliolus) Trlaspls

Neallolus Uroslgalphus

Triaspls Schlz (Mulrlella)

Schlzoprymnus Schlzoprymnus

Uroslgalphus Polydegmon

Schlz (Mulrlella) Foersterla

Nlteobrachls Poy1degmon

Nlteobrachis •

Fig.2A. PAUP most parsimonious tree No. 120. Hypothesis of relationships. •

• f .-::> ~ j !!! .. .. 1 if ::> .. ~ a:J m.. ~ .. i .!! ~ ::> .a ~ .. .. ! j .~ &J.. l .. '2 ~ ::> Il ::> .2 1 0 a:J W W m w c • ~ 2·,.0 11: 1.>0 43: 0->1 ~: 1.>0 42: 00.1 1-32: 2-.' 1.33: 0-., ~4: 0-., ~1: 0-., 10: 0-., ~: 0-.1

g: 0·>1

10: 0-.,

14: 0->1

.:t3: 0->' Q: 0->1

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• 2A lOS •

Fig. 28. PAUP most parsimonious tree No. 120. Hypothesis of relationships. •

• 1 i' 1 -! ~ . 1 l oc• ':fi i i .1 ~ ..èS• • i J îs:; f • -c• ë fQ ~ ~ f Z• ~ ~ ;:) ~ lU lU ! ~ Z 8: \-.0 \-.0 '4: 0-.2 Il: C)..I 2-.0 0..\ 0..1 1·.2 10: 0-.3 1-»0 C)..I 0-.1 0-.1 0-.\ 43: 0-.1 ,3: 0-.1 1-.0 0-.\

'4: 0-.1 1: \.~ 42: 0-.1 2: t-~ 0-.1 43: 0-.,

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• 28

106 •

Fig. 3. Strict consensus tree showing a phylogenetic hypothesis of the relationships of the species of Niteobrachis. The consensus tree was derived from 1000 minimum length trees of 175 steps, using the heuristic • search branch swapping option with the alternative tree-bisection­ reconnection (TBR).

Fig. 4. PAUP minimum length tree No. 522 (cf. Figs. 4A-C) showing a phylogenetic hypothesis of the relationships of the species of Niteobrachis. The tree of 175 steps and a CI of 0.629 was obtained using the heuristic search branch swapping option, alternative tree bisection reconnection (TBR), with a 1000 maximum number of trees saved.

• • • •

~ w

Eubazus(Brachlstes) Eubazus(Allodorus) Eubazus(Calvptus) Eubazus(Brachlstes) Eubazus(Eubazus) Eubazus(Calyptus) Foersterla Eubazus(Eubazus) Eubazus(Aliolus) Foersterla 5chlzoprymnus Polydegmon Trfaspls 8chlzoprymnus Uroslgalphus Trlaspls 8chlz (Mulrlella) Uroslgalphus Neallolus 5chlz (Mulrlella) Polydegmon Neallolus N. gracllls N. gracllls N. n!tldus N. nltldus N.lucldus N. attonltus N. lumlnosus N.mlco N.luna N.candeo N. clarus N.carmenae o- N. venustus ..... N.lctus N. dellcatus N.lumen N. elegans N. amazonlcus N.lnslgnls N. spiendIdus N.luceo N. subtllls - N. serenus N. lumlnosus N. attonltus N.lulsl N.carmenae N.luna N.lulsl N. gustus N. gustus N. clarus N. subtllls N. venustus N.lctus N. elegans N. candeo N.lnslgnls N.lumen N.luceo N. fulgea N. fulgeo N. splendldus N.lucldus N.mlco N. serenus N. amazonlcus N. dellcatus Eubazus(Atiodorus) Eubazus(Allolus) •

Fig. 4A. PAUP minimum length tree No. 522. Hypothesis of relationships. •

• !c ::!! • Z .

1

: •

t

1

9: 0->1

44: ;j!->I

1.32: 2->1

~4: 0->1

13: 0->1

44: 0->2

4A 1-30: 2->0 47: 0->1 NiteObraChls • 1.s2: '->0 108 •

Fig. 48. PAUP minimum length tree No. 522. Hypothesis of relationships. •

• fil :1 fil fil :1 fil fil c: • ~ fil ii ra C 0 ~ :1 .5P 0 2 c ~ u ë :E ! :1 0 .= .3 .3 Q ~ > Cl .5• .= Z Z, Z Z Z Z Z Z Z • 1 J 1 ! 1 1 r L.

,.35. 0-,.

45: 0-,.1

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13: 1->0 • ~7: 1->0

t • 48 109 •

Fig. 4C. PAUP minimum length tree No. 522. Hypothesis of relationships. •

• GJ • ca a • 0 ~ .a ID 0 • •~ c: N if ~ ~ ~ •~ ce • l! ~ ~ E ca c: iIII e 8 3 l! .a ce Z Z . Z Z Z Z Z Z Z

.32: 2->1

.29: t->2

1 19: 0->1 t • ~2: t->2

.29: 2->1

.5: 0->1

1 t_·:I_>O__

• 4C 110 •

Fig. 5. Niteobrachis luns, 9, lateral view; left wings removed. •

• •

10 m.

x 29 1 590 ~ •

t:1ltlt006 • 6-8-96 Bo l te ZEISS DSI19~eA

5

• III •

Fig. 6. Head, anterior view. Niteobrachis serenus.

Fig. 7. • Head, posterior view. Niteobrachis splendidus. • Pedicel Scape

Eye

Frans

Tentona' pit

Clypeus Mandible

6 • Lateral ocellus

Temple

Eye Occipital carina

Foramen magnum Occiput

Hypostomal carina Maxillalry palp

Labial palp • 7 112 •

Fig. 8. Face. Niteobrachis serenus.

Fig. 9. • Head, anterior view. Urosigalphus (Bruchiurosigalphus) bruchi. Arrow painting to lower margin of dypeus.

• • 13 •• 288

S8 .,.

8

• 15 mM x S9

.1'+S815 15-8-96 Bo l te ZEISS .. , . OSM91t8A • 9 113 •

Figs. 10A-B. Head. Eubazus (Allodorus) crassigaster. Arrows showing inter- tentorial distance, acular-tentarial distances and media-ventral taath of • lower margin af clypeus.

• • 18 •• 88

2ee ~.

10A •

• 108 114 •

Figs. 11-14. Head. 11. Niteobrachis venustus, smooth face. 12. N. insignis, punctate face. 13. N. attonitus, rugose face and clypeus. 14. N. graci/is, • punctate clypeus. Arrows showing inter-tentorial and ocular-tentoria1 distances. (scale lines, 1 mm)

Figs. 15-16. Clypeus. 15. 2.5 to 2.7 times wider than long. 16. Clypeus 2.0 times wider than long.

• •

...... -. "... :"::: ...." :.::- :: " ., . ..: :""": :: :". ::. : :.. . ..-- ê~- . ""..: ~ . " ::~ ::': ~:" ï . 11 <8

•

• 15 16 115 •

Fig. 17. Head, posterior view. Nifeobrachis candeo. Arrows pointing te temples.

