Stomata in Marattiaceae (Filicopsida)

Fig. 9. Esquemas de los caracteres de las células oclusivas. A, en superficie; B, en corte transversal. Tomado de Simón (1991).

Análisis con MO y MEB de caracteres epidérmicos foliares en Marattiaceae Sensu Lato

Rolleri C.H., Lavalle M. C. (1997)

Cristina Rolleri, María del Carmen Lavalle,


Revista del Museo de La Plata, n. s., Botánica 14: 433-453 –

Stomata in Christensenia cumingiana (Marattiaceae – Eusporangiopsida)

Epidermis y estomas porociclocíticos en Christensenia cumingiana Christ (Marattiaceae – Marattiales ) Eusporangiopsida)

Rolleri C., Deferrari A. M., Lavalle M. del C. (1994)


Revista del Museo de La Plata, n.s., Botánica No. 98 – 14: 207-219 – ISSN 0376-2149 –


El estoma porocicloCÍtico es considerado aquí una especialización (funcional) de un estoma ciclocítico, tipo generalizado en el grupo.

Stomata in Marattiaceae (Eusporangiopsida)

Epidermis y estomatogenesis en Marattiaceae (Marattiales-Eusporangiopsida)

Rolleri C., Deferrari A. M., Ciciarelli M. de las M.  (1987)



Revista Mus. La Plata, n.s., Bot. 94: 129-147.

(Book: Cristina H. Rolleri, Universidad Nacional de La Plata, Facultad de Ciencias Naturales y Museo) – ISSN 0372-4611 –


Elevated stomata in Hemionitis (Pteridaceae)

Fig. 5. Lamina anatomy of Hemionitis umbrosa – A: paradermal section of abaxial face of epidermis with sinuous anticlinal wallsand anomocytic stomata; B: elevated stoma on abaxial surface (arrowhead)

A new species of Hemionitis (Pteridaceae) from central Brazil

Hirai R. Y., Cruz R., Prado J. (2018)

Hirai Regina Y., Cruz Rafael, Prado Jefferson,

Willdenowia 48(3): 371-380 – doi:



Hemionitis umbrosa (Pteridaceae) is described from the state of Goiás in central Brazil. It resembles H. tomentosa and was previously identified as such; however, it can be distinguished by having rhizome scales serrulate and spores cristate-reticulate. In contrast, H. tomentosa has rhizome scales entire or rarely sparsely denticulate and spores cristate.

The new species is morphologically and anatomically described, illustrated, and a key is provided to the species of Hemionitis with free veins.


Micromorphological and anatomical description —
Lamina. Unistratified epidermis with elevated anomocytic stomata present only on abaxial surface (Fig. 5A, B).

Stomata in Hemionitis (Filicopsida)

Pharmacognostic and Physico-Chemical Investigation of Hemionitis arifolia (Burm.) Moore

Joy J., Bindu A. R. (2017)

Jeethu Joy, Bindu A. R.,

University College of Pharmacy, Mahatma Gandhi University, Cheruvandoor, Ettumanoor-686631, Kottayam, Kerala, India


International Journal of Pharmacognosy and Phytochemical Research 9(8): 1074-1080 – doi: 10.25258/phyto.v9i08.9614,Vol9,Issue8,Article3.pdf

attached at one and with the anticlinal wall of the
epidermal cell .The guard cells are 50×70µm

Stomata of common pteridophytes in Pakistan

Morpho-Anatomical and Ethnobotanical Study of Common Pteridophytes of District Bannu, KPK, Pakistan

Khan R. U., Khan S. U. (2017)

Rehman Ullah Khan, Saad Ullah Khan,
Department of Botany, University of Science and Technology, Bannu, KPK, Pakistan


American-Eurasian J. Agric. & Environ. Sci. 17(6): 482-489 – ISSN 1818-6769 – DOI: 10.5829/idosi.aejaes.2017.482.489


The present study deals with the morphological, anatomical and medicinal uses of common pteridophytes found in district Bannu, KPK, Pakistan. During the survey Morpho-anatomical and medicinal uses of 5 genera and 5 pteridophytes species were reported, which were Adiantum capillus-veneris L., (families
Adiantaceae); Dryopteris affinis Lowe., (Dryopteridaceae); Equisetem arvensis L., (Equisetaceae); Marsilea quadrifoliata L. (Marsileaceae) and Nephrolepis exaltata (L.) Schott. (Nephrolepidiaceae).

It was concluded that the local people were aware of the medicinal uses of these plants. It was reported that Adiantum was used mostly as demulcent, expectorant, diuretic, eye ointment and tonic and is effective against acute bronchitis, fever, mouth blisters, febrifuge, cough and cold.


The epidermis possesses a number of important diagnostic character that offer valuable clues for identification, like size, shape and orientation of stomata,
guard cells and subsidiary cells
, structural peculiarities of epidermal cell walls, distinctive or specialized form of trichomes [20]. Leaf epidermal features like shape of epidermal cells, stomata and trichomes are useful anatomical tools.

Stomata of 3 Asplenium species

Morphology and anatomy of three species of Asplenium L. at Obafemi A Wolowo University ILE-IFE, Nigeria

Oladipo O. T., Philip T. V. , Bamigboye R. A., Oloyede F. A. (2020)

Oladipo O. T. 1 , Philip T. V. 1, Bamigboye R. A. 2, Oloyede F. A. 1,

1 Department of Botany, Obafemi Awolowo University, Ile-Ife, Nigeria.
2 Natural History Museum, Obafemi Awolowo University, Ile-Ife, Nigeria


Ife Journal of Science 22(1): 65-73 – DOI: 10.4314/ijs.v22i1.7


Comparative morphological and foliar anatomical studies were carried out on three species of Asplenium: Asplenium nidus L., Asplenium scolopendrium (L.) Newn and Asplenium barterii Hook with a view to identifying important morphological and anatomical characters that can be employed in the separation of the species.

