Stomata in Gnetum (Gymnospermae)

Photo credit: Google

Gnetum gnemon

The Stomata of Gnetum

by Maheshwary P., Vasil V. (1961)

in Ann. Bot 25: 313-319.


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In both Gnetum gnemon and G. ula the stomatal development in the collars and axes of the male and female cones and the outer two envelopes of the ovule is of the haplocheilic type.

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In the leaf some stomata do show parallel subsidiary cells, but they arise from the surrounding epidermal cells and do not seem to have a common origin with the guard cells. This conflicts with earlier accounts and is of interest in relation to the types of stomata met with in other gymnosperms, living and fossil.

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See the text: Ann. Bot.

Stomata in Podocarpus

Photo credit: NCBI

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)

in Ann Bot. 2012 Oct;110(5):1057-66.

doi: 10.1093/aob/mcs179. Epub 2012 Aug 7.



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.


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.

See the full text: Annals of Botany


Anderson A. A. (1897) – Stomata on the bud scales of Abies pectinata – Bot. Gaz. 1897: 94-95 – (On our blog :

Barone Lumaga M.R. , Moretti A., De Luca P. (1999) – Morphological aspects of stomata, cuticle and chloroplasts in Ceratozamia kuesteriana Regel (Zamiaceae) – Plant Biosystems, Volume 133, Issue 1, 1999. – – – (On our blog :

Bender O., Rudnik T. I. (2008) – Variability of needle structure in Siberian stone pine in provenance plantations – Annals of Forest Research 51(1): 165-168 – – (On our blog :

Brodribb T., Hill R. S. (1997) – Imbricacy and stomatal wax plugs reduce maximum leaf conductance in Southern Hemisphere conifers – Australian Journal of Botany 45: 657–668 – – (On our blog : )

Campbell R. (1972) – Electron microscopy of the epidermis and cuticle of the needles of Pinus nigra var. maritima in relation to infection by Lophodermella sulcigena – Ann. Bot. 36: 307-314 – (On our blog :

Chabot J. F., Chabot B. F. (1977) – Ultrastructure of the epidermis and stomatal complex of balsam fir (Abies balsamea). – Can J Bot 1977, 55:1064-1075. -DOI: 10.1139/b77-125.(Publisher Full Text) – View Article – – (On our blog :

Chaturvedi S. (1995) –  Micromorphology and vegetative anatomy of leaves of Taxodiaceae. – Int. J. Mendel, 12: 81. (Abstract not found – Who can send us one ?)

Chen L. Q., Li C. S. (2004) – The epidermal characters and stomatal development of Ginkgo biloba. – Bull. Bot. Res. 24: 417-422. – – (On our blog :

Coiro M. (2015) – The unusual stomatal apparatus of Stangeria reveals a familiar tale – – (On our blog :

Davies W. J., Kozlovski T. T., Lee K. J. (1974) – Stomatal characteristics of Pinus resinosa and Pinus strobus in relation to transpiration and antitransparant efficiency – Can. J. For. Res. 4: 571-574 – – – (On our blog :

Denniston K. (2015) – Stomatal Bloom – Northwest Conifer Connections JULY 15, 2015 – – (On our blog :

Florin R. (1931) – Untersuchungen zur Stammesgeschichte der Coniferales. Morphologie und Epidermisstruktur der Assimilationsorgane bei den rezenten Koniferen – K. Svenska Vet. Akad. Handl. 10: 1-588.

Florin R. (1933) – Studien über die Cycadales des Mesozoikums. Erörterungen über die Spaltöffnungdapparate der Bennettitales.  K. Svenska Vet. Akad. handl. 12: 1-134.

Florin R. (1934) – Die Spaltöffnungsapparate von Welwitschia mirabilis Hook. f. – Svensk. Bot. Tidskr. 28: 265-284 – (Article not found)

Florin R. (1958) – Studien über die Cycadales des Mesozoikumus nebst Erörterungen über die Spaltöffnungsapparate der Bennttitales. K. Svenska Vetensk. Akad. Handl., ser. 3, 12.

