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The Bull Kelp Durvillaea Antarctica FACULTAD DE CIENCIAS DEL MAR UNIVERSIDAD CATÓLICA DEL NORTE DOCTORADO EN BIOLOGÍA Y ECOLOGÍA APLICADA “Factors influencing the potential of rafting dispersal - analysis of patterns and causal mechanisms along the geographic range of intertidal species on the coast of Chile” Boris Alejandro López Arriagada Profesor Guía: Dr. Martin Thiel COQUIMBO, 2018 FACULTAD DE CIENCIAS DEL MAR UNIVERSIDAD CATÓLICA DEL NORTE DOCTORADO EN BIOLOGÍA Y ECOLOGÍA APLICADA “Factors influencing the potential of rafting dispersal - analysis of patterns and causal mechanisms along the geographic range of intertidal species on the coast of Chile” Por: Boris Alejandro López Arriagada Departamento de Biología Marina Fecha: 17‐10‐2018 Aprobado Comisión de Calificación ____________________________ Dr. Juan Macchiavello Armengol Decano Facultad de Ciencias del Mar ________________ Dr. Martin Thiel Profesor Guía _______________________ ________________ Dra. Fadia Tala Dr. Marcelo Rivadeneira Comité Tutorial Comité Tutorial ____________________ ____________________ Dr. Moisés Aguilera Dr. Erasmo Macaya Comité Tutorial Profesor Externo Tesis entregada como un requisito para obtener el título de Doctor en Biología y Ecología Aplicada en la Facultad de Ciencias del Mar. Universidad Católica del Norte. Sede Coquimbo. 2018 ii FACULTAD DE CIENCIAS DEL MAR UNIVERSIDAD CATÓLICA DEL NORTE DOCTORADO EN BIOLOGÍA Y ECOLOGÍA APLICADA Departamento de Biología Marina “Factors influencing the potential of rafting dispersal - analysis of patterns and causal mechanisms along the geographic range of intertidal species on the coast of Chile” Actividad de Titulación presentada para optar al Título de Doctor en Biología y Ecología Aplicada Boris Alejandro López Arriagada Coquimbo, Octubre de 2018 iii FACULTAD DE CIENCIAS DEL MAR UNIVERSIDAD CATÓLICA DEL NORTE DOCTORADO EN BIOLOGÍA Y ECOLOGÍA APLICADA DECLARACIÓN DEL AUTOR Se permiten citas breves sin permiso especial de la Institución o autor, siempre y cuando se otorgue el crédito correspondiente. En cualquier otra circunstancia, se deberá solicitar permiso de la Institución o el autor. Boris Alejandro López Arriagada Firma 2018 iv RESUMEN Varios estudios, usando evidencia molecular y ecológica, han confirmado que el rafting contribuye a la conectividad poblacional y la expansión de rangos geográficos de muchas especies. Sin embargo, también hay indicios de que la efectividad del transporte de especies vía rafting es limitada al momento de establecerse en nuevos hábitats. Esto sugiere que el transporte puede no ser el factor limitante para la dispersión de especies, pero que otros factores (ecológicos o fisiológicos) podrían restringir la colonización potencial de estas especies. Asimismo, la dispersión efectiva de una especie podría variar dentro de su rango geográfico. El objetivo de este estudio fue evaluar los patrones de dispersión del alga flotante Durvillaea antarctica (Chamisso) Hariot 1892 y sus epibiontes asociados a lo largo de la costa continental de Chile (28°S-42°S), así como examinar cuales mecanismos afectan la dispersión y conectividad de sus poblaciones dentro del rango geográfico del clado continental de D. antarctica. Se propuso que los factores ecológicos son más importantes en la dispersión efectiva de individuos de D. antarctica en poblaciones centrales de su rango geografico, mientras que en poblaciones periféricas características reproductivas y funcionales de las especies serán más importantes; desviaciones a partir de esta predicción de dispersión efectiva dentro del rango geográfico podría deberse a condiciones oceanográficas locales. La variabilidad espacio-temporal de ejemplares de D. antarctica varados en playas (Capítulo 1), mostró que los varamientos y características de los individuos variaron fuertemente de acuerdo a los distritos biogeográficos a lo largo de la costa de Chile durante tres años consecutivos. El estudio bimensual de varamientos en tres playas (32°S, 36°S y 39°S) durante tres años consecutivos (Capítulo 2) confirmó el patrón espacial de que altas biomasas varadas fueron observadas en playas del norte y sur en comparación a playas de la zona central. Además, se observó un fuerte patrón estacional que estuvo relacionado con la intensidad de tormentas locales. Una prospección a gran escala de ejemplares dispersados por rafting (28°S-42°S) y una a menor escala de ejemplares bentónicos (31°S-32°S) (Capítulo 3) evidenció que la proporción de machos y hembras que viajan juntos en discos coalescentes de D. antarctica fue baja (entre 5% - 17%). Asimismo, el análisis de capacidades funcionales indicó que no hubo diferencias en las concentraciones de pigmentos y florotaninos según el sexo de los individuos. La v evaluación espacio-temporal de la diversidad de