Stomata in Podocarpus

The stomatal complex of Podocarpus observed in cross-section using cryofracture: a preliminary study

by Whiting M., Mill R. R., Jeffree C. E. (2017)

In Edinburgh Journal of Botany 74(3): 345-364 –


Cryofracture of living material and fracture at room temperature of herbarium material were used to obtain cross-sections of the stomatal complexes of four species of Podocarpus (Podocarpaceae) for scanning electron microscopy. Cross-sections of the stomata of one species in Podocarpus subgenus Foliolatus, section Foliolatus (Podocarpus rubens),

one in Podocarpus section Globulus (Podocarpus beecherae), one in Podocarpus subgenus Foliolatus section Longifoliolatus (Podocarpus insularis) and one in subgenus Podocarpus section Australis (Podocarpus nivalis) were studied. The architecture of the stomatal complex, including the wax plug, is described.

It was found that the wax plug sits high in the stomatal antechamber in Podocarpus rubensP. beecherae and Podocarpus decipiens and about halfway up the chamber in P. nivalis. A ridge, which appears to correspond to the crease where the guard cells meet, exists on the underside of the wax plug in Podocarpus beecheraeP. decipiens and P. rubens; its presence in P. nivalis requires confirmation. In addition, ridges within the stomatal antechamber were observed when viewing the cross-sections of Podocarpus decipiens and P. rubens, the internal surface of the cuticle of P. decipiensPodocarpus teysmanniiP. insularis and Podocarpus milanjianus, and the external surface of the cuticle of Podocarpus chinensisPodocarpus macrophyllus and Podocarpus pilgeri. These ridges may consist of wax and be a result of epitaxis.


Stomata in fossil conifers

Zur Unterscheidung fossiler Spaltöffnungen den mitteleuropaischen Coniferen (In German)

by Trautmann W. (1953)

Werner Trautmann, Universität Göttingen (Germany)


In Flora 140: 523-533 – –


Stomata: Pinus, Picea, Taxus, Abies, Larix, Juniperus

Sunken stomata in Pinus (Gymnopsermae)

Photo Google Wikimedia Commons
Cross section: Pinus needle magnification: 400x
A single layered epidermis of lignified cells is covered by a heavy waxy cuticle. Numerous sunken stomata are scattered over the entire epidermal surface. They are bounded by pairs of subsidiary cells, and open internally to a sub-stomatal cavity and externally to a respiratory cavity or vestibule.
The underlying hypodermis consists of several layers of thick-walled sclerenchyma, particularly well-developed at ridges.

10 February 2014, Berkshire Community College Bioscience Image Library

3,264 × 1,840 (5.58 MB)

Stomatal characteristics in Glyptostrobus (Taxodiaceae)

SEM images of stomata. Scale bars = 50 µm. Fig. 11. Stomata of outer surface of a linear leaf collected from Hangzhou. Figs 12, 13. Stomata of inner surface of linear leaves collected from Hangzhou.



Screen Shot 2018-06-04 at 23.47.51
Stomata from linear-subulate leaves collected from Hangzhou. Arrows indicate pits. Scale bars = 30 µm.


Epidermal structures and stomatal parameters of Chinese endemic Glyptostrobus pensilis (Taxodiaceae)

by Ma Q.-W., Li C.-S., Li F.-L., Vickulin S. V. (2004)

Qing-Wen MA, Cheng-Sen LI, Feng-Lan LI, SergeiI V. VICKULIN,


in Botanical Journal of the Linnean Society 146(2): 153–162 –


Glyptostrobus pensilis K. Koch, the only living species, is endemic to southern China. Epidermal structures of Gpensilis have been studied on leaves collected from Guangzhou, southern China, the native locality of the species, and from Hangzhou, eastern China, the cultivated locality.Leaves are linear, linear-subulate and scale-like. Epidermal cells are rectangular and elongate parallel to the mid-vein on areas lacking stomata, and short, with rounded corners, on intrastomatal areas.

Stomatal bands lie parallel to the mid-vein on both surfaces of leaves. Commonly the stomata have five or six subsidiary cells. Stomatal parameters (density and index) of the same surfaces of linear leaves from Guangzhou and Hangzhou show no statistically significant differences (P > 0.05).

Considering the stomatal parameters of the same surfaces of linear-subulate leaves between the two localities, the stomatal index of the abaxial surfaces reveals no significant differences (P > 0.05), while the stomatal index of the adaxial surfaces and the stomatal density of both surfaces exhibit significant differences (P < 0.05). Intra-individual variation in stomatal index is smaller than that in stomatal density based on the coefficient of variability of stomatal parameters of the same areas of leaves.

When studying the correlation between stomatal parameters of G. pensilis and atmospheric CO2 concentrations, the stomatal parameters of linear leaves are mostly significant, and stomatal index is more useful than stomatal density.

Leaf anatomy and stomatal structures of six genera of Taxaceae



Leaf anatomy and its implications for phylogenetic relationships in Taxaceae s. l.

by Ghimire B., Lee C., Heo K. (2014)

Department of Applied Plant Science and Oriental Bio-herb Research Institute, Kangwon National University, Chuncheon, 200-701, Korea.


in J Plant Res. 127(3):373-388 – doi: 10.1007/s10265-014-0625-3 – Epub 2014 Feb 5 –


The comparative study on leaf anatomy and stomata structures of six genera of Taxaceae s. l. was conducted. Leaf anatomical structures were very comparable to each other in tissue shape and their arrangements. Taxus, Austrotaxus, and Pseudotaxus have no foliar resin canal, whereas Amentotaxus, Cephalotaxus, and Torreya have a single resin canal located below the vascular bundle. Among them, Torreya was unique with thick-walled, almost round sclerenchymatous epidermal cells. In addition, Amentotaxus and Torreya were comprised of some fiber cells around the vascular bundle. Also, Amentotaxus resembled Cephalotaxus harringtonia and its var. nana because they have discontinuous fibrous hypodermis. However, C. fortunei lacked the same kind of cells.

