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Half Century in Collembola (Hexapoda) Systematics and Phylogeny

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Half Century in Collembola (Hexapoda) Systematics and Phylogeny MEDIO SIGLO EN LA SISTEMÁTICA Y FILOGENIA DE COLLEMBOLA (HEXAPODA) Half Century in Collembola (Hexapoda) Systematics and Phylogeny José G. Palacios-Vargas. Laboratorio de Ecología y Sistemática de Microartrópodos, Dpto. Biología, Facultad de Ciencias, UNAM, 04510 México, D. F., México. E-mail: [email protected] Palabras Clave: Collembola, sistemática, filogenia, actualización. Introducción. Los colémbolos son un grupo de artrópodos que anteriormente eran considerados como insectos apterigotos, y ahora se consideran como una clase separada de los insectos, dentro de la superclase Hexapoda. Son entognatos, es decir tienen las mandíbulas y las maxilas dentro de una cavidad bucal, carecen de alas, miden por lo general menos de 2 mm de longitud, son comunes y abundantes en diferentes microhábitats, y con frecuencia se registran densidades hasta de más de 100,000 individuos por m2 (Hopkin 1997, Palacios-Vargas y González, 1995). Su alta capacidad de dispersión les ha permitido conquistar diferentes ambientes con climas extremos tales como los desiertos y las regiones polares (Hopkin 2002), así como también las zonas tropicales y templadas. Los ambientes en los que se encuentran son: suelo, hojarasca, musgo, corteza, guano, epífitas y los suelos suspendidos en el dosel de los bosques y selvas (Hopkin 1998; Palacios-Vargas, et al. 2000). Sin embargo hasta la fecha sólo se han descrito cerca de 8,000 especies a nivel mundial. La clasificación ecológica del grupo es interesante ya hay formas altamente modificadas al ambiente en que viven (Cuadro. 1). Los colémbolos son importantes por su influencia sobre la estructura edáfica. La mayoría de los suelos tienen millones de sus heces, mismas que pueden ser benéficas al retardar la liberación de nutrientes esenciales a las raíces de las plantas, así como servir de sustrato para el desarrollo de gran cantidad de microorganismos. Por otro lado, los colémbolos son presa de muchos insectos, en particular de hormigas y escarabajos, así como de numerosos ácaros depredadores, y aunque poco conocido también pueden servir de alimento para algunas aves, reptiles y hasta peces, lo que les confiere gran importancia como elemento fundamental de las cadenas tróficas de todos los ecosistemas, incluyendo biotopos como la superficie de los remansos de ríos, lagunas, y litorales marinos. Los únicos ambientes que no han podido colonizar son las aguas profundas y el mar abierto sin embargo podemos proponer la siguiente clasificación para las formas asociadas a la superficie del agua (Fig. 2). De cerca de 8,000 especies de colémbolos que se conocen en el mundo, en México se tienen registros de 714 especies de colémbolos ubicadas en poco más de 107 géneros y 22 familias (Castaño-Meneses, 2005; Palacios-Vargas et al. 2004, 2007). Al analizar la cantidad de colémbolos que han sido descritos de México desde la primera especie que se describió en 1898, se nota que en el último medio siglo, de 1968 hasta el 2008 se ha descrito la mayor cantidad de especies, y que en la década de 1978 a 1987 es cuando se describieron más especies (Cuadro. 3). 918 Fig. 1. CLASIFICACION ECOLÓGICA DE LOS COLÉMBOLOS Categoria/Caracteristicas Ambientes y ejemplos Ojos Pigmento Antenas Patas Fúrcula Faneras Habitat/ Habitos y Familias Epiedáficas 8 + L L L - dosel Salina(Paronellidae), Deuterosminthurus (Bourletiellidae) Hemiedáficas Normal 8 + C C C - hojarasca Ceratophysella, Xenylla (Hypogastruridae) Xeromorfa 8 ó + - C C C - musgos, líquenes menos Friesea (Neanuridae) 8 ó + - C-L C C - superficie agua Epineústicas menos Sminthurides(Sminthrididae), Ballistura (Isotomidae) Litorales 8 ó - + C C C + arena menos Actates (Actaletidae) , Archisotoma (Isotomidae) Euedáficas 0 - C C C-R - Suelo y cuevas Onychiurus (Onychiuridae) Mesaphorura (Tullbergiidae) Sinecomorfas 0 - L L L + hormigueros y termitas Cyphoderus (Cyphoderidae) Troglomorfas 0 - L L L + cuevas y grutas Arrhopalites (Arrhopalitidae), Pseudosinella (Entomobryidae) Epiedáficas 8 + L L L - dosel Salina (Paronellidae), Deuterosminthurus (Bourletiellidae) (C = cortas, L = largas, R = reducida) Fig. 2. Categorías de colémbolos acuaticos ______________________________________________________________________________ 1. Epineústicos. Son los que viven sobre la capa superficial del agua dulce, como Sminthurides. Pueden caminar y hasta brincar sobre el agua, generalmente no se encuentran en otros ambientes. 2. Epineusfílicos. Frecuentemente se encuentran en la superficie del agua dulce, ya que viven muy cerca de la orilla de los lagos y estanques, como Ballistura. 3. Epineusxénicos. Son los que en realidad viven en otros ambientes, nunca en el agua. Si ellos caen en la superficie del agua pueden morir, ya que no pueden caminar sobre ella. Ceratophysella. 