Stomata in Sequoia sempervirens and their variation in different plantations in Chile

Características de los estomas, densidad e índice estomático en secuoya (Sequoia sempervirens) y su variación en diferentes plantaciones de Chile –

Stomata Characteristics, stomatal density and stomatic index in redwood (Sequoia sempervirens) and its variation in different plantations in Chile

Toral M., Manríquez A., Navarro-Cerrillo R., Tersi D., NaulinP. (2010)

Manuel Torala*, Adelina Manríquez a, Rafael Navarro-Cerrillo b, Denise Tersi a, Paulette Naulin a,

Universidad de Chile, Facultad de Ciencias Forestales y Conservación de la Naturaleza, casilla 9206, Santiago, Chile
Universidad de Córdoba, Departamento de Ingeniería Forestal, Córdoba, España.


Bosque 31(2): 157-164 –


Several authors suggest that the stomata of Sequoia sempervirens (redwood) may present morphological changes when this species grows in areas without the presence of crypto-precipitation. This work studied the characteristics of the stomata, density and stomata index in leaves of redwood in terms of precipitation level, soil, slope exposition and location in the forest stand. The anatomical study was made by diaphanization and staining techniques of the leaf samples. The main results of this work indicate that the sequoia stomata, in the populations studied, are smaller than those from natural populations, with an average length between 31.4 and 37.7 µm and an average width between 12.6 and 14.9 µm; mainly found in the abaxial epidermis. Their density varied between 81.3 and 111.4 stomata mm-2, with an average stomatic index between 15.5 and 21.1 depending on the locality studied. These values are in ranks lower than those found for the species by other authors; indicating a certain degree of acclimation for the species in Chile. While there was significant variation among localities in Chile, changes in the stomatal complex were not so evident in response to the site environmental characteristics. However, in areas with more restraining climatic conditions (lower atmospheric humidity and precipitation), combined with edaphic conditions (soil with greater apparent density or less depth), stomata showed smaller and less dense, revealing a significant degree of phenotypic plasticity of the species for the character studied, which allows adaptation processes to contrasting environmental situations.


Varios autores sugieren que los estomas de secuoya (Sequoia sempervirens) pueden presentar cambios morfológicos cuando esta especie crece en áreas con ausencia de criptoprecipitación. En este trabajo se estudiaron las características de los estomas, densidad e índice estomático en hojas de secuoya en función del nivel de precipitación, suelo, exposición y ubicación del árbol en el rodal. El estudio anatómico se realizó mediante técnicas de diafanización y teñido de las muestras de hojas. Los principales resultados indican que los estomas de secuoya en las poblaciones estudiadas son pequeños en comparación con las poblaciones naturales, con longitud media entre 31,4 y 37,7 µm y ancho medio entre 12,6 y 14,9 µm, y se presentan principalmente en la epidermis abaxial. Su densidad media varió entre 81,3 y 111,4 estomas mm-2, con un índice estomático medio entre 15,5 y 21,1 según la localidad estudiada. Estos valores están en los rangos inferiores a los encontrados para la especie por otros autores, lo que indica cierto grado de aclimatación para la especie en Chile. Si bien hubo variación significativa entre localidades de Chile, los cambios en el aparato estomático no fueron tan marcados en respuesta a las características ambientales de los sitios. Sin embargo, en localidades con condiciones climáticas más limitantes (menor humedad atmosférica y precipitación), en combinación con las condiciones edáficas (suelos con mayor densidad aparente o menor profundidad), mostraron estomas más pequeños y en menor densidad, revelando un importante grado de plasticidad fenotípica de la especie para el carácter evaluado, que permite procesos de aclimatación a situaciones ambientales contrastantes.

Stomata in Zamiaceae (Gymnospermae)

The leaf epidermis of three species in Zamiaceae

Xia S. J., Liao F., Huang Y. Y., Lin J. Z. (2003)

S.J.n Xia, Liao Fen, Huang Yu Yuan, Lin Jian Zhao,


Acta Bot. Yunn. 25(5): 596-602 – ISSN/ISBN: 0253-2700 –


The leaf epidermal characters of 3 species (Dioon holmgreii, Macrozamia moorei and Zamia furfuracea) in Zamiaceae were observed under the light microscope. To some extent, they proved that the division of three genera was natural. Stomatal cluster was discovered in Z. furfuracea.

