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Neevia Docconverter Capítulo I Neevia docConverter1 5.1 INTRODUCCIÓN GENERAL La familia Rhabdiasidae Railliet, 1915 La familia Rhabdiasidae Railliet, 1915 está compuesta por siete géneros y presenta una distribución cosmopolita (Baker, 1978). Sus especies en la fase adulta son hermafroditas y parásitas típicamente de los pulmones de diversas especies de anfibios y reptiles (Anderson y Brain, 1982). El género Rhabdias fue establecido por Stiles y Hassall, 1905, indicando a Rhabdias (=Ascaris nigrovenosa) bufonis (Schrank, 1788) como especie tipo, sin embargo, los autores no presentaron una diagnosis para el género (figura 1a). Rhabdias bufonis se encontró alojado en el pulmón de Bufo bufo Linneo, 1768 (=Bufo vulgaris Laurenti, 1768) (Anura). Travassos (1930), Yamaguti (1961) y posteriormente Baker (1978) presentaron una diagnosis del género Rhabdias. Hasta antes del 2006, se habían descrito alrededor de 50 especies del género (tabla 1). En 1927, Pereira adicionó un nuevo género a la familia Rhabdisidae, Acanthorhabdias Pereira, 1927, y designó a Acanthorhabdias acanthorhabdias parásito de Liophis miliaris subespecie miliaris Linneo, 1758 (=Coluber miliaris Linnaeus, 1758) (Serpentes) como especie tipo; éste es un género monotípico distribuido en Brasil. Pereira (1927) y posteriormente Yamaguti (1961), presentaron una diagnosis del género. En 1974, Fernandes y de Sousa, realizaron una redescripción de Acanthorhabdias acanthorhabdias, a partir del material recolectado también de los pulmones de Liophis miliaris. Este género se diferencia principalmente de Rhabdias, en la forma de la región anterior del cuerpo; Acanthorhabdias a diferencia de Rhabdias, presenta entre 8 y 10 estructuras cuticulares piramidales (“protuberancias”) circumorales, seguido por un pequeña cápsula bucal (Pereira, 1927) (figura 1b). Tres años después, se erigió al género Entomelas Travassos, 1930, parásito pulmonar típicamente de saurios (Agamidae y Anguidae), con Entomelas entomelas (Dujardin, 1845), Travassos, 1930 como especie tipo. Travassos (1930) incluyó en el género Entomelas a tres especies que habían sido registradas dentro del género Rhabdias: Entomelas entomelas (Dujardin, 1845), Travassos, 1930, E. dujardini (Maupas, 1916) Travassos, 1930, y E. chamaeleonis (Skrjabin, 1916). Dentro del género Entomelas se han incluido a siete especies, cuatro de ellas distribuidas en la región Paleártica (Travassos, 1930; Sharpilo y Vakker, 1972; Crusz y Sanmugasunderam, 1973; Sharpilo, 1976; Lewin, 1990; Kuzmin, 1996; Kuzmin y Sharpilo, 2000); 1 en la región Oriental (Baker, 1980); 1 especie en la región Etiópica (Baker, 1982), y Neevia docConverter2 5.1 sólo una especie se ha descrito en la región Neotropical (Bursey y Goldberg, 2006) (figura 1c, tabla 1). El género Entomelas difiere de Rhabdias porque presenta una cápsula bucal de grandes dimensiones y dientes (onchia) en la base de ésta (Yamaguti, 1961). Travassos (1930), Yamaguti (1961) y Baker (1980), han presentado diagnosis de este género. En 1943, Yamaguti describió una especie nueva de Rhabdias, R. horigutii Yamaguti, 1943, parásito de Rhabdophis tigrina Boie, 1826 (=Natrix tigrina Stejneger 1907) (Serpentes). En ese trabajo Yamaguti propuso al subgénero Ophiorhabdias Yamaguti, 1943, basándose en la aparente ausencia de una cápsula bucal, cuya característica estaba asociada exclusivamente a aquellas especies de Rhabdias que parasitan serpientes, nombrando así a la especie nueva como R. (Ophiorhabdias) horigutii Yamaguti, 1943. Sin embargo, la ausencia de una cápsula bucal evidente y la aparente relación de esta característica con la especificidad hospedatoria8 hacia serpientes, fue refutada posteriormente, debido a que existen algunas especies del género Rhabdias que presentan esta misma condición morfológica y que son parásitos de anfibios y por otro lado, existen algunas que presentan una cápsula bucal no muy evidente y que son parásitas de reptiles (Sharpilo, 1976). Ballantyne y Pearson (1963) transfirieron a Pneumonema tiliquae Johnston, 1916 de la familia Rictulariidae a la Rhabdiasidae debido a la similitud morfológica y las afinidades biológicas entre las especies y otros rhabdiásidos (figura 1f). Pneumonema tiliqua es parásito pulmonar de Tiliqua scincoides White, 1790 (=Lacerta scincoides White, 1790) (Sauria). Este género monotípico se diferencia del resto de los géneros descritos hasta ese momento para la familia Rhabdisidae, con base en la presencia de espinas cuticulares. Sharpilo (1974) transfirió a las especies de Hexadontophorus ophisauri Kreis, 1940 de la familia Strongylidae a la Rhabdiasidae. La similitud morfológica entre Hexadontophorus ophisauri Kreis, 1940 y Entomelas spp. se mencionó en dicho trabajo, sin embargo, la posición genérica de las especies no fue modificada y no proporcionó una diagnosis que logre diferenciar al género Hexadonthophorus Kreis, 1940 de Entomelas. El género Kurilonema fue propuesto en 1969 por Szczerbak y Sharpilo, designando a K. markovi Szczerbak y Sharpilo, 1969, parásito pulmonar de Eumeces latiscutatus Stejneger, 1907 (=Plestiodon latiscutatus Hallowell, 1861) (Sauria) como especie tipo (figura 1d). El género se distingue de Rhabdias por la presencia de una evidente y gran cápsula bucal, mientras que de Entomelas se diferencia por la ausencia de dientes en la base de la cápsula bucal. Hasta el momento, solo se ha descrito una especie para este género. 