4. Litorales. Viven en las costas del mar, generalmente en las rocas. Ellos tienen adaptaciones para vivir en el agua marina y pueden ser sumergidos por las olas, pero sobreviven. Los machos tienen antenas prensiles para sujetar a las hembras. Actaletes. 5. Psamnóticos. Están adaptados para vivir en la arena de las playas. Su cuerpo es alargado y no se encuentran en otros ambientes.Isotogastrura. ______________________________________________________________________________ Al analizar la cantidad de especies que se han registrado para cada estado de la república mexicana (Fig. 2), se observa que en algunos sólo se conocen unas cuantas y que los que mayor cantidad tienen son Veracruz (129), Estado de México (107) y Guerrero (95); sin embargo, esto 919 está relacionado con los estudios que se han llevado a cabo y no con la verdadera riqueza del lugar. Fig. 1. Crecimiento de especies por decadas Fig. 2. Total de especies registradas por estado La variación en la sistemática del grupo ha variado mucho en medio siglo, no sólo de su posición dentro de los Arthropoda, sino también la cantidad de familias que incluye. En antiguos libros de Entomología (de hace medio siglo) se consideraban cuatro o cinco familias (Poduridae, 920 Onychiuridae, Entomobryidae, Sminthuridae y Neelidae). Actualmente se han elevado varios niveles taxonómicos y en página de colémbolos del mundo (Bellinger et al., 2009) se hace un arreglo de los colémbolos en 30 familias actuales (Cuadro. 3). Cuadro. 3. Total de especies registradas por estados ORDEN Poduromorpha Entomobryomorpha Neelipleona Symphypleona Superfamilia Neanuroidea Tomoceroidea Neelidae Sminthurididoidea Familia Neanuridae Oncopodurida Mackenziellidae Familia Brachystomellidae Tomocerida Sminthuridida Familia Odontellidae Superfamilia Poduroidea Isotomoidea Katiannoidea Familia Poduridae Isotomidae Katiannidae Superfamilia Hypogastruroidea Entomobryoidea Spinothecidae Familia Gulgastruridae Entomobryidae Arrhopalitidae Familia Hypogastruridae Paronellidae Familia Pachytullbergiidae Cyphoderida Familia Paleotullbergiidae Microfalculidae Sminthuridae Bourletiellidae Superfamilia Onychiuroidea Paronellidae Dicyrtomoidea Familia Onychiuridae Dicyrtomidae Familia Tullbergiidae Superfamilia Isotogastruroidea Actaletoidea Familia Isotogastruridae Actaletidae Familia Coenaletidae Algunos autores han propuesto que hasta el momento se conoce sólo el 13% de la fauna de insectos existentes, al extrapolar el número de especies citadas de colémbolos para México, se puede pensar que la diversidad en el país podría alcanzar las 5,000 especies (Palacios-Vargas, 2000). Los colémbolos son interesantes desde varios puntos de vista, pueden estar entre los primeros artrópodos colonizadores de la tierra firme y su relaciones filogenéticas con otros taxa aún está en debate (Giribet et al., 2006; Grimaldi y Engel, 2005; Machida, 2006). A pesar de esto su relaciones filogenéticas internas son poco conocidas. La sistemática de los colémbolos ha tenido una base principalmente intuitiva (Cassagnau, 1974), y la pequeña cantidad de trabajos hechos sobre el grupo puede deberse a las dificultades para su estudio, que incluyen su pequeño tamaño (con frecuencia menor de 1 mm). Las aproximaciones filogenéticos en este grupo son recientes. Incluyen estudios basados en la evidencia morfológica (Lee, 1985; Deharveng, 1989; Deharveng y Bedos, 1991; D'Haese y Weiner, 1998; Simón Benito, et al, 2005), datos moleculares (Dell' Ampio, et al., 2002; D' Haese, 2002, Frati y Carapelli, 1999; Frati y Dell' Ampio, 2000; Frati, et al. 1997), o en la evidencia total (Lee y Thibaud, 1998; Lee, et al., 1995, 1996). Estas aproximaciones han cubierto a menudo cantidades limitadas de especies dentro de géneros seleccionados tales como Friesea (Deharveng y Bedos, 1991) o Willemia (D'Haese y Wiener, 1998), raramente incluyendo la mayoría de las especies de un género dado (D'Haese, 2000), o a niveles taxonómicos más altos (D'Haese, 2003a, 921 2003b). Los estudios basados en datos moleculares del RNA mitocondrial se han vuelto más frecuentes en la actualidad para tratar de ver las relaciones filogenéticas entre los colémbolos (Xiong, et al.). Literatura Citada. Bellinger, P.F., K.A. Christiansen & F. Janssens. 1996-2009. Checklist of the Collembola of the World. http://www.collembola.org Cassagnau, P. 1974. Chétotaxie et phylogénie chez les Collemboles Poduromorphes. Pedobiologia, 14 (2/5):300-312. Castaño-Meneses, G. 2005. Catálogo de los colémbolos (Hexapoda: Collembola) de Sian Ka’an, Quintana Roo, México (Reseña). Rev. Mex. Biodiversidad, 108:107-108. Deharveng, L. 1989. In Dallai, R. [Ed.]. The problem of homoplasies in Neanurinae (Insecta: Collembola): microcomputer assisted phylogeny of the genus Deutonura. 3rd International Seminar on Apterygota,
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