Stomata in some gymnospermae species leaves

Comparative anatomical study of some gymnospermae species leaves

Bercu R., Broasca L., Popoviciu R. (2010)

Rodica Bercu, Livia Broasca, Razvan Popoviciu,

Department of Botany, Faculty of Natural and Agricultural Sciences “Ovidius” University, Constanta, Romania


Botanica Serbica 34: 21-28 –


Engrossed stomata, with large (Cedrus species) or
small (Abies and Juniperus species) substomatal cavities
occur. Th ey are numerous for the leaves of Abies and
Juniperus species and few for Cedrus species.

Amorphous epicuticular wax crusts disturbing the normal gas exchange

Epicuticular wax on stomata of damaged silver fir trees (Abies alba Mili.)

Bačić T., Krstin L., Roša J., Popović Z. (2005)

Tomislav Bačić, Ljiljana Krstin, Jadranka Roša, Željko Popović,


Acta Societatis Botanicorum Poloniae 74: 159-166 (eissn: 2083-9480) –

Wax tubules on the epistomatal rims of stomata

Epicuticular wax on stomata of damaged silver fir trees (Abies alba Mili.)

Bačić T., Krstin L., Roša J., Popović Z. (2005)

Tomislav Bačić, Ljiljana Krstin, Jadranka Roša, Željko Popović,


Acta Societatis Botanicorum Poloniae 74(2) : –


Condition of epistomatal wax on the abaxial surface of the current and previous-year needles of damaged silver fir trees (Abies alba Mill.), both from the polluted Risnjak and “clean” Donja Dobra sites in Gorski Kotar region, both influenced by pollutants coming from Europe, during two years, three times a year, were examined with Scanning Electron Microscope.

In the course of time the wax tubules on the epistomatal rims of stomata in polluted, but also in “clean” needles surface, become fused and agglomerated rapidly to various extents of morphologically different types of amorphous wax crusts, primarily compact and particulate ones. This process begins very early, especially in polluted Risnjak site, and may be interpreted as a possible result of air pollution.

However, the recrystalization, or production of new tubules, also appears relatively quickly in mostly cases. Quantitative estimations indicate a very large total amount of amorphous wax crusts in the current-year needles, and a very high percentage of the same wax in previous-year needles.

Amorphous wax crusts cover stomatal pores, as well as the rims, disturbing the normal gas exchange. Statistically there is a signicant tendency of increase in wax degradation in the needles of the polluted site in comparison with those of the unpolluted one, but there is an insignificant wax degradation among the needles of damaged trees within each site.

These results confirmed most of the research done in our preliminary report.

Stomata and SEM and LM foliar epidermal micro-morphology of gymnosperms

Taxonomic importance of SEM and LM foliar epidermal micro-morphology: A tool for robust identification of gymnosperms

Khan R., Abidin S. Z. U., Ahmad M., Zafar M., Liu J., Lubna, Jamshed S., Kiliç Ö. (2019)

Raees Khana-b, Sheikh Zain Ul Abidinc, Mushtaq Ahmadc, Muhammad Zafarc, Jie Liud, Lubnac, Shayan Jamshedc, Ömer Kiliçe,

a Department of Plant Sciences, Quaid-i-Azam University Islamabad, Pakistan

b Zoological Survey of Pakistan, Ministry of Climate Change, Pakistan

c Department of Plant Sciences, Quaid-i-Azam University Islamabad, Pakistan

d Kunming Institute of Botany, Chinese Academy of Sciences, Yunnan, PR China

e Department of Park and Garden Plant, Bingol University, Turkey


Flora 255: 42-68 – ISSN 0367-2530 –


• Variation in stomata, silica bodies (SiO2.nH2O), macrohairs, microhairs, hooks, papillae, prickles.

