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ABSTRACT
STEBBINS, G. L., and G. S. KHUSH. (U. California, Davis.) Variation in the organization of
the stomatal complex in the leaf epidermis of monocotyledons and its bearing on their phylogeny.
Amer. Jour. Bot. 48 (1): 51-59. Illus, 1961.-Using macerated pieces of epidermal tissue from
living plants and herbarium specimens, stomatal complexes of 192 species belonging to 49 families
of monocotyledons were studied. Four categories are recognized, 2 with 4 or more subsidiary cells
surrounding the guard cells, I with 2 subsidiaries, and I with none. Development of the 2-subsidi·
ary type, studied in acetocarmine preparations of Juncus elfusus and Sagittaria montevidensis,
resembles that in Gramineae previously described. No correlation was found between type of
stomatal complex and either leaf shape or type of xylem vessel, but some correlation exists between
this character and type of seed germination, vascular anatomy of seedling, growth habit of mature
plant, and geographic distribution. Types with 4 or more subsidiaries are most commonly phanero-
phytes with tropical distribution, many vascular bundles in the cotyledon, and hypogeal germina-
tion. Complexes with 2 subsidiaries occur in many families of a diverse nature, but the most
primitive plants with this type are hydrophytes or helophytes with tropical or temperate distribu-
tion, I vascular bundle in the cotyledon, and epigeal germination. Stomatal complexes without
subsidiaries are almost confined to the Liliales and their more specialized derivatives. These
plants are predominantly geophytes with temperate or tropical distribution, 2 bundles in the
cotyledon, and epigeal germination. Reasons are advanced for suggesting that the type with many
subsidiaries is the most primitive and the other 2 types have been derived from it independently
by reduction of the number of subsidiary cells.
ALTHOUGH most systems of classification of Constance (1955) of these and other studies has
angiosperms emphasize chiefly floral morphology, justly emphasized the importance of basing hypo.
information from other botanical disciplines is be- theses about phylogeny on evidence derived from
coming increasingly useful. Thus, pioneer works of as many characteristics as possible.
Bailey and his associates on the vascular anatomy The stomatal cell complex in angiosperms was
of dicotyledons and of Cheadle and his students on first studied by Strasburger (1866) and particularly
the vascular anatomy of monocotyledons have made by Vesque (1889) who recognized 4 broad cate-
significant contributions to our understanding of gories based on the presence and arrangement of
the phylogeny of angiosperms. The review by accessory cells as well as their mode of develop.
ment. Vesque, however, studied only dicotyledons.
, Received for publication April 18, 1960. More recently Florin (1931) has made a detailed
This research was supported by National Science Foun- study of stomatal complexes in gymnosperms, in
dation Grant #G3737. We are greatly indebted to Drs.
Herbert L. Mason, Curator of the Herbarium, University of which he presented some data on development,
California, Berkeley, and John L. Tucker of the Herbarium while Dehnel (1957) has amplified our knowledge
at Davis for the loan of the specimens and for permission about the dicotyledons, presenting a full review of
to remove small bits of leaf tissue. We are equally in- the literature to date. There exists, however, no
debted to Drs. Herbert Baker, Director of the Botanical comparable study of stomatal complexes in mono-
Garden, University of California, Berkeley, and R. E. Baker,
director of the Strybing Arboretum, San Francisco, for some cotyledons.
of the living material of unusual species. The use of information about stomatal complexes
52 AMERICAN JOURNAL OF BOTANY [Vol. 48
in studying phylogeny should be particularly en- in green material. Dried material was boiled in
lightening in view of our detailed knowledge re- water for 10-15 min., and, in most cases, with a
garding their development. Our observations are little manipulation the epidermis can be easily
at variance with those of Tognini (1897) who peeled off. IIi some species like Astelia and palms,
states that several modes of development of sto- treatment with 5% NaOH at oven temperature of
mata may occur in different organs of the same 60°F. for 2 hr. was very helpful. Small pieces of
plant. In all the material studied by us, the devel- epidermis separated from the rest of the mesophyll
opmental sequences are precisely the same for a tissue were mounted and stained with acetocarmine.
