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REVISIÓN TAXONÓMICA DE LAS HORMIGAS Tapinoma Förster (HYMENOPTERA: FORMICIDAE: DOLICHODERINAE) EN LA REGIÓN NEOTROPICAL

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REVISIÓN TAXONÓMICA DE LAS HORMIGAS Tapinoma Förster (HYMENOPTERA: FORMICIDAE: DOLICHODERINAE) EN LA REGIÓN NEOTROPICAL UNIVERSIDAD CENTRAL DE VENEZUELA FACULTAD DE CIENCIAS POSTGRADO EN ZOOLOGÍA REVISIÓN TAXONÓMICA DE LAS HORMIGAS Tapinoma Förster (HYMENOPTERA: FORMICIDAE: DOLICHODERINAE) EN LA REGIÓN NEOTROPICAL Tesis Doctoral presentado ante la ilustre Universidad Central de Venezuela por el Ldo. Roberto José Guerrero Flórez, para optar al título de Doctor en Ciencias Tutor (es): Juan Carlos Navarro, Ph. D. Fernando Fernández, Ph. D (ICN-UN, Colombia) Caracas – Venezuela Octubre de 2017 2 3 RESUMEN Las hormigas del género Tapinoma son un elemento conspicuo adentro de la subfamilia Dolichoderinae. Estas hormigas son cosmopolitas, exhibiendo una mayor riqueza de especies en las regiones paleotropicales, no obstante, la región Neotropical alberga una fauna considerable de especies de Tapinoma. Por primera vez y tomando en consideración las especies de la región Neotropical, se revisan taxonómicamente las especies del género Tapinoma. El análisis morfológico, datos de distribución y en algunos casos la integración de información ecológica, respaldan la delimitación de 10 especies distribuidas en cuatro grupos de especies (Grupo Litorale, Grupo Melanocephalum, Grupo Ramulorum y Grupo Sessile), de los cuales Litorale y Ramulorum albergan el mayor número de especies Neotropicales. El esquema taxonómico es el siguiente: Tapinoma amazonae Wheeler, W.M. 1934, Tapinoma atriceps Emery, 1888 (=Tapinoma atriceps breviscapus Forel, 1908), Tapinoma inrectum Forel, 1908 (estatus revisado y revivido), T. litorae Wheeler, 1905 (=litorale cubaensis Wheeler, W.M. 1913, nuevo sinónimo; =panamense Wheeler, W.M. 1934, nuevo sinónimo), T. melanocephalum (Fabricius, 1793) (=T. luffae (Kuriam 1955), nuevo sinónimo; =T. melanocephalum coronatum Forel, 1908, nuevo sinónimo; =T. melanocephalum malesianum Forel, 1913, nuevo sinónimo), T. opacum Wheeler, W.M. & Mann, 1914, T. ramulorum Emery, 1896 (=T. ramulorum var. saga Forel, 1912, nuevo sinónimo; =T. ramulorum subsp. anellatum Wheeler, W. M., 1934, nuevo sinónimo; =T. ramulorum var. satullum Wheeler, W. M., 1934, nuevo sinónimo; =T. ramulorum subsp. toltecum Wheeler, W. M., 1934, nuevo sinónimo; =T. gibbosum Stitz, 1933, nuevo sinónimo), T. rasenum Smith, D.R. y Lavigne, 1973, T. sessile Say, 1836. Una nueva especie es descrita, Tapinoma RJGF-01 sp. nov., pero el epíteto específico es omitido siguiendo consideraciones del código de nomenclatura zoológica. La diagnosis y descripción del género para la obrera, macho y hembra reproductora es modificada ligeramente de la propuesta por Shattuck (1992). Se describen las obreras, machos y hembras reproductoras de casi todas las especies, excepto en T. rasenum donde se desconoce la hembra reproductora; se presentan nuevos caracteres taxonómicos para la delimitación de las especies, y se discute la variación morfológica de cada una de las especies a lo largo de su distribución. Cuatro especies son de distribución muy restringida a ambientes continentales o endémicos de las 4 islas del Caribe; otras son de amplia distribución en toda la región Neotropical o en biomas específicos como la selva brasileña del Atlántico. Diez y seis lectotipos son designados, y el neotipo de T. melanocephalum es propuesto; Se presenta una clave taxonómica ilustrada para la identificación de las obreras, así mismo se muestran imágenes de alta resolución para cada una de las castas de las diferentes especies de Tapinoma en la región Neotropical. Se discuten aspectos de la biología y se sugieren algunas hipótesis evolutivas derivadas del análisis morfológico de las especies de Tapinoma. PALABRAS CLAVES: Biodiversidad, Distribución geográfica, Imágenes de alta resolución, Insectos, Taxonomía, Variabilidad morfológica. 5 TABLA DE CONTENIDO AGRADECIMIENTOS .......................................................................................................... 12 INTRODUCCIÓN .................................................................................................................. 14 OBJETIVO GENERAL .......................................................................................................... 18 Objetivos específicos ......................................................................................................... 18 MATERIALES Y MÉTODOS ............................................................................................... 19 Ejemplares examinados .................................................................................................... 19 Delimitación de especies ................................................................................................... 