Franich R. A., Wells L. G., Barnett J. R.  (1977) – Variation with tree age of needle cuticle topography and stomatal structure in Pinus radiata D. Don. – Ann. Bot. 41: 621-626 – – – (On our blog :

Ghimire B., Lee C., Heo K. (2014) – Leaf anatomy and its implications for phylogenetic relationships in Taxaceae s. l. – J Plant Res. 127(3):373-388 – doi: 10.1007/s10265-014-0625-3 – Epub 2014 Feb 5 – – (On our blog : )

Hamidipour A., Radjabian T., Charlet D. A., Zarrei M. (2011) – Leaf anatomical investigation of Cupressaceae and Taxaceae in Iran – Wulfenia 18: 95 –111 – – (On our blog : )

Hanover J. W., Reicosky D. A. (1971) – Surface wax deposits on foliage of Picea pungens and other conifers – Amer. J. Bot. 58(7): 681-687 – (On our blog :

Hildebrand,  (1860) – Der Bau der Coniferenspaltöffnungen und einige Bemerkungen uber die Vertheilhung derselben. – Bot. Zeit. (18): 149-152 – (On our blog :

Inamdar J. A., Bhatt D. C. (1972) – Epidermal structure and ontogeny of stomata in vegetative and reproductive organs of Ephedra and Gnetum – Ann. Bot. 36: 1041-1046. – (On our blog :

Jeffree C. E., Johnson R. P. C., Jarvis P. G. (1971) – Epicuticular wax in the stomatal antechamber of sitka spruce and its effect on the diffusion of water vapour and carbon dioxide – Planta 98: 1-10 -DOI: 10.1007/BF00387018 – – (On our blog :

Johnson R. W., Riding R. T. (1981) – Structure and ontogeny of the stomatal complex in Pinus strobus L. and Pinus banksiana Lamb. – Am J Bot 1981, 68: 260-268. – 10.2307/2442858. -(Publisher Full Text) – View Article – (On our blog :

Kanis A. W., Karstens K. H. ( 1963) On the occurrence of amphistomatic leaves in Ginkgo biloba L. – Acta Botanica Neerlandica 12: 281-286. – DOI: 10.1111/j.1438-8677.1963.tb00121.x – – (On our blog :

Karatela Y.Y., Gill, S. L. (1984) –  Epidermal structures and stomatal ontogeny in some Gymnosperms from Nigeria. – Feddes Rep. 95(5-6): 351-354. – (No abstract found – Who can send us one ?)

Kausik S. B. (1974) – Ontogeny of the stomata in Gnetum ula Brongn. – Bot. J. Linn. Soc. 68 (2): 143-151 – (On our blog :

Kausik S. B. (1974) – The stomata of Ginkgo biloba L., with comments on some noteworthy features – Bot. J. Linn. Soc. 69: 137- 146. – (On our blog :

Kausik S. B. (1976) – A contribution to foliar anatomy of Agathis dammara, with a discussion of the transfusion tissue and stomatal structure – Phytomorphology 26: 263-273 – (Article not found)

Kouwenberg L. L. R. , McElwain J. C. , Kurschner W. M., Wagner F., Beerling D. J., Mayle F. E., Visscher H. (2003) – Stomata frequency adjustment of four conifer species to historical changes in atmospheric CO2 – Am. J. Bot. 90. 610-619.- doi: 10.3732/ajb.90.4.610 – – (On our blog : – No. of stomata per mm of needle length. Tsuga, Picea, Larix.

Lehela A. R., Day R. J., Koran Z. (1972) – A close up of the stomatal regions of white spruce and jack pine – For. Chron. Feb. 1972: 32-34 – – (On our blog :

Locosselli G. M., Ceccantini G. (2012) – Plasticity of stomatal distribution pattern and stem tracheid dimensions in Podocarpus lambertii: an ecological study – Ann Bot 110(5): 1057–1066 -doi:  10.1093/aob/mcs179 – PMC3448432 – – (On our blog : )

Ma Q.-W., Li C.-S., Li F.-L., Vickulin S. V. (2004) – Epidermal structures and stomatal parameters of Chinese endemic Glyptostrobus pensilis (Taxodiaceae)  – Botanical Journal of the Linnean Society 146(2): 153–162 – – – (On our blog : )

Maäcz J. G. (1956) – Über die Blattepidermis von Ginkgo biloba L. – Acta Biol. Szeged 4: 403-410. – (Article not found)

MacDonald G. M. (2002) – Conifer Stomata in: Tracking Environmental Change Using Lake Sediments: Volume 3: Terrestrial, Algal and Siliceous Indicators edited by J. P. Smol, H. J. Birks, W. M. Last – Kluwer 2002: 33-47. – – (On our blog :

