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Coleoptera: Scarabaeidae: Scarabaeinae) REVISIÓN TAXONÓMICA Y ANÁLISIS FILOGENÉTICO DE LA TRIBU ATEUCHINI (Coleoptera: Scarabaeidae: Scarabaeinae) TESIS QUE PRESENTA FERNANDO ZAGURY VAZ DE MELLO PARA OBTENER EL GRADO DE DOCTOR EN CIENCIAS SISTEMÁTICA Xalapa, Veracruz, México 2007 2 3 4 "... a gente quer passar um rio a nado, e passa; mas vai dar na outra banda é num ponto muito mais embaixo, bem diverso do em que primeiro se pensou. Viver nem não é muito perigoso?" (Guimarães Rosa: Grande Sertão: Veredas, 1956) AGRADECIMIENTOS A Silvia, Léo y Nuno, por el apoyo, paciencia y resignación. A mis padres y abuelos por el apoyo a la educación sensu lato, y por nunca pedir justificaciones ni reportes. A mis amigos (presentes y ausentes) y compañeros (todos) por el apoyo, aún en forma de "deja esto, te estás poniendo loco…". A Pablo, Nico, Abundis, Esther, Hugo y Pedro, por presentarme un nuevo país, mejor dicho varios. A Gonzalo y Violeta Halffter, por el apoyo a todos mis caprichos que llevaron al resultado aquí presentado. A Mario Zunino, Juanjo Morrone, Alejandro Espinosa de los Monteros y demás jurados, por la (espero) sinceridad de sus críticas. A David Edmonds por la paciencia y buenas ideas. A Júlio Louzada, Everardo y Paschoal Grossi, Ayr Bello y Cristiano Lopes-Andrade, por el apoyo en todo esto, desde el planteamiento. Ojalá les haya gustado. A todos los coleccionistas, colectores y curadores que prestaron o cedieron material examinado a lo largo de estos últimos años, por la diversión proporcionada. A Maru Rivas, Fer Escobar, Rocío Rodríguez, Elizabeth Avila, Ana Morales y Alejandra Valencia, por el apoyo burocrático – yo no hubiese podido sin Uds. A François Génier por las calurosas discusiones sobre el tema de la tesis, con el apoyo indudable de la maravillosa industria cervecera québécois. A David Martínez y a Uroš Ušćebrka, que me enseñaron que es posible ser más gruñón que yo. A los que tuvieron la paciencia y la piedad de leer partes de este trabajo. A mucho más gente que no menciono aquí por falta de espacio, pero que ayudó en los más diversos asuntos, ¡Uds. saben quienes son! A la Fundação CAPES (Ministério da Educação, República Federativa do Brasil), por la beca y facilidades (supongo que tengo que citar el numerito: BEX 1208-020); a CONABIO por el apoyo al conocimiento de la fauna mexicana y más directamente a partes de este trabajo. Al Instituto de Ecología, A.C. (principalmente División de Posgrado y Departamento de Biodiversidad y Ecología Animal), por la oportunidad y apoyo. A las instituciones que me posibilitaron el acceso al material, bibliografía e investigadores. A muchas personas e instituciones que no me quisieron ayudar o que me crearon dificultades, gracias por las clases de superación personal. ¡Uds. también saben quienes son! "Ah, vai vir um tempo, em que não se usa mais matar gente..." (Guimarães Rosa: Grande Sertão: Veredas, 1956) RECHAZO PARA FINES DE NOMENCLATURA ZOOLÓGICA Este trabajo, en su actual presentación (tésis de grado), no debe considerarse como una publicación válida para fines de nomenclatura zoológica. Este es el disclaim / dénégation mencionado en el Código Internacional de Nomenclatura Zoológica (edición 1999), capítulo tres, artículos 8.2 y 8.3. A mi familia (presente y ausente), por todo. 5 6 7 8 ÍNDICE Lista de Tablas 11 Lista de Figuras 12 Lista de apéndices 15 Resúmen general 16 Presentación 17 Capítulo I. Introducción general 19 La larga y confusa historia de la taxonomía supragenérica de los Scarabaeinae 22 La corta y decepcionante historia de los estudios filogenéticos de relaciones entre las tribus de 28 Scarabaeinae Problemas nomenclaturales 29 Objetivos del presente trabajo 31 Literatura citada 31 Capítulo II. Synopsis of the new subtribe Scatimina (Coleoptera: Scarabaeidae: Scarabaeinae: 47 Ateuchini), with descriptions of twelve new genera Abstract 49 Resúmen 50 Resumo 51 Introduction 52 Materials and Methods 56 Results and Discussion 57 Acknowlegdements 115 Literature cited 116 Figure Captions 125 Capítulo III. The genera formerly placed in the tribe Ateuchini (Coleoptera: Scarabaeidae: 147 Scarabaeinae) and their phylogenetic position Introduction 149 Methods 151 Results and Discussion 152 Acknowledgements 165 Literature cited 165 Figure Captions 173 Appendix 1 174 Appendix 2 186 9 Capítulo IV. A new dung beetle genus with two new species from Chile 195 (Coleoptera: Scarabaeidae: Scarabaeinae) Abstract 197 Resúmen 197 Introduction 197 Specimens Studied 198 Tesserodoniella gen. n. 198 Biogeography of Tesserodoniella 204 Acknowledgements 205 References 206 Capítulo V. Revision and phylogeny of the dung beetle genus Zonocopris Arrow 1932 (Coleoptera: 207 Scarabaeidae: Scarabaeinae), a phoretic of land snails Abstract 209 Résumé 209 Material and Methods 210 Results and Discussion 211 Acknowledgements 215 References 216 Appendix 217 Capítulo VI. Discusión general 219 Análisis autocrítico de los cuatro capítulos centrales 221 La taxonomía supragenérica de los Scarabaeinae: problemas para trabajos ecológicos 223 Caracteres, árboles, e iluminación recíproca 225 Tribus y subtribus de Scarabaeinae: ¿que grupos siguen con problemas? 