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Conferencias Magistrales CONFERENCIAS MAGISTRALES LAS AGALLAS DE LOS ENCINO: UN ECOSISTEMA EN MINIATURA QUE HACE POSIBLE ESTUDIOS MULTIDISCIPLINARES Juli Pujade-Villar. Universitat de Barcelona, Facultat de Biologia, Departament de Biologia Animal. Avda. Diagonal 645, 08028-Barcelona, Spain. E-mail: [email protected] RESUMEN. Desde que en el siglo XVII el italiano Marcelo Malpighi (Crevalcore, 1628 - Roma, 1694) descubriera la relación causa-efecto entre un insecto y su agalla, numerosos naturalistas y científicos han centrado sus esfuerzos en estas estructuras vegetales, hasta hacer de la Cecidología (Ciencia que se ocupa del estudio de las agallas de las plantas) una ciencia de ámbito multidisciplinar que se asienta en estudios ecológicos, morfológicos y estructurales, etiológicos, taxonómicos, faunísticos, histológicos, fisiológicos, genéticos, etc. En este estudio se hará un repaso de que son las agallas y de los distintos estudios que pueden realizarse a partir de las agallas producidas por los Cynipidae (Hymenoptera). Palabras Clave: agallas, encinos, ecosistema, estudios. The oak galls: a miniature ecosystem which makes possible multidisciplinary studies ABSTRACT. Since the Italian Marcelo Malpighi (Crevalcore, 1628 - Roma, 1694) discovered the cause and effect relationship between an insect and its gall in the XVII century, many naturalists and scientists have focused their efforts in these plant structures to make for the Cecidology (science that deals with the study of plants galls) a multidisciplinary science based on different branches: ecologic, morphological, structural, etiologic, taxonomic, faunistic, histologic, physiologic, genetic, etc. In this work a review of what the galls are and the diverse studies which can be carried out from the Cynipidae (Hymenoptera) galls is made. Key words: galls, oaks, ecosystem, studies. Introducción Numerosos paseantes, naturalistas, estudiantes, agrícolas, etc. han observado, en algún momento de su vida, agallas o deformaciones vegetales inducidas por otros organismos. No obstante son incapaces no solo de identificarlas sino que, debido a las formas caprichosas que presentan o a su abundancia en un huésped determinado, son a menudo confundidas con estructuras propias de los vegetales o consideradas como enfermedades. Otras muchas agallas pasan desapercibidas por su escasa temporalidad o por el pequeño tamaño que presentan. Unas pocas, son bastante conocidas para el público en general, ya que pueden afectar a cultivos y aparecen con frecuencia en parques y jardines. Las agallas vegetales han llamado la atención de observadores y estudiosos del medio natural desde tiempos antiguos. De hecho, ya desde la época grecorromana existen escritos que documentan su utilización para diversos fines: desde usos médicos, a usos industriales (curtientes de pieles y en la fabricación de tintas), pasando por usos lúdicos o decorativos e incluso como alimento. Numerosas y variadas formaciones vegetales pueden incluirse dentro del concepto de agallas o cecidias. Alrededor de 15.000 especies de organismos (virus, bacterias, algas, hongos, protozoos, rotíferos, nematodos, ácaros e insectos) pueden inducir su formación. Las agallas producidas por artrópodos (ácaros e insectos) se denominan zoocecidias y la mayoría de ellas son producidas por especies de insectos. Las zoocecidias, a diferencia de otros modelos de agallas, presentan una forma y un tamaño relativamente constante por lo que con solo la visualización de dicha estructura podemos conocer cuál es la especie causante de la malformación vegetal. 2 Entre los artrópodos existen alrededor de 14.000 especies gallícolas (más de 13.000 producidas por insectos), en su mayoría asociadas a plantas angiospermas, capaces de inducir su formación y que se pueden dividir en dos grandes grupos; uno en el que la formación de la agalla está relacionada con su actividad alimenticia (artrópodos picadores: Tisanópteros, Hemípteros y ácaros) y otro, para los cuales la agalla es una consecuencia directa de la puesta y el desarrollo de las larvas (Coleópteros, Lepidópteros, Dípteros e Himenópteros). Un rasgo característico de los insectos gallícolas es su especificidad frente al género o especie de planta hospedante y la parte atacada de la planta, de modo que una determinada especie gallícola está asociada únicamente con una especie o grupo relacionado de especies botánicas e induce sus agallas de manera constante y exclusiva sobre un único órgano de la planta. Cualquier órgano y parte de la planta es susceptible de la acción del artrópodo incluyendo raíces, tallos, yemas, hojas (predominantemente), flores y frutos. Múltiples y diversos estudios pueden realizarse con estas estructuras deformantes, las cuales además constituyen un ecosistema en sí mismos en los