particular complex. Thus in as different genera as Observations can be made immediately after mount-
[uncus and Sagittaria, which fall into the same ing, but after 10-12 hr. the remains of the nuclei
category as regards their stomatal complex, the stain more clearly. Measurements of guard cells
differentiation of the stomatal complex takes place and epidermal cells were made with an ocular
in similar sequences. Similarly, developmental micrometer. In each case, 10 cells were measured
modes are constant, even as to minute details, from at random and the mean length calculated. In some
organ to organ within the same plant. Thus, in species a few stomatal rows are interspersed with
barley the present authors followed the same de- rows of epidermal cells which lack stomata. Length
velopmental sequences in as different organs as of epidermal cells in such rows is longer than the
coleoptile, leaf and awn of the lemma. length of epidermal cells found between stomatal
Another point which makes the study of the rows. In that case 5 cells from each region were
stomatal complexes a desirable tool in taxonomic measured. In species where leaves are absent as in
research is that genera and even families show great some species of Scirpus, [uncus, and Leptocarpus,
constancy for their possession of a particular com- preparations were made from peels of stems. Cam.
plex, yet there is considerable variability from one era lucida drawings were made 10-12 hr. after
higher taxon to another. Thus at the level where mounting and are reproduced at a magnification
classical methods of cytology and genetics cannot of X240.
be applied, this study of stomatal complexes should For studying development, the meristematic bases
help us to understand true evolutionary relation- of developing leaves were fixed in a mixture of 3
ships of monocotyledons. parts absolute alcohol-l part glacial acetic acid,
MATERIALS AND METHODs.-Stomatal complexes mounted whole in a solution of acetocarmine diluted
were examined in 192 species belonging to 49 of to lh strength with 50% acetic acid, and heated to
the 68 families of monocotyledons recognized by boiling for a short time. The mounts, sealed in a
Hutchinson. In addition, more than 50 genera of paraffin-beeswax mixture and kept in a refrigerator,
Gramineae have been examined by the senior au· lasted for several weeks, and the details of the
thor, either directly or through published figures. epidermal cells could be seen clearly enough by
The arrangement of families is according to Hutch- focussing on the upper epidermis with the oil im-
inson (1934), and his system is followed strictly mersion lens. Camera lucida drawings are repro-
throughout the descriptive section. Representatives duced in fig. 1--4 at a magnification of X 535.
of the other 18 families are either aquatic and thus DESCRIPTION OF RESULTs.-The various types of
lack stomata or are aphyllous and hence unsuitable stomatal complexes in monocotyledons can be di-
for investigation. Investigations were made on vided into 4 broad categories. (1) Two guard cells
either dried material from herbarium specimens or and 4--6 subsidiary cells around the guard cells in
preserved material from living plants. Herbarium all 4 directions (fig. 13, 21, 22); (2) Two guard
specimens were obtained from the herbarium of cells and 4--6 subsidiary cells 2 of which are round.
the University of California, Berkeley, and from ish and smaller than the rest and at the ends of
the Botany Department herbarium, University of guard cells, the rest being lateral to guard cells (fig.
California, Davis. Leaves from living plants were 10, 12, 16, 24); (3) Two guard cells and 2 sub.
fixed in 3: 1 alcohol-acetic acid and stored in 70% sidiary cells lateral to the guard cells. This, the
alcohel. All of the numerous specimens taken from most common type of stomatal complex, is distrib-
Botanical Garden, U. C., Berkeley, and Strybing uted over many families (fig. 5-9, n, 17, 23);
Arboretum, Golden Gate Park, San Francisco, were (4) Two guard cells without any subsidiary cells;
already labelled and the rest were determined with the second most common type of stomatal complex
the help of appropriate manuals. Bailey's (1949) (fig. 18, 19, 20) .
manual was used to determine the identity of the The first type is found in the families Commelina-
cultivated plants. The generic identity of all speci- ceae, Araceae, Bromeliaceae, Musaceae, Strelitzi,
mens was verified by the senior author. A table aceae, Zingiberaceae and Cannaceae (table 3).
has been prepared listing the names and source of Tradescantia, Rhoeo and Zebrina of Commelinaceae
material of all of the 192 species studied, but it have 4 subsidiary cells, 1 on each of the 4 sides of
proved to be too long to include in the present paired guard cells and forming a beautiful square
paper. Mimeographed copies of it will be sent by (Stebbins and Jain, 1960). Commelina, of the same
the senior author upon request. family, has 6, 2 additional being present on the
There is no difficulty in peeling off the epidermis sides (fig. 21). In Araceae, 2 genera, Scindapsis
January, 1961] STEBBINS AND KHUSH-STOMATA IN MONOCOTYLEDONS 53
TABLE 1. Families with no subsidiary cells as the predomi- It would be inappropriate to discuss the results
nant condition. Genera deviating from the modal con- without giving a brief description of the develop-
dition are listed separately under their family, the mental patterns of the complexes discussed above.
