Historical development of the present classification of morphological types of stomates

There is a long-standing confusion between morphologic and ontogenetic classifications of stomates. The earliest scheme, by Vesque (1889) was proposed as basically ontogenetic, but it was soon widely applied to mature leaves. The ontogenetic distinction between haplocheilic and syndetocheilic stomates in gymnosperms, proposed by Florin (1931, 1933) soon suffered a similar fate. Continuing studies over the past half-century have shown that stomatal ontogeny is only poorly correlated with the mature morphology. Efforts to combine ontogenetic and morphologic features in a single scheme have led to classifications so complex as to be impractical. The explicitly morphological classification by Metcalfe and Chalk (1950) is the foundation for the most widely used present scheme, in which some 14 morphological types are recognized. The distinctions among these types are conceptually useful, though often arbitrary in practice.     

Observations on the structure and ontogeny of stomata and trichomes on developing and mature pericarps ofCassia occidentalis L

Paracytic, anisocytic, anomocytic, transitional forms, tetracytic, cyclocytic stomata and partly and completely amphicyclic forms are found, often on the same surface, in nine combinations. The most frequent type is paracytic. A few morphological variations in the basic types and eight types of abnormalities in stomata are recorded. The stomatal ontogeny may be mesogenous, mesoperigenous or perigenous. Trichomes are multicellular glandular club-shaped and unicellular eglandular. The taxonomic significance of stomata is indicated.     

On the laterocytic stomatotype in angiosperms

The laterocytic type of stomatal apparatus in angiosperms is considered. Investigation of the stomatal apparatus of 18 species of Hamamelidaceae, representing 15 genera, showed that in addition to the anomocytic, paracytic, and encyclocytic stomatotypes previously known in the family, the laterocytic type is found in several genera (Dicoryphe, Exbucklandia, and others). Study of 15 species of Chloranthaceae, representing all five genera, showed that laterocytic stomates occur inChloranthus andSarcandra and sometimesHedyosmum, along with stomates of other types.Barbeya oleoides (Barbeyaceae) and the four investigated species ofBalanops (Balanopaceae) have exclusively (or inB. oliviformis mainly) laterocytic stomates. Laterocytic stomates are present also inKadsura andSchisandra of the Schisandraceae, along with the previously known paracytic type. In addition to the foregoing genera, laterocytic stomates are known in some members of the Buxaceae, Celastraceae, Crypteroniaceae, Hydrangeaceae, Icacinaceae Platanaceae, Tetracentraceae and Trochodendraceae.     

A classification of stomatal types

In 1950 Metcalfe & Chalk showed the need for new technical terms 'devoid of taxonomic or ontogenetic implications' to replace those of Vesque. Their new terms and some others, proposed by Metcalfe and Stace, are now generally used. All previous classifications of stomatal types included only those of Angiosperms and Gymnosperms. The eight new forms described in this and a previous paper (Van Cotthem, 1968) include five known only from the Filices and bring the number of known stomatal types to 15, of which two have been subdivided.     

HELICOCYTIC AND ALLELOCYTIC STOMATA: UNRECOGNIZED PATTERNS IN THE DI COTYLEDON AE

Two new mesogenous stomatal patterns are described for the Dicotyledonae: helicocytic, with a helix of four or more subsidiary cells surrounding the guard cell pair, and allelocytic, with an alternating complex of three or more C-shaped subsidiary cells of graded sizes. The latter pattern may have guard cells developed parallel to the subsidiary cells (parallelocytic) or at right angles to them (diallelocytic). For both types, the complex development of regular sequences of mesogene subsidiary cells with particular attributes of size and arrangement provides strong probability of correlated ontogeny and adult pattern. They are related to mesogenous forms of the anisocytic, paracytic and diacytic patterns, but these latter may also be developed in different ways. The use of stomatal pattern data for taxonomic and evolutionary studies of dicote is discussed.     

