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Coleoptera, Curculionidae) UNiVERSIDAD COMPLUTENSE DE MADRID FACULTAD DE CIENCIAS BIOLÓGICAS DEPARTAMENTO DE BIOLOGIA ANIMAL 1 (ENTOMOLOGIA) TESIS DOCTORAL UU~1111111~IDHM 530982374 UNIVERSIDAD COMPLUTENSE ESTUDIO MORFOLÓGICO Y TAXONÓMICO DEL GÉNEROSitonaGerfllar,1817 (Coleoptera, Curcul¡onidae) Antonio Jesús Velázquez de Castro González Madrid, 1997 UNIVERSIDAD COMPLUTENSE DE MADRID FACULTAD DE CIENCIAS BIOLÓGICAS DEPARTAMENTO DE BIOLOGIA ANIMAL 1 (ENTOMOLOGIA) TESIS DOCTORAL ESTUDIO MORFOL ÓGICO Y TAXONÓMICO DEL GÉNERO Sitona Germar, 1817 (Coleoptera, Curculionidae) Firmas Memoria presentada por: Antonio Jesús Velázquez de Castro González, para optar al grado de doctor en Ciencias Biológicas A> y0 B0 de los directores de la Tesis: Dr. Miguel Angel Alonso Zarazaga Dpto. Biodiversidad y Biología Evolutiva (Entomolo’ Museo Nacional de Ciencias Naturales Dr. Raimundo Outerelo Dominguez Departamento de Biología Animal 1 (Entomologia) Facultad de Ciencias Biológicas Universidad Complutense de Madrid Madrid, 1997 ÍNDICE Agradecimientos L INTRODUCCION 7 1. Historia, nomenclatura ~ clasificación de Sitonini .9 1.1. Desarrollo histórico 9 1.2. Monograflas de Sitona 11 1.3. Posición de la tribu Sitomm 12 2. Escuelasiaxonónhicas 13 2.1. Taxonomía evolutiva o tradicional 14 2.2. E~cuelafenética 14 2.3. Escuela cladista 15 IL OBJETIVOS -21 HL MATERIAL Y MÉTODOS 25 1 Especies estudiadas . 27 1.1. Selección de especies 27 1.2. Identificación de los ejemplares 130 2. Obtención y preparación de las muestras . 33 2.1. Reblandecimiento de los ejemplares .33 2.2. Aclaramiento . .. 34 2.3. Disección -. 34 2.4. Montaje de los ejemplares 35 2.5. Preparaciones microscopicas 36 3. Obtencíónde datos 38 3.l.Datosmorfológicos -38 3.2. Datos biológicos 39 4. Preparación de ilustraciones 40 4.1. Dibujos 41 4.2.Fotografla 41 4.3. Montaje 41 5. Preparación de descripciones 42 5.1. Descripción de las estructuras 42 5.2. Descripción de las especies . 42 1 6. Métodos taxononucos . 43 61. Estudio filogenético .43 6.2. Estudio fenético 46 IV. RESULTADOS Y DISCUSIÓN 49 1. Descripción de las estructuras -. 49 ilLacabeza 51 111. La cápsula cefálica 51 1.1.2. Los ojos 52 1.1.3. Las piezasbucales 57 - 1.4. Las antenas 77 1.2. El tórax - . .... 77 1.2.1. El protórax.. 77 1.2.2. El mesotórax 79 1.2.3. El metatórax SO 1.2.4. El metendoslenuto . .. — 81 1.2.5. Las alas 91 1.2.6. Las patas 97 1.3. El abdomen 97 1.3.1. Los lerguitos 98 1.3.2. Los eslernitos 101 1.4, Revestimientos cuticulares . 101 3.4.1. Las escamas 101 3.4.2. Las sedas elitrales y las sedas oculares 100 1.5. El proventriculo 105 1.6, El aparato reproductor 113 1.6.1. Aparato reproductor fememno 113 1.6.2. Laarinaduradelsacoinlernn 118 2. La planta huésped 129 3. Análisis taxonomico 146 .,. 1. Análisis filogenético de los géneros de Siton¡m 146 3.1.1. Seleccióndegrupo externo 146 3.1.2. Génerosvespecies incluidos en el grupo interno 146 3.1.3. Caracteres utilizados 147 3,1.4. Análisis filogenético 148 3.2. Análisis fenético de las especies y subgéneros de Silona 149 3.2.1. Caracteres utilizados 149 3.2.2. Agnipamiento de las OTUs mediante UPGMA 151 3.3. Análisis filogenético de las especies y subgénerosde Sitona 159 3.3.1. Caracteres utilizados 159 3.3.2. Cladograma resultante 