number of subsidiaries is given in parentheses, fol-
Bunning and Biegert (1953) followed by Stebbins
lowed by the number of species studied. Orders which
appear in 2 tables are marked with an asterisk (.). and Jain (1960) have investigated thoroughly the
developmental sequences in Allium. This genus is
NUMBER STUDIED a good representative of type 4, which lacks sub-
ORDER FAMILY Genera Species sidiary cells. The differentiation of the guard
LILIALES· Liliaceae 34 40 mother cell from an ordinary epidermal cell occurs
Astelia (2) -5 spp. by an asymmetrical division. Before division, the
TecophiIaceae 3 3 nucleus migrates to the distal end of the meriste-
Trilliaceae 1 1 matic cell. The cytoplasm at the distal end is much
Smilacaceae 1 1 denser than at the proximal end. As the result of
ALSTROEMERIALES Alstroemeriaceae 1 1 asymmetrical division, the smaller cell, which has a
Philesiaceae 1 1
higher density of cytoplasm, divides once again to
ARALES· Lemnaceae 1 1
AMARYLLlDALES Amaryllidaceae 10 10
give rise to 2 paired guard cells. Bunning and Bie-
'Iridaceae 4 4 gert (1953) found that by centrifuging, that is, by
DIOSCOREALES Dioscoreaeceae 2 2 changing the position of nuclei in the cells, differen-
AGAVALES Xanthorrhoeaceae 6 8 tiation could be inhibited. Thus differentiation is
Xanthorrhoea (2) -2 spp. strictly epigenetic depending upon the organization
Agavaceae 11 15 of the cytoplasm and the position of the nucleus
Doryanthes (2+) -1 sp, prior to division.
HAEMODORALES· Apostasiaceae 1 1 Developmental sequences of the third type, with
Taccaceae 1 2 2 subsidiary cells, were studied by Campbell (1881)
BURMANNIALES Burmanniaceae 1 1
in corn, Porterfield (1937) in bamboo, Flint and
ORCHlDALES Orchidaceae 4 4
Moreland (1946) in sugar cane, by Kaufman
(1957) in rice, and recently verified in other tribes
genera of Cyclanthaceae studied have 4 subsidiary of Gramineae by Stebbins and Shah (1961).
cells. The present authors investigated the development
Worth mentioning is Scheuchzeria palustris (fig. of this type of complex in [uncus and Sagittaria.
10) , belonging to the unrelated family Scheuchzeri- Initial differentiation of the guard mother cell is
aceae, which has a similar pattern, with 4 subsidiary as in Allium, except that it occurs at an earlier
cells but the end cells are not very well defined, as stage, when all of cells are in the meristematic con-
in Palmae and Pandanaceae.
The third type is spread consistently over 24 TABLE 2. Families with 2 subsidiary cells as the predomi-
families out of 50 examined (table 2) and is nant condition
present in occasional members of other families,
for example, the genus Astelia (fig. 9) of Liliaceae, NUMBER STUDIED
Xanthorrhoea (fig. 7) of Xanthorrhoeaceae, Dory- ORDER FAMILY Genera Species
anthes (fig. 23) of Aogayaceae and Sararanga of BUTOMALES Bu.omaceae 2 2
Pandanaceae. It was found in all species studied of ALISMATALES· Alismataceae 2 2
the orders Butomales, Alismatales (except Scheuch- JUNCAGINALES J uncaginaceae 1 1
zeria), Iuncales, Graminales, Cyperales, Typhales, Liliaceae 1 1
POTAMOGETONALES Potamogetonaceae 1 1
Haemodorales (except families Apostasiaceae and
COMMELINALES· Flagellariaceae 2 2
Taccaceae}, Juncaginales, Potamogetonales, Apona- Mayacaceae 1 1
getonales, Eriocaulales, Xyridales, familes Flagel- XYRlDALES Xyridaceae 1 1
lariaceae and Mayacaceae of Commelinales, Maran- Rapataeaceae 1 1
taceae and Lowiaceae of Zingiberales, and Ponte- ERIOCA ULALES EriocauIaceae 3 3
dericeae of Liliales. ZINGIBERALES· Lowiaceae 1 1
The fourth type is limited only to the orders Marantaceae 2 2
closely related to Liliales (table 1, 2) and seems to LILIALES· Pontederiaceae 2 2
have arisen only once in evolutionary history. It oc- TYPHALES Sparganiaceae 1 3
curs in Liliales (except family Pontederiaceae and Typhaceae 1 1
HAEMODORALES· Haemodoraceae 10 10
genus Astelia of Liliaceae), Dioscoreales, Alstroe-
Hypoxidaceae 2 2
meriales, Amaryllidales, Iridales, Agavales (except Velloziaceae 1 1
genera Doryanthes and Xanthorrhoea of families PhiIydraceae 1 1
Agavaceae and Xanthorrhoeaceae respectively) . Bur- JUNCALES Juncaceae 2 3
manniales and Orchidales are characterized by this Centrolepidaceae 2 2