Fig. 18. • Head and mesoscutum, posterior view. Nifeobrachis /una. Arrows pointing to temples.

• ." _... • 23 •• x 195

190 loi.

:1=158822 18-12-6 Bo l te ZEISS DSMSlt9A 17

• Il 16 •• -x 188

188 .,•

• 1 ....883 6-8-96 80 l te , ~ ZEISS 'f DSt1SlteA t---- • 18

116 •

Fig. 19. Head, posterior and lateroposterior view. Niteobrachis splendidus. Arraws painting ta temples.

Fig. 20. Head, lateroposterior view. Triaspis kurtogaster. Arrows painting ta • occipital carina.

• ...., ~ • 20 •• x 82 1 213e loi-

#158825 18-12-6 Bolte 5§~~~8A 19

• .63kV 1'+ ••

138

• 20 117 •

Figs.21-21A. Head, lateroposterior view. 21. Foersteria laeviuscula. Arrows pointing to occipital carina. 21 A. Cenocoelius saperdae. Arrow pointing to subgenal carinae. •

• . __"V 2... ••

S0 l-Im

#1 ... 6023 21-8-96 Bo l te ·ZE 1 SS 21

• V. . ...,"""V ~... 16 mm

S0 1 200 l-Im

#1lt600lt 21-8-96 Bo l te ZEISS • 21A 118 •

Fig. 218. Head, dorsal view. He/con tardator. Arrew painting te ocellus on the frontal cavity.

Fig. 22. • Mesosoma (pronotum omitted) dorsal view, Niteobrachis venustus.

• .~:pr

••• ':.. # • • .:Cl 17 •• =2: .. ~~::~~.. ,.. .;-. ~.. . "t.- .: 58 \. -~ ~... - .... 288 ....

• 1..5882 15-8-98 Bo l te ZEISS DSnS"'A • 218

Longitudinal carina

Median lobe of mesoscutum

Lateral lobe of mesoscutum

Notaulus

Scutellar sulcus

Metanotum

Propodeum • 22 119 Figs.23-24. Mesosoma (pronotum omitted), dorsal view. Niteobrachis serenus. Arrows pointing to longitudinal carina of median lobe of mesoscutum. 24. Niteobrachis candeo. Arrow pointing to margin median lobe of mesoscutum. •

•• • --

.1..282. 31-"-88 Boit. 11~1i'-- 23 •

eN CD~ =11= )( cnm 00) ~ ~ :1.... -1 (JI CI) (-

Fig. 25. Head and Mesoscutum, dorsal view. Niteobrachis carmenae. Arrows pointing to raised carina-like margin (cf. Fig. 24) and size of crenulation • of notaulus.

Fig. 26. Mesoscutum. Niteobrachis amazonicus. Arrows pointing to presence of longitudinal carina.

• •

25

m~

o~ ~ -1 CD· ...0) ~ IDI œ CD CD • JI:' • en CD •C

Il. .', ~.~ 23 -.- .x 98 :.... ~ 1 1 ; ."" 100

• 26 121 •

Fig. 27. Mesoscutum and metanotum, dorsal view. Niteobrachis luna. Arrow pointing to absence of longitudinal carina.

Fig. 28. • Mesoscutum, dorsal view. Niteobrachis lucidus. Arrows painting to absence of longitudinal carina and size of crenulation.

• • 15 ••

209 l-'M

~:M: 1 4lt 0 0it 6-8-96 Bo l te ·~·2E 1 SS lOIR :DSM9ltu 27 • 13 ••

x 98 299 ..,.

, .

"'~. ~\.1lt2812 . '31-7-96 .~~Bo l te "\ ZEISS __ .... ~ ...... • 28 122 •

Fig. 29. Head, posterior view, and mesoscutum, dorsal view. Polydegmon foveolatus. Arrow painting to size of crenulation of nataulus (cf. Fig. • 25).

Fig. 29A. Metanotum. Urosigalphus (Bruchiurosigalphus) bruchi. Arrows painting ta scutellar sulei.

• •

29 CJN aJ~ )( 01T1 o~ ~ 2 ..... -1 CD- (Den ~ClO .j çen fDl ( CD ~ , •~ m !I• CD • 4 • . "~I(,, 28 •• x SE 1 2ee ...

• 29A 123 •

Fig. 30. Mesosoma, partial lateral view showing mesopleuron. Niteobrachis serenus. Arrow pointing ta absence of crenulation.

Fig. 31. • Pronotum, partial lateral view, and mesospleuron. Niteobrachis insignis. Arrows pointing ta presence of crenulation on sternaulus.

• • , 13 •• 76

=l=1lt2819 3 1-7-96 8 o l te ZEISS 30 DSnSlt8A • 11 •• \':--;;~~ 8~

'_~J"'~~X~.:I!:. '. .,' -,..IIIIII!II--- J~,"~. ',288 ~ • .'f..: .- ...... J) ,t}:'-.lI''::'.~' _:( .~~ .. J ~•... ,....' . ',t // ::?~f)'~ . :.>~. ') l • /;. \\ ' -" ..------;r;'~' ~ . .... - ~ .---- J..:_ ') .1 ..2 8 15 ;'. iI~lT;98 ç ~~!IS. -- • 31 124 •

Fig. 32. Mesosoma, partial lateral view. Eubazus (Srachistes) salicicola. Arrow pointing to epicnimial carina.

Fig. 33. • Mesosoma, partial lateral view. Eubazus (Eubazus) pallipes. Arrows pointing to epicnimial carina.

• 15 •• • 86

,l~~ .~.\~>.1lf.6889 -2B 1-8-96 '" 0 l te -'-:2E~. 1 SS " ..n,...."' •. A_ 32 • 1.. iU

200

• 33 125 •

Fig. 34. Pretarsus. Niteobrachis elegans. Arrow pointing to lobe of claw.

Fig. 35. Pretarsus and apical tarsomere of Schizoprymnus (Schizoprymnus) • ambiguus. Arrow pointing to claw (scale line, 1 mm).

Fig. 36. Pretarsus and apical tarsomere of Urosigalphus (Bruchiurosigalphus) bruchi. Arrow pointing ta claw (scale line, 1 mm).

• • f IDKY • •• 158

28 .,.

·:.1~2986 &'''-:';~3 1-1-96 Bo l te ./~~!~~ftft 34 •

35 36 • 126 •

Figs.37A-B. Hind coxa of Foersferia laeviuscula. Arrows painting ta carina. •

• • 12 •• 168

188 ~.

1 ..581 .. 15-8-96 Balte

37A ~§AI~8A •

378

• 127 •

Fig. 38. Forewing of Niteobrachis species (scale line, 1 mm).

Fig. 39. Hind wing of Niteobrachis species (scale line, 1 mm). •

• 3RS M+CU 1M 1m-eu 2RS r R1a R1b • (RS+M)a

A 1a 1CUa 38 2cu-a (RS+M)b cu M • 1CUb

C+Sc+R R SC+R RS M+CU 1r-m R1

39

• A cu-a 1M 2M 128 •

Figs. 40A-B. Measurements taken in the wing venation of Niteobrachis species. 40A. Angle variations. 408. Ratio of length variations (scale line, 1 mm). •

• 2RS • 2CUa

(RS+M)b

2cu-a 40A

• 1M R 1 • 1....' 1 1 ,.1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 ~ 1 1 ~;:;~~~~~I1~~~~ 1

1M 1r-m • 408 129 Figs. 41-44. Forewings of Niteobrachis species (scale lines, 1 mm). •

• •

41. amazonicuS

/-- -!