Maheshwari P., Vasil V. (1961) – The stomata of Gnetum – Ann. Bot 25: 313-319 – – (On our blog :

Menendez C. A., Caccavari M. A. (1966) – Estructura epidermica de Araucaria nathortstii Dus. de Terciario de Pico Quemado, Rio Negro – Ameghiniana 4: 195-197 – (Article not found)

Nautiyal D. D., Singh S., Pant D. D. (1976) – Epidermal structure and ontogeny of stomata in Gnetum gnemom, G. montanum and G. ula – Phytomorphology 26: 282-296. – (On our blog :

Pant D. D., Basu N. (1977) – A comparative study of the leaves of Cathaya argyrophylla Chun&Kuang and three species of Keteleeria Carrière – Bot. J. Linn. Soc. 75: 271-282 – (On our blog :

Pant D. D., Mehra B. (1963) – Epidermal Structure and Development of Stomata in Ephedra foliata Boiss. – The New Phytologist Vol. 63, No. 1 (Mar., 1964), 91-95 – DOI: 10.1111/j.1469-8137.1964.tb07362.x – – (On our blog :

Pant D. D., Mehra B. (1964) – Development of stomata in leaves of three species of Cycas and Ginkgo biloba L. – Bot. J. Linn. Soc. 58: 491-497. –DOI: 10.1111/j.1095-8339.1964.tb00917.x – – (On our blog :

Pant D. D., Mehra B. (1964) – Development of stomata in leaves of three species of Cycas and Ginkgo biloba L. – Bot. J. Linn. Soc. 58: 491-497 – (On our blog :

Pant D. D., Nautiyal D. D., Singh S. (1978) – Cuticular studies of reproductive organs of Gnetum and Ephedra – Phytomorphology 28(2): 141-150 – (On our blog :

Pant D. D., Srivastava G. K. (1968) – On the cuticular structure of Araucaria (Araucarites) cutchensis (Feistmantel) comb. nov. from the Jabalpur series, India – Bot. J. Linn. Soc. 61: 201-206  – (On our blog :

Pant D. D., Verma B. K. (1974) – Taxonomy of the genus Ephedra. Significance of stem and leaf epidermis and cuticle – Bot. J. Linn. Soc. 69: 287-308 – (On our blog :

Pisaric M. F. J., Szeicz J. M., Karst T., Smol J. P. (2000) – Comparison of pollen and conifer stomates as indicators of alpine treeline in northwestern Canadian lake sediments – Can. J. Bot. 78: 1180–1186 – – (On our blog :

Reed J. E., Smith W. K. (2012) – Stomatal Frequency, Distribution, and Needle Hydrophobicity in Cloud Forest Spruce and Fir, Southern Appalachian Mountains – RURALS: Review of Undergraduate Research in Agricultural and Life Sciences: 7(1):  Article 3 – – (On our blog : )

Rhine J. B. (xxxx) – The clogging of stomata in conifers – Thesis (M.S.)–University of Chicago, Department of Botany – – (No abstract found – Who can send us one ?)

Rhine J. B. (1924) – Clogging of Stomata of Conifers in Relation to Smoke Injury and Distribution – Botanical Gazette Vol. 78, No. 2 (Oct., 1924), pp. 226-232 – – (On our blog :

Riding R. T., Aitken J. (1982) – Needle structure and development of the stomatal complex in cotyledons, primary needles, and secondary needles of Pinus radiata D. Don – Bot. Gaz. 143: 52-62 – (On our blog :

Rook D. A., Hellmers H., Hesketh J. D. (1971) – Stomata and cuticular surfaces of Pinus radiata needles as seen with the scanning electron microscope – J. Ariz. Acad. Sci. 6: 222-225 – (Article not found)

Rudall P. J., Bateman R. M. (2019) – Leaf surface development and the plant fossil record: stomatal patterning in Bennettitales – Biological Reviews 94(3) – – – (On our blog : )

Rudall P. J., Rice C. L. (2019) – Epidermal patterning and stomatal development in Gnetales – Annals of Botany, mcz053 – – – (On our blog : )

Rudall P. J., Rowland A. R., Bateman R. M. (2012) – Ultrastructure of stomatal development in Ginkgo biloba. – International Journal of Plant Sciences 173: 849–860. – DOI: 10.1086/667230 – – (On our blog :

