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“Patrones Filogeográficos En El Gastrópodo Marino Acanthina

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“Patrones Filogeográficos En El Gastrópodo Marino Acanthina ;6<=6 !"# $ %&'(()*+ ,##&- -. / $$023 4 56567 6 899:2 Agradecimientos Quiero agradecer a las personas que han aportado en mi formación de profesional y más como persona, y que han hecho que este proceso responda a mis expectativas y culmine de buena manera a través de la presente tesis de investigación. En primer lugar quiero agradecer de forma especial a mi núcleo familiar por su apoyo incondicional en todos mis actos (papá, mamá y Tania), destacando que en gran parte, gracias a ellos eh logrado cumplir este proceso como una de mis metas... a ellos dedico este trabajo y dedicare muchos mas. Por el lado académico agradecer y destacar a Leyla por ser la guía principal de mi tesis y en esta ultima etapa de formación como profesional ayudarme a desarrollar nuevas temáticas y actitudes personales, además mención especial para el espacio físico y apoyo logístico que me otorgo, y que permitieron el desarrollo de esta tesis de investigación, a Roger por su activa participación directa, como copatrocinante y en mis primeras asistencias a congresos, y a Antonio por sumarse a mi tesis, colaborando y mostrando siempre buena disposición a pesar de la distancia. También de cerca destaco el apoyo de mi polola (Katty), de mi círculo cercano de primos y mis amigos de barrio, y a mis compañeros de laboratorio: Daniela, Chalo, Jano y Zambra con los cuales afrontamos esta etapa en tiempos similares apoyándonos mutuamente. A todos... Gracias... 2 >; > 6 2 ! " # $%& " ' &"% ( "& ) &*%* +, -./""& 1* 2, *13*& ! 2+ *13*"".$*" 2+ & !4** 22 &"%! *" 2) "%"* #5 &$& " '5 ! " 55 $%& " 5) 6*" (, 3 >?'2 @A >?8 !" #$$%& ! '& !(& ) ! *& @A >? B + ,- !+,+& ./0 !..1+1.& + 0 ,- !/+,+& 2 3 4 % @( >?) , - !56 7/& ) !89 : $ $%& @( >? @2 - - * ; *< *=< = 0 - @C 4 D-' ? - - ./0 = )= A9 D-8 - - * @& 2& 3& 4& %& A& A' D-B @$ * - * = A8 D- ) - !7 *& - ! & ./0 )= !.. . < + B < . & AB D-@ 5 !( @C@$$$& - !"& ?2 9 $ %4D . 9 $ $$@ A) 5 D-A ? * 2%D 4A * = * - 0 E = ' * !& 7 @ !7@& 7 2 !72& A@ D-( + - * !+& 5GC 1@ H$@ 9 $ @$ !I& . . 5GC 1@ 2D@ : $ @$ !& B 5GC 12 434 9 $ $@ !& . 5GC $ 22D : $ @$ 0 9 G - - 9 AA D-C +& , J I& - !E - & 9 - = = A( D-: ,- E ! & AC 6 D- '9 0 ' - !* @& 9 - 9 , ? 9 / ? = " 9 " 5 A: 7 RE SU M E N RESUMEN . Recientes avances en ecología molecular de especies marinas en otros océanos han mostrado que las bases históricas de diversidad necesitan ser reconocidas y entendidas (i.e. diferentes trayectorias de evolución de las especies). Además a nivel contemporáneo, varios estudios genéticos en especies marinas han demostrado la presencia de subdivisión poblacional, limites a la dispersión y barreras a la conectividad en el ambiente marino. Esto limita nuestro entendimiento de los procesos (históricos y contemporáneos) que conducen a la presente distribución de las especies. A lo largo del Pacífico Sureste (PSE) numerosos gastrópodos marinos de la familia Muricidae son endémicos de esta costa y constituyen especies claves en las comunidades intermareales de la región. Acanthina monodon (Pallas, 1774), conocido localmente como caracol con diente, se distribuye en el intermareal y submareal somero, desde los 28º a 55º S, posee desarrollo holobentonico, con ausencia de una etapa larval de dispersión en la columna de agua. En este trabajo se estudia, a través de marcadores moleculares y morfometría geométrica, el patrón geográfico de distribución de la diversidad genética y morfológica de A. monodon. Dada la extensa distribución de A. monodon esta espacie atraviesa todas la zonas biogeográficas propuestas para el PSE (Provincia Peruana, Área Intermedia, Provincia Magallánica) de aquí que, basados en una aproximación filogeográfica, se evalúa el efecto de los quiebres biogeográficos en esta especie. Los resultados indican que este gastrópodo marino presenta una alta diversidad genética a nivel de especie, con fuerte disminución de ésta hacia los extremos del rango de distribución y 8 con marcada estructuración genética-poblacional asociada a la biogeografía de la zona de estudio, sugiriendo influencia de la historia de vida de la especie y eventos históricos del área. Los patrones morfológicos demostraron variabilidad a través del rango estudiado, con variación morfológica asociada a las diferentes localidades y un quiebre morfológico concordante con un cambio en la conformación física de la costa del PSE. 9 ABSTRACT. Recent advances in molecular ecology of marine species in