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The Stem and Root Anatomy of Sanmiguelia lewisii, and a Comparison with
Extant Dicots and Monocots

Presentation · March 2003

DOI: 10.13140/RG.2.1.3232.3446

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The Stem and Root Anatomy of Sanmiguelia lewisii,

and a Comparison with Extant Dicots and Monocots
by Bruce Cornet

page 1

Abstract. Very well preserved remains of Sanmiguelia lewisii were found in the
upper Trujillo Formation (Late Triassic: late Carnian-Norian) of northwest Texas in
1980-1981, and published on in 1986 and 1989. Because Sanmiguelia was preserved
in growth position, buried in stages while living, its fossils preserve many if not most
of the important morphological and anatomical characteristics of this plant. Its stem
and root anatomy, as well as its overall growth habit, are described. Its unusual
combination of monocot and dicot characteristics, some of which are no longer found
in either Class today, are compared with herbaceous monocots and dicots, as well as
with vines and lianas. It is concluded that Sanmiguelia was either 1) a primitive
monocot evolving towards a dicot, or 2) a primitive dicot evolving towards a
monocot, or more likely 3) an extinct type of angiosperm that occupied an
intermediate position between monocots and dicots. Sanmiguelia had to undergo
significant morphological and anatomical changes in order to give rise to the types of
dicots that were involved in the Cretaceous radiation of angiosperms. Finally,
Sanmiguelia is placed into morphologic and geologic perspective in order to show
why pre-Cretaceous angiosperms were not more abundant and diverse.

Table of Contents

Page 1

Preparation for Discussion

Sanmiguelia Stem Anatomy

Growth Habit

Habitat and Adaptation

Comparison of Sanmiguelia with Non-Arborescent Angiosperms

Herbaceous Rosettes (extant)

1
Herbaceous Erects (extant)

Vines and Lianas (extant)

Glossary

Page 2

Comparison of Sanmiguelia with Arborescent Monocots

What are the shared characteristics?

Hypothetical Transformation of Sanmiguelia

Burger's 1981 hypothesis that monocots came first

Discussion

Conclusions

References

Glossary

Source publications

http://bcornet.tripod.com/evoltheo/Slewisii.htm
http://www.sunstar-solutions.com/sunstar/evoltheo/slewisii.htm

http://bcornet.tripod.com/evtrend/Sanmig2.htm
http://www.sunstar-solutions.com/sunstar/evtrends/sanmig2.htm

Preparation for Discussion

A leaf gap is an area along the stem where phloem and xylem diverge from the
vascular cylinder and enter the base of a leaf. This process results in a temporary gap
in the vascular cylinder, called a leaf gap.

2
From Esau (1961: p. 209).

Angiosperms possess several types of leaf gaps or nodal anatomy, ranging from
unilacunar and trilacunar, to multilacunar.

Dicot stem anatomy.

3
From Esau (1966: p. 212).

Sanmiguelia Stem Anatomy

No extant angiosperm possesses all three conditions: 1) a woody cylinder of
secondary xylem like that found in dicots, 2) a multilacunar node, and 3) parallel-
veined leaves with clasping/sheathing leaf bases. Sanmiguelia is unique in that it
possessed all three conditions. Magnolia spp. possess a woody cylinder (#1) and
multilacunar nodes (#2), but have simple pinnately-veined leaves with
petioles. Monocots possess multilacunar nodes (#2) and parallel-veined leaves with
4
clasping leaf bases (#3), but no secondary xylem or woody cylinder. According to the
new taxonomy (Fig. 7: Doyle and Endress, 2000), the Monocotyledonae and
Magnoliales must chronologically post-date the combination of conditions found in
Sanmiguelia. In the Magnoliales (MAG) condition #3 was lost, while in the
Monocotyledonae (MON) condition #1 was lost. Note that in monocots, the
Xanthorrhoea, Yucca, and Dracaena (Xanthorrhoea, Agavaceae, and Dracaenaceae)
evolved a unique type of secondary thickening cambium with secondary "wood"
containing crowded vascular bundles, but structurally it is not homologous with the
secondary wood of Sanmiguelia or of dicots.

Stylized cross section of Sanmiguelia lewisii stem anatomy, based on Cornet (1986).