Mature healthy plants of each species were collected from Erin Ijesa Waterfall and nursed in the Fern Garden of Natural History Museum at Obafemi Awolowo University, Ile-Ife. Morphological observations and measurements were documented for each species.

The transverse sections of the petiole and stem foliar sections were prepared by cutting at 10 µm thickness using Reichert Sliding Microtome and microphotographs were taken.

Quantitative data obtained from this study were subjected to one-way Analysis of Variance with Duncan Multiple Range Test for mean separation. Findings revealed that the predominantly irregular epidermal cells on the adaxial surface of A. barterii is diagnostic of the species.

The presence of tetracytic stomata in addition to anisocytic type also separates A. nidus from the other taxa. Similarly, A. scolopendrium had a significantly higher guard cell area compared to the other taxa.

Morphological and foliar anatomical features in the genus can be used to delimit the three species while the possession of two vascular bundles broadly admit the species into the genus Asplenium.

Stomata in Drymoglossum

Morphology, Physiology, and Anatomy of Penny Fern (Drymoglossum phyloselloides) and Its Effect on Cocoa

Yuliasmara F., Ardyani F. (2013)

Fitria Yuliasmara

Indonesian Coffee and Cocoa Research Institute

Fitria Ardyani

Indonesian Coffee and Cocoa Research Institute


Pelita Perkebunan (a Coffee and Cocoa Research Journal) 29(2): –


This study aims to determine the anatomy, physiology and morphology of penny fern (Drimoglosum phylloseloides) and its effect on cocoa. Morphological observation of penny fern used microscope to observe the roots, stems, leaves and spores. Physiology of penny fern was observed based on number of stomata and stomatal conductance using stomata printing method, while the amount of chlorophyll based on spectrophotometric method and rate of transpiration used cobalt chloride paper.

Penny fern anatomy on cross-sectional and longitudinal in roots, stems and leaves. Penny fern growth was observed based the length of tendrils once a week during rainy and dry season. While the effect of penny fern invasion was observed based on variable leaf area with gravimetric method, the cross-section of attacked cacao branch using microtom and microscope and chlorophyll content by chlorophyll meter. Results showed that penny fern is a epiphytic weed which was crassulaceae acid metabolism plants that have the ability to absorb carbon dioxide at night and carry out photosynthesis during the day with closed stomata. Penny ferns reproduce using spores. The growth rate of penny fern 2.18 cm/week during the dry season and while in rainy season 3.89 cm/week. Penny fern leaf contains 0.0212 mg/g chlorophyll.

Penny fern stomata density was 18.33/mm 2 with a width of opening stomata at night 26.3 µm which caused a veryslow rate of transpiration of 0.69 mm 2 /seconds. The existence penny fern on cocoa decreased leaf area and chlorophyll content decreased crop productivity which was indicated by decreasing in number of flowers, number of small, medium fruit, and large pods. However it had no effect on the number of leaves on one side flush cocoa.

New molecular and physiological evidence for the presence of active stomatal control in ferns

Evolutionary conservation of ABA signaling for stomatal closure

Cai S., Chen G., Wang Y., Huang Y., Marchant B., Wang Y., Yang Q., Dai F.,
Hills A., Franks P. J., Nevo E., Soltis D. E., Soltis P. S., Sessa E., Wolf P. G., Xue D., Zhang G., Pogson B. J., Blatt M. R., Chen Z. H. (2017)

Shengguan CaiGuang ChenYuanyuan WangYuqing HuangD. Blaine MarchantYizhou WangQian YangFei DaiAdrian HillsPeter J. FranksEviatar NevoDouglas E. SoltisPamela S. SoltisEmily SessaPaul G. WolfDawei XueGuoping ZhangBarry J. PogsonMichael R. BlattZhong-Hua Chen

Plant Physiol 174(2): 732-747 –


Abscisic acid (ABA)-driven stomatal regulation reportedly evolved after the divergence of ferns, during the early evolution of seed plants approximately 360 million years ago. This hypothesis is based on the observation that the stomata of certain fern species are unresponsive to ABA, but exhibit passive hydraulic control. However, ABA-induced stomatal closure was detected in some mosses and lycophytes. Here, we observed that a number of ABA signaling and membrane transporter protein families diversified over the evolutionary history of land plants. The aquatic ferns Azolla filiculoides and Salvinia cucullata have representatives of 23 families of proteins orthologous to those of Arabidopsis (Arabidopsis thaliana) and all other land plant species studied. Phylogenetic analysis of the key ABA signaling proteins indicates an evolutionarily conserved stomatal response to ABA. Moreover, comparative transcriptomic analysis has identified a suite of ABA-responsive genes that differentially expressed in a terrestrial fern species, Polystichum proliferum. These genes encode proteins associated with ABA biosynthesis, transport, reception, transcription, signaling, and ion and sugar transport, which fit the general ABA signaling pathway constructed from Arabidopsis and Hordeum vulgare. The retention of these key ABA-responsive genes could have had a profound effect on the adaptation of ferns to dry conditions. Furthermore, stomatal assays have shown the primary evidence for ABA-induced closure of stomata in two terrestrial fern species Pproliferum and Nephrolepis exaltata. In summary, we report, to our knowledge, new molecular and physiological evidence for the presence of active stomatal control in ferns.