Stomatal structure and stomatogenesis in Azolla (Filicopsida)

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Azolla pinnata at Belconnen,


Stomatal structure and stomatogenesis in Azolla pinnata P. Brown.

by Sen U. (1983)

Dep. of Bot., Kalyani Univ., Kalyani 741 235, West Bengal, India


in Ann. Bot. 52: 201-204 – ISSN 0305-7364 –


A stoma in A. pinnata consists of a unicelled binucleate guard cell with a pore, and one or more subsidiary cells towards the proximal side.

Stomatal development is of the polocytic or mesoperigenous anomocytic type except that the guard-cell mother cell fails to form 2 distinct guard cells because of restricted cytokinesis.

Similarities and differences between the stoma of A. pinnata and that of the fossil lycopod Zosterophyllum myretonianum are discussed.


Stomata in Pteridium caudatum (Dennstaedtiaceae) – (ferns)

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Lacy bracken fern (Pteridium aquilinum var. caudatum)


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Scanning electron micrographs of the lower epidermal surface of frond laminae in Pteridium. Fig. 1. LINN 1246.14. Scale bar = 200 µm. Fig. 2. LINN 1246.15. Scale bar = 200 µm. Fig. 3. P. caudatum, Venezuela, NSW 361276. Scale bar = 50 µm. Fig. 4. P. arachnoideum, Venezuela, NSW 361275. Scale bar = 50 µm. Fig. 5. P. arachnoideum, Surinam, NSW 360580. Scale bar = 50 µm. Fig. 6. P. caudatum, Costa Rica, NSW 420297. Scale bar = 25 µm. Fig. 7. LINN 1246.14. Scale bar = 25 µm. Fig. 8. P. arachnoideum, Costa Rica, NSW 420304. Scale bar = 25 µm.

Clarification of the taxonomic status and relationships of Pteridium caudatum (Dennstaedtiaceae) in Central and South America.

by Thomson J., Alonso-Amelot M. E. (2002)

John A. Thomson, National Herbarium of New South Wales, Royal Botanic Gardens, Mrs Macquaries Road, Sydney, NSW 2000, Australia

Miguel E. Alonso-Amelot

in Botanical Journal of the Linnean Society 140: 237–248 –

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Light microscopy of false-indusial segment margin (Fig. 9–12) and stomatal guard cells (Figs 13–16) after ruthenium red staining of epidermal peels of species of Pteridium. Scale bar = 25 µm for all micrographs. Fig. 9. LINN 1246.14. Fig. 10. LINN 1246.15. Fig. 11. P. caudatum, Mexico, NSW 360620. Fig. 12. P. arachnoideum, Brazil, NSW 360588. Fig. 13. LINN 1246.14. Fig. 14. LINN 1246.15, photographed through dense covering of long vein hairs. Fig. 15. P. caudatum, Peru, NSW 360578. Fig. 16. P. arachnoideum, Costa Rica, NSW 420304.


Contemporary systematic treatments of the Central and South American bracken ferns in the genus Pteridium Gled. ex Scop. recognize morphotype caudatum as either a full species or a variety of P. aquilinum (L.) Kuhn.

Geographically representative sporophytes of morphotype caudatum, including the type in the Linnaean Herbarium, are shown using spore size, guard-cell length and morphology of the cells of the false indusium to be tetraploid (based on 4n = 208).

DNA fingerprinting of field-collected Venezuelan samples supports the generalization that morphotype caudatum is a fertile allotetraploid containing genomic elements otherwise distinctive of the southern hemisphere diploid P. arachnoideum (Kaulf.) Maxon, together with elements characteristic of northern hemisphere diploids including the North American P. aquilinum var. pubescens Underw. and P. aquilinumvar. pseudocaudatum (Clute) A. Heller.

Evidence of genetic isolation from taxa with overlapping distributions, as well as morphological, biochemical and ecological data, validate recognition of P. caudatum (L.) Maxon at species level.

Heterogeneity observed within P. caudatum is consistent with multiple origins through independent hybridization events.