5
 The path of primary xylem bundles can be seen along a pith cast (highlighted in the

image on the right). The leaf gaps are preserved along the
outer "surface" of the pith cast; they are lenticular and quite long (8 cm). The top and
bottom of adjacent leaf gaps are visible in the pith cast. The vascular traces appear to
arise from primary xylem bundles progressively higher (centrifugally) as the leaf gap
widens. Thus, the central or primary veins of a leaf arose first along with any branch
traces, while traces that supply the lateral veins of Sanmiguelia leaves arose
progressively higher in the leaf gap, consistent with the anatomy of sheathing leaf
bases.

The central pith began small at the base of vertical (aerial) stems, and grew in width
upwards as the thickness of secondary xylem decreased. After about 14-15 cm the
pith occupied the entire width of stem casts (see below), indicating that secondary
xylem growth decreased upwards. This fact is important, because it corresponds to
the distribution of functional leaves along the upper part of vertical stems (see below;
cf. Tidwell et al., 1977).

The leaf and branch traces are circular, have a small central pith, and a ring of
xylem. They possessed an outer ring of phloem (although deformed by constriction
within the leaf gap), as well as a circular cambium, which were separate from any
cambium and phloem associated with the outer woody cylinder (Paleozoic
pteridosperms, such as Medullosa, possessed multiple or sequential cambia: Taylor
and Taylor, 1993). The rationale for saying this is the ability of the circular xylem
traces to divide within the multiseriate rays. It would also be necessary for the branch
and leaf traces to grow and stretch to at least some degree with increase in stem
diameter. Whereas leaves in most dicots dehisce and fall off before secondary growth
of branches begins, that was not the case for Sanmiguelia. It required a stiff support
structure for its large leaves, which persisted until their vascular supply was cut off by
secondary xylem growth in the outer ring. The fact that this secondary wood
consisted mostly of thick-walled fiber cells indicates that its function was primarily
for support rather than water conduction.

6
 The thin sections below were taken from secondary wood preserved in situ,
between a pith cast and the cast of a long sheathing leaf base. The specimen
illustrated below came from near the base of an aerial (vertical) stem, where the pith
cast was narrow and preserved secondary wood development thickest. The brown
color of the wood is natural. The space between the sheathing leaf base and stem
apparently widened following the death of the axis, allowing sediment to infiltrate
during early stages of decomposition, sedimentary compaction, and diagenesis.

Tangential cross section.

7
8
 The secondary wood consisted of rows of tracheids, fiber cells, and
parenchyma. Uniseriate rays were common. Multiseriate rays occurred at regular
intervals in the wood, and functioned as leaf gaps for the multiple leaf traces in
sheathing leaves.

9
Enlargement of secondary xylem showing a uniseriate ray and thick-walled fiber cells.

10
 In tangential section, the secondary xylem reveals the presence of numerous cell
inclusions, which are interpreted as fungal spore resting bodies. Arrows below point
out circular-bordered pits in tracheids.

11
 Either they are fungal spores/cysts, or some of cell inclusions may represent
tyloses, which are specialized growths of adjacent (companion) parenchyma cells into
tracheid lumina that seal off the wood and make it non-water-conducting. Tyloses in
extant angiosperms occur only in vessels, and form in older wood in order to seal off
water-conducting vessels. Tylosis development produces heartwood, which has the
property of being more water resistant (used in wooden barrel making). If tracheid
inclusions in Sanmiguelia can be shown to represent tyloses (as opposed to fungal
contaminants), either the origin of tyloses in angiosperms would predate vessel
evolution, or the ancestor of Sanmiguelia possessed vessels.

Once a secondary xylem cylinder began to enlarge, vascular traces to leaves and
branches would eventually be stretched to the breaking point, causing the leaves and
branches on the older part of the stem to die. Evidence for this can be seen in the
preserved axes of Sanmiguelia, which lack apparent branching, except where damage
has occurred (see pith cast below and illustration 'a' - S4 below). The fact that vertical
axes branch at the rhizome instead of higher on those axes is another indication that
branching probably did not occur on the lower woody parts of vertical axes
(illustration 'b', below).

The condition as it exists in extant dicots.