42. attonitus

• f- --J 1

43. candeo

• 44. carmenae f- -1 130 •

Figs. 45-48. Forewings of Niteobrachis species (scale lines, 1 mm). •

• •

45. clarus

• 46. delicatus

47. elegans

• 48. fulgeo 131 •

Figs. 49-52. Forewings of Niteobrachis species (scale lines. 1 mm). •

• •

49. gracilis

• 50. gustus

51. ictus

• 52. insignis 132 •

Figs. 53-56. Forewings of Niteobrachis species (scale lines, 1 mm). •

• j j j • j j j j j j 53. Iuceo j j j j j j j j j 54. IUCidUS j j • j j j j j j j 55. IUisi j j j -1 j j j j j j 56. lumen j j • j j j IJJ j j j j j •

Figs.57-60. Forewings of Niteobrachis species (scale lines, 1 mm). •

• •

57. luminasus

• 58. luna

59. mica

• 60. nitidus 134 •

Figs.61-64. Forewings of Niteobrachis species (scale lines, 1 mm). •

• •

61. serenus

• 62. splendidus

63. subtilis

• 64. venustus 135 •

Fig. 65. Farewing af Helcanini. Arraw painting ta vein r-m (scale line, 1 mm).

Figs. 66-67. • Farewings of Brachistini. Arrows pointing ta vein 3RS (scale line, 1 mm).

Fig. 68. Forewing of Brachistini. Arraw painting to break at juncture of veins C+Sc+R with pterostigma (scale lines, 1 mm).

• •

65. He/con yukonensis

• 66. Triaspis kurlogaster

67. Schizoprymnus (Schizoprymnus) ambiguus

• 68. Urosiga/phus (Bruchiurosigalphus) bruchi 136 •

Fig. 69. • Forewing of Brachistini. Arrow pointing at juncture of veins C+Sc+R with pterostigma (scale lines, 1 mm).

Figs. 70-71. Forewings of Brachistini. Arrow pointing to absence and presence of vein 2cu-a (scale line, 1 mm).

• •

69. Schizoprymnus (Muiriella) concisa

• 70. Eubazus (Calyptus) chinensis After van Achterberg (1990)

71. Eubazus (Brachistes) lapponicus • After van Achterberg (1990) 137 •

Figs. 72-75. Hind wings of Niteobrachis species (scale Iines, 1mm). •

• •

72. amazonicus

• 73. attonitus

74. candeo

• 75. carmenae 138 •

Figs. 76-79. Hind wings of Niteobrachis species (scale lines, 1 mm). •

• •

76. clarus

• 77. delicatus

78. elegans

• 79. fulgeo 139 •

Figs. 80-83. Hind wings of Niteobrachis species (scale lines, 1 mm). •

• •

80. gracilis

• 81. gustus

82. ictus

• 83. insignis 140 •

Figs.84-87. Hind wings of Niteobrachis species (scale lines, 1 mm). •

• •

84. /uceo

• 85. lucidus

86. luisi

• 87. lumen 141 •

Figs. 88-91. Hind wings of Nifeobrachis species (scale lines, 1 mm). •

• •

88. luminosus

• 89. luna

90. mica

• 91. nifidus 142 •

Figs. 92-95. Hind wings of Niteobrachis species (scale lines, 1 mm). •

• •

92. serenus

• 93. splendidus

94. subtilis

• 95. venustus 143 •

Fig. 96. Hind wing of Brachistini (scale line, 1 mm). • Figs. 97-98. Propodeum, dorsal view. 97. Niteobrachis serenus. 98. N. insignis.

• • Cell R

96 Urosigalphus (Sruchiurosigalphus) bruchi.

Median longitudinal carina

Median areola Anterior transverse carina Lateral longitudinal ­ carina

• 97

o. Posterior transverse carina • 98 144 •

Fig. 99. Metanotum and propodeum. Niteobrachis venustus. Arrows pointing to curvature of lateral longitudinal carina and division of median areola.

Fig. 100. • Mesosoma (pronotum omitted), dorsal view. Niteobrachis splendidus.

• .tj~ICU • 1 1 •• 125 1 188 " •

....

99 •

\ CJN tu.. :1= x (1)," OCX) .. .. :1.... -1 (JI

Fig. 101. Mesosoma (pronotum omitted), dorsal view. Niteobrachis candeo.

Fig. 102. Propodeum and metasomal tirst median tergite. Niteobrachis /una. Arrow • painting to division of median areola and sigmoideal lateral longitudinal carina.

• •

101

eN m~ =If: X enlT1 00) ~ ~ :3.... -1 (JI CS) -c. (Den ~~ 0) .en R)~ CS) =) ~ 1 ~ ~ ~ , 2) en ç ç • • • • < 16 ••

x 118

190 ~.

=ll=11t1t88S 6-8-96 Bo l te ZEISS DSt19't8A • 102 146 •

Figs. 103-104. Propodeum. dorsal view. 103. N. c/arus. Arrow pointing to division of median areola. 104. He/con tardator. Arrow pointing to median • longitudinal carina.

• • x 228

103

tt 2" SI••

... 104

147 •

Figs. 105-106. Metasoma, dorsal view. 105. Niteobrachis splendidus. 106. N. candeo. •

• • Lateral longitudinal carina First median tergite

Articulation

Second median tergite

Suture

Third median tergite

105 •

: ( r···J (0-:.. # .. 1'..;) ~ fT1 00:' ~ .... -1 U1 r-.J œ Cf) co ~I -~ Cf) ..... ~ ~ 1 t'V oc ~ a ~ 0') W 59: ~ 9 C • 106 148 •

Fig. 107. Metasoma, dorsal view. Nealiolus rufus. Arrow pointing to articulation. •

• •

• m~ )( 'N ~ /)m OUI en . 1 .... -1 c du) ~CXli :1 :-U) QI c ~ (JI , 107 .. CD ç • .. ~ en • •

• 149 •

Figs. 108-112. First median tergite. 108. Niteobrachis serenus. 109. N. lucidus. 110. N. venustus. 111. N. splendidus. 112. N. insignis. Arrows • showing distances of posterior border of first median tergite between and lateral to laterallongitudinal carinae.

