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Biodiversidad De Pinophyta (Coníferas) En México

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Biodiversidad De Pinophyta (Coníferas) En México Revista Mexicana de Biodiversidad, Supl. 85: S126-S133, 2014 126 Gernandt y Pérez-de la Rosa.- BiodiversidadDOI: 10.7550/rmb.32195 de coníferas Biodiversidad de Pinophyta (coníferas) en México Biodiversity of Pinophyta (conifers) in Mexico David S. Gernandt1 y Jorge A. Pérez-de la Rosa2 1Departamento de Botánica, Instituto de Biología, Universidad Nacional Autónoma de México. Apartado postal 70-367, 04510 México, D. F., México. 2Departamento de Botánica y Zoología, Instituto de Botánica, Universidad de Guadalajara. Apartado postal 139, 45110 Zapopan, Jalisco, México. [email protected] Resumen. Las coníferas (Pinophyta) son árboles o arbustos con hojas simples y estructuras fértiles arregladas en conos polínicos simples y conos ovulados compuestos, excepto en Taxaceae. Las coníferas son los componentes dominantes de diversos tipos de vegetación. En México crecen desde el nivel del mar hasta por encima de los 4 000 m; la mayor diversidad se encuentra en los bosques montañosos de la Sierra Madre Occidental y Sierra Madre Oriental. Están representadas por 4 familias: Pinaceae (4 géneros y 61 especies), Cupressaceae (4 géneros y 29 especies), Podocarpaceae (1 género y 3 especies) y Taxaceae (1 especie). De las 94 especies de coníferas mexicanas, 43 son endémicas del país, de éstas 18 tienen un rango de distribución limitado a 3 o menos estados. Palabras clave: coníferas, conos, especies, distribución. Abstract. Conifers (Pinophyta) are woody trees or shrubs with simple leaves and reproductive structures arranged in simple pollen cones and (except in Taxaceae) compound seed cones. They are dominant components of several vegetation types. In Mexico conifers grow from sea level to above 4 000 m, with the greatest diversity concentrated in montane forests of the Sierra Madre Occidental and Sierra Madre Oriental. Four families have natural distributions in the country: Pinaceae (4 genera and 61 species), Cupressaceae (4 genera and 29 species), Podocarpaceae (1 genus and 3 species), and Taxaceae (1 species). Of the 94 Mexican conifer species, 43 are endemic to the country, 18 with ranges restricted to 3 or fewer states. Key words: conifers, cones, species, distribution. Introducción y de 2 a muchos microesporangios ubicados distalmente en la cara abaxial. Los granos de polen en algunas Las coníferas (Pinophyta o Coniferophyta) son árboles especies presentan sacos aeríferos. Los conos ovulados o arbustos dioicos o monoicos con la corteza rugosa o lisa, están formados por un complejo bráctea-escama que se en placas grandes y gruesas con fisuras o en tiras largas arregla helicoidalmente alrededor de un eje central; sin y delgadas (Figs. 1-4). Las ramas laterales están bien embargo, en Podocarpaceae y Taxaceae están típicamente desarrolladas. Las hojas son simples y pueden ser en forma reducidos a 1 o 2 escamas con un solo óvulo. Estos conos de aguja, escama, lineares, lanceoladas, a veces oblongas o son típicamente leñosos, pero en Juniperus son fibrosos o falcadas. Por lo general las hojas son persistentes por más carnosos, y en Podocarpaceae y en Taxaceae, los óvulos por de un año, pero a veces son deciduas. Su madera posee un lo general están cubiertos por estructuras estériles, carnosas xilema compacto compuesto principalmente de traqueidas y de colores comúnmente llamativos (presuntamente una con paredes gruesas y poros uniseriados o multiseriados. adaptación para la zoocoria) denominadas epimacios y En contraste con angiospermas, el xilema carece de vasos. arilos, respectivamente; los conos se desarrollan durante Frecuentemente presentan canales resiníferos en su madera, un año en la mayoría de especies, aunque en Pinus y en la corteza, las hojas o los conos. Los estróbilos o conos, algunas especies de Juniperus tardan 2 o hasta 3 años. El son monoesporangiados. Los conos microesporangiados complejo de bráctea-escama puede ser libre o fusionado o polínicos son simples, formados de microesporófilas y presentar 1 o muchos óvulos en su cara adaxial, que arregladas en forma de hélice alrededor de un eje central estarán expuestos al momento de la polinización; este es un carácter propio de todas las gimnospermas (las coníferas, las cícadas, Ginkgo y las gnetofitas). El micrópilo en Recibido: 08 abril 2011; aceptado: 25 marzo 2013 algunos grupos está orientado hacia la parte distal y en Revista Mexicana de Biodiversidad, Supl. 85: S126-S133, 2014 DOI: 10.7550/rmb.32195 127 Figuras 1-4. Ejemplos de coníferas mexicanas. 1, Juniperus deppeana. 2, Taxus globosa. 3, Pinus pinceana. 