12
From Essau (1966: p. 244).

Leaves are regularly sacrificed (deciduous) with stem growth in dicots, but not
branches. That does not appear to be the case with Sanmiguelia, which had sheathing
leaves. Sheathing leaves in monocots do not have an abscission zone which allows a
leaf to separate from the plant. The combination of monocot and dicot vegetative and
anatomical features gave Sanmiguelia a hybrid-like appearance, but also limited its
potential variability. Most of the Cretaceous radiation of angiosperms was a dicot
radiation, which makes one wonder whether the evolution of the monocot type of leaf
was unrelated to the Cretaceous radiation.

13
 Growth Habit
Sanmiguelia was clonal, monoecious, and bore separate unisexual male and female
inflorescences on the same plant or on separate vertical axes arising from a common
underground rhizome (Cornet, 1986). This interpretation is based on the fact that only
one individual plant (vegetative colony) was excavated, which had both types of
inflorescences in close proximity to in situ axes or had leaves, stems, and flowers in
organic connection. Branches were restricted to the upper region of the main stem,
and functioned mainly in reproduction: That is, inflorescences were ephemeral or
seasonal. Thus, those types of branches could be sacrificed in Sanmiguelia once the
reproductive season was over.

14
Figure 8 from Cornet (1986), showing Axelrodia burgeri (female) on left and Synangispadixis
tidwellii (male) on right.

A large apocarpous flower or fruit containing seeds (Nemececkigone fabaforma

Cornet) was found caught (preserved) between the nested rachises of Cladophlebis

15
macrophylla fronds, which radiated outwards from a common rhizome, indicating not
only that the fern grew alongside Sanmiguelia, but that it was probably situated
directly below an inflorescence at the time the fruit dropped.

16
17
Above: Cladophlebis macrophylla, which grew alongside Sanmiguelia.

We know that the growth form (habit) of Sanmiguelia was both sympodial and
monopodial. The rhizomes and stolons branched by dichotomy. One branch could
generate a bipolar axis that developed into aerial stem and primary root
system. Branches on aerial axes came from the axils of leaves or bracts (i.e. axillary
branches: see Cornet, 1986; 1989a). And yet axillary branching was limited, and
seems to have functioned mainly in reproduction or in the repair of broken stems (see
below).

Sanmiguelia may have resembled an herbaceous version of arborescent monocots

such as the palms, or of arborescent Carboniferous lycopods such as Lepidodendron,
where the leaves and branches were clustered at the tops of solitary stems, which
arose from an underground rhizome (see Cornet, 1986 and figures below).

Sanmiguelia had a primary root system with apical meristem that produced the
aerial axes, and lateral meristems that produced the rhizome (underground stem) and
stolons (thinner runners). This can be seen in the specimens preserved in growth
position (illustrated below). Aerial stems arose from the primary root, while roots
extended and branched below it. No evidence for a single taproot was found,
however. One stem appeared to have two primary roots (each with secondary roots)
extending below it, although one of them could have belonged to another stem
concealed in the outcrop. Stolon or rhizome meristems could generate a new primary
root with bipolar growth (illustrated below).

The rhizome and primary root apex apparently had the ability to generate new
aerial axes if adjacent ones became broken or damaged (see below). However, if an
aerial axis broke above a leaf gap, a secondary or axillary branch could develop and
restore the lost part of the stem (see diagram below).

Some of the roots below the central stem to the right of 'a' in Fig. 2 of Cornet
(1986) below were silicified in places. Thin sections show tracheids varied
considerably in width up to the size of small vessels. Sidewall pitting was
crowded. An end plate to a vessel-like element was found in a cross section. It had
scalariform pitting (Cornet, 1986: Plate 3-e).

18
 Pith cast showing a broken or terminated axis being extended through an axillary
branch, which looks like a stem dichotomy.

19
Below: Stem, rhizome, and root casts / petrifactions of Sanmiguelia preserved in
growth position (from Cornet, 1986). See overview of outcrop location.

20
Figure 2 from Cornet (1986).

Vertical axes arose from the dichotomy of an underground rhizome or stolon.

21
Multiple axes originated asynchronously from the same area of a rhizome, indicating
monopodial growth from a persistent meristem.

22
Aerial axes were found connected by rhizomes or stolons in the soil.

23
Functional leaves were concentrated towards the upper parts of vertical stems.

24
Modified after Tidwell et al, 1977.

According to Groff and Kaplan's (1988) classification scheme for clonal growth,
Sanmiguelia was a Class 2 plant. It could not have developed into a Class 1 plant,
characteristic of arborescent dicots.

