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Hymenoptera: Vespomorpha) II. INVENTARIOS Y BIODIVERSIDAD DE INSECTOS FILOGENIA Y SISTEMÁTICA DE LOS HIMENÓPTEROS CON AGUIJÓN EN LA REGIÓN NEOTROPICAL (HYMENOPTERA: VESPOMORPHA) Fernando Fernández C. Resumen Abejas, avispas y hormigas comprenden los llamados himenópteros con aguijón (Aculeata o Vespomorpha), grupo natural en donde el ovipositor de las hembras se ha modificado en aguijón. Vespomorpha comprende en la Región Neotropical 3 superfamilias, 25 familias, 807 géneros y alrededor de 13300 especies descritas. Se ofrece una breve revisión de la sistemática y filogenia del grupo para la región, así como un listado de las familias, subfamilias, tribus, géneros y subgéneros descritos hasta el momento. Palabras clave: Filogenia, Neotropical, Sistemática, Vespomorpha. Filogenia y Sistemática de los Phylogeny and Systematics of Aculeate Hymenoptera in the Neotropical Region himenópteros con aguijón en (Hymenoptera: Vespomorpha) la Región Neotropical (Hymenoptera: Vespomorpha) Abstract Bees, wasps and ants comprise the natural group aculeate Hymenoptera (Aculeata Fernando Fernández C. or Vespomorpha), in which the females have the ovipositor modified in a sting. The Instituto Humboldt group comprises 3 superfamiles, 25 families, 807 genera and already 13300 species Apartado Aéreo 8693 in the Neotropical Region. A short revision of the systematics and phylogeny is Bogotá D.C. offered, with a list of all supraspecific categories described up to date in the Colombia Neotropical region. [email protected] Key words: Neotropics, Phylogeny, Systematics, Vespomorpha. Proyecto de Red Iberoamericana de Biogeografía y Entomología Sistemática PrIBES 2002. Introducción C. COSTA, S. A. VANIN, J. M. LOBO & A. MELIC (Eds.) Los himenópteros con aguijón (Hymenoptera: Vespomorpha o Aculeata) comprenden insectos tan comunes como hormigas, abejas y avispas alfareras y ISBN: 84–922495–8–7 papeleras. Es una división natural dentro de los himenópteros, con más de 140 millones de años de historia y con especies solitarias y sociales. El objetivo de m3m : Monografías Tercer Milenio vol. 2, SEA, Zaragoza, Julio-2002. esta contribución es ofrecer una panorámica sobre el estado actual del pp.: 101–138. conocimiento en filogenia y sistemática de los grupos tradicionalmente considerados como Aculeata o Vespomorpha en la Región Neotropical. Además, el anexo ofrece un listado de todos los géneros y subgenéros tal como R I B E S : Red Iberoamericana de actualmente se aceptan en las familias de aculeados. Como en toda síntesis Biogeografía y Entomología Sistemática. pueden estar ausentes citas, referencias y nombres de taxones, propuestas o http://entomologia.rediris.es/pribes Coordinadores del proyecto: personas; aún así la literatura es relativamente completa y el anexo cubre Dr. Jorge LLorente Bousquets (coord.) cuantos nombres se pudieron encontrar y cotejar con revisiones, monografías, Dra. Cleide Costa (coord. adj.) listados u otras fuentes, todas referenciadas en la literatura. Una breve revisión Coeditores del volumen: Se carece de libros o revisiones que traten la biología, sistemática y filogenia de Sociedad Entomológica Aragonesa -SEA http://entomologia.rediris.es/sea los himenópteros neotropicales; sin embargo se revisan brevemente algunas Avda. Radio Juventud, 37 contribuciones que directa o indirectamente cubren algunos de estos aspectos 50012 Zaragoza (ESPAÑA) para grupos de himenópteros neotropicales. [email protected] Durante mucho tiempo la identificación de himenópteros (a nivel de CYTED— Programa Iberoamericano de familia) dependía de las claves en manuales como Borror et al. (1989) o libros Ciencia y Tecnología para el Desarrollo. similares, pensados para faunas neárticas u holárticas. Esto implicaba un gran Subprograma Diversidad Biológica. Coordinador Internacional: problema en el sentido de que varias familias neotropicales no se encuentran en Dr. Peter Mann de Toledo el Neártico y viceversa; el usuario podía asignar la familia incorrecta al ejemplar bajo estudio. Gauld & Bolton (1988) ofrecen una síntesis sobre biología y evolución del orden Hymenoptera, aunque el tratamiento sistemático está circunscrito a Inglaterra. 