4, Podocarpus matudae. otros está dirigido hacia el eje (los óvulos están invertidos). las gnetofitas comparten varios caracteres de su madera Las semillas de la mayoría de las especies en Cupressaceae y los 2 grupos tienen hojas y conos polínicos simples; y Pinaceae presentan un ala que les facilita la dispersión sin embargo, esta relación estrecha entre los 2 grupos por el viento, o por el agua en Taxodium; la semilla carece de gimnospermas está en conflicto con los resultados de un ala en Juniperus, Podocarpaceae y Taxaceae, y el filogenéticos basados en caracteres morfológicos, los ala es vestigial en los pinos piñoneros, en donde aves u cuales encuentran a las gnetofitas como el grupo hermano otros animales llevan a cabo la dispersión. El número de a las angiospermas (la hipótesis antofita); estos 2 grupos cotiledones es de 2 a 15, excepcionalmente 25 en Pinus comparten los vasos vasculares en la madera y la doble maximartinezii. fertilización. Filogenias moleculares con más caracteres Las coníferas vivientes han sido consideradas como han ubicado a las coníferas como un grupo monofilético un grupo natural por mucho tiempo, aunque Taxaceae y a las gnetofitas cómo un clado independiente (Rai et al., fue clasificado por algunos autores en un orden separado 2008; Lee et al., 2011). (Taxales), sobre todo porque carecen de conos ovulados Entre las coníferas se incluyen algunas de las plantas reconocibles (Florin, 1954). Los análisis filogenéticos más grandes y longevas; el árbol más alto es Sequoia ubican a Taxaceae con las demás coníferas. Adicionalmente, sempervirens Endl. (hasta 115 m), el más masivo es los análisis filogenéticos de secuencias de DNA han Sequoiadendron giganteum (Lindl.) J. Buchholz (>1 establecido una relación estrecha entre las coníferas y 400 m3) y el de mayor edad es Pinus longaeva D. K. las gnetofitas, incluso se han encontrado filogenias en Bailey (de c. 4 700 años). En México, el árbol nacional, las cuales las gnetofitas se derivan de las coníferas, por el “ahuehuete” (Taxodium mucronatum), llega a tener un ejemplo, como un grupo hermano a Pinaceae (la hipótesis tronco de más de 12 m de diámetro (el árbol de Tule gnepine; revisada por Mathews, 2009). Las coníferas y en Oaxaca). Existen además, especies de crecimiento 128 Gernandt y Pérez-de la Rosa.- Biodiversidad de coníferas arbustivo como por ejemplo Juniperus monticola o Pinus Pinaceae, Podocarpaceae y Taxaceae (Figs. 1-4; Cuadro 1). culminicola, que a veces miden menos de 1 m de altura Las 2 familias restantes son la monotípica Sciadopityaceae y árboles que alcanzan hasta 70 m, como Abies religiosa (Sciadopitys verticillata (Thunb.) Siebold et Zucc.) de y Pinus ayacahuite (Farjon y Styles, 1997; Farjon, 2010; Japón y la familia Araucariaceae representada por 3 Ricker y Hernández, 2010). géneros distribuidos en el hemisferio sur. Sólo 10 de los En México las coníferas están presentes en diversos 71 géneros de coníferas están presentes en el país. Hay tipos de vegetación. De acuerdo con Rzedowski (1978) se poco acuerdo en el número de especies de coníferas, por encuentran en el bosque de pino, matorral de pino, bosque ejemplo Eckenwalder (2009) reconoce 546 especies en el de Abies, bosque de Pseudotsuga y Picea, bosque o matorral mundo, mientras que Farjon (2010) reconoce 615. Estas de Juniperus y bosque de Cupressus (= Callitropsis). aportaciones se basan en un concepto amplio de especie; Además, habitan en otros tipos de vegetación como el sin embargo, otras obras al nivel de género adoptaron bosque mesófilo de montaña, bosque mixto (por ejemplo, conceptos más estrechos, citando un mayor número de bosque de pino-encino), matorral xerófilo y bosque de especies, en particular para los géneros más diversos galería (el hábitat de Taxodium mucronatum). como Abies (Liu, 1971), Pinus (Martínez, 1948; Perry, La fase del esporofito domina en las coníferas y esta 1991; Styles, 1993), Callitropsis (Little, 2006) y Juniperus etapa alcanza a durar hasta cientos o miles de años. El ciclo (Adams, 2011). Estimamos que hay 670 especies de reproductivo, el cual comprende desde la iniciación de las coníferas a nivel mundial, de las cuales 94 (14%) tienen yemas hasta la liberación de semillas, dura alrededor de una distribución natural en el país (Cuadro 2). Estas cifras 17 meses en Pseudotsuga menziesii (Allen y Owens, 1972) son mayores que las 77 especies mexicanas reconocidas y de 2 a 3 años en otras especies (Tomlinson y Takaso, por Eckenwalder (2009) o las 74 especies reconocidas 2002). Los primordios de las yemas vegetativas inician por Farjon (2010), pero son menores que las 104 especies un aumento en su actividad, comúnmente a inicios de la reconocidas por Debreczy y Rácz (2011). Algunos clados primavera, con una diferenciación en los conos polínicos de coníferas probablemente sufrieron diversificación y ovulados evidente a finales de la primavera o a inicios reciente, seguidos por migración, contacto secundario del verano. La esporogénesis mediante meiosis dura de e hibridación.
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