25
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Habitat and Adaptation

Sanmiguelia appears to have been well-adapted for its paleoenvironmental
niche. From the excavation of a portion of an in situ plant colony (figured above; see
also Cornet, 1986), vertical axes arose from an underground rhizome (horizontal
26
stem). The habitat was the margin of a shallow oxbow lake on a floodplain (Cornet,
1986), which was apparently frequently flooded until the lake was completely filled
in. Silty to sandy sediment indicative of crevasse splays slowly buried the lower
portions of vertical axes during yearly floods. Sanmiguelia may even have grown in
standing water, similar to Sagittaria sagittifolia, which grows in ephemeral
ponds. And like Sagittaria, its lower leaves may have grown longer and narrower
under water (Arber, 1961: p. 129), explaining the variation in leaf form found at the
Sanmiguelia locality in Texas (Cornet, 1986; Cornet, 1989a).

However, occasionally severe flooding broke some of the stems off. When that
happened, new axes emerged from the underground rhizome. The growth and
development of the outer vascular ring may have helped new axes push up through
layers of sediment, although extant angiosperms with subterranean horizontal stems
(rhizomes), such as Cladium mariscus (Cyperaceae - a grass), can do that quite well
with only leaf meristems in moderately damp soil. Some time later, another flood
would break off those stems at the sediment/water interface. Eventually the rhizomes
could not regenerate vertical axes due to depth of burial, and that portion of the colony
died. Thus, Sanmiguelia was a perennial that lived in a semiaquatic environment
which was subjected to high energy abrasion and stress from silt- and sand-laden
floodwaters. Its stiff outer vascular cylinder would have provided resistance to
bending and breaking, as well as protection to the primary xylem bundles next to the
pith. The amount of woody growth in the stems that survived longest indicates that
Sanmiguelia may have lived for decades before finally succumbing to burial. The
amount of leaf debris, fallen flowers and inflorescences, and seeds in the surrounding
sediments supports that interpretation.

Whether Sanmiguelia's stellar anatomy was molded by the environment, or it lived

there because it could survive those conditions is not known (i.e. did its ancestry pre-
adapted it to that type of environment?). It is because of its habitat that we have so
much of this plant preserved.

Comparison of Sanmiguelia with Non-Arborescent

Angiosperms
The combination of parallel-veined leaves, leaves with long sheathing bases, and a
woody cylinder limited Sanmiguelia's growth form, and therefore its evolutionary
potential. When compared to angiospermous vines, herbaceous rosettes (leaves
concentrated in a basal spiral or whorl), and herbaceous erects (aerial axes with
leaves, branched and unbranched), significant similarities and differences
emerge. The tables below contain selected examples from each of the above
categories. They were selected in order to show the range of variation as well as

27
variation in combinations of identified characteristics. For example: Most herbaceous
monocots have parallel-veined leaves without petioles and with clasping or sheathing
leaf bases. Few herbaceous dicots have parallel venation, although some, such as
Gentiana quinquefolia, approach the monocot condition. A few herbaceous monocots
have leaves which approach the dicot condition: palmately or pinnately lobed and
veined, abundant cross veins with some reticulation, petioles, and clasping rather than
sheathing leaf bases. Most herbaceous monocots have a tuberous or bulbous root
system, or have a rhizome (subterranean stem) if they do not have tubers or bulbs. By
comparison, most herbaceous dicots do not produce tubers or bulbs, while those
which are thought to be more closely related to primitive angiosperms, such as
buttercups (Ranunculaceae) and water lilies (Nymphaeaceae), commonly have
rhizomes (Fernald, 1970; Newcomb, 1977).

Herbaceous angiosperms rarely have woody stems, mainly because they are either
annuals or die back to underground bulbs, tubers, or rhizomes at the end of each
year. Secondary growth is irrelevant or even disadvantageous (energy
inefficient). Those herbaceous angiosperms that do have woody stems are typically
perennials, and this includes some monocots. Yet the type of secondary growth in
monocot stems is very different from that in woody dicots, and is apparently of
secondary origin or evolution (Arber, 1961). Herbaceous monocots with woody
stems are highlighted in red with an asterisk below.

Herbaceous rosettes have their leaves concentrated in a tight spiral or whorl at

ground level, with upright extensions of the stem being limited to inflorescences.