227 La evolución del comportamiento: ¿qué hay de nuevo? 229 Ateuchini, Demarziellini, y qué falta hacer 232 Literatura citada 233 Legendas de las figuras 234 10 LISTA DE TABLAS Capítulo I. Cuadro 1. Géneros-tipo de taxones del grupo de la família en Scarabaeinae y su ubicación taxonómica según distintas síntesis. Capítulo III. Table 1. Species composing the outgroup used for phylogenetic analysis. Tribes in italics in right column are presently considered as synonyms of those in central column. Table 2. Species composing the ingroup used for phylogenetic analysis. Tribes in italics in right column are presently considered as synonyms of those in central column. Central column: tribes marked with an asterisk indicate that such genera had been considered a priori as more closely related to some genus of Ateuchini than to the tribes they were included in. Right column: tribes marked with an asterisk indicate that such genera were included in Ateuchini in a sense different than that used by Montreuil (1998), and equivalent to Dichotomiini. Capítulo V. Table 1. Matrix used for phylogenetic analysis. 11 LISTA DE FIGURAS Capítulo I. Figuras 1-4. Representaciones gráficas publicadas de hipótesis filogenéticos en Scarabaeinae. 1. Paulian (1933); 2. Zunino (1983); 3. Zunino (1983) según Philips et al. (2004); 4. Montreuil (1998). Figuras 5-7. Representaciones gráficas publicadas de hipótesis filogenéticos en Scarabaeinae. 5. Montreuil (1998) reanalisada por Philips et al. (2004); 6. Villalba et al. (2002); 7. Philips et al. (2004). Figuras 8-9. Representaciones gráficas publicadas de hipótesis filogenéticos en Scarabaeinae, de Ocampo & Hawks (2006). Figuras 10-11. Representaciones gráficas publicadas de hipótesis filogenéticos en Scarabaeinae, de Ocampo & Hawks (2006). Capítulo II. Figures 1-4. Pedaridium hirsutum, apex of right anterior tibia and tarsus, respectively dorsal and ventral views: 1-2. m#; 3-4. f#. Figures 5-11. Left epipleuron and pseudoepipleuron, ventral view. 5. Trichillum externepunctatum; 6. Onoreidium cristatum; 7. Scatrichus bicarinatus; 8. Pedaridium hirsutum; 9. Silvinha unica; 10. Besourenga minutus; 11. Degallieridium lilliputanum. Scale = 1.0 mm. Figures 12-14. Spermatheca (receptaculum seminis), and attached part of spermathecal duct, and spermathecal gland. 12. Pedaridium hirsutum; 13. Trichillum externepunctatum; 14. Eutrichillum hirsutum. Scale = 0.2 mm. Figures 15-22. Dorsal views. 15-16: Besourenga: 15. B. sp.; 16. B. horacioi ; 17. Bradypodidium sp. ; 18. Degallieridium lilliputanum; 19. Eutrichillum sp.; 20. Feeridium woodruffi; 21-22: Genieridium: 21. G. bordoni; 22. G. cryptops. Figures 23-30. Dorsal views. 23. Leotrichillum louzadaorum; 24-25: Martinezidium: 24. M. fulgens; 25. M. maya; 26. Nunoidium argentinum; 27. Onoreidium cristatum; 28. Onoreidium bottimeri; 29. Pedaridium hirsutum; 30. Pereiraidium almeidai, m#. Figures 31-38. Dorsal views. 31. Pereiraidium almeidai, f#; 32. Scatimus ovatus; 33. Scatrichus bicarinatus; 34. Silvinha unica; 35. Trichillidium quadridens; 36. Trichillidium brevisetosum; 37. Trichillum heydeni; 38. Trichillum pauliani. Figures 39-46. 39-40: Besourenga minutus: 39. head; 40. parameres; 41. B. amarillai, head; 42. B. horacioi, m# front tarsus; 43-44: Bradypodidium bradyporum: 43. head; 44. parameres; 45-46: B. adisi; 45. head; 46. m# front tarsus. Figures 47-54. 47-48: Degallieridium lilliputanum: 47. head; 48. parameres; 49-51: Eutrichillum hirsutum: 49. head; 50. parameres; 51. m# front tarsus; 52-54. Feeridium woodruffi: 52. head; 53. parameres; 54. ventral view of hind legs and abdomen. Figures 55-62. Genieridium. 55-56: G. bidens: 55. head; 56. parameres; 57-59: G. bordoni: 57. head; 58. parameres; 59. pronotum, lateral view; 60-61: G. cryptops: 60. head; 61. parameres; 62. G. margareteae, head. 12 Figures 63-70. Genieridium. 63. G. margareteae, parameres; 64-67: G. medinae: 64. head; 65. parameres; 66. m# hind tibia; 67. m# hind tibia detail; 68-69: G. paranense: 68. head; 69. parameres; 70. G. zanunciorum, head. Figures 71-78. 71. Genieridium zanunciorum, parameres; 72-73: Leotrichillum louzadaorum: 72. head; 73. parameres; 74-76: Martinezidium: 74-75: M. galileoae: 74. head; 75. parameres; 76. M. maya, head; 77- 78: Nunoidium argentinum: 77. head; 78. parameres. Figures 79-86. 79-83: Onoreidium: 79-81: O. cristatum: 79. head; 80. parameres; 81. middle tibia; 82. O. howdeni, head; 83. O. aff. bottimeri, head;
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