Acta Botanica Sinica
植 物 学 报
2004, 46 (2): 242－252
http://www.chineseplantscience.com
Structure, Delimitation, Nomenclature and Classification of Stomata
Malvey PRABHAKAR
(Plant Anatomy and Taxonomy Laboratory, Department of Botany, Osmania University, Hyderabad-500 007, India)
Abstract: 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.
Key words: classification; definition; plant stomata
The early history of stomatal nomenclature dates back
to Prantle (1872; 1881). Subsequently several classificatory
systems have come into existence, which are mostly based
on ontogeny ( Florine, 1931; 1933; Pant, 1965; Payne, 1970;
1979), or combination of structure and ontogeny (Paliwal,
1969b; Fryns-Claessens and van Cotthem, 1973; Stevens
and Martin, 1978). Though several classificatory systems
and review works (Vesque, 1889; Francey, 1936; Metcalfe
and Chalk, 1950; Metcalfe, 1961; Stace, 1965; van Cotthem,
1970b; Dilcher, 1974; Patel, 1979; Rasmussen, 1981; Kidwai,
1981; Inamdar et al., 1986; Baranova, 1987; 1992) have been
published, we lack precise definition for subsidiaries and
stomatal types.
Francey (1936) recognised eight categories and 34 structural stomatal types based on the number, position and
size of the subsidiaries. Though his classificatory system
was fairly good but it was not used by subsequent workers.
Survey of the literature reveals that the most widely accepted and used stomatal classificatory system based on
the mature stomatal structure is that of Metcalfe and Chalk’
s
(1950), Metcalfe (1961) and Stace (1965). While using their
classificatory systems some strictly followed the definition
of the subsidiaries and delimit the stomata, while others do
not (Prabhakar and Leelavathi, 1992 ). For example, though
the stomata of Lathyrus (Simola, 1968), which may normally be considered as paracytic on the basis of orientation of the two surrounding cells in relation to the guard
cells, following strictly the definition of Metcalfe and Chalk
(1950), they have been classified under anomocytic by
Simola (1968), because of the indistinct nature of the abutting cells. However, Stace (1965) has named such stomata
Received 10 Oct. 2002
Accepted 7 Jul. 2003
as paracytic. Metcalfe and Chalk (1950) also did not seem
consistent in applying their definition of subsidiaries
throughout their book. For example the stoma represented
in Figs.1788, 2220, 3098 (Metcalfe and Chalk, 1950) are
surrounded by two parallel indistinct cells and hence should
be assigned to anomocytic stomata sen. str. Metcalfe and
Chalk (1950). The authors however described them as
paracytic stomata. While others classified such stomata
having four or more abutting cells into paracytic stomata
(Paliwal, 1961; Inamdar, 1968; 1970; Fryns-Claessens and
van Cotthem, 1973; Wilkinson, 1979; Kidwai, 1981) and a
stoma having only two abutting cells/subsidiaries parallel
to the guard cells have been designated as hemiparacytic,
parallelocytic Eupara-twi-monocyclic, obliparatwimonocyclic, duplo-para-tetra-bicyclic, para-polycyclic,
para-spirocyclic, etc. (Patel, 1979; Wilkinson, 1979). The
classificatory systems provided subsequent to Metcalfe
and Chalk (1950), Metcalfe (1961) and Stace (1965), I do not
feel at all an improvement over the concept of Metcalfe and
Chalk (1950). In these new classifications several new terms
which the authors themselves were not able to use them in
strict sense in their subsequent publication for describing
the stomata have been introduced.
The draw back of all these classificatory systems including that of Metcalfe and Chalk (1950 ) mainly lies in the
definitions of subsidiaries and partly the definitions of stomatal types. Further several researcher have provided different definitions for a single type of stomata. Some of these
definitions for paracytic stomata are listed below:
Vesque (1889): “Stomata with two subsidiary cells parallel to the long axis of the pore-Rubiaceous”(*Figs.1－10).
Malvey PRABHAKAR: Structure, Delimitation, Nomenclature and Classification of Stomata
Solereder (1908): “The pair of the guard cells are accompanied on either sides (i.e. to the right and left) by one or
more subsidiary cells, which are placed parallel to the poreRubiaceous type”(*Figs.1－5, 7, 9, 10, 13 －16).
Francey (1936): Stoma surrounded by two parallely oriented cells which may be equal or unequal, designated as
types 3 and 4 respectively (*Figs.1－3, 11, 12 ).
