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Paleobioindicadores Lacustres Neotropicales El libro cuenta con once capítulos sobre los grupos de paleoindicadores biológicos (unicelulares, vegetales y zoológicos) más utilizados en reconstrucciones ambientales y climáticas, en los que se compila información detallada de cada uno de ellos, técnicas y PALEOBIOINDICADORES metodologías de trabajo, así como la presentación de diversos casos de estudio y P referencias bibliográficas actualizadas. Rápidamente el lector obtendrá un panorama de los ALEOBIOINDICADORES LACUSTRES NEOTROPICALES LACUSTRES distintos registros biológicos que se utilizan en la Paleolimnología Neotropical. Los dos primeros capítulos describen los bioindicadores unicelulares donde el lector NEOTROPICALES quedará fascinado con la gran cantidad de información que se puede obtener con tan pequeños organismos, posteriormente se presentan los capítulos relacionados con los paleobioindicadores vegetales como polen, fitolitos, pigmentos sedimentarios y material Editores carbonizado con los cuales se pueden realizar inferencias de cambios ambientales Liseth Pérez terrestres en la cuenca de estudio. Avanzando en los niveles de organización, se despliegan Julieta Massaferro los capítulos sobre los paleobiondicadores zoológicos (crustáceos, dípteros y moluscos) Alexander Correa-Metrio más utilizados dada su alta sensibilidad y preferencias ecológicas. Finalmente, con el objetivo de que el lector pueda comprender y posteriormente explorar con la práctica el campo y laboratorio de la paleolímnología se presenta un capítulo enfocado a los métodos estadísticos más utilizados en las reconstrucciones paleoambientales. CODIGO ISBN CODIGO ISBN Paleobioindicadores lacustres neotropicales Editado por Liseth Pérez Instituto de Geología, Universidad Nacional Autónoma de México Ciudad Universitaria, 04510, Ciudad de México, México [email protected] Julieta Massaferro CONICET, CENAC/APN Fagnano 244, Bariloche, Argentina [email protected] Alexander Correa-Metrio Instituto de Geología, Universidad Nacional Autónoma de México Ciudad Universitaria, 04510, Ciudad de México, México [email protected] Asistente de edición Karla Zurisadai Rubio Sandoval Posgrado en Ciencias Biológicas, Universidad Nacional Autónoma de México Ciudad Universitaria, 04510, Ciudad de México, México [email protected] Editorial (UNAM) ISBN: 978-607-30-0209-7 Tapa: Florencia Crocco INDICADORES ZOOLÓGICOS CAPÍTULO 10: Mollusca Capítulo 10: Mollusca Denisse Álvarez Sanhueza, Facultad de Ciencias, Universidad Santo Tomás, Concepción, Chile, [email protected] Claudio G. De Francesco, Instituto de Investigaciones Marinas y Costeras (IIMyC), CONICET- Universidad Nacional de Mar del Plata, Juan B. Justo 2550 1ºPiso, 7600, Mar del Plata, Argentina, [email protected] Palabras clave: Gasterópodos, Bivalvos, Cuaternario, 18O. 1. Introducción Los moluscos (Phylum Mollusca) se encuentran entre los bioindicadores macroscópicos más comúnmente hallados en los sedimentos continentales cuaternarios. Están representados exclusivamente por las clases Gastropoda (caracoles, babosas) y Bivalvia (almejas, mejillones). La fauna dulceacuícola neotropical está representada por al menos 14 familias de gasterópodos, (Strong et al. 2008) y por aproximadamente 10 familias de bivalvos (Bogan 2008). Habitan casi todo tipo de ambientes acuáticos como ríos, arroyos, canales, manantiales, cascadas, lagos, esteros y pantanos. La mayoría de las especies fósiles corresponden a especies que se encuentran en la actualidad, lo que permite una aproximación uniformitarianista directa para inferir paleoambientes. Los moluscos cuaternarios fueron utilizados en todo el mundo para reconstruir ambientes locales y condiciones climáticas en combinación con otros registros biológicos y sedimentológicos. Dado que proveen información sobre la dinámica hidrológica de los cuerpos de agua, resultando potencialmente útiles como bioindicadores paleoambientales (De Francesco 2013). Un gran número de estudios se focalizó en el análisis de los isótopos estables (18O y 13C) presentes en las valvas como un mecanismo indirecto para inferir temperaturas, procesos de evaporación y niveles de productividad (Bonadonna et al. 1999; Leng et al. 1999; Jones et al. 2002; De Francesco y Hassan 2013; Álvarez et al. 2015). Más recientemente han comenzado a usarse para comprender los procesos de fosilización en aguas continentales (tafonomía) para lo cual se ha llevado adelante estudios de tafonomía actualista (Brown et al. 2005; Kotzian y Simões 2006; Martello et al. 2006; Cristini y De Francesco 2012; Tietze y De Francesco 2012, 2014). En la zona neotropical el conocimiento sobre la malacofauna acuática continental fósil es fragmentario, debido a que las investigaciones desarrolladas con anterioridad al presente siglo se han limitado