8 A lo largo de la tesis se manejarán los términos especificidad hospedatoria y preferencia hospedatoria como indistintos. Neevia docConverter3 5.1 Singh y Ratnamala (1975) erigieron un nuevo género de rhabdiásido, Shortia, con Shortia shortii Singh y Ratnamala (1975) como especie tipo (en Singh y Ratnamala (1977). Este género se diferencia de Rhabdias por la presencia de glándulas hipodermales (“areolas”). Otro género, Paraentomelas Sharpilo, 1976 fue erigido para incluir a dos especies, P. dujardini (Maupas, 1916) y P. kazachstanica (Sharpilo y Vakker, 1972) las cuales previamente habían sido incluidas dentro del género Entomelas. Este autor consideró que la presencia de una cobertura cuticular ensanchada a lo largo del cuerpo y la forma de la cola (“forma de aguja”), son caracteres válidos para diferenciarla del género Entomelas. En 1980, Baker realizó una revisión taxonómica de la familia Rhabdiasidae y en particular sobre la taxonomía del género Entomelas. En su trabajo, Baker (1980) propone la sinonimia de los géneros Kurilonema, Hexadontophorus y Paraentomelas con el género Entomelas. Adicionalmente, Paraentomelas dujardini, P. kazachstanica y H. ophisauri fueron sinonimizados con E. entomelas. Por otra parte, el subgénero Ophiorhabdias y el género Shortia se consideraron como sinónimos del género Rhabdias, mientras que Entomelas chamaeleonis (Skrjabin, 1916) Travassos, 1930 se transfirió a Rhabdias. Esta propuesta se basó principalmente en características del desarrollo larvario tardío en la fase parasítica observadas en Rhabdias (Baker, 1979) y dicha característica fue extrapolada al desarrollo en Entomelas spp. Aunado a ésto, se tomó en cuenta la información de la forma y tamaño de la cápsula bucal, la presencia o ausencia de dientes (onchia), número de dientes, la cubierta cuticular, etc. En resumen, Baker (1980) consideró como válidos a 4 géneros en la familia Rhabdiasidae: Acanthorhabdias Pereira, 1927; Entomelas Travassos, 1930; Pneumonema Johnston, 1916 y Rhabdias Stiles y Hassall, 1905. La propuesta fue aceptada por Anderson y Bain (1982). En 1989, Hasegawa estableció un nuevo género en la familia Rhabdiasidae, Neoentomelas Hasegawa, 1989, designando a N. asatoi Hasegawa, 1989, parásito de Ateuchosaurus pellopleurus Hallowell, 1861 (Sauria) como especie tipo (Hasegawa, 1989) (figura 1e). La especie se distingue de Entomelas Travassos, 1930 (sensu Baker 1980), por presentar una forma “aberrante” de la región anterior con 4 pseudolabios (2 dorsales y 2 ventrales), y por un marcado ensanchamiento de la región cefálica, formando lóbulos hacia la base de la región cefálica. El sistema presentado por Baker (1980) fue modificado por Kuzmin (1996) y Kuzmin y Sharpilo (2000, 2002), quienes realizaron investigaciones sobre el desarrollo larvario para 4 especies del género Entomelas; a partir de esta información, se han considerado como válidas a E. dujardini, E. kazachstanica y H. ophisauri. Estos autores consideraron que la presencia de Neevia docConverter4 5.1 dientes en la cápsula bucal de Entomelas dujardini, E. kazachstanica, H. ophisauri, E. entomelas y en E. cruszi Baker, 1980, es una característica diagnóstica para el género Entomelas. Con base en esta afirmación se considera válida la sinonimia de Paraentomelas y Hexadonthophorus con Entomelas (Kuzmin, 1996; Kuzmin y Sharpilo, 2000; 2002). Por otro lado, Kuzmin y Sharpilo (2000; 2002) restablecieron a Kurilonema como un género válido, debido a que éste carece de dientes en la base de la cápsula bucal. Lhermitte-Vallarino et al. (2005) describieron el último género para la familia Rhabdiasidae, Chabirenia Lhermitte-Vallarino et al., 2005 con una sola especie, Chabirenia cayennensis, la cual se encontró en la mucosa de la cavidad bucal de lagartijas (figura 1g). Además de diferir de otros géneros de la familia por su inusual hábitat, la especie nueva posee crestas cuticulares longitudinales (similar a un sínlofo, característico descrito para algunos nemátodos tricostróngilidos). Acorde con la información recapitulada, la familia Rhabdiasidae Railliet, 1916 está compuesta por 7 géneros (Tabla 1): Rhabdias Stiles y Hassall, 1905; Pneumonema Johnston,
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