Herbaceous Rosettes (extant)

secondar sheathin
leaf
y leaf venation petiol phyllotax g- bulb or
Species Class teet
growth - type type e y clasping tuber
h
woody base
Acorus monoc no sword- no parallel no tight yes no,
calamus ot shaped basal rhizome
spiral
Allium monoc no long no parallel no tight yes bulb
canadense ot very basal
narrow spiral
Arisaema monoc no pinnate no pinnate yes tight yes, tuberous
atrorubens ot 3-lobed basal sheathin
spiral g
Cacalia dicot no elliptical no palmatel yes tight no tuberou
tuberosa y basal s

28
 parallel spiral
Cladium monoc no long no parallel no tight yes, culm
mariscus ot very basal sheathin short
narrow spiral g triangula
r
Dodecathe dicot no elliptic no pinnate yes tight no no
on oblong basal
meadia spiral
Draba dicot no elliptical no pinnate no tight no no,
verna basal stolons
spiral
Dicentra dicot no pinnately no pinnate yes tight no granulat
cucullaria dissected basal e
spiral bulb
Erythroniu monoc no narrow no parallel no tight yes, scaly
m ot lance- basal sheathin bulb
albidum shaped spiral g
Galax dicot no round- yes palmate yes tight no no,
aphylla chordate basal rhizome
spiral
Goodyera dicot no elliptical no pinnately yes tight no no
pubescens palmate basal
spiral
Hemerocall monoc no long no parallel no tight yes, tuberous
is ot sword- basal sheathin
fulva shaped spiral g
Hydrocotyl dicot no round no palmate yes tight no no
e basal
umbellata spiral
Lobeila dicot no narrow no pinnate no tight no no
dortmanna or basal
cylindric spiral
al
Nuphar dicot no round- no palmate yes tight no tuberou
variegatum chordate basal s
spiral
Pinguicula dicot no elliptical no pinnate no tight no no
vulgaris basal
spiral
Plantago dicot no elliptical no palmatel yes tight no no
major y basal
parallel spiral

29
 Pyrola dicot no ovate or yes pinnate yes tight no no
asarifolia egg- basal
shaped spiral
Ranunculus dicot no dissected no palmatel yes tight no bulb
bulbosus 3-pinnate y basal
pinnate spiral
Ranunculus dicot no very no pinnate yes tight no no,
reptans narrow basal stolons
lance- spiral
shaped
Sagittaria monoc no arrow- no pinnatel yes tight yes, tuberous
latifolia ot shaped y basal sheathin ,
parallel spiral g stolons
Saxifraga dicot no lance- yes pinnate yes tight no no
pensylvanic shaped basal
a spiral
Spiranthes monoc no lanceolat no parallel no tight yes, thick
cernua ot e basal sheathin tuberous
spiral g
Spiranthes monoc no elliptical no parallel no tight yes, thick
tuberosa ot basal sheathin tuberous
spiral g
Talinum dicot no slender no pinnate no tight no no
teretifolium cylindric basal
al spiral
Trifolium dicot no pinnate yes pinnate yes tight no no,
repens 3-lobed basal stolons
spiral
Viola dicot no kidney- yes palmate yes tight no no
renifolia shaped basal
spiral
Yucca monoc yes* long no parallel no tight yes, tuberous
filamentosa ot lance- basal sheathin
shaped spiral g

Herbaceous erects have their leaves distributed along a vertical stem, but in
addition can also have leaves in a tight spiral or whorl at ground level. Flowers and
inflorescences are born terminally at the end of the main stem, or in the axils of leaves
along the stem. In addition, the stem can be branched or unbranched.