102 Fernando Fernández Fig. 1. Filogenia de Orussidoidea + Apocrita, indicando Fig. 2. Filogenia de las superfamilias de Vespomorpha. la posición de Aculeata (basado en Whitfield, 1998). El primer libro que trata todas las familias del orden, Filogenia en Vespomorpha y las familias y subfamilias de Aculeata es el editado por Los himenópteros con aguijón (Aculeata o Vespomorpha) Goulet & Huber en 1993. Ésta es una excelente contribu- conforman una agrupación monofilética dentro de Hyme- ción, gracias a un buen glosario, claves profusamente noptera, aparentemente como grupo hermano de las avispas ilustradas y elaboradas en un sentido didáctico. Cada grupo parasitoideas Ichneumonoidea (Fig. 1; Rasnytsin, 1988; se presenta con un tratamiento básico de diagnosis, biología Sharkey & Wahl, 1992). Aculeata a su vez se divide en tres y referencias (escrito por especialistas en el grupo), así superfamilias monofiléticas: Chrysidoidea, Apoidea y como un habitus de cada una de las familias. Los datos de Vespoidea (Fig. 2). Brothers (1975), Brothers & Carpenter números de géneros y especies básicamente se limitan a la (1993) y Brothers (1999) estudian la filogenia de Aculeata. fauna neártica. En 1995 Hanson & Gauld editan el libro Se considera a Vespomorpha como grupo monofilético, Hymenoptera of Costa Rica. La primera gran ventaja de aunque no hay sólidas sinapomorfias identificadas para el este libro es ofrecer unos capítulos introductorios sobre grupo (Tabla I). De los trabajos de Brothers & Carpenter biología, morfología, evolución y aspectos económicos del sólo se considera que las hembras hayan perdido los cerci. orden con enfoque hacia la Región Neotropical. El trata- Se han propuesto dos sinapomorfias para las superfamilias miento sistemático y claves, aunque referidos a Costa Rica, Apoidea y Vespoidea (los aculeados sensu stricto), hem- en general cubren Centroamérica y en algunos casos toda la bras con 12 y machos con 13 antenómeros (número que se Región. Por cada familia se ofrece diagnosis, guía de ha reducido secundariamente en varios linajes, hasta 4 identificación y claves para subfamilias, tribus o géneros antenómeros incluso) y hembras con el séptimo tergo del para Costa Rica o Centroamérica. Este es el primer libro metasoma oculto y no esclerotizado. Las tablas I a IV (por lo menos en tiempos recientes) que puede considerarse presentan las sinapomorfias propuestas para las superfami- útil para los estudiantes de la fauna neotropical. Un primer lias, familias y algunas subfamilias de Vespomorpha. Las problema es que las claves por debajo de familia o subfami- figuras 3 a 8 muestran cladogramas propuestos al interior lia y las guías de géneros no incluyen Suramérica. Además, de cada superfamilia, dentro de cada familia y el algunas el costo del libro lo hace prohibitivo para la mayoría de tribus y géneros de diversos grupos. usuarios latinoamericanos. La literatura sobre evolución y filogenia en Hymenop- Superfamilia Chrysidoidea tera ha sido más rica. Ronquist (1999) ofrece un buen balance de las propuestas sobre filogenia de todo el orden. Superfamilia basal de Aculeata. Filogenia estudiada en Ronquist et al. (1999) ofrecen una reinterpretación cladísti- Carpenter (1986a, 1999) y Brothers & Carpenter (1993). Se ca de las propuestas de Rasnytsin en 1988. Nieves-Aldrey han propuesto tres sinapomorfias para la superfamilias & Fontal-Cazalla (1999) revisan las propuestas sobre (Tabla II), siendo la más fácil de observar la reducción en filogenia y evolución realizadas en las últimas décadas por la venación alar. La figura 3 muestra algunas de las pro- diferentes investigadores y diversos criterios. puestas de filogenia al interior de este grupo. Chrysidoidea Para Hymenoptera Aculeata el trabajo ya clásico es el comprende siete familias con algo más de 1.000 especies en de Brothers (1975) donde se propone la monofilia del grupo el Neotrópico. Todas sus especies son solitarias, con y se divide en tres superfamilias. Alexander (1992), algunos grupos con hembras especializadas en un tipo Alexander & Michener (1995), Roig-Alsina & Michener particular de presa. (1993), Carpenter (1986a, 1999), Brothers (1999), Brothers Plumariidae es el grupo basal dentro de la superfami- & Carpenter (1993), Shimizu (1994), Kimsey (1991, 1992), lia, con distribución gondwánica y alrededor de 18 especies Shattuck (1992), Brandão et al. (1999), Baroni Urbani et al. en el Neotrópico. Claves en Roig-Alsina (1994). (1992), Grimaldi et al. (1997) y Grimaldi & Agosti (2000) Scolebythidae es una pequeña familia descrita para el exploran la filogenia de varias familias y subfamilias dentro Neotrópico y Australia con dos géneros y tres especies en de Aculeata. el Neotrópico (Evans, 1963; Azevedo, 1999). Hymenoptera Vespomorpha en la Región Neotropical 103 Bethylidae es la familia más grande de la superfami- Tabla I. Sinapomorfias propuestas para Vespomorpha lia, con más de 3.000 especies en el Mundo. Algunas y Apoidea + Vespoidea especies son ectoparásitas de coleópteros, microlepidópte- Sinapomorfia de Vespomorpha: ros y avispas esfécidas. Se conocen menos de 500 especies Hembras con cerci ausentes. en la Región Neotropical (Evans, 1964). Sinapomorfias de Apoidea + Vespoidea (Aculeata s. str.): La segunda familia en número de especies es Chrysi- Machos con 13 antenómeros y hembras con 12. Hembras con el séptimo tergo metasomal oculto y didae (“avispas cuclillo”).
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