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Dendroctonus Pseudotsugae HOPKINS (COLEOPTERA: CURCULIONIDAE: SCOLYTINAE)

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Dendroctonus Pseudotsugae HOPKINS (COLEOPTERA: CURCULIONIDAE: SCOLYTINAE) INSTITUTO POLITÉCNICO NACIONAL ESCUELA NACIONAL DE CIENCIAS BIOLÓGICAS SECCIÓN DE ESTUDIOS DE POSGRADO E INVESTIGACIÓN HISTORIA DEMOGRÁFICA Y FILOGEOGRAFÍA DE Dendroctonus pseudotsugae HOPKINS (COLEOPTERA: CURCULIONIDAE: SCOLYTINAE). T E S I S que como uno de los requisitos para obtener el grado de DOCTOR EN CIENCIAS QUIMICOBIOLÓGICAS PRESENTA: ENRICO ALEJANDRO RUIZ CASTILLO MÉXICO, D. F., JUNIO DE 2009. El presente trabajo se realizó en el Laboratorio de Variación Biológica y Evolución de la Escuela Nacional de Ciencias Biológicas del Instituto Politécnico Nacional bajo la dirección del Dr. Gerardo Zúñiga. La presente investigación formó parte del proyecto "Estructura genética poblacional de tres especies del género Dendroctonus (Coleoptera: Curculionidae: Scolytinae) por medio de marcadores moleculares", CGPI 2004978, CGPI 20051200 y CONACyT 44887-Q. El sustentante fue becario del Instituto Politécnico Nacional (Beca Institucional) de Febrero a Diciembre de 2005; del Consejo Nacional de Ciencia y Tecnología (CONACyT), de Febrero de 2006 a Diciembre de 2008 y del Programa Institucional De Formación De Investigadores (PIFI) del Instituto Politécnico Nacional, de Septiembre de 2006 a Diciembre de 2008. HISTORIA DEMOGRÁFICA Y FILOGEOGRAFÍA DE Dendroctonus pseudotsugae HOPKINS (COLEOPTERA: CURCULIONIDAE: SCOLYTINAE). Ruiz Castillo Enrico Alejandro. _________________________________________________________________________________________________________________________ RESUMEN El propósito de este estudio fue la estimación de diferentes aspectos de la genética y la biología evolutiva del escarabajo descortezador Douglas-fir, Dendroctonus pseudotsugae: el efecto del aislamiento geográfico sobre la estructura genética; la estimación del status taxonómico de sus subespecies; y la historia demográfica y filogeografía de las poblaciones de D. pseudotsugae. Debido a que las respuestas a estas preguntas derivadas de los tres aspectos a evaluar requieren la implementación de muchos métodos diferentes (que a su vez parten de bases conceptuales diferentes), el presente estudio fue dividido en tres capítulos. Todos ellos estuvieron basados en un fragmento de 550 pb de la Citocromo Oxidasa I (COI) del DNA mitocondrial. Para examinar el efecto del aislamiento geográfico sobre la estructura genética (capítulo I) se analizaron 60 haplotipos del gen COI (172 secuencias de 550 pb). Los resultados de los análisis molecular de varianza (AMOVA y SAMOVA), estadísticos F, y regresiones lineales sugieren que la estructura genética de D. pseudotsugae está fuertemente influenciada por la distancia geográfica. También se encontró que la distancia p está correlacionada con las distancia. Las diferencias genéticas observadas entre el norte (Canadá y USA) y el sur (México) en D. pseudotsugae confirman que estos dos conjuntos de poblaciones corresponden a las subespecies previamente asignadas. En el capítulo II, se estimó la validez de la nueva subespecie D. barragani, debido a que su descripción estuvo basada en una única población conocida en México. Para probar si más poblaciones apoyan la existencia de dos subespecies, se realizó una revaluación taxonómica combinando secuencias de COI, los caracteres morfológicos usados en la descripción original, y caracteres morfológicos recientemente descritos. El análisis de máxima parsimonia con 89 haplotipos confirmó que las poblaciones mexicanas son distintas de aquellas provenientes de las localidades del norte (Canadá y USA). Por i HISTORIA DEMOGRÁFICA Y FILOGEOGRAFÍA DE Dendroctonus pseudotsugae HOPKINS (COLEOPTERA: CURCULIONIDAE: SCOLYTINAE). Ruiz Castillo Enrico Alejandro. _________________________________________________________________________________________________________________________ otro lado, mientras que al menos siete caracteres de la cabeza, el pronoto y los élitros permiten distinguir consistentemente entre las poblaciones de Canadá y USA por un lado, y las de México por otro. Finalmente, en el capítulo III se evaluó si la distribución actual en Norteamérica de D. pseudotsugae ha sido moldeada por las glaciaciones pasadas del periodo Cuaternario. Por otro lado, se cree que el efecto de dichas glaciaciones no fue tan severo en las Montañas Rocosas del sur y en el norte de México. En este trabajo, se pusieron a prueba distintas hipótesis acerca de la localización de refugios del pasado en el Pacifico Noroeste y en las Montañas Rocosas, y se probó también si las poblaciones del sur de Norteamérica y México son más diversas y han experimentado tamaños poblacionales constantes a lo largo del tiempo, en oposición a aquellas de distribución más boreal. Las hipótesis de expansión demográfica