• •

100 J.1rn 200 J.1m 108 :-----1 109 f----1

•

200 J.1m 110 111

200 J.ln1 • 112 150 •

Fig. 113A. Metasoma, lateral view. Niteobrachis splendidus (scale line = 1 mm). •

• Female First median tergite • Second median tergite lateral longitudinal carina Third median tergite

First laterotergite

Second laterotergite

Sternum Ovipositor • Thircllaterotergite Male

• 113A 151 •

Fig. 1138. Metasoma, dorsal view. Nifeobrachis spp. Darsh line indicating outline of sterna beneath carapace (cf. Figs. 105 & 113A) (scale line = 1 mm). •

• • Female

• Ovipositor • Male

• 1138 152 •

Figs. 113C-O. Metasoma, Istaral view. Niteobrachis candeo. Arrow in Fig. 113C painting ta absence of crease between third median tergite and laterotergite (cf. Fig. 1148). •

• 1 1 •• x 111 • -~.~ -----.~.- 11e l-'-

:tI=1a.211a 31-7-96 Bo l te -:~ ---ZE 1 SS OSM9Lt8A 113C •

113D • 153 •

Figs. 114A-B. Metasoma, lateral view. Nea/io/us rufus. Arrow in Fig. 114A pointing to articulation; in Fig. 1148 painting to presence of crease between third median tergite and laterotergite. •

• ."~KV 12 • J: - •• x 55 1 2ee ..,.

=1:11+5886 15-8-96 8 o l te ZEISS 114A DSnSlt8A •

1148 • 154 •

Figs.115A-B. Eubazus (A/io/us) crabilli. Arrow in Fig. 115A painting te articulation; in Fig. 1158 pointing ta suture between second and third laterotergite and posterior margin of third median tergite. •

• ~. 14 mm • x 66

200 ~m

#"146013 '2 '1-8-96 B.:. 1. te

ZEISS 1 OSM940A· 115A •

1158 • lSS •

Figs. 116A-B. Metasama, lateral view. Foersteria laeviuscula. Arrow in Fig. 1168 painting ta sterna. •

• • 28 •• ...... x .. 1 288 ~ •

.(- .. '

.158828 18-12-6 80 l te ZEISS DSMS ..8A 116A •

1168 • 156 •

Fig. 117A. Metasoma, lateral view. Triaspis kurlogaster. Arrow pointing to suture between tirst and second median tergites.

Fig. 1178. • Laterotergites. Triaspis kurlogaster. Arrow painting to second and third laterotergites.

• • 19 •• x 37

1 se8 ..,.

*158829 18-12-6 Bo l te ~~ISS Ma 1. IIlIIn 117A •

•

1178 • 157 •

Figs. 118A-B. Metasama, lateral view. Polydegmon foveo/atus. Arrows painting to lateral margin of third median tergite. •

• • 21t •• x 58

288 ~.

_ .....~~--'..-i,

_illQiiiOo.:-o;-- _.11t6821 21-8-96 Bo l te ZEISS 118A OSM91t8A •

1188 • 158 •

Figs.119A-B. Metasoma, lateral view. Eubazus (Srachistes) salicico/a. Arrows in Fig. 119A pointing ta absence of creases between second and third median tergites with respect to their laterotergites; in Fig_ 1198 painting to • painting to lack of carapace.

• • ~ ~x16 •• ------68 J J 1 1 288 loi.

*11tS818 21-8-96 Bo l te ZEISS 119A DS"91t8A •

1198 • 159 •

Figs. 120A-B. Metasoma. lateral and dorsal view. Che/ostes vardzia (scale line = 1 mm).

Figs. 121 A-B. • Metasoma. lateral and dorsal view. Schizoprymnus (Muiriella) concisa. Arrow in Fig. 121A pointing to denticles; in Fig. 121 B pointing to notch (scale line = 1 mm).

• •

120A 1208 •

t

121A 1218 • l60 •

Fig. 122. Metasoma, lateral view. Eubazus (Allodorus) crassigaster. Arrows cf. Figs. 113C, 1148 and 1158 (scale line = 1 mm). •

• •

• 122

• 161 •

Figs.123A-B Metasoma. lateral and dorsal view. Urosigalphus (Bruchiurosigalphus) bruchi. Arrow in Fig. 1238 painting ta third median tergite posterior • margin projections.

• .'+~I

2e0 lJm

=1:146003 21-8-96 Bolte ZEISS 123A

;.,..,....~..._al!t.~~ . . ..,... • 22 •• x 68 2ae .., ...

=ll=11t6882 21-8-96 Bo l te ZEISS OSM9't8A • 1238 162 •

Figs. 124A-B. Metasoma, lateral and dorsal view. Schizoprymnus (Schizorpymnus) ambiguus. Arrows in Fig. 124A pointing ta presence and absence of sutures; Fig. 124B painting to third median tergite posterior margin natch. •

• 13 •• • x 16 288 .,.

----+ ':"~-.-. _ ~_~_. r_ - - ______- ""--.4 • --. - ...... :-._-----

.11t6811 21-8-96 Ba l te ZEISS OSn91tiA 124A • 23 ••

125 1 188 l-I-

·~~.11t6825 21-8-96 Bolte ZEISS DSnSlt8A • 1248 163 •

Fig. 125. Ovipositor length compared to body length. •

• •

1 1 Body length 1 1 1 1 1 le --- --'~--' 1 1 1 1 1 1 J 1 1 1 1 1 1 1 1 1 1 1 1 1 __-.r-- J 1 1 1 1 1 1 1

1 r 1 1 1 1 1 1 1 1 • 1 1 f 1 1 J 1 1

1 ~...------f r • 1 1 1

: Ovipositor length 1:

125 • 164 •

Maps 1-2. Composite distribution of Niteobrachis species. 1. In Costa Rica and Panama. 2. In the Neotropical Region. •

• •

1

~~ ." ~ •.... :, • e::- .. • .•~

• 2 165 •

Maps 3-6. Known distribution of Niteobrachis species. •

• •

3. delicatus 4. serenus

• ~s;:r ~~ •...... ". .... • CIl .~ • --=- .-• .~•

.~ • •.~

• 5. lucidus 6. fulgeo 166 -.

Maps 7-10. Known distribution of Nifeobrachis species. •

• •

7. /uceo 8. insignis

• ~:ti? •.... • e:-. '!... .~

10. venustus • 9. e/egans 167 •

Maps 11-14. Known distribution of Niteobrachis species. •

• •

• 11. clarus 12. gustus ~~ . -

• 13.luna 14. luisi 168 •

Maps 15-18. Known distribution of Nifeobrachis species. •

• •

• 15. luminosus 16. subtilis

• 17. splendidus 18. amazonicus

169 •

Maps 19-22. Known distribution of Niteobrachis species. •

• •

• 19. lumen 20. ictus ~:s:;r ._" . . ~ . !.... "

• 21. carmenae 22. candeo

170 •

Maps 23-26. Known distribution of Niteobrachis species. •

• ~:s? •. '. • • ca ... ~ li .~

• 23. mico 24. attonitus

• 25. nitidus 26. gracilis

171 •

Map 27. Known distribution of Foersteria Szépligeti in the Palearctic Region compared ta Niteobrachis Gen. Nov. in the Neotropical Region. •