30
 Herbaceous Erects (extant)
secondar sheathin
venatio bulb
y leaf leaf petiol phyllotax g-
Species Class n or
growth - type teeth e y clasping
type tuber
woody base
Acnida dicot no lance- no pinnate yes alternate no no
cannabina shaped
Asclepias dicot no lance- no pinnate yes opposite no no
viridiflora shaped and
no
Asparagus monoc no scale- no parallel no spiral yes, no,
officinalis ot like clasping rhizom
e
Anthemis dicot no dissecte no pinnate no alternate no no
arvensis d
Cardamine dicot no ovate or wavy palmate yes alternate no bulb
bulbosa lance- toothe
shaped d
Coreopsis dicot no lance- no pinnate yes whorled no no
major shaped
Epipactis monoc no lance- no parallel no spiral yes, no
helleborine ot shaped clasping
Gentiana dicot no lance- no parallel no opposite yes, no
quinquefolia shaped clasping
Gerardia dicot no pinnatel no pinnate yes opposite no no
virginica y
lobed
Helianthus dicot no lance- yes pinnate yes opposite no no
divaricatus shaped
Hibiscus dicot no egg- yes palmate yes alternate no no
palustris shaped
or 3-
lobed
Hieracium dicot no elliptical yes pinnate no alternate no no
gronovii or
egg-
shaped
Lactuca dicot no deeply yes pinnate yes alternate no no
biennis lobed and

31
 no

Lilium monoc no lance- no parallel no spiral yes, bulb

tigrinum ot shaped clasping
Lilium monoc no lance- no parallel no whorled yes, bulb
superbum ot shaped clasping
Mertensia dicot no chordate no pinnate yes alternate no no
virginica or and
heart- no
shaped
Mitchella dicot no ovate or no pinnate yes opposite no no,
repens roundish stolons
Polygonatu monoc no lance- no parallel no spiral yes, no,
m ot shaped alternate clasping rhizom
pubescens e
Ranunculus dicot no palmatel yes palmate yes alternate no no
hispidus y
lobed
Rumex dicot no lance- no pinnate yes alternate no no
orbiculatus shaped
Silene dicot no lance- no pinnatel yes whorled no no
stellata shaped y
parallel
Sparganium monoc no linear no parallel no spiral yes, bulb
eurycarpum ot sword- sheathin
shaped g
Stellaria dicot no ovate or no pinnate yes opposite no no
media roundish
Typha monoc no linear no parallel no spiral yes, tuberou
latifolia ot sword- sheathin s
shaped g
Uvularia monoc no lance- no parallel no spiral yes, no,
perfoliata ot shaped alternate clasping rhizom
e
Veratrum monoc no large no parallel no spiral yes, bulb
viride ot lance- sheathin
shaped g
Verbascum dicot no lance- no pinnate no alternate yes, no
thapsus shaped clasping
Xerophyllu monoc no very no parallel no spiral yes, tuberou
m ot narrow sheathin s

32
asphodeloid sword- g
es shaped

Now take a look at angiospermous vines and lianas in the table below. Dicot vines
are much more common than monocot vines. Those monocots that have a vine-type
growth habit (e.g. Dioscorea and Smilax) are unique. Not only are the leaves of these
plants dicot-like in shape and major venation, but they have petioles and lack
sheathing or clasping leaf bases typical of most monocots. The leaves of Smilax
dehisce (fall off) during winter, like those of deciduous dicots. These monocot plants
delivered or produced a unique type of secondary wood or stem thickening (Arber,
1961). Smilax rotundifolia (common greenbrier) even produces thorns like that of
some dicot shrubs and vines. Secondary growth and tendrils in dicot vines are not
necessary, however, as the table indicates.

Vines or Lianas (extant)

secondar sheathin
leaf
y leaf venation petiol g- tendril
Species Class teet stipules
growth - type type e clasping s
h
woody base
Aristolochia dicot yes chordate- no palmate yes no no no
durior ovate
Bignonia dicot yes lance- no pinnate yes no no yes
capreolata ovate
paired
Campsis dicot yes pinnately yes pinnate yes no no aerial
radicans compoun roots
d
Cynanchum dicot no lance- no pinnate yes no no no
nigrum ovate
Dioscorea monoco yes* chordate no palmatel yes no no no
villosa t or y
heart- parallel
shaped
Echinocystis dicot no palmatel no palmate yes no no no
lobata y
3-lobed
Ipomoea dicot no heart- no palmate yes no no no
lacunosa shaped
or 3-
lobed
Lonicera dicot yes oblong no pinnate- no no no no

33
sempervirens paired palmate
Menispermu dicot no peltate no palmate yes no no no
m 3-7 lobed
canadense
Rhus dicot yes palmatel yes palmatel yes no no aerial
radicans y y roots
3-pinnate pinnate
Parthenociss dicot no palmatel yes palmatel yes no no yes
us y y w/discs
quinquefolia 5-pinnate pinnate
Rubus dicot no pinnately yes pinnate yes yes no no
pubescens compoun
d
Smilax monoco yes* peltate no pinnatel yes modifie no stipula
rotundifolia t y d r
palmate
Solanum dicot no palmatel no pinnate yes no no no
dulcamara y
lobed
Vitis dicot yes 3-lobed yes palmate yes no no yes
labrusca

In comparison to extant angiosperms, Sanmiguelia comes closest to the habit of

Veratrum viride and V. album (Liliaceae) in overall size and form (Tidwell et al,
1977), but Veratrum has a false stem (pseudostem or pseudo-axis) constructed from
sheathing leaf bases (Arber, 1961).