Variation in the structure and development of foliar stomata in the Euphorbiaceae

The structure and ontogeny of foliar stomata were studied in 50 species of 28 genera belonging to 17 tribes of the family Euphorbiaceae. The epidermal cells are either polygonal, trapezoidal, or variously elongated in different directions and diffusely arranged. The epidermal anticlinal walls are either straight, arched or sinuous. The architecture of cuticular striations varies with species. The mature stomata are paracytic (most common), anisocytic, anomocytic and diacytic. Occasionally a stoma may be tetracytic, cyclocytic or with a single subsidiary cell. The ontogeny of paracytic stomata is mesogenous dolabrate or trilabrate, mesoperigenous dolabrate; that of diacytic stomata is mesogenous dolabrate, whereas that of anisocytic stomata is mesogenous trilabrate; rarely an anisocytic stoma may be mesoperigenous. Hemiparacytic stomata are mesoperigenous unilabrate; tetracytic stomata are mesoperigenous dolabrate and anomocytic stomata perigenous. Abnormalities encountered include four types of contiguous stomata, stomata with a single or both guard cells aborted and persistent stomatal initials. Cytoplasmic connections between the guard cells of two adjacent stomata or the guard cell of a stoma and an adjacent epidermal/subsidiary cell, or both types occurring in a species, were noticed. The stomatal development, distribution, diversity and basic stomatal type with reference to systematics are discussed.     

Development of Normal and Abnormal Stomata in some Araliaceae

The present investigation describes the ontogeny of normal andabnormal stomata for three species of the Araliaceae. The maturestomata are mostly anisocytic, occasionally paracytic, and rarelyanomocytic. Several aberrent patterns of stomatal developmenthave been noticed in anisocytic stomata and a few in paracyticones. The ontogeny of anisocytic and paracytic stomata conformsto the syndetocheilic or mesogenous type, while that of anomocyticis haplocheilic or perigenous.     

Structure, distribution and taxonomic importance of foliar stomata in some Indian Amaranthaceae

The structure, distribution and taxonomic importance of foliar stomata in 45 taxa belonging to 19 genera have been studied. In all, six stomatal types have been recognized viz., anomocytic, anisocytic, diacytic, paracytic, hemiparacytic and brachyparacytic. The majority of the taxa are amphistomatic whereas hypostomatic leaves are confined to only three taxa. Stomatal diversity is common but most of the taxa show either dominance or codominance. Stomatal distribution is helpful in distinguishing the three tribes of the Amaranthaceae. The tribe Celosieae shows exclusive presence of anomocytic and anisocytic stomata whereas Amarantheae and Gomphreneae show other stomatal types viz., paracytic and diacytic in addition to anomocytic and anisocytic stomata. Further, the latter two tribes are each distinguishable into two subtribes on the basis of stomata.     

Observations on the cotyledonary and hypocotyledonary stomata and trichomes in some Caesalpiniaceae with a note on their taxonomic

The structure and ontogeny of stomata on cotyledons and hypocotyls and the trichomes on hypocotyl are accounted for in eighteen species of Caesalpiniaceae. Trichomes are eglandular, bi- rarely tri-celled, smooth walled or walls wavy with cuticular striations or tubercles. Anomocytic, haplocytic, paracytic, diacytic, transitional, tetracytic, tricytic and cyclocytic stomata occur in different combinations even on the same surface of the cotyledon. In all, there are fourteen combinations. Inspite of the diversity, the most frequent type is anomocytic in most of the species and paracytic in some species of Cassia and Delonix (abaxial) but rarely it is haplocytic or anisocytic. In hypocotyls it is anomocytic. Ontogenetically anomocytic, tetracytic and cyclocytic stomata are perigenous, whereas other types are mesogenous or mesoperigenous. There is an increase in the number of subsidiary cells by their division or the neighbouring perigenes assuming their shapes. About eight such types are described. A pair of stomata has a common subsidiary cell. Twelve types of guard cell and stomatal approximation abnormalities are described. A range in the number, size and shape of the nuclei in guard cell are recorded. Megastomata (giant stomata) are observed in Parkinsonia and Tamarindus. The taxonomic significance of the stomata is also discussed.     