• •

•

•

172 APPENDIX 1 • CHARACTER FILE OF DELTA ·SHQW SPECIES OF NITEOBRACHIS. July 29, 1997 ·CHARACTER LIST

#1. ! #2. antenna with ! flagellomeres! #3. temple ! 1. raised laterally (Fig. )! 2. evenly convex (Fig. )! #4. clypeus ! 1. 2.5 to 2.7 times wider than long (Fig. )1 2. 2.0 times wider than long (Fig. )1 #5. clypeus ! • 1. punctate (Fig. )1 2. rugose or heavily pitted (Fig. )1 #6. clypeus 1 1. very convex (Fig. )1 2. fiat ta weakly convex (Fig. )1 #7. face ! 1. smooth (Fig. )1 2. punctate (Fig. )1 3. rugose (Fig. )1 #8. median lobe of mesoscutuml 1. without carina on margin (Fig. )1 2. with carina on margin (Fig. )1

• 173 #9. longitudinal median carina / • 1. complete (Fig. )/ 2. present only posteriorly (Fig. )/ 3. absent (Fig. )/ #10. notaulusl 1. completely crenulate (Fig. )/ 2. smooth anteriorly crenulate posteriorly (Fig.)1 #11. crenulae / 1. medium (Fig. )/ 2. large (Fig. )/ #12. sternaulus/ 1. with crenulae (Fig. )/ 2. without crenulae (Fig. )/ #13. posterior transverse carinae of propodeuml 1. present (Fig. )/ • 2. absent (Fig. )1 #14. lateral longitudinal carinae / 1. straight (Fig. )/ 2. curved (Fig. )1 #15. median areola 1 1. divided (Fig. )/ 2. undivided (Fig. )1 #16. forewing veins: 1m-cul 1. straighU 2. curvedl #17. 2RS/ 1. straight or nearly sol 2. curvedl • 174 #18. 2cu-a with an angle of 1 degrees with respect to 2CUaJ • #19. anterior hait of 1cu-a with an angle of 1 degrees with respect to AI #20. 2RS with an angle of / degrees with respect to (Rs+M)bl #21. Hind wing veins: cu-al 1. straightl 2. curved at least in posterior 1.51 #22. ratio of length of 1r-m to 1M from/ 1. 1.2 to 1.5/ 2. 0.9 to 1.11 3.0.7 to 0.81 #23. ratio of length of R ta 1M troml 1. 0.2 to 0.31 2.0.4 to 0.51 3.0.6 to 0.71 • #24. tirst median tergitel 1. with striations (Fig. )1 2. without striations (Fig. )1 #25. <1 st m. T.> posterior border between lateral longitudinal carinael 1. shorter than posterior border lateral to lateral longitudinal carinae/ 2. longer than posterior border lateral to laterallongitudinal carinae/ 3. nearly equal to posterior border lateral to laterallongitudinal carinael #26. ovipositorl 1. equal ta or longer than body length (Fig. )/ 2. shorter than body length (Fig. )/ • 175 #27. ! #28. ! • #29. ! #30. ! #31_ / #32. / #33. /

•

• 176 APPENDIX2 • TOKEY FILE OF DELTA

*SHOW Translate into KEY format *HEADING Niteobrachis species. Data converted July. 29, 1997. *LISTING FILE tokey.lst *INPUT FILE specs *TRANSLATE INTO KEY FORMAT *EXCLUDE CHARACTERS 1-227-33 *USE NORMAL VALUES

*CHARACTER RELIABILITIES 3,54,7 5-6,57,38,109,8 10-11,5 12,7 13,3 14,2 15,1 16-20,721,322-23,724,1 25-26,5

*KEY OUTPUT FILE kchars • *INPUT FILE chars *KEY OUTPUT FILE kitems *INPUT FILE items

• 177 • • •

Abbreviatlons: 0 =primitive; 1-4 =derived states; ? =unknown; - =mlssing. ml'rl. t 1 2 3 4 5 1 1 1 1 10 11 12 13 14 15 Frontal Frontal anterlor Medial, Texture Inter tel Texture Surface Wldlh 0 lower n Number Numbtr TemDlet OccIDlta Forewln! 1 ANCESTOR 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 2 Cenocoellus 0 1 0 0 ? 1 ?? 0 1 1 0 0 1 0 3 Brull.1a 1 1 0 1 '7 1 1 0&2 0 1 0 0 1 4 Blacus(Blacus) 1 0 1 1 ? 0 "? " 1 0 1 1 0 0 1 5 Dlospllus 1 0 1 1 ? 0&1 '7 "? 1 0 0 1 0 0 1 EublZul(Aliodorus) 1 1 1 , ? 1 ? ? 0 1 1 1 0 0 1 •7 Eubazus(Brachlstes) 1 0&1 1 1 ? 0&1 ?? 0 0 1 1 0 0 1 Eubazus(Calyptus) 1 0&1 1 1 ? 0 ? ? 1 0 1 1 0 0 1 • Eubazul(Eubazus) 1 0&1 1 1 ? 0&1 ? 1 0 1 1 0 0 1 10• FoelSterla 1 1 1 1 ? 0 ? "? 1 0 1 1 0 2 1 11 Eubazus(Aliolus) 1 0&1 1 1 '7 0&1 ?? 0&1 0 1 1 0 0 1 12 Schlzoprymnus 1 1 1 1 ? 1 ? ? 0&1 0 0 1 0 0 1 13 Nul/olus 1 0 1 1 ? 0 ? ? 0&1 0 0 1 0&1 0 1 14 Polydegmon 1 0 1 1 ? 0 ?? 1 0 1 1 0 0 1 ~~ 15 TrIaIpJs 1 0&1 1 1 ? 0&1 ?? 0&1 0 1 1 0 0 1 ,. Urosigalphus , 1 0 1 0&1 ? ? 1 3 0 1 1 1 1 ~;I 17 Schlz (Mulrlella) 1 0 1 1 "? 1 ? 0 0 , 0 1 1 ~ I:Z - 0&1&2 0 0&1 0&1 0&1 0 1 , 0&1 00 Nlteobrachls 1 1 1 1 " " 0 1 ~5Z 11" N.grac:1l1a 1 1 1 1 0 0 1 0 1 0 1 1 1 0 1 20 N. nltidui 1 1 1 1 0 0 0 0 1 0 1 1 1 0 1 21 N.lucldus 1 1 1 1 0 0 0 1 0 0 1 1 1 0 1 ~= 22 N. lumlnosus 1 1 1 1 2 0 1 1 1 0 1 1 0 0 1 23 N.luna 1 1 1 1 1 0 0 1 0 0 1 1 0 0 1 24 N.c1arus 1 1 1 1 2 0 1 1 1 0 1 1 0 0 1 25 N. venustus 1 1 1 1 2 0 1 1 1 0 1 1 0 0 1 21 N. dellçalus 1 1 1 1 2 0 1 1 0 0 1 1 0 0 , 27 N. elegans 1 1 1 1 2 0 1 1 1 0 1 1 1 0 1 21 N...Jenus 1 1 1 1 1 0 1 1 0 0 1 1 1 0 1 21 N. a"onllus 1 1 1 1 0 0 0 0 1 0 1 , 1 0 1 30 N.carmenae 1 1 1 , 0 0 1 1 1 0 1 1 1 0 1 31 N.lulsl 1 1 1 1 2 0 1 1 0 0 1 1 0 0 1 32 N. gustus 1 1 1 1 2 0 1 1 0 0 1 1 0 0 1 33 N. subtllls 1 1 1 , 0 0 1 1 1 0 t 1 1 0 1 34 N, Ictus 1 1 1 1 0 0 , 1 1 0 1 1 1 0 1 35 N. candeo 1 1 1 1 0 0 0 1 1 0 1 1 1 0 , 3. N, lumen 1 1 1 1 0 0 0 0 0 0 1 1 1 0 1 37 N. fulgeo 1 1 1 1 2 0 1 1 1 0 1 1 1 0 1 31 N.lnslgnls 1 1 1 1 0&1 0 1 1 1 0 1 1 1 0 1 31 N. splendidus 1 1 1 1 0 0 0&1 1 1 0 1 1 1 0 1 40 N. mlco 1 1 1 1 0 0 0 0 1 0 1 1 1 0 1 41 N.luceo 1 1 1 1 1 0 1 0 1 0 1 1 0 0 1 42 N. amazonlcus 1 1 1 , 0 0 0 0 1 0 1 1 , 0 1 • • •