２４３
Metcalfe and Chalk (1950): “Stoma accompanied on either sides by one or more subsidiary cells (cells distinct
from other epidermal cells) parallel to the long axis of the
pore and guard cells-Paracytic type”(*Figs.1－5, 7, 9, 10,
13－16).
Stace (1965): “Stoma with two subsidiary cells lying parallel to the guard cells-Paracytic”(Figs.1－10).
Figs. 1－62. 1－24. Diagrammatic representation of the so called paracytic stomata. 25－36. Stomata surrounded by distinct or indistinct
subsidiaries. 25, 26. Stomata with distinct subsidiaries. 27, 28. Stomata with indistinct subsidiaries. 29－33. Subsidiaries dicyclic. 34, 35.
Stomata with distinct crystalliferous subsidiaries. 36. Stomata with one distinct and the other indistinct subsidiary. 37－62. Pericytic
stomata. 37, 38. Stomata with distinct or indistinct subsidiary. 39, 40. Subsidiary transversly oriented to guard cells. 41, 42. Subsidiary
parallel to guard cells. 43, 44. Subsidiary oblique to guard cells. 45－51. Subsidiaries 1 1/2 cyclic. 52－59. Subsidiaries dicyclic. 60－62.
Subsidiaries tricyclic.
*, figures cited here fit into the definition given by the respective authors.
２４４
Cronquist (1968): “Stoma with two equal subsidiary cells
flanking the guard cells-Paracytic stomata”(*Fig.1).
Fahn (1969): “Each guard cell is accompanied by one or
more subsidiary cells, the longitudinal axis of which are
parallel to that of the guard cells and aperture-Rubiaceous
or Paracytic type”(*Figs.1－5, 7, 9, 10, 13－16).
Paliwal (1969a): “Two or more subsidiary cells flank the
stomata parallel with long axis of the guard cells-Paracytic
stomata”(*Figs.1－5, 13, 14, 16 ).
Rajagopal (1973): “Abutting subsidiaries two, parallel
to the guard cells, distinct or indistinct, with or without
conjoint walls towards one or both poles-a Diacytic type”
(*Figs.1－5, 7, 9－12, 15, 18－21 ).
Note: When the two subsidiaries are without conjoint
walls towards one or both poles of the guard cells, it is
bound to leave a space which will be abutted by one or
more cells as also illustrated by the same author (Figs.18－
21).
Dilcher (1974): “One or two cells adjacent to the guard
cells with their long axis parallel to the long axis of the
guard cells-Paracytic type”(*Figs.1－4, 7, 9－12, 15, 17－
22).
Cutter (1978): “One or more (often two) subsidiary cells
are present, with their long axis parallel to the guard cellsParacytic”(*Figs.1－10, 13－17).
Esau (1979): “One or more subsidiary cells border the
stoma parallel with the long axis of the guard cells- Paracytic
type”(*Figs.1－10, 13－17 ).
Wilkinson (1979): “Stoma accompanied on either side
by one or more subsidiary cells parallel to the long axis of
the pore and guard cells. The subsidiaries may or may not
meet over the poles and may or may not laterally elongateParacytic type”(*Figs.1－5, 7, 9, 10, 13－16, 19, 20, 23, 24).
Following strictly the above definitions the stomata represented in Figs.1－24 can be accommodated under the so
called paracytic ones.
A New Classification and Modified Definitions
It is clear from the above introduction that the definitions provided by various workers for different stomatal
types, are ambiguous. In order to remove the ambiguity the
author felt the necessity to redefine the subsidiaries and
stomatal types which is presented below based on his observations on more than 500 species of angiosperms (for
details of the species c.f. Anna Mani & Prabhakar, 1991a;
1991b; 1993; 1994a; 1994b; 1995; Anna Mani et al., 1993;
Bhatia et al., 1986; 1988; Ferzana et al., 1991; 1994a; 1994b;
Jelani and Prabhakar, 1991; 1992; 1993; Jelani et al., 1990;
1991; 1993; Koteswar Rao et al., 1988; Leelavathi and
Ramayya, 1975; Leelavathi et al., 1980; 1981; 1984a; 1984b;
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1985; 1988a; 1988b; Padma Rao et al., 1988; Prabhakar and
Anna Mani, 1995; 1996; Prabhakar and Leelavathi, 1989;
1991;1992; Prabhakar and Ramayya, 1975; 1979; Prabhakar
et al., 1984; 1985;1986; 1988a; 1988b; 1988c; 1990; Ramayya
and Prabhakar, 1973; 1975; Ramayya et al., 1983; Rao et al.,
1991; 1992a; 1992b; 1992c, 1993a; 1995; Verma et al., 1989;
1991; 1992; Vijay Kumar et al., 1986) and also the information available in the literature.