Stomatogenesis in the genus Hibiscus L. (Malvaceae)

In Hibiscus , stomata are anisocytic, anomocytic, paracytic and tetracytic, the first type being the most frequent and occurring in all plant parts in the ten species studied, whereas the others are scarce and have a limited organographic distribution. The stem, petiole, pedicel, staminal tube, ovary and style are stomatiferous; the leaf-blade, stipule, bracteole and sepals are amphistomatic and petals hypostomatic in the species investigated.
The stomatal types are often homoplastic, the anisocytic being either mesogenous trilabrate or mesoperigenous dolabrate, the anomocytic, mesoperigenous dolabrate or mesogenous trilabrate, and the tetracytic, mesoperigenous dolabrate or mesoperigenous trilabrate. But the typical paracytic stomata (with the subsidiaries completely enclosing the poles) are constantly mesogenous dolabrate and therefore probably indicate mesogenous dolabrate development. Although several patterns of stomatogenesis are encountered in any specific organ, only one of them is found to be dominant. A new subcategory of stomatal ontogeny, mesoperigenous trilabrate, is proposed in Hibiscus. No significant stomatal variation involving reduction in the divisive capacity of the meristemoid has been observed from the vegetative to floral parts; and stomata functioning as hydathodes have not been noticed in the latter, thus indicating that florogenic factors have no effect on the stomata.     

Studies on the structural and developmental variation and distribution of stomata in the Rubiaceae

The structure and ontogeny of the stomata has been studied in 26 species of Rubiaceae, in relation to their organographic distribution. The stomata are mostly paracytic on the leaves, but vary on other parts, particularly on the floral disk, where they are exclusively anomocytic. The foliar stomata are mesogenous and paracytic, but may be either dolabrate or trilabrate in origin. The disk stomata are perigenous and are probably derived from the mesogenous paracytic ones by the loss of divisive potential of their meristemoid due to anthogenetic factors. They are regarded to be hydathodal in function. If the dendroid habit is regarded as primitive, in the family, trilabrate stomata were quite possibly derived from dolabrate ones, which are mostly associated with the dendroid habit. Evidence from stomatal ontogeny is shown to be taxonomically important in the Rubiaceae.     

Epidermal structure and development of stomata in vegetative and floral organs ofHybanthus enneaspermus (Linn.)F. Muell

The present paper deals with the epidermal structure and ontogeny of stomata in vegetative and floral organs ofHybanthus enneaspermus. The epidermal cells are either polygonal or elongated with straight, sinuous or arched thick anticlinal walls. The surface of the cuticle shows parallel striations radiating from the guard cells or hair bases. Unicellular and uniseriate bicellular trichomes with verrucose margins have been observed on all organs. The mature stomata are anisocytic, paracytic, anomocytic and transitional between anisocytic and paracytic. The ontogeny of anisocytic and paracytic stomata is syndetocheilic or mesogenous, anomocytic is haplocheilic or perigenous, while that of the transitional type is mesoperigenous. Four types of stomata have been observed on all the vegetative and floral organs and their ontogeny from organ to organ of this plant is constant. Stoma with a single guard cell is the result of disintegration of one of the guard cells before or after pore formation. Contiguous stomata are also occasionally noticed.     

Structure and Development of Stomata in Vegetative and Floral Organs of Three Species of Kalanchoe

The structure and ontogeny of stomata in vegetative and floralorgans of three species of Kalanchoe is described. The matureanisocytic stomata are mono-cyclic or completely or incompletelyamphicyclic, rarely paracytic, transitional between paracyticand anisocytic and with a single subsidiary cell. The developmentof all the types is syndetocheilic or mesogenous from organto organ but the mature stomatal apparatus varies from organto organ as regards the number and arrangement of subsidiarycells. Abnormal stoma with a single guard cell and arresteddevelopment were observed on all organs. An abnormal stoma witha single guard cell develops directly from the meristemoid.     

A new stomatal type inChenopodiaceae

A new stomatal type—paracytic mesoperigenous—which has not been separated from the paracytic mesogenous type in previous studies of theChenopodiaceae, is described. The frequent occurrence of this previously unknown paracytic mesoperigenous type in this family is demonstrated. As a result a new phylogenetic pathway between anisocytic mesoperigenous and paracytic mesogenous types may be drawn.     