complex. In addition it is present in the families Restionaceae 3 3
Apostasiaceae and Taccaceae of Haemodorales and CYPERALES Cyperaceae 4 9
Lemnaceae of Arales. GRAMINALES Gramineae 50 50
January, 1961] STEBBINS AND KHUSH-STOMATA IN MONOCOTYLEDONS 55
with no subsidiaries, on the other hand, have pre- the evolution of the monocotyledons, and should
dominantly 2 vascular bundles in the cotyledon, be regarded as a character of minor importance in
and although type A germination is the most com- delimiting families. Nearly all of the family realign-
mon, types Band C are not infrequently found. ments which Hutchinson made on the basis of this
Finally, the group with many subsidiaries contains assumption are supported by the stomatal evidence.
mostly families with many vascular bundles in the The evidence from stomatal types also supports
cotyledon, the only deviating families being the in general the conclusion of Hutchinson that the
Commelinaceae with 2 and the Bromeliaceae with development of a biseriate perianth occurred rela-
1 bundle. Furthermore, hypogeal germination of tively few times in the phylogeny of the group, and
type C is the most common, with type B next in therefore should be given greater importance. All
frequency, while type A is found in this group only of the families placed by Hutchinson in his Division
in some of the Bromeliaceae and Zingiberaceae. Calyciferae have stomatal complexes with 2 sub.
An equally good or even better correlation exists sidiary cells except for the Bromeliales and many
between stomatal type and habit of growth. When Zingiberales, which have more than 2. Thus, the
the plants in tables 1-3 are classified according to hypothesis that the Zingiberales are parallel to and
life form, as recognized by Raunkiaer (1937), those not related or ancestral to the Orchidales is fully
with 2 subsidiary cells are found to consist almost supported by the present evidence.
entirely of hemicryptophytes, helophytes, and aquat- Finally, the present evidence supports the concept
ic types. Geophytes are completely lacking, and of Engler and Gilg (1924) and particularly of Hal-
phanerophytes are found only in the grasses (bam- lier (1912) regarding the series Spadiciflorae, con-
boos, Arundo, ete.}, the Velloziaceae, and Flagel- taining the Araceae, Lemnaceae, Palmaceae, Pan-
lariaceae. Among the species having no subsidiaries, danaceae, and Cyclanthaceae. All of these families,
on the other hand, geophytes are in the majority, except for the Lemnaceae, agree in having stomatal
with phanerophytes next in frequency. Hemicrypto- complexes with many subsidiaries. Furthermore
phytes are found only in some Liliaceae and Or- they agree with each other in the small size of their
chidaceae, while helophytes and hydrophytes are epidermal cells, except for certain probably derived
completely lacking from this group, except for the genera of Araceae with large, non-evergreen leaves,
much reduced Lemnaceae. Finally, the group with such as Lysichiton and Arisaema. In this latter re-
more than 2 subsidiaries contains a majority of spect they differ from the Commelinaceae and to a
phanerophytes, a few hemicryptophytes in Com- lesser extent the Bromeliacae, the 2 other families
melinaceae, Zingiberaceae, Cannaceae, and Araceae, with many subsidiaries, but do resemble the anom-
geophytes in Zingiberaceae and Araceae, and a alous, apparently primitive genus Scheuchzeria. The
single helophyte, Scheuchzeria. This correlation present authors, therefore, favor the retention of
with growth habit must be seriously considered in the Spadiciflorae as an interrelated group of orders,
any discussion of the phylogenetic progression of rather than scattering its components over various
stomatal types. branches of the Division Corolliferae, as is done by
CONNECTIONS BETWEEN STOMATAL TYPE AND CER- Hutchinson. 1£ this change is made, then the re-
TAIN REPRODUCTIVE CHARACTERISTIcs.-In discuss- maining members of the Corolliferae, except for
ing the association between type of stomatal com- some groups which will be discussed below, agree
plex and characteristics of the flower and inflores- in possessing stomatal complexes without subsidiary
cence, 4 characters should be considered, since they cells.