epibiontes en discos de ejemplares varados de D. antarctica (Capítulo 4) evidenció que especies sésiles fueron más frecuentes. La riqueza taxonómica de epibiontes fue más alta en los distritos biogeográficos del sur (33°S- 42°S) que en aquellos del norte (28°S-33°S), incrementándose en verano en comparación a invierno. Especies formadoras de hábitat (e.g. mitílidos, algas no flotantes) favorecieron las co-ocurrencias con otros epibiontes dentro de un disco de D. antarctica, estando asociadas a expansiones de rangos geográficos de especies de epibiontes, particularmente en la zona sur (33°S-42°S). Los patrones filogeográficos de dos especies de algas epifitas del género Gelidium, frecuentes en discos de ejemplares varados de D. antarctica fueron contrastantes (Capítulo 5). Gelidium lingulatum tuvo alta diversidad genética, pero su estructura genética no siguió un claro patrón geográfico, lo que puede ser explicado por dispersión vía rafting de individuos flotantes de D. antarctica. Por otro lado, G. rex tuvo menor diversidad genética con una débil estructura genética, y un quiebre filogeográfico que coincide con la discontinuidad filogeográfica descrita para la región entre los 29°S-33°S, lo que sugiere eventos de dispersión a corta distancia. Los resultados indicaron que D. antarctica posee una importante potencial de dispersión mediante ejemplares flotantes y de transporte de especies asociadas, pero existirían fuertes fluctuaciones espacio-temporales en la disponibilidad de ejemplares flotantes que retornan a la costa que afectarían su eventual inmigración exitosa en otras áreas. El ciclo anual de crecimiento de las poblaciones bentónicas de D. antarctica, la disponibilidad de sustrato primario, la explotación de sus bancos naturales, y características oceanográficas y topográficas a escala regional (i.e. surgencias, plumas de ríos), y de mayor escala (i.e. el fenómeno del Niño) podrían explicar los patrones observados en este estudio. En general, los resultados sugieren que las características reproductivas y funcionales de los individuos flotantes no serían limitantes en la capacidad de dispersión efectiva y consecuente baja conectividad de sus poblaciones bentónicas, en comparación con factores oceanográficos locales y demográficos de las poblaciones bentónicas residentes. Por lo tanto, este estudio muestra que la efectividad de la dispersión por rafting mediante algas flotantes en zonas templadas varía fuertemente a lo largo del rango geográfico de las especies, así como en escalas temporales. Corrientes locales y regímenes de vientos generan zonas de retención para suministros flotantes que limitan el arribo hacia otras áreas. En general, las resultados de esta tesis sugieren que en vi los límites de baja latitud del rango de una especie, el stress fisiológico suprime el potencial de dispersión de algas flotantes, mientras que en los límites de alta latitud del rango, las fluctuaciones de la disponibilidad de individuos flotantes y las interacciones ecológicas con comunidades bentónicas establecidas afectan la conectividad poblacional. Palabras Claves: algas flotantes; biogeografía marina; cochayuyos; dispersión por rafting; varamientos. vii ABSTRACT Several studies, using both molecular and ecological evidence, confirm that rafting contributes to population connectivity and range expansion of many species. However, there is also indication that the effectiveness of rafting transport of species to establish in new habitats is limited. This suggests that transport might not be the limiting factor for the dispersal of species but that other factors (ecological or physiological) may restrict the colonization potential of these species. Also, this effective dispersal of a species could vary within its geographic range. The aim of this study was to assess the dispersal patterns of the floating bull kelp Durvillaea antarctica (Chamisso) Hariot 1892 and its associated epibionts along the continental coast of Chile (28°S-42°S) as well as to examine which mechanisms affect dispersal and connectivity of their populations within the geographic range of the continental clade of D. antarctica. It was proposed that ecological factors are more important in the effective dispersal of individuals of D. antarctica in central populations of its geographic range, while in peripheral populations reproductive and functional characteristics of the species are more important; deviations from the predictions of the effective dispersal within the geographic range could be due to local oceanographic conditions. The spatio-temporal variability of stranded bull kelps on beaches (Chapter 1), showed that strandings and morphometric characteristics of rafts varied strongly according to biogeographic districts along the coast of Chile during three consecutive
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