Structure, distribution and taxonomic importance of stomata in some Indian Tephrosia Pers.(Fabaceae)

SUBBA RAO, J.V.& SHANMUKHA RAO, S. R.(1994).Structure, distribution and taxonomic importance of stomata in some Indian Tephrosia Pers.(Fabaceae).Structure, distribution and taxonomic importance of stomata in all the vegetative parts of 18 taxa of Indian Tephrosia including five species endemic to south India are described. The stomata are anisocytic, anomocytic or paracytic. In addition, brachyparacytic stomata have been recorded for the first time in this taxon. This is the first attempt to assess subgeneric treatment in the light of stomatal characteristics and it suggests certain realignments at the infrageneric level in Indian Tephrosia.     

Epidermal Structure and Stomatal Ontogeny in Some Polygonales and Centrospermae

The epidermal structure and ontogeny of stomata in 19 speciesof Centrospermae and two of Polygonales are described. The cellsof the epidermis are polygonal, isodiametric, or elongated invarious directions and arranged irregularly. The anticlinalepidermal walls are thick, sinuous, straight, or arched. Eleventypes of glandular and eglandular trichomes have been observed.Six types of stomata: anomocytic, paracytic, stomata with asingle subsidiary cell, diacytic, anisocytic, and transitionalbetween diacytic and paracytic, have been noticed in the speciesinvestigated. The ontogeny of anomocytic stomata is haplocheilicor perigenous, while that of the other five types is syndetocheilicor mesogenous. Abnormal stomata with a single guard cell, unequalguard cells, aborted guard cells, and arrested development arecommon. Groups of stomata are also frequent but contiguous stomataare rather rare.     

Structure,Delimitation,Nomenclature and Classification of Stomata

The paper reviews stomatal types observed in 500 species of angiosperms besides those described in the literature and deals with the problems of their structure, delimitation, nomenclature and classification. In view of the varied definitions available in the literature for subsidiaries, stomatal types and,the definition and delimitations being variously interpreted by different workers, a modified definition for the subsidiaries and stomata is presented. In accordance with the international code of nomenclature for plants, the names of the stomata widely in use are retained (rule of priority). They have been presently classified as pericytic, desmocytic, paracytic, diacytic, anisocytic, anisotricytic, isotricytic, tetracytic,staurocytic, anomocytic, cyclocytic and a good number of varieties under each type are presented. These stomatal types are recognised on the basis of their structure rather than its ontogenetic pathways.     

Terminology and classification of stomata and stomatal development—a critical survey

The literature on terminology of stomata and stomatal development is reviewed and the terminology rationalized. The classification of developmental types and of the developing cells should not be combined with the morphological classification of mature stomatal complexes. The cells involved in the development should be distinguished on the basis of their origin and position in the developing stomatal complex, and not on the basis of their future form and appearance. It is unsound to distinguish any kind of cell only on the basis of a presumed future division by which it is replaced by its two daughter cells. Development of stomata begins with the formation of a stomatal meristemoid by an unequal division of a protodermal cell. A meristemoid may divide unequally to produce a new meristemoid and a mesogene cell. Stomatal meristemoids eventually function as guard-cell mother-cells. The adjective perigene is restricted to those cells that have arisen by divisions of protodermal cells surrounding the future stoma. The undivided cells surrounding protodermal cells should be termed agene cells, and not neighbouring cells, a term which should be restricted to morphological terminology.     

Structure, Delimitation, Nomenclature and Classification of Stomata

The paper reviews stomatal types observed in 500 species of angiosperms besides those described in the literature and deals with the problems of their structure, delimitation, nomenclature and classification. In view of the varied definitions available in the literature for subsidiaries, stomatal types and, the definition and delimitations being variously interpreted by different workers, a modified definition for the subsidiaries and stomata is presented. In accordance with the international code of nomenclature for plants, the names of the stomata widely in use are retained (rule of priority). They have been presently classified as pericytic, desmocytic, paracytic, diacytic, anisocytic, anisotricytic, isotricytic, tetracytic, staurocytic, anomocytic, cyclocytic and a good number of varieties under each type are presented. These stomatal types are recognised on the basis of their structure rather than its ontogenetic pathways.