Stomatal pore: An opening in the epidermis surrounded
by a pair or more guard cells.
Stoma: Stomatal pore and a pair of guard cells.
Stomatal complex: Stoma surrounded by subsidiaries
(stomatal types).
Subsidiary cells*: Cells surrounding a pair of guard cells
in one (Fig. 25) or more cycles (Fig.26); subsidiaries of the
cycle abutting on stoma may (Figs.25, 26, 34－36) or may
not be distinct from the adjacent epidermal cells ( Figs. 27,
28 ), but of the other cycle(s) when present are morphologically distinct (Fig. 26). The subsidiaries of the outer cycle
may (Fig. 26) or may not coincide with those of the abutting
cycle in number, arrangement and orientation, etc. (Figs.
29－33). The term subsidiary is restricted to mature stomata
only. The subsidiaries of the abutting as well as other cycle
(s) may be perigynous or mesogenous in origin and can be
referred as perigene and mesogen cells (while describing
the ontogeny) depending upon their origin. A subsidiary
derived from stomatal meristemoid need not be distinct at
maturity from other epidermal cells. Similarly a subsidiary
not derived from stomatal meristemoid ( i.e. perigene ) may
be distinct from other epidermal cells or other subsidiary
cells.
Pericytic stomata: Stoma (pair of guard cells) surrounded
*, cells abutting the stoma has been designated as subsidiaries/auxillary
cells when they are distinct from other epidermal cells and as annexes/surrounding cells/accessory cells/contact cells/ordinary cells
when they are indistinct (Francey, 1936; Eames and Mc Daniels,
1947; Metcalfe and Chalk, 1950; Mc Lean and Cook, 1951; FrynsClaessens and van Cotthem, 1973; Tomlinson, 1974; Patel, 1978;
Cutter, 1978; Rasmussen, 1981). However the criteria to judge the
distinctness has been mostly left to the subjective judgement. In
retrospect the criteria used for distinctness by various workers are
morphological (shape, size, anticlinal walls and surface), cytological
(density of cytoplasm, size of the nucleus, presence or absence of
plastids and ergastic substances) (Figs.34－36), number, position,
orientation, physiological and ontogenetical (Patel, 1978; Stevens
and Martin, 1978). Tracing cytological, physiological and
ontogenetical characters in the herbarium and fossil materials is
impracticable. All the cells abutting the guard cells are distinct from
the other epidermal cells by virtue of their position (i.e. abutting
nature to the guard cells) hence prefer to call them subsidiaries.
Malvey PRABHAKAR: Structure, Delimitation, Nomenclature and Classification of Stomata
by a single distinct or indistinct subsidiary cell (Figs. 37－
44). The subsidiary cell may be variously oriented to the
guard cell and may be monocyclic ( Figs.37－44 ), 11/2 cyclic ( Figs.45－51 ), di-, tri- to polycyclic ( Figs. 52－62 ).
Synonyms: Adetostomy, amphipericytic, co-pericytic,
２４５
coperi-desmo-twi-bicyclic, coperi-polohemibicyclic, copoloperi-hemibicyclic, duplopericytic, double floating,
duploperi-twi-bicyclic, floting, pericytic, pyrosia type, perihaplo-monocyclic, Ringzellen-type, stomata libera,
unizyklisch (Prantle, 1881; Maroti, 1961; Mickel, 1962;
Figs. 63－122. 63－90. Desmocytic stomata. 63. Stomata with distinct subsidiary. 64. Stomata with indistinct subsidiary. 65, 66.
Subsidiary transversely oriented to guard cells. 67, 68. Subsidiary parallelly oriented to guard cells. 69, 70. Subsidiary obliquely oriented
to guard cells. 71－78. Subsidiaries 1 1/2 cyclic. 79－89. Subsidiaries dicyclic. 90. Subsidiaries tricyclic. 91－122. Paracytic stomata. 91.
Stomata with distinct subsidiaries. 92. Stomata with indistinct subsidiaries. 93－96. Subsidiaries parallel to the guard cells. 97－101.