Stomata were arranged in two stomatal bands separated by a mid-vein. The most unique stomatal structure was of Taxus with papillose accessory cells forming stomatal apparatus and of Torreya with deeply seated stomata covered with a special filament structure. Some morphological and molecular studies have already been discussed for the alternative classification of taxad genera into different minor families.

The present study is also similar to these hypotheses because each genus has their own individuality in anatomical structure and stomata morphology.

In conclusion, these differences in leaf and stomata morphology neither strongly support the two tribes in Taxaceae nor fairly recognize the monogeneric family, Cephalotaxaceae. Rather, it might support an alternative classification of taxad genera in different minor families or a single family Taxaceae including Cephalotaxus.

In this study, we would prefer the latter one because there is no clear reason to separate Cephalotaxus from the rest genera of Taxaceae. Therefore, Taxaceae should be redefined with broad circumscriptions including Cephalotaxus.

Stomata in different Ephedra species



[SEM observation on leaf epidermis of different Ephedra species] (in Chinese)

by Wu J. L., Niu J. Y., Yan Z. Z., Li S., Gao Y. H., Jiang H. Y. (2007)

College of Life Sciences & Technology, Gansu Agricultural University, Lanzhou 730070, China.


in Zhongguo Zhong Yao Za Zhi. 32(18):1854-1857 – PMID: 18051888



Characters of stem epidermis, leaf epidermis and stoma could be used as important microcosmic morphological characteristic when inheritance trend is studied in Ephedra breeding and identification.


The stomatic density, stoma major axis and mimor axis, stomatic morphylogy, characters of leaf and stem epidermis of 6 Ephedra plants’ stems were examined by SEM.


The stomatic density and characteristic of leaf epidermis and stem epidermis in six Ephedra species was differenc, there were no obvious morphological differences in stoma shape and size. The guard cells were covered with heavy cuticle and sunken stomata, which were the typical characteristics of xerophytes. The stomas of leaf lower epidermis were oblong or hexagon, but the stomas of steam epidermis were narrowed-oblong or dumbbell-shape, they all belonged to anomalous type.


The stoma type and characters of Ephedra plants is stable and conservative, there was no obvious morphological differences in stoma shape and size between species, so it is difficult to distinguish different species by the variance of stomas, but that can be applyed to distinguish Ephedra from others at plant taxonomy.

Stomatal distribution pattern in Podocarpus (Gymnospermae)

Micrographs of the abaxial epidermis of P. lambertii. (A) A view of stomatal distribution in longitudinal rows. Between the rows of stomata there are always sclereids beneath the epidermis, indicating that where there are sclereids there are no stomata (scale bar = 100 µm). (B) A view of the epidermis without sclereids (scale bar = 100 µm). (C) A detail of paratetracytic stomata with four subsidiary cells (scale bar = 50 µm).


Plasticity of stomatal distribution pattern and stem tracheid dimensions in Podocarpus lambertii: an ecological study

by Locosselli G. M., Ceccantini G. (2012)

Universidade de São Paulo, Instituto de Biociências, Departamento de Botânica, Laboratório de Anatomia Vegetal, Rua do Matão, 277, 05508-090, Sao Paulo, SP, Brazil


in Ann Bot 110(5): 1057–1066 – doi:  10.1093/aob/mcs179 – PMC3448432 –


Background and Aims

Leaf and wood plasticity are key elements in the survival of widely distributed plant species. Little is known, however, about variation in stomatal distribution in the leaf epidermis and its correlation with the dimensions of conducting cells in wood. This study aimed at testing the hypothesis that Podocarpus lambertii, a conifer tree, possesses a well-defined pattern of stomatal distribution, and that this pattern can vary together with the dimensions of stem tracheids as a possible strategy to survive in climatically different sites.


Leaves and wood were sampled from trees growing in a cold, wet site in south-eastern Brazil and in a warm, dry site in north-eastern Brazil. Stomata were thoroughly mapped in leaves from each study site to determine a spatial sampling strategy. Stomatal density, stomatal index and guard cell length were then sampled in three regions of the leaf: near the midrib, near the leaf margin and in between the two. This sampling strategy was used to test for a pattern and its possible variation between study sites. Wood and stomata data were analysed together via principal component analysis.

Key Results

The following distribution pattern was found in the south-eastern leaves: the stomatal index was up to 25 % higher in the central leaf region, between the midrib and the leaf margin, than in the adjacent regions. The inverse pattern was found in the north-eastern leaves, in which the stomatal index was 10 % higher near the midrib and the leaf margin. This change in pattern was accompanied by smaller tracheid lumen diameter and length.


Podocarpus lambertii individuals in sites with higher temperature and lower water availability jointly regulate stomatal distribution in leaves and tracheid dimensions in wood. The observed stomatal distribution pattern and variation appear to be closely related to the placement of conducting tissue in the mesophyll.