ml'rl. 2 11 17 18 1. 20 21 22 23 24 25 21 27 21 2. 3D Farewlnt Farewln! FW. lAI Farewln· Forewln- Forewln· FW 2A: ant. hall Forewln, Forewln. Ralla of Ratio 01 Hlnd wh Lana. rit Crenula' 1 ANCESTOR 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 2 Cenocoellus 0 0 1 0 0 0 ? ? 0 1 ?? 0 1&2 0 3 Brunela 0 0 1 0 0 0 ? ? 0 0 ? ? 0 0&1 2 4 Blacus(Btacus) 0 0 0 0 t 1 ?? 0&1 t ?? 0 0 2 5 DIOsPllul 0 0&1 0 0 0 0 ?? 0 0 ? ? 0 0 2 1 Eubazus(Allodorus) 0 0 0 0 1 0 ? ? 0 0 ? ? 0 0 2 7 EubazuI(Brachlstes) 0 0 0 0 1 0 ? ? 0 0 ? ? 0 0 2 Eubazul(Calvptus) 0 0 0 0 1 1 ?? 0&1 0&1 ? ? 0 0 2 • Eubazus(Eubazus) 0 0 0 0 1 0 ?? . 0 0 ? ? 0 0 2 10• Foell,erla 0 0 0 0 1 0 ? ? 0 1 ?? 0 0 0 11 Eublzus(AlIolus) 0 0 0 0 1 0 ? ? 0 0 ? ? 0 0 2 12 Schlzoprymnus 1 0 0 0 1 0 ? ? 1 1 ? ? 0 0 2 13 Neallolus 0 0 0 0 t 0 ? ? 1 1 ? ? 0 0&1&2 0&1&2 14 Polvdeamon 0 0 0 0 1 0 ? ? 1 1 ? ? 0 0 2 11 TrIupIs 1 0 0 0 1 0 ? ? 1 1 ? ? 0 0 2 1. Uroslgalphus 0 0 0 0 1 0&1 ?? 0&1 0&1 ?? 0 0 2 17 SChlz (Mulrlella) 0 0 0 0 1 0 ? ? 1 1 ? ? 0 0 0 Nlt80brachll 0 0&1 0 0&1 1 0 ~1&2&3c 0&1&2 0 0 0&1&2 0&1&2 0&1 0&1&2 0&' l'18 N. gracilis 0 0 0 0 1 0 2 0 0 0 2 2 0 2 0 \C 20 N. nltldus 0 0 0 1 1 0 2 1 0 0 1 2 1 1 0 " 21 N.luc/dus 0 0 0 0 1 0 2 0 0 0 1 1 0 0 0 22 N, fumlnosus 0 0 0 0 1 0 2 3 0 0 1 1 0 1 0 23 N.luna 0 0 0 0 1 0 2 0 0 0 2 1 0 0 0 24 N. clarus 0 0 0 0 1 0 2 0 0 0 0 0 0 2 0 25 N. venustus 0 0 0 0 1 0 2 0 0 0 0 2 0 2 0 21 N. dellcatus 0 0 0 0 1 0 1 3 0 0 0 0 0 1 0 27 N. elegans 0 0 0 0 1 0 2 0 0 0 1 1 0 2 0 21 N. serenus 0 1 0 0 1 0 2 2 0 0 0 2 1 2 1 28 N. attanltus 0 0 0 1 1 0 2 0 0 0 0 2 1 2 0 30 N.cannenae 0 0 0 1 1 0 2 1 0 0 2 1 0 1 1 31 N.lulsl 0 0 0 0 1 0 1 1 0 0 2 0 0 1 0 32 N. gUltus 0 0 0 0 1 0 0 1 0 0 1 1 0 2 0 33 N. subtllls 0 0 0 0 1 0 1 0 0 0 1 2 0 2 0 34 N. Ictus 0 0 0 0 1 0 4 0 0 0 1 1 1 1 0 35 N. candeo 0 0 0 1 1 0 1 0 0 0 1 1 1 2 0 31 N.lumen 0 0 0 0 1 0 2 1 0 0 2 1 0 1 0 37 N. turgea 0 0 0 0 1 0 3 2 0 0 2 1 0 2 0 31 N.lnslgnls 0 0 0 0 1 0 3 1 0 0 2 2 0 2 0 3' N. splendldus 0 0&1 0 O. 1 0 3 0 0 0 2 1 0 2 0&1 40 N. mlco 0 0 0 1 1 0 3 0 0 0 1 2 1 0 0 41 N.luceo 0 0 0 0 1 0 2 1 0 0 2 2 0 2 0 42 N. amazonlcus 0 0 0 0 1 0 4 1 0 0 2 2 0 1 0 • • •