162 3.3 3 Comparación entre la evolución de Sitona y de su planta huésped 163 3.4. Implicaciones taxonomicas 177 3.4.1. Implicaciones conjuntas del estudiofenético y filogenético 177 3.4.2. Sistemática propuesta para el género Sitona 177 .4. Descripción de los taxones 183 4.1. Tribu Sitonini 183 4.1.1. Género Cecracres 183 4.1.2. Género Schelopius 185 4.1.3. Género Eugnarhus 187 4.1.4. Género Sitona 193 4.2. Tribu Alophini 263 4.3. Tribu Pachvrhvnchini 265 4.4. Tribu Tanvmecím 266 4.5. Tribu Naupactím. 267 V. CONCLUSIONES 425 VL BIBLIOGRAFIA 431 VIL ANEXOS 445 Anexo 1: Material estudiado 447 Anexo II: Clave de detenninación de las especies paleárticasoccidentales del género Sitona 465 Anexo III: Matrices usadas con NTSYS 485 Anexo IV: Índice de figuras 489 3 Agradecimientos: Esta Tesis nunca se hubiera llevado a término sin la colaboración y el apoyo de un numeroso grupo de personas que, desde diversos ámbitos han contribuido a su realización. En primer lugar quiero agradecer a mis directores de Tesis, Dr. Miguel Ángel Alonso Zarazaga y Dr. Raimundo Outerelo Domínguez, por su asesoramiento y por la confianza depositada en mi y su apoyo y ánimo continuo durante todos estos años, no sólo en la Tesis, sino desde que en mis inicios empecé a preguntarme qué es un gorgojo. Agradezco su ayuda a todas aquellas personas e instituciones que contribuyeron con el préstamo e intercambio de ejemplares, e incluso con la donación de los mismos en aquellos casos en que les fue posible. Todos han contribuido de forma amable y desinteresada, aunque tengo un recuerdo especial para dos personas que ya no están entre nosotros: Sr. Manuel Ortego (Madrid) y Dr. Lothar Dieckmann (Eberswalde, Alemania). Las siguientes personas pusieron a mi disposición material de museos e instituciones: Dr. L. Behne (Deutsches Entomologisehes Institut, Alemania), Dra. 1. Izquierdo (Museo Nacional de Ciencias Naturales, Madrid), Dr. C. Lyal (British Museum Natural History), Dr. O. Merkl (1-lungarian Museum of Natural History), Dr. M. de los Mozos (Estación Experimental de Albaladejito, Cuenca), Dra. M. J. Sanz y Dra. P. Gurrea (Universidad Autónoma de Madrid), Dr. A. Sendra (Museo del Medio Ambiente, Valencia), y Dra. P. Syrett (Lincoln University, Nueva Zelanda). Las siguientes personas aportaron ejemplares de sus colecciones paniculares: Sr. J. Blasco (Pina de Ebro), Ing. R. Borovee (Rep. Checa), Dr. E. Colonnelli (Italia), Sr. F. Dorado (Madrid), Dr. G. Israelson (Suecia), Sr. J. de Ferrer (Algeciras), Dr. J. L. Forest (Francia), Dr. y. Karasjov (Bielorrusia), Dr. B. Korotyaev (Rusia), Dr. M. Kostál (Rep. Checa), Sr. M. Morales (Sta. Cruz de Tenerife), Dr. J. Pelletier (Francia), Sr. A. Pérez Onteniente (Valencia), Dr. R.W.J. Read (Reino Unido), Sr. M. Sánchez Ruiz (Madrid), Dr. N. Talwar (India), Sr. J. Trautner, (Alemania), y Dr. X. Vázquez (Barcelona). 