de los haplogrupos (identificados a partir de 136 haplotipos de 331 secuencias del gen COI) fueron examinadas a través de pruebas de neutralidad contra crecimiento poblacional, mismatch distribution, y Bayesian Skyline Plots. Los resultados mostraron que los haplogrupos del noroeste de Norteamérica y la Sierra Madre Occidental han experimentado eventos de expansión demográfica, pero no los haplogrupos del suroeste de Norteamérica y de la Sierra Madre Oriental. Los tiempos de divergencia entre pares de haplogrupos fueron estimados entre el Pleistoceno temprano al Pleistoceno medio, mucho tiempo antes del último periodo glacial. Finalmente, el análisis de clados anidados también aportó evidencia de que los cambios demográficos estuvieron acompañados de eventos de expansión de área continua en el noroeste de Norteamérica, mientras que en las poblaciones de Arizona y el norte de México experimentaron fragmentaciones alopátricas y reducción de flujo génico histórico. Los resultados del presente estudio proveen ahora de una base para probar si existe congruencia entre la variación genética de D. pseudotsugae y la correspondiente a la de su huésped. ii HISTORIA DEMOGRÁFICA Y FILOGEOGRAFÍA DE Dendroctonus pseudotsugae HOPKINS (COLEOPTERA: CURCULIONIDAE: SCOLYTINAE). Ruiz Castillo Enrico Alejandro. __________________________________________________________________________________________________________________________ ABSTRACT The purpose of the present study was to assess different aspects of genetics and the evolutionary biology of the Douglas-fir beetle Dendroctonus pseudotsugae: the effect of geographic isolation on genetic structure; the assessment of the taxonomic status of subspecies; and the historical demography and phylogeography of D. pseudotsugae populations. Because answering specific questions raised by all three aspects require many different methods (and therefore, different assumptions), this study was divided in three chapters. All of them are based on a fragment of 550 bp of mtDNA Cytochrome Oxidase I (COI). To examine the effect of geographic isolation on genetic structure (chapter I), we analyzed 60 haplotypes of COI gene (172 sequences). Results of analyses of molecular variance (AMOVA and SAMOVA), F-statistics, and linear regressions suggest that the genetic structure of D. pseudotsugae is strongly influenced by geographic distance. We also found that the p-distance is correlated with geographic distance. The observed genetic differences between north (Canada-USA) and south (Mexico) on D. pseudotsugae confirm that these two sets of populations correspond to previously assigned subspecies. In the chapter II, the validity of a new subspecies of D. pseudotsugae was assessed. This was done because the description of D. p. barragani was based on only one known Mexican population. To test if additional populations support the existence of two subspecies, a taxonomic reassessment combining COI sequences, morphological characters used in the original description, and newly described morphological characters was performed. Maximum Parsimony analysis of 89 haplotypes confirms that the Mexican populations are distinct from those from northern locations (in USA and Canada) sampled. On the other hand, at least seven characters on the head, pronotum, and elytra consistently discriminate among Canada-USA and Mexico populations. Finally, in chapter III we evaluated if contemporary northern distribution iii HISTORIA DEMOGRÁFICA Y FILOGEOGRAFÍA DE Dendroctonus pseudotsugae HOPKINS (COLEOPTERA: CURCULIONIDAE: SCOLYTINAE). Ruiz Castillo Enrico Alejandro. __________________________________________________________________________________________________________________________ of D. pseudotsugae has been shaped by past glaciations of the Quaternary period. It is hypothesized that past glaciations were not that severe in southern Rocky Mountains and Northern Mexico. In this study, we test hypotheses about location of ancient refugia at the Pacific Northwest and Rocky Mountains, and test if southern North American and Mexican populations are more diverse and experienced constant population sizes trough time than northern ones. In this study, 136 haplotypes out of 331 sequences were identified. Hypothesis of demographic expansion of haplogroups was examined trough neutrality tests against population growth, mismatch distribution, and Bayesian Skyline Plots. Results showed that Northwestern North America and Sierra Madre Occidental haplogroups have experienced demographic expansion events while Southwestern North America and Sierra Madre Oriental haplogroups did not. Divergence times between pairs of haplogroups were estimated from early to middle Pleistocene, long before last glacial maxima. Finally, the nested clade analysis provided evidence that the demographic change were accompanied by continuous range expansion
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