m.trlx S 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 Number sternaul Median Lateral r1DOSI. trs medlan Median claws Flrst thl Flrst thl Flrsllhl LaI. lonl Lateral 1lst med Strletlon 1 ANCESTOA 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 2 CenocoeUus 1 1 0 0 0&1 0 0 1 0 0 0 0 0 · 0 1 . 0&1 3 BruUela 0 1 1 0 0 0 0 0 0 · 1 4 Blacus(B1aeus) 0&1 0 0 0 0 0&1 0 0 0 0 0 1 0 · 0&1 5 OlolpUus 1 1 1 1 0 0 0 1 0 0 0 0 0 · 0 Eubazus(AlIodorus) 1 2 0 0 0 0 0 0 0 0 0 0 0&1 · 0 •7 Eub8Zua(Brachlstes) 1 2 0 0 0 0 0 0 0 0 0 0 1 0 0 Eubazus(Calyptus) 1 2 0 0&1 0 0 0&1 0 0 0 0 0 1 0&1 0 •1 Eubazus(Eubazus) 1 2 0 0 0 0 1 0 0 0 0 0 0&1 0 0 10 Foersterla 1 2 0 1 0&1 0 0&1 1 1 0 0 0&1 0 · 0 11 Eubazus(AUolus) 1 1&2 0 0&1 0&1 0 0&1 0 1 0 0 0 0&1 · 0 12 Schlzopryrnnus 1 1&2 0&1 0 0 0 1 0 1 1 1 0 0 · 0 13 NeaUolus 1 1&2 1 0&1 0&1 0 0&1 1 1 0 0 0 1 0 0&1 14 Polydegrnon 1 2 1 1 0&1 1 0 1 0 0 1 · 0 15 TrlaspIs 1 1 1 0 0 0 1 1 1 1 1 0 0&1 · 0 1. Uroslgalphus 0 1 1 0 0 0 1 0 1 1 1 0 0 · 0 17 Schlz (Mulrlella) 0 1 0 1 0 0 1 0 1 0 1 0 0 · 0 11 Nlteobrachls 1 1&2 1 0&1 0&1 0 0&1 1 1 0 0 0 1 0&1&2 0&1 11 N. gllelill 1 1 1 0 0 0 1 1 1 0 0 0 1 1 1 -~ 20 N. nltldu8 1 2 1 0 0 0 1 1 1 0 0 0 1 1 1 21 N.lucldus 1 1 1 1 0 0 1 1 1 0 0 0 1 2 0 22 N. lurnlnosu8 1 1 1 1 0 0 0 1 1 0 0 0 1 1 0 23 N,luna 1 1 1 1 0 0 0 1 1 0 0 0 1 1 0 24 N,clarus 1 1 1 1 0 0 0 1 1 0 0 0 1 0 1 25 N. vanuatus 1 1 1 1 0 0 0 1 1 0 0 0 1 1 1 21 N. dellcatus 1 2 1 0 0 0 1 1 1 0 0 0 1 0 0 27 N. elegans 1 1 1 1 0 0 0 1 1 0 0 0 , 0 0 21 N. grenus 1 2 1 0 0 0 1 1 1 0 0 0 1 2 0 21 N. al10nltus 1 1 1 0 0 0 1 1 1 0 0 0 1 1 1 30 N.carmenae 1 2 1 0 1 0 0 1 1 0 0 0 1 1 1 31 N.lulsl 1 1 1 1 0 0 0 1 1 0 0 0 1 1 1 32 N. gustus 1 1 1 0 0 0 0 1 1 0 0 0 1 1 0 33 N. subtilis 1 1 1 0 1 0 1 1 1 0 0 0 1 1 0 34 N. Ictus 1 2 1 0 0 0 1 1 1 0 0 0 1 1 1 35 N. candeo 1 2 1 0 0 0 1 1 1 0 0 0 1 2 1 31 N. lumen 1 1 1 0 0 0 1 1 1 0 0 0 1 1 1 37 N. fulgeo 1 1 1 1 0 0 1 1 1 0 0 0 1 1 0 31 N.lnslgnll 1 1 1 0 1 0 0 1 1 0 0 0 1 0 0 31 N. Iplendldus 1 1&2 1 0 0 0 1 1 1 0 0 0 1 1 1 40 N, rnlco 1 1 1 0 0 0 1 1 1 0 0 0 1 1 0 41 N,luceo 1 1 1 1 1 0 0 1 1 0 0 0 1 0 0 42 N. arnazon/cus 1 1 1 0 0 0 1 1 1 0 0 0 , , , • • •

mltrlx 4 46 47 48 49 50 51 52 53 54 55 58 57 58 Median Striation Artlcul. artlcul. 1Median Strlatlon Suture 1Post an, Post. m. Shaoe 1 Sulure 1LaterolE OVIDosl' 1 ANCE5TOA 0 0 0 0 0 0 0 0 0 0 0 0 0 2 Cenocoellus 2 1 0 0 0 1 0 0 0 0 0 0 ? 3 BNllela 2 0 0 0 0 1 0 0 0 0 0 0 ? 4 Blacus(Blacus) 0&1&2 1 0 0 0 1 0&1 0 0 0 0&1 0 ? 5 Dlospllus 2 1 0 0 0 1 0 0 0 0 0 0 ? Eubazus(Allodorus) 2 0 0 1 0 0&1 1 0 0 0 0 0 1 •7 Eubazus(Brachlstes) 2 1 0 0 0 1 0 0 0 0 0 0 ? 8 Eubazus(Calyptus) 2 1 0 0 0 1 0 0 0 0 0 0 ? 1 Eubazus(Eubazus) 2 1 0 0 0 1 0 0 0 0 0 0 ? 10 Foeraterla 1 0&1 0 0 0 0&1 1 0 0 1 1 1 0&1 11 Eubazus(Aliolus) 0 0 0 1 1 0 1 0 0 1 0 0 1 12 Schlzoprymnus 0 0 1 1 0 2 ? 2 2 1 0 1 13 Neallolus 0 0 0 0 1 0&1 1 0 0 1 1 0 0&1 14 Polydegmon 1 0 0 0 0 0 1 1 0 1 1 1 0&1 15 Trlaspls 0 0 1 1 0 1 0 0&2 2 1 0 1 1. Urosloalphus 0 0 1 1 0 0 1 0&1&2 2 1 0 1 17 Schlz (MulrleUa) 0 0 1 1 0 0 1 1 ? 0 0 1 1. Nlteobrachls 0 1 0 0 0 1 0 0 0 1 1 0 0&1 00 19 N. gracills 0 1 0 0 0 1 0 0 0 1 1 0 0 20 N. nltldus 0 1 0 0 0 1 0 0 0 1 1 0 0 21 N.lueldus 0 1 0 0 0 1 0 0 0 1 1 0 0 22 N. lumlnosus 0 1 0 0 0 1 0 0 0 1 1 0 0 23 N.luna 0 1 0 0 0 1 0 0 0 1 1 0 1 24 N.c1aNs 0 1 0 0 0 1 0 0 0 1 1 0 0 25 N. venustus 0 1 0 0 0 1 0 0 0 1 1 0 0&1 2. N. dellcatus 0 1 0 0 0 1 0 0 0 1 1 0 0 27 N. elegsns 0 1 0 0 0 1 0 0 0 1 1 0 0 2. N. serenus 0 1 0 0 0 1 0 0 0 1 1 0 1 29 N. anonltus 0 1 0 0 0 1 0 0 0 1 1 0 0 30 N.cannenae 0 1 0 0 0 1 0 0 0 1 1 0 0 31 N.luls1 0 1 0 0 0 1 0 0 0 1 1 0 0 32 N. guslus 0 1 0 0 0 1 0 0 0 1 1 0 0 33 N. subtills 0 1 0 0 0 1 0 0 0 1 1 0 0 34 N. Ictus 0 1 0 0 0 1 0 0 0 1 1 0 1 35 N. candeo 0 1 0 0 0 1 0 0 0 1 1 0 0 38 N.lumen 0 1 0 0 0 1 0 0 0 1 1 0 1 37 N. rulgea 0 1 0 0 0 1 0 0 0 1 1 0 1 3. N.lnslgnls 0 1 0 0 0 1 0 0 0 1 1 0 0 31 N. splendldus 0 1 0 0 0 1 0 0 0 1 1 0 0&1 40 N. mlco 0 1 0 0 0 1 0 0 0 1 1 0 . 41 N.lueeo 0 1 0 0 0 1 0 0 0 1 1 0 0 42 N. amazonlcus 0 1 0 0 0 1 0 0 0 1 1 0 1 APPENDIX4

