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: https://doi.org/10.3372/wi.48.48305



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


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): – https://doi.org/10.22302/iccri.jur.pelitaperkebunan.v29i2.60



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 – http://doi.org/10.1104/pp.16.01848



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.

Stomata in Osmunda

Stomatal traits in Iberian populations of Osmunda regalis (Osmundaceae, Polypodiopsida) and its relationship with bioclimatic variables

Lavilla M., Seral A., Murciano A., Molino S., Fuente P. D. L., Galan J. M. G. Y. (2017)

Mikel Lavilla, Andrea Seral, Antonio Murciano, Sonia Molino, Pablo de la Fuente, Jose María Gabriel y Galán,

Acta Botanica Malacitana 42: 5-13 – DOI: https://doi.org/10.24310/abm.v42i1.3029



Stomata are very conserved structures in plants, which allows and control the  gas exchange. This ecophysiological fact appears to be critical in the ecology  and adaptation of plants to environment. Plant individuals can, among other  issues, adjust size and density of the stomata to adapt themselves to hydric,  thermic and light regimes. In turn, this led to the reduction of transpiration and  control of water losses, which is crucial in areas with Mediterranean climate. The  fern  Osmunda regalis  has populations in both biogeographical regions of the  Iberian Peninsula (Eurosiberian and Mediterranean regions), but when locations  are highly continental the populations are scarce and isolated. The objective of  this study is to characterize the stomatal morphological traits in both regions, to  detect possible adaptations in individuals occurring in Mediterranean locations.  26 individuals of 17 different populations were sampled. Applying microscopic  techniques, 4447 observations of length and width, and 234 of density and PCI  were done. The sampling spots were characterized by two bioclimatic indices of  Emberger and Gorzynski, and also mean maximum and minimum temperature  values were obtained. All the information was included in a dataset that was  statistically analyzed with the software SPSS. Our results show that  Osmunda  regalis  fronds have higher PCI and density values in the Eurosiberian region,  as expected. Gorczynski continentality index (K), and the mean maximum and  minimum annual temperatures influences over stomatal traits. This is clearly  informing that in the Mediterranean region, the temperature is an important  factor that triggers stomatal adaptations of  O. regalis  to continental locations with  higher levels of environmental stress. On the contrary, Emberger pluviometric  index (Q) seems to be less explicative by its own. Perhaps this is related with  the habitat of  O. regalis  in soils with high freatic level, fact which could lead to a  more independent stomata from rainfall.

Stomata in Microgramma (Polypodiaceae)

On the histochemical diagnostic characters of leaves from Microgramma squamulosa (Kaulf.) Sota (Polypodiaceae)

Jaime G. S., Barboza G., Vattuone M. A. (2007)

Gloria S. Jaime1 , G. Barboza2, M.A. Vattuone3,
1 Cátedra de Botánica y Cátedra de Fitoquímica

2 IMBIV.UNC. Argentina

3 Instituto de Estudios Vegetales. Fac. Bioq.Qca.y Farm. UNT. Ayacucho 471. San
Miguel de Tucumán.


Boletín Latinoamericano y del Caribe de
Plantas Medicinales y Aromáticas

Bol. Latinoam. Caribe Plant. Med. Aromaticas 6(5):195-196 – ISSN: 0717-7917 – https://www.redalyc.org/pdf/856/85617508034.pdf


En este trabajo se realiza el estudio exo-endomorfológico e histoquímico de las frondes de Microgramma squamulosa (Kaulf.) Sota,
“calaguala del mate”, especie empleada en medicina popular como antiulcerosa. Se obtuvieron parámetros micrográficos diagnósticos de M. squamulosa:
hojas hipostomáticas, estomas estaurocíticos, índice estomático = 5.76, hipodermis colenquimática en lámina y esclerenquimática en nervio medio; en
pecíolo esclerenquimática, con desarrollo de esclereidas en parénquima peciolar de hojas maduras que permiten reconocer la droga cruda ante eventuales
controles de calidad. Los tests histoquímicos demostraron la presencia de compuestos fenólicos y flavonoides de interés farmacológico.


The aim of this work was to study leaf, and petiole anatomical and histochemical characters of Microgramma squamulosa (Kaulf.) Sota, a fern belonging to Polypodiáceas (Pteridofitas). This species, popularly known as “calaguala del mate” in Argentina, is used in our folk medicine as antiulcer. In this contribution we describe the histological aspects that could help to identify the crude drug. Diagnostic micrographic parameters were obtained of this species: hipostomatic leaf, staurocytic stomata, stomata index = 5.76, collenchymatous hypodermis in blade and midrib with sclerenchyma; alone sclerenchymatic in petiole of mature leaves.