Subsidiaries transverse to guard cells. 102－103. Subsidiaries oblique to guard cells. 104－112. Subsidiaries 1 1/2 cyclic. 113－120.
Subsidiaries dicyclic. 121. Subsidiaries hemitricyclic. 122. Subsidiaries tricyclic.
２４６
Mickel and Lersten, 1967; van Cotthem, 1970a, 1970b;
Probst, 1971; Dilcher, 1974; Patel, 1979).
Desmocytic: A stoma completely surrounded by a single
distinct or indistinct cell with a conjoint wall. The subsidiary cell may be variously oriented to the guard cells and
the conjoint wall may be variable in position. The subsidiaries may be monocyclic ( Figs.63－70 ), 1 1/2 cyclic, di-, trito polycyclic (Figs.71－90).
Synonyms: Codesmoperi-twi-bicyclic, codesmo-polohemibicyclic, copolo-desmo-hemibicyclic, desmocytic,
desmo-haplo-monocyclic, duplodesmo-twi-bicyclic, hanging or suspended stomates, stomata suspensa (Mickel and
Lersten, 1967; van Cotthem, 1970a; 1970b; Dilcher, 1974;
Patel, 1979; Prantle, 1881) or indistinct (Figs. 91－96 ), equal
or unequal (Figs. 91－96 ), parallel (Figs. 91－96 ), transverse
(Figs. 97－101 ) or obliquely oriented (Figs. 102,103 ) to the
guard cells, but their conjoint walls are towards the poles
of the guard cells. The subsidiaries may be mono- to polycyclic (Figs. 91－122 ).
Note: Francey (1936) recognised two types of stomata
under this category based on the size of subsidiaries. Type
1 is represented by more or less equal sized subsidiaries
(Figs.91, 92). Type 2 by unequal sized subsidiaries (Figs.
91,92). These can be now recognised as varieties under the
type paracytic only. Similarly the stoma having distinct or
indistinct subsidiaries, parallel oblique or transversely oriented subsidiaries can be recognized into different varieties.
Synonyms: Allelocytic, amphiparacytic, bicyctic
paracytic, duplo-para-tetra-bicyclic, eupara-twimonocyclic,
laterocytic oblipara-twimonocyclic, paracytic, parallelocytic,
para-polycyclic, paraspirocyclic, rubiaceous (Vesque, 1889;
Metcalfe and Chalk, 1950; Payne, 1970; Guyot, 1971; Dilcher,
1974; Hortog and Bass, 1978; Patel, 1979).
Diacytic stomata: A stoma completely surrounding by
only two distinct or indistinct subsidiaries (Figs.123, 124),
equal or unequal (Figs.123－126), parallel (Figs. 127, 128),
transverse (Figs. 129, 130) or obliquely oriented (Figs. 131,
132 ) to the guard cells, but the conjoint wall of the abutting
subsidiaries are lateral to the guard cells. The subsidiaries
may be monocyclic to polycyclic ( Figs. 123－147 ).
Note: Francey (1936) recognised two types of stomata
under this category also, based on equal ( Figs. 123, 124 ) or
unequal sized subsidiaries (Figs.125, 126). These can be
recognised as varieties under the diacytic stomata only.
Similarly a stoma having distinct or indistinct subsidiaries,
transversely, obliquely or parallelly oriented subsidiaries
can be recognised into different varieties.
Synonyms: Amphidiacytic, axillocytic, bicyclic diacytic,
caryophyllaceous, coaxilocytic, coparirtocytic, copolocytic,
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diacytic, diallelocytic, diapolocyclic, diapolycyclic, diaspirocyclic, diatwi-monocyclic, duplodia-tetra-bicyclic,
labiateous, oblidia-monocyclic, perietocytic, polocytic,
polohaplo-acyclic, polycytic, pteris type, stomata applicata,
stomata adnata (Prantle, 1881; Vesque, 1889; Giesenhagen,
1901; Metcalfe and Chalk, 1950; Payne, 1970; van Cotthem,
1970a; 1970b; Guyot, 1971; Probst, 1971; Dilcher, 1974; Patel,
1979).
Anisocytic stomata: A stoma completely is surrounded
by only three subsidiaries, variable in position and shape,
but one of the subsidiary is distinctly small (Figs.148－156).
The subsidiaries may be mono- to polycyclic (Figs.148－
164).