• Taxonomic applications of leaf epidermal SEM and Light morphology.

• Establishing key based on foliar epidermal micromorphological characters.


The current study reports the implication of scanning electron microscopy (SEM) and light microscopy (LM) epidermal micro-morphological characters in the identification of gymnosperms of Pakistan.

A number of characters were found to be of great taxonomic significance and can be used to distinguish different species of gymnosperms with its help. Here, the foliar cuticular features of 44 gymnosperm species were assessed, in which 32 were hypostomatic, 6 were epistomatic and 6 were amphistomatic. Four different types of guard cells were observed, 27 species having crescent shaped, 10 species having rectangular, 5 species having dome-shaped, and 2 species having triangular guard cells.

Sunken type stomata were very frequent in all gymnosperm species. 27 species have different types of phytoliths, while in the remaining 17 species no phytoliths were seen. In the current study, we have recognized seven types of Florin rings, named as Type A, Type B, Type C, Type D, Type E and Type F. Three types (Type I, Type II and Type III) of epicuticular wax were also observed in the investigated species.

The quantitative character showed diversity and was useful in the complete assessment and key making. A taxonomic key based on these micro-morphological characters was prepared. The study characters (stomatal index, stomatal density, epidermal cell density and stomatal area) were also analyzed by different statistical methods. SEM and LM foliar epidermal studies were found very useful in taxonomic identification. Based on current results we recommend such studies for accurate taxonomic identification and complete phylogeny.

The structure of the stomatal complex (the shape and arrangement of the subsidiary cells) was different in each Pinus species

Figure 2. Stomatal variables: Aa, stomatal width; La, stomatal length; Ab, upper woody lamellar width; Lb, upper woodylamella length; lc, distance between the external limits of the medial lamellae borders measured at the centre; ld, distancebetween the external limits of the medial lamellae borders measured at the point at which both meet to form the stem;e, medial lamellae border width; Lt, stem length; At, stem width; a, angle of attachment of the upper woody lamella; b,angle between the stem and medial lamellae border; coef_a = Aa/La, stomatal width ratio. Terminology based on that ofFlorin (1931), Trautmann (1953) and Hansen (1995) (Appendix).

The value of leaf cuticle characteristics in the identification and classification of Iberian Mediterranean members of the genus Pinus

Garcia Alvarez S., Garcia-Amorena I., Rubiales J. M., Morla C. (2009)


Unidad Docente de Botánica, Escuela Técnica Superior de Ingenieros de Montes, Universidad Politécnica de Madrid, Ciudad Universitaria s/n, 28040 Madrid, Spain


Botanical Journal of the Linnean Society 161: 436–448 –

Figure 3. Stomatal rows of Pinus nigra subsp. salzmannii.

This study reports the value of leaf cuticle characteristics in the identification and classification of Iberian Mediterranean species of the genus Pinus (P. nigra subsp. salzmannii, P. pinaster, P. pinea and P. halepensis), with the aim of using these characters to identify isolated cuticles and stomata in palynology slides.

Preparations were made of the cuticles of pine needles belonging to one natural Iberian population of each of the above species. A number of epidermal morphological characteristics were then recorded with the aim of distinguishing these species from one another.

The structure of the stomatal complex (the shape and arrangement of the subsidiary cells) was different in each species. The aperture of the epistomatal chamber was significantly smaller in P. pinea than in the other species examined, and the variables recorded for the thickening of the guard cells provided relationships that clearly distinguished all four taxa.

The width and length of the stomata and the upper woody lamellae, the central distance between the external limits of the medial lamellae borders and the length of the stem were the most useful variables in this respect.

The present results contribute to the ongoing discussion regarding the taxonomic classification of the members of Pinus, and provide valuable clues for the identification of Iberian Mediterranean pine species from small pine needle fragments or isolated stomata.

After validation of the present results for multiple populations, these results could also be used to help identify fossil leaf macroremains and the scattered/isolated stomata commonly observed in palaeopalynological samples.