Hypodermal tissue and the occurrence of stomata at the abaxial face are typical characteristics of xeromorphic leaves

Leaf dimorphism of Microgramma squamulosa (Polypodiaceae): a qualitative and quantitative analysis focusing on adaptations to epiphytism

Rocha L. D., Droste A., Gehlen G., Schmitt J. L. (2013)

Ledyane Dalgallo Rocha1, Annette Droste1, Günther Gehlen1 , Jairo Lizandro Schmitt1

Rev. biol. trop 61(1) –  San José Mar. – Revista de Biología Tropical – On-line version ISSN 0034-7744 – Print version ISSN 0034-7744 –



The epiphytic fern Microgramma squamulosa occurs in the Neotropics and shows dimorphic sterile and fertile leaves. The present study aimed to describe and compare qualitatively and quantitatively macroscopic and microscopic structural characteristics of the dimorphic leaves of M. squamulosa, to point more precisely those characteristics which may contribute to epiphytic adaptations. In June 2009, six isolated host trees covered by M. squamulosa were selected close to the edge of a semi-deciduous seasonal forest fragment in the municipality of Novo Hamburgo, State of Rio Grande do Sul, Brazil. Macroscopic and microscopic analyzes were performed from 192 samples for each leaf type, and permanent and semi-permanent slides were prepared. Sections were observed under light microscopy using image capture software to produce illustrations and scales, as well as to perform quantitative analyses. Fertile and sterile leaves had no qualitative structural differences, being hypostomatous and presenting uniseriate epidermis, homogeneous chlorenchyma, amphicribal vascular bundle, and hypodermis. The presence of hypodermal tissue and the occurrence of stomata at the abaxial face are typical characteristics of xeromorphic leaves. Sterile leaves showed significantly larger areas (14.80cm2), higher sclerophylly index (0.13g/cm2) and higher stomatal density (27.75stomata/mm2) than fertile leaves. The higher sclerophylly index and the higher stomatal density observed in sterile leaves are features that make these leaves more xeromorphic, enhancing their efficiency to deal with limited water availability in the epiphytic environment, compared to fertile leaves.


El helecho epífito Microgramma squamulosa se encuentra en el Neotrópico y tiene hojas estériles y fértiles dimorfas. El objetivo de este estudio fue describir y comparar cuantitativa y cualitativamente la  organización estructural de las hojas de la M. squamulosa, investigando las características morfológicas y anatómicas, y señalando los factores que contribuyen a la adaptación al  ambiente epífito. Los análisis macroscópicos y microscópicos se realizaron a partir de 192 muestras de hojas de cada tipo. Láminas permanentes y semi-permanentes fueron  preparadas y las secciones fueron observadas en el microscopio de luz. Hojas fértiles y estériles no mostraron diferencias estructurales  cualitativas, son hipostomáticas, presentan epidermis uniseriada, haz vascular anficrival e hipodermis. La presencia de hipodermis y aparatos estomáticos en la nsuperficie abaxial de la epidermis son típicas de hojas xeromórficas. Hojas estériles presentaran  áreas significativamente mayores, tasa más alta de esclerofilia y mayor densidad de estomas que hojas fértiles. La mayor tasa de esclerofilia y mayor densidad de estomas son características que hacen las hojas estériles más xeromórficas, aumentando su eficiencia para hacer frente a la disponibilidad de agua limitada en el ambiente epífito, en comparación con las hojas fértiles.

Stomata in the Lateral-Line Aerenchyma in Bracken Ferns

Structure and Development of the Lateral-Line Aerenchyma in Bracken Ferns (Pteridium: Dennstaedtiaceae)

Barton D. A., Overall R. L., Thomson J. A. (1979)

Deborah A. BartonRobyn L. OverallJohn A. Thomson,

International Journal of Plant Sciences 176(7): –



Premise of research. Bracken ferns have a worldwide distribution. Plants consist of aerial fronds with petioles (stipes) that extend underground to specialized subterranean stems (rhizomes). A narrow continuous surface strip of specialized aerenchyma on each side of rhizomes and stipes, referred to as the lateral line, forms a pathway for gases. Developmental changes in the lateral line appear to affect gas exchange and influence water relations, soil exploitation, dormant survival, and disease resistance in these plants. An integrated account of the ontogeny and cytochemistry of the lateral line in the rhizomes and stipes of Pteridium has not been published.

Methodology. LM with fluorescence cytochemistry, supported by SEM, was used to compare developmental changes in tissues of the lateral-line aerenchyma in the stipes (petioles) and rhizomes (stems) of representative bracken taxa.

Pivotal results. The lateral lines of the rhizome and stipe in Pteridium aquilinum and Pteridium esculentum are a continuous organ. Developmental changes in the lateral lines of stipe and rhizome are similar. Cell walls of the closed lateral line but not the rind of the mature rhizome and the stipe belowground are rich in tannins. Intercellular protuberances extending from the walls of parenchyma cells below the lateral lines are primarily composed of pectin and polyphenols and contain tannin only late in development. Stomata of the lateral lines of the stipe above ground remain functional until frond senescence.

Conclusions. The lateral-line aerophores in the stipe and rhizome of bracken ferns are continuous, with similar anatomy, development, and likely roles in aeration and resistance to waterlogging.