have figured prominently in various phylogenetic GEOGRAPHIC DISTRIBUTION OF THE STOMATAL
schemes proposed for the monocotyledons. These TYPES.-The geographic distribution of plants with
are separate carpels vs, syncarpous ovary, hypogyny the 3 types of stomatal complex shows a regular
vs. epigyny, uniseriate vs. biseriate perianth, and progression. The orders with no subsidiaries are
the development of a large spathe or bract sub- predominantly temperate, except for Amaryllidales,
tending the inflorescence. Dioscoreales, Burmanniales, and Orchidales. These
The first point to note is that all monocotyledons are obviously the most specialized families in the
having apocarpous gynoecia have stomatal com- group. Furthermore, the Amaryllidaceae are well
plexes with subsidiary cells. In most of these groups, represented in temperate as well as in tropical re-
such as the Alismatales and Butomales, their num- gions, while the bulk of tropical orchids, being
ber is 2, but Scheuchzeria has usually 4, and some epiphytic, are in growth habit the most specialized
of the palms, perhaps including those with nearly of their family. In this group, therefore, the trend
free carpels, have the same number. Nearly free of specialization among modern forms appears to
carpels are found also in primitive Liliaceae (To. go from temperate to tropical types.
fieldia) , which lack subsidiaries. The orders with 2 subsidiaries are of 3 types,
In respect to hypogyny vs, epigyny, which served helophytes or temperate hydrophytes, such as the
as a major diagnostic characteristic in the system Butomales, Alismatales, Juncaginales, and Potamo-
of Bessey (1915), the evidence from stomatal com- getonales; tropical or southern hemisphere families
plexes supports the conclusion of Hutchinson like Flagellariaceae, Xyridales, Eriocaulales, Zingi-
(1934) that epigyny has arisen repeatedly during bel'ales, and Haemodorales; and 2 widespread
.58 AMERICAN JOURNAL OF BOTANY [Vol. 48
families, Cyperaceae and Gramineae. The Grami- obstacle to the hypothesis that the group with no
neae are almost certainly tropical in origin (Bews, subsidiaries is monophyletic, and had its origin in
1929), and the same may be true of the Cyperaceae. primitive Liliales in the general affinity of the
'The marsh and water plants, although world wide, Heloniadeae and Narthecieae.
.are somewhat better represented in the tropics and On the other hand, the types with 2 subsidiaries
the southern hemisphere than in the north, so that and with more than 2 are much more closely inter-
this group shows a recognizable progression of connected. In the Zingiberaceae, Araceae, Palmae,
.specialization from tropical to temperate types. and Pandanaceae both types are found, and with
Finally, the group with many subsidiaries con- the relatively small sampling which has been made
.sists almost entirely of tropical or predominantly of these families their relative abundance cannot yet
tropical families, the only exception being Scheuch- be estimated. The data available, however, do sug-
zeria. This group, therefore, almost certainly origi- gest that the types with 2 subsidiaries are relatively
nated in the tropics, and only a few members of the advanced within their family. Amomum and Zingi-
Commelinaceae and Araceae have been successful ber both belong to the tribe Zingibereae, which
in temperate regions. Hutchinson places last in his synopsis of the Zingi-
The information now presented can afford a basis beraceae; Arisaema is in the Areae, the last of the
for deciding which of the types of stomatal develop- tribes of Araceae; while in the Palmae Calamus is
ment should be considered the most primitive, and a specialized climber and Caryota occupies a rela-
what have been the principal lines of phylogenetic tively advanced position in the family. This indi-
progression with respect to this character. The first cates that types with 2 subsidiaries are in general
point to emphasize is that, with a few exceptions, derived by reduction from those with several. Fur-
the orders belonging to the group which lacks ther evidence in favor of such a trend is the high
-subsidiary cells are all clearly related to each other epidermal/guard cell ratio of many of the types
.and, in respect to floral characteristics, can be re- with 2 subsidiaries. This indicates a greater amount
garded as one or a few related and parallel lines of of cellular differentiation in the epidermis.
increasing specialization starting with the more The types with more than 2 subsidiaries appear
primitive Liliales. The exceptions are as follows. likewise to be more primitive than those with none.