que diversos artrópodos y vertebrados pueden interaccionar en ellas. Por otro lado, algunas de ellas han sido usadas o se usan aún como ornamentos, confección de tintes, juegos, o constituyen objetos culinarios, por mencionar algunos usos. Las agallas son en definitiva estructuras coevolutivas entre organismos animales y vegetales que además presentan estrategias disuasorias para impedir el ataque de los parasitoides, al mismo tiempo que éstos presentan contraestrategias para posibilitar la puesta en su interior. Qué son las agallas Las agallas p f mp m m ‘ v ’ p p presencia, bajo la acción nutricia o de puesta, de virus, bacterias, algas, hongos, protozoos, rotíferos, nematodos, ácaros e insectos (Pujade-Villar, 1986). No obstante, una definición más estricta es considerar a las agallas como estructuras anormales de partes de tejidos u órganos de plantas que se desarrollan por la acción específica a la presencia o actividad de un organismo inductor (Meyer, 1987; Shorthouse y Rohfritsch, 1992). En las agallas coexisten dos fenómenos que las diferencian de otras anormalidades que pueden detectarse en los vegetales: la hipertrofia (crecimiento anormal de células) y la hiperplasia (multiplicación anormal de células); a menudo, en las agallas más complejas puede acontecer un tercer fenómeno, la neogénesis (aparición de tejidos no específicos en el vegetal sano como el tejido nutricio); la nutrición es proporcionada exclusivamente por la capa de tejido más interna que rodea la larva, que representa generalmente una parte muy pequeña del volumen total de la agalla. Por lo mencionado, la acción de los insectos minadores de hojas, de los barrenadores de madera o de los enrolladores de hojas, por ejemplo, no pueden ser considerados como agallas a pesar de ser o parecer estructuras deformantes vegetales. Las agallas pueden corresponder a invaginaciones más o menos complejas de zonas del haz o del envés foliar, simples torsiones o enrollamientos foliares, bolsones englobadores ya sean abiertos o cerrados, hinchazones más o menos aparentes y estructuras altamente complejas provistas de distintas capas de tejidos organizados. En los áfidos (N. Pérez, per. com.) se utiliza el término pseudoagalla para aquellas deformaciones que constituyen simples pliegues de la hoja y que no se rompen para dejar salir a los alados, simplemente se despliegan (como en Eriosoma ulmi o Forda marginata) en contraposición al termino de agalla para aquellas deformaciones que estás cerradas (o casi 3 perfectamente cerradas) y que rompen su pared para dejar salir a las formas emigrantes (Geoica utricularia, Eriosoma lanuginosum, Pemphigus populi, por ejemplo); no obstante, en algunos casos, esta distinción no es clara ya que hay especies como Pemphigus bursarius, P. immunis, P. spyrothecae, entre otras, que están a medio camino. Dónde buscarlas Cualquier especie vegetal es susceptible de tener agallas, no obstante, la mayor parte de ellas se encuentran en angiospermas y más concretamente en las dicotiledóneas. Cualquier parte vegetal es también susceptible de tener agallas (hojas, flores, yemas, tallo, raíces y frutos) pero las hojas tienen preferencia en el computo global (Mani, 1964) si exceptuamos a los lepidópteros que prefieren los tallos y las yemas. Es importante por tanto observar atentamente todas las partes del huésped puesto que, salvo que las agallas sean perennes y de gran tamaño, nos podrían pasar desapercibidas. Por otro lado, hay agallas caducas por lo que el período de observación se reduce enormemente. También hemos de considerar que cada especie ataca, en la mayor parte de los casos, un órgano concreto y un tipo de huésped específico. Por todo ello el conocimiento del huésped se hace imprescindible. Quién las produce Los grupos implicados en la formación de las agallas son múltiples y variados tanto de origen animal (zoocecidias) como de otros orígenes (fitocecidias) contabilizándose más de 15.000 especies. Las fitocecidias son producidas por virus, bacterias y hongos, aunque algunos autores incluyen también en este grupo a algunas especies parásitas de fanerógamas (Ozenda y Capdebon, 1979). Estas agallas suelen ser tumores vegetales sin un tamaño constante y sin un aspecto bien determinado. Las zoocecidias son producidas por animales rotíferos, nematodos y artrópodos (ácaros e insectos). Se conocen unas 13.000 especies gallícolas de insectos y unas 500 especies de ácaros. Dichas agallas presentan un tamaño y un aspecto relativamente constante y definido. Según se menciona en Nieves-Aldrey (1998), a nivel mundial Felt (1940) puso de manifiesto las diferencias biogeográficas en la composición de la fauna gallícola. Así, en la región Holártica predominan las agallas en Fagales, Salicales, Asterales y Fabales producidas por cecidómidos, tentredínidos
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