Synonyms: Amphianisocytic, anisocytic, aniso-trimonocyclic, axillocytic, coaxillocytic, cruciferous,
helicocytic, helico-spirocyclic obli-haplo-acyclic (Vesque,
1889; Metcalfe and Chalk, 1950; Payne, 1970; Dilcher, 1974;
Patel, 1979).
Anisotricytic stomata: A stoma completely surrounded
by only three subsidiaries, variable in position and shape,
but one of the subsidiary is distinctly large. The subsidiaries may be mono- to polycyclic (Figs.165－170).
Synonyms: Anisocytic, anomocytic, helicocytic,
diacytic ( Metcalfe and Chalk, 1950; Stace, 1965; Wilkinson,
1979 ).
Isotricytic stomata: A stoma completely surrounded by
only three subsideries, variable in position and shape,but
the three subsidiaries are more or less of equal size. The
subsidiaries may be mono- to polycyclic ( Figs.171－174 ).
Synonyms: Anomocytic (Metclafe and Chalk, 1950;
Stace, 1965).
Note: Francey (1936) recognised seven types under the
above three categories (i.e. stoma surrounded by three
cells), based on the size and position of the subsidiary
cells. The definitions of some of these types and figures
represented are overlapping. Hence presently they are divided into three categories of stomata which are structurally distinct from one another, i.e. anisocytic, isotricytic
and anisotricytic. Under each of these categories a good
number of varieties can be recognised based on the shape,
size, structure of inner and outer cycles of the subsidiaries.
Tetracytic stomata: Stomata completely surrounded by
only four subsidiaries, of variable size and shape, of which
two are polar and two lateral in position. The subsidiaries
may be mono- to polycyclic (Figs.175－186).
Synonyms: Anphibrachiparatetracytic, amphiparatetracytic, angiopteris type, bicytic paracytic, brachyparahexacytic, brachyparatetracytic, cyclocytic, duplo-para-tetraacyclic, grass type, hemidiahaplo-acyclic, hemiparacytic,
Malvey PRABHAKAR: Structure, Delimitation, Nomenclature and Classification of Stomata
hemipara-haplo-acyclic, hexacytic, hexa-a-hemibicyclic,
hexa-bhemibicyclic, obli-tri-acyclic, obli-twi-acyclictetracytic, palm type, paracytic, parahexacytic,
paratetracytic, para-twi-acyclic, quadricytic, rubaceous,
staurocytic, stomata quadricellulaires, tetra cyclocytic,
tetracytic, tetramonocyclic, tricyclic (Vesque, 1889; Metcalfe
and Chalk, 1950; Maroti, 1958; Prat, 1960; Metcalfe, 1961;
２４７
Stace, 1965; van Cotthem, 1970b; Guyot, 1971; Dilcher, 1974;
Tomlinson, 1974; Patel, 1979; Wilkinson, 1979).
Staurocytic stomata: Stoma completely surrounded by
only four subsidiaries, variable in shape and size but two of
their conjoint walls polar, while the other two are lateral to
guard cells. The subsidiaries may be mono to polycyclic
(Figs.187－194 ).
Figs. 123－174. 123－147. Diacytic stomata. 123, 124. Stomata with distinct and indistinct subsidiaries respectively. 125, 126. Stomata
with unequal subsidiaries. 127, 128. Subsidiaries parallel to guard cells. 129, 130. Subsidiaries transverse to guard cells. 131, 132.
Subsidiaries oblique to guard cells. 133, 134, 140－142, 145, 146. Subsidiaries 1 1/2 cyclic. 135－139, 143, 144, 147. Subsidiaries dicyclic.
148－164. Anisocytic stomata. 148. Stomata with indistinct subsidiaries. 149, 150. Stomata with distinct subsidiaries. 151－156. Stomata
with one or two distinct subsidiaries. 157. Subsidiaries 1 1/2 cyclic. 158－164. Subsidiaries dicyclic. 165－170. Anisotricytic stomata.
165. Stomata with indistinct subsidiaries. 166, 167. Stomata with distinct subsidiaries. 168, 169. Stomata with one or two distinct
subsidiaries. 170. Subsidiaries 1 1/2 cyclic. 171－174. Isotricytic stomata. 171, 172. Stomata with indistinct and distinct subsidiaries
respectively. 173, 174. Stomata with one or two distinct subsidiaries.