Pteridium caudatum is strikingly analogous to the tropical Asian/Australasian allotetraploid P. semihastatum (N. Wallich ex J. G. Agardh) S. B. Andrews [=P. yarrabense (Domin) N. A. Wakef.].


Knowledge of ploidy level is of considerable potential value in systematic and evolutionary studies, but attains significance only when generalization to an entire taxonomic unit becomes possible. Methods for unequivocal direct assessment of ploidy, including chromosome counts, determination of nuclear DNA content or molecular genome and isozyme analyses, typically require fresh or specially preserved material and are inapplicable to routine dried herbarium specimens. Correlates of ploidy level, especially stomatal guard-cell length and spore size (Barrington, Paris & Ranker, 1986), must therefore be used to generalize results obtained on a few individuals by direct methods to the taxon as a whole. The relationship between guard-cell length and ploidy level has been established for P. arachnoideum and P. caudatum (Thomson, 2000a) by (i) measurement of guard-cell length in specimens of P. arachnoideum and a presumed hybrid from the Galapagos Islands that showed chromosome complements of 2n = 104 and 4n = 208, respectively (Jarrett, Manton & Roy, 1968), and (ii) measurement of nuclear DNA content (Table 1;2c = 16.5 pg for P. arachnoideum and 4c = 30.9 pg for P. caudatumTan & Thomson, 1990). SEMs of the guard cells of P. caudatum (NSW 420297, Costa Rica) are shown in Figure 6 for comparison with those of Linnaeus’ specimen 1246.14 (Fig. 7) and P. arachnoideum (NSW 420304, Fig. 8). Guard cells of the type of P. caudatum (LINN 1246.15) could not be visualized with the SEM using the limited tissue available for examination due to the dense overlying indumentum in this specimen (Fig. 2, cf. Fig. 1). Light micrographs of guard cells in LINN 1246.14 and LINN 1246.15 are shown in Figures 13 and 14, respectively, with P. caudatum (NSW 360578) from Peru (Fig. 15) and P. arachnoideum (NSW420304) from Costa Rica (Fig. 16) for comparison. Consistent with a difference in ploidy level between them, the larger size and less crenulate margins of the epidermal cells in P. caudatum (including the Linnaean specimens) compared with those of P. arachnoideum are evident in both the SEM photographs of Figures 6–8and the light micrographs of Figures 13–16.

Mean guard-cell lengths, based on measurements for 30 stomata for each accession, are shown in Table 1. These data greatly extend the preliminary observations reported by Thomson (2000a,b) and suggest that all specimens assigned to P. arachnoideum on morphological grounds are diploid, and are consistent with the conclusion that all accessions of P. caudatum, including LINN 1246.14 and the type (LINN 1246.15) are tetraploid. Within both P. arachnoideum and P. caudatum there is significant between-accession variation in guard cell length. The ranges of the means for the two groups do not overlap. The smallest difference between mean guard-cell lengths for any pair of P. caudatumand P. arachnoideum specimens represented in Table 1 is significant at P < 0.001. In overview, accessions of P. arachnoideum have a mean guard cell length less than 34 µm; those of P. caudatummore than 38 µm. Guard-cell length in the North American diploid brackens (vars latiusculumpseudocaudatum and pubescens) is significantly greater than in the southern diploid P. arachnoideum(Thomson, 2000a), limiting the value of this character as an independent indicator of ploidy level. Similar differences in the relationship of ploidy to guard-cell length have been reported for other fern taxa (Barrington et al., 1986), and presumably reflect the adaptive significance of stomatal morphology in gas and water economy.

The normal and the abnormal stomata in Pteridium (fern)

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Pteridium aquilinum (bracken fern)


The abnormal stomata on the diploid and the tetraploid bracken ferns.

by Takahashi C. (1962)

Chihiro Takahashi, Biological Laboratory, Department of General Education, Nagoya University

in Cytologia, 27: 151–157 –

The normal and the abnormal stomata on the leaf of Pteridium aquilinum var. latiusculum were studied.