Salt A. (2019) – Epidermal patterning and stomatal development in Gnetales – Botany One May 30, 2019 – – (On our blog : )

Schwabach E. (1902) – Zur Entwicklung der Spaltöffnungen bei Coniferen – Ber. Deutsch. Bot. Ges. 20: 1-7 – (On our blog :

Sikarwar R., Rajawat B. S., Sharma K. R. (2014) – Studies on relationship between Stomatal density and oleoresin yield in Chirpine (Pinus roxburghii Sargent) – Int. Journ. Adv. Res. (2014), Volume 2, Issue 3, 751-758 –  – (On our blog :

Spjut R. W. (2014) – A new variety of Taxus brevifolia from the Pacific Northwest of North America – The World Botanical Associates, The IV International Workshop, Oct 23–25, 2014. – (No abstract found – Who can send us one ?)

Stockey R. A., Ko H.  (1990) – Cutical micromorphology of Dacrydium (Podocarpaceae) from New Caledonia. – Bot. Gazette 151: 138-149 – (ISSN: 0006-8071)DOI : 10.1086/337813 – – – (On our blog :

Stockey R. A., Frevel B. J., Woltz P. (1998) – Cutical micromorphology of Podocarpus, subgenus Podocarpus, section Scytopodium (Podocarpaceae) of Madagascar and South Africa. – Int. J. Plant Sci. 159: 923-940 – DOI: 10.1086/297613 – – (On our blog :

Stockey R. A., Taylor T. N. (1978) – Cuticular features and epidermal patterns in the genus Araucaria De Jussieu – Bot. Gaz. 139(4): 490-498 – (On our blog :

Su J. X., Liao F., Huang Y. Y. (2003) – The leaf epidermis of three species in Zamiaceae. – Acta Bot. Yunn. 25: 596-602. – (No abstract found – Who can send us one ?)

Sweeney C. A. (2000-2004) – Conifer stomata analysis in Late Quaternary paleoecolgy in Scandinavia – AMAP Project Portal – – (On our blog :

Sweeney C. A. (2004) – A key for the identification of stomata of the native conifers of Scandinavia – Review of Palaeobotany and Palynology, vol. 128, no. 3–4, pp. 281 – 290, 2004. – – – (On our blog :

Takeda H. (1913a) – Development of stomata in Gnetum gnemon – Ann. Bot. 27: 365-366 – (Article not found)

Takeda H. (1913b) – Some points in the anatomy of leaves of Welwitschia mirabilis – Ann. of Bot. 27: 347-375 – – (On our blog :

Wan H., Shen J., Tang L., Li C. (2010) – Morphology of stomata of Pinaceae and Cupressaceae from northwestern China – – (On our blog : )

Whiting M., Mill R. R., Jeffree C. E. (2017) – The stomatal complex of Podocarpus observed in cross-section using cryofracture: a preliminary study – Edinburgh Journal of Botany 74(3): 345-364 – – (On our blog : )

Wilhelm K. (1883) – Ueber eine Eigentumlichkeit der Spaltöffnungen bei Coniferen. – Ber. dtsch. bot. Ges. (I): 325-30 – (Article not found)

Wu J. L., Niu J. Y., Yan Z. Z., Li S., Gao Y. H., Jiang H. Y. (2007) – [SEM observation on leaf epidermis of different Ephedra species] (in Chinese) – Zhongguo Zhong Yao Za Zhi. 32(18):1854-1857 – PMID: 18051888 – – (On our blog : )

Xu B.-s., Zhang R.-h. ( 2002) – Study on chlorophyll content and stomata morphology of Pïnus massoniana – Chemistry and Industry of Forest Products 22(3): 59-61  – – (On our blog :

Stomata in Pseudotsuga (Gymnospermae)

Photo credit: Gardens Online

Douglas Fir (Pseudotsuga menzieszii)

Morphology and Stomatal Function of Douglas Fir Needles Exposed to Climate Change: Elevated CO2 and Temperature

by Apple M.E., Olszyk D.M., Ormrod D.P., Lewis J., Southworth D., Tingey D.T. (2000)

in Int. J. Plant Sci. 2000 Jan;161(1):127-132.


Climate change may have an impact on the productivity of conifer trees by influencing the morphology (size and surface characteristics) and function (capacity for gas exchange) of conifer needles.

