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Morfología Y Anatomía Del Cefalio De Cephalocereus Senilis (Cactaceae)

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Morfología Y Anatomía Del Cefalio De Cephalocereus Senilis (Cactaceae) Anales del Jardín Botánico de Madrid 62(2): 153-161 www.rjb.csic.es Morfología y anatomía del cefalio de Cephalocereus senilis (Cactaceae) por Monserrat Vázquez-Sánchez1, Teresa Terrazas1 & Salvador Arias2 1 Programa de Botánica, Colegio de Postgraduados, Montecillo, Estado de México, 56230, México. [email protected] (M. Vázquez-Sánchez), [email protected] (T. Terrazas) 2 Jardín Botánico, Instituto de Biología, Universidad Nacional Autónoma de México. 70-614 México D.F. 04510, México. [email protected] Resumen Abstract Se estudió la morfología y anatomía de la zona florífera de Ce- We study the morphology and anatomy of Cephalocereus seni- phalocereus senilis (Haw.) Pfeiff. con la finalidad de caracterizar- lis (Haw.) Pfeiff. flowering zone to describe and compare it with la y compararla con la información disponible para otras espe- the information available for other species. We described in de- cies. Se describen con detalle las ramas vegetativas y reproducti- tail the vegetative and reproductive branches. Results show that vas. Los resultados mostraron que existen cambios en la morfo- there are morphological changes linked to anatomical ones af- logía relacionados con cambios en la anatomía después de la ter the flowering zone is differentiated. The flowering zone in formación de la zona florífera. La reducción de los espacios inte- this species is termed lateral cephalium by its interareolar space rareolares, el incremento en el tamaño de las aréolas y una den- reduction, areole size increment, and high production of long sa producción de cerdas y tricomas largos en las aréolas de la bristles and trichomes in the areoles. Moreover, periderm deve- zona florífera permiten reconocerla como un cefalio lateral. Tres lopment near the apical meristem, lack of chlorenchyma and a cambios anatómicos se asocian a la presencia del cefalio lateral: delay in xylematic fibers differentiation are traits related with the el desarrollo de la peridermis cercana al meristemo apical, la au- lateral cephalium. The lateral cephalium of C. senilis shows mo- sencia de clorénquima y el retardo en la formación de fibras en difications previously reported for the Espostoa species with la- el xilema secundario. El cefalio lateral de C. senilis presenta las teral cephalium, except for the presence of crystals in the tricho- modificaciones reportadas para las especies del género Espos- mes, the addition of new ribs to the cephalium, and the growth toa, también con cefalio lateral, excepto por la presencia de cris- in xylem and phloem parenchyma. tales en los tricomas, la incorporación de nuevas costillas al ce- falio y la dilatación del parénquima xilemático y floemático. Palabras clave: cactáceas, cefalio lateral, Cephalocereus, clo- Keywords: cacti, Cephalocereus, chlorenchyma, columnar ha- rénquima, hábito columnar, Hidalgo, paedomorfosis, perider- bit, flowering zone, Hidalgo, lateral cephalium, paedomorpho- mis, pseudocefalio, viejito, zona florífera. sis, periderm, pseudocephalium, viejito. Introducción de zona florífera. Las zonas floríferas han sido deno- minadas indistintamente cefalios o pseudocefalios La mayoría de las cactáceas no muestran diferen- por diferentes autores (Buxbaum, 1964a; Rauh, 1979; cias morfológicas entre las ramas vegetativas o juveni- Barthlott & Hunt, 1993; Anderson, 2001), a pesar de les y aquellas maduras que portan las estructuras re- que Buxbaum (1964a,b) define a un cefalio como productivas (Anderson, 2001). Sin embargo, en algu- aquella zona donde las aréolas de la zona florífera se nas especies de las tribus Cereeae, Pachycereeae y Tri- modifican y no continúan con su función vegetativa, y chocereeae, se presentan cambios en la morfología a un pseudocefalio, cuando las aréolas de la región flo- cuando la planta inicia su reproducción al llegar a la rífera continúan con su función vegetativa después de fase adulta (Terrazas & Arias, 1999). El área de las ra- la floración. Además las han llamado cefalio o pseu- mas o tallos asociados a la reproducción en el que se docefalio anular y semi-periférico (Bravo-Hollis, producen los cambios morfológicos recibe el nombre 1978; Rauh, 1979; Taylor & Zappi, 1989; Barthlott & 154 Anales del Jardín Botánico de Madrid 62(2) 2005 Hunt, 1993; Terrazas & Loza-Cornejo, 2002). Las zo- Biosfera de Metztitlán (Hidalgo, México) y están de- nas floríferas pueden desarrollarse en posición lateral positados en CHAPA (T. Terrazas et al. 640, 640A). o apical. En la zona florífera apical, el crecimiento de La morfología de la rama vegetativa se describió sin todas las costillas es interrumpido periódica o perma- hacer ninguna disección (Fig. 1a), mientras que en la nentemente (Barthlott & Hunt, 1993), mientras que la reproductiva se eliminaron de las