34
35
Modified from Arber (1961: p. 57).

Continued on page 2
References on page 2

Glossary
amphivasal: amphi for around or on both sides; vas L. for vessel or vasculature.

angiosperm: Any plant of the class (Angiospermae) having the seeds in an enclosed
ovary.

arborescent: Resembling a tree in growth structure or appearance; tree-like.

bulb: A bud, usually underground, consisting of a short, thick stem sending out roots
from below, and beqring overlapping, scalelike leaves, as in the lily, onion, or tulip; a
fleshy tuber or corm resembling a bulb, as in a dahlia bulb.

bundles: A general reference to vascular tissue organized into discrete circular to

elliptical or lunate-shaped (cross section) clusters of fluid-conducting cells (xylem and
phloem) that pass like "rods" vertically through the stem.

cambium: A meristem with products of division arranged orderly in parallel files.

Preferably applied only to the two lateral meristems, the vascular cambium and the
cork cambium or phellogen. Consists of one layer of initials and their undifferentiated
products, or derivatives.

corm: A short, bulblike, underground, upright stem, invested with a few thin
membranes or scale leaves, as in the crocus or gladiolus.

cutin: A wax-like highly complex fatty substance present in plants as an impregnation

of epidermal walls and as a separate layer, the cuticle, on the outer surface of the
epidermis. Makes the walls more or less impervious to water.

dioecious: A condition in which ovule- and pollen-bearing structures are borne on

separate plants; all dioecious plants have unisexual reproductive structures (contrast
monoecious).

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dicotyledonous (dicot): An angiosperm embryo having two cotyledons (first leaves); a
member of the dicots.

disulcate: A pollen type having two (normally distal) sulci.

ephemeral: Short-lived, transient.

eudicot: First or earliest record of higher extant dicots.

floodplain: The flat land on either side of a river or stream which has been shaped by
water erosion and sedimentary deposition.

habit: The form a plant takes, such as a tree, shrub, erect herb, rosette herb, aquatic
herb, etc.

herbaceous: A plant having little or no secondary development; nonwoody.

interfascicular cambium: Vascular cambium giving rise to secondary xylem and

interfascicular rays.

intrafascicular cambium: Vascular cambium restricted to a vascular bundle, or

fascicle; not connected with cambia of other vascular bundles.

leaf gap: In the nodal region of a stem. A region of parenchyma in the vascular
cylinder occurring where a leaf trace is bent away from the vascular system of the
stem toward the leaf.

leaf trace: A vascular bundle in the stem extending between its connection with a leaf
and that with another vascular unit in the stem. A leaf may have none or more traces.
Sometimes the whole complex of traces of one leaf is called a leaf trace.

meristem: Tissue primarily concerned with protoplasmic synthesis and formation of

new cells by division - responsible for stem, root, and leaf growth.

microphyllous: A plant having small bract-shaped leaves characteristic of arid and

semiarid habitats.

monocotyledonous (monocot): An angiosperm embryo having only one cotyledon

(first leaf); a member of the monocots.

monoecious: Bearing both pollen- and ovule-producing organs on the same plant; the
organs can be either unisexual or they can be bisexual (both types in a single
fructification )(contrast dioecious).
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monopodial: Stem or main axis arising from a terminal meristem.

multilacunar node: multi L. for many, more; lacuna L. for cavity, hollow, cavern; in
reference to the number of spaces or gaps in a vascular cylinder for leaf and branch
vascular traces.

niche: A unique environmental area that provides all of the essential physical and
chemical elements for a plant or animal to survive, grow and reproduce.

node (nodal): That part of the stem at which one or more leaves are attached. Not
clearly delimited anatomically.