5 Quiero agradecer a D. José Luis García Alonso las muchas horas de colaboración en el diseño y montaje de las ilustraciones, y el poner a mi disposición todos los medios informáticos de la compañía Síntesis Gráfica C.B. También fue imprescindible la ayuda del Dr. Ximo Baixeras, y en general, del personal del departamento de Entomología de la Facultad de Biología de la Universidad de Valencia. Gracias a ellos pude disponer del material óptico para poder realizar observaciones, dibujos y fotografias. A Pilar Gurrea y Maria José Sanz, colegas, amigas y expertas en la biología de Curculionidae ibéricos, por sus comentarios y ayudas con la bibliografia A todos mis compañeros, con los que me he encontrado haciendo Tesis o Tesinas, gracias por el buen ambiente y los buenos ratos, ya en el “Museo”, ya en la “Facultad”, o en los diversos congresos a los que hemos asistido. Gracias a José Miguel Vela y Gloria Bastazo por sus comentarios sobre aspectos que comparten Crisomélidos y Curculiónidos, a José María Hernández y Manolo Sánchez por la ayuda con el programa de taxonomía NTSYS-pc, a Milagro Coca por los sugerencias sobre herramientas taxonómicas. A mi familia por su compañía y apoyo constante. A mi esposa, Chelo, por compartir con cariño tantos buenos y malos ratos de la Tesis. 6 L INTRODUCCIÓN 7 Introducción 1. HISTORIA, NOMENCLATURA Y CLASifICACIÓN DE SITONINI. 1.1. Desarrollo histórico En este apartado se expone la historia de la descripción y clasificación de las especies de la tribu Sitonini. Es una referencia histórica para comprender mejor la clasificación actual. Se ha dividido en tres secciones, correspondientes cada una a un siglo. Siglo XVIII: El punto de partida obligado es la décima edición de la obra de LII4NAEUS, Systema Na/urae. en 1758. De entre las 86 especies de Curculionoidea descritas por él, se encuentra sólo una especie de Sitona, £ lineatus, descrita, como la mayoría de Curculiónidos, bajo el género Curculio. Durante el resto del siglo se describieron otras 22 especies, todas igualmente dentro del género Curculio, de las cuales sólo ocho se aceptan hoy día como válidas. El gran entomólogo FABRICIUS es autor de tres: 8. griseus (F., 1775), 8. hispidulus (E., 1776) y 8. gressor¡us (F., 1792). Siglo XIX: Durante la primera mitad del siglo XIX se describe un gran número de especies, y ya no son solamente europeas, sino también africanas, asiáticas y amencanas. A principios de siglo GERMAR (1817) crea el género Sitona, con la indicación “46. Sitona nob. Curc. gressorius, lineatus, hispidulus”, si bien poco después (1824) este autor ofrece una descripción del género e incluye en él diez especies. La especie tipo, Curculio lineatus L., 1758 había sido anteriormente propuesta por SCHONI-IERR (1823). Este último autor describe en 1826 el subgénero Charagmus, si bien este detalle pasará inadvertido durante más de un siglo. SCHONI-IERR publica además entre 1833 y 1847 una obra de trascendental importancia en sistemática de Curculionoidea, Genera et species Curculionidum, en cuyo segundo volumen, (1834) además de incluir una docena de nuevas descripciones de Sitona por parte de su colaborador Gyllenhal, describe un nuevo género de Sitonini, Eugnathus Schónherr (fig. lb), para incluir la única especie conocida hasta entonces, E. virídianus Uyll., 1834. El sexto volumen (SCHONIIERR, 1840), merece especial atención. En él se describen otros dos géneros de Sitonirxi: Ceerqetes (fig. la) y Catachaenus, y además la especie Tanymecusplan 4lrons Fáhraeus para la que más adelante, DESBROCHERS (1871) creará el género Sehelopius (fig. le). 9 Introducción Sin embargo SCJ-IÓNHERR no sitúa ninguno de estos tres géneros junto al género Sitona.
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