The normal stoma on the diploid or the tetraploid plant is formed as follows: a stomatal initial appears in the dermatogen; dividing once or twice it gives rise to a stomatal mother cell; a stomatal mother cell divides into a pair of cells; they develop into a pair of guard cells differentiated typically.

The persistent stomatal mother cell is rarely observable on all diploid and tetraploid plants. It remains undivided, circularly or elliptically shaped, and not thick-walled.

The epidermized stoma is rarely observable on the tetraploid plants. In this type a stomatal mother cell divides into a pair of cells but they develop erratically into the epidermis-like cells. They are not raised above the level of the epidermis. Each cell of the epidermized stoma has the wavelike shape, no thick wall and no pore between two cells.

The existence of the poreless multicellular stoma is restricted to three of the tetraploid plants. This interesting stoma is characterized by the following features: it is constructed out of many cells (three to more than ten) caused by the further divisions of two cells originating from a stomatal mother cell; guard cells are not differentially thick-walled; no pore is formed; guard cells are stained not so well with Lugol solution; the chloroplast number is variable and sometimes very few; not a normal stoma in structure and function is formed on these three plants.

The tetraploid plants bearing this stoma exhibit many other deviations from the normal morphology and physiology. However, such deviations are not specific to three plants but appear on the other tetraploid plants.

Also the causes of the occurrence of these anomalous stomata were discussed.

Stomata in Pteridium aquilinum (ferns)

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Bracken fern (Pteridium aquilinum)

Anomalous stomata on polyploid bracken.

by Takahashi C. (1960)

Chihiro TAKAHASHI, Biological Laboratory, Department of General Education, Nagoya University

in Bot. Mag. Tokyo 73. 160 –

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Stomata in Gleichenia and Dicranopteris (ferns)

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Gleichenia gigantea

Stomatogenesis in Gleichenia gigantea, Dicranopteris linearis var. montana and D. splendida.

by Sen U.,  Bhunia S. (1984)

in Blumea 30: 13-16 –

Photo credit Google – Dicranopteris splendida –


This is the first ever study of the development of stomata in Gleichenia gigantea Wallich ex Hook. (subg. Diplopterygium), Dicranopteris linearis (Burm. f.) Underw. var. montana Holttum and D. splendida (Hand.-Mazz.) Tagawa in detail.

In these ferns, as in many polypodioid members, the polocytic stomata far outnumber the mesoperigenous anomocytic type.

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Stomatal development in Davallia (ferns)

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Davalliaceae: Davallia denticulata

The evidence of stomatal development on the relationships between Davallia and genera associated with it by recent authors.

by Sen T. (1986)

Tuhinsri Sen

Kalyani University, Kalyani, India

in Ann. Bot. 58: 663-677 – –


The structure and ontogeny of stomata occurring on the fronds of Davallia and allied genera, as arranged by Copeland (1947) and Holttum (1947), are investigated.

Of the nine types of stomata recorded, seven are new reports for these ferns. All the nine types are mesoperigenous in origin, and pass through either the polocytic or the copolocytic condition during their formative stages.

In the hemiparacytic stomata, the long axis of the nuclear spindle in the dividing guard-cell mother cell is always at right angles to the wall first formed by the stomatal meristemoid, and not parallel to it as in all other types.

Stomatal structure supports Holttum’s scheme, in which Rumohra (placed in Aspidiaceae by Copeland) is associated with Davallia, and also his separation of the genera Nephrolepis, Oleandra, Arthropteris and Psammiosorus as a distinct group; among the latter, Nephrolepis rather than Oleandra (Pichi Sermolli, 1977), is shown to be peculiar. It is also shown that the type species of Paradavallodes (Ching, 1966) does not differ from Araiostegia.

Stomata in Isoetes (ferns)

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The genus Isoetes in India

by Pant D. D., Srivastava G. K. (1962)

Allahabad University, India

Divya Darshan Pant, Gopal Krishna Srivastava,

in Proc. Nat. Inst. Sci. India 28: 242-280 –

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