In order to test the responses of needles to climatic variables, Douglas fir (Pseudotsuga menziesii [Mirb.] Franco), saplings were grown in sunlit controlled environment chambers at ambient or elevated (+200 parts per million above ambient) CO2 and at ambient or elevated temperature (+4 degrees C above ambient).

Needle characteristics, including length, width, area, stomatal density (stomata per mm2), percentage of stomatal occlusion, and the morphology of epicuticular wax, were evaluated.

Needle function was evaluated as stomatal conductance to water vapor and transpiration. Needle length increased significantly with elevated temperature but not with elevated CO2. Neither elevated CO2 nor elevated temperature affected stomatal density or stomatal number in these hypostomatous needles.

Epicuticular wax was less finely granular at elevated than at ambient temperature and was similar in appearance at elevated and ambient CO2. Stomatal conductance and transpiration increased with elevated temperature and associated increased vapor pressure deficit; however, neither conductance nor transpiration was affected by elevated CO2.

These results indicate that simulated climate change influences Douglas fir needle structure and function.

See the text: NCBI-PubMed

Stomata in Ceratozamia (Gymnosperms)

Photo credit: Google

Ceratozamia kuesteriana

Morphological aspects of stomata, cuticle and chloroplasts in Ceratozamia kuesteriana Regel (Zamiaceae)

by Barone Lumaga M. R., Moretti A.De Luca P. (1999)

in Plant Biosystems, Volume 133, Issue 1, 1999


Light and scanning electron microscopy were utilised to study stoma and cuticle morphology whereas transmission electron microscopy was used to observe plastid ultrastructure in Ceratozamia kuesteriana Regel (Zamiaceae).

Results show that in C. kuesteriana a diperigenous‐type stoma (or a derivation of a diperigenous type) occurs and that protein crystalloids and prolamellar bodies are simultaneously present in the chloroplast.

Read the text: Taylor & Francis Online

Stomata in Taxus (Gymnosperms)

Photo credit: World Botanical Association

Taxus globosa var. floridana, showing stomata band with 7 rows of stomata, and midrib and marginal cells in mid region of leaf on abaxial surface

A new variety of Taxus brevifolia from the Pacific Northwest of North America

by Spjut R. W. (2014)

©The World Botanical Associates

The IV International Workshop, Oct 23–25, 2014,

The Taxus wallichiana Subgroup  in North  America is characterized by leaves having tall rectangular to nearly quadrangular epidermal cells as seen in cross section (illus. above map), stomata in 12 rows/band, and a papillose abaxial midrib (top two rows of illus.).

There are two species, Taxus  brevifolia, subdivided into four varieties, var.brevifolia, var. klamathensis, var. polychaeta, and var. reptaneta, and T. globosa subdivided into two varieties, var. floridana and var. globosa.

The two species are distinguished by number of stomata rows in a stomata band, 4–7 (-9) in T. brevifolia, and 7–11 in T. globosa as plotted on the map above from plant specimens in herbaria and collected in the field, and by the shape of the marginal cells that border stomata bands, appearing ± rectangular in T. globosa, irregular  and inflated in T. brevifolia.

Taxus canadensis, which also occurs in the Mediterranean  Region (Spjut , is classified in the Taxus cuspidata Alliance (Spjut 2007b) of the Taxus baccata Group.  It is recognized by the absence of papillae across the midrib and marginal cells on the abaxial surface (Spjut 1992, 1993, 2000, 2007b).

Stomata of fossil Pinus (Gymnosperrms)



by C. A. Froyd

in Ecology 86:579–586.

The analysis of fossil stomata reveals the early postglacial presence of Pinus sylvestris at two sites in the Scottish Highlands, 1600 and 600 years prior to the arrival times indicated by traditional palynological methods.

Fossil stomata provide unambiguous evidence of past local presence for Pinus sylvestris, which produces abundant and widely dispersed pollen, revealing its presence when pine pollen frequencies are as low as 0.4%, considerably below the commonly adopted minimum frequency threshold of 20%. Thus a species may be present for hundreds to thousands of years before expansion of the local population is registered in the palynological record. This has significant implications, not only for the initial spread of pine throughout the British Isles, but more generally for analyses of the continental-scale migration of temperate and boreal forest taxa based on palynological data.

Failure to differentiate effectively among the processes of arrival, establishment, and expansion in analyses of plant migration rates and patterns means that many existing reconstructions of postglacial colonization may, in actuality, represent the expansion of populations over time, rather than the initial spread of species.

Read More: ESA Ecology