• Phylogenetic analysis of the tribe Brachistini to establish the monophyly and placement ofthe genus Niteobrachis:

Branch-and-bound search completed:

Shortest tree found =212 Number of trees retained =120

The following taxa have been deleted: N. delicatus N. serenus N.lucidus N. fulgeo N.luceo N. insignis N. elegans N. venustus N. clarus N. gustus N.luna • N. luisi N. luminosus N. subtilis N. splendidus N. amazonicus N. lumen N. ictus N.carmenae N.candeo N. mico N. attonitus N. nitidus N. gracilis

Oesignated outgroup taxa: ANCESTOR

Oesignated ingroup taxa: Cenocoelius Brulleia Blacus (Blacus) • 182 Diospilus Eubazus (Allodorus) Eubazus (Brachistes) • Eubazus (Calyptus) Eubazus (Eubazus) Eubazus (Aliolus) Nealiolus Triaspis Urosigalphus Schizoprymnus (Sehizoprymnus) Schizoprymnus (Muiriella) Polydegmon Foersteria Niteobrachis

Current status of ail eharaeters:

Character Type Inform? Status Weight States 1.Frontal ca Unord y 1 01 2.Frontal ri Unord y 1 01 3.anterior 0 Unord y 1 01 4.Medial pro Unord y 1 01 5.Texture of Unord N 1 012 • 6.1nter tent Unord y 1 01 7.Texture of Unord N 1 01 S.Surface of Unord N 1 01 9.Width ofe Unord y 1 01 10.Lower marg Unord y 1 0123 11.Number of Unord y 1 01 12.Number of Unord y 1 01 13.Temples Unord y 1 01 14.0ccipital Unord y 1 012 15.Forewing v Unord y 1 01 16.Forewing v Unord y 1 01 17.Forewing v Unord y 1 01 18.FW._(RS+M Unord y 1 01 19.Forewing_v Unord N 1 01 20.Forewing_v Unord y 1 01 21.Forewing_v Unord y 1 01 22.FW_2RS_an Unord N 1 01234 23.ant._half Unord N 1 012 24.Forewing v Unord y 1 01 25.Forewing v Unord y 1 01 • 183 26.Ratio of H Unord N 1 012 27.Ratio_of_H Unord N 1 012 28.Hind_wing_ Unord N 1 01 • 29.Long. ridg Unord y 1 012 30.Crenulate Unord y 1 012 31.Number of U:1ord y 1 01 32.stemaulus Unord y 1 012 33.Median Ion Unord y 1 01 34.Lateral 10 Unord y 1 01 35.post. tran Unord y 1 01 36.median are Unord y 1 01 37.Median are Unord y 1 01 38.Claws Unord y 1 01 39.First thre Unord y 1 01 40.First thre Unord y 1 01 41 .First thre Unord y 1 01 42.Lat. long Unord y 1 01 43. Lateral 10 Unord y 1 01 44.1 st med. T Unord y 1 012 45.Striations Unord y 1 01 46.Median ter Unord y 1 012 47.Striations Unord y 1 01 48.Articul.b Unord y 1 01 49.articul.b Unord y 1 01 • 50.Median ter Unord y 1 01 51 .Striations Unord y 1 01 52.Suture bet Unord y 1 012 53. Post and 1 Unord y 1 01 54.Post. marg Unord y 1 012 55.Shape_of_1 Unord y 1 012 56.Suture be Unord y 1 01 57.Lateroterg Unord y 1 01 58.0vipositor Unord y 1 01

Phylogenetic analysis of the species of Niteobrachis to establish their relationship within their genus:

Heuristic search completed:

Length of shortest tree found =175 Number of trees retained = 1000 • 184 The following taxa have been deleted: • ANCESTOR Cenocoelius Brulleia Blacus (Blacus) Diospilus Niteobrachis

Oesignated outgroup taxa: Eubazus (Allodorus) Eubazus (Srachistes) Eubazus (Calyptus) Eubazus(Eubazus) Foersteria Eubazus (Aliolus) Schizoprymnus (Schizoprymnus) Nealiolus Polydegmon Triaspis Urosigalphus Schizoprymnus (Muiriella) • Oesignated ingroup taxa: N. delicatus N. serenus N. lucidus N. fulgeo N.luceo N. insignis N. elegans N. venustus N. clarus N. gustus N.luna N. luisi N. luminosus N. subtilis N. splendidus N. amazonicus N. lumen N. ictus N. carmenae • 18:5 N. candeo N.mico N. attonitus • N. nitidus N. gracilis

Current status of ail eharacters:

Charader Type Intorm? Status Weight States 1.Frontal ca N Constllgn 1 1 2.Frontal ri Unord y 1 01 3.Anterior 0 Unord N Ignored 1 01 4.Medial pro N Constllgn 1 1 5.Texture of Unord y Excluded (1 ) 012 6. Inter tent Unord y 1 01 7.Texture of Unord y Excluded (1 ) 01 S.Surface of Unord y Excluded (1 ) 01 9.Width ofe Unord y 1 01 10.Lower marg Unord N Ignored 1 013 11.Number of Unord y 1 01 12.Number of N Constllgn 1 1 13.Temples Unord y 1 01 14.0ccipital Unord y 1 012 • 15.Forewing v N Constllgn 1 1 16.Forewing v Unord y 1 01 17.Forewing v Unord y 1 01 18.FW._(RS+M N Constllgn 1 0 19.Forewing_v Unord y 1 01 20.Forewing_v N Constllgn 1 1 21.Forewing_v Unord y 1 01 22.FW_2RS_an Unord y Excluded (1 ) 01234 23.ant._half_ Unord y Excluded (1 ) 0123 24.Forewing v Unord y 1 01 25.Forewing v Unord y 1 01 26.Ratio of H Unord y Excluded (1 ) 012 27.Ratio_of_H Unord y Excluded (1 ) 012 28.Hind_wing Unord y 1 01 29.Long. ridg Unord y 1 012 30.Cranulate Unord y 1 012 31. Number of Unord y 1 01 32.sternaulus Unord y 1 12 33.Median Ion Unord y 1 01 34.Lateral 10 Unord y 1 01 • 186 35.post. tran Unord y 1 01 36.median are Unord N Ignored 1 01 37.Median are Unord y 1 01 • 38.Claws Unord y 1 01 39.First thre Unord y 1 01 40.First thre Unord y 1 01 41 .First thre Unord y 1 01 42.Lat. long. Unord y 1 01 43.Lateral 10 Unord y 1 01 44.1 st med Unord y 1 012 45.Striations Unord y 1 01 46.Median ter Unord y 1 012 47.Striations Unord y 1 01 48.Articul.b Unord y 1 01 49.articul.b Unord y 1 01 50.Median ter Unord y 1 01 51.Striations Unord y 1 01 52. Suture bet Unord y 1 012 53.Post and 1 Unord y 1 01 54.Post. marg Unord y 1 012 55.Shape_of_1 Unord y 1 012 56.Suture be Unord y 1 01 57. Lateroterg Unord y 1 01 • 58.0vipositor Unord y 1 01

• 187