en su mayoría a hacer mención de especies en localidades discretas, sin una visión integradora que permita comprender el origen y los patrones de diversificación de las especies durante el Cenozoico (véase del Río et al. 2007 para una síntesis del estado del conocimiento). Además, dentro de los 192 PALEOBIOINDICADORES LACUSTRES NEOTROPICALES escasos estudios que se han llevado adelante existe un sesgo importante hacia los depósitos aluviales (e.g., Prieto et al. 2004; De Francesco et al. 2007) siendo escasos los estudios desarrollados sobre moluscos cuaternarios en sedimentos lacustres (De Francesco et al. 2013). El objetivo de este capítulo es realizar una síntesis de las principales líneas de investigación desarrolladas en América Latina, en relación a la reconstrucción paleoambiental y paleoclimática basada en moluscos lacustres. 2. Biología Los moluscos se caracterizan por presentar una concha externa de carbonato de calcio compuesta por una (gasterópodos) o dos piezas (bivalvos), que es generalmente la única estructura que se preserva en el registro fósil. Por lo tanto, la identificación taxonómica se realiza a partir de los caracteres diagnósticos presentes en la concha (Figura 1). Esto resulta confiable sólo en casos donde existe una relativa estabilidad morfológica, mientras que se vuelve muy dificultoso en aquellos casos en que las conchas exhiben convergencia morfológica entre distintas especies y plasticidad fenotípica, como por ejemplo los caracoles de la superfamilia Rissooidea (De Francesco 2007). Figura 1. Características morfológicas de las conchas de gasterópodos (izquierda) y bivalvos (derecha). Modificado de De Francesco (2013). Diversidad y Taxonomía La taxonomía de los moluscos dulceacuícolas se halla al presente siendo revisada sustancialmente a nivel mundial, debido a la incorporación de nuevos caracteres anatómicos, ultraestructurales y moleculares (véase Bouchet y Rocroi 2005). Además de esta situación, en América Latina existe escasa información para muchas regiones, lo que vuelve la actualización del conocimiento taxonómico aún más complicada. 193 INDICADORES ZOOLÓGICOS CAPÍTULO 10: Mollusca En la región neotropical los bivalvos están representados por un total de 9 familias, 52 géneros y 226 especies, incluidos en tres subclases: Pteriomorpha, Paleoheterodonta y Heterodonta (Tabla 1). La subclase Pteriomorpha solamente está representada por una especie exótica, el mejillón dorado Limnoperna fortunei, que ha invadido recientemente diferentes áreas de América Latina, causando importantes problemas ecológicos y ambientales (Darrigran y Pastorino 2004). La subclase Paleoheterodonta incluye el orden Unioniformes, uno de los grupos con mayores radiaciones en agua dulce, estando representado por las familias Hyriidae (con 9 géneros), Mycetopodidae (12 géneros) y Unionidae (20 géneros), estos últimos, constituidos por especies exóticas. Los bivalvos de las familias Hyriidae, Mycetopodidae y Sphaeriidae dan cuenta de la mayoría de la diversidad faunística autóctona de América Latina (con un total de 135 especies) (Bogan 2008). Tabla 1. Clasificación taxonómica y número de géneros y especies de bivalvos presentes en ambientes dulceacuícolas de la región neotropical. Datos tomados de Bogan (2008), Rumi et al. (2008) y De Francesco (2013). *indica taxa exóticos. Clase Subclase Orden Familia Géneros/ Ejemplos Especies Bivalvia Pteriomorpha Mytiloida Mytilidae* 1/1 Limnoperna fortunei Paleoheterodonta Unioniformes Hyriidae 9/55 Diplodon chilensis Mycetopodidae 12/39 Anodontites trapesialis Unionidae* 20/85 Elliptio complanata Heterodonta Veneroida Corbiculidae* 2/2 Corbicula fluminea Sphaeriidae 5/41 Musculium argentinum Myoida Erodonidae 1/1 Erodona mactroides Teridinidae 1/1 Nausitora dunlopei Lyonsiidae 1/1 Anticorbula fluviatilis Por su parte, los gasterópodos neotropicales, con una diversidad mayor que la de los bivalvos (Tabla 2), están dominados por dos clados principales, que son los Caenogastropoda y los Heterobranchia (Bouchet y Rocroi 2005). También hay algunos representantes (~10 especies) de la familia Neritidae (dentro del clado de los Neritimorpha) que fueron previamente incluidos como un subgrupo de los ‘arqueogasterópodos’. Los Caenogastropoda (antiguamente conocidos como Prosobranchia) comparten unos pocos caracteres primitivos (probablemente ancestrales) como la respiración a través de branquias y la presencia de un opérculo. Usualmente son de sexos separados. En la región neotropical están representados por 7 familias, de las cuales la más diversa es la de los Cochliopidae (fundamentalmente por el género Heleobia), seguida por los Ampullariidae. Este grupo ha invadido los ambientes de agua dulce en varias oportunidades a lo largo de la historia de la vida, al menos tantas veces como
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