aréolas cerdas y tri- posición lateral está confinada a una o varias costillas comas (Fig. 1b). En ambas se midieron el diámetro, debajo del ápice y también puede ser permanente o número de costillas, altura y ancho de la costilla, dis- periódica. La presencia de zonas floríferas ha sido tancia entre aréolas, largo y ancho de las aréolas, nú- considerada una característica derivada (Taylor & mero de espinas radiales y centrales, y se documentó Zappi, 1989) que fue adquirida independientemente su forma y color. Cada rama se cortó desde el ápice en las tres tribus (Terrazas & Arias, 2002). Terrazas & hasta la base en segmentos de 10 cm de grosor (Fig. Loza-Cornejo (2002) señalaron la necesidad de reali- 1a,b). En las ramas adultas se tomaron segmentos de zar estudios para verificar si el pseudocefalio es la si- la zona florífera (Fig. 1 b1) y del lado opuesto, que co- tuación intermedia entre la total ausencia de especia- rresponde a la región no florífera o vegetativa (Fig. 1 lización y una muy acusada. Uno de los miembros de b2). Estos segmentos se dividieron en pequeñas sec- Pachycereeae con zona florífera es Cephalocereus se- ciones que incluían desde las aréolas hasta la médula. nilis (Haw.) Pfeiff., especie endémica de la zona árida Los segmentos se fijaron en formalín-ácido acético- de los estados de Hidalgo y Veracruz, en México. La alcohol e incluyeron en parafina para su cortado (Jo- zona florífera de esta especie ha sido llamada indis- hansen, 1940). tintamente cefalio y pseudocefalio (Bravo-Hollis, 1978; Gibson & Nobel, 1986; Anderson, 2001), aun- Resultados que no existe ningún trabajo estructural que describa Morfología los cambios asociados a su formación. En este trabajo abordamos el estudio morfológico y anatómico de la Cephalocereus senilis es una planta de hábito co- zona florífera de C. senilis con el fin de caracterizarla lumnar, monopódica, de hasta 15 m de altura y 45-50 y compararla con las ramas juveniles. cm de diámetro en su parte más ancha, con ramas es- casas que se consideran basítonas porque se originan Materiales y métodos de las aréolas de la base (Fig. 2a); cuando ocasional- mente hay ramas en la parte superior, éstas se originan La morfología de C. senilis se describe a partir de la como respuesta al daño del meristemo apical. Ramas observación de ramas juveniles en estado vegetativo y verdes, con 32-34 costillas poco prominentes, de ramas maduras en estado reproductivo. Los ejempla- 9-12 mm de altura; aréolas de 6,5-7,0 mm de largo y res estudiados fueron recolectados en la Reserva de la 5,0-5,4 mm de ancho, de circulares a elípticas, separa- das entre sí 12-14 mm (Fig. 2d); no hay espinas radia- les ni centrales distinguibles, pero en cada aréola se a b observan 3-5 espinas y numerosas cerdas largas y fle- xibles de color blanco; cerdas de 3-5 cm de longitud en las aréolas jóvenes, pero de más de 40 cm de longi- tud en las aréolas más viejas en la base de la rama. Las b1 ramas juveniles están cubiertas con abundantes cer- das de color blanco (Fig. 2c). Estas cerdas son muy notorias en los individuos o ramas jóvenes (Fig. 2a-c); b2 por ello, esta especie recibe el nombre vernáculo de “viejito”. Cuando la planta alcanza la madurez se inicia el de- sarrollo de una zona florífera lateral diferenciada (Fig. 2b). Esta zona florífera, que crece cada año por incor- poración de nuevas aréolas a la reproducción, cubre menos de la mitad del tallo en longitud. En sus prime- Fig. 1. Ramas de Cephalocereus senilis donde se indica con lí- ros estadios sólo dos o tres costillas se diferencian, neas verticales y horizontales los segmentos para el estudio ana- tómico. a, rama juvenil; b, rama reproductiva con cefalio (b1) al pero en las fases más avanzadas ocupa cerca de la que se han eliminado tricomas y cerdas, y zona vegetativa (b2) mitad del diámetro del tallo o de la rama (Figs. 2a,b, opuesta al cefalio. 3b). Las costillas modificadas tienen aréolas de M. Vázquez-Sánchez & al.: Cefalio de Cephalocereus senilis 155 8,7-12,5 mm de largo y 6,9-10,3 mm de ancho, espa- uno o más cristales prismáticos en todas las células ciadas entre sí 6-8 mm, y conservan su disposición li- epidérmicas (Fig. 4a). Los estomas son paracíticos neal, pero con los podarios prominentes (Fig. 3b-d). con tres pares de células subsidiarias y las células oclu- Además, las costillas se cubren por una densa capa de sivas se localizan al mismo nivel que el resto de las cé- tricomas de color beige y hasta 8 cm de longitud. En lulas epidérmicas. En la zona florífera son más abun- las aréolas también hay algunas cerdas largas (ca 8 dantes los estomas en las depresiones de las costillas. cm), muy flexibles y similares en color a los tricomas La epidermis de la base de la aréola da origen exclusi- (Fig. 3a). Tricomas y cerdas cubren casi completa- vamente a tricomas largos con cristales y cerdas igual- mente a las flores hasta que los botones alcanzan su ta- mente largas en la zona florífera (Fig.
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