palmate: Branching outwards from a common point, like fingers on a hand.

periderm: Secondary protective tissue derived from the phellogen (cork cambium) and
replacing the epidermis, typically in stems and roots. Consists of phellem (cork),
phellogen, and phelloderm.

petiole: The slender stem that supports the blade of a foliage leaf; a leafstalk.

phellem (cork): Protective tissue composed of nonliving cells with suberized walls.
Replaces the epidermis in one-year and older stems and roots of many plants and
formed by the phellogen (cork cambium). Part of the periderm.

phelloderm: A tissue formed by the phellogen in the opposite direction of the cork.
Resembles cortical parenchyma. Part of the periderm.

phellogen (cork cambium): A lateral meristem forming the periderm, a protective

tissue common in stems and roots of dicotyledons and gymnosperms. Produces
phellem (cork) toward the surface of the plant, phelloderm toward the inside.

phloem: The principal food-conducting tissue of the vascular plant, basically

composed of sieve elements, parenchyma cells, fibers, and sclereids.

phylogeny: The race history of an animal or plant type (contrast ontogeny).

phyllotaxy: The arrangement of leaves on a stem.

pinnate: Branching at right angles as in a fern frond.

procambium: Primary meristem or meristematic tissue which differentiates into the

primary vascular tissue. Also called provascular tissue.

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primary xylem: Xylem tissue differentiating from procambium during primary growth
and differentiation of a vascular plant. Commonly divided into the earliest protoxylem
and the later metaxylem. Not differentiated into axial and ray systems.

protoxylem: The first formed elements of the xylem in a plant organ. First part of the
primary xylem.

radiation: An evolutionary/genetic process by which diversity rapidly increases and

new species, genera, and families of animals or plants evolve from one or more
closely closely-related ancestors; if the radiation continues long enough (millions of
years), new orders, subclasses, and classes of animals and plants can evolve.

rhizome: horizontal stem, whether lying on the ground (prostrate) or growing below
ground.

riparean: stream banks.

scalariform pitting: Elongate pits arranged parallel so s to form a ladder-like

(scalariform) pattern.
selfing: The reproductive process by which a plant fertilizes itself.

sessile: Attached directly by the base; not raised upon a stalk or petiole.

sheathing base: Applied to the base of a leaf, either sessile or petiolate, when the leaf
base encircles the stem.

stelar: Relating to or of the stele.

stele: conceived by Van Tieghem as a morphological unit of the plant body

comprising the vascular system and the associated ground tissue (pericycle,
inerfascicular regions, and pith). The central cylinder of the axis (stem and root).

stolon: a runner or rootstock used to propogate certain grasses.

suberin: The same definition as for cutin with which it is closely related.

suberization (suberized): Impregnation of the wall with suberin or deposition of

suberin lamellae in the wall.

sympodial: The apparent main stem does not develop from a terminal bud, but is
made up of successive secondary axes, each of which represented one branch of a
dichotomy, the other branch being a weaker growth that gives rise to specialized axes
such as inflorescences.
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 tracheary element: General term for a water-conducting cell, tracheid or vessel
member.

trilacunar node: tri L. for three; lacuna L. for cavity, hollow, cavern; in reference to
the number of spaces or gaps in a vascular cylinder for leaf and branch vascular
traces.

tuber: A potato-shaped swelling of a root, which serves as a storage organ for

nutrients.

unilacunar node: uni L. for one; lacuna L. for cavity, hollow, cavern; in reference to
the number of spaces or gaps in a vascular cylinder for leaf and branch vascular
traces.

vascular cylinder: A term of convenience applied to the vascular tissue and associated
ground tissue in stem or root. Refers to the same part of stem or root that is designated
stele but without the theoretical implications of the stelar concept. Same as central
cylinder.

vascular: Possessing specialized water-conducting cellular elements.

vascular trace: water- and food-conducting tissue and associated ground tissue passing
from the stem to a leaf or branch.

xylem: The principal water-conducting tissue in vascular plants characterized by the

presence of tracheary elements. The xylem may be also a supporting tissue, especially
the secondary sylem (wood).

The glossary was created with the help of glossaries and definitions in the following
books:
Anatomy of Seed Plants by Katherine Esau (1966)
Composition of Scientific Words by Roland W. Brown (1978)
Webster's New Collegiate Dictionary (1961)

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