other oceans have shown that the historical foundations of diversity needs to be recognized and understanding (i.e. different paths of evolution of species). In addition to contemporary level, several genetic studies on marine species have demonstrated the presence of population subdivision, limits to dispersal and barriers to connectivity in the marine environment. Throughout the Southeast Pacific (SEP) Acanthina monodon (Pallas, 1774) (Gastropoda: Muricidae) is distributed in the intertidal and shallow subtidal, from 28 º to 55 º S, this species has holobenthic development with absence larval dispersal stage in the water column. In this study we analyzed, through molecular markers and geometric morphometrics, the geographical pattern of distribution of genetic diversity and morphology of A. monodon. Given its wide distribution, this species crosses all biogeographic zones proposed for the SEP, and therefore based on a phylogeographical approach; we evaluated the effect of biogeographical breaks in this species. The results indicate that this marine gastropod has a high genetic diversity at the species level, with sharp decline of this towards the ends of the range of distribution and with and strong population genetic structure correlated with the biogeography pattern of the study area. The morphological patterns showed variability across its range distribution associated with different localities and a morphological break consistent with a change in the physical conformation of the coast along SEP. 10 INTRODUCCIO N. La filogeografía es el campo de estudio relacionado con los principios y procesos que gobiernan la distribución geográfica de linajes genéticos, sobre todo aquellos entre y dentro de especies cercanamente relacionadas (Avise et al., 2000). Se ha situado a la filogeografía como punto de unión o intersección entre la macroevolución y la microevolución (Avise et al., 1987). La filogeografía enfatiza aspectos históricos de la distribución espacial actual de linajes de genes utilizando información temporal (histórica) que ha sido posible obtener a partir del uso y análisis de la información contenida en las moléculas de ADN. Por supuesto, para el análisis e interpretación de las distribuciones de linajes se requiere de la contribución teórica, metodológica y/o conceptual de campos de conocimiento muy diversos, como la genética de poblaciones, genética molecular, demografía, etología, filogenia, paleontología, geología y geografía histórica (Avise et al., 2000). Parte de la historia del surgimiento de la filogeografía viene entrelazada no sólo con el desarrollo de técnicas moleculares, sino también con el conocimiento de la molécula de ADN mitocondrial (mtADN) a mediados de la década de los 70’s. Sus propiedades más interesantes en términos filogeográficos son: (i) su alta tasa de evolución (sustitución) a nivel de secuencias de nucleótidos; (ii) su prácticamente nula recombinación; (iii) su gran variación intraespecífica y; (iv) su herencia estrictamente materna (con algunas excepciones). Una de las aplicaciones más amplias de los estudios filogeográficos ha sido el poder determinar el grado de estructuración poblacional de las especies a lo largo de su área de distribución, así como descifrar cuáles han sido los 11 procesos que han determinado dicha distribución. Aplicaciones relevantes de los estudios filogeográficos (Eguiarte et al., 2007) corresponden a: (1) Inferir eventos históricos a través de la identificación de patrones geográficos de variación genética y la reconstrucción de rutas de migración de especies. (2) Interpretar las respuestas bióticas a los cambios paleoclimáticos y paleoecológicos dando lugar a interpretaciones biogeográficas con valor predictivo de frente al inminente escenario de cambio climático acelerado. (3) Comparar los resultados de especies co-distribuidas a fin de comprender los mecanismos de origen de la biodiversidad a una escala regional. (4) Realizar predicciones acerca de los patrones filogeográficos esperados bajo nuevas condiciones, en el pasado o futuro, o en áreas a ser exploradas por su diversidad. (5) Determinar el lugar de origen y las vías de dispersión de especies consideradas plagas o invasoras. (6) Reconocer poblaciones en riesgo de extinción. (7) Brindar datos teóricos para establecer prioridades de conservación de la biodiversidad, entre otros. (8) También frecuentemente abordados a través de la filogeografía han sido aspectos de sistemática y taxonomía. En los últimos años, el estudio de patrones filogeográficos se ha extendido al estudio de múltiples
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