21 Análisis de caracteres morfológicos y morfométricos .................................................... 22 Imágenes a color ................................................................................................................ 25 Mapas de Distribución ...................................................................................................... 25 RESULTADOS....................................................................................................................... 27 Historia taxonómica del genero Tapinoma ..................................................................... 27 Diagnosis ............................................................................................................................ 27 Distribución ....................................................................................................................... 28 Sinopsis taxonómica de las especies de Tapinoma .......................................................... 29 Grupo Litorale .................................................................................................................. 29 Grupo Melanocephalum ................................................................................................... 30 Grupo Ramulorum ........................................................................................................... 30 Grupo Sessile ..................................................................................................................... 31 Especies neotropicales vivientes no asignadas a un grupo ............................................ 31 Nombres excluidos de Tapinoma ..................................................................................... 31 Clave para las especies de Tapinoma de la región Neotropical, basada en obreras .... 37 Sinopsis de las especies ..................................................................................................... 44 Tapinoma amazonae Wheeler, W.M. 1934 ...................................................................... 44 Tapinoma atriceps Emery, 1888 ....................................................................................... 49 Tapinoma inrectum Forel, 1908 ........................................................................................ 67 Tapinoma litorale Wheeler, W.M. 1905 ........................................................................... 81 Tapinoma melanocephalum (Fabricius, 1793) .............................................................. 100 Tapinoma opacum Wheeler, W.M. & Mann, 1914 ....................................................... 121 Tapinoma ramulorum Emery, 1896 ............................................................................... 132 Tapinoma rasenum Smith, D.R. & Lavigne, 1973 ........................................................ 157 6 Tapinoma sessile (Say, 1836) .......................................................................................... 166 Tapinoma RJGF-01 sp. nov. ........................................................................................... 181 DISCUSIÓN ......................................................................................................................... 195 REFERENCIAS BIBLIOGRÁFICAS .................................................................................. 199 MAPAS ................................................................................................................................. 213 7 LISTADO DE FIGURAS Figura 1. Variables morfométricas registradas en la cabeza de la casta obrera, hembra reproductora y macho de Tapinoma spp. ................................................................................ 24 Figura 2. Vista frontal de la cabeza (A) y perfil corporal (B) de la obrera Lectotipo de Tapinoma amazonae (Pará, Brasil: USNM). Fotografías tomadas por Roberto J. Guerrero. 49 Figura 3. Imágenes de la obrera de Tapinoma atriceps. A y B) Vista frontal de la cabeza y perfil corporal de la obrera Lectotipo de Tapinoma atriceps (Rio Grade do Sul, Brasil: MSGH); C y D) Vista frontal de la cabeza y perfil corporal de la obrera Lectotipo de Tapinoma atriceps breviscapum (São Paulo, Brasil: MHNG). Imágenes tomadas de www.atnweb.org (Tapinoma atriceps CASENT0904029; Tapinoma atriceps breviscapum CASENT0909768); todas las fotografía tomada por Zach Lieberman. .................................. 62 Figura 4. Casta de obrera, reina y macho en Tapinoma atriceps. A y B) Vista frontal de la cabeza y perfil corporal de la obrera (Paraná, Brasil: MZSP); C y D) Vista frontal de la cabeza y perfil corporal de una reproductora alada (Misiones, Argentina: CWEM); E y F) Vista frontal de la cabeza y perfil corporal de una reproductora sin alas (São Paulo, Brasil: MZSP), note la diferencia en la longitud de los escapos entre ambas reproductoras; G y H) Vista frontal de la cabeza y perfil corporal de un macho (Minas Gerais, Brasil: MZSP). Fotografías tomadas
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