'The Lemnaceae are probably a much reduced The reasons for this assumption are as follows.
derivative of the Arales, which have lost their sub- First, they occur in a number of distantly related
-sidiary cells along with most of their other types of groups, some of which are obviously of a relictual
specialized cells. The Pontederiaceae, the only character (Scheuchzeria) and have primitive floral
family possessing subsidiaries which Hutchinson structures. Second, they are associated with a low
places in the Liliales, probably belong more cor- epidermaljguard cell ratio, and hence with a smaller
rectly next to the Commelinaceae, where they are amount of differentiation in the epidermis as a
placed by Engler and Gilg (1924) and perhaps whole. Third, the plants having several subsidiaries
should be included in the order Commelinales. are mostly evergreen phanerophytes, which are
Erdtman (1953) states that their pollen is most generally regarded as representing the most primi-
'similar to that of the Xyridales. Perhaps, therefore, tive type of life form (Raunkiaer, 1937). Fourth,
the Pontederiaceae should be placed in an order by they are chiefly tropical in distribution. Recent
themselves, between the Commelinales and Xyri- discussions of angiosperm phylogeny have empha-
dales. At any rate, they deserve further study. sized that available evidence strongly supports a
'Three genera of Liliales and Agavales, Astelia of tropical origin for most families of dicotyledons
the Liliaceae, Xanthorrhoea of the Xanthorrhoea- (Axelrod, 1952), and a similar origin for the mono-
-ceae, and Doryanthes of the Agavaceae, differ from cotyledons might be expected. Finally, the mono-
other genera of their families in possessing subsidi- cotyledonous stomatal type with several subsidi-
ary cells. When this fact was discovered, a particu- aries and a low epidermaljguard cell ratio is
lar effort was made to examine genera placed near closely similar to the haplocheilic type of stomatal
to them in Hutchinson's system, but all those ex- complex described by Florin (1931, 1933) in a
.amined turned out to have stomata of the Lilialean variety of gymnosperms, including Ephedra, Gink-
type, without subsidiaries. The systematic position go, and Cycadales as well as conifers. He considers
of these 3 genera, one of widespread antarctic dis- this to be the most primitive type of stomatal com-
tribution and the 2 others endemic to Australia, plex in the class. The writers believe, therefore,
deserves further investigation. Finally, the Apostasi- that the haplocheilic type in primitive monocotyle-
aceae and Taccaceae, placed by Hutchinson in the dons is directly descended from the same type in
Haemodorales, are without subsidiaries, while the some as yet unidentified pre-angiosperm ancestor,
remainder of the Haemodorales possess them. This rather than from any form similar to moderildi-
·suggests that the arrangement of Engler and Gilp;, cotyledons.
which places the Apostasiaceae as an aberrant off- The present evidence suggests also that the types
·shoot of Orchidaceae and the Taccaceae next to the with 2 and with no subsidiaries are independently
Dioscoreaceae, may be more nearly correct. None derived from the primitive type with several. The
-of these exceptions, therefore, is an insuperable principal evidence for this hypothesis is their asso-
January, 1961] STEBBINS AND KHUSH-STOMATA IN MONOCOTYLEDONS 59
ciation with quite different life forms. One might value. On the other hand, the frequent association
suggest that the type with 2 well-defined subsidi- of no subsidiaries with geophytes and xeric rosette
aries, as found in Alismataceae, Eriocaulaceae, plants like the Agavaceae and Aloinae suggests
Xyridaceae, Juncaceae, Cyperaceae, and Gramineae, that this type of stomatal complex may have origi-
became first established in association with the nated under relatively dry conditions. In both
marsh or aquatic habit, in which it predominates. instances, the specialized stomatal complex, once
The function of the subsidiary cells is not known, developed, probably acquired a selective value
but their consistent presence in these families sug- which made it difficult to modify by mutation and
gests that they must serve some function, particu- selection, so that it persisted after the evolutionary
larly in view of the fact that stomata without sub- line bearing it had changed its adaptation radically
sidiary cells are rather easily induced in these types from the habitat in which the initial evolution of
through abnormal environments during ·develop- the stomatal type took place.
ment (Stebbins et aI., unpubl.}, and might be ex- DEPARTMENT OF GENETICS
pected to occur naturally "through mutation unless UNIVERSITY OF CALIFORNIA
such mutants had. a strongly negative selective DAVIS, CALIFORNIA
LITERATURE CITED
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