２４８
Synonyms: Type 15, stauro-cyclocytic, staurocytic,
stauro-tetra-monocylic (Francey, 1936; van Cotthem, 1970a;
1970b; Dilcher, 1974; Patel, 1979; Wilkinson, 1979 ).
Anomocytic stomata: Stoma surrounded by four or more
subsidiaries, variable in position, shape and size ( other
than tetracytic and staurocytic types; Figs.195－206 ).
Synonyms: Actinocytic, amphibrachyparacytic,
anomocytic-polycytic, brachyparacytic, hemiparacytic,
latero-twi-acyclic, oblitwi-acyclic, paracytic, stephanocytic,
termino-twi-acyclic (Dilcher, 1974; Patel, 1979; Wilkinson,
1979; Baranova, 1987).
Cyclocytic stomata: Stoma is surrounded by four or more
subsidiaries, variable in position, shape and size but arranged in a narrow ring ( Figs.207－212 ).
Ａｃｔａ　Ｂｏｔａｎｉｃａ　Ｓｉｎｉｃａ　植物学报　Ｖｏｌ．４６　Ｎｏ．２　２００４
Note: A desmocytic (Fig. 37), pericytic (Fig. 63), paracytic
(Fig. 91), diacytic (Fig.123 ), anisocytic (Fig. 149 ), isotricytic
(Fig.172 ), anisotricytic ( Fig.167 ), staurocytic ( Fig.187)
and tetracytic stoma (Fig.175), having one to four subsidiaries arranged in a narrow ring are not included under
cyclocytic stomata.
Synonyms: Amphicyclocytic, angiopteris type,
cyclocytic, cyclo-multi-monocyclic, encyclocytic, hexamonocytic, multi-bicyclic, multi-polycyclic, polycytic
(Shtromberg, 1956; Stace, 1965; Probst, 1971; Dilcher, 1974;
Patel, 1979).
Actinocytic, stephanocytic, actino-multi-monocyclic: I
feel that identification of actinocytic stomata ( so also with
other types like stephanocytic ) is a difficult problem
Figs. 175－212. 175－186. Tetracytic stomata. 175, 176. Stomata with distinct or indistinct subsidiaries respectively. 177－181.
Stomata with one to three distinct subsidiaries. 182, 183. Subsidiaries 1 1/2 cyclic. 184－186. Subsidiaries dicyclic, hemitricyclic and
tricyclic respectively. 187－194. Staurocytic stomata. 187, 188. Stomata with distinct and indistinct subsidiaries respectively. 189－191.
Stomata with one to three small subsidiary cells. 192－194. Stomata with one to three large subsidiary cells. 195－206. Anomocytic
stomata. 195, 196, 202, 203. Stomata with one, three or more radiating subsidiaries. 197, 199. Stomata with one distinct parallel
subsidiary. 200. Stomata with two distinct parallel subsidiaries. 201. Subsidiaries hemidicyclic. 204. Subsidiaries parallel to guard cells.
205. Subsidiaries transverse to guard cells. 206. Some subsidiaries parallel to guard cells while other radiating. 207－212. Cyclocytic
stomata. 207－211. Subsidiaries monocyclic. 212. Subsidiaries dicyclic.
Malvey PRABHAKAR: Structure, Delimitation, Nomenclature and Classification of Stomata
because in a number of angiosperms taxa I have come across
stomata in which some of the subsidiaries are arranged in a
radiating fashion while others are otherwise (Figs.195,196,
202－206). Further in a number of publications the stomata
with radiating subsidiaries are also described as anomocytic
stomata. For example, Fig. 10.3a represented by Wilkinson
(1979) is more like an actinocytic stomata by definition but
has designated as anomocytic stomata, and Fig.6a by
Baranova (1987) is an anomocytic type but has been described as stephanocytic one. In order to avoid these confusions the present author prefer to categorise them as
anomocytic stomata only and do not recognise actinocytic,
stephanocytic stomata, etc.
Cycles: Depending upon the cycles of subsidiaries the
stoma may be described as monocyclic (Figs.53, 54, 63, 64),
1 1/2 cyclic ( hemidicyclic; Fig.77), dicyclic (Figs.55, 78 －
83), hemitricyclic (Figs.185), tricyclic and so on. However,
the above definitions of stomata are not applicable to the
abnormal stomata viz., stoma having more (Fig.87) or less
than a pair of guard cells (Fig.88), abutting stoma (Figs.89,
90), etc.
２４９
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